The cone snails of Cape Verde: marine endemism at a terrestrial scale

10 Cape Verde in the Eastern Atlantic is typical of many island groups in supporting a wealth of endemic 11 species both terrestrial and marine. Marine gastropod molluscs of the genus Conus, commonly 12 known as cone snails, occur in coastal tropical waters throughout the globe, but in Cape Verde their 13 endemism reaches its apogee with 53 out of 56 species occurring nowhere else, the majority of 14 which are restricted to single islands and frequently to single bays. However, Cape Verde is rapidly 15 moving to a tourism-based economy with a projected boom in infrastructure development often 16 coincidental with the shallow-water habitat of many range-restricted Conus . The conservation 17 assessment of Conus to standards of the International Union for the Conservation of Nature (IUCN) 18 Red List of Endangered Species, found that 45.3% of 53 species assessed from Cape Verde are 19 threatened or near-threatened with extinction compared to 7.4% of 579 species in the rest of the 20 world. The only three Conus species globally assessed as Critically Endangered and on the cusp of 21 extinction are all endemic to Cape Verde. Our analysis of Conus species distribution, together with 22 spatial data of coastal protected areas and tourism development zones, identify important areas for 23 future research and new marine protection. Our findings show that endemism with its associated 24 risks for Conus in Cape Verde has worldwide parallels with many non-marine taxa, while our 25 proposed strategy for Conus conservation extends beyond the confines of the country and this 26 taxonomic group. 27

ries together constituting only 9% 48 (Nshimyumuremyi & Simpasa, 2015). Cape Verde has few natural resources apart from marine 49 products and services and the land is generally unsuited to agriculture, such that around 80% of food 50 is imported (de Carvalho, 2013). 51 Tourism is now considered C V primary economic force and including directly associated 52 sectors, is responsible for 40% of gross domestic product (2014), forecast to increase to 49% by 53 2025 with visitor numbers expected to reach nearly 701 000 by 2025 (World Travel & Tourism  54 Council, 2015). In addition to attracting foreign investment tourism also drives the construction 55 sector (AfDB et al. 2013) including new harbour facilities at Porto Grande, São Vicente, and 56 international airports on Boa Vista and São Vicente to augment those already on Sal and Santiago. 57 Plans have also been agreed for the development of a large international casino on the islet of Santa 58 Maria off the southern coast of Praia, Santiago (Semedo & Gomes, 2015). 59 To support and develop tourism while protecting the natural environment, Integrated Tourism 60 Development Zones (ZDTI) have been delineated (Cabo Verde, 1994) and selected for geographical 61 location and landscape suitability (Fig. 2). Tourism Reserve and Protection Zones serving as buffers 62 to ZDTIs offer some natural protection from development, although incursion from ZDTIs into these 63 zones has made their value questionable (GEF/UNDP, 2013). Prior to development for each ZDTI a 64 management plan is required which must then undergo a full environmental impact assessment. Verde (Tables S1 & S2) was to be the establishment of a Protected Area Autonomous Authority (PAAA). The project was 80 scheduled for completion by December 2014, however, the latest report indicates that lack of 81 financing for the PAAA together with failure of enforcement of environmental legislation threatens 82 its sustainability (Laurie & Benchimol, 2013). 83 Currently, 27 PAs have been scheduled that encompass coastal elements of which one, Baía da 84 Murdeira, is exclusively marine (Fig. 2, Table S1). Protection of the shoreline, although typically 85 planned as a constraint on development in order to preserve the landscape, should coincidentally 86 reduce disturbance to taxa that dwell in the shallows. Unfortunately however, the PA network was 87 not created in a scientifically structured way with many areas selected on the basis of features such 88 as their landscape merit, bird nesting sites or recreational appeal without consideration to an overall 89 conservation objective (Vasconcelos et al., 2012). Additionally, they are subject to little in the way of 90 planned wildlife inventories or biodiversity monitoring programmes (Vasconcelos et al., 2012 has not been recorded for several years and has probably always been scarce. There are two other 263 NT species, namely C. navarroi and C. saragasae. As with C. curralensis and C denizi both occupy 264 highly restricted ranges in shallow water where they are at risk from pollution, over-gathering and 265 habitat loss, although not yet at a level where they are at immediate risk. 266

São Nicolau 267
The island only hosts C. kersteni, which is restricted to the southwest. This species has been 268 assessed as NT on a precautionary basis owing to its highly restricted range. regonae has its habitat extending to the north and south of the port. Both of these range-restricted 278 shallow water species are threatened because of risk to their habitat from marine pollution, in 279 particular the accidental discharge of oil from boat traffic including tankers and other commercial 280 vessels using the port of Palmeira. 281 C. mordeirae, with its population restricted to the bay that bears its name, has been observed to be 282 in decline, with the highest density of taxa occurring adjacent to resort developments. Similarly, C. 283 felitae occurs solely in the north of the bay where plans have been mooted to extend development. 284 Under such eventuality and in the absence of special conservation measures, this species may 285 require re-categorisation from VU to CR. 286

