Sites for priority biodiversity conservation in the Caribbean Islands Biodiversity Hotspot

The Caribbean Islands Biodiversity Hotspot is exceptionally important for global biodiversity conservation due to high levels of species endemism and threat. A total of 755 Caribbean plant and vertebrate species are considered globally threatened, making it one of the top Biodiversity Hotspots in terms of threat levels. In 2009, Key Biodiversity Areas (KBAs) were identified for the Caribbean Islands through a regionallevel analysis of accessible data and literature, followed by extensive national-level stakeholder consultation. By applying the Vulnerability criterion, a total of 284 Key Biodiversity Areas were defined and mapped as holding 409 (54%) of the region’s threatened species. Of these, 144 (or 51%) overlapped partially or completely with protected areas. Cockpit Country, followed by Litchfield Mountain Matheson’s Run, Blue Mountains (all Jamaica) and Massif de la Hotte (Haiti) were found to support exceptionally high numbers of globally threatened taxa, with more than 40 such species at each site. Key Biodiversity Areas, building from Important Bird Areas, provide a valuable framework against which to review the adequacy of existing national protectedarea systems and also to prioritize which species and sites require the most urgent conservation attention.


INTRODUCTION
The Caribbean Islands Biodiversity Hotspot is exceptionally important for global biodiversity conservation, due to high levels of species endemism and threat.The Caribbean is home to approximately 11,000 plants species, of which 72% are endemic to the region.The vertebrates are also characterized by extremely high levels of species endemism: 100% of 189 amphibian species, 95% of 520 reptile species, 74% of 69 mammal species and 26% of 564 species birds are unique to the Caribbean Islands.In terms of endemism at the genus level, it ranks third among the world's 34 Biodiversity Hotspots with 205 plants and 65 vertebrate genera endemic to the islands (Smith et al. 2004).Species restricted to the Caribbean Islands Biodiversity Hotspot represent 2.6% of the world's 300,000 plants species and 3.5% of the world's 27,298 vertebrate species (Wege et al. 2010).
The high level of biological diversity in the Caribbean is due to several factors.During the early Cretaceous (120 to 140 million years b.p.), a chain of volcanic islands (called Proto-Antilles) began to emerge along the eastern edge of the Caribbean Plate in the Pacific Ocean.The plate drifted eastward serving as a stepping-stone route for limited exchange of terrestrial organisms between the Nearctic and Neotropical regions.By the Eocene (58 million years b.p.), the core of the Greater Antilles achieved their present positions (Brown & Lomolino 1998).The Lesser Antilles are the active remnants of an ancient volcanic chain, and are younger than the Greater Antilles.Several islands have particularly rugged and mountainous landscapes separated by large stretches of sea, which resulted in the isolation of populations and eventually to speciation.
The Caribbean has suffered from high levels of habitat loss since the arrival of Europeans in the 1490s.This destruction has reduced the hotspot's original estimated 229,549km 2 of natural vegetation to just 22,955km 2 (or just 10%).The loss of native habitat combined with other threat factors, such as introduced (alien invasive) species, has resulted in severe and widespread degradation of the Caribbean's unique biodiversity.Currently, 755 plants and vertebrate species are at risk of extinction, making the region one of the biodiversity hotspots holding the most globally threatened species.
National governments and donor agencies have primarily invested in developing protected areas systems to halt biodiversity loss.However, these are rarely comprehensive in their overlap with unique species and habitats, are frequently inadequately managed, and often fail to protect important places for biodiversity.Key Biodiversity Areas (KBAs) in the Caribbean can be used as a tool for reviewing the efficacy of existing national protected-area systems.KBAs provide a site-based framework against which gaps in protected-area coverage can be identified and candidate sites for expansion.This paper has three objectives.First, to explain the identification process for Caribbean KBAs that was conducted within the context of developing the Caribbean Islands Ecosystem Profile for the Critical Ecosystem Partnership Fund.Second, to evaluate how the identified KBAs are represented in the region's existing protected-area systems, therefore highlighting key gaps in them, and to prioritize among the KBAs for conservation action.Finally, to evaluate the effectiveness of the KBA approach in guiding conservation priorities in the Caribbean Islands Hotspot.

