SPECIES STATUS ASSESSMENT Marron bacora (Solanum conocarpum)

VERSION 1.0

Photo by: USFWS

April 2019

U.S. Fish and Wildlife Service Southeast Region Caribbean Ecological Service Field Office Boquerón, Puerto Rico

Solanum conocarpum Species Status Assessment 2019

This document was prepared by Jaime Yrigoyen, Omar Monsegur, Maritza Vargas, José Cruz-Burgos, and Marelisa Rivera (U.S. Fish and Wildlife Service - Caribbean Ecological Services Field Office) with assistance from Andreas Moshogianis, Kaye London and Erin Rivenbark (U.S. Fish and Wildlife Service - Southeast Regional Office).

Valuable peer review of a draft of this document was provided by Jennifer Possley (Conservation Program Manager, Fairchild Tropical Botanic Garden).

Suggested citation:

U.S. Fish and Wildlife Service. 2019. Species status assessment of Marron bacora (Solanum conocarpum), Version 1.0. U.S. Fish and Wildlife Service Southeast Region, Caribbean Ecological Services Field Office, Boquerón,

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Solanum conocarpum Species Status Assessment 2019

EXECUTIVE SUMMARY

Marron bacora is a rare, tropical dry forest shrub that was once thought to be only found on St. John in the U.S. Virgin Islands; an additional population was discovered on Tortola in the British Virgin Islands in 2018. When the species was petitioned to be listed in 1996, it was listed as a candidate species, Solanum conocarpum, February 22, 2011 (76 FR 9722).

When the US Fish and Wildlife Service (Service) was petitioned to list the species, there were only two known wild individuals, one located in the Virgin Islands National Park (VINP) and the other on private land. By 2011, there were 198 wild individuals documented at 8 locations and propagation of the species led to the augmentation of 5 of those locations with 243 individuals, for a total of 441 individuals. Since 2011 the populations have been assessed several times revealing declining numbers at all locations. In 2017, population assessments revealed that the species was then known by 127 wild and 67 introduced individuals, for a total of 194 at 7 locations. The known number of individuals had been reduced by 56%, 3 of the original populations had become extirpated and two new populations were recently discovered. Post Hurricanes Irma and Maria assessments in 2018 revealed a slight increase in the numbers of individuals to 238 (mainly juveniles). Although virtually no natural recruitment had been documented in the wild since 2011, 40 seedlings were observed at the largest known population on St. John. In addition to the documenting 40 seedlings, post hurricane assessments in the British Virgin Islands led to the discovery of 46-48 individuals of marron bacora. The total known individuals of marron bacora in 2018 was 324 at 8 locations, 7 locations in the USVI and 1 in the BVI.

Marron bacora has a fragmented distribution due to land clearing and development that has occurred since the 1700s. Although most of the known locations are on National Park Service (NPS) land the largest population may be threatened by development and habitat fragmentation in the near future, as neighboring habitat extend to private lands. Feral ungulates, including white-tailed deer, goats, pigs, and donkeys have been introduced to many of the islands in the past. White-tailed deer were introduced to St. John beginning in the early 1700s and up through the early 1900s. The white-tailed deer in particular are at very high densities on St. John. Although the deer do not heavily graze on marron bacora, it is hypothesized that the depredation of their fruit by these feral ungulates has led to low levels of natural recruitment across its range. In addition, other threats to the species include; sap sucking insects, small population sizes, fragmented/isolated populations and a vulnerability to stochastic events.

Marron bacora is a dioecious obligate outcrosser and thus self-incompatible so it is important to take the ratio of mature male to female plants into consideration when evaluating the viability of a population. Under greenhouse conditions marron bacora may reach sexual maturity in two years, however, this is not common in the wild populations.

To evaluate the viability of the species, which is the likelihood of persistence over the long term,

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we utilized the concepts of resilience, redundancy and representation. Marron bacora is known from a small geographic area with little connectivity and therefore has low representation. Redundancy is low due to an apparent reduction on the range of the species and the low number of known viable populations. All known populations are most likely below the estimated minimum viable population level, thus, resilience is low.

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Solanum conocarpum Species Status Assessment 2019

Table of Contents

Executive Summary 3 Table of Contents 5 List of Tables and Figures 9 Chapter 1. Introduction and SSA Framework 11 Chapter 2. Species Life History and Ecology 13 2.1 Morphology 13 2.2 Taxonomy 13 2.3 Genetic Diversity 14 2.4 Phenology and Reproduction 14 2.5 Survival Rates, Lifespan, and Demographic Trends 17 2.6. Habitat 18 2.7 Geographic Range 19 2.7.1 Historical Range 19 2.7.2 Current Range 20 Chapter 3. Population Level Current Conditions 22 3.1 Nanny Point (St. John, USVI) 22 3.2 Friis Bay (St. John, USVI) 25 3.3 John’s Folly (St. John, USVI) 26 3.4 Brown Bay Trail (St. John, USVI) 27 3.5 Reef Bay Trail (St. John, USVI) 28 3.6 Reef Bay Valley (St. John, USVI) 29 3.7 Europa Ridge (St. John, USVI) 29 3.8 Sabbat Point (St. John, USVI) 29 3.9 Base Hill (St. John, USVI) 29

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3.10 Brown Bay Ridge (St. John, USVI) 29 3.11 Sabbath Hill (Tortola, BVI) 30 3.2 Range of Estimated Potential Habitat on St. John, USVI 30 Chapter 4. Summary of Individual, Population and Species Requirements 31 4.1 Requirements of Individuals 31 4.1.1 Habitats 31 4.1.2 Reproduction 32 4.1.3 Precipitation 32 4.2 Requirements of Populations 33 4.2.1 Minimum Viable Population 33 4.2.2 Size of Habitat Patch 34 4.3 Requirements of Species 34 Chapter 5. Factors Affecting the Survival of marron bacora: Threats 36 5.1 Factors Affecting the Viability of the Species 36 5.1.1 Demographic consequences of small population size and density 36 5.1.2 Genetic consequences of small population size 36 5.1.3 Human Induced Fires 36 5.1.4 Climate Change and Hurricanes 37 5.1.5 Invasive Species 37 5.1.6 Insect Pests and Pathogens 40 5.1.7 Phenology and Breeding System 41 5.1.8 Recreation 41 5.1.9 Conservation Measures 42 5.1.10 Summary 43 5.2 Climate Change 44

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5.2.1 Background 44 5.2.2 Temperature, Precipitation and Drought Stress 46 5.2.3 Life Zones 49 5.2.4 Storm Frequency and Ferocity 51 5.2.5 Sea Level Rise 53 5.2.6 Potential Threats to Ecosystems 53 Chapter 6. Future Climatic Scenarios 55 6.1 Precipitation and Drought Conditions 55 6.2 Life Zones 55 6.3 Summary 55 6.4 Future Scenarios 56 6.4.1 Scenario I Description 56 6.4.2 Scenario II Description 56 6.4.3 Scenario III Description 57 6.4.4 Likelihood of Scenarios 57 6.5 Future Resilience 57 6.5.1 Nanny Point (St. John, USVI) 59 6.5.2 John’s Folly (St. John, USVI) 60 6.5.3 Brown Bay Trail (St. John, USVI) 61 6.5.4 Friis Bay (St. John, USVI) 62 6.5.5 Base Hill (St. John, USVI) 62 6.5.6 Reef Bay Trail (St. John, USVI) 63 6.5.7 Brown Bay Ridge (St. John, USVI) 63 6.5.8 Sabbat Point (St. John, USVI) 64 6.5.9 Reef Bay Valley (St. John, USVI) 64 6.5.10 Europa Ridge (St. John, USVI) 64

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6.5.11 Sabbath Hill (Tortola, BVI) 64 6.5.12 Future Propagation and Reintroductions 65 6.6 Future Resilience 65 6.7 Future Redundancy 66 6.8 Future Representation 66 VIII. Literature Cited. 67 Appendix A. Population Resilience Tables. 73

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List of Figures 1. Figure 1. SSA Framework’s Three Basic Stages 12 2. Figure 2. Drawings of (a) pistillate flower and (b) staminate flower 15 3. Figure 3. Photos of (a) pistillate flower and (b) staminate flower 16 4. Figure 4. Distribution map of marron bacora 21 5. Figure 5. Map of Nanny Point 24 6. Figure 6. Photo of downed marron bacora after Hurricane Irma 25 7. Figure 7. Trail Cam photos: Pre and post Hurricane Irma at Nanny Point 26 8. Figure 8. Map of John’s Folly 27 9. Figure 9. Map of Brown Bay Trail 28 10. Figure 10. Habitat Suitability Map of St. John 31 11. Figure 11. Average rainfall chart for St. John, USVI 33 12. Figure 12. Rapid View of SSA Assessment 35 13. Figure 13. Photo, Nanny Point before and after 2017 hurricanes 38 14. Figure 14. Distribution map of invasive plants at Nanny Point 39 15. Figure 15. Photos of guinea grass encroaching on marron bacora habitat 39 16. Figure 16. Trail Cam photos of deer in Nanny Point population 40 17. Figure 17. Photos of Jacaranda insects on marron bacora 41 18. Figure 18. Future projected temperature changes 46 19. Figure 19. Projected precipitation changes for the US Caribbean 47 20. Figure 20. Temperature projections for Puerto Rico under 3 scenarios 48 21. Figure 21. Projection for number of days above 90° in Puerto Rico 49 22. Figure 22. Holdridge Life Zones for St. John 50 23. Figure 23. Forest Types of St. John 50 24. Figure 24. Projections for life zone distribution changes in Puerto Rico 51 25. Figure 25. Historic Caribbean hurricane tracks 52

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Solanum conocarpum Species Status Assessment 2019

List of Tables 1. Table 1. Marron bacora population estimates 44 2. Table2. Description of habitat and population factors for resilience 59 3. Table A-1. Nanny Point resilience scores under 3 scenarios 74 4. Table A-2. John’s Folly resilience scores under 3 scenarios 75 5. Table A-3. Brown Bay Trail resilience scores under 3 scenarios 76 6. Table A-4. Friis Bay resilience scores under 3 scenarios 77 7. Table A-5. Base Hill resilience scores under 3 scenarios 78 8. Table A-6. Reef Bay Trail resilience scores under 3 scenarios 79 9. Table A-7. Brown Bay Ridge resilience scores under 3 scenarios 80 10. Table A-8. Sabbat Point resilience scores under 3 scenarios (extirpated) 81 11. Table A-9. Reef Bay Valley resilience scores under 3 scenarios (extirpated) 81 12. Table A-10. Europa Ridge resilience scores under 3 scenarios (extirpated) 82 13. Table A-11. Sabbath Hill resilience scores under 3 scenarios 83 14. Table A-12. Resilience scores under 3 scenarios for all populations 84

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Chapter 1. Introduction and SSA Framework

Marron bacora (Solanum conocarpum) is a dry-forest, flowering shrub of the family of Solanaceae, or Nightshade family, which grows to approximately 3 meters (m) (9.8 feet (ft)) in height. The species was originally described by Dunal in 1813 from a specimen collected from Coral Bay by French Botanist, Louis M. Richard, who visited St. John in 1787 (Acevedo- Rodríguez 1996, p. 415). Marron bacora was thought to be extinct in the early 1900s until it was once again described by Pedro Acevedo-Rodríguez in 1992 (Vilella and Palumbo 2010, p. 1). Previously thought to be endemic to St. John in the U.S. Virgin Islands, the range of the species has expanded to include Tortola in the British Virgin Islands with a confirmed record reported in 2018 (Heller 2018, pp. 1-10).

Marron bacora has been a U.S. Fish and Wildlife Service (Service) candidate for listing since 2011 (76 FR 9722, entire). We are now reviewing the species for listing under the Endangered Species Act (Act). This Species Status Assessment (SSA) is a comprehensive status review of marron bacora that evaluates the best available science to inform the listing decision and guide future conservation efforts. This SSA will be a living document to be easily updated as new information becomes available and will support all functions of the Endangered Species Program from Candidate Assessment to Listing to Consultations to Recovery.

The SSA framework (USFWS 2016b, entire) summarizes the information assembled and reviewed by the Service, incorporating the best available scientific and commercial data, to conduct an in-depth review of a species’ biology and threats, evaluate its biological status, and assess its resources and conditions needed to maintain long-term viability. For the purpose of this assessment, we define the viability of marron bacora as its ability to sustain populations in the wild beyond the end of a specified time period. Using the SSA framework, we consider what the species needs to maintain viability through an assessment of its resilience, redundancy, and representation (Figure 1).

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• Resilience refers to the population size necessary to endure stochastic environmental variation (Shaffer and Stein 2000, pp. 308-310). Resilient populations are better able to recover from losses caused by random variation, such as fluctuations in recruitment (demographic stochasticity), variations in rainfall (environmental stochasticity), or changes in the frequency of anthropogenic activities.

• Redundancy refers to the number and geographic distribution of populations or sites necessary to endure catastrophic events (Shaffer and Stein 2000, pp. 308- 310). As defined here, catastrophic events are rare occurrences, usually of finite duration, that cause severe impacts to one or more populations. Examples of catastrophic events include tropical storms, floods, prolonged drought, and unusually intense wildfire. Figure 1. Species Status Assessment Framework Measured by the number of populations, their resiliency, and their distribution (and connectivity), redundancy gauges the probability that the species has a sufficient margin of safety to withstand or recover from catastrophic events (such as a rare destructive natural event or episode involving many populations).

• Representation refers to the genetic diversity, both within and among populations, necessary to conserve long-term adaptive capability (Shaffer and Stein 2000, pp. 307-308). Representation can be measured by the breadth of genetic or environmental diversity within and among populations and gauges the probability that a species is able to adapt to environmental changes. The more representation, or diversity, a species has, the more it is capable of adapting to changes (natural or human caused) in its environment and to colonizing new sites.

In summary, this SSA is a scientific review of the available information related to the biology and conservation status of marron bacora. Importantly, it does not provide or pre-determine the Service’s decision that this species does, or does not, warrant protection under the Act. The Service will make that decision after reviewing this document, along with the supporting analyses, other relevant scientific information, and all applicable laws, regulations, and policies, and the results of the decision will be announced in the Federal Register.

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Chapter 2. Species Life History and Ecology

This chapter compiles the background information about marron bacora upon which our viability assessment is based. Research on the species’ reproductive biology, population genetics and habitat modeling has been conducted by various authors (Anderson et al. 2015, p. 471; Palumbo et al. 2016 p. 1; Stanford et al. 2013, p. 173). The 2011 species’ 12-month finding (76 FR 9722, entire) in conjunction with the 2016 USFWS Species Assessment and Listing Priority record for marron bacora (USFWS 2016a, entire) were used as a baseline document for the completion of this assessment. In addition, assessments of the largest known populations of marron bacora were conducted by the Caribbean Ecological Services Field Office (CESFO) in collaboration with NPS in 2017 and 2018. These research and analysis provide the best scientific and commercial data available, and we refer to it throughout the SSA Report.

2.1 Morphology

In 1996, Acevedo-Rodriguez (p. 415) described marron bacora as a dry-forest shrub of the Solanaceae family that may attain 3 meters (m) (or 9.8 feet (ft)) in height. Its leaves are oblong- elliptic or oblanceolate (broader at the distal third than the middle), ranging in size from 3.5 to 7 centimeters (cm) [or 0.62 to 1.5 inches (in)] wide. The leaves are coriaceous (leathery texture) and glabrous (no hairs), and have a conspicuous yellowish mid-vein. The flowers are usually paired in nearly sessile (not stalked) lateral or terminal cymes (flat-topped flower cluster). The corolla consists of five separate petals that are light violet, greenish at the base, and about 2 cm (0.78 in) wide. The fruit, a berry, is ovoid-conical (teardrop shaped), 2 to 3 cm (0.78 to 1.2 in) long, and turns from green with white striations to golden yellow when ripe (Acevedo-Rodríguez 1996, p. 415). The tallest specimen recorded was a cultivated individual of approximately 5 meters (m) with a basal diameter of around 12 cm (USFWS 2017a, p. 4). Little is known about the natural history, reproductive biology, and the effects of herbivory on the species (Ray and Stanford 2003, p. 3) and current knowledge of the ecology of the genetic diversity of Virgin Islands rare flora is sparse (Stanford et al. 2013, p. 173). This information is discussed in later sections.

