Geo-political genetics: Claiming the commons through species mapping

Geoforum 41 (2010) 897–907

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Geo-political genetics: Claiming the commons through species mapping Lisa M. Campbell a,*, Matthew H. Godfrey a,b a b

Nicholas School of Environment, Duke University, 135 Duke Marine Lab Road, Beaufort, NC 28516, USA North Carolina Wildlife Resources Commission, 1507 Ann Street, Beaufort, NC 28516, USA

a r t i c l e

i n f o

Article history: Received 6 May 2009 Received in revised form 15 April 2010

Keywords: Science Sea turtles Ocean commons Political ecology Conservation genetics Conservation territories

a b s t r a c t Genetic techniques are increasingly employed in the field of conservation biology; our understanding of sea turtle biology, and particularly of sea turtle migrations and population structures, has increased through genetic analyses that ‘match’ turtles found in various and often widely distributed habitats (e.g. nesting beaches, foraging grounds, migratory corridors). This relatively recent technological development has implications for how sea turtles are conceived, both as resources and as objects of conservation. Traditionally, sea turtle populations have been identified with nesting beaches, and most conservation efforts have been focused on these discrete geographic locations and undertaken by the state. The more complete understanding of relationships among turtles found in geographically disparate areas, achieved via genetic analysis, can take conservation beyond the beaches and territorial waters of individual states; foraging populations can now be linked to nesting populations sometimes hundreds of kilometers distant. In this paper, we explore the implications of genetic analysis for sea turtle conservation, the scale at which it is undertaken, and the variety of actors with competing interests in it. We focus on the case of hawksbill sea turtles in the Caribbean, where genetic data are invoked in conservation conflicts. We are particularly interested in the way genetic data can support the scaling-up of sea turtle conservation, creating new ‘conservation territories,’ and we draw on political ecology and common pool resource theory to explore the implications thereof. Ó 2010 Elsevier Ltd. All rights reserved.

1. Introduction Contemporary wildlife conservation involves debates and decisions about rights to wildlife as resources (Brockington et al., 2006), the scale at which wildlife should be managed (Brosius and Russell, 2003), and the role of science in dictating conservation policy (Gray and Campbell, 2009; Lackey, 2007). The issues can be resolved, reconciled, or prioritized in multiple ways, with varied ‘conservation territories’ resulting. Zimmerer (2006, p. 67) defines conservation territories as the ‘‘human-designed spaces of nature protection and resource management”, thus invoking elements of both physical space and governance. In this paper, we use the case of sea turtle conservation to explore the interaction of rights, scale, and science in informing conservation territories, and how arguments made on one issue can further goals on others. We specifically consider the implications of recent advances in genetic analysis, a scientific advancement, for the scale at which conservation is undertaken, and what this implies for the rights of various stakeholders to determine how sea turtles should be managed. Sea turtle conservationists have increasingly called for the * Corresponding author. Tel.: +252 504 7628; fax: +252 504 7648. E-mail addresses: [email protected] (L.M. Campbell), [email protected] (M.H. Godfrey). 0016-7185/$ - see front matter Ó 2010 Elsevier Ltd. All rights reserved. doi:10.1016/j.geoforum.2010.06.003

consideration of sea turtles as regionally or internationally shared resources, to be managed as such (e.g. Frazier, 2000; MTSG, 1995), but until recently there were few data available to describe the precise nature of such sharing. Genetics has changed this and, by extension, changed related debates about how to best conserve sea turtles. While genetic analyses of sea turtle populations – and discussion of the conservation implications – are now widespread (e.g. Naro-Maciel et al., 2007), we focus on the case of hawksbill sea turtles (Eretmochelys imbricata) in the Caribbean Sea, where conflicts over their conservation have spurred genetics research and where genetic data have been applied in attempts to resolve those conflicts. Specifically, we look at efforts by Cuba to trade hawksbill shell with Japan, a case that has been the focus of international debate in the sea turtle conservation community for over a decade. We explore how efforts to define ownership of sea turtles (rights), and in particular to move away from local or national and towards regional or international management (scale), can be supported using genetic data (science). We draw on common pool resource theory and political ecology to explore these transitions and their implications for how we conceptualize conservation territories. Specifically, we: (1) situate genetics in sea turtle conservation generally; (2) describe the role of genetics in the Cuban hawksbill debate; (3) illustrate various possible ways of conceptualizing

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conservation territories that result from the genetic data on Caribbean hawksbills; and (4) consider how other ways of conceptualizing conservation interact with or are displaced by genetic arguments. While our focus is on sea turtles, our argument is more widely relevant. Genetic analyses have profound implications for the study and practice of conservation biology, and there is discussion in the community itself on merits and drawbacks of the ‘genetic’ turn in the field (DeSalle and Amato, 2004; DeYoung and Honeycutt, 2005).1 Conservation of various terrestrial species has been influenced by genetic data, from African elephants (Okello et al., 2008) to Japanese flowering cherry trees (Tsuda et al., 2009). In the USA, the National Marine Fisheries Service relies heavily on genetic analyses to determine management units, and management units are the basis of recovery action plans under the Endangered Species Act (e.g. Krahn et al., 2004; NMFS and USFWS, 2008). An analysis of ‘geopolitical genetics’ may be particularly relevant for contemporary debates about how to best conserve marine resources. The current ‘crisis’ in marine conservation has led to renewed efforts to rethink how common pool marine resources, both within and outside of national jurisdictions, should be conceptualized and managed (Steinberg, 2008; Gray, in press). Given the emphasis of these efforts on clarifying rights and establishing territories through tools like marine protected areas and marine spatial planning (Gray, in press; Mims et al., 2009), the case explored here is timely and may offer insights into some of the issues that will arise as the conservation community seeks new approaches to marine conservation and governance.

