Botanical Journal of the Linnean Society

IUCN Red List assessment of the endemic flora: towards a Global Strategy for Conservation in Macaronesia

Journal: Botanical Journal of the Linnean Society

Manuscript ID: BOTJLS-May-2015-2139-OM Manuscript Type: OriginalPDF Manuscript Proof Date Submitted by the Author: 22-May-2015

Complete List of Authors: Romeiras, Maria; Faculty of Sciences, University of Lisbon, Biosystems and Integrative Sciences Institute (BioISI); Tropical Research Institute (IICT), Tropical Botanical Garden (JBT) Catarino, Sílvia; Tropical Research Institute (IICT), Tropical Botanical Garden Gomes, Isildo; Instituto Nacional de Investigação e Desenvolvimento Agrário de Cabo Verde (INIDA), Fernandes, Claudia; Ministerio do Ambiente, Habitação e Ordenamento do Território (MAHOT), Costa, Jose; ISA, University of Lisbon, Landscape Architecture, Biodiversity and Conservation Group (LEAF) Caujapé Castells, Juli; Jardín Botánico Canario , Dept. of Molecular Biodiversity and DNA Bank Duarte, Maria; Tropical Research Institute (IICT), Tropical Botanical Garden; University of Porto, Centre in Biodiversity and Genetic Resources (CIBIO/InBIO)

endemic < Conservation, island biology < Phytogeography, conservation < Conservation, IUCN Red List Categories < Conservation, Keywords: plant conservation < Conservation, dispersal < Ecology, geographical distribution < Phytogeography, systematic < Systematics, threatened plants < Conservation, chorology < Phytogeography

