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DEPARTMENT OF COMMERCE complete the required fields, and enter determination, we consider first or attach your comments. whether a group of organisms National Oceanic and Atmospheric • Mail: Submit written comments to constitutes a ‘‘species’’ under the ESA, Administration Chelsey Young, NMFS Office of then whether the status of the species Protected Resources (F/PR3), 1315 East qualifies it for listing as either 50 CFR Parts 223 West Highway, Silver Spring, MD threatened or endangered. Section 3 of [Docket No. 141219999–5133–01] 20910, USA. the ESA defines a ‘‘species’’ to include Instructions: You must submit ‘‘any subspecies of fish or wildlife or RIN 0648–XD681 comments by one of the above methods plants, and any distinct population to ensure that we receive, document, segment of any species of vertebrate fish Endangered and Threatened Wildlife and consider them. Comments sent by or wildlife which interbreeds when and Plants; Proposed Rule To List the any other method, to any other address mature.’’ On February 7, 1996, NMFS Tanzanian DPS of African Coelacanth or individual, or received after the end and the U.S. Fish and Wildlife Service as Threatened Under the Endangered of the comment period, may not be (USFWS; together, the Services) adopted Species Act considered. All comments received are a policy describing what constitutes a AGENCY: National Marine Fisheries a part of the public record and will distinct population segment (DPS) of a Service (NMFS), National Oceanic and generally be posted for public viewing taxonomic species (the DPS Policy; 61 Atmospheric Administration (NOAA), on http://www.regulations.gov without FR 4722). The DPS Policy identified two Commerce. change. All personal identifying elements that must be considered when identifying a DPS: (1) The discreteness ACTION: Proposed rule; 12-month information (e.g., name, address, etc.), of the population segment in relation to petition finding; request for comments. confidential business information, or otherwise sensitive information the remainder of the species (or SUMMARY: We, NMFS, have completed a submitted voluntarily by the sender will subspecies) to which it belongs; and (2) comprehensive status review under the be publicly accessible. We will accept the significance of the population Endangered Species Act (ESA) for the anonymous comments (enter ‘‘N/A’’ in segment to the remainder of the species African coelacanth (Latimeria the required fields if you wish to remain (or subspecies) to which it belongs. As chalumnae) in response to a petition to anonymous). stated in the DPS Policy, Congress list that species. We have determined You can obtain the petition, status expressed its expectation that the that, based on the best scientific and review report, the proposed rule, and Services would exercise authority with commercial data available, and after the list of references electronically on regard to DPSs sparingly and only when taking into account efforts being made our NMFS Web site at http://www.nmfs. the biological evidence indicates such to protect the species, L. chalumnae noaa.gov/pr/species/petition81.htm. action is warranted. Section 3 of the ESA defines an does not meet the definition of a FOR FURTHER INFORMATION CONTACT: endangered species as ‘‘any species threatened or endangered species when Chelsey Young, NMFS, Office of evaluated throughout all of its range. which is in danger of extinction Protected Resources (OPR), (301) 427– throughout all or a significant portion of However, we determined that the 8491 or Marta Nammack, NMFS, OPR, Tanzanian population of the taxon its range’’ and a threatened species as (301) 427–8469. one ‘‘which is likely to become an represents a significant portion of the SUPPLEMENTARY INFORMATION: taxon’s range, is threatened across that endangered species within the portion, and is a valid distinct Background foreseeable future throughout all or a population segment (DPS). Therefore, significant portion of its range.’’ We On July 15, 2013, we received a interpret an ‘‘endangered species’’ to be we propose to list the Tanzanian DPS of petition from WildEarth Guardians to L. chalumnae as a threatened species one that is presently in danger of list 81 marine species as threatened or extinction. A ‘‘threatened species,’’ on under the ESA. We are not proposing to endangered under the Endangered the other hand, is not presently in designate critical habitat for this DPS Species Act (ESA). This petition danger of extinction, but is likely to because the geographical areas occupied included species from many different become so in the foreseeable future (that by the population are entirely outside taxonomic groups, and we prepared our is, at a later time). In other words, the U.S. jurisdiction, and we have not 90-day findings in batches by taxonomic primary statutory difference between a identified any unoccupied areas that are group. We found that the petitioned threatened and endangered species is essential to the conservation of the DPS. actions may be warranted for 27 of the the timing of when a species may be in We are soliciting comments on our 81 species and announced the initiation danger of extinction, either presently proposal to list the Tanzanian DPS of of status reviews for each of the 27 (endangered) or in the foreseeable future the coelacanth as threatened under the species (78 FR 63941, October 25, 2013; (threatened). ESA. 78 FR 66675, November 6, 2013; 78 FR When we consider whether species DATES: Comments on our proposed rule 69376, November 19, 2013; 79 FR 9880, might qualify as threatened under the to list the coelacanth must be received February 21, 2014; and 79 FR 10104, ESA, we must consider the meaning of by May 4, 2015. Public hearing requests February 24, 2014). This document the term ‘‘foreseeable future.’’ It is must be made by April 17, 2015. addresses the findings for one of those appropriate to interpret ‘‘foreseeable ADDRESSES: You may submit comments 27 species: The African coelacanth L. future’’ as the horizon over which on this document, identified by NOAA– chalumnae. Findings for seven predictions about the conservation NMFS–2015–0024, by either of the additional species can be found at 79 FR status of the species can be reasonably following methods: 74853 (December 16, 2014). The relied upon. The foreseeable future • Electronic Submissions: Submit all remaining 19 species will be addressed considers the life history of the species, electronic public comments via the in subsequent findings. habitat characteristics, availability of Federal eRulemaking Portal. Go to We are responsible for determining data, particular threats, ability to predict www.regulations.gov/ whether species are threatened or threats, and the reliability to forecast the #!docketDetail;D=NOAA-NMFS-2015- endangered under the ESA (16 U.S.C. effects of these threats and future events 0024. Click the ‘‘Comment Now’’ icon, 1531 et seq.). To make this on the status of the species under

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consideration. Because a species may be also required to make listing not warranted or contribute to forming susceptible to a variety of threats for determinations based solely on the best the basis for listing a species as which different data are available, or scientific and commercial data threatened rather than endangered. The which operate across different time available, after conducting a review of two basic criteria established by the scales, the foreseeable future is not the species’ status and after taking into PECE are: (1) The certainty that the necessarily reducible to a particular account efforts being made by any state conservation efforts will be number of years. Thus, in our or foreign nation to protect the species implemented; and (2) the certainty that determinations, we may describe the (16 U.S.C. 1533(a)(1)). the efforts will be effective. We consider foreseeable future in general or In making a listing determination, we these criteria, as applicable, below. We qualitative terms. first determine whether a petitioned re-assess the extinction risk of the NMFS and the USFWS recently species meets the ESA definition of a species in light of the existing published a policy to clarify the ‘‘species.’’ Next, using the best available conservation efforts. interpretation of the phrase ‘‘significant information gathered during the status If we determine that a species portion of the range’’ (SPR) in the ESA review for the species, we complete a warrants listing as threatened or definitions of ‘‘threatened’’ and status and extinction risk assessment endangered, we publish a proposed rule ‘‘endangered’’ (76 FR 37577; July 01, across the range of the species. In in the Federal Register and seek public 2014). The policy consists of the assessing extinction risk, we consider comment on the proposed listing. following four components: the demographic viability factors Status Review (1) If a species is found to be developed by McElhany et al. (2000) endangered or threatened in only an and the risk matrix approach developed We conducted a status review for the SPR, the entire species is listed as by Wainwright and Kope (1999) to petitioned species addressed in this endangered or threatened, respectively, organize and summarize extinction risk finding (Whittaker, 2014), which and the ESA’s protections apply across considerations. The approach of compiled information on the species’ the species’ entire range. considering demographic risk factors to biology, ecology, life history, threats, (2) A portion of the range of a species help frame the consideration of and conservation status from is ‘‘significant’’ if its contribution to the extinction risk has been used in many information contained in the petition, viability of the species is so important of our status reviews, including for our files, a comprehensive literature that without that portion, the species Pacific salmonids, Pacific hake, walleye search, and consultation with experts. would be in danger of extinction or pollock, Pacific cod, Puget Sound We also considered information likely to become so in the foreseeable rockfishes, Pacific herring, scalloped submitted by the public in response to future. hammerhead sharks, and black abalone our petition finding. The draft status (3) The range of a species is (see http://www.nmfs.noaa.gov/pr/ review report was also submitted to considered to be the general species/ for links to these reviews). In independent peer reviewers; comments geographical area within which that this approach, the collective condition and information received from peer species can be found at the time USFWS of individual populations is considered reviewers were addressed and or NMFS makes any particular status at the species level according to four incorporated as appropriate before determination. This range includes demographic viability factors: finalizing the draft report. those areas used throughout all or part Abundance, growth rate/productivity, The status review report provides a of the species’ life cycle, even if they are spatial structure/connectivity, and thorough discussion of demographic not used regularly (e.g., seasonal diversity. These viability factors reflect risks and threats to the particular habitats). Lost historical range is concepts that are well-founded in species. We considered all identified relevant to the analysis of the status of conservation biology and that threats, both individually and the species, but it cannot constitute an individually and collectively provide cumulatively, to determine whether the SPR. strong indicators of extinction risk. species should reasonably be expected (4) If a species is not endangered or We then assess efforts being made to to respond to the threats in a way that threatened throughout all of its range protect the species, to determine if these causes actual impacts at the species but is endangered or threatened within conservation efforts are adequate to level. The collective condition of an SPR, and the population in that mitigate the existing threats. Section individual populations was also significant portion is a valid DPS, we 4(b)(1)(A) of the ESA requires the considered at the species level, will list the DPS rather than the entire Secretary, when making a listing according to the four demographic taxonomic species or subspecies. determination for a species, to take into viability factors discussed above. We considered this policy in consideration those efforts, if any, being The status review report is available evaluating whether to list the coelacanth made by any State or foreign nation to on our Web site (see ADDRESSES as endangered or threatened under the protect the species. We also evaluate section). The following section ESA. conservation efforts that have not yet describes our analysis of the status of Section 4(a)(1) of the ESA requires us been fully implemented or shown to be the African coelacanth, L. chalumnae. to determine whether any species is effective using the criteria outlined in endangered or threatened due to any the joint NMFS/USFWS Policy for Species Description one or a combination of the following Evaluating Conservation Efforts (PECE; Latimeria chalumnae, a fish five threat factors: The present or 68 FR 15100, March 28, 2003), to commonly known as the African threatened destruction, modification, or determine their certainty of coelacanth, belongs to a very old lineage curtailment of its habitat or range; implementation and effectiveness. The of bony fish, the class Sarcopterygii or overutilization for commercial, PECE is designed to ensure consistent lobe-finned fishes, which includes the recreational, scientific, or educational and adequate evaluation of whether any coelacanths, the lungfish, and very early purposes; disease or predation; the conservation efforts that have been tetrapods. Most species of lobe-finned inadequacy of existing regulatory recently adopted or implemented, but fish are extinct. Among the lobe-finned mechanisms; or other natural or not yet demonstrated to be effective, fishes, L. chalumnae is one of only two manmade factors affecting its continued will result in improving the status of the living species belonging to the order existence (16 U.S.C. 1533(a)(1)). We are species to the point at which listing is Coelacanthiformes. The belief that the

