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Vol. 6: 173–184, 2008 ENDANGERED RESEARCH Printed December 2008 doi: 10.3354/esr00102 Endang Species Res Published online August 21, 2008

Contribution to the Theme Section ‘The IUCN Red List of Threatened Species: assessing its utility and value’ OPENPEN ACCESSCCESS Challenges of assessing cetacean population recovery and conservation status

Milton M. R. Freeman*

Canadian Circumpolar Institute, University of Alberta, Edmonton, Alberta T6G 2E1, Canada

ABSTRACT: This paper examines the extent to which depleted populations have recovered (or not) and their perceived current conservation status and prognosis for continued survival, as repre- sented in the International Union for Conservation of Nature and Natural Resources (IUCN) Red Lists. It is concluded that current hypothetical and untested predictions of risk, while in many cases drawing attention to justifiable conservation needs, may seriously undervalue the resilience of species that have evolved to live in a dynamic and ever-changing reality including cen- turies of heavy exploitation by humankind. The paper questions the appropriateness of Red Listing criteria for long-lived highly mobile ocean-dwelling species that are scarcely affected by the princi- pal threats upon terrestrial or aquatic species living in relatively restricted areas subject to habitat fragmentation and/or loss. This analysis draws attention to the problems associated with objectively assessing the conservation status of charismatic species and the value conflicts that may override evidence-based scientific conservation assessments.

KEY WORDS: · · Conservation · Population recovery · Predicting extinction · Red Listing · Charismatic species

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INTRODUCTION of the mysticetus; both are believed to number in the tens or low hundreds at this The history of whaling provides one of the best- time. There are also populations whose princi- known examples of over-exploitation of living pal or exclusive range is in rivers, or in coastal or resources. However, no whale species has become nearshore areas where fishery bycatch and boat traffic extinct in historic or modern times, despite the pres- are causes of high mortality/injury. Thus, there is a far sures of whaling over many centuries, although uncer- greater risk of extirpation or extinction than is likely to tainty exists as to whether pre-modern whaling in the exist for wide-ranging cetacean populations. Many 17th century contributed to the extinction of the North whale species are wide-ranging, with populations Atlantic (Gerber et al. 2000, Clapham & numbering in the tens or hundreds of thousands, and Link 2006). It was only in 2007 that the first known their habitat is expansive and non-fragmented; how- human-caused extinction of a cetacean species is ever, some of these species are shown on the Interna- believed to have occurred, when the last few indivi- tional Union for Conservation of Nature and Natural duals of the world’s rarest species, the Resources (IUCN) Red List as being in serious danger Yangtze Lipotes vexillifer, appear to have of extinction. finally succumbed to boat collisions, dam construction, The 1991 IUCN Red List provides a benchmark to pollution and bycatch in local fisheries (Turvey et al. assess changes in cetacean conservation status at later 2007). There are examples of severely depleted popu- points in time, although the listing criteria and mem- lations of some widely distributed whale species, bership in the Cetacean Specialist Group also changes including the west Pacific population of the gray whale over time. In the 1991 Red List (Klinowska 1991), 5 spe- Eschrichtius robustus and the Spitsbergen population cies are listed as Endangered, 7 as Vulnerable, 1 as

*Email: [email protected] © Inter-Research 2008 · www.int-res.com 174 Endang Species Res: 6:173–184, 2008