2.2 Taxonomy

Solanum conocarpum belongs to the Solanaceae (also known as the nightshade family, and is in the same family as potatoes and tomatoes). The genus Solanum includes approximately 1,400 species, mostly herbs and shrubs, but also includes small trees and woody vines with a worldwide distribution (Acevedo-Rodríguez 1996, p. 415). Twenty-four species of Solanum have been recognized from what was once described as the Puerto Rican bank (St. Thomas, St. John, Tortola, Guana, Greater Camanoe, Necker Cay, Virgin Gorda and Cayo Diablo) (Liogier and Martorell 2000, p. 331), with 8 species occurring on the island of St. John (Acevedo- Rodríguez 1996, p. 415). Marron bacora was originally described in 1813 from a type specimen collected by L.C. Richard at Coral Bay, St. John, U.S. Virgin Islands (USVI), in 1787 (Acevedo-

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Rodríguez 1996, p. 415). The best available scientific and commercial information recognizes that S. conocarpum is a valid, distinct species. The common name, marron bacora, was given by Dunal in the original species description (Acevedo-Rodríguez 1996, p. 415), and is used throughout this SSA document as an accepted common name.

2.3 Genetic Diversity.

The genetic diversity of marron bacora has remained relatively high [(Hspe ) = 0.262] at the species level (species heterozygosity), and the variability falls within the natural range of variation that is common in wild plants (Stanford et al. 2013, p. 176). However, genetic variation at the population level (mean population heterozygosity) for this species falls at the low end of the natural range of variation seen in plants, with Htot = 0.0789, and Pp = 20.9 %, primarily because most populations are extremely small (Stanford et al. 2013, p. 176). Nonetheless, these authors suggest that overall genetic variability is comparable to that found in Solanum polygamum, a species native to the Virgin Islands, Hispaniola and Puerto Rico, and widely distributed on the island of St. John. The highest diversity of marron bacora is found in the Nanny Point population located within the VINP. Small populations are unlikely to reproduce successfully since the species is self-incompatible (Stanford et al. 2013, p. 178), they have genetic mechanisms that prevent self-pollination. It is important to highlight that additional wild individuals were identified at John’s Folly, Friis Bay, and Reef Bay Trail on the island of St. John, which may be sampled to expand the above research on the species’ genetic diversity. Thus, there is a potential for a broader genetic variability among these populations.

2.4 Phenology (life-cycle events that are influenced by climate and seasonal change) and Reproduction.

Based on the information available at the time of the species’ 12-month finding (76 FR 9722), it was suggested that marron bacora was functionally dioecious (requiring male and female flowers from different plants to reproduce) (Figure 2 and 3). The outcrossing nature of the species was also suspected due to the absence of offspring in small populations (Stanford et al. 2013, p.174). The findings by Anderson et al. (2015, p. 479) support the hermaphroditic and dioecious biology of marron bacora. It was found that crossing of pollen to the stigma of other male flowers or transferred to the stigma of the same flower resulted in no pollination, and thus, confirmed the auto self-incompatibility of the species. Additional evidence of the self-incompatibility of the species is the absence of fruits on wild and greenhouse plants bearing male (pistillate) flowers (Anderson et al. 2015, p. 478). Nonetheless, Eleanor Gibney (pers. comm., 2017), horticulturist and specialist in the flora of St. John, has recorded fruit production on isolated plants, suggesting the species still has mechanisms for self-pollination . Her observations are supported by the record of flowers and fruits in a solitary wild plant discovered by Acevedo-Rodríguez (pers. comm., 2002) in the White Cliff area (in the general vicinity of Reef Bay), and further suggested that marron bacora may have less reproductive fitness due to self-pollination. The species’ self- incompatibility may be the result of male flowers that are non-functional due to stigmatic

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differences of the pistils (e.g., a lack of sufficient stigmatic exudate) rather than genetic-based self-incompatibility (Anderson et al. 2015, p. 478). Thus, there is a chance that some isolated flowers may be able to be self-pollinated.

The sex ratio of marron bacora is 1:1 and a much longer time is needed for female plants to flower for the first time (from the seedling stage) compared with the males (Anderson et al. 2015, p. 475). This may explain the rarity of the species in the landscape as only half of the wild individuals (based on the 1:1 ratio) have the potential to produce fruits and viable seeds, and thus highlights the importance of introducing an adequate number of plants into the wild (Anderson et al. 2015, p. 482). Nonetheless, there is no available information regarding the seed dormancy or long-term storage (orthodox) potential for marron bacora.

As plant populations become reduced and spatially segregated, important life-history needs provided by pollinators and seed dispersers may be compromised (Kearns and Inouye 1997, p. 305). The fragmented distribution of marron bacora on St. John can be attributed to historical

Figure 2. Drawings of marron bacora flowers: (A) pistillate flower and (B) staminate flower (Anderson et al. 2015, p. 475)

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Solanum conocarpum Species Status Assessment 2019

Figure 3. Photos of marron bacora flowers: (A) pistillate flower and (B) staminate flowers (USFWS, 2017)

habitat degradation. Based on the hermaphroditic and dioecious biology of marron bacora, the species requires cross-pollination. However, pollinators have not been recorded visiting the flowers after dozen of hours of field work (Anderson et al. 2015, p. 475). Nonetheless, recent surveys by the Service (May 2017) recorded carpenter bees (Xylocopa mordax), honey bees (Apis mellifera) and bananaquits (Coereba flaveola) visiting the flowers of marron bacora at Nanny Point (USFWS 2017a, p. 7). In fact, about 92 percent of the 75 marron bacora natural individuals in this area were observed in flower (USFWS 2017a, p. 7). Work by Caraballo-Ortiz and Santiago-Valentín (2011, p. 26) showed that honey bees and bananaquits are effective pollinators of the endangered Matabuey (Goetzea elegans; Solanaceae).

At this point, the natural dispersal mechanism of marron bacora remains unknown, but fruit predation is suspected as the explanation of lack of natural recruitment in the wild (76 FR9722, p. 9726). Although predators may also disperse the species, it is likely that the seeds have not adapted to passing through the gastrointestinal tracts of the exotic mammals currently occurring in the island of St. John (e.g., white-tailed deer, feral hogs, donkeys). The native hermit crab (Coenobita clypeatus) has also been documented depredating marron bacora fruit (Ray and Stanford 2005, p. 18; Vilella and Palumbo 2010, p. 1) and although there are several species of fruit eating bats on St. John (Artibeus jamaicensis, Brahyphylla cavernarum and Stenoderma rufum) there have been no studies to document their possible role in the life history of marron bacora if any. Also, it is possible that natural fruit dispersers of marron bacora had targeted other food sources as the populations of this shrub became increasingly patchy, as a result of deforestation and introduction of exotic plant species. The patchy distribution of this species may suggest that its natural disperser is extinct or that the populations of the plant are too small to attract the disperser (Roman 2006, p. 82). Despite this, the Service recently recorded prolific fruit production on several individuals within the Nanny Point population (i.e., about 70 fruits developing on a single plant); and monitoring of possible fruit dispersal or predation is ongoing by using LTL Acorn (LTL-6310) 12-megapixel scouting cameras (USFWS 2017a, p. 7).

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2.5 Survival Rates, Lifespan, and Demographic Trends.

Little is known of the life history of this plant. Marron bacora is a perennial shrub that may live over two decades. For example, the Nanny Point population was discovered in 2002 (Carper pers. comm., 2005), and at that time the population was already composed mainly of adult individuals and little natural recruitment was recorded. Thus, the current known natural individuals at Nanny Point should be approximately 20 years old. In addition, Eleanor Gibney has marron bacora material under cultivation from the individual rediscovered by Pedro Acevedo-Rodríguez in early 1990s (USFWS 2017a, p. 4). Therefore, these plants would also be over 20 years old. Nonetheless, the species may reach a reproductive maturity after 16 months from germination (under greenhouse conditions) (Anderson et al. 2015, p. 475). However, this period is expected to be greater in the wild as seedlings may require longer periods to grow and individuals may remain suppressed under closed canopy and possible drought conditions.

There is little evidence of natural recruitment in all known populations of marron bacora. The population structure at Nanny Point and John’s Folly is characterized by the absence of individuals smaller than one meter high, with little evidence of seedlings or juveniles (three for Nanny Point and one for John’s Folly) (USFWS 2017a, p. 7). These populations consist primarily of reproductive individuals, as 92 percent and 75 percent of the plants, respectively, were recorded in flower during a recent survey (USFWS 2017a, p. 7). The Nanny Point population size recorded during this survey represents about 52 percent of the estimated number of individuals (144 plants) at the time of the 12-month finding for the species was published (USFWS 2017a, p. 7), thus indicating a population decline. The John’s Folly population was composed of 4 natural adult individuals (reproductive size individuals naturally occurring at this site) or 36 percent of the total (11 plants) estimated at the time of the 12-month finding (USFWS 2017a, p. 9) (see Table 1). The lack of natural recruitment does not seem to be attributed to low seed viability as germination under greenhouse conditions is high, with almost 100% germination (Ray and Stanford 2005, p. 6).

Efforts have been conducted to enhance existing natural populations by planting seedlings (Stanford et al. 2013, p. 178). Stanford et al. planted 128 seedlings (different seed sources) at two localities in the south coast of St. John (Europa Ridge and Reef Bay Valley). Overall survival of these seedlings over a 32 month period was approximately 81.3% in Europa Ridge, and 78.1% in Reef Bay Valley, and irrespective of seed source, survival rate was not significantly different between the two sites (p =0.8828) (Stanford et al. 2013, p. 177). However, growth rates for these sites were recorded as highly erratic and plant material was affected by drought stress and insect herbivory (Stanford et al. 2013, p. 178). Further monitoring of these sites by NPS staff has not located living material of marron bacora, either natural or planted, and these populations are presumed extirpated (McKinley pers. comm. 2017). In fact, an assessment of these areas in 2017 resulted in no finding of the species (USFWS 2017a, p. 11). Additional population enhancements from seedling and cuttings have been conducted at Nanny Point (50), John’s Folly (37) and Brown Bay (36) (76 FR 9722, p. 9724). The current number of surviving 17

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individuals for these sites is 44 (88%), 13 (35%) and 10 (27%), respectively (USFWS 2017a, p. 13).

Similar to marron bacora, mata buey (Goetzea elegans) requires cross pollination for flower and fruit set and its scattered distribution may could prevent outcrossing (Caraballo-Ortiz et al.2011, p. 31). Plant sterility does not appear to be a plausible theory for the lack of recruitment, as, again, germination under greenhouse conditions is highly successful (Ray and Stanford 2005, p. 6). It may be similar to matabuey, which is another species endemic to the Caribbean that shows a conspicuous flowering with showy fruits, but faces problems with its dispersal and recruitment.

2.6 Habitat.

Marron bacora was thought to be endemic to the dry and deciduous forest of the island of St. John (Acevedo-Rodríguez 1996, p. 415), an island with more than 80% of its land surface considered as subtropical dry forest (Stanford et al. 2013, p. 173). However, records of marron bacora in the British Virgin Islands (Acevedo-Rodríguez 1996 p. 415), were recently confirmed in 2018 in the tropical dry forest of Tortola. By 1717, the forested landscape of St. John was parceled into more than 100 estates for agriculture (i.e., sugarcane and cotton) and the majority of this landscape was cleared. Remaining fragments of forested habitat served as refugia for other regional endemics and federally listed species (e.g. Zanthoxylum thomasianum and Calyptranthes thomasianum).

Nonetheless, all known localities of marron bacora are located in areas with evidence of previous vegetation clearing or adjacent to sites with previous intensive agricultural land use, showing the potential of the species to occur on disturbed sites. The newly discovered populations of marron bacora occur on dry and poor soils (Ray and Stanford 2005, p. 6). Carper and Ray (2008, p. 1) reported that the species was locally abundant in exposed topography on sites disturbed by erosion (depositional zones at the toe of the slopes), areas that have received moderate grazing, and around ridgelines as an understory component in diverse woodland communities. According to Acevedo-Rodríguez (pers. comm. 2002), the areas where surviving natural populations have been found (i.e., dense scrubland) may not represent optimum habitat. Research by Ray and Stanford (2005, pp. 11-12) on the forest structure at the Reef Bay area shows a forest community invaded by numerous exotic woody and herbaceous plant species in an area subject to erosion. In the case of Europa Ridge, the area is dominated by black mampoo (Guapira fragrans), a fast- growing species, and the overall area is devoid of woody exotics, suggesting relative lack of perturbation (Ray and Stanford 2005, pp. 11-12). The forest structure at Europa Ridge suggests this area was selectively logged for charcoal, but not subjected to intensive agriculture, probably because of the steep topography. Currently, habitat descriptions are unavailable for the recently described population in Tortola, BVI.

Thus, marron bacora appears to occur naturally in communities with broadly varying suites of associated woody species, and shows little fidelity to any particular suite of community 18

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associates (Ray and Stanford 2005, p. 11). For example, none of the known populations of marron bacora are directly associated or within remnants of other rare endemics (e.g., Eugenia earhartii and Zanthoxylum thomasianum). Marron bacora habitat preference may involve some combination of abiotic factors (Ray and Stanford 2005, p. 13). Therefore, fidelity to a particular assemblage of community associations is apparently not a useful indicator of marron bacora habitat (Ray and Stanford 2005, p. 15).

A spatial habitat model for marron bacora using elevation, slope, soil association, and vegetation types, identified 694.94 ha (1717.23 acres) of high-quality habitat, 1,274.94 ha (3150.45 acres) of moderate-quality habitat, 1,568.53 ha (3875.92 acres) of low-quality habitat, 1,343.16 ha (3319.16 acres) of poor-quality habitat, and 186.88 ha (461.79 acres) of unsuitable habitat (Palumbo et al. 2016, p. 5) on St. John. Within the boundaries of VINP, the model identified 387.43 ha (957.36 acres) of high-quality habitat, 773.77 ha (1912.03 acres) of moderate-quality habitat, 1096.91 ha (2710.52 acres) low-quality habitat, 1106.30 ha (2733.73 acres) of poor- quality habitat and 181.22 ha (447.80 acres) of unsuitable habitat. When adding all hectares of high- and moderate- quality habitat, approximately 32% of the land area of VINP may be suitable habitat for marron bacora (Palumbo et al. 2016, p. 5) (Figure 10). As previously mentioned, the specific habitat requirements of this species remain unknown, but as other species of the genus Solanum, marron bacora may be adapted to poor soils and some sort of natural disturbance (e.g., hurricanes). However, the current rarity and patchy distribution of the species may be the result of the species reproductive biology, the absence of dispersal, suspected fruit predation and the overall disturbance and deforestation that affected the island of St. John.

2.7 Geographic Range. 2.7.1 Historical Range

Marron bacora is a shrub thought to be endemic to the Island of St. John, and was originally known from a type specimen collected by L.C. Richard at Coral Bay, St. John (U.S. Virgin Islands), in 1787 (Acevedo-Rodríguez 1996, p. 415). However, there are no population estimates or details on its abundance from L.C. Richard’s discovery, nor are there any known population estimates or further records of the species prior to 1992.

The species was described again in 1992 by Acevedo-Rodríguez in the area of Europa Ridge, just north of the White Cliff area in the Island of St. John (Ray and Stanford 2003, p. 4). The species was presumed to be near extinction, as two mature plants (two localities) were believed to be the only specimens left in the wild. Acevedo-Rodríguez (1996, p. 415) further mentioned the possibility of the species occurring in Virgin Gorda (British Virgin Islands). However, further surveys on Virgin Gorda did not result in any record of the species (Acevedo-Rodríguez pers. comm., 2002). Thus, Acevedo-Rodríguez (pers. comm., 2002) considered the specimen by Elbert Little (Little 23836) as a possible misidentification from sterile material. Furthermore, extensive habitat assessments at Virgin Gorda have been conducted by staff from the Royal Botanic Garden (KEW) and the British Virgin Islands National Park Trust (NPT) in

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collaboration with Service Biologist Omar A. Monsegur-Rivera (2015, 2016, and 2017), with no marron bacora being observed. Nonetheless, the Island of Virgin Gorda harbors one of the best remnants of native vegetation in the Virgin Islands with populations of rare endemics (e.g. Zanthoxylum thomasianum and Calyptranthes thomasianum) (Barrios et al. 2017, p. 29), and thus, the presence of undetected populations of marron bacora should not be discarded. Recent surveys by staff from the Royal Botanic Garden (KEW) in Tortola identified a plant that is morphologically consistent with marron bacora (Hamilton pers. comm., 2018) (Heller et al. 2018, p.1). In November of 2018 surveys to confirm marron bacora on Tortola succeeded when fertile material (flowers and fruit) was located by staff from KEW and the National Park Trust on marron bacora individuals. The original plant described above could not be located but a population of approximately 40 individuals was identified (Heller pers. comm., 2018).