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coelacanth had gone extinct over 65 1993; Hissmann et al., 1998). However, Habitat off of Tanzania consists of million years ago made the discovery of between 1995 and 2001, catches and rocky terraces occurring between 70– a living specimen off the coast of South observations of coelacanths from the 140 m depth; the water temperature at Africa in 1938 particularly sensational coasts of Kenya (De Vos et al., 2002), coelacanth catch depths is around 20 °C (McAllister, 1971). Latimeria Tanzania (Benno et al., 2006), South (Nyandwi, 2009). A large number (n = chalumnae inhabits coasts along the Africa (Hissmann et al., 2006), and 19) of Tanzanian coelacanths have been western Indian Ocean, while Latimeria Madagascar (Heemstra et al., 1996) caught in the outer reefs near the village menadoensis, commonly known as the suggested that the species was more of Tanga. In this region, some Indonesian coelacanth, observed for the widespread than previously thought, coelacanth catches have been reported first time in 1997, appears to be occupying deep water coastal habitat in to occur at 50–60 m; however, the restricted to Indonesian waters, but several locations throughout the validity of these reports is questionable might also occur along the coastal Western Indian Ocean. The range extent (Benno et al., 2006; Nyandwi, 2009, islands in the eastern Indian Ocean of the coelacanth remains unclear, as Hissman pers. com.). These incidents (Erdmann et al., 1998; Erdmann, 1999; direct observations of established may indicate a shallower depth Springer, 1999; Fricke et al., 2000b, populations rely on dedicated deep preference for Tanzanian coelacanths Hissman pers. com.). Latimeria water canyon surveys, or bycatch than that exhibited by Comoran chalumnae and L. menadoensis are observations from gillnets and artisanal coelacanths; however, more surveys are genetically and geographically distinct handlines (Hissmann et al., 2006). needed to better understand coelacanth (Pouyaud et al., 1999; Holder et al., Today, three established coelacanth habitat use in this region (Benno et al., 1999; Inoue, 2005). While genetically populations have been confirmed by 2006). The benthic substrate off the distinct, the Indonesian and African survey efforts, inhabiting deep-water coast of Tanzania is sedimentary coelacanth species exhibit overlapping caves off the coast of the Comoros, limestone rather than the volcanic rock morphological traits, which makes it South Africa, and the coast of Tanzania. of the Comoros. In this habitat, difficult to differentiate between them The coelacanth is known to inhabit coelacanths are thought to use based on morphology alone. waters deeper than 100m, making submarine cavities and shelves that The coelacanth has a number of surveys difficult and reliant upon have eroded out of the limestone unique morphological features. Most sophisticated technology including composite for shelter. obvious are its stalked dorsal, pelvic, submersibles and remotely operated Coelacanths demonstrate strong site anal, and caudal fins. In the water, vehicles (ROVs), or highly-trained fidelity with relatively large overlapping under camera observation, the body of divers using special gas mixtures. To home ranges, greater than 8 km, as the fish appears iridescent dark blue, date, the best data addressing demonstrated at Comoro and South but its natural color is brown (Hissman coelacanth habitat use come from in situ African sites where expeditions have pers. com.); individuals have white observations of the fish off the steep tracked individual movements using blotches on their bodies that have been volcanic coasts of Grand Comoro Island; ultrasonic transmitters (Fricke et al., used for identification in the field. two decades of coelacanth observation 1994; Heemstra et al., 2006). Surveys off When individuals die, their color shifts there demonstrate that the coelacanth Grand Comoro over 21 years from blue to brown. The name inhabits deep submarine caves and demonstrate that individual coelacanths ‘‘coelacanth’’ comes from the Greek canyons which are thought to provide may inhabit the same network of caves words for ‘hollow’ and ‘spine,’ referring shelter from predation and ocean for decades; for example, 17 individuals to the fish’s hollow oil-filled notochord, currents (Fricke et al., 2011). The fish originally identified in 1989 were re- which supports the dorsal and ventral aggregate in these caves in groups of up sighted in 2008 in the same survey area caudal fin rays (Balon et al., 1988). This to 10 individuals. Retreat into these (Fricke et al., 2011). notochord is composed of collagen caves after nightly feeding activity is Temperature use for the Comoran which is stiffened under fluid pressure most likely a key factor for coelacanth coelacanth, based on survey (Balon et al., 1988). Coelacanth species survival, allowing the fish to rest and observations, was found to be between have a unique intracranial joint conserve energy in a deep-water, low- 16.5 and 22.8 °C (Fricke et al., 1991b). allowing them to simultaneously open prey environment (Fricke et al., 1991a). Surveys of South African coelacanth the lower and upper jaws, possibly an At night, coelacanths occupy deeper habitat off of Sodwana Bay confirm this adaptation for feeding (Balon et al., waters to actively feed, spending the temperature use across a broad portion 1988). Coelacanths undergo majority of their time between 200 and of its range (Hissmann et al., 2006). This osmoregulation via retention of urea 300 m (Fricke et al., 1994; Hissmann et corresponds to estimates of thermal (Griffith, 1991). Their swim bladder is al., 2000). Larger individuals are known requirements based on the temperature- filled with wax-esters used to passively to venture below 400 m, with the dependent oxygen saturation of their regulate buoyancy, allowing the fish to deepest observation at 698 m (Hissmann blood, with an optimum at 15 °C and an reach depths of 700 meters during et al., 2000). upper threshold at 22–23 °C (Hughes et nightly feeding excursions (Hissmann et South African coelacanth habitat has al., 1972). Thus, the coelacanth depends al., 2000). Males and females exhibit also been studied, although to a lesser on cooler waters to help maintain its sexual dimorphism in size, with females extent than in the Comoro Islands oxygen demands. Most likely, the depth larger than males (Bruton et al., 1991b). (Venter et al., 2000; Hissmann et al., distribution of coelacanth depends The natural range of the African 2006; Roberts et al., 2006). In the deep partly on this temperature requirement. coelacanth L. chalumnae was once canyons off the coast of South Africa, The coelacanth’s ecological niche is thought to be restricted to the Comoro suitable coelacanth caves have been likely shaped by this narrow Island Archipelago, located in the found at depths of 100–130 m, whereas temperature requirement, prey Western Indian Ocean between at Grand Comoro Island, most caves are abundance, and the need for shelter and Madagascar and Mozambique. For many in depths of 180–230 m (Heemstra et al., oxygen. years, specimens caught off South 2006). In general, it is thought that the It is thought that sedimentation and Africa, Mozambique, and Madagascar deep overhangs and caves found off the siltation act as a negative influence on were thought to be strays from the shelf of South Africa provide suitable coelacanth distribution. This is Comoro population (Schliewen et al., shelter and refuge for coelacanths. supported by a hypothesis surrounding

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the split between the two living thought to heavily limit their growth Fricke et al. (1991a) and updated to coelacanth species estimated to have rate and productivity due to its control include survey data from 1991 (Fricke et occurred 40–30 million years ago (Mya), over oxygen utilization (Froese et al., al., 1994). The survey area during this corresponding with the collision 2000). Studies of the fish’s blood time covered 9 percent of the projected between India and Eurasia (50 Mya), physiology have demonstrated that the coelacanth habitat along the western which created high levels of siltation oxygen dissociation curve is coast of Grand Comoro (Hissmann et al., and isolated individuals to the east and temperature dependent, and shows an 1998). These estimates showed a west of India (Inoue et al., 2005). This affinity for oxygen at lower relatively stable population ranging hypothesis has been supported by some temperatures (15 °C). Small gill surface between 230–650 individuals (Fricke et surveys off Sodwana Bay where it was area and blood physiology are thought al., 1994). Surveys conducted in 1994 observed that some canyons, despite to influence the coelacanth’s restriction across the southwestern coast of Grand offering suitable habitat requirements, to cold deep water habitat, and may Comoro (the same sample area as in were not occupied by coelacanths; it correlate with their low metabolic rates, earlier surveys) revealed a 68 percent was concluded that the turbidity of the meager food consumption and generally decrease in cave inhabitants and a 32 water in these caves discouraged slow growth and maturation (Froese et percent decrease in the total number of coelacanth habitation, as nearby al., 2000). coelacanths encountered as compared to canyons not affected by turbidity were Population Abundance, Distribution, a 1991 survey that covered the same occupied by coelacanths (Hissmann et and Structure area at the same time of year (Hissmann al., 2006; Roberts et al., 2006). et al., 1998). Three additional surveys of Coelacanths are considered It was once thought that coelacanths the western coast of Grand Comoro ovoviviparous, meaning the embryos are were restricted to the Comoro Island occurred in the 2000s, and are provided a yolk sac and develop inside Archipelago, and that individuals summarized in Fricke et al. (2011). the adult female until they are delivered caught in other locations in the Western These survey methods and area were as live births; coelacanth embryos are Indian Ocean were strays. However, consistent with earlier surveys not surrounded by a solid shell. growing evidence suggests that L. occurring in the late 1980s and 1990s. Embryos remain in gestation for 3 years; chalumnae consists of several During surveys between 2000 and 2009, this period of embryogenesis has been established populations throughout the several marked individuals not sighted determined by scale rings of embryo and Western Indian Ocean (Schartl et al., in 1994 re-appeared, and cave newborn coelacanth specimens (Froese 2005). Two resident and scientifically occupancy rates in these later surveys surveyed coelacanth populations exist et al., 2000). The coelacanth gestation were similar to surveys of the early in waters off South Africa and the period is considered the longest of any 1990s (Fricke et al., 2011). In total, nine Comoro Islands (Hissmann et al., 2006; vertebrate (Froese et al., 2000). It has dedicated coelacanth surveys have Fricke et al., 2011). Increases in been hypothesized that the coelacanth occurred in this area since 1986 (Fricke coelacanth catch off the coast of may live upwards of 40 or 50 years, and et al., 2011). Estimates of population Tanzania during the last decade and even up to 100 years (Bruton et al., abundance along the western coast of genetic analysis of individuals caught 1991a, Fricke et al., 2011, Hissman per. Grand Comoro, based on repeated there demonstrated that an established com.). Coelacanth generation times are surveys over almost 2 decades, are population exists there as well, as long. In fact, they are expected to reach between 300 and 400 individuals, with confirmed by the observance of 9 reproductive maturity between 16 and 145 individuals identifiable via unique 19 years of age (Froese et al., 2000). coelacanth individuals during a 2007 markings (Fricke et al., 2011). The 1994 Coelacanth fecundity is not well known; survey off the Tanzanian coast (Nikaido survey showing population declines is 26 embryos were found within one et al., 2011). Additional coelacanth thought to be an anomaly driven by female caught in 2001 from off of catches have been recorded off higher water temperature, as later Mozambique, and other known Madagascar, Mozambique, and Kenya, surveys demonstrate that the local fecundities are 5, 19, and 23 pups but these regions have not yet been population of western Grand Comoro (Fricke et al., 1992). surveyed (Nulens et al., 2011) so their Coelacanths are extremely slow drift- status is unclear. What is known of the has remained stable since the 1980s hunters. They descend at least 50 to 100 coelacanth’s distribution is largely (Fricke et al., 2011). Some local m below their daytime habitat to feed at based on bycatch data. Thus, the true Comoran fishermen have suggested that night on the bottom or near-bottom, and number of established coelacanth seasonal abundance patterns may exist are thought to consume deep-water populations, and the extent of the for the coelacanth as they do for the prey, or prey found at the bottom of the species’ range across the Western Indian locally-targeted oilfish, but there are ocean (Uyeno et al., 1991; Fricke et al., Ocean remain uncertain. insufficient data to address this 1994). Stomach content analysis has Insufficient data exist to phenomenon (Stobbs et al., 1991). revealed a variety of prey items quantitatively estimate coelacanth Across the coelacanth’s range, including deepwater fishes ranging from population abundance or trends over juveniles (<100 cm) are largely absent cephalopods (including cuttlefish) to time for the majority of its range. from survey and catch data, suggesting eels such as conger eels (Uyeno et al., Population abundance estimates are that earlier life stages may exhibit 1991). The fish exhibits low-energy drift greatly challenged by sampling and differences in distribution and habitat feeding behavior, which is thought to survey conditions wherein deep use (Fricke et al., 2011). Length at birth conserve energy and oxygen for the fish. technical scuba or submersibles are is assumed to be 40 cm (Bruton et al., Metabolic demands have been studied necessary to reach and document the 1991a). Size classes between 40 and 100 in the coelacanth, and demonstrate that coelacanth in its natural habitat. cm are largely absent from surveys of they have one of the lowest resting Quantitative estimates of coelacanth the Comoros, South Africa, and metabolisms of all vertebrates (Hughes abundance have been made only for the Tanzania; these smaller sizes are also et al., 1972; Fricke et al., 2000a). The Comoro Islands. Coelacanth population absent from shallower water, suggesting coelacanth’s gill surface area is much abundance estimates for the western that they inhabit deeper water than smaller than other fishes of similar size; coastline of Grand Comoro were older individuals (Fricke et al., 2011). In this morphological feature is a factor initially made in the late 1980s by general, the distribution and relative