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ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 1 IUCN Red List assessment of the Cape Verde endemic flora: towards a 4 5 2 Global Strategy for Plant Conservation in Macaronesia 6 7 8 3 9 10 11 4 Running Title: Red List assessment of the Cape Verde endemic flora 12 13 5 14 15 6 16 1,2 2 3 4 17 7 Maria M. Romeiras , Silvia Catarino , Isildo Gomes , Claudia Fernandes , Jose C. 18 5 6 2,7 8 Costa , Juli Caujapé-Castells , Maria Cristina Duarte 19 20 9 21 1 22 10 Biosystems and Integrative Sciences Institute (BioISI), Faculty of Sciences, University 23 11 of Lisbon, Campo Grande,PDF 1749-016 Lisbon, Proof Portugal 24 25 12 2 Tropical Research Institute (IICT), Trav. Conde da Ribeira, 1300-142 Lisbon, Portugal 26 3 27 13 Instituto Nacional de Investigação e Desenvolvimento Agrário de Cabo Verde 28 14 (INIDA), São Jorge dos Órgãos. Santiago. Cabo Verde 29 30 15 4 Ministerio do Ambiente, Habitação e Ordenamento do Território (MAHOT), Cidade 31 32 16 da Praia, Santiago. Cabo Verde 33 5 17 Landscape Architecture, Biodiversity and Conservation Group (LEAF-ISA), 34 35 18 University of Lisbon, Tapada da Ajuda, 1349-017 Lisbon, Portugal 36 6 37 19 Jardín Botánico Canario "Viera y Clavijo"-Unidad Asociada CSIC (Cabildo de Gran 38 20 Canaria), Camino al palmeral 15, Tafira Alta 35017 Las Palmas de Gran Canaria, Spain 39 40 21 7 Centre in Biodiversity and Genetic Resources (CIBIO/InBIO), University of Porto, 41 42 22 Campus Agrário de Vairão, 4485-661 Portugal 43 23 44 45 24 46 47 25 Corresponding author details: 48 49 26 Name: Maria Manuel Romeiras 50 51 27 Mailing address: Biosystems and Integrative Sciences Institute (BioISI), Faculty of 52 28 Sciences, University of Lisbon, Campo Grande, 1749-016 Lisbon, Portugal 53 54 29 Email1: [email protected] 55 56 30 Email2: [email protected] 57 31 58 1 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 2 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 32 ABSTRACT 4 5 6 33 7 8 34 Despite endemism richness on islands underlines the outstanding importance of these 9 10 35 enclaves for global conservation, there is no up-to-date synthesis of the conservation 11 12 36 status of the Macaronesian Islands vascular flora, which belongs to one of the world's 13 14 37 hotspot regions. In this study, we review the conservation status and threats to the 15 16 17 38 endemic vascular flora on the Cape Verde Islands, mostly based on the past two decades 18 19 39 of collecting, published literature and herbarium specimens. The application of IUCN 20 21 40 Red List criteria and categories using the RAMAS software reveals that 78% of Cape 22 23 41 Verde’s endemic plants arePDF threatened (29.3%Proof Critically Endangered, 41.3% 24 25 Endangered, 7.6% Vulnerable). Most of these endemics feature a limited geographic 26 42 27 28 43 range, and half of them have Area of Occupancy and Extent of Occurrence of less than 29 30 44 20 km 2 and 200 km 2, respectively. Our data show that, over the last two decades, Cape 31 32 45 Verdean vascular plants have become more threatened, and their conservation status has 33 34 46 declined, mostly as a consequence of the increase in exotic species, habitat degradation 35 36 37 47 and human disturbance. This paper presents the first comprehensive IUCN Red List 38 39 48 data for the plants that are endemic to the Cape Verde Islands, providing an important 40 41 49 step towards the recognition and conservation of its threatened endemic flora at both the 42 43 50 national and global level. It also fills a glaring knowledge gap, as it represents the first 44 45 thorough assessment of the conservation status of the entire endemic flora of a 46 51 47 48 52 Macaronesian archipelago. 49 50 53 51 52 54 ADDITIONAL KEYWORDS: biodiversity hotspot – conservation – oceanic islands – 53 54 55 RAMAS Red List – threatened species – vascular flora 55 56 57 56 58 2 59 60 Botanical Journal of the Linnean Society Page 3 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 57 INTRODUCTION 4 5 58 The continuing decline of plant diversity is the focus of major concerns for researchers, 6 7 8 59 conservation managers, and policy makers (e.g. Tittensor et al., 2014; Pimm et al., 9 10 60 2014). Initiatives to conserve the world’s most threatened diversity have developed over 11 12 61 the last decades, and the IUCN Red List of Threatened Species (www.iucnredlist.org) is 13 14 62 widely recognized as the most objective and comprehensive approach for evaluating the 15 16 17 63 global conservation status of species, and categorizing them according to their estimated 18 19 64 risk of extinction (e.g. Mace et al. , 2008; Jetz & Freckleton, 2015; Maes et al. , 2015). 20 21 65 These Red Lists use quantitative criteria based on population size, rate of decline, and 22 23 66 area of distribution to assignPDF species to oneProof of seven categories of relative extinction 24 25 risk, ranging from ‘Extinct’ to ‘Least Concern’ (IUCN Standards and Petitions 26 67 27 28 68 Subcommittee, 2014). 29 30 69 Assessing the conservation status of endemic plants inhabiting small islands is a key 31 32 70 challenge because of their restricted geographic distribution and high vulnerability to 33 34 71 threats, mainly due to the loss or alteration of their habitats (Caujapé-Castells et al., 35 36 37 72 2010). But the applicability of the IUCN Red List criteria to the exceptionally high 38 39 73 number of endemic vascular plants on most oceanic archipelagos remains to be fully 40 41 74 assessed. Foremost amongst these is the Macaronesian Region (i.e. Azores, Canary, 42 43 75 Madeira, and Cape Verde Islands), that harbors ca. 900 endemic plant species 44 45 (Bramwell & Caujapé-Castells, 2011). Indeed, the Macaronesian vascular flora is one of 46 76 47 48 77 the richest within the Mediterranean biodiversity hotspot (Myers et al. , 2000). 49 50 78 The Macaronesian Region hosts over a quarter of the plant species listed in Annex II of 51 52 79 the Habitats Directive (Sundseth, 2009), despite representing only 0.2% of the European 53 54 80 Union (EU) territory (except the Cape Verde, a non-EU country). Some of these 55 56 57 81 endemics have already been assessed, either in the context of national red lists (i.e. the 58 3 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 4 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 82 Canary Islands, Bañares et al. , 2004; Moreno-Saiz, 2008; Moreno-Saiz et al. , 2015), or 4 5 83 in the European Red List of Vascular Plants (Bilz et al. , 2011). However, these reviews 6 7 84 only cover some of the endemics from the Macaronesian archipelagos that belong to the 8 9 10 85 EU: the Azores and Madeira (Portugal), and the Canaries (Spain). In spite of the utter 11 12 86 rigor of these assessments, a comprehensive Red List for the Macaronesian Region is 13 14 87 still lacking, which has major implications for the conservation of biodiversity in this 15 16 88 hotspot area. 17 18 89 Cape Verde is the only Macaronesian archipelago located in the tropics. 19 20 21 90 Notwithstanding the scientific value of its biota and the existing conservation concerns, 22 23 91 Cape Verde’s biodiversityPDF remains poorly Proof understood. The Flora of the Cape Verde 24 25 92 Islands (Paiva et al. , 1995-1996; Martins et al., 2002) has been an ongoing project for 26 27 93 twenty years, and it has still not been completed (e.g. major plant families like 28 29 30 94 , Cyperaceae, Fabaceae, Malvaceae, and Poaceae, still lack a comprehensive 31 32 95 treatment). Similarly, the preliminary Red List for the archipelago’s flora was published 33 34 96 19 years ago (Leyens & Lobin, 1996), but new endemic taxa have been described or 35 36 97 taxonomically rearranged for the archipelago over the last two decades (e.g. Marrero, 37 38 98 2008; Kilian et al. , 2010; Romeiras et al. , 2011a; Marrero & Almeida, 2012; Knapp & 39 40 41 99 Vorontsova, 2013). Thus, a comprehensive, updated analysis of the available 42 43 100 information regarding population size, distribution, and threats to each endemic species 44 45 101 is urgently required for the conservation of the region’s unique flora. 46 47 102 In this investigation, we assess the conservation status of all vascular endemic plants 48 49 50 103 from the Cape Verde Islands, and we identify the major factors of threat, suggesting 51 52 104 conservation measures to be implemented in this archipelago to further contribute to a 53 54 105 global conservation strategy for the Macaronesian floras. 