Indeterminate, and 65 as Insufficiently Known. New able (IUCN 2007b), that is ‘facing a high risk of extinc- listing criteria were introduced in 1994, and a reassess- tion in the wild’ (IUCN 2007c). The is a ment by the Cetacean Specialist Group using these cri- long-lived, wide-ranging species occurring in every teria listed 2 species as , 6 as ocean and the territorial waters of more than 40 coun- Endangered, 6 as Vulnerable, 1 as Near Threatened, tries from the equatorial to polar regions, yet its classi- 14 as Lower Risk/Conservation Dependent, 33 as Data fication as Vulnerable A1 bd is almost identical to that Deficient (an equivalent category to the ‘Indetermi- of the geoffrensis, listed as nate’ in the 1991 assessments), and 13 as Lower Vulnerable A1cd. Table 3 presents a summary of the Risk/Least Concern (and not included in the 1991 list- factors used to produce these 2 almost identical Red ing; see Table 1). Listings. Furthermore, the Amazon river dolphin has 3 The 1996 list, for the first time also assessed 16 sub- distinct subspecies, each of unknown population num- species and geographical cetacean populations, with 7 bers but thought to collectively total a few thousand of these categorized as Endangered, 5 as Vulnerable, 3 individuals living in a polluted and fragmented habitat as Lower Risk/Conservation Dependent and 1 as Data (with future habitat degradation from mining and Deficient. In many of these listings, reliable population damming, and fishery impacts believed certain to numbers do not exist, and none are mentioned in the intensify). The challenging environmental situation Cetacean Specialist Group Report (IUCN 2007a), and the relatively small and unknown population of where principal importance in several cases is placed river seems to justify a listing as Vulnerable; if on the large numbers of removals during the industrial so, the sperm whale, in comparison, seems much less whaling era, the slow rate of recovery of some popula- ‘threatened with extinction’. Similarly questionable tions, and known or projected environmental threats. listings occur with other formerly heavily exploited whales, e.g. the acutorostrata, the B. bonaeren- RELIABILITY OF WHALE CONSERVATION sis — ‘which may be the most abundant whale ASSESSMENTS species today’ according to the Red List account (IUCN 2007d) — and the B. borealis. Table 4 pro- The current (2007) Red List of cetaceans identifies 15 vides further information on these 4 abundant and species as in danger of extinction in the near future wide-ranging oceanic species taken from the IUCN (i.e. listed as Critically Endangered, Endangered or Red List assessments. Vulnerable). These endangered cetaceans include It is significant that the posited major past, present several riverine or inshore species, some with popula- and future threats to each of these 4 abundant species tions in the low hundreds, together with other more are related to their use for food, scientific, cultural and widely distributed or ubiquitous cetacean species hav- leisure activities, their use in trade, and their vulnera- ing much larger populations (Table 2). Many cetacean bility to fishery-related accidental mortality. As will be population estimates remain uncertain, an example discussed below, the current and probable future being the sperm whale macrocephalus, threat from whale fisheries is in reality very small at whose population estimates in the literature vary from present, and has little realistic chance of expanding in 350 000 (American Cetacean Society, www.acsonline. the future. In the case of the Antarctic minke whale, org/factpack/index.html) to ca. 2 000 000 (www. the current population is believed to range from 70south.com/information/antarctic-animals/whales). 510 000 to 1.14 million (www.iwcoffice.org/conser- The sperm whale appears on the Red List as Vulner- vation/estimate.htm) or from 610 000 to 1.28 million today (www.70south.com/information/ antarctic-animals/ whales). The Red Table 1. The Red List classification of cetaceans in 1991 and 2007 List cites the removal of 98 202 Antarc- tic minke whales taken during the Classification 1991 2007 Notes 1956/57 to 1986/87 whaling seasons as significant. However, assuming a popu- Critically Endangered – 2 CR not used in 1991 (CR) lation of 500 000 minke whales at that Endangered (EN) 5 6 time and a reasonably constant annual Vulnerable (VU) 7 6 rate of removals, that level of harvest Lower Risk (LR) – 28 LR not used in 1991 (98 202 divided by 30 yr = 3274) would Insufficiently Known (K)/ 65 33 K and DD are the same (DD) have represented an annual 0.7% rate Unlisted (U) 1 – U not currently used of removal — a sustainable rate, allow- Indeterminate (I) 1 – I not currently used ing the population to continue growing at >2% each year. If 500 000 is too large Freeman: Cetacean population status and recovery 175

Table 2. Red List of endangered cetaceans (2007). For classification abbreviations see Table 1