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abundance of juveniles across the al., 2013). These studies suggest that L. Because rates of molecular substitution coelacanth’s range remains unknown. chalumnae comprises multiple and evolution are thought to be similar Population estimates have not been independent populations distributed for these two species, the significantly conducted in other parts of the across the Western Indian Ocean. lower diversity measures for L. coelacanth’s range, and it is possible However, based on limited samples, the chalumnae points to smaller that undiscovered populations exist geographic patterns and relatedness populations (as compared to L. across the Western Indian Ocean among coelacanth populations are not menadoensis) or the occurrence of because coelacanths have been caught well understood. Using mitochondrial repeated genetic bottlenecks, rather than (in low numbers) off the coast of DNA analyses, Nikaido et al. (2011) slow evolution rate alone (Inoue et al., Madagascar, Kenya and Mozambique. demonstrated that individuals from 2005, Nikaido et al., 2013). Low Based on current understanding, northern Tanzania differ from those diversity within populations and coelacanth habitat and distribution is from southern Tanzania and the evidence for inbreeding suggest that determined by the species’ need for cool Comoros. In fact, this study estimated populations are independent and small. water and structurally complex caves that a northern Tanzanian population While population structure is not and shelf overhangs for refuge. Using diverged from the rest of the species an clearly resolved across the region, these requirements, Green et al. (2009) estimated 200,000 years ago. Nikaido et available genetic data suggest the conducted a bathymetric survey using al. (2011) hypothesized that following: (1) Oceanographic and data coverage of the Western Indian differentiation of individuals from environmental conditions may cause Ocean in order to identify potential northern Tanzania may relate to uneven gene flow among coelacanth habitat for coelacanth populations, divergence of currents in this region, populations across the region; (2) beyond occupied habitat already where hydrography limits gene flow and populations across the Western Indian identified. The authors identified reduces the potential for drifting Ocean are independent, and do not several locations off Mozambique and migrants. More recent data reflecting a represent strays from the Comoros, or a South Africa that met characteristics of greater number of samples and higher- panmictic population (or a population coelacanth habitat. Lack of adequate resolution population analyses do not in which all individuals are potential data coverage for Tanzania and support a genetic break between mates); (3) Evolutionary rates of Madagascar precluded thorough individuals from north and south coelacanths are extremely slow, and analyses of these regions, so the authors Tanzania. Instead, this more robust lower diversity in L. chalumnae as did not rule out these locations as population-genetics approach reveals compared with L. menadoensis points suitable coelacanth habitat. Although significant divergence among to smaller population sizes and/or this bathymetric study did not lead to individuals from South Africa, genetic bottleneck effects. any additional surveys to confirm its Tanzania, and two populations which Summary of Factors Affecting the findings, the analysis demonstrates the diverged but are co-existing within the African Coelacanth presence of suitable habitat throughout Comoros; the mechanism of divergence Available information regarding the Western Indian Ocean, and thus the between the two co-existing populations potential for yet-undiscovered current, historical, and potential threats of the Comoros remains unclear to the coelacanth was thoroughly coelacanth populations. Based on the (Lampert et al., 2012). All studies are data presented, populations that have reviewed (Whittaker, 2014). Across the consistent in that they demonstrate low species’ range, we found the threats to been surveyed appear to be stable with absolute divergence among populations, unknown abundance and trends the species to be generally low, with which either relates to extremely low elsewhere. isolated threats of overutilization evolutionary rates in L. chalumnae, or Genetic data on coelacanth through bycatch and habitat loss in population structure are limited and recent divergence of populations after portions of its range. Other possible known distribution of coelacanth going through a bottleneck (such as a threats include climate change, populations is potentially biased by founding effect) (Lampert et al., 2012). overutilization via the curio trade, and targeted survey efforts and fishery catch Information derived from unique habitat degradation in the form of data. However, recent whole-genome sequences of mitochondrial DNA pollution; however, across the species’ sequencing and genetic data available support the Comoros as an ancestral full range we classify these threats as for multiple coelacanth specimens can population to other populations low. We summarize information be used to cautiously infer some distributed throughout the Western regarding each of these threats below patterns of population structure and Indian Ocean, because this population according to the factors specified in connectivity across the coelacanth’s appears to have a greater number of section 4(a)(1) of the ESA. Available known range (Nikaido et al., 2011; ancestral haplotypes (Nikaido et al., information does not indicate that Lampert et al., 2012; Nikaido et al., 2011). neither disease nor predation is 2013). Currently, whole-genome All coelacanth populations operative threats on this species; sequences exist for multiple individuals demonstrate the common characteristic therefore, we do not discuss those from Tanzania, the Comoros, and from of low diversity, but the Comoros further here. See Whittaker (2014) for the Indonesian coelacanth L. population is the least diverse (Nikaido additional discussion of all ESA section menadoensis. et al., 2011, Nikaido et al., 2013). 4(a)(1) threat categories. Significant genetic divergence at the Genetic evidence for inbreeding has species level has been demonstrated to been observed in investigations of The Present or Threatened Destruction, exist between L. chalumnae and L. coelacanth mitochondrial DNA and Modification, or Curtailment of Its menadoensis (Inoue et al. 2005) as DNA fingerprinting, where high band- Habitat or Range described above. sharing coefficients showed significant There is no evidence curtailment of Intraspecific population structure has inbreeding effects (Schartl et al., 2005). the historical range of L. chalumnae has been examined using L. chalumnae The species L. chalumnae exhibits occurred throughout its evolutionary specimens from Tanzania, the Comoros, significantly lower levels of genetic history, either due to human and southern Africa (Nikaido et al., divergence than its sister species L. interactions or natural forces. Genetic 2011; Lampert et al., 2012; Nikaido et menadoensis (Nikaido et al., 2013). data and geological history suggest that

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the split between L. chalumnae and its coelacanth habitat may be affected, and in this region provide essential fish Indonesian sister species L. range reduced. However, the nature of nursery habitat and are hot spots for menadoensis occurred 40–30 Mya, and these economic and land use changes, biodiversity (Salm, 1983). Loss of that the genus was previously as well as their direct effect on nearshore coral habitat may negatively distributed throughout the coasts of sedimentation and subsequent impact impact pelagic fish species due to loss Africa and Eurasia (Springer, 1999; on coelacanth habitat, remain highly of nursery habitat; it is highly uncertain Inoue et al., 2005). However, no data are uncertain. how these impacts may affect the prey available to inform an understanding of Pollution of coastal African waters availability for the coelacanth. Dynamite historical changes in the range of the does not currently pose a direct threat fishing in the Comoros was observed species L. chalumnae. Although the to the coelacanth. A review of heavy recently by researchers (Fricke et al., order Coelacanthiformes was deemed to metals in aquatic ecosystems of Africa 2011). While this method is not have become extinct 65 million years showed generally low concentrations, widespread throughout the Comoros, before the 1938 discovery in South close to background levels, and much reduction in the sustainability of Africa, this surprising encounter cannot lower than more industrial regions of nearshore or pelagic fish populations be used as evidence for a curtailment of the world (Biney et al., 1994). Yet, may encourage fishermen to increase the species’ range from historical levels surprisingly, a toxicological study of use of these new methods. Dynamite given lack of any historical data on the two coelacanth specimens detected fishing in Tanzania is widespread, and species prior to its discovery. The lipophilic organochlorine pollutants has led to destruction of nearshore coral species is naturally hidden from human such as polychlorinated biphenyl reefs and disruption of essential fish observation, and therefore, highly (PCBs) and habitat (Wells, 2009). Destructive technical diving, deep water survey dichlorodiphenyltrichloroethane (DDT) fishing practices occur throughout coral equipment, or unique fishing techniques (Hale et al., 1991). Levels ranged from ¥ reefs along the coast of the Western (such as hand lines) are required to 89 to 510 pg kg l for PCB and 210 to Indian Ocean (Salm, 1983). The true ¥l reach the fish’s cavernous, structurally 840 pg kg for DDT concentration, and extent to which the destruction of near complex, and deep habitat; thus, the were highest in lipid-rich tissues such shore coral habitat may affect the contemporary and historical extent of its as the swim bladder and liver (Hale et coelacanth remains uncertain, range remains unclear. al., 1991). The coelacanth has high lipid especially as the fish is thought to Due to its occurrence in deep water content, and its trophic position may consume primarily deep-water prey (>100 meters), the coelacanth may be increase the probability of toxic (Uyeno, 1991; Uyeno et al., 1991). particularly buffered from human bioaccumulation. Insufficient data are disturbance (Heemstra et al., 2006). available to determine the impact of Overutilization for Commercial, Nonetheless, increases in human these toxins on coelacanth health and Recreational, Scientific, or Educational population and development along the productivity. Purposes coastline of the Western Indian Ocean Direct habitat destruction is likely to Bycatch could impart long-term effects on the impact coelacanths off the coast of fish throughout its range. World human Tanga, Tanzania. Plans are in place to Since its discovery in 1938, all known population forecasts predict that the build a new deep-sea port in Mwambani coelacanth catches are considered to largest percentage increase by 2050 will Bay, 8 km south of the original Tanga have been the result of bycatch. be in Africa, where the population is Port. The construction of the Mwambani Particularly in the Comoro Islands, expected to at least double to 2.1 billion port is part of a large project to develop where the highest number of coelacanth (Kincaid, 2010). The result of increased an alternative sea route for Uganda and catches has occurred, researchers have population density on coastal other land-locked countries that have found no evidence of a targeted ecosystems of East Africa may include been depending on the port of coelacanth fishery given that methods increased pollution and siltation, which Mombasa. Development of the port do not exist to directly catch the deep- may impact the coelacanth despite its would include submarine blasting and dwelling fish (Bruton et al., 1991c). The use of a deep and relatively stable channel dredging and destruction of coelacanth meat is undesirable, and environment. known coelacanth habitat in the vicinity thus the fish is not consumed by Human population growth will likely of Yambe and Karange islands—the site humans (Fricke, 1998). lead to increases in agricultural of several of the Tanzanian coelacanth Out of 294 coelacanth catches since production, industrial development, catches (Hamlin, 2014). The new port is its 1939 discovery, the majority of and water use along the coast of the scheduled to be built in the middle of catches (n = 215 as of 2011) have been Western Indian Ocean; these land use a newly-implemented Tanga Coelacanth a result of bycatch in the oilfish, or changes may increase near shore Marine Park. The plans for Mwambani Revettus, artisanal fishery occurring sedimentation, possibly affecting Bay’s deep-sea port construction appear only in the Comoro Island archipelago coelacanth habitat. As described earlier, to be ongoing, despite conservation (Stobbs et al. 1991; Nulens et al. 2011). sedimentation is theorized to negatively concerns. If built, the port would likely The Comoros oilfish fishery uses impact coelacanth distribution disrupt coelacanth habitat by direct unmotorized outrigger canoes (locally (Springer, 1999). The coelacanth has elimination of deep-water shelters, or by called galawas). The fish are caught been shown to avoid caves with turbid a large influx of siltation that would using handlines and hooks close to water, even if other preferred conditions likely result in coelacanth displacement. shore at depths as great as 800m (Stobbs of shelter and food are present Habitat destruction in the form of et al., 1991). This traditional fishery is (Hissmann et al., 2006). Many East nearshore dynamite fishing on coral known locally as maze´ fishing, and African countries are still developing, reefs may indirectly impact the coelacanth catches have only occurred and the population is growing. coelacanth due to a reduction in prey on Grand Comoro and Anjouan Islands Increased food demand may lead to availability, but these impacts are highly (Stobbs et al., 1991). Oilfish are changes in land and water use, and an uncertain. As a restricted shallow-water traditionally caught at night, an act increase in agriculture and thus run-off activity, this destructive fishing would considered locally to be very dangerous and siltation to the coast. It is possible not impact the coelacanth’s deep (+100 (Stobbs et al., 1991). Often, this artisanal that, if increases in siltation occur, m) habitat directly. However, coral reefs fishing is performed only on dark