After 1992, six additional populations of marron bacora were identified on St. John (i.e., Nanny Point, Friis Bay, John’s Folly, Brown Bay Trail, Sabbat Point, and Base Hill) (Figure 4). In addition, the latest population and habitat assessments by the Service (2017) identified new populations, one along the Reef Bay Trail on the southern coast of St. John (USFWS 2017a, p. 11), and a seedling at Brown Bay Ridge on the north east side of the Island. However, during the 2017 surveys, the Service visited the Sabbat Point site along with Eleanor Gibney (botanist who discovered this locality) and found no individuals of marron bacora, and the overall area is extremely disturbed based on the abundance of species like Leucaena leucocephala, Opuntia repens and Bromelia pinguin. According to Eleanor Gibney, this population has not been surveyed in over a decade and the individuals are suspected to be extirpated.

2.7.1 Current Range

As of the date this review was initiated, the species was considered as endemic to the island of St. John, USVI. Based on the best available information, three of the populations discovered since the species’ was re-identified in 1992, are considered extirpated (i.e., Reef Bay, Europa Ridge, and Sabbat Point). There are three populations that contain wild individuals on the northern side of St. John (i.e., Base Hill, Brown Bay Trail and Brown Bay Ridge), and four towards the southeast side of the island (i.e., Nanny Point, Friis Bay, Reef Bay Trail, and John’s Folly). In addition, a seedling of marron bacora was recently identified by Eleanor Gibney and Cecilia Rogers (student, James Madison University) on a new site near the Brown Bay Trail, however further monitoring is needed to confirm if this seedling is effectively marron bacora, and if it is part of a larger population. The Reef Bay Trail population corresponds to a new population located by Service staff in 2017. Of the seven extant populations on St. John, all but one (Friis Bay) occur within the boundaries of the VINP. Nonetheless, the latest available information confirms the range of the species extend to Sabbath Hill in the island of Tortola, British Virgin Islands.

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Figure 4. Distribution map of marron bacora.

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Chapter 3. Population Level Current Conditions

In September 2017, the US and British Virgin Islands were directly impacted by two Category 5 Hurricanes, based on the Saffir-Simpson Scale (sustained winds over 137 knots [kt] or 157 mph). On September 6th, Hurricane Irma directly impacted the northern USVI (St. Thomas and St. John) and the BVI (including Tortola) with sustained wind speeds at 155 kt (178.4 mph) and heavy rain conditions throughout the area (Cangialosi et al. 2018, pp. 3). On September 20th, Hurricane Maria severely impacted the southern island of St. Croix in the USVI when the eye of the storm centered about 25 n mi (28.8 miles) south of the islands with sustained winds at 150 kt (172.6 mph) and heavy rain conditions throughout the area prior to downgrading to a Category 4 (sustained winds between 113 kt and 136 kt) and making landfall in Puerto Rico (Pasch et al. 2018, pp. 2).

Location and topography play major roles in influencing the wind, rain and runoff produced by hurricanes. In general, the forests in the USVI and BVI experienced defoliation and many trees felled. In some areas flash floods and the loss of forest canopies were obvious. For this reason, the following account provides detailed information on the status of the currently known populations of marron bacora including localized impacts caused by Hurricanes Irma and Maria.

3.1 Nanny Point (St. John, USVI)

The largest population of marron bacora is located at Nanny Point in southeast St. John. This population was discovered in 2003, when the owner of the property commissioned a plant inventory of his property and 184 wild individuals were recorded; mainly mature plants with little evidence of natural recruitment (Ray and Stanford 2005, p. 16; 76 FR 9722, p. 9723). The majority of this property was donated and transferred to NPS. Forty individuals that were located along an access corridor were translocated to the adjacent habitat donated to NPS, but failed to get established and later died due to the lack of available water. This event reduced this population to approximately 144 plants in 2010 (76 FR 9722, p. 9723).

The Service conducted a comprehensive assessment of the Nanny Point population in May of 2017 (Figure 5). The population was found to be composed of 75 mature adult individuals, three natural seedlings and 44 planted individuals from a past population enhancement effort. An additional seedling was discovered on a subsequent visit to the population by Service biologist James Yrigoyen and the James Madison University student Cecilia Rogers (Yrigoyen and Rogers 2017, pers. obs.) bringing the total population count to 123 individuals. The population can be characterized as an having an old population structure, with plants showing large size for both basal diameter and height (i.e., all wild individuals were greater than 1 m (3.28 ft) high).The number of adult natural individuals counted during the 2017 survey (i.e., 75) represents approximately 52% of the number of individuals (144 plants) at the time of the 12-month finding (2011) for the species was published (USFWS 2017a, p. 7).

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During the 2017 assessment, about 69 of the natural individuals (92%) were observed in flower, with some individuals already presenting evidence of fruit development (USFWS 2017a, pp. 7). However, despite this evidence of reproduction, only three seedlings were observed. Further fruit development was observed by Cecilia Rogers (student, James Madison University) and Service biologist James Yrigoyen on the weeks following the May 2017 assessment, with about 70 fruits counted on a single plant. The low number of seedlings, despite the relatively high fruit production is consistent with the information already available to the Service indicating that this population continues to show low recruitment (Ray and Stanford 2005 p. 18; 76 FR 9722, p. 9726; USFWS 2017a, p. 7). The distribution of wild/natural individuals appears to be associated with moisture availability at the site as the majority of plants are located along a storm drainage (gut) area. The plants appear to be healthy, although some were observed in stress at the easternmost corner of the population associated to low canopy vegetation and wind exposure (USFWS 2017a, p. 7).

Hurricane Irma, a category 5 hurricane, was considered as one of the most catastrophic and costliest atmospheric events to affect the US Caribbean (Cangialosi 2018, pp. 13-17). In an effort to record possible impacts from this storm on the Nanny Point population, a field camera was installed near the base of a tree facing a marron bacora plant that had been previously documented with 70 fruits. The field camera recorded 6,023 photos between September 2nd and October 21st, 2017. The images captured revealed a flash flood and loss of canopy in the marron bacora population at Nanny Point during Hurricane Irma, and another flash flood caused by Hurricane María (USFWS 2017b, p.3) (Figure 6 and 7).

A rapid assessment of the Nanny Point population was carried out on November 9, 2017 by the Service. Twenty three live individuals were identified with only 15 of those with tags still attached. Some marron bacora had trees or branches on top of them while others were covered with runoff sediment. Four marron bacora tags were discovered in piles of runoff debris and several individuals were observed uprooted. Four individuals were observed with flowers, two of which were still attached to the roots but were lying on the ground (USFWS 2017b, p. 6-8) (Figure 6). However, it is important to highlight this was a partial assessment due to safety as downed trees and branches made it difficult to maneuver through the population.

In June of 2018 a post hurricane assessment of the Nanny Point population was carried out by Island Conservation (IC) with support from the Service. Field objectives for this visit were to assess populations on NPS land, identify potential invasive species threats, install cameras to record impacts from introduced mammals or seed dispersers and lastly to meet with a local horticulturalist that could possibly help with propagation of the species. During this survey the

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Figure 5. Marron bacora population (2017) at Nanny Point pre-hurricane: distribution of the natural individuals (red dots) vs. planted material (yellow dots) and seedlings (green dots) (USFWS 2017a, p. 8)

population was found to be composed of 201 individuals (108 adult individuals, 53 juveniles, 40 seedlings). It is important to point out that for the purpose of this survey; individuals were classified as adults, juveniles and seedlings based on their height (≤ 10.0 cm – seedlings, 10.1 – 50.0 cm – juveniles, ≥ 50.1 cm – adults). One of the main highlights of the IC (2018) is the abundance of natural seedlings (40) and juveniles (53) compared to previous data available to the Service indicating the absence of natural recruitment. It may be suggested that hurricane disturbance created favorable conditions for seedling establishment, and the population is now classified as having a mixed age structure, with 46.3% of the individuals being classified as juveniles and seedlings and 53.7% being classified as adults. However, the survival of the recently recorded seedling remains uncertain and long term monitoring is needed. Of the 44 previously introduced individuals, 29 were found alive, one dead and 14 unaccounted for. Nineteen individuals surveyed and tagged in 2017 could not be located after the hurricanes.

The post hurricane assessment in 2018, 127 individuals (63.2%) exhibited some form of dieback on their leaves and 71 individuals (35.5%) exhibited herbivory impacts, specifically browsing

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potentially from invasive mammals, such as white tail deer (Odocoileus virginianus) (IC Report, 2018, p. 5). Other invasive mammals that have also been described in the VINP are hogs

Figure 6. Adult marron bacora downed by 2017 hurricanes (USFWS 2017b, p. 7)

(Sus scrofa), goats (Capra aegagrus hircus), and donkeys (Equus africanus asinus). The Jacaranda insect, Insignorthezia insignis, initially recorded by USFWS during the 2017 population assessment, was still present during the 2018 survey (IC Report 2018, p. 7) (Figure 17). Potential impacts to marron bacora by these insects are still unknown, but considering the dioecious reproductive biology of marron bacora, such insect pest may have negative impacts on the reproductive output. These pests are also known as ‘sap suckers’ that harm and weaken plants by sucking their sap and leaving them vulnerable to pathogens and molds. Therefore, this may put further stress on the plants and thus reduce their reproductive output, affecting the likelihood of recruitment.

To evaluate the potential impacts of the hurricanes on the population, the orientation of the base of the individuals was recorded as lying or standing. The values of 14.9% and 85.1% were calculated for lying versus standing respectively. These observations were also recorded in the pre-hurricane assessment in 2017 to be 14.2% and 85.8% for lying vs standing (USFWS 2017a). The post hurricane assessment conducted in 2018 also found that 27.8% of adult plants were lying on the ground and 17 adult plants were found dead (IC Report 2018, p. 5).

3.2 Friis Bay (St. John, USVI)

With the discovery of a new population in the British Virgin Islands, this is now the third largest natural population of marron bacora, with an estimated number of 33 individuals (Ray and Stanford 2005, p. 16). The site has not been visited since 2005, thus no current information is available on the status of this population.

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Pre 2017 Hurricanes Flash flood during Hurricane Irma

Post Irma Flash flood during Hurricane Maria Figure 7. Trail camera photos, pre and post hurricanes (USFWS 2017b, p. 3-4)

3.3 John’s Folly (St. John, USVI)

This site is located upslope in a ravine as adjacent to Road 107, and about 700 m northwest of the Nanny Point population. The twelve month finding (76 FR 97220, p. 9724) reported 11 natural individuals and 37 introduced individuals at this site (50 CFR Part 17). A population assessment in 2017 only identified 4 natural individuals and 1 natural seedling, and 13 plants corresponding to planted material from a previous population enhancement with material from the Nanny Point population (USFWS 2017a, p. 7) (Figure 8). The number of natural plants at John’s Folly in 2017 represented about 36% of the known natural plants (11) at the time of the 2010 twelve month finding. Three (75%) of the natural individuals were observed in flower. Despite the evidence of flowering events, natural recruitment appear to be minimal as only one natural seedling was observed. The distribution of the natural individuals is similar to Nanny Point with the majority of the plants at the bottom of the drainage. This site is located along the Park boundaries and the populations appear to be affected by human disturbance such as vegetation clearing for a hiking trail that begins nearby and former evidence of dumping

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(USFWS 2017a, p. 9).

In 2018, a post hurricane assessment of the population reported the population composed of 18 adult individuals with no seedlings or juveniles reported. All individuals documented in this population were found to be over 53.0 cm (20.87 in). Once again, it is important to note that for the purpose of this survey; individuals were classified as adults, juveniles and seedlings based on their height (≤ 10.0 cm – seedlings, 10.1 – 50.0 cm – juveniles, ≥ 50.1 cm – adults) (IC Report 2018, p.2). Average basal diameter was 17.7 mm and none of the plants presented flowers or fruit. All individuals in this population were described as standing (none lying) with three of the individuals (16.7%) exhibiting some form of dieback and 11 plants (61.1%) exhibiting a combination of impacts by herbivorous insects and browsing by potential invasive mammals (IC Report 2018, p. 7).

Figure 8. Distribution of marron bacora at John’s Folly (USFWS 2017b, p.8)

3.4 Brown Bay Trail (St. John, USVI)

The Brown Bay Trail site is located along the Brown’s Bay hiking trail within the VINP, an area of mature secondary dry forest located on the northeastern shore of St. John. The site is located on a slope approximately 60 m (196.85 ft) from shore and included a single natural individual and several planted saplings from material propagated from the Nanny Point population. The wild individual is located on the edge of an NPS maintained hiking trail and showed signs of 27

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direct impacts from trail maintenance activity (i.e., clearing of vegetation) (Palumbo et al. 2016, pp. 6-7). Similar to the other sites visited, there was no direct evidence of natural recruitment (Palumbo et al. 2016, pp. 6-7). In 2017, surveys by Cecilia Rogers (student, James Madison University) located the known natural individual with a basal diameter of 68 mm and about 3.10 m (10.17 ft) high (USFWS 2017a, p. 13). In addition she reported finding 10 planted individuals or 27% of the originally planted material from a previous population enhancement (USFWS 2017a, p. 13).

In 2018, a post hurricane assessment reported that the population was composed of 18 individuals, 17 adults and 1 juvenile. The population here was described as an aged structure, with 94.4% of the individuals being classified as adults. Once again, it is important to note that for the purpose of this survey; individuals were classified as adults, juveniles and seedlings based on their height (≤ 10.0 cm – seedlings, 10.1 – 50.0 cm – juveniles, ≥ 50.1 cm – adults) and not their reproductive ability (IC Report 2018, p. 2). The average basal diameter for adults was 16.7mm and no signs of flowers or fruit were observed in this population. In an effort to evaluate the impacts from hurricanes four individuals (22.2%) were lying on the ground and 14 (77.8%) were described as standing up. Five individuals (27.8%) presented some form of dieback on their leaves, 12 individuals (33.3%) exhibited impacts by herbivorous insects and browsing by potential invasive mammals and all of the plants at this location were described as suffering from severe dry conditions (IC Report 2018, p. 8) (Figure 9).

Figure 9. Distribution of marron bacora at Brown Bay (IC Report 2018, p. 9)

3.5 Reef Bay Trail (St. John, USVI)

The Reef Bay Trail locality is a new population located by Service and NPS staff during the latest population assessments in 2017 (USFWS 2017a, p. 11). The site lies along the NPS hiking trail from Europa Bay to Reef Bay Hiking, approximately 1 km north of White Cliff. A population assessment led by Cecilia Rogers (student, James Madison University) in 2017, 28

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discovered 7 wild individuals, 85% in flower and some individuals producing fruits. Additional habitat surveys may be required for a more thorough assessment of this area. No post hurricane assessments were carried out for this population.

3.6 Reef Bay Valley (St. John, USVI)

This locality is on the southern coast of St. John, along the shore near White Cliffs. In 2005, six wild and 60 introduced individuals were reported at the Reef Bay site (Ray and Stanford 2005, p. 16). However, the information available at the time of the 12-month finding (2011) and information provided by local experts indicates this population was extirpated (76 FR 9722, p. 9724). Further assessments of this area by Service staff along with staff from NPS and other partners were unsuccessful in identifying any individual of marron bacora (USFWS 2017a, p. 11). Thus, the best available information indicates this population is extirpated, and no individuals are known in its proximity.

3.7 Europa Ridge (St. John, USVI)

The Europa Ridge population is a single individual known since the early 1990s, from the time the species was described by Acevedo-Rodriguez in the southern coast of St. John. According to Ray and Stanford (2005, p. 16) the site was composed of one natural individual and 60 planted individuals (population enhancement). However, according to the latest information from NPS staff, and based on the latest habitat assessments by the Service this population is likely extirpated (USFWS 2017a, p. 11).

3.8 Sabbat Point (St. John, USVI)

This population was reported as a single natural individual by Ray and Stanford (2005, p. 16; 76 FR 9722, p. 9724). Service staff was recently guided to the site by Eleanor Gibney (local expert and horticulturalist who discovered this site), however, we were not able to locate the individuals, and the site shows evidence of recent disturbance based on the abundance of Leucaena leucocephala, Opuntia repens and Bromelia pinguin (USFWS 2017a, p. 4). Based on Eleanor’s experience, she considers this population as extirpated.

3.9 Base Hill (St. John, USVI)

The population at Base Hill consists of 1 natural individual as reported by Ray and Stanford (2005, p. 16). There have been no subsequent visits to this population since 2005 and thus, no further data on the status of this individual is known.

3.10 Brown Bay Ridge (St. John, USVI)

In 2017, one wild individual was recently discovered in atop of a ridge approximately 0.25 miles 29

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from the Brown Bay Trail population by Eleanor Gibney (Cecilia Rogers 2017, pers. comm.). Additional habitat surveys may be required for a more thorough assessment of this area and no post hurricane assessments were carried out in this area.