55 56 106 57 58 4 59 60 Botanical Journal of the Linnean Society Page 5 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 107 MATERIAL AND METHODS 4 5 6 108 7 8 109 STUDY AREA 9 10 110 The Cape Verde archipelago encompasses the southernmost islands of Macaronesia, 11 12 111 and is located 1500 km southwest of the Canary Islands and ca. 570 km west of the 13 14 112 African mainland. This archipelago has ten islands distributed in three groups: Santo 15 16 17 113 Antão, São Vicente, Santa Luzia and São Nicolau in the north; Santiago, Fogo, and 18 19 114 Brava in the south; and Sal, Boavista and Maio (the islands with the lowest altitudes) in 20 21 115 the east (Duarte & Romeiras, 2009). The climate of this archipelago is tropical dry, and 22 23 116 altitudinal gradients, togetherPDF with the effects Proof of north-east trade winds, are key factors 24 25 in shaping species distribution (Duarte et al., 2008). The flora of the Cape 26 117 27 28 118 Verde archipelago is currently thought to comprise about 740 taxa, 92 of them endemics 29 30 119 (Romeiras et al. , 2015). 31 32 120 33 34 121 INVENTORY OF THREATENED PLANTS 35 36 37 122 The data on the vascular plants from Cape Verde’s flora have been gleaned mostly from 38 th 39 123 the collections compiled in Portugal in the second half of the 19 century, which are 40 41 124 housed at the LISC herbarium (Instituto de Investigação Científica Tropical), as well as 42 43 125 from specimens collected by the authors of this paper over the last two decades. 44 45 Additional data were obtained from bibliographic references, namely the Flora of the 46 126 47 48 127 Cape Verde Islands (Paiva et al. , 1995-1996; Martins et al., 2002), and other 49 50 128 publications focusing on endemic plants (e.g. Lobin, 1986; Gomes et al. , 1995; 51 52 129 Brochmann et al. , 1997; Gonçalves, 1999; Duarte et al., 2002; Marrero, 2008; Marrero 53 54 130 & Almeida, 2012; Knapp & Vorontsova, 2013). Data bearing on species ecology and 55 56 57 131 distribution in the islands, as well as altitudes, collector’s names and dates of collection 58 5 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 6 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 132 were included in a database that contains ca. 4700 individual records; whenever 4 5 133 possible, the geographical coordinates of the accessions were also considered (only 6 7 134 specimens collected after 1955 could be georeferenced, due to insufficient location 8 9 10 135 information provided on historical specimen labels). A total of 4583 specimens were 11 12 136 georeferenced using 1:25,000 and 1:100,000 cartographic maps, and the data were 13 14 137 compiled in ArcGIS Arcinfo ver. 10.0 (ESRI, 2011). 15 16 138 17 18 139 RED LIST ASSESSMENTS 19 20 21 140 The conservation status of the Cape Verde endemic flora was evaluated following the 22 23 141 IUCN Red List categories,PDF so that each listedProof species could be classified as Extinct 24 25 142 (EX), threatened [i.e. Critically Endangered (CR); Endangered (EN); or Vulnerable 26 27 143 (VU)], Near Threatened (NT), Least Concern (LC), or within a Data Deficient (DD) 28 29 30 144 category for species that are very poorly known. From the five quantitative criteria (A– 31 32 145 E) which are used to evaluate each taxon (IUCN Standards and Petitions Subcommittee 33 34 146 2014), criterion B (i.e. restricted distribution and decline, fluctuations, and/or 35 36 147 fragmentation), and criterion D (i.e. very small or restricted populations) were the most 37 38 148 commonly used. The use of criteria A, C and E was avoided due to a lack of 39 40 41 149 information concerning population trends, present and future population decline and 42 43 150 probability of extinction, respectively. 44 45 151 We calculated the Area of Occupancy (AOO), Extent of Occurrence (EOO), number of 46 47 152 subpopulations, and number of locations for each of the 4583 records of Cape Verdean 48 49 50 153 endemic taxa evaluated. The parameter EOO was estimated using the minimum convex 51 52 154 polygon method, which determines the area contained within the shortest continuous 53 54 155 imaginary boundary that can be drawn to encompass all the occurrences of a taxon, 55 56 156 while AOO was calculated from summing the number of cells occupied by individuals 57 58 6 59 60 Botanical Journal of the Linnean Society Page 7 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 157 in a grid of 1x1km. For both calculations we used the GeoCAT software (Bachman et 4 5 158 al. , 2011). Since sampling efforts were not applied with the same intensity to all species 6 7 159 and in all places, species may have a greater AOO and EOO than known, causing some 8 9 10 160 uncertainty that may result in distribution underestimates for some taxa. The evaluation 11 12 161 of the conservation status was made using the RAMAS Red List software v.2.0 13 14 162 (Akçakaya et al. , 2001), which was successfully applied to the Cape Verde IUCN 15 16 163 extinction risk assessment of reptiles (Vasconcelos et al. , 2013). RAMAS assigns each 17 18 164 taxon to Red List Categories according to the IUCN Red List Criteria, and explicitly 19 20 21 165 handles data uncertainty (Akçakaya et al. , 2000). Considering the uncertainty that 22 23 166 inevitably arises when manyPDF of the parameters Proof are unknown, this software package is 24 25 167 the most recommended by IUCN authorities and allows us testing different values for 26 27 168 the Risk Tolerance (RT). RT ranges from 0 for risk-averse, precautionary; through 0.5 28 29 30 169 for risk neutral; to 1 for risk prone, evidentiary (Akçakaya et al. , 2000). An evidentiary 31 32 170 attitude (RT=0.6) was applied, given the above-mentioned sources of uncertainty. 33 34 171 Finally, major threats for each taxon were assessed using a standardized list (IUCN 35 36 172 Standards and Petitions Subcommittee, 2014) based on information gathered from 37 38 173 fieldwork and published data. 39 40 41 174 42 43 175 RESULTS 44 45 46 176 47 48 177 STATUS OF THREATENED PLANT SPECIES IN CAPE VERDE 49 50 178 Our results revealed that 78% of the assessed endemics (92 taxa) were listed in threat 51 52 179 categories: 27 (29.3%) as CR, 38 (41.3%) as EN, and 7 (7.6%) as VU (Fig. 1). Eight 53 54 180 taxa (8.7%) were classified as NT, one (1.1%) as LC, and five (5.4%) as DD (Table 1). 55 56 57 181 Additionally, six taxa (6.5%) belonging to the Lotus (Fabaceae) were Not 58 7 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 8 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 182 Evaluated (NE), given the wide morphological diversity and considerable taxonomic 4 5 183 uncertainties, that hinder the assignment of collected samples to a particular species. 6 7 184 Apart from the 92 endemics, two species were classified as Extinct (EX) 8 9 10 185 (Stachytarpheta fallax and Habenaria petromedusa ), as they are known only from the 11 12 186 type specimens collected in the 18th century by J.S. Feijó. 13 14 187 Criterion B (geographical range) was the most frequently used for categorization of 15 16 188 threat (73.3% taxa). Most of the endemics display a limited geographic range, with half 17 18 189 of them having areas of occupancy and extents of occurrence of less than 20 and 200 19 20 2 21 190 km , respectively. Approximately 27% of the taxa assessed have simultaneously an 22 2 23 191 AOO and an EOO equal toPDF or less than 10Proof and 100 km , respectively; within these, 18 24 25 192 are single-island endemics (SIEs). The largest AOO and/or EOO values (Table 1) are 26 27 193 displayed by Euphorbia tuckeyana , Cynanchum daltonii , Paronychia illecebroides , and 28 29 30 194 Forsskaolea procridifolia ; these species occur in seven or more islands, and most of 31 32 195 them were evaluated as NT, save for Cynanchum daltonii (LC). The smallest AOO and 33 34 196 EOO values were both for Teline stenopetala subsp. santoantaoi (CR). In CR species, 35 36 197 AOO values ranged mainly from 3 to 10.5 km 2 and in VU species from 27 to 56.5 km 2 37 38 198 whereas EOO values were considerably higher, between 3 to 46.8 km 2 in CR species 39 40 2 41 199 and between 328.3 to 575.8 km in VU species (Fig. 2). 42 43 200 The distribution of threatened species (Fig. 1) shows that the northern and southern 44 45 201 mountain island groups present the highest percentage values (ranging from 71% in São 46 47 202 Vicente to 79.2% in Brava), because both island groups harbor most of the SIEs, which 48 49 50 203 present restricted AOO and EOO, thus potentially qualifying in the highest threat 51 52 204 categories (i.e. CR) under IUCN criterion B (see Table 1). Most of the species 53 54 205 distributed in the Eastern Islands have a large EOO, because they are by and large 55 56 206 widespread in the archipelago. Nonetheless, these results correspond to global 57 58 8 59 60 Botanical Journal of the Linnean Society Page 9 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 207 assessments in the archipelago, and the category of particular species for each island 4 5 208 may be different, depending on the number of populations and the number and intensity 6 7 209 of threats that may affect their survival. 