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moonless calm nights. In general, Trawls have been the mechanism for effort due to sustained global demand subsistence fishing in the region is only 3 total coelacanth catches; minimal (Hutchings et al., 2002). It is reasonable limited by weather conditions, and catch through trawling is thought to to conclude that the use of shark gillnets often disrupted by monsoon or tropical relate to the coelacanth’s preferred will continue or increase in Tanzania storms. This fishery is also limited by a rocky steep cavernous habitat, substrate and will continue to expand throughout tradition of social pressure which not suitable for trawling activity (Benno the Western Indian Ocean; however, restricts fishing to offshore waters et al., 2006). The first confirmed whether this trend will result in an adjacent to each fisherman’s village coelacanth catches using shark gillnets increased threat of coelacanth bycatch is (Stobbs et al., 1991). occurred in Madagascar in 1995 and in uncertain, especially given the Since its discovery in the Comoros (in Tanzania in 2003, although a few earlier uncertainty over the fish’s range and 1938), coelacanth catch rate has been unconfirmed catches in these locations habitat use throughout the coast of East very low, between 2–4 individuals per may have occurred as early as 1953 Africa. year. Coelacanth catch rate in the (Benno et al., 2006). The first Tanzanian Commercial Interest Comoros shows no significant trend catch in 2003 followed the introduction over time; however, it has fluctuated of shark gillnets in the region in 2001 The coelacanth is not desirable historically with changes in fishing (Benno et al., 2006). As of September commercially as a traditional food technology and shifts in the ratio 2003, the capture of coelacanths has source or for artisanal handicrafts. between artisanal and more modern been dominated by those caught in Targeted methods of fishing the pelagic fishing methods (Stobbs et al., Tanzania (Nulens et al., 2011). Since the coelacanth have never been developed, 1991; Plante et al., 1998). From a first 2003 catch in Tanzania, over 60 and local cultures do not value the broader temporal perspective, there was catches via deep water gillnets have coelacanth commercially or for an increasing but insignificant change in been reported, with over 12 fish caught/ subsistence purposes (Fricke, 1998). In the Comoros, the coelacanth has coelacanth catch from the Comoros from year between 2003 and 2008 (Benno et become a source of pride and national 1954 to 1995 (Plante et al., 1998). al., 2006; Nulens et al., 2011). These heritage (Fricke, 1998). However, However, between 1995 and 2008, the shark gillnets are set at depths between cultural interest in the coelacanth does number of galawas in the Comoros has 50 and 150m, and it is thought that not put the fish at risk, and on the declined steadily, corresponding with a accidental coelacanth catches in contrary, may encourage its steady increase in motorized boats Tanzania occur when coelacanths leave conservation. Commercial interest (Fricke et al., 2011). The most recent their caves for nighttime hunting through tourism to the coelacanth’s update of coelacanth catch inventory (Nyandwi, 2009). indicates that catch rates in the Comoro habitat is not a realistic threat either, as archipelago have declined and Expansion of the shark gillnet fishery the deepwater habitat is largely stabilized over the past decade (Nulens across the Western Indian Ocean may inaccessible. In the 1980s there was a et al., 2011). In fact, between 2000 and result in increased bycatch of the rumor that Japanese scientists were 2008, catch rates were the lowest ever coelacanth, as has been observed off the attempting to develop a new anti-aging observed, likely due to the increase in coast of Tanzania, but the potential for serum using the coelacanth notochord motorized boats and decreased artisanal such an increase is uncertain. Available oil. Although these claims made handline fishing over the past decade information suggests that shark fishing international headlines, the rumor has (Fricke et al., 2011). Today, maze´ effort has been increasing off the coast since been rejected. As Fricke pointed fishing is going out of favor in the of east Africa, including the coelacanth out (Fricke, 1998), the unsubstantiated Comoros (Plante et al., 1998; Fricke et range countries of Mozambique, rumor of the ‘fountain of youth’ serum al., 2011); this trend is expected to Madagascar, Kenya, and South Africa had an unexpected result of stirring continue into the future, and reduces (Smale, 2008). Techniques for catching publicity and conservation interest in fishing pressure on the coelacanth in sharks in this region include deep-set the fish. Interest in the coelacanth this region, most likely explaining the shark gillnets, such as those responsible notochord oil for medicinal purposes reduction in coelacanth catch over the for the commencement of coelacanth does not pose a threat to the species, as past decade (Stobbs et al., 1991; Plante bycatch in Tanzania in 2003 (Nulins et claims of its life extending properties et al., 1998; Fricke et al., 2011; Nulens al., 2011). Shark gillnet fishing is used are unsubstantiated. et al., 2011). Fishing mortality has been in other East African countries, such as Interest in coelacanth specimens on determined to be negligible in the Mozambique, where these fisheries are the black market is a possible threat to Comoros population, likely relating to highly profitable, and are driven by the the species. The concern mostly its population stability over time demand for fin exports, with evidence surrounds a curio trade rather than a (Bruton et al., 1991a; Fricke et al., 2011). for frequent illegal export occurring potential aquarium trade. Because the Outside of the Comoros, coelacanths (Pierce et al., 2008). Despite the use of fish is deep-water dependent, it survives have been caught in Tanzania, gillnet fishing practices elsewhere in for only a short period of time at the Madagascar, Mozambique, Kenya, and East Africa, other areas have not shown surface, and thus far, is not maintained South Africa (Nulens et al., 2011). a similar spike in coelacanth bycatch as in aquariums. Several attempts have Historically, far fewer coelacanth has been observed in Tanzania. been made to keep the coelacanth alive catches have occurred outside of the Quantification of effort from the shark in captivity, but these attempts have Comoros Islands. However, over the gill net fishery in South Africa has been demonstrated that the deep water fish is past decade, the trend in coelacanth challenging due to high levels of illegal fragile and that it has been shown to catches shows a drastic increase in or unreported fishing occurring; for survive at the surface for less than 10 catch rate off Tanzania via shark gillnets example, as little as 21 percent of the hours (Hughes et al., 1972); the cause of (Fricke et al., 2011; Nulens et al., 2011). actual catch for shark gillnet and seine death is thought to be a combination of Hand line maze´ fisheries are absent fisheries may be reported in South capture stress and overheating resulting outside of the Comoros, thus catches Africa (Hutchings et al., 2002). in asphyxiation. Comment threads across the rest of the Western Indian Nonetheless, shark fisheries in this found on the popular Web site Monster Ocean have occurred using different region are thought to be overexploited, Fish Keepers, a forum for private gear—deep-set shark gillnets and trawls. which may lead to an increase in future aquarium and fish hobbyists, reveal

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widespread knowledge of the (Suzuki et al., 1985; Hamlin, 1992). The coelacanth is typically observed coelacanth’s fragility; these hobbyists These attempts were not successful; it at 15–20 °C, with upper thermal express general understanding that the was determined that coelacanth cannot preferences of 22–23 °C (Hughes et al., coelacanth’s life can be sustained at be directly caught, and that they only 1972). The effect of these thermal surface depth no longer than a few survive for a few hours outside of their boundaries on the coelacanth’s hours (Hamlin, 1992; Monsterfish, deep water environments (Hamlin, distribution has been demonstrated by a 2007). Thus, black market trade of the 1992). In the future, larger aquariums 1994 survey of the Comoro Islands, coelacanth for private aquaria is not a may pursue the use of pressurized tanks which revealed a 68 percent decrease in realistic threat. However, the black- to keep the coelacanth alive in captivity, cave inhabitants and a 32 percent market curio trade may be a source of but their success is uncertain given the decrease in the total number of exploitation. The same fish hobbyist challenge of transporting a fish from its coelacanths encountered as compared to forums reveal general interest in the fish native habitat, and then maintaining it a 1991 survey (Hissmann et al., 1998). as a curio specimen, and willingness to in an aquarium environment. Temperature is thought to have directly pay large sums relative to the typical led to this decline in coelacanth Other Natural or Manmade Factors Comoran income for a dead specimen observations; in 1994, temperature of Affecting Their Continued Existence (Monsterfish, 2009). Thus, black market the survey region was 25.1 °C, the curio trade may provide an economic Climate Change warmest ever recorded by researchers incentive for capture of the fish. Coelacanth habitat preference and there (Hissmann et al., 1998). However, However, we did not find data distribution is dictated by specialized it is important to note that individually- suggesting that a black market curio requirements for appropriate shelter identifiable coelacanths had returned to trade is currently active. (caves, caverns, and shelves), prey their previous habitat in subsequent Scientific Interest availability, and a combination of depth surveys (Fricke et al., 2011); this and temperature that meets the fish’s suggests that the warm conditions in Since discovery of the species in 1994 led to a displacement of 1938, international scientists and need for oxygen (relating to optimal blood saturation at 15 °C) (Hughes, coelacanth habitat, but did not lead to researchers have cherished the extirpation of that population, or a coelacanth as the only representative of 1972). Evidence from coelacanth habitation in South Africa is reduction in the population abundance. an important evolutionary branch in the This information suggests that warming tree of life. This has led to a long history particularly useful in demonstrating the trade-offs among these important may impact coelacanth distribution, but of surveys to better understand the fish’s there may be suitable habitat to ecology, habitat, distribution, and characteristics: There, coelacanths accommodate a displacement of evolution. A tissue library from occupy depths of 100–140 m. The populations, where warming may not bycaught specimens is maintained at the optimal temperature for the uptake of ° lead to decreases in population sizes or Max Planck Institute in Germany, which oxygen (15 C) occurs at lower depths of extirpation of populations. Despite deep provides the opportunity for scientific 200 m, where fewer caves exist. It is water warming that has occurred over use of samples derived from these thought that the occupation of shallower the last decade, the surveyed coelacanth accidental coelacanth catches (Fricke, depths is a trade-off between the need population in the Comoros is described 1998). Coelacanth specimens have been for shelter and optimal oxygen uptake; as stable, and not declining (Fricke et used by more than 30 laboratories. In increases in oceanic temperature as is al., 2011). earlier years of coelacanth research, a expected in connection with climate reward of US$300–400 was offered to change may disrupt the tight balance Based on the majority of climate fishermen for each coelacanth caught between coelacanths’ metabolic needs model predictions, it is likely that (Fricke, 1998). However, those rewards and the need for refuge (Roberts et al., current coelacanth habitat will reach have not been offered for decades. Prior 2006). temperatures exceeding the fish’s to strict regulations on coelacanth trade, Across the globe, ocean temperature is thermal preferences by 2100 (IPCC, the global museum trade offered increasing at an accelerated rate (IPCC, 2013). It is unlikely that the low- between US$400 and US$2000 for each 2013). The extent of this warming is diversity fish with long generation times specimen caught. Today, trade of the reaching deeper and deeper waters will physiologically adapt to withstand coelacanth is prohibited by the (Abraham et al. 2013). Increase of global the metabolic stress of a warming ocean. Convention on International Trade in mean surface temperatures for 2081– However, the fish may be able to move Endangered Species (CITES) because the 2100 relative to 1986–2005 is projected to suitable habitat outside of its current coelacanth is listed as an Appendix I to likely be in the ranges derived from range, thus adapting its range to avoid species; however, some transfer of the concentration-driven CMIP5 model the warming deep water conditions. If specimens for scientific study is simulations by the Intergovernmental the fish is displaced based on its need permitted. We did not find any evidence Panel on Climate Change (IPCC), that is, for cooler waters, but complex cave that targeted coelacanth catch for 0.3 °C to 1.7 °C (RCP2.6), 1.1 °C to 2.6 shelters are not available, local scientific purposes is occurring. Thus, °C (RCP4.5), 1.4 °C to 3.1 °C (RCP6.0), extirpation or range restriction may the demand for specimens for scientific or 2.6 °C to 4.8 °C (RCP8.5) (IPCC, 2013). occur. However, currently, these research is not considered a threat. While these predictions relate to surface impacts and responses are highly In the future, scientific interest and ocean temperatures, evidence from uncertain. Thus, it is reasonable to study may be used as a basis for the deep-water ocean measurements and conclude that a warming ocean may public display of the coelacanth. The models suggest that heat flux to the impact the fish’s distribution, but the public display of the fish would be of deep ocean has accelerated over the last impact of warming on the future high commercial value, and efforts to decade (Abraham et al., 2013). If deep- viability of the species is uncertain. Due keep the coelacanth in captivity have water warming continues to keep pace to the coelacanth’s temperature- already been made. In the late 1980s with (or exceed the pace of) surface dependent oxygen demand, coupled and early 1990s, American and Japanese warming, even the most conservative with a highly specific need for deep aquariums attempted to directly capture IPCC scenarios may mean a warming of structurally complex cave shelter, and bring the coelacanth into captivity current coelacanth habitat. warming oceanic waters may pose a