3.11 Sabbath Hill (Tortola, BVI)

In 2018, surveys conducted by staff from the Royal Botanic Garden (KEW) in Tortola identified a plant morphologically consistent with marron bacora, near Sabbath Hill. On a follow-up trip to confirm marron bacora in the area, a population of approximately 46 to 48 individuals were identified. Forty of the plants were described as small plants [<1m (3.28 ft) tall, stem diam <1 cm (0.39 inch) and 6-8 as large plants 1.5-2.2 m (4.92-7.22 ft) tall, stem diam. >3 cm 1.18 inches)]. Three of the large plants were described as fertile, with one having flowers with no fruit, another having flowers and immature fruit, and the last having fruit but no flowers. Cuttings were collected for DNA analysis. The habitat was described as having open vegetation compared with the surrounding forest and containing a lot of non-native annuals and Acacia riparia encroaching. Feral animal droppings were also noted in the area and grazing of marron bacora also noted (Heller et al. 2018, entire).

3.2 Range of Estimated Potential Habitat on St. John, USVI.

Marron bacora was thought to be endemic to St. John in the USVI until 2018 when a population was described on the island of Tortola in the British Virgin Islands. A GIS habitat suitability model for marron bacora was created utilizing NPS and USDA databases that include information on vegetation cover, soil associations, digital elevation models and VINP boundaries. This information was then used to generate a model for the species using environmental characteristics from 8 locations on St. John. For each location values for elevation, slope, soil association and vegetation were extracted. The weighted linear combination (WLC) method was used to create composite maps. Model outputs included geographic representation of the spatial distribution of marron bacora habitat and an estimate of area in hectares of each suitability category. Overall, the model identified 1,929.87 ha (4,768.81 acres) of highly suitable and moderately suitable habitat, and of these, 1161.20 ha (2,869.39 acres) (60.2%) occurring within the boundaries of Virgin Islands National Park. The results of the spatial habitat model for marron bacora identified 694.94 ha (1,717.23 acres) of high-quality habitat, 1,274.94 ha (3,150.45acres) of moderate-quality habitat, 1,568.53 ha (3,875.92 acres) of low-quality habitat, 1,343.16 ha (3,319.02 acres) of poor-quality habitat and 186.88 ha (461.79 acres) of unsuitable habitat. Within the boundaries of VINP, the model identified 387.43 ha (957.36 acres) of high-quality habitat, 773.77 ha (1,912.03 acres) of moderate-quality habitat, 1,096.91 ha (2,710.52 acres) low-quality habitat, 1,106.30 ha (2,733.73 acres) of poor-quality habitat, and 181.22 ha (447.80 acres) of unsuitable habitat. When adding together the hectares of high- and moderate- quality habitat, approximately 32% of the land area of VINP may be suitable habitat for marron bacora (Palumbo et al 2016, p. 5) (Figure 10).

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Figure 10. Map of St. John including habitat suitability model ranks, locations of sampled populations of marron bacora, other known population locations at that time and boundaries of the Virgin Island National Park (Palumbo et al., 2016, p. 6)

Chapter 4. Summary of Individual, Population, and Species Requirements

This section summarizes the requirements of individuals, populations, and the species, based on the information presented in Section II.

4.1 Requirements of Individuals.

4.1.1 Habitats.

Acevedo-Rodríguez (1996, p. 415) described the habitat of marron bacora as a dry, deciduous forest, and the species was reported to occur on dry, poor soils (Ray and Stanford 2005, p. 6). In addition, the species was further described as locally abundant in exposed topography, on sites disturbed by erosion (depositional zones at the toe of the slopes), areas that have received moderate grazing, and around ridgelines as an understory component in diverse woodland communities (Carper and Ray 2008, p. 1). A habitat suitability model suggests that the vast majority of marron bacora habitat is found in the lower elevation (<85m, 278.87 ft ) coastal scrub forest and that about 32% of the land area of the VINP harbors suitable habitat for the species (Vilella and Palumbo 2010, p. 10). The largest natural populations (almost all individuals of the species) are located along a single gut (valley) in the sub-watershed of Drunk Bay, a part of the Coral Bay Watershed. The Coral Bay Watershed is characterized by its steep slopes, highly erodible soils, and high runoff volumes during average rain events (Coral Bay Watershed Plan 2008, p. 6). After several re-introductions, only 54 individuals out of approximately 243 have 31

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survived (22.2%) and two populations that had 120 of those 243 re-introductions are now considered extirpated.

4.1.2 Reproduction.

The system of breeding in marron bacora is very likely to be that of an obligate outcrosser with self-incompatibility (Stanford et al. 2013, pp. 174; Anderson et al. 2015, pp. 479). Recent findings support the hermaphroditic and dioecious biology of marron bacora (Anderson et al. 2015, p. 479). Nonetheless, Eleanor Gibney (horticulturist and specialist in the flora of St. John) has recorded fruit production on isolated plants, suggesting the species still has mechanisms for self-pollination (pers. comm. 2017). Therefore, any restoration efforts must ensure that both sexes are proportionally present and in sufficient quantities to secure good seed set (Anderson et al. 2015, p. 484).

The greatest present threat to marron bacora appears to be the lack of natural recruitment (Palumbo et al. 2016, p.2). Depredation of marron bacora fruit by feral ungulates may be largely responsible for the low levels of seedlings recruitment. Thus, the Service anticipates that successful recruitment may require low densities of ungulate browsers. As suggested by Vilella and Palumbo (2010, p. 12) exclosure experiments may be developed to better understand the seed and seedling predation hypothesis. Currently (2018) there are cameras in the field to help collect this data; however, camera imagery is hard to verify what is actually being consumed. Native hermit crabs (Coenobita clypeatus) have also been observed depredating fallen marron bacora fruit (Ray and Stanford 2005, pp. 18; Vilella and Palumbo 2010, p. 18; Yrigoyen and McKinley 2017, pers. obs.) and could possibly be natural dispersers. No observations of fruit depredation by bats have been documented.

4.1.3 Precipitation.

The majority of the marron bacora habitat lies within the subtropical dry life zone, which is characterized by low annual rainfall and a high evapotranspiration ratio (Ewel and Whitmore 1973, p.10). In fact, more than 80% of the St. John surface is considered as subtropical dry forest (Stanford et al. 2013, p.173). According to Ewel and Whitmore (1973, p. 10), the vegetation in the subtropical dry life zone tends to form a complete ground cover and is almost completely deciduous. As an endemic to the Virgin Islands (St. John and Tortola), marron bacora is adapted to these environmental conditions, and the species phenology is synchronized with rainy season. Recent assessments by the Service coincided with a massive flowering event following a rainy event at St. John (USFWS 2017a, p. 7). Most of the yearly rainfall on St. John occurs between May and December with official hurricane season from June 1 through November 30 (Figure 11).

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Figure 11. Current average rainfall on St. John (www.weather-us.com/en/united-states-virgin-islands-usa/saint-john-climate)

4.2 Requirements of Populations.

4.2.1 Minimum viable population size and density.

The marron bacora minimum viable population (smallest population size that has a high probability of surviving a prescribed time) is unknown. Currently, population sizes range from one to 201 individuals reported at the seven known locations on St. John, however, the ratio of male to female that make up the natural populations is unknown. According to Stanford et al. (2013, p.178) the genetic diversity within the species remains relatively high and further highlights that low genetic diversity may not be a serious barrier to survival in this species. Nonetheless, the healthiest population at Nanny Point is currently characterized by an old population structure with little evidence of natural recruitment. In addition this population showed a decline of about 48 % of the number of known wild plants when comparing to the information available at the time of the species 12-month finding in 2011 (USFWS 2017a, p. 7). The smallest population known to produce fruit is at Reef Bay Trail where there are only seven individuals; however this site shows no evidence of natural recruitment (USFWS 2017a, pp. 11). The best available information suggests extirpation for at least three populations; Reef Bay Valley, Europa Ridge and Sabbat Point (USFWS 2017a, p. 11). Therefore, based on the currents threats at St. John, and the fact that the biggest natural population (Nanny Point) was steadily declining over the last decade we can infer that viable populations should contain at least 200 reproductive individuals, along with evidence of recruitment. Currently the species is under ex- 33

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situ conservation at the Fairchild Tropical Botanic Garden (Florida), the St. George Village Botanical Garden (St. Croix) and the J.R. O’Neal Botanic Garden (Tortola).

4.2.2 Size of habitat patches.

As plant populations become reduced and spatially segregated, important life-history needs provided by pollinators and seed dispersers may be compromised (Kearns and Inouye 1997, p. 299). Marron bacora flowers lack floral nectaries, thus, the only reward to pollinators is the pollen itself (Anderson et al., 2015, p. 472) and in 2015 Anderson et al. (2015, p. 475) reported observing no pollinators on St. John at any time over dozens of hours in the field. Anderson et al. (2015, pp. 475) also explains that the pollinator guild(s) and behavior remain to be studied. Little is known regarding the natural history (i.e., pollinators and seed dispersal) and basic ecology of marron bacora, including effects of herbivory (Palumbo et al., 2016, p. 1). As previously mentioned, by 1717 the forested landscape of St. John was parceled into more than 100 estates for agriculture and the majority of the landscape was deforested (Stanford et al. 2013, p. 173). Despite the current protection of about 56% of the island as part of the Virgin Islands National Park (Stanford et al. 2013, p. 173), populations of narrow endemic species (i.e., Solanum conocarpum, Zanthoxylum thomasianum and Calyptranthes thomasianum) were fragmented due to former deforestation as part previous agricultural land use in St. John. Therefore, considering the reproductive biology of marron bacora, maintaining forest connectivity and habitat corridors between known populations is critical for the long term conservation of the species, and to ensure that ecological interaction with native pollinators and dispersers remains functional.

4.3 Species Requirements.

The viability of a species is assessed in terms of the three conservation biology principles of resilience, redundancy, and representation (Shaffer and Stein 2000, pp. 307–310). Resilience often refers to population sizes which influence the capacity for a population to sustain itself over time; larger populations are more likely to persist than smaller ones. Redundant populations increase the species’ chances of surviving catastrophic events. Representation refers to the breadth of genetic diversity necessary to conserve long-term adaptive capability. With regard to resilience, additional data would be necessary to estimate a minimum of individuals necessary for a viable population and in this case with a minimum ratio of males to females that constitute that minimum. The best available information does not indicate what the minimum viable degree of representation and redundancy should be; it is reasonable to conclude that more is better. In order to ensure the survival of the species, all populations should be brought up to sizes large enough to allow outcrossing, and ex-situ conservation should be employed (Stanford et al. 2013, p. 173).

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Needs of Current Condition of Solanum concarpum Solanum conocarpum

INDIVIDUALS NEED suitable habitat of: INDIVIDUAL CONDITIONS: . Vary between populations. Dioecious obligate Caribbean dry, deciduous forest. Occurs on dry outcrossers and self-incompatible. Unknown poor soils in lower elevation (˂ 85 m) coastal scrub pollinators, unknown dispersers. forests. Shows little fidelity to any particular suite of POPULATION CONDITIONS: community associations. Preference may involve abiotic factors. All populations declining, low viability with Reach maturity in 2 years under greenhouse potential for extirpation. conditions but not in wild. 324 known individuals, 201 located at 1 of 8 locations. POPULATIONS NEED: RANGE-WIDE CONDITIONS:

Continuous suitable habitat in available Very low to no natural recruitment. 3 of 8 dry forest, preferably on NPS land, to support populations have over 18 indv., 2 populations have large, resilient populations. 18 indv. and 3 populations have less than 7 indv. RANGE-WIDE SPECIES NEED: insufficient Small and isolated populations with little to no connectivity. Multiple resilient populations are needed to Exotic mammals and insect pests. increase connectivity, thus, increasing redundancy across the range.

Primary Stressors Future Condition (Viability) of Solanum conocarpum Solanum conocarupum

MAIN STRESSOR: HABITAT RESILIENCY IS LOW

Significant reduction of suitable habitat due to Populations too small to be resilient and are early land clearing and development. No highly vulnerable to future extirpation. connectivity. Climate change No natural recruitment. Hurricanes increase in frequency Feral ungulates and herbaceous pests present and ferocity. Unknown pollinators and dispersers. Sea Level Rise is minimal threat. Drought conditions predicted to MAIN SOURCES: increase. Heat stress.

REDUNDANCY IS LOW Mammalian species (Goats, deer, donkeys, pigs, etc.). Invasive plant threats (Guinea grass, Only 1 population with over 46 individuals. Tan-tan, introduced vines). (populations are too small and isolated and have a Genetic consequences of small population low probability of persistence). (limited gene flow). REPRESENTATION IS LOW Small populations only 1 individual. Habitat loss may include loss of pollinators and dispersers. Genetic diversity at species level is normal only Threats of development. in the largest population. Loss of habitat may continue into the Genetic diversity is low at the population level. future, important to establish populations on protected land. OVERALL SPECIES VIABILITY IS LOW.

Figure 12. Rapid View of the SSA Assessment for marron bacora. 35

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Chapter 5. Factors Affecting the Survival of marron bacora: Threats and Vulnerabilities

5.1 Factors Affecting the Viability of the Species

The following list describes factors that affect the continued survival of marron bacora. These largely interrelated factors are not listed in order of severity.

5.1.1 Demographic consequences of small population size and density.

Small populations are less likely to recover from losses caused by random environmental changes (Shaffer and Stein 2000, pp. 308–310), such as fluctuations in recruitment (demographic stochasticity) and variations in rainfall due to severe drought (environmental stochasticity). The majority of marron bacora natural populations are small and do not show evidence of natural recruitment. The Service’s 2011 12-month finding (76 FR 9722) indicates that habitat modification may result in irreversible damage to the species’ natural habitat, decreasing the number of individuals in already small populations. Marron bacora is also threatened by the lack of natural recruitment, lack of connectivity between populations, and habitat modification by exotic mammal species and may also be threatened by the absence of dispersers. Thus, small populations may be easily wiped out by natural disasters (e.g., hurricanes) or exotic pests. The species reproductive biology (functionally dioecious) further reduces the cross pollination, fruit production and likelihood of natural recruitment to allow populations to recover and results in a downward spiral toward the extirpation of small populations.

5.1.2 Genetic consequences of small population sizes.

Along with a decreasing population size, negative impacts of anthropogenic habitat fragmentation may result in erosion of genetic variation through the loss of alleles by random genetic drift (Honnay and Jacquemyn 2007, p. 823). Johansson et al. (2005, p. 1676) describes a high degree of gene flow in a continuous landscape compared to a fragmented landscape. Although population genetic analysis revealed that species level diversity in marron bacora remains high, the majority of the species diversity is confined to a single population (i.e., Nanny Point). Decrease in population size may lead to genetic drift in small populations resulting from a loss of alleles occurring at a low frequency (Mhemmed et al. 2008, p. 751). Moreover, the research by Anderson et al. (2015, p. 479) confirm the hermaphroditic and dioecious biology of marron bacora which may result in the functional extinction of small isolated populations, and thus the loss of unique alleles contained in these populations.

5.1.3 Human-induced fires.

In the Caribbean, native plant species, particularly endemics with limited distribution, may be vulnerable to man-made events such as human-induced fires. Fire is not a natural component of subtropical dry forest in Puerto Rico and the Virgin Islands; thus, most species found in this type 36

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of forest are not fire adapted and are not likely to withstand frequent fire events (76 FR 9722, p. 9726). Marron bacora is associated with lower elevation dry forests. This habitat may be susceptible to forest fires, particularly on private lands, where fire could be accidentally ignited. Furthermore, regenerating forests, such as the ones prevalent in St. John, are prone to wildfires promoting the development of persistent shrub land dominated by introduced tree species and grasses (Wiley and Vilella 1998, p. 340). Studies conducted within the Guánica Forest in southern Puerto Rico indicate that some exotic tree species can remain as a dominant canopy species for at least 80 years (Wolfe 2009, p. 83). Given the growth habit of marron bacora, it is unlikely that individuals would survive a fire even of moderate intensity (Vilella and Palumbo 2010, p. 15), and, therefore, the species might be outcompeted from the habitat intrusion by exotics plants. However, a site visit to St. John to evaluate the threats to the species, found no substantial evidence indicating fires posed an imminent threat to the species (Monsegur, pers. obs. 2010). The site that is more vulnerable to fires is John Folly, due to its proximity to a road and the accumulation of debris associated with a former house (Monsegur, pers. obs. 2010).