METHODS
The Caribbean Islands Hotspot (Image 1) includes the biologically and culturally diverse islands of The Bahamas (Image 2), Greater Antilles (Puerto Rico (Image 5), Jamaica (Image 3), Cuba (Image 2) and Hispaniola [comprising the countries of the Dominican Republic and Haiti] (Image 4)), Virgin Islands, Cayman Islands, Lesser Antilles (Image 5) and the Netherlands Antilles (Image 6).It does not include Trinidad and Tobago.So defined, the hotspot represents a complex geopolitical region of 12 independent nations, and six British and three U.S. overseas territories, two French overseas départements, two French overseas collectivités, three special municipalities of the Netherlands and two constituent countries within the Kingdom of the Netherlands.The hotspot encompasses more than four million km 2 of ocean (not included in the analysis) as well as c. 230,000km² of land area, with the four islands of Cuba, Hispaniola, Jamaica and Puerto Rico making up about 90% of this.The coastal area included in the analysis embraces territorial waters up to 12 nautical miles.The elevational range spans from over 3,000m above sea level to 40m below sea level and encompasses a diverse array of habitats and ecosystems.The Caribbean KBAs were identified as an integral part of developing the Caribbean Islands Hotspot Ecosystem Profile for the Critical Ecosystem Partnership Fund during 2009 (Wege et al. 2010).For the Caribbean Islands Hotspot, only the Vulnerability criterion was applied to select KBAs, with the confirmed presence of globally threatened species triggering the definition of a KBA.The Irreplaceability criterion was not applied due to lack of quantitative data for other taxa in the hotspot.This criterion is currently only used to define KBAs for birds, as this is the only group for which the concept of restricted-range species has been quantitatively defined: species with global breeding ranges of less than 50,000km 2 (Stattersfield et al. 1998).However, to prevent a bias toward site priorities for birds, KBAs in the Caribbean Islands Hotspot are identified only based on the Vulnerability criterion.
The KBA identification process comprised a deskbased, regional-level analysis of accessible data and literature, followed by national-level stakeholder consultations coordinated by BirdLife International, in collaboration with the Durrell Wildlife Conservation Trust, the University of Bath and the New York This foundation of IBAs was supplemented by applying the Vulnerability criterion to non-avian taxa to define terrestrial KBAs based on the occurrence of globally threatened species as categorized on the 2008 IUCN Red List (a global, standardized assessment of species threat status).The taxonomic groups used to define KBAs were land-based mammals, birds (through the IBA process previously mentioned), amphibians, reptiles and plants.Sea turtle nesting beaches were included in the analysis where more than 100 crawls annually had been recorded (Dow et al. 2007).KBAs were delineated using a geographic information system (GIS) and by taking into consideration the distribution of available habitat for the globally threatened species, and also land/ protected area management units.The degree of protection of the KBAs was analyzed against the 2010 World Database on Protected Areas (WDPA).
National profile coordinators in The Bahamas (Bahamas National Trust), Dominican Republic (Grupo Jaragua), Haiti (Société Audubon Haïti), Jamaica (BirdLife), Lesser Antilles (BirdLife in Barbados) and Puerto Rico (Sociedad Ornitológica Puertorriqueña) facilitated a review of priorities within their own countries.In Cuba, it was not possible to conduct the analysis for taxonomic groups other than birds.National workshops were held in Dominican Republic, Haiti and Jamaica during June 2009, with a region-wide workshop held in July 2009 on Antigua as a formal part of the 17 th Regional Meeting of the Society for the Conservation and Study of Caribbean Birds.