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threat to the coelacanth and and national mechanisms, including the levels of diversity reflect low adaptive displacement of populations, but the Kyoto Protocol of 2007. Because climate and evolutionary potential, making the impact of this threat on the future change-related threats have not been coelacanth particularly vulnerable to viability of the species is highly clearly or mechanistically linked to environmental change and episodic uncertain, and climate change threats decline of coelacanths, the adequacy of events. These events may reduce have not been clearly or mechanistically existing or developing measures to diversity further, and result in a linked to any decline in coelacanth control climate change threats is not significant change or loss of variation in populations. possible to fully assess, nor are life history characteristics (such as sufficient data available to determine reproductive fitness and fecundity), Inadequacy of Existing Regulatory what regulatory measures would be morphology, behavior, or other adaptive Mechanisms needed to adequately protect this characteristics. Due to their low CITES Appendix I regulates trade in species from the effects of climate diversity, coelacanth populations may species in order to reduce the threat change. While it is not possible to be at an increased risk of random international trade poses to those conclude that the current efforts have genetic drift and could experience the species. The coelacanth is included in been inadequate such that they have fixing of recessive detrimental genes CITES-Appendix I. Appendix I contributed to the decline of this that could further contribute to the addresses those species deemed species, we consider it likely that species’ extinction risk (Musick, 2011). threatened with extinction by coelacanth will be negatively impacted While demographic factors increase international trade. CITES prohibits by climate change given the predictions the coelacanth’s vulnerability, the status international trade in specimens of of widespread ocean warming (IPCC, review classified the risk of threats these species except when the purpose 2013). across its range as low or very low of the import is not commercial, meets (Whittaker, 2014). We found that, in criteria for other types of permits, and Extinction Risk general, the coelacanth is largely can otherwise be legally done without In general, demographic buffered from habitat impacts due to its affecting the sustainability of the characteristics of the coelacanth make it occurrence in deep water. Thus, the population, for instance, for scientific particularly vulnerable to exploitation. threats of dynamite fishing, pollution, research. In these exceptional cases, While coelacanth abundance across its land-use changes, and sedimentation are trade may take place provided it is entire range is not well understood, it is considered low-risk. The direct loss of authorized by the granting of both an likely that population sizes across the coelacanth habitat may occur if the deep import permit and an export permit (or Western Indian Ocean are small, as port of Mwambami Bay is developed off re-export certificate). We found no described in Whittaker (2014). The the coast of Tanzania. However, evidence of illegal trade of the likelihood of low abundance makes whether plans to build this port will coelacanth. Trade is limited to the coelacanth populations more vulnerable come to fruition remains uncertain, and transfer of specimens for scientific to extinction by elevating the impact of the effects will impact a small portion purposes. There is no evidence that stochastic events or chronic threats of the coelacanth’s range. The threat of CITES regulations are inadequate to resulting in coelacanth mortality. Their port development does not represent a address known threats such that they growth rate and productivity is widespread threat to the species, and are contributing to the extinction risk of extremely limited. The coelacanth has the port of Mwambami Bay is the only the species. one of the slowest metabolisms of any large coastal development project (that The coelacanth is also listed as vertebrate, and this relates to their we found) that would directly impact Critically Endangered on the meager demand for food, slow swim the fish. International Union for the speed and passive foraging, need for As for impacts from overutilization, Conservation of Nature’s (IUCN) Red refuge to rest, and small gill surface area bycatch has historically been thought to List. The IUCN is not a regulatory body, which limits their absorption of oxygen. pose the greatest threat to the and thus the critically endangered In addition, their gestation period is coelacanth, but survey data show there listing does not impart any regulatory longer than any vertebrate (3 years), is no observed link between coelacanth authority to conserve the species. although their fecundity is moderate. bycatch and population decline. A The threat to the coelacanth stemming They are long-lived species, with long decade ago, the Comoros oilfish fishery from anthropogenic climate change generation times. The extremely long was responsible for the highest rate of includes elevated ocean temperature gestation period and late maturity coelacanth bycatch. Historically, the reaching its deep-water habitat and makes the coelacanth particularly Comoran fishery was responsible for resulting in decreased fitness or vulnerable to external threats such as catch rates of about 3 fish per year, and relocation of populations based on bycatch, possibly impeding recovery is not thought to have contributed to elimination of suitable habitat, which from mortality events (Froese et al., declines in population abundance. may become restricted due to the tight 2000). Genetic data suggest that the While the Comoran oilfish fishery has interaction between the coelacanth’s coelacanth comprises independent and seen recent declines in effort and has thermal requirements and need for isolated populations, originating in the never contributed to population decline highly complex cave shelter and prey. Comoros, but fully established around of the coelacanth, a greater threat of Impacts of climate change on the marine the Western Indian Ocean. The small bycatch has emerged in Tanzania over environment are already being observed and isolated nature of coelacanth the last decade. As evidenced by high in the Indian Ocean and elsewhere populations, only three of which are rates of coelacanth bycatch via the shark (Hoerling et al., 2004; Melillo et al., confirmed to exist, increases gillnet fishery, which began in 2001 in 2014) and the most recent IPCC vulnerability by preventing their Tanzania, this fishing method has the assessment provides a high degree of replacement and recovery from external potential to impact the coelacanth. certainty that human sources of threats and mortality events, and Since 2003 in Tanzania, coelacanth greenhouse gases are contributing to increases the potential for local catch rates have been more than 3 times global climate change (IPCC, 2013). extirpations. Finally, the species greater than ever observed in the Countries have responded to climate exhibits extremely low levels of Comoros, at over 10 fish per year. It is change through various international diversity (Schartl et al., 2005). Low unclear whether this catch rate is

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unsustainable due to limited organization the Coelacanth Yet, it is unclear whether these have information on trends and abundance of Conservation Council (CCC) has been been used at sea, their success is the Tanzanian population. While the primary body advocating for unproven, and it is unknown whether traditional Comoran handline fishing is coelacanth conservation over the years the method has been adopted by local no longer the most pressing bycatch since 1987. fishermen. threat to the fish, data suggest that the The CCC has its headquarters in Ongoing scientific research on the expansion of a shark gill net fishery Moroni, Comoros, and the Secretariat is coelacanth may play a role in throughout the Western Indian Ocean currently in Grahamstown, South Africa coelacanth conservation, as could result in additional coelacanth with branches in Canada, the United management of the species can improve bycatch. The reduction of sustainable Kingdom, the United States, Germany with a more complete understanding of fisheries throughout the east African and Japan. The CCC has set forth general its biology and natural history. In 2002, and South African coastline may objectives of promoting coelacanth South Africa instituted its African encourage shifts to alternative fishing research and conservation, along with Coelacanth Ecosystem Programme, methods, such as gillnets, or trawling establishing an international registry of which has coordinated an extensive closer to shore, both of which could coelacanth researchers and the array of research including bathymetric increase the probability of coelacanth compilation of a coelacanth inventory surveys, taxonomic studies, and bycatch. Bycatch in Tanzania is an and bibliography, which were published observational expeditions. This program ongoing threat, and potential for for the first time in 1991 and recently is funded by the Global Environment additional coelacanth bycatch across the updated in 2011 (Bruton et al., 1991b; Facility of the World Bank and it is in fish’s range poses a potential but Nulens et al., 2011). its third phase, taking an ecosystem- uncertain threat to the fish’s persistence Several conservation initiatives were based approach to understanding into the foreseeable future. Coelacanth implemented in the Comoros in the coelacanth distribution and habitat population abundance in Tanzania, and 1990s to reduce coelacanth bycatch. For utilization across the Western Indian whether current bycatch rates are instance, fishing aggregation devices Ocean, and providing deep-water sustainable, is unknown. Thus, the risk were installed to encourage pelagic research tools and resources for this of bycatch across the species’ entire fishing and reduce pressure on the research. range is generally low. There is no real coelacanth from nearshore handline Local efforts for marine conservation indication that overutilization for fishing. During this time, the use of exist in the Comoros. For example, the scientific purposes, public display, or motorized boats was encouraged for the Mohe´li Marine Park takes a co- the curio trade is occurring; thus we do same purpose, in order to direct fishing management approach to stop some not consider these factors as off-shore and reduce the use of artisanal destructive fishing and conserve marine contributing a risk to the future handlines. Initially, there were some habitat using a series of no-take persistence of the species across its challenges, including lack of reserves. The park encompasses 212 range. infrastructure preventing the repair of km2, and was set up during a 5-year Because threats are low across the motors. However, the fishing trend biodiversity conservation project which species’ range, we have no reason to today in the Comoros shows a clear shift began in 1998, funded by the World consider regulatory measures to motorized pelagic fishing, and Bank’s Global Environment Facility; the inadequate in protecting the species. reduced interest in traditional handline goals of the project were to address the Regarding other natural or manmade fishing; this trend is occurring due to a loss of biodiversity in Comoros and factors, the threat of climate change via natural shift in social perspectives and develop local capacity for natural ocean warming may work local economic trends. resource management (Granek et al., synergistically to enhance all other A supporter of coelacanth 2005). However, no alternative revenue- threats to the coelacanth across its conservation and member of the U.S. generating activities have been range, but the nature of these impacts is Explorer Club, Jerome Hamlin, author provided, making life difficult for some highly uncertain as described in and curator of the Web site fishermen. The World Bank’s Global Whittaker (2014). The extent of this DINOFISH.com, has encouraged the use Environment Facility biodiversity impact on the coelacanth remains of a ‘Deep Release Kit’ for coelacanth management project in the Park ended uncertain, and there has been no clear conservation when bycaught. The Deep in 2003, and there has been no source or mechanistic link between climate Release Kit was created in response to of additional financing to continue the change or temperature warming and the ‘Save the Coelacanth Contest’ resource co-management. The Moheli coelacanth population declines. Thus, sponsored by DINOFISH.com (Hamlin, Park has brought together some key the threat of climate change poses a low 2014). The kit consists of a barbless institutions to encourage sustainable risk to the coelacanth. hook attached to a sack. The fisherman management and monitoring of marine Overall, the fish’s demographic puts some of his sinker stones in the habitat of the Comoros; however, factors make it particularly vulnerable sack, places the hook in the lower jaw specific laws have not been enacted, to ongoing and future threats, but of the fish he has just caught with the and existing legislation has not been existing threats pose a generally low shank pointing down to the sack, and enforced (Ahamada et al., 2002). No risk. Thus, we find that the coelacanth releases the fish to the bottom where it coelacanths have ever been caught off is at a low risk of extinction due to frees itself. The purpose of the Deep the island of Moheli, so the park’s current and projected threats to the Release procedure is to get the fish impact on bycatch of the species is not species. quickly to the cold bottom water with applicable. no further exertion on its part. A surface Other conservation efforts in the form Protective Efforts release (in theory) leaves the fish of marine parks distributed throughout Since its discovery, much debate has without the strength to get back down the Western Indian Ocean may benefit surrounded the need to conserve the to depth. Hundreds of these devices the coelacanth by reducing habitat coelacanth, as an evolutionary relic and have been distributed in the Comoros destruction and improving prey for its value to science. The long history and Tanzania. These kits are some of the availability; however, the direct impacts of this debate was summarized by only direct coelacanth conservation of these conservation efforts on the Bruton (1991). The international measures in the Comoros or Tanzania. species is difficult to evaluate. Efforts to