5.1.4 Climate Change and Hurricanes

Hurricanes and tropical storms frequently affect the islands of the Caribbean and, thus native plants should be adapted to such disturbance. In fact, successional responses to hurricanes can influence the structure and composition of plant communities in the Caribbean islands (Van Bloem et al. 2005, p. 576). However, climate change is predicted to increase the frequency and strength of tropical storms, but also cause severe droughts (Hopkinson et al. 2008, p. 255). Climate model simulations indicate an increase in global tropical cyclone intensity in a warmer world, as well as an increase in the number of very intense tropical cyclones, consistent with current scientific understanding of the physics of the climate system (USGCRP 2018, 2: 8). The vulnerability of species to climate change is a function of sensitivity to changes and exposure to those changes, and the adaptive capacity of the species (Glick et al. 2011, p. 1). Within natural conditions it is likely that marron bacora is well-adapted to these atmospheric events. However, the cumulative effects of severe tropical storms and increased sediment runoff may jeopardize the establishment of seedlings along drainage areas usually associated with suitable habitat for marron bacora (Ray and Stanford 2005, p. 2) (Figure 13). In fact, the Service recently recorded evidence of direct impacts to the Nanny Point population due to the flash flood event associated to Hurricane Irma (USFWS 2017b, p.3)

5.1.5 Invasive Species

Feral exotic mammal browsers are found throughout the range of marron bacora on St. John Island. These include feral goats (Capra aegagrus hircus), pigs (Sus scrofa), white-tailed deer (Odocoileus virginianus), and donkeys (Equus asinus) (Vilella and Palumbo 2010, p. 5; Monsegur, pers. obs. 2010). Nonetheless, there have been control efforts targeting problem areas such as Reef Bay Valley, and feral hogs populations within NPS land are currently low (NPS

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Figure 13. Nanny Point pre 2017 hurricanes Nanny Point post 2017 hurricanes

2008, p. 2). However, hogs continue to be a problem at the Reef Bay area as they uproot the vegetation searching for food and water (76 FR 9722, p. 9727). The Service conducted a field assessment that confirmed the presence of exotic mammal species within marron bacora habitat, highlighting the abundance of the white-tailed deer and herds of feral goats (Figure 16) (76 FR 9722, p. 9727). Observations by Monsegur (2010) coincide with reports of high abundance of white-tailed deer within the range of marron bacora by Ray and Stanford (2005, p. 5), and with reports from the NPS that describe deer populations as increasing (NPS 2008, p. 2). Despite the reports of the intrusion of free-roaming ungulates within marron bacora populations (Ray and Stanford 2005, p. 5), there is a lack of information regarding the specific adverse effects of these exotic animals on the marron bacora. Nonetheless, the population assessment by Service staff in 2017 identifies individuals of marron bacora with rut marks (deep scars on the bark) likely from deer (USFWS 2017a, p. 7). In addition, these exotic mammals are probably modifying the structure of the vegetation that may result in changes in microhabitat conditions that may affect seed germination and seedling recruitment of marron bacora. Feral animal droppings were also documented in the area where marron bacora was described on Tortola, BVI (Heller et al. 2018, p. 4).

Camera traps were installed in the Nanny Point population in 2017 in an attempt to gather information on the effects of feral ungulates, birds and crabs on the population (Figure 16). Unfortunately, the damage caused by the two hurricanes in 2017 made it impossible to continue monitoring. In June of 2018, during a post hurricane assessment it was noted that 71 of 127 plants surveyed exhibited herbivory impacts from potential invasive mammals and deer or goat feces were observed throughout the population site (IC Report 2018, p. 5). During this assessment, two invasive plant species were also identified in the Nanny Point population, Guinea grass (Megathyrsus maximus) and Tan-tan (Leucaena leucocephala) (IC Report 2018, p. 3) (Figures 14 and 15).

Tan-tan had previously been identified by Dr. Ray who described it as an indicator of a high 38

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Figure 14. Distribution of guinea grass (M. maximus) in red and tan-tan (L. leucocephala) yellow at Nanny Point (IC Report, 2018, p. 12)

Figure 15. Marron bacora in guinea grass at Nanny Point (USFWS 2017b) level of recent disturbance; this species is normally replaced in the first four decades of forest recovery from acute disturbance (Ray and Stanford 2005, p. 11). In 2018, the Nanny Point population area was calculated to be approximately 5.4 km² (2.1 mi²) with 19 patches of guinea 39

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grass identified occupying 928.5 m² (9,994.3 ft²) and tan-tan occupying 80.4 m² (865.4 ft²) . The total are occupied by both invasives was approximately 1.0 km² (0.39 mi²) or 18.7% (IC Report 2018, p. 11) (Figure 14).

Figure 16. Deer at Nanny Point Foraging off of the ground near marron bacora

5.1.6 Insect Pests and Pathogens

Although known marron bacora populations are relatively protected, the small size of populations coupled with the effects of insect pests or pathogens could cause local extirpation. Stanford et al. (2013 p. 178) recorded the presence of an unknown pathogen affecting the Reef Bay population (currently extirpated). More recently, in 2018 Island Conservation reported 63.2% of the marron bacora individuals at Nanny Point show some sort of stem dieback, however it is not clear if this is due to some pest or disease (2018, p.5) . Nonetheless, observations by Service staff indicates that dieback is clustered mainly to the eastern corner of the Nanny Point population, associated to edge vegetation, vines and shrub land vegetation exposed to salt spray.

In addition, the assessment by Service staff in 2017 recorded the presence of the Jacaranda bug (Insignorthezia insignis) at the Nanny Point population (Figure 17), and Praelongorthezia praelonga (Douglas) and Insignorthezia insignis on plants at the gardens of the NPS facilities (USFWS 2017a, p. 14). The Jacaranda bug is a sap-feeding insect in the Orthezidae family. This insect is a pest of coffee but also feeds on foliage, shoots and fruit of citrus and herbaceous and woody hosts, and was identified as the main threat to a threatened species in Kenya, causing the death of approximately 400 of the 2,000 of remaining specimens of Commidendrum robustum (Schabel 2006, p. 99). The scale insect (Praelongorthezia praelonga) can also damage plants directly by sucking their sap, or indirectly by injecting toxic salivary secretions which may attract ants, transmit pathogens and encourages growth of sooty molds (Ramos et al. 2018, p. 273). However, no studies have been carried out to ascertain the extent of potential impacts by these pests on marron bacora. 40

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Figure 17. Jacaranda bug on marron bacora plant (Photo by Yrigoyen 2017) and fruit (IC Report, 2018, p.7) on marron bacora plant exhibiting loss of bark and yellowing of the leaves.

5.1.7 Phenology and Breeding System

As previously mentioned, Anderson et al. (2015, p. 479) confirmed the hermaphroditic and dioecious biology of marron bacora, as crossing of pollen to the stigma of other male flowers or transferred to the stigma of the same flower resulted in no pollination. Anderson et al. (2015, p. 475) also found a 1:1 sex ratio and a much longer time for marron bacora female plants to flower for the first time (from the seedling stage) compared with the males. At this point the natural disperser of marron bacora remains unknown, and possible fruit predation is suspected as the explanation of lack of natural recruitment in the wild (76 FR 9722, p.9725). It is possible that natural fruit dispersers of marron bacora have targeted other food sources as the populations of this shrub became increasingly patchy, as a result of deforestation and introduction of exotic plant species. The absence of a fruit disperser may also indicate that the disperser of the species is extinct or that the populations are too small to attract the disperser (Roman 2006, p. 82). The above information highlights the vulnerability of extirpation of relatively small populations of marron bacora as they may become functionally extinct and cannot support recovery or rescue of neighboring populations, limiting their value for redundancy and species resiliency.

5.1.8 Recreation.

Some evidence of damage consistent with trail maintenance was recorded on the individual along Brown Bay trail, and further habitat disturbance was observed at the John Folly site (park boundary) (76 FR 9722, p. 9724). Also, site disturbance (vegetation clearing) was recorded in 2017 at the John Folly population, where for example, one seedling on the middle of the trail is susceptible of being trampled by hikers (USFWS 2017a, p. 9). However, considering the remoteness of the marron bacora habitat and given that the majority of the populations are within NPS land, recreational uses have a low probability of affecting the survival of the species. 41

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5.1.9 Conservation Measures

Eleanor Gibney, horticulturist, propagated marron bacora by cuttings and reproduced them sexually by dusting the flowers (B. Kojis and R. Boulon, pers comm., 1996). She then distributed marron bacora specimens to various places in the Virgin Islands (Ray and Stanford 2005, p. 6). Gary Ray and Alice Stanford (2003) developed an implementation plan to conduct shade-house propagation of marron bacora using both seedlings and cuttings for reintroduction within the NPS land in St. John. A Nanny Point landowner funded and implemented a propagation program of marron bacora through germination and cloning of adult individuals to enhance natural populations of the species in Nanny Point, Brown Bay Trail and Johns Folly (Ray and Carper 2009, p. 6).

The National Park Service (NPS) has its own regulatory mechanisms to protect the species within the Virgin Islands National Park on St. John. The NPS is responsible under the Organic Act (16 U.S.C. § 1; NPS 2006) for managing the national parks to conserve the scenery and natural and historic objects and the wildlife. The National Park Omnibus Management Act of 1998 (Pub. L. 105-391, Sec. 1(a), Nov. 13, 1998, 112 Stat. 3497; NPS 2006), Title II, “National Park System Resource Inventory and Management” giving a research mandate to the NPS to support resource management decisions (16 U.S.C. 5936; NPS 2006). This law affects not only the NPS, but other federal agencies, universities, and other entities that conduct research in the National Park Systems. Currently, the NPS has implemented its resource management responsibilities through its Management Policies, Section 4.4, which states that “it will maintain as part of the natural ecosystems of parks all plants and animals native to the park ecosystem”. Section 207 of the Omnibus Management Act of 1998 allows NPS to withhold from the public information related to the nature and specific location of endangered, threatened, or rare species unless disclosure would not create an unreasonable risk of harm to the species (16 U.S.C. § 5937; NPS 2006).

The Territory of the U.S. Virgin Islands currently considers marron bacora as endangered under the Virgin Islands Indigenous and Endangered Species Act (V.I. Code, Title 12, Chapter 2), and has amended an existing regulation (Bill No. 18-0403) to provide for protection of endangered and threatened wildlife and plants by prohibiting the take, injury, or possession of indigenous plants.

In 2017, Island Conservation was awarded funding through the USFWS Coastal Program to 1) propagate at least 100 marron bacora individuals to enhance the largest known population at Nanny Point, 2) introduce propagated materials to the Nanny Point population , 3) assess the extent of invasive mammal species impacts to marron bacora and its habitat, 4) assess the extent of invasive mammal species to additional sites identified for marron bacora introduction, and 5) provide management recommendations for invasive mammals in order to significantly advance the recovery of marron bacora (IC Report 2018, p. 1).

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5.1.10 Summary.

Marron bacora is currently threatened by its overall low numbers of individuals, low numbers of populations, and low numbers of individuals at each population site. Thus, we deem the resiliency, redundancy and representation of this species are low. There is a lack of knowledge regarding the abundance and roles of dispersers and pollinators at the marron bacora population and species levels. Current knowledge of the ecology and genetic diversity of Virgin Islands rare flora is sparse (Stanford et al. 2013, p. 173). While the genetic diversity at the species level of marron bacora is relatively high, the majority of its diversity is confined to the largest population at Nanny Point (Stanford 2013, p. 178). Due to historic habitat destruction and modification,, the current fragmented population distribution of the species also attributes to the lack of genetic exchange between populations, which may eventually increase the risk of genetic drift. Another main threat to all the occurrences of marron bacora is the lack of natural recruitment most likely due to depredation of its fruits by feral ungulates; however, additional research would be necessary to evaluate this threat fully. As of November 2018, private property surrounding the largest population, at Nanny Point, is currently up for sale and urban development on this property may alter the habitat further. A previous plan to build a resort on the land was originally announced in 2016 (Manes 2016, blog), however, it never materialized. According to St. John Tradewinds (2017), the sale of the Concordia Eco-Resort had been cancelled due to an unnamed reason (Roberts 2017, website).

YEAR 2010- 2018 (post 2005 2014 2015 2016 2017 Hurricane 2011 Irma/Maria) LOCATION 144˟ 144˟ 144˟ 75 ˟ 108° Nanny Point ᶷ 184* 40^ 144˟ 44˘ 53ᵜ 50† 50† 50† 4ᶱ 40ᶱ 11 ˟ 11 ˟ 4 ˟ John’s Folly ᶷ 37† 11° 37† 11° 13 ˘ 18° 1ᶱ 1 ˟ 1 ˟ 1 ˟ 17° Brown Bay Trail ᶷ 1* 1 ˟ 1 ˟ 36† 36† 10 ˘ 1ᵜ

Friis Bay 33* 33 ˟ 33 ˟ 33 ˟ 33 ˟ 33 ˟ 33 ˟

Base Hill ᶷ 1* 1 ˟ 1 ˟ 1 ˟ 1 ˟ 1 ˟ 1 ˟ 43

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YEAR 2010- 2018 (post 2005 2014 2015 2016 2017 Hurricane 2011 Irma/Maria) LOCATION

Reef Bay Trail ᶷ 7 ˟ 7 ˟ Sabbat Point 1* 1 ˟ 1 ˟ 1 ˟ 1 ˟ X X 6* 6 ˟ X 6 ˟ 6 ˟ Reef Bay Valley ᶷ 60† 60† ? † 60† X X X ? † 1* 1 ˟ 1 ˟ Europa Ridge ᶷ 1 ˟ 1 ˟ 60† 60† 60† X X Brown Bay Ridge ᶷ 1* 1*

Sabbath Hill, Tortola, 46* BVI

227* 198 ˟ 181˟ 198 ˟ 187 ˟ 122 ˟* 230 ˟°* 120† 243† 11° 243† 11° 67 ˘ 54ᵜ Population Estimate 50† 5ᶱ 40ᶱ (347††) (441††) (242††) (441††) (198††) (194††) (324††) Table 1. Marron bacora poplulation estimates. Sources: Ray and Stanford 2005; USFWS 2010-2017, and IC Report-2018 Symbols: * wild, † introduced, ˟ natural, ^ lost due to translocation, ˘ planted, ° adult individual, ᵜ juvenile, ᶱ seedling, †† total, ᶷ NPS (Public Land), X reported extirpated Year Data Collected: 2005, 2010, 2014, 2017, 2018.

5.2 Climate Change

5.2.1 Background

According to the 2018 U.S. Global Change Research Program (USGCRP) report, the impacts of climate change are already influencing the environment and more frequent and intense extreme weather and climate-related events, as well as changes in average climate conditions, are expected to continue to damage ecosystems. Rising water and air temperatures are leading to changes in precipitation and intensifying droughts, increasing heavy downpours, reducing snow pack, and causing declines in surface water, with varying impacts across the US (USGCRP 2018, summary). The Caribbean is one of the regions considered threatened by drought, flooding and sea level rise. Climate change is also projected to alter the geographic range and distribution of

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pests and diseases (USGCRP 2018, summary). These stressors account for indirect and direct effects at some level to all life stages and across the species’ range.

Additive climate change stressors projected for the future include: a) increased number and intensity of strong storms, b) increased temperatures, and c) shifts in the timing and amounts of seasonal precipitation patterns. Recent works on climate change models for tropical islands, predict that, for example, by mid-21st century Puerto Rico will be subject to a decrease in rainfall, along with increase drought intensity (Khalyani et al. 2016 p. 265). Moreover, the work by Khalyani et al (2016) anticipates shifts in the life-zones with potential loss of subtropical rain, moist, and wet forest, and the appearance of tropical dry and very dry forests. Thus, considering the proximity of Puerto Rico to St. John, and that these islands belong to the same biogeographical unit (Puerto Rican Bank), we believe these model predictions could also extend to the USVI (including St. John). Given the low number of known populations and individuals of marron bacora, it potentially does not have the genetic breath to adapt to these predicted conditions. Nonetheless, it should be noted that the three main characteristics of a species that pose a particular challenge to predicting climate change vulnerability assessments are present with this species. There is little knowledge of its life history (e.g., fruit/seed dispersers and germination requirements in the wild), it has a restricted known range (e.g., mainly St. John), and its habitat is fairly degraded due free ranging populations of feral animals (e.g., deer and goats) (Foden and Young 2016, p. 22).