Species Classification Occurrence Population estimateSource

Vaquita CR Local, riverine <1000 WWFa Phocoena sinus Yangtze river dolphin CR Local, riverine <100 WWFa Lipotes vexillifer Indus river dolphin EN Local, riverine ca. 1000 Martin (1990) Planista minor Ganges river dolphin EN Local, riverine <2000 WWFa Planista gangetica Hector’s dolphin EN NZ coastal waters ca. 7400 WWFa hectori Amazon river dolphin EN Local, riverine Low–10s of thousands IUCNb Inia geoffrensis Harbour VU Wide-ranging coastal >600000 IUCNb phocoena Beluga VU Wide ranging coastal >200000 ACSc Delphinapterus leucas N. Pacific EN Localized, coastal ca. 150 ACSc Eubalaena japonica N. Atlantic right whale EN Localized, coastal ca. 350 WWFa Eubalaena glacialis EN Wide-ranging, oceanic N. Atlantic: 32 200 IWCd Balaenoptera musculus US west coast: 1900 Gerber et al. (2000) S. Indian Ocean: 5000 Gerber et al. (2000) S. Hemisphere: 1260 Gerber et al. (2000) EN Wide-ranging, oceanic N. Hemisphere: 40 000 ACSc Balaenoptera physalus S. Hemisphere: 85 000 Gerber et al. (2000) Sei whale EN Wide-ranging, oceanic N. Atlantic: ca. 10 000 WWFa Balaenoptera borealis N. Pacific: 22 000 – 37 000 WWFa S. hemisphere: 24000 WWFa Sperm whale VU Wide-ranging, oceanic 200 000 – 1.5 million NMFSe Physeter macrocephalus VU Wide-ranging, N. Atlantic: 10 600 Gerber et al. (2000) Megaptera novaengliae coastal – offshore N. Pacific: 6000 – 8000 Gerber et al. (2000) S. Hemisphere: 17 000 Gerber et al. (2000) aWorld Wildlife Fund: www.panda.org/about_wwf/what_we_do/species/about_species/species_factsheets/ bInternational Union for Conservation of Nature and Natural Resources: www.iucnredlist.org/search-basic cAmerican Cetacean Society: www.acsonline.org/factpack/ dInternational Whaling Commission: www.iwcoffice.org/conservation/estimate.htm eNational Marine Fisheries Service: www.nmfs.noaa.gov/pr/species/mammals/cetaceans/spermwhale.htm

an assumed population level for 1956/57, the level of ongoing whale monitoring and protection, and an offtake was evidently still sustainable as demonstrated always existing need to deploy resources to protect by that 1956/57 population increasing to the current truly threatened or (Brownell et higher population level. al. 1989, Tyack 1989, Gerber et al. 2000). Among the questions that should be asked in regard to whale conservation (and which should likely be kept in mind when considering Red Listing) are the follow- RECOVERY FROM HISTORIC ing: (1) Can depleted whale stocks recover in the OVER-EXPLOITATION medium-term future? (2) How do we assess whether conservation goals are achieved? (3) How can we On biological grounds, it might appear that whales ensure that stocks are safeguarded against biologi- are inherently vulnerable because, although many of cally-significant depletion? These are important pol- the larger species are long-lived (e.g. bowheads live icy-relevant questions, given the high cost of ensuring for more than 200 yr), some species require 10 yr to 176 Endang Species Res: 6:173–184, 2008