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improve marine resource management guidance from the IUCN has encouraged not lead us to conclude that the species and conservation in developing nations plans for community-based and meets the definition for either an of east Africa have increased in the past adaptive park management (Harrison, endangered species or a threatened decade. Today, 8.7 percent of the 2010). species based on the rangewide continental shelf in Kenya, 8.1 percent Applying the considerations analysis. Thus, under the final in Tanzania, and 4.0 percent in mandated by our PECE policy, we Significant Portion of Its Range (SPR) Mozambique have been designated as determine that the implementation and policy announced in July 2014, we must marine protected areas (Wells et al., enforcement of the park’s regulations go on to consider whether the species 2007). Many of these parks intersect and goals are unclear and untested; may have a higher risk of extinction in with known coelacanth habitat, or are in further, there are several reasons to a significant portion of its range (79 FR range countries where coelacanths have believe that infrastructure, funding, and 37577; July 1, 2014). been caught and potential populations park management may not be adequate The final policy explains that it is exist. However, in many areas, ongoing to fully prevent coelacanth bycatch necessary to fully evaluate a portion for socioeconomic challenges have within the park’s boundaries: For one, potential listing under the ‘‘significant precluded effective management of illegal fishing off the coast of Tanzania portion of its range’’ authority only if these regions (Francis et al., 2002). is high (Tobey et al., 2006; Hempson, information indicates that the members Analysis of east African Marine 2008; Wells, 2009). Widespread poverty of the species in a particular area are Protected Area (MPA) management has and other regional socio-economic likely both to meet the test for biological demonstrated that socio-economic challenges in the region have reduced significance and to be currently barriers make it more difficult to reach the effectiveness and implementation of endangered or threatened in that area. conservation goals (Tobey et al., 2006). other east African marine parks, and it Making this preliminary determination Because of this, much effort has gone is likely that the Tanga Coelacanth triggers a need for further review, but into creating community-based Marine Park will face similar challenges does not prejudge whether the portion conservation planning in recent years (Toby, 2006; Wells, 2012). Although actually meets these standards such that (e.g., Harrison (2010)). Management recommendations and goals are set in the species should be listed: place to increase tourism to the Park as constraints still remain. First, there are To identify only those portions that large gaps in ecosystem knowledge an economic offset for stricter fishing regulations, the economic infrastructure warrant further consideration, we will surrounding these marine parks; for determine whether there is substantial instance, many vital habitats and and incentives needed for this shift are information indicating that (1) the portions species are not yet fully represented by not in place or have not yet been proven may be significant and (2) the species may be MPAs in place today (Wells et al., 2007). to be effective. Next, there are plans to in danger of extinction in those portions or Next, monitoring is not widely build a new deep-sea port in Mwambani likely to become so within the foreseeable implemented and data are not available Bay, just 8 km south of the original old future. We emphasize that answering these questions in the affirmative is not a to determine whether biodiversity or Tanga Port, which would include submarine blasting and channel determination that the species is endangered socio-economic goals are being met or threatened throughout a significant (Wells et al., 2007). dredging and destruction of known coelacanth habitat in the vicinity of portion of its range—rather, it is a step in A new marine park in Tanga, Yambe and Karange islands—the site of determining whether a more detailed analysis of the issue is required. Tanzania has been put in place, and was several of the Tanzanian coelacanth prompted by increases in coelacanth catches. The new port is scheduled to be 79 FR 37586. catch in the region. The Tanga built in the middle of the Tanga Thus, the preliminary determination Coelacanth Marine Park is located on Coelacanth Marine Park. The that a portion may be both significant the northern coastline of Tanzania, construction of Mwambani port is part and endangered or threatened merely extending north of the Pangani River of a large project to develop an requires NMFS to engage in a more estuary 100 km along the coastline alternative sea route for Uganda and detailed analysis to determine whether towards Mafuriko village just north of other land-locked countries which have the standards are actually met (Id. at Tanga city. The park covers an area of been depending on the port of 37587). Unless both are met, listing is 552 km2, of which 85 km2 are terrestrial Mombasa. The plans for Mwambani not warranted. The policy further and 467 km2 are marine. The plans for Bay’s deep-sea port construction appear explains that, depending on the the park were announced in 2009, and to be ongoing, despite conservation particular facts of each situation, NMFS a general management plan published in concerns. It is unclear whether this port may find it is more efficient to address 2011 (Parks; MPRU, 2011). The goal of will be built, but its presence would the significance issue first, but in other the Tanga Coelacanth Marine Park is to negate many of the benefits (even now, cases it will make more sense to conserve marine biodiversity, resource unproven) of the Park. The general examine the status of the species in the abundance, and ecosystem functions of management plan for the park will be potentially significant portions first. the Park, including the coelacanth and fully evaluated every 10 years, with a Whichever question is asked first, an its habitat; and enable sustainable mid-term review every 5 years. The affirmative answer is required to livelihoods and full participation of effectiveness of Tanga Coelacanth proceed to the second question. Id. (‘‘[I]f local community users and other key Marine Park is not yet known, and for we determine that a portion of the range stakeholders. The plans for the park, reasons described above, we do not is not ‘‘significant,’’ we will not need to specific to the coelacanth, are to restrict consider this park to provide certain determine whether the species is fishing within its boundaries, including conservation measures that would endangered or threatened there; if we fishing with deep-set shark gillnets, the alleviate extinction risk to the species. determine that the species is not primary source of coelacanth bycatch in endangered or threatened in a portion of the area. Additional restrictions against Significant Portion of Its Range Analysis its range, we will not need to determine destructive fishing and development As noted above, we find that the if that portion was ‘‘significant.’’). Thus, practices have been set forth in the species is at a low risk of extinction if the answer to the first question is park’s 2011 general management plan throughout its range. In other words, our negative—whether that regards the (MPRU, 2011). Partnership and range-wide analysis for the species does significance question or the status

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question—then the analysis concludes population comprises a significant threats; the population is not thought to and listing is not warranted. portion of the range of the species maintain connectivity with other After a review of the best available because this portion’s contribution to populations, and thus has no source for information, we identified the the viability of the African coelacanth is replacement of individuals lost outside Tanzanian population of the African so important that, without the members of its own reproductive processes. Fast- coelacanth as a population facing in this portion, the African coelacanth moving currents along the Eastern coast concentrated threats because of would be likely to become in danger of of Africa are thought to prevent increased catch rates in this region since extinction within the foreseeable future, connectivity among populations in the 2003, and the threat of a deep-water port throughout all of its range. region (Nikaido et al., 2011). This may directly impacting coelacanth habitat in Because the Tanzanian population of be particularly true for Tanzania. We this region. Due to these concentrated the coelacanth was determined to consider current evidence for the threats, we found that the species may represent a significant portion of the Tanzanian population’s high isolation be at risk of extinction in this area. range of the species, we performed an from the rest of the species to contribute Under the policy, if we believe this extinction risk assessment on the to a moderate risk of extinction, as these population also may constitute a Tanzanian population by evaluating are natural factors (relevant under ‘‘significant’’ portion of the range of the how the demographic factors section 4(a)(1)(E)) that may increase African coelacanth, then we must go on (abundance, productivity/growth rate, vulnerability of this population by to a more definitive analysis. We may spatial structure/connectivity, and preventing its replacement and recovery either evaluate the extinction risk of this diversity) of the species would be from external threats and mortality population first to determine whether it impacted by the ESA section 4(a)(1) events, and increase the potential for is threatened or endangered in that factors, considering only those factors extinction. portion or first determine if it is in fact affecting the Tanzanian population. Genomic analyses of individuals from ‘‘significant.’’ Ultimately, of course, Coelacanth abundance across its the Tanzanian population and other both tests have to be met to qualify the entire range is not well understood, and representatives of the species reveal that species for listing. no abundance estimates exist for the divergence and diversity within and We proceeded to evaluate whether Tanzanian population. Based on general among populations is very low (Nikaido this population represents a significant knowledge of the African coelacanth, et al., 2013). Low levels of diversity portion of the range of the African the Tanzanian population is likely reflect low adaptive and evolutionary coelacanth. The Tanzanian population associated with very restricted and potential, making the Tanzanian is one of only three confirmed specific habitat requirements and low population particularly vulnerable to populations of the African coelacanth, growth rates. We conclude that it is environmental change and episodic all considered to be small and isolated. likely that the population size of the events. These events may reduce Because all three populations are Tanzanian population is small for the diversity further, and result in a isolated, the loss of one would not same reasons described above for the significant change or loss of variation in directly impact the other remaining species as a whole: It exhibits low levels life history characteristics (such as populations. However, loss of any one of diversity (Nikaido et al., 2013), long reproductive fitness and fecundity), of the three known coelacanth generation times, and restricted habitat morphology, behavior, or other adaptive populations would significantly (Hissmann et al., 2006; Fricke et al., characteristics. Due to the Tanzanian increase the extinction risk of the 2011). The likelihood of low abundance population’s low diversity, this species as a whole, as only two small makes the Tanzanian population more population may be at an increased risk populations would remain, making vulnerable to extinction by elevating the of random genetic drift and could them more vulnerable to catastrophic impact of stochastic events or chronic experience the fixing of recessive events such as storms, disease, or threats resulting in coelacanth mortality. detrimental genes that could further temperature anomalies. Tanzanian and Growth rate and productivity for the contribute to the species’ extinction risk Comoran populations are approximately Tanzanian population is thought to (Musick, 2011). 1,000 km apart, ocean currents are exhibit similar characteristics to other Regarding habitat threats to the thought to have led to their divergence populations of the species. The species Tanzanian population, loss and over 200,000 years ago, and connectivity as a whole has one of the slowest degradation of coelacanth habitat can between them is not thought to be metabolisms of any vertebrate. The take the form of pollution, dynamite maintained (Nikiado et al., 2011). The extremely long gestation period and late fishing, sedimentation, and direct loss South African population is separated maturity makes the Tanzanian through development. Future human from the Comoran and Tanzanian population particularly vulnerable to population growth and land use changes populations by hundreds of miles. The external threats such as bycatch, off the coast of Tanzania increase these Tanzanian population exhibits the possibly impeding recovery from threats to the Tanzanian population, but greatest genetic divergence from the mortality events (Froese et al., 2000). their trends and impacts are highly other populations, suggesting that it The Tanzanian population is thought uncertain. In general, the coelacanth is may be the most reproductively isolated to represent a single isolated population largely buffered from habitat impacts among them (Lampert et al., 2012). of the species. It has been estimated that due to its occurrence in deep water, and Potential catastrophic events such as this population diverged from the rest of general effects of pollution and storms or significant temperature the species 200,000 years ago (Nikaido development are similar to those changes may affect the Comoran and et al., 2011). Differentiation of described for the rest of the species. Tanzanian populations simultaneously, individuals from the Tanzanian However, specifically related to the due to their closer geographic population may relate to divergence of Tanzanian population, direct loss of proximity. The South African currents in this region, where habitat is likely to occur if the deep port population, while not as genetically hydrography limits gene flow and of Mwambami Bay is developed. The isolated, may experience isolated reduces the potential for drifting port is planned to be built just 8 km catastrophic events due to its geographic migrants. The isolated nature of the south of the original old Tanga Port, and isolation. This reasoning supports our Tanzanian population lowers the this would include submarine blasting conclusion that the Tanzanian potential for its recovery from external and channel dredging and destruction of