Given the uncertainty of predicting future conditions past 25 years, we consider climate change as a stressor that could realistically determine the long term viability of marron bacora. However, given the little knowledge on some aspects of the species’ life history, it is difficult to predict its vulnerability to climate change stressors. Nonetheless, data from controlled propagation show the species maintains high rates of viability, including a high germination rate (seed viability) and reaching a reproductive size in a short period (less than two years) under nursery conditions. Therefore, it is possible to make some assumptions about the potential adverse impacts from climate change on this species’ viability. In order to gauge potential future conditions affecting marron bacora and its habitat, three scenarios were examined based on local climate change predictions for the USVI and the Caribbean. The scenarios focus on a range of conditions predicted based on climate change scenarios and are meant to provide the uncertainty always associated with the actual species response to factors of climate change. The range of what is likely to happen in each scenario will be described based on the current condition and how resilience, representation and redundancy would be expected to change. Some stressors are anticipated to remain relatively stable and are not expected to change significantly from the current condition including NPS forest management practices, urban development, and feral ungulate presence. If any of these stressors are found to be increasing or placing greater stress on the species, it would have to be re-assessed. For example, if the land surrounding the largest known population at Nanny Point (where 72% of the known individuals reside) were to be slated for development, a reassessment of the species condition would be 45

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necessary. 5.2.2 Temperature, precipitation and drought stress

Temperatures in the U.S. Caribbean have fluctuated over the last 100 years, however, since 1950 temperatures have increased by about 1.5°F (16.94°C) in Puerto Rico (USGCRP 2018, 20: 819). Some climate projections (from 1960-2099) indicate a 4.6 to 9 °C (20.3 to 48.2 oF) temperature increase for Puerto Rico (Khalyani et al. 2016, p. 275) indicating a general consensus on a continued warming trend into the future amongst climate modeling studies for the entire US Caribbean including the USVI. Thus, temperature across the Caribbean is expected to continue increasing over the next century. Global climate models project about a 1.5°F (16.94°C) to 4°F (15.56°C) increase in average temperatures for the U.S. Caribbean in 30 years (year 2050) with end of the century (2100) estimates showing increases as high as about 9°F (12.78°C) under higher emission scenarios (USGCRP 2018, 20: 819) (Figures 18, 19, 20, 21). Precipitation is also projected to decrease, influenced by warming, and it will tend to accelerate the hydrological cycles, resulting in wet and dry extremes (Jennings et al. 2014, p. 4; Cashman et al. 2010, p. 1). It is reported that the Caribbean is expected to get more frequent and severe droughts from reduced precipitation and an increased evapotranspiration ratio due to climate change with an increase in the amount of precipitation produced during hurricane events (Herrera et al. 2018, p. 1). However, the majority of models predict future decreases in precipitation are likely (Carter et al. 2014, p. 399). Subtropical dry forests are already subject to water deficit for ten months of the year (Miller and Lugo, 2009, p. 86) and are expected to become drier in the future, especially in regions like the U.S. Caribbean where oceans have the largest influence on local precipitation (USGCRP 2018, 20: 820) (Figure 19). Climate model results consistently project significant drying in the U.S. Caribbean region by the middle of the century ((USGCRP 2018, 20: 820).

Figure 18. Observed and projected temperature changes are shown compared to the 1951-1980 average. Observed data are for 1950-2017, and the range of model simulations for the historical period is for 1950-2005. The range of projected temperature changes from global climate models is shown for 2006-2100 under lower and higher emissions scenarios. Projections from two regional climate models are shown for 2036- 2065, and they align with those from global models for the same period (USGCRP 2018, 20: 820). 46

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Figure 19. Projected Precipitation Change for the US Caribbean. This figure shows the projected percent change in annual precipitation over the U. S. Caribbean region for the period 2040-2060 (lower figure) compared to 1985-2005 (upper figure) based on the results of two regional climate model simulations. These simulations downscale two global models for the higher scenario and show that within-island changes are projected to exceed a 10% reduction in annual rainfall. Uncertainty remains as to the location of the largest reductions within the islands. Projections of precipitation change for the U.S. Virgin Islands are particularly uncertain because of model limitations related to resolving these smaller islands (Bowden et al. 2018 cited in USGCRP 2018, 20: 821).

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Figure (20). Temperature projections for Puerto Rico under three GCM scenarios. (Khalyani et al. 2016, p. 277)

Major consequences of warming, also include significant increases in the number of days in the Caribbean with temperatures over 90°F (32.2°C). For example, since 1970, the average annual number of days exceeding 90°F (32.2°C) has gone up an average of 0.5 days per year since 1970 (USGCRP 2018, 20: 821) (Figure 21).

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Figure 21. Days Above 90°F in Puerto Rico. This figure illustrates the deviation from the long-term (1970-2016) average annual number of days exceeding 90°F, based on data from eight climate stations in Puerto Rico (UPR cited in USGCRP 2018, 20: 837)

5.2.3 Life Zones The boundaries of life zones sensu the Holdridge System are based on three climatic measurements: annual precipitation, bio temperature and ratio of potential evapotranspiration to annual precipitation (Holdridge 1947, entire). Figure 22 and Figure 23 illustrate the distribution of the two major life zones (tropical moist and tropical dry forests) and forest distribution on St. John respectively. Dramatic shifts in the life zones with potential loss of subtropical rain, moist, and wet forest, and the appearance of tropical dry, and very dry forests are anticipated (Khalyani et al. 2016, p. 275). Although predicted life zone distribution illustrations are not available for the US Virgin Islands, Figure 24 shows the predicted life zone distribution changes in Puerto Rico (Khalyani et al. 2016, p. 277). Thus, we can predict such conditions extending to the USVI (including St. John). Such shifts will be the result of the conditions explained above under the “Temperature, precipitation and drought stress” section, and the capacity of marron bacora to adapt to such conditions is expected to be reduced due to the small number of populations and individuals.

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Figure 22. Holdridge Life Zones of St. John (DRYFLOR 2010, p. 27)

Figure 23. Forest type distribution on St. John, US Virgin Islands (DRYFLOR 2010, p. 24) 50

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Figure (24). Projections for life zone distribution changes in the Puerto Rico under three GCM scenarios. (From: Khalyani et al. 2016, p. 279)

5.2.4 Storm Frequency and Ferocity Reconstruction of the past 5,000 years of intense hurricane activity in the western North Atlantic

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suggests that hurricane variability has been strongly modulated by El Niño during this time, and that the past 250 years has been relatively active in the context of the past 5,000 years (PRCCC 2013, p. 31). Extreme events such as hurricanes, floods and droughts are projected to increase in frequency and intensity, particularly in the Caribbean Sea (USGCRP 2018, 20: 127) and Gulf of Mexico, which have the highest ocean heat capacity, and thus greater potential for storm development. The upward trend in intensity is related to an increase in sea-surface temperature. Tropical storms and hurricanes have become more intense during the past 20 years, and hurricane winds speeds and rainfall associated with hurricanes are likely to increase as the climate continues to warm. It is stated that hurricanes may play an important role shaping the forest structure of dry forests within the Caribbean by promoting the multiple-stemmed condition prevalent within pristine or relative undisturbed areas (Van Bloem et al., 2005 p. 571). However, the increasing hurricane intensity and frequency coupled with species showing reduced populations, low number of individuals, habitat degradation and fragmentation may have adverse consequences for native species and their habitat. Although heavy rainfall associated with storms is expected to increase, shifting weather patterns have caused total rainfall to decrease in the Caribbean, thus resulting in more pronounced seasonal droughts (EPA 2016, p. 1).

Figure 25. Historical Atlantic Hurricane Tracks through Caribbean and USVI (NOAA 2019)

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5.2.5 Sea Level Rise As the ocean warms the sea level is likely to rise one to three feet in the next century (EPA 2016, entire). In 2012, NOAA conducted a review of available research on global sea level rise projections, and concluded that there is a high confidence that the mean sea level rise will be at least 8 inches (0.2 m) but no more than 6.6 feet (2.0 m) by 2100 (Lindsey 2018, p. 5). Rising sea levels combined with hurricanes may produce storm surges in the Caribbean that will inundate low-lying areas and may threaten marron bacora populations adjacent to the sea (e.g., Nanny Point and Brown Bay Trail). Such exposure to hurricane winds, drought stress and increased soil salinity may be adverse for these populations. For example, Florida Keys have shown increased mortality of larger stem diameter trees due to increased salinity and water stress following hurricane induced storm surge events (Sah et al., 2010, p. 12). Same patterns have been observed along coastal ecosystems (e.g., dunes and mangrove ecosystems) in Puerto Rico following Hurricane María (2017) (Monsegur pers. obs., 2018). However, the remaining known populations of marron bacora (including the population recently located at Tortola, BVI) are located in areas that should not be affected by sea level rise because they are located a higher elevations. 5.2.6 Potential Threats to Ecosystems Globally, tropical dry forests are one of the most threatened forest types. Limited land use and high population pressure has resulted in the loss of over 60% of these forests and unfortunately little is known of the ecological responses of tropical dry forests to climate change (Nelson et al. 2018, p. 1337). Insular dry tropical forests are distinct from continental forests due to their high species-level of endemism and failure to protect these forests would result in major losses of unique species diversity (DRYFLOR 2016, p. 1385). Some populations of marron bacora, such as Brown Bay Trail, are already exhibiting signs of heat stress and lack of water (IC Report 2018, p. 9). These already stressed systems will be exposed to additional stressors that could result in unfavorable changes to the structure and function of existing habitats. For example, the expected increase in severe droughts may pose several stressors to marron bacora due to reduced seedling survival (changes in microhabitat conditions) and from increased predation by feral browsers. Thus, it is anticipated that some ecosystems and species will be able to acclimate to change in environmental conditions than others. As stated above, one of the biggest threats of climate change to the future viability of marron bacora are the predicted shifts in the tropical moist and dry forest life zones. The boundaries of life zones in the Holdridge System are based on three climatic measurements: annual precipitation, bio temperature and ratio of potential evapotranspiration to annual precipitation (Holdridge 1947, p. 367-378). Figure 22 shows the distribution of the two major life zones in St. John, tropical moist and tropical dry forests. In general, dramatic changes are projected in the life zones throughout the Caribbean. This shift from wet and dry zones to drier conditions may have acute impacts on the ecosystem where marron bacora occurs, hence on its survival.

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It has been concluded that the geographic ranges of many terrestrial and freshwater plant and animal species have moved over the past several decades in response to warming; approximately 17 km (10.5 mile) per decade poleward, and 11 m (36 ft) up in altitude per decade (IPCC 2018, p. 218). Such shifts in species distribution may be critical in the Caribbean, where habitats are already limited due to the size of the islands and habitat fragmentation due to land use history. Pollinators such as bees have been shown to retain significantly greater geographic ranges under 1.5° C (34.7° F) global warming as compared with 2°C (35.6° F) (IPCC 2018, p. 218). As the climate changes, there is also the potential that highly adaptive invasive species may be able to establish in new areas or in some cases when some species (pests or diseases) move into areas which become climatically suitable, they may become invasive or harmful to the system (IPCC 2018, p. 244). In fact, species such as the exotic guinea grass (Megathyrsus maximus) are naturalized within the Caribbean and has become very effective colonizing dry forest areas and promoting habitat conditions favorable for human induced fires. Such human changes in the vegetation structure and the vulnerability to human induced fires alter the natural ecosystem and microhabitat conditions necessary for marron bacora.

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Chapter 6. Future Climatic Scenarios These future scenarios are based on available climate change data for the US Virgin Islands and the Caribbean region. Although there is increasing research on climate change impacts on other Caribbean ecosystems, such as beach and coral reef habitats, a literature review in 2018 of the ecological response of tropical dry forests to climate change found only two such studies (Nelson et al. 2018, p. 1338). Therefore, this assessment of future viability includes qualitative descriptions of the likely impacts of climate change and takes into consideration the uncertainty in marron bacora’s response to climate stressors. 6.1 Precipitation and Drought Conditions Herrera et al. (2018) reported that the Caribbean is expected to get more frequent and severe droughts from reduced precipitation, and increased evaporative demand due to climate change with an increase in the amount of precipitation produced during hurricane events. Anthropogenic warming accounted for approximately 15-17% of the drought’s severity and 7% of its spatial extent. This research also suggests that warming has already increased and there is a drought risk for the Caribbean (Herrera et al. 2018, p. 1). In accordance with an overall drying trend, an increasing drought risk is projected for Small Island Developing States (SIDS) and moderate to extreme drought conditions are projected to be about 9% longer on average for 2°C (35.6° F) versus 1.5° C (34.7° F) for islands in this region (IPCC 2018, p. 234). 6.2 Life Zones In terrestrial ecosystems, impacts to habitats due to climate change began in the past decades. Risks to unique and threatened terrestrial ecosystems are generally higher under warming of 2°C as compared with 1.5°C. In the case of unique ecosystems, these warming trends may lead to biome shifts and species range loss due to the species inability to adapt to these levels of warming (IPCC 2018, p. 251). The number of species projected to lose over half of their climatically determined geographic range (about 18% insects, 16% plants, 8% vertebrates) is reduced by 50% (plants, vertebrates) or 66% (insects) at 1.5°C versus 2°C of warming (IPCC, 2018, pp. 179). A recent meta-analysis of 27 studies concerning a total of 976 species found that 47% of local extinctions (extirpations) reported across the globe during the 20th Century could be attributed to climate change and it is significantly higher for tropical regions (IPCC 2018, p. 2018). In this context, the expected future Tropical Very Dry Forest extending to St. John (USVI) may include higher temperatures, reduced precipitation and pronounced drought stress conditions, thus reducing the capacity of marron bacora to adapt to such conditions and likely resulting in the extirpation of the species. 6.3 Summary In general, global carbon emissions will continue to rise, temperatures will continue to rise, and lower precipitation rates are expected. These predicted changes will most likely bring about a loss of forests and species having to adapt to higher latitudes/altitudes and a shift in the overall 55

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biome. In addition to these predictions, shifts in the timing of phenological events, pollinator movements to higher altitudes, highly adaptive invasive species and unregulated feral ungulates could lead to a major loss of marron bacora habitat on St. John.

6.4 Future Scenarios We have considered the best scientific and commercial data on the biological and ecological needs of the marron bacora, including the factors driving the historical and current conditions of those needs. In this Chapter, we now consider how those factors may affect future conditions for the species, relative to its continued viability as a species.

Thus, considering the above, the following three hypothetical scenarios are proposed for marron bacora on a 10, 25, and 50 year time frame. This period, considers the species habitat requirements, its reproductive biology, patterns of natural recruitment for known populations (e.g., Nanny Point), and ongoing threats that we anticipate will extend and will increase into the future. The proposed scenarios are as follows; Scenario I, Scenario II, and Scenario III. Due to the amount of uncertainty in predicting local factors over time, predictions of future conditions beyond 50 years are based on climate change, sea level rise and increase in stochastic events.

6.4.1 Scenario I

Scenario I is based on the prediction that conditions continue along their current trajectory. Threats to the habitat and populations remain as currently over the projected time. Numbers of known individuals remain the same, and threats from development, feral ungulates, and insect depredation remain present. Population sizes in the areas where threats are occurring are currently in a state of decline, and will typically continue along this trend if populations remain unmanaged and not artificially augmented.

6.4.2 Scenario II

Scenario II is based on the prediction that habitat and environmental conditions improve over time. Under this ideal scenario, populations occurring on private lands or along the boundaries of NPS lands are effectively protected thru conservation easements or land-owner’s agreements (e.g.,) through consultation and technical assistance (Section 7 of the Act). In addition, an effective program for the management and control of feral ungulates (e.g., deer and goats) is implemented in order to minimize predation on marron bacora. Additionally, populations that show low number of individuals and no recruitment are augmented to self-sustainable levels by establishing a robust propagation and planting program. Current predictions of climate change models (Khalyani et al. 2016, entire) fail to reach the expected threshold on rainfall reduction and temperature increment, thus, not exposing marron bacora to conditions that further preclude its natural recruitment. Ongoing surveys in neighboring islands (e.g., Tortola and Virgin Gorda) identify new populations with unique genetic variability.

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6.4.3 Scenario III

Scenario III is based on the prediction that habitat conditions continue to deteriorate over time. Under this scenario, no reintroductions or population enhancement occurs, and urban development and associated disturbance are occurring on the populations (e.g., habitat fragmentation and encroachment by exotic plants). In addition, populations of feral ungulates (e.g., deer and goats) remain unmanaged and continue increasing, maximizing the threat predation threat on marron bacora. As predicted by climate change models, the islands of the Caribbean face a reduction on the average rainfall and an increase in temperatures towards the mid-21st century, resulting on shifts of current life-zones that will potentially lead to a “Tropical Very Dry Forest” (Khalyani et al. 2016). These climate projections will likely minimize seedling establishment and recruitment due to drought stress, and will likely aid in the establishment of invasive/exotic plants, increasing the potential for predation by feral ungulates during severe droughts.

6.4.4 Likelihood of Scenarios

Considering the current status of the resources, governmental priorities and conservation efforts, Scenario I is the most likely scenario for the future. After Scenario I, the likelihoods of Scenario II and Scenario III are contingent upon decisions, resources and management priorities, which are difficult to predict. Scenario III is likely if funding and conservation efforts are not directed to an ongoing propagation program, habitat protection/restoration, and feral ungulate/pest management efforts. Scenario II is likely if liberal funding is directed toward initiatives described under this scenario, and climate change projections do not reach expected results.