8 9 10 210 11 12 211 RED LIST CHANGES IN THE LAST TWO DECADES 13 14 212 A comparison between the current conservation assessment of the endemic taxa and the 15 16 213 one carried out in 1996 (Table 1, Fig. 3) shows that, overall, the Cape Verdean plants 17 18 214 are more threatened and their conservation status has declined over the last two decades. 19 20 21 215 Although about one quarter of the endemic vascular flora was not evaluated by Leyens 22 23 216 & Lobin (1996), it is notedPDF that the three Prooftaxa previous classified as CR ( 24 25 217 schlechtendalii, Carex antoniensis and C. paniculata subsp. hansenii ) still remain in 26 27 218 this threat category. Recent field surveys have revealed that C. schlechtendalii is 28 29 30 219 restricted to a small population in São Nicolau, and the two Carex are only found in 31 32 220 very small populations in Santo Antão (Ribeira do Paul). Moreover, the categories 33 34 221 ‘Undetermined’ and ‘Rare’ applied by Leyens & Lobin (1996) are no longer considered 35 36 222 by the IUCN (for further details see information in Fig. 3), and almost all the taxa under 37 38 223 these categories were now classified as CR or as EN due to the small fragmented and 39 40 41 224 restricted populations. On the other hand, of the 15 taxa assessed as VU in 1996, only 42 43 225 Euphorbia tuckeyana was downlisted from VU to NT due to its widespread distribution 44 45 226 in the archipelago, and to the fact that some populations with a significant number of 46 47 227 individuals were recently found in Santo Antão (namely in Tope Coroa). 48 49 50 228 Despite our results point to an increase in extinction risk during the last two decades, 51 52 229 recent field surveys allowed us to rediscover several species reported as extinct by 53 54 230 Leyens & Lobin (1996). A first example is glauca , only recorded in Boavista 55 56 231 in 1851 (leg. Schmidt; type collection), and considered extinct by Brochmann et al., 57 58 9 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 10 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 232 (1997) until it was collected in 2013 by one of us (MCD). Also during recent fieldwork, 4 5 233 scattered trees of Dracaena draco subsp. caboverdeana were found in Santiago and 6 7 234 Brava, thus supporting the contention by Marrero & Almeida (2012) that currently this 8 9 10 235 species only has natural populations in Santo Antão, São Nicolau and Fogo, growing 11 12 236 sub-spontaneously also in Santiago and Brava. Finally, Marrero & Pérez (2013) 13 14 237 reported the re-discovery in Brava of the native species Eulophia guineensis Lindl. 15 16 238 (Orchidaceae). 17 18 239 19 20 21 240 MAIN THREATS 22 23 241 Following the IUCN ThreatsPDF Classification Proof Scheme (Version 3.2), the most pervasive 24 25 242 threats reported in Cape Verde were, by decreasing order of importance (Fig. 4): i) 26 27 243 gathering plants for intentional use; ii) invasive alien species; and iii) nomadic grazing. 28 29 30 244 Natural disasters, specifically recent volcanic events (with the most recent eruption 31 32 245 occurring in 2014) have an impact on species that occur above 1600 m a.s.l. on Fogo 33 34 246 Island, whereas tourism and recreation areas are especially significant for the taxa 35 36 247 distributed in lowland coastal areas. Most of the threats were recorded between 400 and 37 38 248 1200 m a.s.l. and were especially associated with strong anthropic disturbances, namely 39 40 41 249 nomadic grazing, gathering endemic plants for intentional use, and invasive species 42 43 250 (Fig. 4). 44 45 251 46 47 48 252 DISCUSSION 49 50 253 51 52 254 Our study presents the most comprehensive Red Data List for the endemic vascular 53 54 255 plants of the Cape Verde Islands, providing an important step towards the recognition 55 56 57 256 and conservation of the threatened flora of this archipelago at both national and global 58 10 59 60 Botanical Journal of the Linnean Society Page 11 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 257 level. Internationally, the conservation of endemic Cape Verde flora is of great 4 5 258 importance; according to a recent review by Caujapé-Castells et al. (2010), its flora is 6 7 259 one of the most threatened in the Macaronesian archipelagos. Our results, pointing to 8 9 10 260 78% of threatened taxa, unequivocally confirm the uppermost position followed by 11 12 261 Azores, Madeira and the Canary Islands with 63%, 49% and 30% of threatened taxa, 13 14 262 respectively (Caujapé-Castells et al. , 2010). 15 16 263 A high extinction risk was recently documented in the Red List of the endemic 17 18 264 monocotyledons from Morocco (Rankou et al. , 2015), which is a West African country 19 20 21 265 mainly characterized by a semi-arid climate, like the Cape Verde Islands. In both cases 22 23 266 70% of the assessed endemicPDF flora was classifiedProof in high-risk categories (CR or EN). 24 25 267 Therefore, the higher vulnerability of the Cape Verde endemic flora could be mainly 26 27 268 explained by its tropical dry climate and by the increasing aridity that affects the 28 29 30 269 islands, especially at lower altitudes, which could have led to population reductions and 31 32 270 restrictions on distribution ranges of the taxa. 33 34 271 Nevertheless, the high proportion of threatened species revealed in our study might be 35 36 272 also influenced by some drawbacks during the conservation assessments, mostly 37 38 273 because the sampling efforts were not uniformly distributed in all Cape Verde 39 40 41 274 regions/islands, and species may have greater areas than recognized. Thus, the 42 43 275 inadequacies in taxonomic and distributional data, the so-called Linnean and Wallacean 44 45 276 shortfalls (Whittaker et al. , 2005), which are recognized to increase in more remote 46 47 277 areas such as oceanic islands (Ladle & Whittaker, 2011), constitute two of the most 48 49 50 278 pressing problems for the thorough conservation of the Cape Verde flora. To reduce the 51 52 279 referred shortfalls, it is essential to study different biodiversity units, which can range 53 54 280 from genes to landscapes. Therefore, the conservation of intraspecific genetic diversity 55 56 281 at archipelago scale, together with the protection of species and habitats, will be of 57 58 11 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 12 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 282 particular importance to underpin conservation programmes. Recently, studies focusing 4 5 283 on the Macaronesian flora (García-Verdugo et al. , 2015; Patiño et al ., 2015; Romeiras 6 7 284 et al., 2015) revealed that some plant lineages are genetically more diverse than 8 9 10 285 previously recognized. These findings further highlight the need to conserve insular 11 12 286 populations, as many of them are not as “genetically depauperated” as previously 13 14 287 thought (García-Verdugo et al. , 2015). Nevertheless, the concrete application of genetic 15 16 288 data to the design of protected areas at the archipelago scale remains largely unexplored 17 18 289 in insular biodiversity hotspots (Buerki et al. , 2015). 19 20 21 290 The present global assessment of the conservation status of Cape Verdean endemic 22 23 291 vascular plants shows thatPDF more than three-quarters Proof of species are classified in one of 24 25 292 the threat categories (i.e. CR, EN, and VU) under IUCN Red List standards. A further 26 27 293 five species are listed as DD, and it is extremely likely that these endemic taxa are 28 29 30 294 threatened. Nevertheless, and as suggested in these situations (Akçakaya et al. , 2000), 31 32 295 our recommendation is that Cape Verdean DD species should be assigned the same 33 34 296 degree of protection as threatened species until more information is available. Such 35 36 297 scenario would add new species to the threat categories, and have consequences for 37 38 298 conservation prioritization. Additionally, our results illustrate the importance of an 39 40 41 299 accurate prior to a conservation assessment in insular endemic plants, thus 42 43 300 reinforcing Mace’s (2004) opinion that more collaboration is needed among 44 45 301 conservation biologists and high-profile taxonomists. This was particularly relevant 46 47 302 regarding the genus Lotus , which was not evaluated due to the wide morphological 48 49 50 303 diversity that hinders the clear taxonomic delimitation of the species. 