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known coelacanth habitat in the vicinity reasons described above for the rest of boundaries within which differences in of Yambe and Karange islands—the site the species. control of exploitation, management of of several of the Tanzanian coelacanth Overall, the Tanzanian population’s habitat, conservation status, or catches. The new port is scheduled to be demographic factors make it particularly regulatory mechanisms exist that are built in the middle of the Tanga vulnerable to ongoing and future significant in light of section 4(a)(1)(D)’’ Coelacanth Marine Park. The threats, which pose a moderate risk to of the ESA (61 FR 4722; February 7, construction of Mwambani port is part the species. Based on the best available 1996). of a large project to develop an information, threats of bycatch to the Significance: If a population segment alternative sea route for Uganda and Tanzanian population appear to be is found to be discrete under one or both other land-locked countries that have persistent, and the potential of the above conditions, then its been depending on the port of development of a deep port within this biological and ecological significance to Mombasa. The plans for Mwambani population’s habitat could be the taxon to which it belongs is Bay’s deep-sea port construction appear catastrophic to the population in the evaluated. This consideration may to be ongoing, despite conservation foreseeable future. Thus, we find that include, but is not limited to: (1) concerns, and thus it is reasonable to the Tanzanian population is at a ‘‘Persistence of the discrete population conclude that it poses a likely threat to moderate risk of extinction due to segment in an ecological setting unusual the species. Whether plans to build this current and projected threats. or unique for the taxon; (2) evidence port will come to fruition remains Therefore, we conclude that the that the loss of the discrete population uncertain, but if built, the deep port Tanzanian population is at moderate segment would result in a significant could significantly impact the risk of extinction in a significant portion gap in the range of a taxon; (3) evidence Tanzanian population of coelacanths by of the African coelacanth’s range of the that the discrete population segment destroying habitat directly. For the species. represents the only surviving natural Tanzanian population, the construction Distinct Population Segment Analysis occurrence of a taxon that may be more abundant elsewhere as an introduced of this deep-water port could be In accordance with the SPR policy, if catastrophic, and it is clear that the population outside its historic range; a species is determined to be threatened and (4) evidence that the discrete boundaries of the new Tanga Marine or endangered in a significant portion of Park are insufficient in halting plans for population segment differs markedly its range, and the population in that from other populations of the species in the port’s development. significant portion is a valid DPS, we As for impacts from overutilization, its genetic characteristics’’ (61 FR 4722; will list the DPS rather than the entire February 7, 1996). bycatch has historically been thought to taxonomic species or subspecies. pose the greatest threat to the Because the Tanzanian population Discreteness coelacanth. While survey data from the represents a significant portion of the The Tanzanian population cannot be Comoros show there is no observed link range of the species, and this population differentiated from other populations between coelacanth bycatch and is at a moderate risk of extinction, we based on its morphology. In fact, no population decline, since 2003 in performed a DPS analysis on that coelacanth population exhibits Tanzania, coelacanth catch rates have population. significant distinguishing morphological been more than 3 times greater than ever As defined in the ESA (Sec. 3(15)), a characteristics, and morphological observed in the Comoros, at over 10 fish ‘‘species’’ includes any subspecies of differences within the Latimeria genus per year. It is unclear whether this catch fish or wildlife or plants, and any as a whole have been debated (Pouyad rate is sustainable due to limited distinct population segment of any et al., 1999, Holder et al., 1999; information on trends and abundance of species of vertebrate fish or wildlife Erdmann et al., 1999). No unique the Tanzanian population. The further which interbreeds when mature. The behavioral, physical, or ecological expansion of a shark gill net fishery in joint NMFS–U.S. Fish and Wildlife characteristics have been identified for Tanzania, as has been observed over the Service (USFWS) policy on identifying the Tanzanian population to set it apart last decade, could result in additional distinct population segments (DPS) (61 from the rest of the taxon. Only a single coelacanth bycatch. Bycatch in FR 4722; February 7, 1996) identifies dedicated survey of the Tanzanian Tanzania is an ongoing threat. While two criteria for DPS designations: (1) population is available; thus, future direct data assessing Tanzanian The population must be discrete in surveys may reveal distinguishing coelacanth population decline are not relation to the remainder of the taxon ecological features of the population. available, the relatively high and (species or subspecies) to which it As stated above, genetic data on persistent catch rate in this region has belongs; and (2) the population must be coelacanth population structure are the potential to deplete this small and ‘‘significant’’ (as that term is used in the limited and known distribution of isolated population, which has life context of the DPS policy, which is coelacanth populations is potentially history characteristics that greatly different from its usage under the SPR biased by targeted survey efforts and impede its recovery and resiliency to policy) to the remainder of the taxon to fishery catch data. However, recent mortality. which it belongs. whole-genome sequencing and genetic We consider the threat of Discreteness: A population segment of data available for multiple coelacanth overutilization for scientific purposes, a vertebrate species may be considered specimens can be used to cautiously public display, or for the curio trade as discrete if it satisfies either one of the infer some patterns of population low for reasons described above, as they following conditions: (1) ‘‘It is markedly structure and connectivity across the apply to the rest of the species. separated from other populations of the coelacanth’s known range (Nikaido et We consider the threat of inadequate same taxon as a consequence of al., 2011; Lampert et al., 2012; Nikaido regulatory mechanisms as low for the physical, physiological, ecological, or et al., 2013). Intraspecific population Tanzanian population for the same behavioral factors. Quantitative structure has been examined using L. reasons described above for the rest of measures of genetic or morphological chalumnae specimens from Tanzania, the species. Additionally, we classify discontinuity may provide evidence of the Comoros, and southern Africa the risk of climate change as low for the this separation’’; or (2) ‘‘it is delimited (Nikaido et al., 2011; Lampert et al., Tanzanian population for the same by international governmental 2012; Nikaido et al., 2013). These

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studies suggest that L. chalumnae depth, prey, temperature, and shelter of extinction from the aforementioned comprises multiple isolated and requirements are remarkably similar threats within the foreseeable future. In reproductively independent populations among the known coelacanth our consideration of the foreseeable distributed across the Western Indian populations (Hissman et al., 2006). We future, we evaluated how far into the Ocean, only three which have been found no evidence to suggest that future we could reliably predict the confirmed (inhabiting waters off of differences in the ecological setting of operation of the major threats to this Tanzania, the Comoros, and South the Tanzanian population have led to population, as well as the population’s Africa). any adaptive or behavioral response to those threats. We are While population structure of the characteristics that set the population confident in our ability to predict out taxon, described earlier, is not fully apart from the rest of the taxon, or several decades in assessing the threats resolved, all genetic data available contribute significant adaptive diversity of overutilization and habitat suggest that the Tanzanian population to the species. destruction, and their interaction with represents a single isolated population The Tanzanian population is one of the life history of the coelacanth, with of the species. Multiple genetic studies only three known populations within its lifespan of 40 or more years. With corroborate a significant divergence the species. Although it is not the only regard to habitat destruction, we between Tanzanian individuals, and surviving natural occurrence of the evaluated the likelihood of the deep individuals from the South African and taxon, we determined that loss of this water port being constructed. If the port Comoros populations (Nikaido et al.; population segment would result in a is to be developed, the results could 2011, Lampert et al., 2012). This significant gap in the taxon’s range for significantly impact the Tanzanian includes evidence from both nuclear the following reasons: Although coelacanth population. Evidence and mitochondrial DNA (Nikaido et al., coelacanth populations are not thought suggests that the plans for its 2011, Lampert et al., 2012, Nikaido et to maintain reproductive connectivity, construction are moving forward; its al., 2013). The Tanzanian population is loss of one population would make the construction is not certain, but likely. If the most diverged of all coelacanth other two populations more vulnerable built, the construction of the port would populations (Lampert et al., 2012). to catastrophic events, as explained likely occur within the next decade. Differentiation of individuals from the earlier. The extent of the Tanzanian With bycatch, and its interaction with Tanzanian population may relate to population’s range is not known, but the fish’s demographic characteristics, divergence of currents in this region, given the existence of only three known we feel that defining the foreseeable where hydrography limits gene flow and coelacanth populations considered to be future out to several decades is reduces the potential for drifting small and isolated, loss of the appropriate. Based on this information, migrants (Nikaido et al., 2011). All Tanzanian population would constitute we find that the Tanzanian population available data suggest that the a significant gap in the range of the is at a moderate risk of extinction within Tanzanian population does not likely taxon, and thus we consider this the foreseeable future. Therefore, we maintain connectivity with other population to be significant to the taxon consider the Tanzanian population to be populations, and likely has no source as a whole. threatened. for replacement of individuals outside We determined that the Tanzanian In accordance with the our SPR of its own reproductive processes. population is discrete based on policy, if a species is determined to be The Tanzanian population is evidence for its genetic and geographic threatened or endangered across a geographically isolated from the isolation from the rest of the taxon. The significant portion of its range, and the Comoran and South African population also meets the significance population in that significant portion is populations. The Tanzanian population criterion set forth by the DPS policy, as a valid DPS, we will list the DPS rather is approximately 1,000 km away from its loss would constitute a significant than the entire taxonomic species or the Comoran population and over 4,000 gap in the taxon’s range. Because it is subspecies. Based on the best available km away from the South African both discrete and significant to the scientific and commercial information population, with oceanic currents taxon as a whole, we identify the as presented in the status report and this further reducing their potential for Tanzanian population as a valid DPS. finding, we do not find that the African connectivity. While it is thought that the Proposed Determination coelacanth L. chalumnae is currently in Comoran population is the source of danger of extinction throughout all of its other populations along the Western We assessed the ESA section 4(a)(1) range, nor is it likely to become so in the Indian Ocean, the Tanzanian and South factors and conclude that the species, foreseeable future. However, because African populations may have been viewed across its entire range, the Tanzanian population represents a established as many as 200,000 years experiences a low risk of extinction. significant portion of the range of the ago, as genetic data suggest (Nikaido et However, we determined that the species, and this population is al., 2011). Tanzanian population constitutes a threatened, we conclude that the Based on genetic evidence, and the significant portion of the range of the African coelacanth is threatened in a clear geographic isolation of the species, as defined by the SPR policy significant portion of its range. Because Tanzanian population, we determined (79 FR 37577; July 1, 2014). The the population in the significant portion that the Tanzanian population of L. Tanzanian population faces ongoing or of the range is a valid DPS, we will list chalumnae is discrete from other future threats from overutilization and the DPS rather than the entire populations within the species. habitat destruction, with the species’ taxonomic species or subspecies. natural biological vulnerability to Therefore, we propose to list the Significance overexploitation exacerbating the Tanzanian DPS of the African The Tanzanian population does not severity of the threats. The Tanzanian coelacanth as threatened under the ESA. persist in an ecological setting unusual population faces demographic risks, or unique for the taxon. Although the such as population isolation with low Similarity of Appearance Tanzanian individuals are thought to productivity, which make it likely to be The petition requested that, if the inhabit limestone ledges rather than influenced by stochastic or depensatory African coelacanth were listed under volcanic caves where Comoran and processes throughout its range, and the ESA, the Indonesian coelacanth also South African individuals are found, the place the population at an increased risk be listed due to its ‘‘similarity of

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appearance.’’ The ESA provides for Effects of Listing longer necessary. Section 4(a)(3)(A) of treating any species as an endangered Conservation measures provided for the ESA (16 U.S.C. 1533(a)(3)(A)) species or a threatened species even if species listed as endangered or requires that, to the maximum extent it is not listed as such under the ESA threatened under the ESA include prudent and determinable, critical if: (1) Such species so closely resembles recovery plans (16 U.S.C. 1533(f)); habitat be designated concurrently with the listing of a species. However, critical in appearance, at the point in question, concurrent designation of critical a species which has been listed habitat shall not be designated in habitat, if prudent and determinable (16 pursuant to section 4 of the ESA that foreign countries or other areas outside U.S.C. 1533(a)(3)(A)) and consistent enforcement personnel would have U.S. jurisdiction (50 CFR 424.12(h)). with implementing regulations; Federal substantial difficulty in attempting to The best available scientific data as agency requirements to consult with differentiate between the listed and discussed above identify the NMFS under section 7 of the ESA to unlisted species; (2) the effect of this geographical area occupied by the ensure their actions do not jeopardize substantial difficulty is an additional species as being entirely outside U.S. the species or result in adverse threat to the listed species; and (3) such jurisdiction, so we cannot designate modification or destruction of critical treatment of an unlisted species will critical habitat for this species. We can habitat should it be designated (16 substantially facilitate the enforcement designate critical habitat in areas in the and further the policy of the ESA. U.S.C. 1536); and, for endangered United States currently unoccupied by While the African and Indonesian species, prohibitions on taking (16 the species, if the area(s) are determined species exhibit morphological U.S.C. 1538). Recognition of the species’ by the Secretary to be essential for the similarities, they are clearly plight through listing promotes conservation of the species. Based on geographically and genetically conservation actions by Federal and the best available information, we have separated. Enforcement personnel state agencies, foreign entities, private not identified unoccupied area(s) in would have no difficulty in groups, and individuals. U.S. water that are currently essential to differentiating between the Tanzanian Identifying Section 7 Conference and the species proposed for listing. Thus, DPS of the African coelacanth and the Consultation Requirements as we discussed above, we will not Indonesian coelacanth because of propose critical habitat for this species. similarity of appearance because their Section 7(a)(2) (16 U.S.C. 1536(a)(2)) Identification of Those Activities That geographic separation (in the Western of the ESA and NMFS/USFWS Would Constitute a Violation of Section Indian Ocean and Indo-Pacific, regulations require Federal agencies to 9 of the ESA respectively) should facilitate regulation consult with us to ensure that activities of taking. The species experience no they authorize, fund, or carry out are not On July 1, 1994, NMFS and FWS overlap in range and catch of both likely to jeopardize the continued published a policy (59 FR 34272) that species is relatively low, and well- existence of listed species or destroy or requires NMFS to identify, to the documented. We do not deem ESA adversely modify critical habitat. maximum extent practicable at the time protection for the Indonesian coelacanth Section 7(a)(4) (16 U.S.C. 1536(a)(4)) of a species is listed, those activities that to be advisable at this time, as the clear the ESA and NMFS/USFWS regulations would or would not constitute a genetic and geographic differences also require Federal agencies to confer violation of section 9 of the ESA. between the two species set them apart with us on actions likely to jeopardize Because we are proposing to list the in a way that allows for easy the continued existence of species Tanzanian DPS of the African identification, regardless of their similar proposed for listing, or that result in the coelacanth as threatened, no appearance. destruction or adverse modification of prohibitions of Section 9(a)(1) of the Because we are proposing to list the proposed critical habitat of those ESA will apply to this species. Tanzanian DPS as a threatened species species. It is unlikely that the listing of this DPS under the ESA will increase Protective Regulations Under Section under the ESA, we also considered any 4(d) of the ESA potential similarity of appearance issues the number of section 7 consultations, that may arise in differentiating between because the DPS occurs outside of the We are proposing to list Tanzanian the proposed DPS and other populations United States and is unlikely to be DPS of the African coelacanth, L. of the species. No morphological affected by Federal actions. chalumnae as threatened under the ESA. In the case of threatened species, characteristics separate the Tanzanian Critical Habitat DPS from other populations of the ESA section 4(d) leaves it to the species. However, we do not conclude Critical habitat is defined in section 3 Secretary’s discretion whether, and to that listing the South African or of the ESA (16 U.S.C. 1532(5)) as: (1) what extent, to extend the section 9(a) Comoran populations based on The specific areas within the ‘‘take’’ prohibitions to the species, and similarity of appearance is warranted. geographical area occupied by a species, authorizes us to issue regulations First, outside of Tanzania, coelacanth at the time it is listed in accordance necessary and advisable for the catches are infrequent, and well with the ESA, on which are found those conservation of the species. Thus, we documented. Second, the three known physical or biological features (a) have flexibility under section 4(d) to coelacanth populations do not overlap essential to the conservation of the tailor protective regulations, taking into geographically. Differentiation between species and (b) that may require special account the effectiveness of available the African and Indonesian coelacanth, management considerations or conservation measures. The 4(d) and likewise between the Tanzanian protection; and (2) specific areas outside protective regulations may prohibit, DPS and other populations of the the geographical area occupied by a with respect to threatened species, some species, could potentially pose a species at the time it is listed upon a or all of the acts which section 9(a) of problem for enforcement of section 9 determination that such areas are the ESA prohibits with respect to prohibitions on trade, should any be essential for the conservation of the endangered species. These 9(a) applied. However, that issue is species. ‘‘Conservation’’ means the use prohibitions apply to all individuals, addressed, at least with respect to of all methods and procedures needed organizations, and agencies subject to imports and exports, by the inclusion of to bring the species to the point at U.S. jurisdiction. We will consider coelacanth in CITES Appendix I. which listing under the ESA is no potential protective regulations