2. Conserving sea turtles: common property and political ecology perspectives on the question of scale As with other species categorized as charismatic mega-fauna, sea turtle conservation is highly politicized (Bjorndal and Bolten, 2003; Broderick et al., 2006; Meylan, 1998; Mrosovsky, 1983, 1997; Webb and Carrillo, 2000). This is in part because sea turtles hold wide public appeal and are valued in a variety of ways (Campbell, 2003; Campbell and Smith, 2006). Historically, and up until as recently as the 1970s, sea turtles were widely used for their meat, eggs, shell, skins, and oils (Campbell, 2003; Frazier, 2003). A shift towards greater protection of sea turtles began in the 1960s and 70s, based on concerns about perceived population declines for sea turtles specifically (Bjorndal, 1981) and with the emergence of new forms of environmentalism in North America and Europe (McCormick, 1989). Through a variety of international agreements, such as the 1975 Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), and national laws such as the 1973 USA Endangered Species Act, the legal use of and trade in sea turtles and their products was curtailed in many parts of the world. Restrictions on use have continued through more recent agreements such as the 2002 Inter-American Convention for the Protection and Conservation of Sea Turtles (IAC) (Campbell et al., 2002), pressure on particular nations to live up to existing agreements, and incentive programs designed to encourage alternative uses of sea turtles, through activities like ecotourism (Troëng and Drews, 2004). In spite of conservation efforts made over the past 40 years, six of seven species of sea turtles are currently classified as facing some risk of extinction on the International Union for Conservation 1 Some of the relevant areas in conservation that can be informed by genetic data include resolving population structure, defining management units within species, forensics, pedigree analysis, and estimating population size and sex ratio (for a general overview, see DeSalle and Amato (2004)). However, the application of genetics data to management actions is most powerful in combination with behavioral and demographic information (DeYoung and Honeycutt, 2005).

of Nature’s (IUCN) Red List.2 Although the Red List aims to provide ‘‘the most objective, scientifically-based information on the current status of globally threatened biodiversity” (www.iucnredlisit.org), the red listing process has been controversial for sea turtles (Godfrey and Godley, 2008; Mrosovsky, 2003; Seminoff and Shanker, 2008). Of interest here is the question of scale, and whether or not a global categorization of endangerment is appropriate for widely distributed species. Leatherback sea turtles (Dermochelys coriacea), for example, are experiencing different regional population trends: nesting is declining in the Pacific, while nesting is increasing in the Atlantic, (Spotila et al., 2000; Turtle Expert Working Group, 2007). For the wider region and species of interest in this paper, the situation is also mixed. Some hawksbill populations are believed to be declining (e.g. Caribbean Costa Rica, Troëng et al., 2005) while others are believed to be increasing (e.g. Brazil, Marcovaldi et al., 2007), and nesting trends can vary among beaches within countries. In this context, global assessments of threat may not be the most informative. The question of the appropriate scale at which sea turtle populations should be conceptualized is central to debates about sea turtle conservation more generally (Campbell, 2007a). In policy, a tension exists in statements made by organizations like the Marine Turtle Specialist Group (MTSG) of the IUCN that on one hand advocates regional or international approaches, while on the other encourages community-based approaches (Campbell et al., 2002; Frazier, 1999, 2000). In practice, contemporary conservation varies across nation states. Although the international focus of sea turtle conservation has moved increasingly towards strict protection (Campbell, 2002a), sea turtles are still used consumptively, both as adults and eggs, in many nations where turtles nest or are found in territorial waters (Campbell, 2003, 2007b). In regions like the Caribbean, where the jurisdictions of many nations co-exist and sometimes overlap in a relatively small geographic area, the full spectrum of approaches to sea turtle conservation can be found, ranging from strict protection (e.g. US Virgin Islands), to smallscale fisheries (e.g. Turks and Caicos Islands), to larger-scale fisheries (e.g. Caribbean Nicaragua), to a commercial sea turtle farm in Cayman Islands (Campbell et al., 2009). In this context, the contrasting approaches to conservation at the international, regional, national, and local levels are striking, and can result in conflict when the actions of one nation are perceived as impacting on the rights of another. This is exacerbated by the highly migratory nature of sea turtles, which often traverse hundreds of kilometers between nesting and foraging areas, across social–political and geographic scales (Campbell, 2007a). In common pool resource theory, the appropriate scale at which to manage resources and concerns about the potential for scalar mismatches have emerged as important themes (Berkes, 2006; Cash et al., 2006; Cumming et al., 2006). Scalar mismatches occur when the scale of the ecological process or resource extraction occurs at wider or finer scales than associated governance institutions (Cumming et al., 2006). Mismatches are particularly problematic for highly migratory marine resources, where the problem of exclusion, i.e. the inability of resource users to exclude others, is amplified (Berkes, 2006). Scalar mismatches are a central concern for sea turtle biologists, who question the ability of any one state to account for and manage these highly migratory animals (Campbell, 2000, 2002a). According to Berkes (2006), for migratory marine species, local and even national oversight is not enough: ‘‘an international agreement becomes necessary to solve the scale mismatch problem.” Berkes recognizes, however, that attempts to resolve one set of scalar mismatches can create others. For example, that a resource is highly migratory does not negate the need for local level user groups to be involved in

2

The seventh is listed in the ‘data deficient’ category.