6.5 Future Resilience, Population Level

All factors (e.g., propagation efforts, management of feral ungulates, evidence of natural recruitment, habitat intrusion by exotic plants, insect pests and potential impacts from climate change) that may influence the predicted future resilience of a species must be considered. For 10, 25 and 50 year projections, we may predict the resiliency of populations by summing together scores for each habitat or population metric. However, when population trends suggest extirpation is likely, though uncertain, resiliency for the population is forced Low or Extirpated, with a corresponding numerical score of -4. This numerical value demonstrates that although extirpation is not predicted with certainty, the condition of the population is expected to be worse than others receiving a low resilience score (no other scores below -2). Due to high degrees of uncertainty about all of the considered factors further into the future, the only factor for later resilience is due to climate change related impacts.

We generated resiliency scores for marron bacora by combining scores for 3 habitat metrics (Protection/Development Risk, Feral Ungulates, and Pest Depredation), and one population metric (Population Size and/or Trend, dependent on availability). The best available information 57

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for each population was gathered from the literature and species experts. Each metric was weighted equally; with the overall effect that habitat (i.e., Protection/Development Risk, Feral Ungulates, and Pest Depredation) weighted higher than population size/trend (1 metric). There have been no periodic monitoring established to estimate marron bacora population sizes and trends, and surveys since 2005 have sporadic. Survey methodology and reporting has varied from population to population, with survey results given as estimated abundances, estimated densities, and convenience sampling. Even with the same methodology and reporting, survey success can differ based on external factors like weather conditions, surveyor experience, etc. All of the above-described factors in combination contribute to high levels of uncertainty in what current and past population sizes truly are/were, and how they have changed over time. This is why resilience classifications relied more heavily on habitat conditions than population size and trend estimates. For each metric, populations were assigned a score of -1, 0, or 1, as described below in Table 2.

Habitat Metrics Population Metric

Habitat Protection/ Feral Pest Presence/ Population Score Development Risk Ungulates Depredation Size/Trend

-1 Relatively low Habitat not Exotic Pests population size protected, at risk of Mammals Present and/or declining being developed High High trend

0 Relatively moderate Some habitat population size Pest protected, some at and stable trend, Unknown or Present risk of being OR High degree of Moderate Moderate developed (Both) uncertainty in population size/trends

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1 Exotic Relatively high Habitat protected mammals Pests population size absent Absent and/or growth

Table 2. Description of how habitat and population factors were scored to determine marron bacora population resilience.

The scores for each population across all metrics were summed, and final population resilience categories were assigned as follows:

Low Resilience: -4 to -2 Moderately Low Resilience: -1 Moderate Resilience: 0 Moderately High Resilience: 1 High Resilience: 2 to 4

Below, we describe each population and the conditions contributing to their resilience classifications.

6.5.1 Nanny Point (Current Resilience – Low)

Under future Scenario I, resilience at Nanny Point is expected to be very low in 10 to 25 years and extirpated in 50 years. Since 2002 the Nanny Point marron bacora population has been described as the healthiest and largest. Nonetheless, after 15 years of monitoring by local experts and the Service, the overall population shows a clear decline in the number of individuals along with an old population structure, and little or no evidence of seedling recruitment. It was not until recent (following Hurricane Irma in 2017) that substantial seedling occurrence was recorded, probably associated to extreme rain events. However, those seedlings have been affected by predation by feral ungulates (e.g., deer) (IC Report 2018, p. 7). Therefore, further long term monitoring is needed to effectively determine if those seedlings are recruited into the population with positive long-term impact on the species. Despite the parcel that contains the marron bacora population has been transferred to NPS and set for conservation, the surrounding habitat is on private lands and remains vulnerable to urban and tourism development (76 FR 9722, p. 9723). In fact, development plans have been advertised for the neighboring properties, so the area is still considered threatened by development in the near future. Further habitat modification and fragmentation at Nanny Point may adversely affect the genetic exchange (cross-pollination) with other natural populations (e.g., Johns Folly), and may further reduce suitable habitat needed for seedling recruitment. Under Scenario I, the vulnerability to hurricane storm surges and the current population size/trend remain the same as the current condition.

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Under Scenario II, we consider the habitat can be effectively protected through consultation and technical assistance (section 7 of the Act), and by providing habitat conservation measures to private landowners (e.g., minimizing the deforestation to the footprint of the development and by maintenance of forested corridors). In addition, other conservation mechanisms may include land acquisition or conservation easements. Scenario II also considers the management or control of populations of feral ungulates, along with pest management and ongoing population monitoring. Under this scenario the marron bacora population grows or remains stable, increasing resiliency from very low to moderate in 50 years. Under Scenario III, development takes place, which can destabilize habitat at higher elevations above Nanny Point; increase runoff through the population, limit population expansion, and feral ungulates and pests would be still present causing dramatic marron bacora population declines. All these adverse effects can lead this population to extirpation. Therefore, the predicted resiliency of this population is very low to extirpated in 25 years.

A major hurricane (e.g., Irma and Maria, 2017) coupled with 0.35 m (1.15 ft) of SLR could have catastrophic effects on low-lying populations. The Nanny Point marron bacora population is relatively close to the shore and with an increment on SLR it can be directly affected by the storm surge and may result in hypersaline soils. For example, Hurricane Irma cause surge that reached this population, but since it also was impacted by flash flood, it is difficult to determine the magnitude of the impacts from storm surge. Therefore, we believe that SLR and hurricanes would likely lead to extirpation of marron bacora at Nanny Point. Scenarios are summarized for Nanny Point in Table 3, Appendix A.

6.5.2 John’s Folly (Current Resilience – Low)

Under Scenario I, the John’s Folly population is expected to be extirpated in 50 years. The population at John’s Folly is located just along the boundary of NPS land. Over the past decade this population has been artificially sustained by a population enhancement effort, and from the originally 11 known natural individuals (76 FR 9722, p. 9724), only about four remain alive according to one of the latest assessments (USFWS 2017a, p. 9). There are currently about 14 (78%) individuals corresponding to planted material that are located within NPS lands, the remaining 4 natural individuals (22%) are located at private lands. This population was augmented by planting approximately 37 individuals between 2010 and 2011, thus, the currently surviving 14 plants represent a roughly 38% survival. Natural recruitment has been virtually absent at this population with only one seedling being identified since 2005 (USFWS 2017a, p. 9). Thus, current trends suggest this population will be extirpated in the foreseeable future between 10 and 50 years. Under this scenario the vulnerability to hurricane storm surges and the current population size/trend would remain the same as the current condition.

Under Scenario II, we consider the habitat can be effectively protected thru consultation and technical assistance (section 7 of the Act), and by providing habitat conservation measures to private landowners (e.g., minimizing the deforestation to the footprint of the development and by 60

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maintenance of forested corridors). For example, due to the proximity of this population to suitable habitat that harbors populations of other federally listed species (e.g., Zanthoxylum thomasianum), land acquisition or conservation easements may be mechanisms for the conservation of this site. Scenario II considers the management or control of populations of feral ungulates, pest and invasive plant management, and ongoing population monitoring. However, considering the current precarious status of this population, even after enhancement efforts it's the resiliency will remain low. Under Scenario III, development takes place leading to an increased runoff through the population, intrusion of exotic plant species,feral ungulates, and pests, causing population declines, limiting natural recruitment, and potentially leading to extirpation. Thus, this population is predicted to be extirpated in 10 to 25 years.

The John’s Folly marron bacora population is located at a high elevation protected from storm surge and SLR. Therefore, we believe storm surge from hurricanes and SLR will not likely lead to extirpation of marron bacora at this location. In fact, available information indicates that the most recent hurricane (i.e., Irma) did not cause direct mortality to individuals (IC Report 2018, entire). However, since this population is located on a steep drainage, it may be affected by erosion and sedimentation resulting from flash floods. Scenarios are summarized for John’s Folly in Table 4, Appendix A.

6.5.3 Brown Bay Trail (Current Resilience – Low)

Under Scenario I, the Brown Bay Trail population is expected to be extirpated in 10 to 25 years. According to Ray and Carper (2009, p. 3), the population consisted of one large individual, which was augmented by planting 36 additional individuals, totaling 37 individuals in 2011. As of 2017, the population was reported to consist of 18 individuals located along a hiking trail within NPS lands. The largest individual (natural plant) shows evidence of pruning due to trail maintenance (76 FR 9722, p. 9724). During surveys in 2018, some of the individuals of this population appeared to be stressed by lack of water and excessive heat (IC Report, p. 9). Natural recruitment has not been reported at this population despite enhancement efforts, thus we expect the population to be extirpated within the foreseeable future (i.e., less than 50 years). Under this scenario, the vulnerability to hurricane storm surges and the current population size/trend will remain the same as the current condition. Under Scenario II, an effective trail management plan that accounts for marron bacora is developed and implemented, and thus trail maintenance does not adversely affect the species. In addition, feral ungulate and pest management are underway, along with population monitoring. However, considering the current status and the lack of natural recruitment after enhancement efforts, the population may slightly grow, but resiliency will remain low. Under Scenario III, feral ungulates and pests remain present causing dramatic population declines, limiting natural recruitment and potentially leading to extirpation. Thus, the population is predicted to be extirpated in 10 to 25 years.

The Brown Bay Trail population is located at an elevation protected from storm surge. Therefore, we believe SLR and hurricanes will not likely lead to extirpation of marron bacora at

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this location. In fact, available information indicates that the most recent hurricanes Irma and Maria did not cause direct mortality to individuals; neither were there reported losses due to storm surge caused by these hurricanes (IC Report 2018, entire). Therefore, we believe that SLR and hurricanes will not likely lead to extirpation of the marron bacora population at Brown Bay Trail. Scenarios are summarized for Brown Bay Trail in Table 5, Appendix A.

6.5.4 Friis Bay (Current Resilience – Unknown)

The most recent assessment of this population was in 2005. With no subsequent visits, no current data regarding its status or threats are available. Given the declining status of the species, and assuming that threats to this location are consistent with all other population locations, we predicted scenario outcomes.

Under Scenario I, the Friis Bay population is expected to be extirpated in 10 to 25 years. This population is located on a private property, and in 2005 consisted of 33 individuals. Its vulnerability to hurricane storm surges and the current population size/trend would remain the same as the current condition under Scenario I.

Under Scenario II, conservation easements would be in place, feral ungulate, pests, and invasive plant species are managed, and the population is monitored. Although the population is expected to grow under this scenario, its resiliency is expected to remain low due to a lack of knowledge on how to create a self-perpetuating population. Under Scenario III, no conservation easements are awarded, feral ungulates and pests are present causing dramatic population declines, limiting natural recruitment and potentially leading to extirpation. Under these circumstances this marron bacora population is predicted to be extirpated in 10 to 25 years.

The Friis Bay population is located at an elevation protected from storm surge. Although we have no recent information from this population, we believe SLR and hurricanes will not likely lead to extirpation of marron bacora at this location because it is found at a high elevation. Also, based on the previously known number of individuals, we neither believe that a flash flood would extirpate this population. Scenarios are summarized for Friis Bay in Table 6 Appendix A.

6.5.5 Base Hill (Current Resilience – Unknown)

The most recent assessment of this population was in 2005. With no subsequent visits, no current data regarding its status or threats are available. Considering a single individual was observed at this site in 2005, that the species is self-incompatible, and the prevailing lack of natural recruitment across the known populations, the likelihood this population still persists is very low. Given the overall declining status of the species, and assuming that threats to this location are consistent with all other population locations, we predicted scenario outcomes. Under Scenario I, the Base Hill population is expected to be extirpated in 10 years (given there is no current data available, this population may already be extirpated). The vulnerability to 62

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hurricane storm surges and the current population size/trend would remain the same as the current condition. Under Scenario II, feral ungulate and pest management is underway, population monitoring is ongoing, and invasive plants are managed. Hence, the population is expected to grow due to population enhancements, but its resiliency will remain very low because of the low rate of survival for introduced individuals, the lack of these introduced individuals to reach sexual maturity in the wild and the lack of knowledge on the optimum ratio of male to female individuals necessary for a self-perpetuating population. Under Scenario III, feral ungulates, invasive plants and pests are present causing dramatic population declines, limiting natural recruitment and potentially leading to extirpation; predicted population is extirpated in 10 years or less.

Available information indicates that the marron bacora Base Hill population is located at an elevation protected from storm surge. Although we have no recent information from this population, we believe SLR will not likely lead to extirpation of this location because it is found at a high elevation. However, given only one individual was known from this location, we believe a major hurricane could extirpate this locality. Scenarios are summarized for Base Hill in Table 7, Appendix A.

6.5.6 Reef Bay Trail (Current Resilience – Low)

Under Scenario I, the Reef Bay Trail population is predicted to be extirpated in 10 to 25 years. This population was discovered in 2017 and consists of 7 individuals located on protected NPS public lands. The vulnerability to hurricane storm surges and the current population size/trend would remain the same as the current condition. Under Scenario II, feral ungulate and pest management are underway and population monitoring ongoing, the population grows due to population enhancements but its resiliency will remain low due to the low rate of survival for introduced individuals, the lack of introduced individuals to reach sexual maturity in the wild (flower production) and the lack of knowledge of the optimum ratio of male to female individuals necessary for a self-perpetuating population. Under the Scenario III, feral ungulates and pests are present causing dramatic population declines, limiting natural recruitment and potentially leading to extirpation; predicted population is extirpated in 10 years.

The Reef Bay Trail population is located at an elevation protected from storm surge. Therefore, we believe SLR is not a threat to this population. Also, although the population is found on steep terrain, we have no information about flash floods or landslides caused by recent atmospheric events. Scenarios are summarized for Reef Bay Trail in Table 8, Appendix A.

6.5.7 Brown Bay Ridge (Current Resilience – Low) Under the Scenario I, the Reef Bay Trail population is expected to be extirpated in 10 years. This population was discovered in 2017 and consists of 1 individual located on protected NPS public lands. The vulnerability to hurricane storm surges and the current population size/trend remain the same as the current condition. Under Scenario II, feral ungulate and pest 63

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management are underway and population monitoring ongoing, the population consists of a single individuals and may expand due to population enhancements, however, its resiliency will remain very low because of the low rate of survival of introduced individuals, the lack of introduced individuals to reach sexual maturity and the lack of knowledge on the optimum ratio of male to female individuals necessary for a self-perpetuating populations , so it is still expected to be extirpated in 10 years. Under Scenario III, feral ungulates and pests are present causing dramatic population declines’, limiting natural recruitment and potentially leading to extirpation; predicted populations is extirpated in 10 years.

Available information indicates that the marron bacora Brown Bay Ridge is located at an elevation protected from storm surge. However, given only one individual was known from this location, we believe a major hurricane could extirpate this locality. Scenarios are summarized for Brown Bay Ridge in Table 9, Appendix A.

6.5.8 Sabbat Point (Extirpated)

This population is extirpated and originally consisted of a single wild individual. A table with metrics for Sabbat Point is summarized in Table 10, Appendix A.

6.5.9 Reef Bay Valley (Extirpated)

This population is extirpated and was originally described as consisting of 6 wild individuals with 60 introduced in 2005. A table with metrics for Reef Bay Valley is summarized in Table 11, Appendix A.

6.5.10 Europa Ridge (Extirpated)

This population is extirpated and originally consisted of a single individual with 60 introduced in 2005. A table with metrics for Europa Ridge is summarized in Table 12, Appendix A.

6.5.11 Sabbath Hill (Tortola, BVI) This population was first described in the British Virgin Islands in November of 2018. There are no previous data on this population, thus, the status of threats to this population are unknown. Given the declining overall status of the species, and assuming that threats to this location are consistent with all other population locations, we predicted scenario outcomes. Under Scenario I, the Sabbath Hill population is expected to remain low in the near future but to be extirpated in 50 years. The population is located on private property. The vulnerability to hurricane storm surges and the current population size/trend would remain the same as the current condition. Under Scenario II, conservation easements are in place, with feral ungulate and pest management underway and population monitoring ongoing, the population grows due to population enhancements, but resiliency remains low due to the low survival rate of introduced individuals, the lack of introduced individuals to reach sexual maturity in the wild and the lack of 64

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knowledge on the optimum ratio of male to female individuals necessary for a self-perpetuating population. Under Scenario III, no conservation easements are awarded; feral ungulates and pests are present causing dramatic population declines, limiting natural recruitment and potentially leading to extirpation; predicted populations are extirpated in 25 years.

Available information indicates that the marron bacora is located at an elevation protected from storm surge. Scenarios are summarized for Sabbath Hill in Table 13, Appendix A.

6.5.12 Future Propagation and Reintroductions

New research should be conducted in the areas of species phenology, genetics and habitat requirements in order to support future reintroductions efforts. New sites should be selected based on habitat factors that favor self-sustaining populations of marron bacora, along with persistence and population growth, including habitat availability, connectivity, and protection, absence of feral ungulates and pests, and safety from rising sea levels. Assuming that sites are selected well, and reintroduced populations are monitored to identify threats that may affect the planted individuals, all new reintroduced populations are expected to attain high resilience under these scenarios. Under Scenario I, 1 new population is introduced (currently funded), under Scenario II, new populations are introduced at a rate of 1 (representative of the genetically unique currently known population) per decade, and under Scenario III, no new populations are reintroduced. Ideally, sites for new reintroductions will be selected so that the elevation and topography of the site protects future marron bacora populations so they are at minimal risk from rising sea levels, hurricanes and drought conditions.