51 52 304 The IUCN Red List system has been refined over the 20 years since the first Cape 53 54 305 Verdean plant Red List was published; given the significant differences in assessment 55 56 306 methodologies it seems in order to compare with the previous list. Although our data 57 58 12 59 60 Botanical Journal of the Linnean Society Page 13 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 307 point towards an increase in extinction risk during the last two decades, this result might 4 5 308 be also related to: i) the use of new more accurate data, which generate more realistic 6 7 309 assessments; ii) the application of IUCN available tools (e.g. the RAMAS software) 8 9 10 310 instead of ‘expert opinion’; iii) the inclusion of new analyses on the threats impacting 11 12 311 the Cape Verdean plants; iv) the enlargement of the assessment to all the endemic flora, 13 14 312 with a considerable number of taxa being categorized for the first time and being 15 16 313 assigned to thread categories; and v) changes in the taxonomy of some groups that 17 18 314 produced an upward surge of critically endangered species due to taxonomic splitting. 19 20 21 315 This may have considerable conservation implications since some clades were divided 22 23 316 (e.g. Echium stenosiphon PDFs.l., see Romeiras Proof et al. , 2011a), with the new taxa presenting 24 25 317 more restricted AOO and EOO. 26 27 318 The high percentage of threatened taxa in Cape Verde is alarming and, as in other 28 29 30 319 insular ecosystems (e.g. Caujapé-Castells et al. , 2010; Kueffer et al. , 2010), habitat 31 32 320 degradation, human disturbance (e.g. intentional use for agriculture or traditional uses; 33 34 321 Romeiras et al. , 2011b) and introduction of exotic species since the beginning of islands 35 36 322 colonization (Romeiras et al., 2014) are among the main threats. Furthermore, the recent 37 38 323 volcanic activity on Fogo Island could lead to population extinctions, particularly 39 40 41 324 among the single-island endemics (SIEs: Echium vulcanorum , Erysimum 42 43 325 caboverdeanum and Verbascum cystolithicum ) that mainly occur above 1600 m in Chã 44 45 326 das Caldeiras. 46 47 327 In general, these threats have a great negative impact in the Cape Verde flora: most 48 49 50 328 endemics display a limited geographic range (both in terms of AOO and EOO), thus 51 52 329 being more susceptible to extinction. To prevent taxa from going extinct, several 53 54 330 conservation actions were undertook in the last years by the Cape Verde authorities, 55 56 331 namely establishing a system of Protected Areas (PA) to safeguard the archipelago’s 57 58 13 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 14 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 332 natural heritage (MAAP, 2004). Presently, the National PA network encompasses 47 4 5 333 different regions, totalizing ca. 63,067 ha of land area (see 6 7 334 http://www.areasprotegidas.gov.cv) that include four main protection categories: Nature 8 9 10 335 Reserve, Natural Park, Natural Monument, and Protected Landscape. Among these, the 11 12 336 Natural Parks encompass sensitive areas for biodiversity conservation, where the 13 14 337 majority of the endemic plants occur (e.g. Tope de Coroa – Santo Antão; Monte Gordo 15 16 338 – São Nicolau; Monte Verde – São Vicente; Serra da Malagueta – Santiago; Chã das 17 18 339 Caldeiras - Fogo). These National Parks are found in mountain regions of the northern 19 20 21 340 and southern island groups, where most of the endemics occur as small and isolated 22 23 341 populations, mostly in thePDF northeast-exposed Proof slopes above 400 m. In these mountain 24 25 342 areas, greater floristic affinities are shared with the other Macaronesian archipelagos, 26 27 343 especially with the Canaries and Madeira. Among these affinities we find several 28 29 30 344 endemics that belong to some of the biggest plant radiations in Macaronesia and 31 32 345 worldwide (e.g. Aeonium ; Echium ; Euphorbia ; Micromeria ; Sonchus ; Tolpis ), thus 33 34 346 posing a compelling need to conserve the whole extent of their natural ranges as a key 35 36 347 objective to the informed conservation of the Macaronesian biodiversity. 37 38 348 On the other hand, the eastern islands (i.e. Sal, Boavista, and Maio) are lower in 39 40 41 349 altitude, they undergo long periods of severe drought, and they have poor vegetation 42 43 350 contents, with fewer endemics for which urgent conservation actions are also needed. 44 45 351 However, the implementation of the PA was focused mainly on marine resources or 46 47 352 fauna species, like see birds or turtles (Mauremootoo, 2012). Threats to the endemic 48 49 50 353 plant species are mainly driven by habitat loss and anthropogenic disturbance, in 51 52 354 particular related with touristic infrastructures and urban development. Especially on the 53 54 355 coastal sands and dunes of Sal and Boavista, tourism growth has caused dramatic 55 56 356 habitat changes, with already noticeable negative impacts on the endemic flora (for 57 58 14 59 60 Botanical Journal of the Linnean Society Page 15 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 357 instance, in the surrounding areas of Praia de Santa Maria on Sal Island, where a very 4 5 358 small population of Pulicaria burchardii subsp . longifolia is undergoing fast decline). 6 7 359 The information in this study should be used to provide guidance for future 8 9 10 360 management and conservation efforts to ensure a survival of these threatened species in 11 12 361 these islands. However, because designating new PA will be a complex task (i.e. there 13 14 362 are several competing land-use options and considerable socio-economic costs 15 16 363 associated with PA implementation), a species prioritization procedure is mandatory. 17 18 364 19 20 21 365 FINAL REMARKS 22 23 366 Internationally, the four MacaronesianPDF archipelagosProof ratified through their respective 24 25 367 countries the Convention on Biological Diversity, and are included in the Mediterranean 26 27 368 “Biodiversity Hotspot”, which implies that concerted actions aimed at preserving the 28 29 30 369 profuse biodiversity of the Macaronesian region should be taken. At the regional level, 31 32 370 policy frameworks have been implemented to guide biodiversity conservation in the 33 34 371 Cape Verde Islands (National PAs network), and it is urgent to develop new field 35 36 372 surveys to fill glaring gaps in species distribution data, and to monitor population sizes, 37 38 373 their sensitivity to disturbances, and other threats that may affect them. Recent 39 40 41 374 inventories allowed the rediscovery in some islands of species reported extinct, thus 42 43 375 further underscoring the importance of increased field collection efforts. Intensifying 44 45 376 prospection, especially in the hardly accessible mountain areas, would probably increase 46 47 377 the number of records of both old-described and undercollected species, while refining 48 49 50 378 the knowledge of their distribution ranges. 51 52 379 Conservation projects are time-sensitive, and research funding opportunities in 53 54 380 developing countries like Cape Verde are becoming increasingly restricted, but our 55 56 381 ability to embrace informed, integrative approaches to biodiversity science is always 57 58 15 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 16 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 382 contingent on the availability of scientifically sound data. This Red List provides a first 4 5 383 comprehensive framework for identifying and prioritizing threatened species, thus 6 7 384 constituting a crucial step towards a better strategy to conserve the endemic flora in the 8 9 10 385 southernmost archipelago of Macaronesia. 11 12 386 13 14 387 ACKNOWLEDGMENTS 15 16 17 388 We are grateful to many Cape Verde collaborators for their assistance during fieldwork 18 19 389 [e.g. S. Gomes (Santiago); H. Diniz (Fogo); E. Ramos, A. Fortes; G. Monteiro (S. 20 21 390 Antão); C. Monteiro (S. Vicente); A. Fernandes, H. Santos (Sal); I. Duarte (Boavista; S. 22 23 391 Nicolau)] and to L. CarvalhoPDF and M. Borges Proof (Director General of Environment of Cape 24 25 Verde) for collecting permits. We are also grateful to our colleagues P. Beja, AF. Filipe, 26 392 27 28 393 MF. Magalhães for helpful discussions with the application of RAMAS, and also to J. 29 30 394 Alves, L. Tavares and J. Farminhão. Special thanks go to M. Carine (NHM, London) 31 32 395 for all the fruitful discussions and collaboration during the fieldwork in Cape Verde 33 34 396 with MMR. This study was supported by the Portuguese Foundation for Science and 35 36 37 397 Technology (FCT) and European Social Funds through project PTDC/BIA- 38 39 398 BIC/4113/2012; and by the “The Mohamed bin Zayed Species Conservation Fund”, 40 41 399 grant number MBZ: 12255026 to MMR. The field surveys were conducted under a 42 43 400 protocol established between the IICT/Portugal and the Ministry of Environment 44 45 (MAHOT) of Cape Verde. 46 401 47 48 402 49 50 403 51 52 404 53 54 405 55 56 57 406 58 16 59 60 Botanical Journal of the Linnean Society Page 17 of 31 Botanical Journal of the Linnean Society