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management; while local involvement in international agreements is theoretically possible, weak vertical linkages between scales can limit this potential (Berkes, 2006).3 The scale issue is particularly complex in the Caribbean, where the dense concentration of many nations makes most marine resources shared and supports calls for international collaboration and management (Chakalall et al., 2007), but where the nature of governance institutions and limited potential for state-led top-down enforcement requires greater collaboration between states and resource users (Charles et al., 2007). The question of scale is further complicated when we problematize the concept itself, an issue of considerable interest to geographers and political ecologists. As Swyngedouw and Heynen (2003, p. 913) describe it: ‘‘The continuous reorganisation of spatial scales is an integral part of social strategies to combat and defend control over limited resources and/or a struggle for empowerment.” Brown and Purcell (2005) call for more attention to the ‘politics of scale’ in particular; rather than taken as ontologically given, concepts of local, national, and international should be seen as socially constructed, both fluid and fixed, and as relational ideas. McCarthy (2005) illustrates how conservation actors strategically reinforce or challenge traditional concepts of scale, depending on the argument being made. Both Brown and Purcell (2005) and McCarthy (2005) highlight that questions about socio-political scale should not (and perhaps cannot) be separated from biophysical scale; scale is produced ‘‘simultaneously by social processes and by biophysical processes that elude human control” (McCarthy 2005, p. 736). However, neither McCarthy (2005) nor Brown and Purcell (2005) problematize our understanding of biophysical processes and ecological scale, or the ways that these can also be constructed (rather than given) in support of particular scalar politics. For example, Campbell (2007a) shows how scientists selectively employ ecological data to support conservation goals at particular socio-political scales. Thus, although both common pool resource theory and political ecology have interests in scale and scalar politics, here we call attention to the role of science (i.e. the understanding of biophysical processes, ecology, and biology of sea turtles) in such scalar politics. 3. What we know, how we know it, and what we do with it: the biology, ecology, and politics of sea turtle conservation Biological understanding of sea turtles, and the ability of such understanding to inform conservation policy, has traditionally been limited by sea turtle life history characteristics. Turtles are most accessible on the nesting beaches as adult females, hatchlings, and eggs. Our ability to study juveniles of both sexes and adult male turtles has been constrained by the difficulties of conducting in-water studies and because of the long distance migrations turtles make between nesting and foraging grounds. Thus, one of the challenges for understanding sea turtle ecology has been the difficulty of linking individual turtles found at different life stages and using distinct habitats to larger populations. Advances in tagging turtles, particularly using satellite telemetry, have greatly increased our understanding of individual turtles and their movements, but small sample sizes and the absence of tracking data on hatchlings means that conventional tagging is limited in how much it can reveal about sea turtle populations (Godley et al., 2008). This question of the relationships between animals found at various life stages in various places is one that genetic data can help to answer. For sea turtles, genetic studies have largely focused on maternally inherited mitochondrial DNA (mtDNA), and look for unique 3 Campbell et al. (2002) illustrate the difficulties of integrating local communities into decision making in the Inter-American Convention for the Conservation of Sea Turtles.

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markers (or haplotypes) along specific regions of mtDNA sequences. The elucidation of different haplotypes (or different ratios of haplotypes) from specific sea turtle nesting rookeries is a powerful tool that has greatly increased our understanding of sea turtle biology and behavior. For example, genetic data have proved a long-standing hypothesis that adult female sea turtles return to breed and lay eggs on or close to the beach where they were born, a process known as natal homing (Dutton et al., 1999; FitzSimmons et al., 1997; Meylan et al., 1990). Comparisons of haplotype diversity among sea turtle nesting assemblages has led to the conceptualization of different rookeries as independent management units (e.g. Encalada et al., 1998; FitzSimmons et al., 1997; Schroth et al., 1996), a concept central to our analysis in Section 4.4 Nevertheless, the full impact of genetic analyses for sea turtles has been limited due to gaps in the understanding of general demographic information for all sea turtle species, including growth rates, age at sexual maturity, and survival rates (Heppell et al., 2003). That genetics are used to describe ‘management units’ speaks to their importance in conservation practice; what genetic data tell us matters for how sea turtles are used and for the people who use them. And, as with other examples of charismatic species, the science and politics of sea turtle conservation have proven difficult to separate (see Thompson (2002, 2004) on the case of African Elephants). Concerns about the use of misuse of science have been expressed on a variety of issues related to sea turtle conservation, including the use of genetics in distinguishing between different species of sea turtles. For example, there has long been debate about whether or not the green turtle (Chelonia mydas) found in the Pacific is a distinct species (Chelonia agassizi) (Mrosovsky, 1983). In an exchange published in Conservation Biology, Pritchard (1999) argued that morphological differences between Pacific and Atlantic green turtles were enough to warrant the distinct species status, and that such status would be beneficial to conservation. Bowen and Karl (1999) and Karl and Bowen (1999) opposed this based on their genetic analysis. While definitions of a species have long plagued biologists (Rojas, 1992), what is most interesting about the exchange is the extent to which implications for conservation were explicitly addressed in a published forum. The establishment of the Pacific green turtle as a distinct species essentially divides global green turtle population numbers. Thus, both the green and the black turtle might be categorized as more highly endangered since, as separate species, there would be fewer individuals of each. Karl and Bowen (1999) argued that the black turtle was a ‘geopolitical’ rather than taxanomic species, and reported they had been pressured ‘‘to downplay or ‘reinterpret’ genetic data on the evolutionary distinctiveness of the black turtle” (Bowen and Karl, 1999, p. 1014) to serve conservation purposes. In a more recent attempt to redefine a sea turtle species, a petition was submitted to the USA Federal government in 2002 to officially designate loggerhead sea turtles (Caretta caretta) nesting from northern Florida through North Carolina as a distinct subpopulation of loggerheads nesting along the US SE coast, based in part on genetic discreetness. Under the ESA, the larger population is threatened; a reclassification likely would have made the northern subpopulation endangered (FWS and NMFS, 2003). The US Federal government ruled in 2003 that there was not sufficient genetic evidence to support the recognition of a separate subpopulation north of Florida, and therefore the classification of this species was unchanged (FWS and NMFS, 2003). In 2008, a new US Recovery Plan for loggerheads in the NW Atlantic was published, and it

4 Although there is evidence of male-mediated gene flow among these different management units (Roberts et al., 2004), the argument for the dominance of mtDNA data in defining management units is that natal homing exhibited by adult females implies that a rookery will not be re-established if all females are extirpated (Bowen and Karl, 2007).

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recognized separate management units for loggerheads within the SE USA, based in part on genetic discreteness (NMFS and USFWS, 2008). The northern management unit is essentially the same as that proposed in the rejected 2002 Petition. While the status of loggerheads has not changed under the ESA, an explicit spatial distinction between loggerheads nesting on the Northern and Southern parts of the US SE coast has been made. In the analysis that follows, we explore how genetic data can be used to support claims to sea turtles outside of national jurisdictions and in the ocean commons. In contrast to the cases above, genetic data were used to argue that all hawksbills in the Caribbean were genetically related, and thus should be managed as a single, regional conservation territory.