6.6 Future Resilience Summary

The future resilience for marron bacora is summarized in the table below (Table 14). After evaluating the future condition of the species using the scenarios presented, all populations on St. John are predicted to become low to extirpated by 2043. Even with conservation efforts to prevent extirpation, they are not expected to become highly resilient because of the number of the threats facing them and the lack of knowledge on the life history of the species. The population with the highest current resilience, Nanny Point, contains 62% (n = 201) of the known individuals of the species and has been declining since 2005 with low natural recruitment. The status of the second largest population is still unknown, being discovered in November of 2018. Two populations (i.e., John’s Folly and Brown Bay Trail) contain only 18 individuals (unknown ratio of male to female), and population on Reef Bay Trail contains only 7 individuals. As for the marron bacora population in Tortola, BVI, it was described as consisting of 46-48 individuals. All these three populations are thought to be declining and predicted only under Scenario II of having a low to moderate future resiliency. Two populations (i.e., Friis Bay and Base Hill) have not been assessed since 2005. Given the declining state of all of the other populations, Base Hill (single individual) is predicted to be extirpated under all scenarios, and Friis Bay (33 individuals on private property) predicted to be extirpated under Scenarios I and 65

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III. Resiliency only remains low under Scenario II for these two populations. Given the threats these populations are facing, the most effective way to increase their overall future resilience range-wide is to gain a better understanding of the life history of the species and to apply lessons learned to an ongoing propagation program for the reintroduction of new populations in quality protected habitat, with the optimum ratio of male to female individuals and with ongoing monitoring. In fact, conservation of existing populations of marron bacora and their habitat on NPS land, via population augmentation, and habitat restoration is still important to contribute to redundancy and representation for the species.

6.7 Future Redundancy

Redundancy of populations with low resilience under Scenario I is predicted to diminish with populations made up of low numbers of individuals becoming extirpated. The only population that remains under the Scenario I is Nanny Point which currently contains the highest number of individuals of the species (Table 14, Appendix A). Under Scenario III, all populations become extirpated, resulting in extinction of the species. Scenario II is the only scenario that improves redundancy with the introduction of three new populations that have become self-sustaining and the augmentation of the existing populations for a total of 9-10 populations.

6.8 Future Representation

Currently, marron bacora is known from eight locations with populations reported at five locations and single individuals occurring at two of the locations. Future representation for the species depends on the establishment of additional self-sustaining populations in preferred habitat. Genetic diversity within the species remains high, however, since most of the individuals are located in a single population at Nanny Point, ex situ conservation of this population is essential (Stanford et al. 2013, p. 178).

Although Scenario I includes one reintroduction, with all other locations besides Nanny Point becoming extirpated, Representation remains low. Under Scenario II, Representation improves with the three additional reintroductions and the augmentation of existing populations, which would bring up the total populations to approximately 9. In order to improve redundancy within representative units, other factors such as geographic proximity to other locations, sex ratios in populations, feral ungulate and pest management and presence of pollinators should be considered for site selections. Isolated populations with no gene flow between them would be expected to diverge genetically over time, and could at some point be considered distinct representative types. Moreover, locations with single individuals would be expected to become extirpated. Due to small population sizes, much of this genetic divergence is likely to occur in the form of non-adaptive loss of alleles from genetic drift.

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Appendix A. Population Tables

Table A-1. Current and future resilience scores under 3 future scenarios for Nanny Point. Nanny Point Habitat Metrics Population Totals Metric

(201 indv.) Habitat Feral Insect Population Resilience Ungulates Depredation Size/Trend

Scenario Protected, Both High, High, Description Summary or Development Moderate or Moderate or Risk Absent Absent

Current Condition Both (0) High (-1) High (-1) Declining (-1) Low (-3)

10 Year Both (0) High (-1) High (-1) Declining (-1) Low/ Scenario I Extirpated (-3)

25 Year Both (0) High (-1) High (-1) Declining Low/Extirpated Scenario I (-1) (-3)

50 Year Both (0) High (-1) High (-1) Declining Extirpated Scenario I (-1) (-3)

10 Year Both (0) High (-1) High (-1) Stable (0) Low (-2) Scenario II

25 Year Protected (+1) Moderate(0) Moderate (0) Stable (0) Low Scenario II (+1)

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Moderate (+3) Scenario II

10 Year Development High (-1) High (-1) Declining Low/Extirpated (- Scenario III (-1) (-1) 2)

25 Year Development High (-1) High (-1) Declining Extirpated (-4) Scenario III (-1) (-1)

50 Year Development High (-1) High (-1) Declining Extirpated (-4) Scenario III (-1) (-1)

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Table A-2. Current and future resilience scores under 3 future scenarios for John’s Folly. John’s Folly Habitat Metrics Population Metric

(18 indv.) Habitat Feral Insect Population Resilience Ungulates Depredation Size/Trend

Scenario Protected, Both or Present, Present, Description Summary Development Risk Moderate or Moderate or Absent Absent

Current Both (0) High (-1) Unknown (0) Declining Low (-2) Condition (-1)

10 Year Both (0) High (-1) High (-1) Declining (-1) Low/ Scenario I Extirpated (-3)

25 Year Both (0) High (-1) High (-1) Declining Low/ Scenario I (-1) Extirpated (-3)

50 Year Both (0) High (-1) High (-1) Declining Extirpated Scenario I (-1) (-3)

10 Year Both (0) High (-1) High (-1) Stable (0) Low Scenario II (-2)

25 Year Protected (+1) Moderate (0) Moderate (0) Stable (0) Low (+1) Scenario II

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Low (+4) Scenario II

10 Year Area Development High (-1) High (-1) Declining Low/ Scenario III (-1) (-1) Extirpated (-4)

25 Year Area Development High (-1) High (-1) Declining Extirpated Scenario III (-1) (-1) (-4)

50 Year Area Development High (-1) High (-1) Declining Extirpated Scenario III (-1) (-1) (-4)

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Table A-3. Current and future resilience scores under 3 future scenarios for Brown Bay Trail. Brown Bay Habitat Metrics Population Trail Metric

(18 indv.)[YJ1] Habitat Feral Insect Population Resilience Ungulates Depredation Size/Trend

Scenario Protected, Both Present, Present, Moderate Description Summary or Development Moderate or or Absent Risk Absent

Current Protected (+1) High (-1) High (-1) Declining Low (-3) Condition (-1)

10 Year Protected (+1) High (-1) High (-1) Declining Low/ Scenario I Extirpated (-3)

25 Year Protected (+1) High (-1) High (-1) Declining Extirpated (-3) Scenario I (-1)

50 Year Protected (+1) High (-1) High (-1) Declining Extirpated (-3) Scenario I (-1)

10 Year Protected (+1) High (-1) High (-1) Stable (0) Low (-1) Scenario II

25 Year Protected (+1) Absent (+1) Moderate (0) Stable (0) Low (+2) Scenario II

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Low (-3) Scenario II

10 Year Protected (+1) High (-1) High (-1) Declining Low/ Scenario III (-1) Extirpated (-2)

25 Year Protected (+1) High (-1) High (-1) Declining Extirpated (-2) Scenario III (-1)

50 Year Protected (+1) High (-1) High (-1) Declining Extirpated (-2) Scenario III (-1)

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Table A-4. Current and future resilience scores under 3 future scenarios for Friis Bay. Friis Bay Habitat Metrics Population Totals Metric

(33 indv. In Habitat Feral Insect Population Resilience 2005) Ungulates Depredation Size/Trend

Scenario Protected, Both or High, High, Description Summary Development Moderate Moderate Risk or Absent or Absent

Current Unprotected (-1) High (-1) Unknown (0) Status Unknown/ Unknown/ Condition Declining (0) Low (-2)

10 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Low/ Scenario I Extirpated (-4)

25 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Low/ Scenario I Extirpated (-4)

50 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario I

10 Year Unprotected (-1) High (-1) High (-1) Stable (0) Low/ Scenario II Extirpated (-3)

25 Year Protected (+1) Moderate (0) Moderate (0) Stable (0) Low (+1) Scenario II

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Low (+3) Scenario II

10 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario III

25 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario III

50 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario III

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Table A-5. Current and future resilience scores under 3 future scenarios for Base Hill. Base Hill Habitat Metrics Population Totals Metric

(1 indv.)[YJ1] Habitat Feral Insect Population Resilience Ungulates Depredation Size/Trend

Scenario Protected, Both High, High, Moderate Description Summary or Development Moderate or or Absent Risk Absent

Current Protected (+1) High (-1) Unknown Single Unknown Condition Individual/ Unknown

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated Scenario I (-2)

25 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario I

50 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario I

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Low/ Scenario II Extirpated (-2)

25 Year Protected (+1) Moderate (0) Moderate (0) Stable (0) Low/ Scenario II Extirpated (+1)

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Low (+3) Scenario II

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-3) Scenario III

25 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-3) Scenario III

50 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-3) Scenario III

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Table A-6. Current and future resilience scores under 3 future scenarios for Reef Bay Trail. Reef Bay Trail Habitat Metrics Population Totals Metric

(7 indv.)[YJ1] Habitat Feral Insect Population Resilience Ungulates Depredation Size/Trend

Scenario Protected, Both High, High, Description Summary or Development Moderate Moderate Risk or Absent or Absent

Current Protected (+1) High (-1) Unknown (0) 7 Individuals/ Low (-1) Condition Declining (-1)

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Low/ Scenario I Extirpated (-2)

25 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario I

50 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario I

10 Year Protected (+1) High (-1) High (-1) Stable (0) Low (-1) Scenario II

25 Year Protected (+1) Moderate (0) Moderate (0) Stable (0) Low (+1) Scenario II

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Low (+3) Scenario II

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario III

25 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario III

50 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario III

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Table A-7. Current and future resilience scores under 3 future scenarios for Brown Bay Ridge. Brown Bay Population Ridge Habitat Metrics Metric Totals

(1 indv.) Insect Population Habitat Feral Ungulates Depredation Size/Trend Resilience

Scenario Protected, High, Moderate High, Description Summary Both or or Absent Moderate Development or Absent Risk

Current Protected (+1) High (-1) Absent (+1) Single Low (0) Condition Individual/ Declining (-1)

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated Scenario I (-2)

25 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario I

50 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario I

10 Year Protected (+1) High (-1) High (-1) Stable (0) Low/ Scenario II Extirpated (-1)

25 Year Protected (+1) Moderate (0) Moderate (0) Stable (0) Low/ Scenario II Extirpated (+1)

50 Year Protected (+1) Absent (+1) Moderate (0) Stable (0) Low/ Scenario II Extirpated (+2)

10 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario III

25 Year Protected (+1) High (-1) High (-1) Declining (-1) Extirpated (-2) Scenario III

50 Year Protected High (-1) High (-1) Declining (-1) Extirpated (- Scenario III (+1) 2)

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Solanum conocarpum Species Status Assessment 2019

Table A-8. Current and future resilience scores under 3 future scenarios for Sabbat Point. Sabbat Point Habitat Metrics Population Totals Metric

(0 indv.) Habitat Feral Insect Population Resilience Ungulates Depredation Size/Trend

Scenario Protected, Both or High, High, Description Summary Development Risk Moderate Moderate or Absent or Absent

Current Unprotected (-1) High(-1) Unknown Extirpated Extirpated Condition

Scenario I

Scenario II

Scenario III

Table A-9. Current and future resilience scores under 3 future scenarios for Reef Bay Valley. Reef Bay Valley Population Habitat Metrics Metric Totals

(0 indv.) Feral Insect Population Habitat Ungulates Depredation Size/Trend Resilience

Scenario Protected, Both High, High, Moderate Description Summary or Development Moderate or Absent Risk or Absent

Current Protected (+1) High (-1) Unknown Extirpated Extirpated Condition

Scenario I

Scenario II

Scenario III

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Solanum conocarpum Species Status Assessment 2019

Table A-10. Current and future resilience scores under 3 future scenarios for Europa Ridge.

Europa Ridge Population Habitat Metrics Metric Totals

(61 indv.) Feral Insect Population Habitat Ungulates Depredation Size/Trend Resilience

Scenario Protected, Both High, High, Description Summary or Development Moderate Moderate Risk or Absent or Absent

Current Protected (+1) Present (-1) Unknown Extirpated Extirpated Condition

Scenario I

Scenario II

Scenario III

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Solanum conocarpum Species Status Assessment 2019

Table A-11. Current and future resilience scores under 3 future scenarios for Sabbath Hill. Sabbath Hill Population Habitat Metrics Metric Totals

(46 indv.) Feral Insect Population Habitat Ungulates Depredation Size/Trend Resilience

Scenario Protected, Both or High, High, Description Summary Development Moderate Moderate Risk or Absent or Absent

Current Unprotected (-1) High (-1) Unknown (0) 46 Individuals/ Low (-2) Condition Unknown Status/ Declining (0)

10 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Low (-4) Scenario I

25 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Low/ Scenario I Extirpated (-4)

50 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario I

10 Year Unprotected (-1) High (-1) High (-1) Stable (0) Low (-3) Scenario II

25 Year Protected (+1) Moderate (0) Moderate (0) Stable (0) Low (+1) Scenario II

50 Year Protected (+1) Absent (+1) Moderate (0) Growth (+1) Low (+3) Scenario II

10 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Low/ Scenario III Extirpated (-4)

25 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario III

50 Year Unprotected (-1) High (-1) High (-1) Declining (-1) Extirpated (-4) Scenario III

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Solanum conocarpum Species Status Assessment 2019

Table A-12. Summary table of future resilience for marron bacora populations in 2043 under 4 scenarios. Values reflecting good conditions for marron bacora are shaded green (darker green for better conditions), while values reflecting unfavorable conditions are shaded red (darker red for worse conditions), and moderate values are shaded orange. 25 25 Year 10 Year 25 Year 50 Year 10 50 Current 10 Year 50Yr Year Population Scenario I Scenario Scenario Scenario Year Year Status Scenario I Scenario I Pess. II II II Pess. Pess.

L L/X L/X X L L/M M L/X X X Nanny Point (-3) (-3) (-3) (-3) (-2) (+1) (+3) (-2) (-4) (-4) L L/X X X L L L L/X X X (-2) John’s Folly (-3) (-3) (-3) (-2) (+1) (+4) (-4) (-4) (-4)

L L L/X X X L L L/X X X Brown Bay (-3) (+2) (-3) (-3) (-3) (-1) (+3) (-2) (-2) (-2) Trail Unk X X L/X L L L/X X X (-2) L/X Friis Bay (-4) (-4) (-3) (+1) (+3) (-4) (-4) (-4) (-4) Unk X X X L/X L/X L X X X Base Hill (0) (-2) (-2) (-2) (-2) (+1) (+3) (-3) (-3) (-3)

Reef Bay L L/X X X L L L X X X Trail (-1) (-2) (-2) (-2) (-1) (+1) (+3) (-2) (-2) (-2) Brown Bay L X X X LX LX LX X X X Ridge (0) (-2) (-2) (-2) (-1) (+1) (-2) (-2) (-2) (-2)

Sabbat Point X X X X X X X X X X Reef Bay X X X X X X X X X X Valley

Europa Ridge X X X X X X X X X X

L L L/X X L L L L/X X X (-2) Sabbath Hill (-4) (-4) (-4) (-3) (+1) (+3) (-4) (-4) (-4)

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Solanum conocarpum Species Status Assessment 2019

New None None 1 2 None populations Species Extirpated: Extirpated: Extirpated: Extirpated: Extirpated: Extirpated: Extirpated: Extirpated: Extirpated: Extirpated: Summary 11 5 9 11 3 3 3 6 11 11 Low/Ex.: 0 Low/Ex 5 Low/Ex.: 2 Low/Ex.: 0 Low/Ex.: 2 Low/Ex.: 2 Low/Ex.: 1 Low/Ex.: 5 Low/Ex.: 0 Low/Ex.: 0 (Overall) Low: 0 Low: 1 Low: 0 Low: 0 Low: 5 Low: 6 Low: 6 Low: 0 Low: 0 Low: 0 Moderate: 0 Moderate: 0 Moderate: 0 Moderate: 0 Moderate: 0 Moderate: 0 Moderate: 1 Moderate: 0 Moderate: 0 Moderate: 0 High: 0 High: 0 High: 0 High: 0 High: 0 High: 0 High: 0 High: 0 High: 0 High: 0 Unknown: 2

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