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10 556 11 12 557 13 14 15 558 FIGURE LEGENDS 16 17 18 559 Figure 1. Distribution of Cape Verde endemic species by threatened categories 19 20 560 [Critically Endangered (CR); Endangered (EN); Vulnerable (VU)] in the archipelago 21 22 561 (left) and in each island (right). Island abbreviations: Santo Antão (ANT); São Vicente 23 PDF Proof 24 562 (VIC); Santa Luzia (LUZ); São Nicolau (NIC) (Northern Group); Sal (SAL); Boavista 25 26 27 563 (BOA); Maio (MAI) (Eastern Group); Santiago (SAN); Fogo (FOG); Brava (BRA) 28 29 564 (Southern Group). 30 31 565 32 33 566 Figure 2. Species geographic range - AOO (area of occupancy) and EOO (extent of 34 35 36 567 occurrence) - for the 72 endemic species classified under threatened categories 37 38 568 [Critically Endangered (CR); Endangered (EN); Vulnerable (VU)]. 39 40 569 41 42 570 Figure 3. Status change in IUCN Red List of the Cape Verdean endemic flora from 43 44 571 1996 (Leyens & Lobin previous inventory) to 2015 (present assessment). Species 45 46 47 572 classified in 1996 as Undetermined (applied when it was not possible to accurately 48 49 573 classify a species into any of the threatened categories CR, EN or VU) or Rare (species 50 51 574 restricted to isolated populations, and to which there was not enough information to 52 53 575 determine their conservation status, but corresponding most likely to CR, EN or VU) 54 55 56 576 were considered here as Threatened. 57 58 23 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 24 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 577 4 5 578 Figure 4. Incidence along elevation classes of the main threats to the Cape Verdean 6 7 579 endemic plants. Dimension of circles proportional to the number of species affected in 8 9 10 580 the corresponding elevation class. Classification of threats as defined by IUCN. 11 12 581 (http://www.iucnredlist.org/technical-documents/classification-schemes/threats- 13 14 582 classification-scheme) 15 16 583 17 18 19 584 TABLE : 20 21 585 Table 1. Some parameters used for the assessment of conservation status of the Cape 22 586 Verde endemic plant taxa, and Red List categories and criteria. 23 PDF Proof 24 25 587 26 AOO EOO Red List Criteria Family Taxon Islands * 2 2 27 (km ) (km ) 1996** 2015 2015 28 29 Crassulaceae Aeonium gorgoneum J. A. Schmidt 3 38 224,26 LR EN B1ab(ii,iii)+2ab(ii,iii) 30 31 Poaceae Aristida cardosoi Cout. 10(9) 53 848,37 NE NT 32 Asteraceae Artemisia gorgonum Webb 3 37 260,40 VU VU B1ab(ii,iv)+2ab(ii,iv) 33 34 Asparagaceae Asparagus squarrosus J. A. Schmidt 7 50 924,03 LR NT 35 36 Asteraceae daltonii (Webb) Walp. subsp. daltonii 1 11 66,73 EN EN D 37 Asteraceae Asteriscus daltonii subsp. vogelii (Webb) Greuter 7 91 1151,37 LR NT 38 39 Asteraceae Asteriscus smithii (Webb) Walp. 1 4 4,00 EN CR B1ab(iii)+2ab(iii) 40 Brachiaria lata (Schumach.) C. E. Hubb. subsp. 41 Poaceae 4 41 569,44 VU VU B1ab(iii)+2ab(iii) caboverdeana Conert & Ch. Köhler 42 43 Campanulaceae Campanula bravensis (Bolle) A. Chev. 3 44 224,99 LR EN B1ab(ii,iv)+2ab(ii,iv) 44 45 Campanulaceae Campanula jacobaea C. Sm. ex Webb 4 74 514,36 NE VU B1ab(ii)+2ab(ii) 46 Plantaginaceae Campylanthus glaber Benth. subsp. glaber 6 66 1001,20 VU EN B1ab(ii)+2ab(ii) 47 48 Campylanthus glaber Benth. subsp. spathulatus (A. Chev.) Plantaginaceae 1 11 143,13 NE EN B1ab(ii)+2ab(ii); D 49 Brochmann, N. Kilian, Lobin & Rustan 50 51 Cyperaceae Carex antoniensis A. Chev. 1 3 3,00 CR CR B1ab(ii)+2ab(ii); D 52 Cyperaceae Carex paniculata L. subsp. hansenii Lewej. & Lobin 1 3 3,00 CR CR B1ab(ii)+2ab(ii); D 53 Centaurium tenuiflorum (Hoffmanns. & Link) Fritsch 54 Gentianaceae 3 11 207,63 NE CR D 55 subsp. viridense (Bolle) A. Hansen & Sunding 56 57 58 24 59 60 Botanical Journal of the Linnean Society Page 25 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 Asteraceae Conyza feae (Bég.) Wild 6(5) 76 617,39 EN EN B1ab(ii,iv)+2ab(ii,iv) 4 Asteraceae Conyza pannosa Webb 5 22 166,08 EN EN B1ab(ii,iv)+2ab(ii,iv) 5 6 Asteraceae Conyza schlechtendalii Bolle 1 3 3,00 CR CR D 7 8 Asteraceae Conyza varia (Webb) Wild 5 45 258,96 EN EN B1ab(ii,iv)+2ab(ii,iv) 9 Asclepiadaceae Cynanchum daltonii (Decne. ex Webb) Liede & Meve 7 105 1534,62 NE LC None 10 11 Diplotaxis antoniensis Rustan 1 16 187,84 NE VU D1+2 12 13 Brassicaceae Diplotaxis glauca J. A. Schmidt 2 10 103,56 VU CR D 14 Brassicaceae Diplotaxis gorgadensis Rustan subsp. brochmannii Rustan 1 3 3,00 VU CR B1ab(ii)+2ab(ii) 15 16 Brassicaceae Diplotaxis gorgadensis Rustan subsp. gorgadensis 1 12 171,77 NE EN B1ab(ii)+2ab(ii) 17 18 Brassicaceae Diplotaxis gracilis (Webb) O. E. Schulz 1 10 146,12 VU EN B1ab(iii)+2ab(iii) 19 Brassicaceae Diplotaxis hirta (A. Chev.) Rustan & Borgen 1 25 212,58 NE EN B1ab(ii)+2ab(ii) 20 21 Brassicaceae Diplotaxis sundingii Rustan 1 3 3,00 R CR B1ab(ii)+2ab(ii) 22 23 Brassicaceae Diplotaxis varia Rustan PDF Proof2 25 214,53 I EN B1ab(ii)+2ab(ii) 24 Brassicaceae Diplotaxis vogelli (Webb) Cout. 1 6 43,61 I CR B1ab(ii)+2ab(ii) 25 Dracaena draco (L.) L. subsp. c aboverdeana Marrero Rodr. 26 Asparagaceae 6(3) 16 53,00 NE CR B1ab(ii,iv) 27 & R. Almeida 28 Dryopteridaceae Dryopteris gorgonea J.P. Roux 3 3 3,00 NE DD 29 30 Boraginaceae Echium hypertropicum Webb 2 34 222,65 EN EN B1ab(ii)+2ab(ii) 31 Echium stenosiphon Webb subsp. glabrescens (Pett.) 32 Boraginaceae 1 29 105,60 LR EN B1ab(ii)+2ab(ii) 33 Romeiras & Maria C. Duarte 34 Echium stenosiphon Webb subsp. lindbergii (Pett.) Boraginaceae 1 42 284,83 I EN B1ab(ii)+2ab(ii) 35 Bramwell 36 37 Boraginaceae Echium stenosiphon Webb subsp. stenosiphon 1 14 59,48 VU CR B1ab(ii) 38 Boraginaceae Echium vulcanorum A. Chev. 1 21 121,75 EN EN B1ab(ii)+2ab(ii) 39 40 Poaceae Eragrostis conerti Lobin 5 14 71,60 R DD 41 42 Brassicaceae Erysimum caboverdeanum (A. Chev.) Sund. 1 13 50,03 EN CR B1ab(ii)