4. Geo-political genetics: hawksbill sea turtles in the Caribbean One of the more contentious of contemporary sea turtle conservation disputes has revolved around hawksbill sea turtles in the Caribbean. Hawksbills are listed as critically endangered by the IUCN (www.iucnredlist.org), and their depletion has been linked to trade in the scutes that make up hawksbill carapaces, or what is known as tortoiseshell. Trade has been greatly reduced under CITES, but until 1992, trade continued between Japan and Cuba. Hawksbill shell is used by the Japanese Bekko industry to fashion art and cultural artifacts, and Japan continued to trade by maintaining a reservation on hawksbill turtles under CITES. Following increased pressure from the US, Japan announced that it would stop importing shell in 1992 (Donnelly, 1991), and then removed its reservation under CITES in 1994 (Eckert and Eckert, 1994). Subsequently, Cuba sought to trade hawksbill shell legally with Japan through proposals submitted to CITES in 1997 (Prop. 10.60) and again in 1999 (Prop. 11.40, 11.41). These proposals would have transferred hawksbills from CITES Appendix I to II in order to allow for limited and highly regulated trade between Cuba and Japan only. Cuba presented biological studies of its hawksbill population (Carrillo et al., 1999; Moncada et al., 1999), and the proposals included justification of the management strategy, description of a shell registration system, and mechanisms for channeling some profits to local communities and for adapting the harvest in the face of evidence of population decline. Under an approved Prop 11.40, Cuba would continue to harvest 500 hawksbills annually, a voluntary reduction from the 2000 harvested annually that was phased in over 1991 to 1995. Prop 11.41 sought to trade the stockpiled shell in Cuba, with no new harvesting. The Cuban proposals failed to obtain two-thirds majority support in 1997 and again in 1999 (only Prop 11.41 was voted on as Cuba withdrew Prop 11.40). In 2002, Cuba submitted another CITES proposal (Prop. 12.30) to transfer stockpiled shell from Cuba to Japan, but withdrew the proposal three months before the CITES Conference of the Parties in Chile. In January 2008, Cuba announced that it would end its hawksbill harvest, with support from World Wildlife Fund – Canada and the Canadian International Development Agency. While a number of issues surfaced during the hawksbill debate, a key point of contention was the issue of ownership. Given the migratory nature of sea turtles, opponents of the harvest argued that Cuba did not have the ‘right’ to harvest these animals, since doing so impinged on the rights of other countries who share them, countries like Mexico, Bahamas, and Costa Rica that spoke against the Cuban proposals at CITES (Godfrey, 2000, 2002b). During the 1997 CITES meeting, the IUCN Marine Turtle Specialist Group produced a summary of the biology and status of hawksbills in the Caribbean, which concluded that, as a result of harvest of animals on a foraging ground, ‘‘different stocks of marine turtles from distant nesting grounds are likely to be affected” (IUCN/SSC/MTSG, 1997, p. 33). The Caribbean Conservation Corporation (CCC), a

USA-based NGO with a long-term field office in Costa Rica, was one of many NGOs that lobbied against the Cuban proposals with a focus on ownership. In the run up to the 2000 meeting, the CCC stated that hawksbill turtles in Cuban waters are highly migratory and therefore conservation and management actions must involve other stakeholders in the region (Godfrey, 2000, p. 8). In lobbying against Prop 12.3, the CCC ‘‘published and distributed a detailed analysis of the Cuban proposal, including evidence showing that hawksbills killed in Cuban waters migrate from throughout the Caribbean” (Godfrey, 2002b, p. 4). While resolving questions of ownership is not listed as a contribution of genetic analyses in three recent review papers (DeSalle and Amato, 2004; DeYoung and Honeycutt, 2005; Hedrick, 2001), this is the type of argument that genetic data and analyses can inform. Part of Cuba’s argument in favor of a hawksbill harvest was that the harvest targets mostly ‘Cuban’ turtles, i.e. turtles that both nest on Cuban beaches and forage in Cuban waters. This was not the only argument Cuba put forward, and CITES proposals and published research from Cuba acknowledged that some immigration and emigration of hawksbill turtles occurs between Cuban waters and other areas of the Caribbean (Carrillo et al., 1999; Diaz-Fernandez et al., 1999; Mrosovsky and Webb, 1997). Regardless, subsequent research papers examining the implications of the Cuban hawksbill harvest have implied that Cuba claimed to be harvesting and managing a closed population (Bowen et al., 1996; Bowen and Karl, 2007; Heppell and Crowder, 1996). Given the importance of ‘ownership’ in the Cuban hawksbill debate, it is perhaps not surprising that genetic arguments became increasingly important and explicit. General statements about shared ownership were replaced by more specific claims following a publication by Bass (1999). Using >360 genetic samples gathered from eight regional nesting beaches and two foraging sites, Bass (1999) compared genetic profiles of hawksbill turtles found at each, and generated estimates of the relative contribution of source populations (i.e. nesting beaches) to aggregations on foraging grounds, using mixed-stock analysis. Bass (1999) analysis suggested that 25–42% of turtles foraging in Cuban waters originate elsewhere, or that Cuban nesters supply 58–75% of their own foraging stock. In the materials produced describing their opposition to Prop 12.3, NGOs like the CCC and The Species Survival Network referred to genetic data showing Cuba’s turtles are shared. However, they suggested that 30–58% (sometimes rounded up to 60%) of hawksbills in Cuban waters originate from other nesting areas in the Caribbean (Godfrey, 2002a; Species Survival Network, 2002). This is either an exaggeration of Bass (1999) or a misreading, with the estimated minimum percentage of Cuban nesting beach contribution to Cuban foraging grounds (58%) confused with the maximum contributions from other regional nesting beaches to Cuban foraging grounds (42% according to Bass, 1999).5 Regardless, post-1999, opponents of the Cuban harvest were more specific about the shared ownership of Cuban hawksbills, and genetic data fueled their opposition. In 2007, Bowen et al. (2007a) published an updated review of hawksbill genetic studies conducted in the Caribbean, supplemented with new data from their own research. Bowen et al. (2007a) use samples from eight nesting beaches (Cuba, Puerto Rico, US Virgin Islands, Antigua, Barbados, Costa Rica, Belize, and Mexico) and eight foraging grounds (Texas, Bahamas, Cuba [three sites: a, b, d], Dominican Republic, Puerto Rico, and US Virgin Islands) and a total of 962 genetic samples (Table 1; Fig. 1). Based on the results of their mixed-stock analysis, Bowen et al. (2007a, p. 58) claim that:

5 It is possible that the 58% was generated by some other studies, but we have not been able to locate these in the primary published literature.