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pursuant to section 4(d) for the Role of Peer Review Executive Order 13132, Federalism proposed threatened coelacanth DPS. In December 2004, the Office of In accordance with E.O. 13132, we We seek public comment on potential Management and Budget (OMB) issued 4(d) protective regulations (see below). determined that this proposed rule does a Final Information Quality Bulletin for not have significant Federalism effects Public Comments Solicited Peer Review establishing a minimum and that a Federalism assessment is not peer review standard. Similarly, a joint To ensure that any final action required. In keeping with the intent of NMFS/FWS policy (59 FR 34270; July 1, resulting from this proposed rule to list the Administration and Congress to 1994) requires us to solicit independent the Tanzanian DPS of the African provide continuing and meaningful expert review from qualified specialists, coelacanth will be as accurate and dialogue on issues of mutual state and in addition to a public comment period. effective as possible, we are soliciting Federal interest, this proposed rule will The intent of the peer review policy is comments and information from the be given to the relevant governmental to ensure that listings are based on the public, other concerned governmental agencies in the countries in which the best scientific and commercial data agencies, the scientific community, species occurs, and they will be invited available. We solicited peer review industry, and any other interested to comment. We will confer with the comments on the African coelacanth parties on information in the status U.S. Department of State to ensure status review report, including from: review and proposed rule. Comments appropriate notice is given to foreign Five scientists with expertise on the are encouraged on this proposal (See nations within the range the DPS African coelacanth. We incorporated DATES and ADDRESSES). We must base (Tanzania). As the process continues, these comments into the status review our final determination on the best we intend to continue engaging in report for the African coelacanth and available scientific and commercial informal and formal contacts with the this 12-month finding. information. We cannot, for example, U.S. State Department, giving careful consider the economic effects of a References consideration to all written and oral listing determination. Before finalizing comments received. A complete list of the references used this proposed rule, we will consider the in this proposed rule is available upon List of Subjects in 50 CFR Parts 223 comments and any additional request (see ADDRESSES). information we receive, and such Administrative practice and information may lead to a final Classification procedure, Endangered and threatened regulation that differs from this proposal National Environmental Policy Act species, Exports, Imports, Reporting and or result in a withdrawal of this listing record keeping requirements, proposal. We particularly seek: The 1982 amendments to the ESA, in Transportation. (1) Information concerning the threats section 4(b)(1)(A), restrict the Dated: February 25, 2015. to the Tanzanian DPS of the African information that may be considered coelacanth proposed for listing; when assessing species for listing. Based Samuel D. Rauch, III. (2) Taxonomic information on the on this limitation of criteria for a listing Deputy Assistant Administrator for species; decision and the opinion in Pacific Regulatory Programs, National Marine (3) Biological information (life Legal Foundation v. Andrus, 675 F. 2d Fisheries Service. history, genetics, population 825 (6th Cir. 1981), NMFS has For the reasons set out in the connectivity, etc.) on the species; concluded that ESA listing actions are preamble, we propose to amend 50 CFR (4) Efforts being made to protect the not subject to the environmental part 223 as follows: species throughout its current range; assessment requirements of the National (5) Information on the commercial Environmental Policy Act (NEPA) (See PART 223—THREATENED MARINE trade of the species; NOAA Administrative Order 216–6). AND ANADROMOUS SPECIES (6) Historical and current distribution Executive Order 12866, Regulatory and abundance and trends for the Flexibility Act, and Paperwork ■ 1. The authority citation for part 223 species; and continues to read as follows: (7) Information relevant to potential Reduction Act ESA section 4(d) protective regulations As noted in the Conference Report on Authority: 16 U.S.C. 1531–1543; subpart B, § 223.201–202 also issued under 16 U.S.C. for the proposed threatened DPS, the 1982 amendments to the ESA, 1361 et seq.; 16 U.S.C. 5503(d) for especially the application, if any, of the economic impacts cannot be considered § 223.206(d)(9). ESA section 9 prohibitions on import, when assessing the status of a species. take, possession, receipt, and sale of the Therefore, the economic analysis ■ 2. In § 223.102, amend the table in African coelacanth. requirements of the Regulatory paragraph (e) by adding a new entry for We request that all information be Flexibility Act are not applicable to the one species in alphabetical order under accompanied by: (1) Supporting listing process. In addition, this the ‘‘Fishes’’ table subheading to read as documentation, such as maps, proposed rule is exempt from review follows: bibliographic references, or reprints of under Executive Order 12866. This § 223.102 Enumeration of threatened pertinent publications; and (2) the proposed rule does not contain a marine and anadromous species. submitter’s name, address, and any collection-of-information requirement association, institution, or business that for the purposes of the Paperwork * * * * * the person represents. Reduction Act. (e) * * *

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Species 1 Citation(s) for listing determination(s) Critical ESA Common name Scientific name Description of listed entity habitat rules

******* Fishes

Coelacanth, African Latimeria African coelacanth population inhab- [Insert Federal Register citation and NA NA (Tanzanian DPS). chalumnae. iting deep waters off the coast of date when published as a final Tanzania. rule].

******* 1 Species includes taxonomic species, subspecies, distinct population segments (DPSs) (for a policy statement, see 61 FR 4722, February 7, 1996), and evolutionarily significant units (ESUs) (for a policy statement, see 56 FR 58612, November 20, 1991).

* * * * * DATES: Information and comments on endangered or threatened under the [FR Doc. 2015–04405 Filed 3–2–15; 8:45 am] the subject action must be received by ESA, or, in the alternative, delineate six BILLING CODE 3510–22–P May 4, 2015. distinct population segments (DPSs) of ADDRESSES: You may submit comments, the common thresher shark, as information, or data, identified by described in the petition, and list them DEPARTMENT OF COMMERCE ‘‘NOAA–NMFS–2015–0025’’ by either as endangered or threatened. Friends of of the following methods: Animals also requested that critical National Oceanic and Atmospheric • Electronic Submissions: Submit all habitat be designated for this species in Administration electronic public comments via the U.S. waters concurrent with final ESA Federal eRulemaking Portal. Go to listing. 50 CFR Parts 223 and 224 www.regulations.gov/ The petitioner states that the common [Docket No. 141219999–5132–01] #!docketDetail;D=NOAA-NMFS-2015- thresher shark merits listing as an 0025. Click the ‘‘Comment Now’’ icon, endangered or threatened species under RIN 0648–XD680 complete the required fields, and enter the ESA because of the following: (1) or attach your comments. Endangered and Threatened Wildlife; • The species faces threats from historical Mail or hand-delivery: Office of and continued fishing for both 90-Day Finding on a Petition To List Protected Resources, NMFS, 1315 East- the Common Thresher Shark as commercial and recreational purposes; West Highway, Silver Spring, MD (2) life history characteristics and Threatened or Endangered Under the 20910. limited ability to recover from fishing Endangered Species Act Instructions: You must submit comments by one of the above methods pressure makes the species particularly AGENCY: National Marine Fisheries vulnerable to overexploitation; and (3) Service (NMFS), National Oceanic and to ensure that we receive, document, and consider them. Comments sent by there is a lack of regulations that Atmospheric Administration (NOAA), specifically protect the common Commerce. any other method, to any other address or individual, or received after the end thresher shark. ACTION: Notice of 90-day petition finding, request for information, and of the comment period, may not be ESA Statutory Provisions and Policy initiation of status review. considered. All comments received are Considerations a part of the public record and will SUMMARY: We, NMFS, announce the 90- generally be posted for public viewing Section 4(b)(3)(A) of the ESA of 1973, day finding for a petition to list the on http://www.regulations.gov without as amended (U.S.C. 1531 et seq.), common thresher shark (Alopias change. All personal identifying requires, to the maximum extent vulpinus) as either endangered or information (e.g., name, address, etc.), practicable, that within 90 days of threatened under the U.S. Endangered confidential business information, or receipt of a petition to list a species as Species Act (ESA) either worldwide or otherwise sensitive information threatened or endangered, the Secretary as one or more distinct population submitted voluntarily by the sender will of Commerce make a finding on whether segments (DPSs) identified by the be publicly accessible. We will accept that petition presents substantial petitioners. We find that the petition anonymous comments (enter ‘‘N/A’’ in scientific or commercial information presents substantial scientific or the required fields if you wish to remain indicating that the petitioned action commercial information indicating that anonymous). Attachments to electronic may be warranted, and promptly the petitioned action may be warranted comments will be accepted in Microsoft publish the finding in the Federal for the species worldwide. We find that Word, Excel, or Adobe PDF file formats Register (16 U.S.C. 1533(b)(3)(A)). When the petition fails to present substantial only. we find that substantial scientific or scientific or commercial information to FOR FURTHER INFORMATION CONTACT: commercial information in a petition support the identification of DPSs of the Chelsey Young, NMFS, Office of and in our files indicates the petitioned common thresher suggested by the Protected Resources (OPR), (301) 427– action may be warranted (a ‘‘positive 90- petitioners, and, as such, we find that 8491 or Marta Nammack, NMFS, OPR, day finding’’), we are required to the petitioned action of listing one or (301) 427–8469. promptly commence a review of the more of these DPSs is not warranted. SUPPLEMENTARY INFORMATION: status of the species concerned, which Accordingly, we will initiate a review of includes conducting a comprehensive the status of the common thresher shark Background review of the best available scientific at this time. To ensure that the status On August 26, 2014, we received a and commercial information. Within 12 review is comprehensive, we are petition from Friends of Animals months of receiving the petition, we soliciting scientific and commercial requesting that we list the common must conclude the review with a finding information regarding this species. thresher shark (Alopias vulpinus) as as to whether, in fact, the petitioned

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