43 Euphorbia tuckeyana Steud. ex Webb 9(8) 108 1145,26 VU NT 44 45 Zygophyllaceae Fagonia mayana Schlecht. 3 11 91,91 NE DD 46 47 Forsskaolea procridifolia Webb 9 93 1569,67 NE NT 48 Frankenia ericifolia Chr. Sm. ex DC. subsp. caboverdeana Frankeniaceae 3 20 590,48 NE EN B1ab(ii)+2ab(ii) 49 Brochmann, Lobin & Sunding 50 Frankenia ericifolia Chr. Sm. ex DC. subsp. montana 51 Frankeniaceae 1 6 8,19 EN CR B1ab(iii)+2ab(iii) 52 Brochmann, Lobin & Sunding

53 Plantaginaceae Globularia amygdalifolia Webb 5 50 378,28 VU EN B1ab(ii)+2ab(ii) 54 55 Orchidaceae Habenaria petromedusa Webb NE EX 56 57 58 25 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 26 of 31

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 Cistaceae Helianthemum gorgoneum Webb 4 43 490,61 NE EN B1ab(ii,iv)+2ab(ii,iv) 4 Asteraceae Helichrysum nicolai N. Kilian, Galbany & Oberpr. 1 2 2,00 NE CR D 5 6 Kickxia elegans (G. Forst.) D. A. Sutton subsp. Plantaginaceae 4 24 290,46 NE EN B1ab(ii,iv)+2ab(ii,iv) 7 dichondrifolia (Benth.) Rustan & Brochmann 8 Plantaginaceae Kickxia elegans (G. Forst.) D. A. Sutton subsp. elegans 9(8) 56 902,06 NE EN B1ab(iv)+2ab(iv) 9 10 Kickxia elegans (G. Forst.) D. A. Sutton subsp. webbiana Plantaginaceae 1 14 132,63 NE EN B1ab(ii)+2ab(ii) 11 (Sunding) Rustan & Brochmann 12 13 Asteraceae Launaea gorgadensis (Bolle) N. Kilian 3 15 26,29 LR CR B1ab(iii) 14 Asteraceae Launaea picridioides (Webb) Engler 3 56 667,82 LR VU B1ab(ii)+2ab(ii) 15 16 Asteraceae Launaea thalassica N. Kilian, Brochmann & Rustan 1 9 18,36 R CR B1ab(iii)+2ab(iii) 17 18 Lamiaceae Lavandula rotundifolia Benth. 5 95 1060,16 LR NT 19 Plumbaginaceae Limonium braunii (Bolle) A. Chev. 4 24 244,98 NE EN B1ab(ii)+2ab(ii) 20 21 Plumbaginaceae Limonium brunneri (Webb) Kuntze 3 13 59,06 LR CR B1ab(ii,iii,iv) 22 23 Plumbaginaceae Limonium jovibarba (Webb) PDFKuntze Proof2 10 15,58 R CR B1ab(ii); D 24 Plumbaginaceae Limonium lobinii N. Kilian & T. Leyens 1 10 13,58 R CR B1ab(ii); D 25 26 Plumbaginaceae Limonium sundingii Leyens, Lobin, N. Kilian & Erben 1 2 2,00 R CR D 27 Lobularia canariensis (DC.) Borgen subsp. fruticosa 28 Brassicaceae 5 33 192,69 I EN B1ab(ii,iv)+2ab(ii,iv) (Webb) Borgen 29 30 Lobularia canariensis (DC.) Borgen subsp. spathulata (J. A. Brassicaceae 2 11 26,29 I CR B1ab(iii) 31 Schmidt) Borgen 32 33 Fabaceae Lotus alianus J.H. Kirkbr. 2 1 1,00 NE NE 34 Fabaceae Lotus arborescens Lowe ex Cout. 1 4 4,00 R NE 35 36 Fabaceae Lotus brunneri Webb 5 33 354,46 LR NE 37 38 Fabaceae Lotus jacobaeus L. 2 26 305,68 NE NE 39 Fabaceae Lotus latifolius Brand 1 24 283,06 NE NE 40 41 Fabaceae Lotus purpureus Webb 7 58 514,75 NE NE 42 43 Lamiaceae Micromeria forbesii Benth. 5 52 366,77 I EN B1ab(ii,iv)+2ab(ii,iv) 44 Papaveraceae Papaver gorgoneum Cout. subsp. gorgoneum 2 8 39,96 VU CR B1ab(ii)+2ab(ii) 45 Papaver gorgoneum Cout. subsp. theresias Kadereit & 46 Papaveraceae 1 5 14,73 NE CR B1ab(ii)+2ab(ii); D 47 Lobin 48 Caryophyllaceae Paronychia illecebroides Webb 8(7) 103 1518,93 LR NT 49 50 Apocynaceae Periploca chevalieri Browicz 6 61 553,66 EN EN B1ab(ii)+2ab(ii) 51 52 Asteraceae Phagnalon melanoleucum Webb 5 46 255,60 NE EN B1ab(ii)+2ab(ii)