Table 3. Comparison of the similar Red List classification of 2 cetaceans

Amazon river dolphin Sperm whale Inia geoffrensis Physeter marcocephalus

Red List Category: VU A1cd Red List category: VU A1bd Population in 10s of thousands Population in 100s of thousands Distributed in 2 river basins in NE South America Distributed through tropical, temperate and sub-polar oceans in both hemispheres Habitat fragmentation (dams) and more being proposed No habitat fragmentation occurring or likely to occur Mining pervasive, if not rampant, contamination Not immediately threatened, but some regional a concern (including mercury) populations require close evaluation and monitoring Fishermen sometimes kill and maim the dolphins Whaling by one Indonesian village and N. Pacific research whaling together remove ca. 20 to 25 whales annually Growing human population along the rivers, conflicts Indonesia traditional whaling and research whaling certain to intensify unlikely to increase offtake. Ship strikes, net entanglements occur fairly often Major threats listed as: Major threats listed as: habitat degradation and loss; harvesting, international trade in the past; fishing-related mortalities; pollution fishery-related mortalities; poor recruitment, affecting habitat and food reproduction (present and future); other intrinsic factors (present and future) Source: www.iucnredlist.org/search/details.php/10831/all Source: www.iucnredlist.org/search/details.php/41755/all

reach sexual maturity and may have 2 to 3 yr intervals 12 000) in the mid-19th century was likely reduced to between uniparous births. However, low natality rates the low hundreds following about 60 yr of intense com- and associated low predation rates (Ford & Reeves mercial whaling that ended during the first decade of 2008) result in annual net rates of increase greater than the 20th century. Although small numbers of removals 10% in humpback whales Megaptera novaengliae and resulted from continuing community-based whaling, between 6 and 14% in southern right whales Eubal- recovery of this severely depleted population was aena australis (Clapham et al. 1999), although between observed from about 40 to 50 yr after commercial 3 and 4% annual population net growth rates are more whaling ended (Hay et al. 2000). Conservative esti- usual for most whales species. mates place the current population at ca. 7300 bow- It appears that today, and likely for the past few heads (Cosens et al. 2006), which under moderate lev- decades, the majority of depleted whale stocks are re- els of hunting is predicted to achieve full recovery in covering (Aron 1988, Gerber et al. 2000). However, in the next few decades (Dueck & Richard 2006). In con- some cases, although particular stocks or population trast, a historic population of bowhead whales esti- segments of a species are recovering, other regional mated to have numbered between 25 000 and 100 000 stocks of that same species may not be (Clapham et al. in the Spitsbergen region was commercially hunted 2008). In the case of the bowhead, the North Pacific from the 17th to the end of the 19th century, and is now population was thought to be reduced to between 1000 thought to only number in the low tens. This suggests and 2000 individuals from a pre-industrial population that no observed recovery of that remnant population of ca. 22 000, when commercial whaling ceased at the has occurred (Clapham et al. 2008). Interestingly, and end of the first decade of the 20th century. Recovery most would argue appropriately, the IUCN Red List was slow initially, and aboriginal hunting continued to status of bowheads has changed over the past few remove small numbers of bowheads each year. As the decades, from ‘Endangered’ in 1986 to ‘Vulnerable’ in whale population numbers increased, so did the annual 1990 to ‘Lower Risk (conservation dependent)’ in 1996 removals, so that 50 or 60 yr after commercial whaling (and currently) as population recovery has occurred ended the aboriginal hunt was annually removing 70 or over much of the species range where the net rate of more bowheads from a population that had recovered population increase is around 3.2% per year. to more than 10 000; it is currently increasing at a net Reliable population information is also available for rate of more than 3% per year (Gerber et al. 2007). the California gray whale, where the pre-commercial In the Canadian eastern , bowheads have whaling population was believed to have numbered shown a similar pattern of recovery, although a smaller between 20 000 and 25 000 before being reduced to a initial population (believed to have numbered around few hundred (Small 1971, Clapham et al. 1999) or a Freeman: Cetacean population status and recovery 177

Table 4. Major current and/or future threats used to justify Red Listing of commercially hunted whales. Population estimates vary from reasonable certainty (for the common minke) to highly uncertain (for the sperm and sei whales). An extensive web-based search suggested that the sources given in footnotes a to e provided estimates within the range of those available from many available sources. LR/NT: Lower Risk/Near Threatened (close to ‘vulnerable’); LR/CD: Lower Risk/Conservation Dependent (subject to a conservation program, the ending of which would qualify the taxon for a Vulnerable, Endangered or Critically Endangered listing)