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Table 1 Percentage contributions of nesting beaches to foraging grounds of hawksbill sea turtles in the Caribbean, using maximum likelihood analysis,a adapted from Bowen et al. (2007a). Column headings refer to the foraging grounds where hawksbill turtles have been genetically sampled in-water and the row headings refer to nesting beaches. Reading down a column, we see that the origin of hawksbill turtles foraging in Bahamian waters is 37% from Cuban nesting beaches, 4% from Puerto Rico, 25% from USVI, 1% from Antigua, 3% from Costa Rica, and 28% from Mexico. In contrast, hawksbills foraging in Texan waters originate from USVI (7%) and Mexico (93%). Foraging Grounds:

Texas

Bahamas

Cuba a

Cuba b

Cuba d

Dominican Republic

Puerto Rico

USVI

Nesting Beaches Cuba Puerto Rico USVI Antigua Barbados Costa Rica Belize Mexico Unknown

0.00 0.00 6.64 0.00 0.00 0.05 0.60 92.70 0.00

36.90 3.96 25.39 1.19 0.00 2.91 0.00 28.37 1.28

72.52 0.01 6.59 0.00 0.04 11.69 0.00 6.83 2.33

45.81 15.69 25.18 0.15 0.00 2.39 0.00 9.88 0.90

43.18 10.74 7.87 2.72 0.00 19.19 0.00 14.52 1.79

44.71 1.82 19.99 0.01 0.14 24.45 0.00 14.52 2.22

23.45 11.06 39.54 1.03 0.00 11.17 0.00 13.04 0.72

36.98 14.66 15.31 7.18 0.05 10.19 0.00 12.69 2.94

a In Bowen et al. (2007a), the authors also present Bayesian estimates and standard deviations for both types of analysis. The results of the mixed-stock and Bayesian analyses were highly correlated, and while there are subtle differences, they do not impact on our analysis. We use only the mixed-stock analysis results in this paper.

Fig. 1. Nesting and foraging sites providing genetic samples. Nesting sites are located in Cuba (CU), Puerto Rico (PR), US Virgin Islands (USVI) Antigua (AN), Barbados (BD), Costa Rica (CR), Belize (BE), and Mexico (MX). Foraging sites are located in Texas (TX), Bahamas (BA), Cuba (CU, three sites: a, b, d), Dominican Republic (DR), Puerto Rico (PR), and US Virgin Islands (USVI).

Harvest in the Caribbean foraging areas will deplete nesting populations across multiple jurisdictions. . . Therefore, hawksbill conservation embodies a conflict between national and international interests. International cooperation is the last and best hope. Citing Bowen et al. (2007a), Mortimer et al. (2007, p. 18) echo these claims: Harvests in any part of the Caribbean impact the species throughout the region. . . . harvest of hawksbills from any nesting beach could impact multiple foreign feeding grounds, and that harvest on any feeding grounds could impact the nesting populations at multiple sites. Thus, any nation that opens or promotes harvest could be undermining badly needed efforts to conserve the species at other sites. As the above quotes illustrate, Bowen et al. (2007a) and Mortimer et al. (2007) use the understanding gained via genetic analyses to support claims that hawksbill sea turtles are regionally shared and should be managed as such. Further, they have specific views of what such management should entail; harvests will ‘deplete’ or ‘impact’ on regional populations and will undermine conservation. International cooperation (presumably to eliminate harvesting) is needed. In the following section, we use the results of Bowen et al.’s (2007a) mixed-stock analysis to interrogate the veracity of claims

regarding the regional nature of hawksbills in the Caribbean. Making a direct link between the early genetics work of Bass (1999) on decisions by CITES delegates or between Bowen et al. (2007a,b) and Cuba’s ultimate decision to close its fishery in 2008 is beyond the scope of this paper.6 What we have shown is that questions of ownership were critical to opponents of Cuba’s proposals and that the dominant interpretation of the genetic data was that they support regional scale management. Here, we turn to potential alternative claims that could be made based on the same data set. Most of these claims are hypothetical, but those that have circulated in the published literature are noted as such. We then complicate these claims, by considering other issues that need to be taken into account when thinking about conservation in general, and of sea turtles specifically. We do not question the genetic data or the mixed-stock analysis itself. Although the limitations of mixed-stock analysis are recognized (e.g. Waples et al., 2008), we accept its utility for describing the origins of hawksbill turtles found on foraging grounds in the Caribbean.

6 There were undoubtedly many issues at play in this decision, and at CITES this included the amount of attention the hawksbill issue received in the press. In its report on outcomes of CoP-11, CITES notes that ‘‘Proposals which had attracted more attention from the participants and the press, such as . . . 2 proposals of hawksbill turtles . . . were either withdrawn or rejected.” http://www.cites.org/eng/cop/11/ outcome.shtml.

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Fig. 2. Contributions from regional nesting beaches to the foraging stock at Cuba site a. Arrows represent flow from nesting beaches to foraging grounds. Only countries contributing more than five percent to Cuba a’s foraging stock are shown.

4.1. Analyzing resource claims 4.1.1. Cuba harvests its own turtles Of the eight foraging grounds considered by Bowen et al. (2007a), three are in Cuba (sites a, b, d; Fig. 1). Of all eight sites sampled, Cuba site a has the highest percentage of turtles originating from Cuban nesting beaches (over 70 percent; Table 1, Fig. 2). Table 1 shows that Cuba sites b and d are also ‘less mixed’ than most other sites in the region, although over 50 percent of turtles foraging at b and d originate from non-Cuban nesting beaches. These figures could be used by Cuba to support claims to continue harvesting at site a; more than 70 percent of the time, Cuban fishers are likely to be harvesting turtles coming from Cuban nesting beaches.7 4.1.2. Cuba’s harvest decreases foraging populations throughout the region Hawksbill turtles from Cuba’s nesting beaches contribute greatly to regional foraging stocks; they are the largest contributor to stocks at all sites sampled, except for Puerto Rico and Texas (Table 1, Fig. 3). On this basis, Bahamas, Dominican Republic, Puerto Rico, and US Virgin Islands (USVI) could claim that, even if Cuba harvests mostly its own turtles at some sites, the harvest reduces the number of turtles in the population available to migrate to regional foraging areas. The argument might vary depending on the assumptions in place about what promotes migration of Cuban foragers, for example, whether or not this migration is random or driven by density dependence. If density dependence is in place, i.e. if emigration from Cuban waters is driven by high population density there, then harvesting that reduces such density might decrease migration to other foraging grounds in the region. If dispersal is random, the argument is less valid. 4.1.3. Fisheries in USVI or Puerto Rico impact on turtle nesting everywhere Of all foraging aggregations, USVI and Puerto Rico stocks are the most ‘mixed’, with five nesting populations contributing more than ten percent each to their respective foraging stocks (Fig. 4). If USVI or Puerto Rico were to fish turtles, their actions could be contested as drawing on the conservation investments of several different 7 This is a hypothetical scenario. When Cuba reduced its harvest to 500 turtles from 1991 to 1995, it focused its fishing efforts on site b.