53 Arecaceae Phoenix atlantica A. Chev. 4 17 317,03 NE EN B1ab(iii)+2ab(iii) 54 55 Caryophyllaceae Polycarpaea gayi Webb 6 93 920,38 LR NT 56 57 58 26 59 60 Botanical Journal of the Linnean Society Page 27 of 31 Botanical Journal of the Linnean Society

ROMEIRAS ET AL.: Red List assessment of the Cape Verde endemic flora 1 2 3 Asteraceae Pulicaria burchardii Hutch. subsp. longifolia Gamal-Eldin 1 2 2,00 NE DD 4 Asteraceae Pulicaria diffusa (Shuttlew. ex Brunn.) Pett. 5(4) 20 344,42 VU EN B1ab(iv)+2ab(iv) 5 6 Sapotaceae Sideroxylon marginata (Decne.) Cout. 8(5) 24 456,31 EN EN B1ab(ii)+2ab(ii) 7 8 Solanaceae Solanum rigidum Lam. 7(5) 17 396,29 NE VU B1ab(ii)+2ab(ii) 9 Asteraceae Sonchus daltonii Webb 5 44 261,98 I EN B1ab(iv)+2ab(iv) 10 11 Sporobolus minutus Link subsp. confertus (J.A. Schmidt) Poaceae 2 4 4,00 R DD 12 Lobin, N. Kilian & Leyens 13 Verbenaceae Stachytarpheta fallax A.E. Gonç. NE EX 14 15 Teline stenopetala (Webb & Berthel.) Webb & Berthel. Fabaceae 1 1 1,00 NE CR D 16 subsp. santoantaoi Marrero-Rodr. 17 18 Asteraceae Tolpis farinulosa (Webb) Schmidt 5 28 192,45 I EN B1ab(ii)+2ab(ii) 19 Apiaceae Tornabenea annua Bég. 1 28 237,96 VU EN B1ab(ii)+2ab(ii) 20 21 Apiaceae Tornabenea bischoffii J. A. Schmidt 1 20 402,25 VU EN B1ab(ii)+2ab(ii) 22 23 Apiaceae Tornabenea humilis Lobin & K.PDF H. Schmidt Proof1 13 207,73 NE EN B1ab(ii)+2ab(ii) 24 Apiaceae Tornabenea insularis (Parl. ex Webb) Parl. ex Webb 3 22 84,18 LR EN D 25 26 Apiaceae Tornabenea ribeirensis Schmidt & Lobin 1 5 10,46 NE CR B1ab(iii)+2ab(iii) 27 28 Apiaceae Tornabenea tenuissima (A. Chev.) A. Hans. & Sunding 1 8 55,94 VU CR B1ab(ii)+2ab(ii) 29 Crassulaceae Umbilicus schmidtii Bolle 4 23 100,43 R EN B1ab(ii)+2ab(ii) 30 31 Scrophulariaceae Verbascum capitisviridis Hub.-Mor. 6(3) 57 582,22 VU VU B1ab(ii,iii)+2ab(ii,iii) 32 33 Scrophulariaceae Verbascum cystolithicum (B. Petterson) Huber-Morath 1 23 158,42 NE EN B1ab(ii)+2ab(ii) 34 Solanaceae Withania chevalieri A.E. Gonç. 4(3) 6 6,00 NE CR B1ab(ii)+2ab(ii) 35 36 588 *Number of islands where the species is known. In brackets: islands for which was obtained 37 589 data, when different from the known distribution. 38 39 590 **Red List categories according to Leyens & Lobin (1996), but criteria were not obtained at that 40 591 time. 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 27 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 28 of 31

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 Figure 1. Distribution of Cape Verde endemic species by threatened categories [Critically Endangered (CR); 23 Endangered (EN); Vulnerable (VU)]PDF in the archipelag Proofo (left) and in each island (right). Island abbreviations: 24 Santo Antão (ANT); São Vicente (VIC); Santa Luzia (LUZ); São Nicolau (NIC) (Northern Group); Sal (SAL); Boavista (BOA); Maio (MAI) (Eastern Group); Santiago (SAN); Fogo (FOG); Brava (BRA) (Southern Group). 25 291x124mm (300 x 300 DPI) 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Botanical Journal of the Linnean Society Page 29 of 31 Botanical Journal of the Linnean Society

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 PDF Proof 24 25 26 27 28 29 30 31 32 33 34 35 36 37 Figure 2. Species geographic range - AOO (area of occupancy) and EOO (extent of occurrence) - for the 72 38 endemic species classified under threatened categories [Critically Endangered (CR); Endangered (EN); 39 Vulnerable (VU)]. 40 117x107mm (300 x 300 DPI) 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Botanical Journal of the Linnean Society Botanical Journal of the Linnean Society Page 30 of 31

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 PDF Proof 24 25 26 27 28 29 30 31 32 33 34 Figure 3. Status change in IUCN Red List of the Cape Verdean endemic flora from 1996 (Leyens & Lobin 35 previous inventory) to 2015 (present assessment). Species classified in 1996 as Undetermined (applied 36 when it was not possible to accurate ly classify a species into any of the threatened categories CR, EN or VU) 37 or Rare (species restricted to isolated populations, and to which there was not enough information to 38 determine their conservation status, but corresponding most likely to CR, EN or VU) were considered here as Threatened. 39 162x131mm (300 x 300 DPI) 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Botanical Journal of the Linnean Society Page 31 of 31 Botanical Journal of the Linnean Society

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 PDF Proof 24 25 26 27 28 29 30 Figure 4. Incidence along elevation classes of the main threats to the Cape Verdean endemic plants. 31 Dimension of circles proportional to the number of species affected in the corresponding elevation class. 32 Classification of threats as defined by IUCN. (http://www.iucnredlist.org/technical-documents/classification- 33 schemes/threats-classification-scheme) 34 171x118mm (300 x 300 DPI) 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Botanical Journal of the Linnean Society