Species Population estimate Listing Threat category

Common minke 209 800a LR/NT Harvesting (hunting/gathering) – food – Balaenoptera acutorostrata subnational/national/regional/ international trade (past & ongoing). Cultural/scientific/leisure activities Accidental mortalities – bycatch – fisheries-related (ongoing) Antarctic minke >600 000b LR/CD Harvesting (hunting/gathering) – food – Balaenoptera bonaerensis subnational/national/regional/ international trade (past & ongoing). Cultural/scientific/leisure activities Accidental mortalities: bycatch – fisheries-related (ongoing) Sei whale S. Hemisphere 24 000c EN Harvesting (hunting/gathering) – food – Balaenoptera borealis N. Atlantic 10 000 A1abd regional/international trade (past) N. Pacific 22 000 – 37 000 Sperm whale 200 000 – 1.5 Milliond VU Harvesting (hunting/gathering) – food – Physeter macrocephalus 350 000e A1bd regional/international trade (past). Accidental mortalities: bycatch – fisheries-related – entanglement (ongoing). Intrinsic factors: poor recruitment/ reproduction/regeneration (present, future) – other (present, future)

awww.iwcoffice.org/conservation/estimate.htm; bwww.70south.com/information/antarctic-animals/whales; cwww.antarcticconnection.com/antarctic/wildlife/whales/sei.shtml; dwww.nmfs.noaa.gov/pr/species/mammals/cetaceans/ spermwhale.htm; ewww.acsonline.org/factpack/index.html

few thousand (Gerber et al. 2000) during historic com- feeding locations (Moore 2004). Both northern and mercial whaling operations. Ongoing hunting resulted southern right whales were each reduced to about 350 in annual removals of ca. 170 gray whales by commer- individuals by over-hunting of a pre-exploitation popu- cial and aboriginal whalers until ca. 1990, after which lation believed to number ca. 60 000 whales in each case. 140 to 150 whales have been removed annually by However, southern right whales have now increased to aboriginal whalers. It appears that full recovery of the about 8000 and show high rates of increase in several re- population occurred around the year 2000; at this time, gions, notably South Africa (6.6%), Argentina (7.6%) the population peaked at around 26 600, followed by and off the west coast of Australia (11.7 to 13%) (Perry et natural die-off over the next few years to what appears al. 1999). Other stocks are not re- to be a more sustainable population level (given cur- covering at such high rates, and the northern right whale rent conditions) of ca. 23 000. population of about 350 animals remains quite static. It Other whale populations have experienced even has been suggested, on the basis of recent DNA research greater levels of depletion than the Pacific bowhead and (Lippsett 2005), that an uncounted population, repre- the eastern Pacific gray whale. The most notable ex- senting a further 10 to 15% males, may account for the ample is the northern right whale Eubalaena glacialis, northern right whales’ uneven rate of reproductive suc- which currently numbers in the low hundreds and is con- cess (e.g. 26 calves in 2001, the largest number counted sidered the most endangered whale species. It has been since the 1980s; Pittman 2001). Ship collisions and net- suggested that non-recovery may result from difficulty in entanglements are the main causes of mortality in the locating mates or the genetic consequences of small pop- nearshore waters where northern right whales occur ulation size (Gerber et al. 2000), or shortage of suitable (Clapham et al. 1999). 178 Endang Species Res: 6:173–184, 2008