countries, and undermining those countries’ claims to the resources. Since there are no legal fisheries in USVI or Puerto Rico, such claims have not been articulated. However, there is a small sea turtle fishery in British Virgin Islands (BVI), immediately adjacent to USVI (Campbell et al., 2009). Assuming that genetic diversity sampled in USVI does not differ significantly from that in BVI, opponents of BVI’s harvest could claim the harvest relies on and impacts nesting populations in five other countries. In contrast to the Cuban case, BVI has few nesting turtles of its own and can be portrayed as capitalizing on conservation investments made at nesting beaches elsewhere. 4.1.4. Reduced harvesting in Cuba has led to increased nesting in Barbados Beggs et al. (2007) describe increased nesting in Barbados based on data gathered from 1997 to 2004. They suggest that the reduction of Cuba’s harvest over 1991–1995, from 2000 to 500 turtles, helps to explain increased nesting seen in Barbados. While the authors are aware of Bowen et al.’s (2007a) work, and cite it elsewhere in their paper, Bowen et al.’s analysis does not support this claim. The mixed-stock analysis of Cuba’s foraging grounds finds minimal evidence of migration from Barbadian beaches to Cuban waters (i.e. Barbados nesters contribute 0.04 percent to the foraging stock at Cuba site a, and zero percent to the other two sites); thus, based on the existing genetic information, the reduced Cuban harvest is unlikely to impact nesting numbers in Barbados. 4.2. Complicating genetically based claims The claims outlined above, both hypothetical and real, are straightforward, based on percentages associated with the mixed-stock analysis. The meaning of the percentages, though, is open to interpretation, as we suggest above. The meaning becomes even more complicated when other factors are taken into account, for example, nesting trends in source populations, genetic diversity, population dynamics, and conservation investments. 4.2.1. Claims in light of nesting trends While hawksbill populations are considered greatly reduced it the Caribbean from an historical standpoint, there is evidence of contemporary increases based on nesting trends on some of the region’s beaches. Of the nesting beaches (source populations) sampled in Bowen et al. (2007a), annual nest numbers in Puerto Rico

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Fig. 3. Contributions from Cuban nesting beaches to foraging populations in other countries. With the exceptions of the Texas foraging site, where no Cuban contributions are detected, and the Puerto Rico foraging site, where USVI is the highest contributor, Cuban beaches are the most important source population for the foraging sites sampled, including the three foraging sites within Cuban waters.

Fig. 4. Contributions from regional nesting beaches to the USVI foraging population, the most ‘mixed’ of the eight foraging stocks sampled.

(Van Dam et al., 2008), Antigua (Richardson et al., 2006), and Barbados (Beggs et al., 2007) are increasing. Nest trends in Mexico are fluctuating (increasing according to Garduño-Andrade et al. (1999); decreasing according to Abreu-Grobois et al. (2005)), and decreasing in Costa Rica (Troëng et al., 2005). Nest trends in Cuba and Belize are unknown. Overlaid with genetic data, nesting trends complicate resource claims. For example, while Puerto Rico contributes to all three of Cuba’s foraging stocks, its own nesting population is increasing (Van Dam et al., 2008). While recently documented increases may be associated with the decrease in the Cuban harvest in 1995, the post-1995 harvest does not appear to be impairing nesting recovery in many places. In contrast, even though Costa Rica contributes only 12 percent of Cuba site a’s mixed stock, decreased nesting in Costa Rica could support arguments that any take of these turtles, however low the percentage, impacts negatively on Costa Rica’s nesting populations. In the latter case, a major decline appears to have occurred 50 years ago, with low level nesting since then (Troëng et al., 2005).

4.2.2. Claims in light of genetic diversity Nesting populations are genetically distinctive; thus their utility for defining management units. In the nesting populations considered in Bowen et al. (2007a), most are dominated by a single haplotype, with two or three others present in very small numbers. For example, of 70 samples from Cuba’s nesting populations, 62 are haplotype A, 5 are haplotype c, and there are one each of haplotype F, Cu3, Cu4. In contrast, Puerto Rico has greater diversity (of 35 samples, 13 are haplotype F, 12 are N, there are 2 each of J, L, M, O and one each of A, and K). Some haplotypes (J, K, L, M and O) are rare in the foraging populations throughout the region, so one could claim that regional foraging grounds where these haplotypes are found need to be better protected. Bowen et al. (2007a) argue that small contributions are relevant to conservation, an argument made more generally in conservation genetics; the loss of unique haplotypes can combine with declining trends in population sizes to reduce genetic variation within a population, and this in turn may lead to a decreased ability of a population to adapt