RESULTS
A total of 284 Key Biodiversity Areas were defined for all the countries and territories contained within the Caribbean Islands Biodiversity Hotspot (Table 1).The 284 KBAs in the Caribbean Islands Hotspot cover just over 50,000km 2 , or roughly 22% of the terrestrial and coastal portion of the hotspot.Half of the KBAs are either fully or partially protected according to the 2010 WDPA (Images 1-6).
In total, 409 globally threatened plants and vertebrate species triggered the Vulnerability criterion to define KBAs (hereafter referred as trigger species).No data were available for the other 346 globally threatened species, most of which were plants, at the time of this analysis.Plants had the highest number of globally threatened species, while for vertebrate species groups, amphibians and then birds were the most abundant groups (Table 2).However, globally threatened birds were responsible for defining the largest number of KBAs, followed by amphibians and plants (Table 3).The majority of KBAs were defined by the presence of multiple globally threatened trigger species, with almost half of all KBAs (141 or 49.64%) supporting two to 10 trigger species and 28 (or 9.86%) by 11 to 44 species.Cockpit Country in Jamaica supports an incredible 60 globally threatened species, followed by Litchfield Mountain-Matheson's Run, Blue Mountains (both with 44) and Massif de la Hotte in  The percent coverage by KBAs can be over 100% as some KBAs include both land and water area, while the total area for each country is calculated based solely on land area.Cuban key biodiversity areas comprise only sites (IBAs) identified as important for globally threatened birds.Haiti (with 42).On the other hand, 114 (or 40%) KBAs were identified for a single trigger species (Table 4).A total of 173 (or 42.29%) of the trigger species have distributions confined to just one KBA (see Table 5).However, 25 KBAs hold populations of more than one of these "single-site" species with Cockpit Country, Blue Mountains and Massif de la Hotte each supporting populations of more than 20 such species.An important result of the KBA process was the identification of 56 KBAs which contained the only record of a globally threatened species in the Caribbean Islands Biodiversity Hotspot.No fewer than 19 of these KBAs were identified in 2010 as global Alliance for Zero Extinction (AZE) sites (Tables 4 and 5)-defined by the presence of Critically Endangered or Endangered species confined to just a single site.One hundred and sixty nine (169 or 41.32%) of the trigger species occur only in two to five KBAs and the other 67 (16.38%) trigger species occur in more than six KBAs (Table 4).