Slow recovery has also been observed in Antarctic One difficulty associated with assessing recovery of stocks of the blue whale Balaenoptera musculus. This depleted whale populations arises from establishing southern hemisphere stock feeds almost exclusively on acceptable estimates of pre-exploitation population a single prey species (krill Euphausia superba), and its levels. The currently accepted method of doing this slow rate of recovery may result from unsuccessful relies heavily on analysis of whaling ship logbooks and competition with other krill-feeding species — includ- industrial records of barrels of oil and sheets of baleen ing the Antarctic minke whale, crabeater seals Lobo- plates entering the market (Tillman & Donovan 1983, don carcinophagus, leopard seals Hydrurga leptonyx Smith & Reeves 2006). However, recent DNA-based and antarctic fur seals Arctocephalus gazella — all of research has produced pre-exploitation population which experienced significant population increases estimates many times larger than those obtained following the decline in blue whale numbers (Boyce through currently accepted log-book and market- 2000). However, blue whale populations in the North report analyses for whales in the North Atlantic Atlantic and the eastern North Pacific appear to be (Roman & Palumbi 2003) and the North Pacific (Alter et increasing (Gerber et al. 2000). al. 2007). The new estimates were obtained by measur- ing the diversity of mutations in the DNA of living whales, and comparing these findings to the rate at MEETING WHALE CONSERVATION GOALS which such mutations are thought to occur in whales. The results suggested a pre-exploitation North Have conservation goals been met? Seeking the Atlantic humpback whale population of 240 000 — a answer to this question has policy importance, as men- large increase over the currently accepted pre-whal- tioned earlier, as well as other practical and ethical ing population estimate of 40 000. The North Atlantic considerations that flow from the ongoing contribution fin whale Balaenoptera physalus population, using of whales to some peoples’ nutritional, cultural, and genetic analytic methods, is 360 000, compared to a socio-economic needs. Unnecessarily restricting ac- currently accepted estimate of between 30 000 and cess to these important resources through unwar- 50 000 (Roman & Palumbi 2003). ranted protection raises ethical and legal issues of Geneticists and whale biologists are critical of these human rights and distributive justice (Ward 1993). The new elevated population estimates, noting the World Conservation Strategy, Agenda 21, the Conven- untested methodology and a number of questionable tion on Biological Diversity, and policies adopted and assumptions being used (Clarke 2003, Lubick 2003, actions taken by IUCN all stress the importance of Mitchell 2004, Palsbøll et al. 2007). For example, social equity and the sustainable use of natural genetic diversity is a product of mutations continuously resources for achieving successful and enduring con- occurring in the aggregate population of all whales of servation outcomes (Edwards 1995, Hutton & Leader- a given species over millions of years of existence (e.g. Williams 2003, Kinne 2006). 6 to 10 million yr for the humpback whale) at unknown Some whale populations may be increasing, and rates that might have varied at different periods of evo- some may not. However, there remains uncertainty lutionary time in populations whose numbers also var- about the reliability of historic and current population ied over those millions of years. In addition, estimates level estimates (see next paragraphs), making it diffi- of genetic diversity are complicated and error-prone, cult to decide whether conservation goals have been such that subtle and small changes in the assumed met in each case. Furthermore, there is very little infor- mutation rates can lead to massive yet spurious mation allowing a useful understanding of non-anthro- changes in the calculated results (Clarke 2003). pogenic marine environmental changes that might influence whale abundance over decadal or century- long intervals. In addition, the high level of current ESTABLISHING VALID AND CREDIBLE public interest in whales encourages biologists and CONSERVATION GOALS managers to be cautious in suggesting that whale recovery has occurred or is underway, unless the evi- Setting conservation goals is more often a question dence for it is quite certain. Currently, a steady stream of culture or politics than of science (Gerber et al. 2000, of reports predicting elevated risks of extinction due to Heazle 2004). If the culturally determined goal is to re- climate warming over the next few decades has caused establish marine ecosystems having several hundred many scientists to express uncharacteristic certitude thousand more large-bodied whales, this raises serious when predicting species’ increased risk of extinction in fisheries management questions in a world that might the next few decades (IUCN 2007e; however, see expect to at least partially meet the future global food- cautions in Hilborn 2006, Krebs & Berteaux 2006, security needs of a growing human population from a Botkin et al. 2007). diverse array of exploitable marine resources. How- Freeman: Cetacean population status and recovery 179