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in response to ecological challenges (DeSalle and Amato, 2004). However, conservation biologists have debated the relative importance of population dynamics versus genetic diversity; for most species, the processes at stake in genetic extinction are likely to be less important than more immediate population pressures. 4.2.3. Claims in light of population dynamics While population dynamics of sea turtles are not fully understood, Godfrey et al. (2007) challenge Bowen et al.’s (2007a) conclusion that genetic data show harvesting in one country will be detrimental to conservation in another. Godfrey et al. (2007) base their argument on a fundamental principle of wildlife management, namely that target populations can be harvested or culled at some level without causing a decline in the overall population, due to density-dependent growth rates (Getz and Haight, 1989). While the question of what levels of harvesting can be withstood without harm to the overall population remain, Godfrey et al. (2007) point to some evidence of density dependence in sea turtles (e.g. Bjorndal et al., 2000), and suggest that continued nesting increases in the Caribbean indicate current hawksbill harvests are likely below some maximum threshold past which they will no longer be sustainable. ‘‘Although the new phylogenetic data provided by Bowen et al. (2007a,b) illuminate hawksbill migratory behavior, they do not specifically inform us about sustainable levels of mortality of sea turtle populations” (Godfrey et al. 2007, p. 3511). In response, Bowen et al. (2007b) argue that they do not make any claims about sustainable mortality in their paper, and clarify their key assumption: any turtle taken in a harvest is not available to migrate to other foraging areas in the region. 4.2.4. Claims in light of conservation investment Reflecting the traditional focus of biological studies on nesting females, conservation efforts have also focused on nesting beaches and sometimes on adjacent waters.8 While there have long been debates about how best to protect sea turtles (Frazer, 1992; Mrosovsky, 1983), population modeling is useful in ranking the relative importance of different life stages to the status of a sea turtle population (Crouse et al., 1987). In particular, elasticity analyses suggest that management actions aimed at reducing mortality of large juveniles and adult turtles will have the greatest positive impact on sea turtle populations (Crowder et al., 1994). While modeling work has not curtailed efforts to eliminate mortality at other life stages, it does suggest that some conservation efforts are ‘worth more’ in terms of species productivity and conservation; protecting adult females on and around nesting beaches appears to be a good conservation investment. However, nesting beach protection is economically costly, involving both direct costs (land purchase, protection, and monitoring) and the opportunity costs associated with removing coastal land and its resources from development. Thus, investments by countries like Costa Rica, where the nesting beach at Tortuguero has been protected since the 1970s (site CR in Fig. 1) could be used to ‘weight’ resource ownership claims. 5. Discussion and conclusions Opponents of Cuba’s proposals to CITES invoked ‘shared ownership’ of Caribbean hawksbills as a central part of their argument, and based on the most recent analysis, Bowen et al. (2007a) and Mortimer et al. (2007) make the case for broadly based regional or international collaboration, given the stock mixing evident in the Caribbean. While we agree that Bowen et al.’s (2007a) analysis 8 Since the 1990s, there have been significant resources devoted to reducing the indirect negative impacts of fisheries on sea turtles (e.g. the incidental capture of sea turtles when fishing for other things).

shows foraging stocks are often mixed, in some cases there are few countries involved, and smaller scale ‘regionalisms’ are likely at stake. For example, genetic data show that only nesting beaches in USVI, Costa Rica, and Mexico contribute more than five percent of the foraging population at site a in Cuba (Fig. 2). These same three countries contribute 15–25 percent of the foraging populations at Cuba sites b and d (Table 1). In theory, a compromise agreement to eliminate harvesting at sites b and d while continuing a harvest at site a could address concerns about the impacts of Cuban harvesting on nesting populations in USVI, Costa Rica, and Mexico. Whether or not this would be politically possible is another matter. The point is that the genetic data do not imply the whole region need be involved in this discussion, at least not from the standpoint of genetics (there may be other reasons for wider collaboration). Regardless of what ‘regionalisms’ are involved, here we turn to what is achieved through this particular science of sea turtles in terms of rights to resources and the scale at which conservation takes place. In this case, rights to resources are intimately tied to the issue of scale; if sea turtles can be shown to be shared among states, then traditional rights of those sovereign states to manage sea turtles independently when they are on nesting beaches or in territorial waters are more easily challenged. The focus of sea turtle conservation becomes international, and the conservation territory proposed in this case is Caribbean wide. While the idea that sea turtles are a shared resource is not new, the science that illuminates the specifics of such sharing is. General claims about shared resources can be replaced by specific claims by specific countries, and given greater weight. Although a number of other issues were debated during the course of the Cuban hawksbill dispute, including overall numbers of nesting turtles in the region and historical declines, the questions of ownership and Cuba’s right to exploit a shared resource dominated, and the emerging new information on genetics was increasingly cited as a means to answer such questions. The ‘scaling-up’ of conservation, and the ecological rationale for it, has been explored and interrogated by others (e.g. Brosius and Russell, 2003; Zimmerer, 2000). Large scale, international conservation territories are not just about space; they have implications for governance. Here, we are interested in what happens among states, as international environmental regimes can both strengthen and weaken territorial sovereignty (Bulkeley, 2005). Bowen et al.’s (2007a) work is used to strengthen some state claims, like those of Costa Rica; data showing where turtles nesting and protected on Costa Rican beaches go can support Costa Rica’s desires to influence conservation in those places. Bowen et al.’s (2007a) work is also invoked to weaken claims by states like Cuba where, contrary to trends in international sea turtle conservation, the state has continued to harvest sea turtles and has petitioned for the right to trade sea turtle products internationally. What we have tried to show is that the genetic data themselves can support a variety of ownership claims; those that are strengthened and promoted are those that fall in line with goals of the international sea turtle conservation regime. The January 2008 announcement by Cuba that it would end its sea turtle harvest after years of considerable international pressure illustrates the reach of the global regime into sovereign state space (cf Bulkeley, 2005). Similar to the implications of genetics for resource ownership claims, the implications for conservation are also subject to interpretation. For example, an argument in favor of Cuba’s right to harvest hawksbill turtles at Cuba site a requires accepting that approximately 30 percent of the time, turtles captured there may not come from Cuba’s nesting colony. It implies something about diversity, as there is a chance that some of the more rare haplotypes, e.g. those contributed by Costa Rica’s source population, will be caught. It involves accepting that using turtles might be part of a