DISCUSSION
The countries with the greatest numbers of Key Biodiversity Areas are the large islands of the Greater Antilles and multi-island countries such as The Bahamas.This is to be expected as the principles of island biogeography dictate that the larger (and older) the island, the greater the species diversity.Higher species diversity on each of the Greater Antilles, combined with greater ecosystem, habitat and altitudinal diversity, has led to larger numbers of endemic species and consequently higher numbers of globally threatened taxa.Small islands in archipelagos such as The Bahamas often result in taxonomic isolation and the presence here of globally threatened species occupying very small ranges (often confined to a single island) has in turn led to the definition of relatively large numbers of KBAs.In Cuba, the KBAs included only IBAs as it was not possible to incorporate the results of analyses of other taxonomic groups or consultations with experts for the definition of other sites.
The amount and quality of available data on the distribution of globally threatened species among Caribbean KBAs vary between taxonomic groups but the sites identified will almost certainly be important for other groups for which data are not currently available.However, there are likely to be additional  threatened, and no KBAs were defined for this group.
In addition to the occurrence of globally threatened species (the Vulnerability criterion), KBAs can also be defined on the basis of the presence of restricted-range species (the Irreplaceability criterion): their inclusion as a next step may result in a better coverage of the poorly represented taxonomic groups mentioned above.Sites regularly supporting significant populations of restricted-range species are global conservation priorities because there are few or no other sites in the world where conservation action for these species can be taken.However, there are no quantitative data for restricted-range species (other than birds) in the hotspot and thus this criterion could not be applied.
The main threats to the terrestrial biodiversity of the insular Caribbean, as prioritized during the national ecosystem profiling workshops in order of highly significant regional threat or impact are: invasive species; residential and commercial development; severe weather events and global climate change; agricultural expansion and intensification; overexploitation of natural resources; mining and energy production; pollution; transportation; and geological events.There is a complex mix of interacting socioeconomic, political, cultural and environmental factors that are driving environmental change and threatening biodiversity (and thus the KBAs) in the insular Caribbean.Principal among these are the increasing human population and material consumption, poverty and inequitable access to resources, the inherent economic and environmental vulnerability of the islands to external forces, such as changes in global trade regimes, and climate change.Some of these, such as poverty, are local or national issues, while others, such as climate change, require attention at the global level.All these drivers can be either exacerbated or mitigated by public policies and institutional arrangements, at national, regional and international levels.
A number of constraints need to be overcome to address the environmental threats and achieve more effective conservation of biodiversity and ecosystem services.The main ones discussed during the national workshops and consultations in order of highly significant regional barrier are: weak and ineffective policy; poor land-use planning; limited capacity and resources for biodiversity conservation; inefficient institutional frameworks; poor participation by stakeholders; limited technical and scientific knowledge for decision-making; and lack of awareness of biodiversity and ecosystem services (particularly their value) among decision-makers and the general public.
Most countries have significantly updated, or are in the process of updating (e.g.Haiti and St.Vincent), their policies and legislation on biodiversity, environmental management and sustainable development while their obligations under international agreements have helped drive this process (Brown et al. 2007).However, there exists significant variation among countries with regard to comprehensiveness and effectiveness, particularly concerning the protection of threatened biodiversity and ecosystems (BirdLife International 2008), and there is a need for specific analyses of "gaps" in legislation and policies, which very few countries have undertaken (an exception being Jamaica [NEPA 2003]).Overall, national public policy frameworks for environmental management remain largely oriented toward control, regulation and a reactive approach to environmental issues, although new approaches and instruments, including environment service markets, have begun to be promoted by some donors, governments and NGOs as means of changing destructive patterns of behavior.
Environmental policy in the Caribbean tends mostly to address environmental issues and impacts rather than their underlying root causes/drivers, such as human population increase.In the case of climate change, Caribbean countries do not consider themselves to be net contributors and therefore policy responses are largely limited to adaptation.All the countries in the hotspot are active participants in the main multilateral environmental agreements.All are signatories to the three "Rio Conventions"-the UN Convention on Biological Diversity, UN Convention to Combat Desertification, and the UN Framework Convention on Climate Change -and most are members of the other key biodiversity related agreements, such as the Ramsar Convention, the World Heritage Convention and the Convention on International Trade in Endangered Species, but not the Convention on Migratory Species.At the regional level, the main agreement is the Convention for the Protection and Development of the Marine Environment of the Wider Caribbean Region (Cartagena Convention) and its three Protocols.These together constitute the only legal instrument for regional cooperation on environmental issues for the wider Caribbean, although not all signatory countries are properly meeting their commitments, e.g.required legislation not enacted, management plans not developed, and biodiversity action plans not in place.
Key Biodiversity Areas can achieve the following: (i) Help Caribbean nations honour their commitments to multilateral environmental agreements (ii) Provide the basis for protected area gap analyses (as is the case in Haiti where KBAs are being used in the development of the national system for protected areas plan) (iii) Provide a prioritized framework within which to monitor the status of biodiversity in the region.

Sierra del Rosario
Accipiter gundlachi EN

Xenoligea montana VU
Parque Nacional del Este Amazona ventralis VU

Xenoligea montana VU
Massif de la Selle Amazona ventralis VU

Image 1 .
Partially or completely protected (in green) and unprotected (in red) Key Biodiversity Areas [2010] in the Caribbean Islands Biodiversity Hotspot.Image 2. Partially or completely protected (in green) and unprotected (in red) Key Biodiversity Areas [2010] in the Bahamas and Cuba.

Image 3 .
Partially or completely protected (in green) and unprotected (in red) Key Biodiversity Areas [2010] in Jamaica.Image 4. Partially or completely protected (in green) and unprotected (in red) Key Biodiversity Areas [2010] in Hispaniola.Botanical Garden, and with technical support from Conservation International.Important Bird Areas (IBAs), documented by BirdLife partners and collaborating organizations in 2008 (BirdLife International 2008), were used as a scientifically robust starting point for KBA identification.

Image 6 .
Partially or completely protected (in green) and unprotected (in red) Key Biodiversity Areas [2010] in Aruba and Bonaire.Image 5. Partially or completely protected (in green) and unprotected (in red) Key Biodiversity Areas [2010] in Puerto Rico and the Lesser Antilles.

Table 4 . Total frequency distributions of the number of trigger species per KBA, and the number of KBAs per trigger species in the Caribbean Islands sites
holding globally threatened species that have not been identified during this process.This is because reptiles, plants (especially cacti and orchids) and bats have not been systematically assessed against Red List criteria.Neither have freshwater fish (of which there are numerous endemics in the region), such that just five have so far been categorized as globally