ever, a more basic question remains: What is the pur- Convention on Biological Diversity, the UN Food and pose or ecological justification for seeking to re-create Agricultural Organization (FAO) and such science- an uncertain hypothetical pre-exploitation population based organizations as the North Atlantic Marine level from centuries earlier when the marine environ- Mammal Commission (NAMMCO), the Pacific Inter- ment at that earlier time (1) no longer exists and can national Council for the Exploration of the Sea (PICES) realistically never be reconstituted (even if we knew its and the Indian Ocean Tuna Commission among others composition), and (2) was never anything other than (Iino & Goodman 2003). Whales, being large-bodied the local marine community existing at a unique, top predators, are widely recognized as a management highly dynamic and ever-changing period in time? consideration in some heavily fished regions today. For This improbable and arguably irrelevant exercise example, in northeast US coastal waters, cetaceans are seems to provide the rationale for much of what passes significant predators, removing 3.5 to 6.6 times the for conservation orthodoxy today, at quantity of taken in commercial fishery catches least when whales are being discussed. As has been (Kenney et al. 1997). In the Pacific Ocean, marine cautioned elsewhere, to manage man-altered ecosys- mammals annually remove about 3 times the quantity tems is a dubious concept because we do not know of fish taken by commercial fishers (Trites et al. 1997). enough about ecosystem dynamics to safely conduct The same concerns about cetacean predation exist in large-scale experiments in nature (Kinne 2006). the North Atlantic fisheries (Vikingsson & Kapel 2000) The IUCN Red List is assumed by many interested and elsewhere (Iino & Goodman 2003) where whales parties to be authoritative, and the listings are cer- are abundant or increasing in numbers. tainly used as such by many, including journalists, edi- Although conservation today is concerned with tors and various interest groups. However, there is a maintaining the integrity of biological communities danger when such information is uncritically used by and sustaining the livelihoods of dependent resource individuals who either do not understand the signifi- users, when it comes to highly politicized charismatic cance of, or choose to ignore, such notations as ‘out of species this contemporary understanding of conserva- date’ or using ‘version 2.3’ (i.e. the 1994 listing criteria, tion seems not to apply, and the debate is much more replaced in 1996 by new and improved criteria). The about special interests and values than science (Butter- 2007 Red List categorizes the fin whale as ‘Endan- worth 1992, Mrosovsky 2000, Heazle 2004, Gerber et gered’ and the humpback whale as ‘Vulnerable’ (using al. 2007). the 1994 listing criteria and out-dated population esti- mates). The Cetacean Specialist Group very likely has access to updated data that allow reassessment SAFEGUARDING WHALE STOCKS according to current criteria and population numbers, which in the case of the southern hemisphere hump- Influenced by theoretical formulations (Clarke 1976), back whales have likely doubled or tripled. Regardless many believed that it was impossible for commercial of whether the updated assessments have not been whale fisheries to ever be sustainable (Small 1971, made by the specialists, or not adopted by the wider Myers 1979). However, bioeconomic analysis (Conrad IUCN process, or just not announced to the public, it is & Bjørndal 1993) and better empirically based under- unfortunate, given the advocacy or other political use standing of small-scale whaling practices and com- made of IUCN listings, that posting outdated informa- mon-property issues (see ‘Sustainability of whale fish- tion fuels conflict in politicized fora where whales or ing’) appear to contradict this belief. Indeed, the whaling (Aron et al. 2000, ICR 2007), or the conserva- scientific committees of the International Whaling tion of other charismatic species (Lapointe 1997, Webb Commission (IWC) and the NAMMCO are among the & Carrillo 2000) are the focus of discussion. scientific bodies agreeing that whale stocks number- ing several thousand or more can sustain a regulated commercial whale fishery. An annual take of 2% of a ACCEPTABLE WHALE POPULATION LEVELS conservative population estimate is considered sus- tainable (Gerber et al. 2000), and the IWC has Science cannot tell us what is an ecologically appro- approved a risk-averse management procedure that priate whale (or other species’) population size, for sci- conservatively sanctions catch levels at one-half of 1% ence cannot predict with any degree of certainty or of the estimated whale stock size. This IWC manage- accuracy what the effect on marine biodiversity would ment scheme, the so-called Revised Management be if large increases in whale numbers occurred (see Procedure (RMP), was exhaustively tested by the IWC Beninca et al. 2008). Today, science-based fisheries Scientific Committee and is considered highly con- management stresses an ecosystem, or multi-species, servative in comparison with all other fishery and approach, as variously endorsed in Agenda 21, the wildlife regimes (Gambell 1995). This adaptive man- 180 Endang Species Res: 6:173–184, 2008