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strategy to conserve them. An interesting feature of this case is that Costa Rica, one of the more outspoken opponents of the Cuban proposals, is recognized as a regional leader in sea turtle conservation and has invested heavily in protection through parks and protected areas and eliminating consumptive use (Campbell, 2002b). Perhaps ironically, Cuba’s sea turtle management program, one based on use, has also been recognized as commendable by the organization that monitors trade in endangered species (Fleming, 2001). It is not that Cuba failed to manage sea turtles, but that it failed to manage them in a way that was compatible with a dominant view of how sea turtles should be conserved. Genetics alone cannot tell us whether or not particular outcomes or conservation strategies are acceptable nor whether or they are sustainable, and other issues come into play in such assessments. The exchange between Godfrey et al. (2007) and Bowen et al. (2007b) on the implications of Bowen et al. (2007a) for sea turtle harvesting was not an isolated event. Many of the authors have opposing views on the potential for sustainable use to contribute to conservation in general; several of them (or the organizations they work for) have taken opposing positions on this particular issue at CITES and in other forums, as active lobbyists. Views of use are in turn influenced by a number of factors (e.g. attitudes to uncertainty, understanding of economics, values attached to turtles, Campbell, 2000, 2002a), but most often debated as questions of science. And what of the science? In reference to biotechnology, Mannion (1993, p. 353) suggests that geographers need not ‘‘engross themselves with the underpinning science” in order to study its implications. In the case of conservation, the increasing role of genetics in directing policy requires just the opposite. It is only when we understand the details of the genetic analyses of hawksbill sea turtles in the Caribbean that we can evaluate the various interpretations of what these analyses imply for conservation territories in the region. In the case of genetics, the science is highly technical and expensive, with a resulting concentration of expertise at a few key labs in the USA and Europe and leaving little room for outside critique.9 While Campbell (2007a) has already discussed how scaling up conservation serves to by-pass communities living with, using, and conserving sea turtles (see also Campbell et al., 2002), these features of genetics further confound the possibility for any ‘vertical linkages’ (Berkes, 2006) that might help avoid scalar mismatches associated with managing highly migratory marine species. For example, how might we combine local and expert knowledge in meaningful ways? While local people often have knowledge of population trends or habitat use, and researchers value and use this knowledge (e.g. Campbell et al., 2009; Godley et al., 2004), it is hard to fathom how local people might contribute to understanding genetic identity of individual turtles and related implications for populations. Our point is that the science of genetics is implicated not only in delineating the physical space of conservation territories, but their governance structures. Science, scale, and rights are tightly coupled. There are some interesting issues arising from the interaction of science and scale in the Cuban hawksbill case that illustrate how biophysical scale is also sometimes constructed. First, genetics focuses on, and almost by default emphasizes, the importance of the individual turtle. Genetics tells us that any one turtle harvested on a foraging ground is potentially related to those nesting in several other countries; the opponents of Cuba’s harvest argue that such turtles are part of a regionally ‘shared’ population. Genetics also tells us that some of the turtles are potentially rare, possessing haplotypes found less frequently in the region; turtles must be pro9 When one of our colleagues critiqued a genetics paper on the basis of limited sample sizes, the paper’s author wrote to tell our colleague he should not be commenting, since he was not a geneticist. This is an excellent example of boundary work (Gieryn, 1995) among scientists themselves.

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tected based on individual uniqueness. This emphasis on the importance of individual turtles as both potentially unique and shared supports a strategy of protecting all individual turtles via a regional conservation territory. Contrary to views of a hawksbill ‘population’ that might be managed as such, the emphasis on the individual turtle eliminates the possibilities for consumptive use as a means of management. Second, this strategy bypasses the national scale, where the majority of conservation investments are made and actions taken. Notably absent from the Cuban hawksbill debate was an argument about how the mixed-stock analysis could be read as strengthening Cuba’s claims of territorial sovereignty. That 70% of turtles foraging at Cuba site a do originate from Cuba’s nesting beaches, and that its other sites are less mixed than other regional foraging grounds, could have been read as vindicating Cuba’s claims to be harvesting ‘mostly’ its own turtles. Rather, having established the ‘closed population’ straw man, opponents used genetic data to knock this down. The mixed-stock analysis was never invoked to describe a foraging ‘stock’ in any way that might inform management of it as such (for example, restricting harvesting to site a in order to reduce the likelihood of turtles originating from Costa Rica, USVI and Mexico being caught). Geneticists themselves warn against using the data in a certain ways: ‘‘It would be inappropriate, for example, to say that exactly 40% of the foraging animals at Puerto Rico originate in the US Virgin Islands. These maximum likelihood estimates give qualitative (rather than precise) estimates of the foraging ground composition” (Bass, 1999, p. 197). Having provided precise numbers (with estimates of error included), geneticists dismiss their precision and argue for a ‘broad’ reading of the results. The Cuba case illustrates that the question of socio-political scale is not just inseparable from biophysical scale, but from the politics thereof. While the science of genetics is somewhat inaccessible, the power in genetic arguments is compelling. Mortimer et al. (2007) pose the question ‘whose turtles are they anyway?’ but we suggest an equally important question is ‘who gets to decide?’ In this case, the genetic identification of individual turtles as linked to particular nesting beaches proved more compelling than other means of claiming resources, e.g. through territorial sovereignty, investments in conservation and management, geographic proximity, or history of use. These latter arguments were all invoked by Cuba in Prop 11.40, without success, against the ‘shared resource’ argument of NGOs, other countries, and scientists. Although we may come to better understand sea turtles and other species through genetics, any decision that this way of knowing is the best, only, or natural way, in essence replacing or at least out-weighing other ways, is ultimately a human and political decision. At a more fundamental level, genetics is set to redefine how we think about nature, working to produce a kind of ‘molecular nature.’ In this molecular scale focus, where the genetic make-up of individual animals is of interest and employed to influence broad scale geopolitics of international conservation, we see an illustration of the ways the construction of ‘the natural’ serves as a ‘socio-spatial’ disciplining mechanism (Bridge et al., 2003). As this paper suggests, defining the molecular as the scale as which we should know and manage nature has profound implications for how we envision rights to resources and the scale at which they should be managed.

Acknowledgements Jens Carlsson and Tom Schultz informed our understanding of conservation genetics and offered helpful literature suggestions. Maps were generated using MapTool on www.seaturtle.org/maptool. Noelle Boucquey, Myriah Cornwell, Gabe Cumming, Noella Gray, N. Mrosovsky, Jennifer Silver and Kelly Stewart provided feedback on various drafts of this paper. The manuscript was

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