Boa Vista 287
Boa Vista has the greatest diversity of Conus with 21 species of which 15 are endemic to the island. 288 All three threatened species occur off the west coast of which C. salreiensis (CR) is only found in the 289 northwest of Boa Vista in the bay at Sal Rei and its adjacent islet. Harbour construction in the early 290 1990s impacted abundance and this species is now mainly found off the islet where it is at risk from 291 pollution and human disturbance. C. crotchii (EN) occurs from Morro de Areia south to Santa 292 Mónica in the centre of the new tourism zone where paved roads and resort hotels are under 293 construction. This places it at high risk from damage to habitat during the construction phase and of 294 continuing disturbance thereafter from holidaymakers. C. teodorae (VU) also occurs around Sal Rei 295 continuing north to Baía Teodora for 4.5 km. Around the southern half of its range it is subject to 296 the same pressures as C. salreiensis. 297 There are seven NT species found off Boa Vista of which five are endemic to the island: C derrubado 298 restricted to just five kilometres of coast in the north; C. diminutus which is found along two 2 299 kilometre sites in the west; C. evorai and C luquei which occur off Baía das Gatas in the northeast 300 with another population of C. evorai at the islet off Sal Rei; and C. trochulus which with C. josephinae 301 occurs along the western shores of Boa Vista adjacent to part of the development zone and 302 continuing north to Sal Rei. There is a sub-population of C. josephinae also on Maio. With the 303 exception of C. trochulus and C. josephinae, all these NT species have highly restricted ranges, and 304 although not at immediate risk as they are sufficiently remote from main centres of tourism, they 305 may become threatened in the future. C atlanticoselvagem (NT) occurs on the João Valente Shoals 306 which are only visited by lobster fishers, and although within SCUBA depths the shoals do not at 307 present attract divers. However, its solitary site and the potential for over-gathering or habitat 308 degradation have placed this species as a candidate for future review. There are no Threatened or Near Threatened Conus species on these islands except where they also 311 occur on other islands (Table 1). However, in the light of recent tourism resort and casino 312 development plans it may be necessary to review their assessments over the short term. 313 3.6. Proposed Priority Areas for Conservation (PPAC) 314 Proposed priority areas for future research (e.g. ground truthing surveys) and conservation (PPACs) 315 occur mostly around Sal, particularly to the southwest, and in the west of Boa Vista (Fig. 4). Other 316 smaller pockets occur around Santo Antão, São Vicente and Santa Luzia. The combined areas range 317 from 35 km 2 to 311 km 2 , depending on which level of PPAC is adopted, representing between 4% 318 and 33% of the entire range of Conus across Cape Verde (Table S3) and between approximately 319 0.12% and 1.11% of Cape Verde's territorial waters. Subject to physical survey, between 36% and 320 93% of species would be represented in any conservation initiative (Table S3) Areas requiring the most urgent research and potential protection (PPAC 1) are located around the 333 islands of Boa Vista, Sal and São Vicente ( Fig. 4  inadequate genetic diversity to ensure a healthy population (Briggs, 1966). 347 There are many who consider marine taxa to be less susceptible to extinction risk than terrestrial 348 species ( strategies. To achieve this we strongly support the establishment of a Protected Area Autonomous 420 Authority (PAAA). We recognise that marine protection can only be effective if it is supported by the 421 populace and in particular by those who are likely to feel disadvantaged by its implementation such 422 as fishers. Consideration therefore needs to be given to the impact on current activity within the 423 proposed areas and how regional authorities, whose responsibility will be to manage and

Role of the funding source 461
The funders played no role in the study, design, collection, analysis and interpretation of data; 462 writing of the report; and in the decision to submit the article for publication. 463    Tables   703   Table S1. Legally  Areas of special ecological and scientific interest. There are two further subsets Partial Natural Reserves and Temporal Natural Reserves. Partial Nature Reserves offer protection to a specific natural resource, whether a single species, group of species or a particular habitat. Uses are permitted that are compatible with the purpose of protection. Temporal nature reserves are small areas established for a limited period of time to allow recovery of the resource or of specific ecological systems.

Integrated Natural Reserve (5 designated)
Areas of special ecological and scientific interest. Integral Natural Reserves offer protection to the entire ecosystem. They restrict further development and human use. National Park (None designated) Areas unaffected by human exploitation and occupation which have a special scientific, socio-economic, educational, recreational or landscape aesthetic interest. Exploitation and human occupation are prohibited beyond visits for recreational or cultural purposes.

Natural Park (11 designated)
Large areas containing predominantly natural habitats, species or representative samples of the country's biodiversity. Within these there may be a traditional local population. Natural Parks aim to conserve, protect and/or restore natural environments and cultural resources: they promote socio-economic development compatible with nature conservation to improve the quality of life for local communities; educational, recreational and scientific use is encouraged. Natural Monument (6 designated) Moderate sized areas which contain at least one natural or cultural element of exceptional value (e.g. rarity, uniqueness, scientific interest, ecological or cultural function). They are protected to safeguard the feature(s) of interest and prohibit activities which changes them.

Protected Landscape (10 designated)
Terrestrial or coastal areas modified by human activity with a particular aesthetic quality or cultural value. Protection focuses on preserving and restoring the characteristics that define them.

Site of Scientific Interest (None designated)
Areas, usually of a small size, which contain natural elements of scientific interest or animal or plant populations threatened with extinction.