agement regime, successfully used in the Norwegian although some have argued that such past examples minke whale fishery since 1993, accommodates all succeeded only because human population size was future oceanic environmental changes, including cli- small and primitive hunting equipment was being mate change and pollution, resulting from known and used. However, early Basque, Dutch, Scottish, Ameri- unknown natural and anthropogenic events. Some can and other nations’ whalers depleted many whale regional whale populations that were reduced to very stocks in a few decades using simple hand-held low levels, and in some cases have not shown recovery, weapons from the 17th century onwards (Ellis 1991), are afforded complete protection from whaling (i.e. suggesting that even simple technology can, in the zero quotas) under current international whaling man- face of a large global market demand, result in serious agement procedures. over-exploitation. Despite the over-exploitation that The global demand for whale products today is characterized industrial whaling in the past, sustain- small, and evidence suggests that it is unlikely to grow able whaling co-existed with unsustainable whaling (Aron et al. 2000, Greenpeace 2007). In 2007, Iceland for centuries, during times when commercial trade in announced that it was suspending its limited commer- whale products flourished, together with subsistence cial whaling operations due to lack of demand for practices (Marquardt & Caulfield 1996, Freeman whale products at home and abroad (Clover 2007). 2001). Even in Japan and Norway (the 2 other, and much Advanced technologies in use today are important larger, commercial-whaling nations), about 40% of for ensuring that whaling is sustainable, more humane people considered whalemeat an unacceptable food and efficient. For example, in regard to monitoring item more than a decade ago (Freeman & Kellert whaling operations for compliance with quotas for 1994), a level of unacceptability likely to increase as example, DNA registers of landed whale products, internationalization continues to transform the con- international observer schemes, and remote sensing of sumer tastes of their growing urban populations. whaling activities (Øen 2005) provide effective means International trade in whale products effectively to deter and detect non-compliance. ended in 1989, and no significant incidents of poaching Some of the critical features of community-based have been discovered since the trade ban was imposed. whaling associated with sustainability include limiting Reports of irregular incidents, involving small amounts whaling access to members of socially defined groups of allegedly unauthorized meat (Baker & Palumbi 1994) dependent upon that resource stock. Whales and have subsequently been accounted for by products whaling often provide multiple benefits to societies lawfully derived from whale strandings, fishery by- depending upon those resources, benefits that vari- catch, and then-remaining stockpiles of legally im- ously possess significant social, symbolic, ceremonial, ported whalemeat (Mills et al. 1997, Phipps et al. 1998). dietary, economic, historical, spiritual and psychic It appears that if illegal taking of whales occurs, it is importance. Related to this, whaling knowledge and neither extensive nor does it involve critically threat- skills, when transmitted inter-generationally, serve to ened whale populations (Clapham et al. 1999). reinforce social solidarity and compliance with prevail- International efforts to control unsustainable whal- ing societal norms that favour the continuing and ing practices before and immediately following World respectful hunting and use of these valued food War II were ineffective, as public indifference in those resources (Young et al. 1994). Now that export markets years resulted in an absence of political will to prevent no longer exist for whale products, commercial whal- serious over-exploitation of whale stocks by economi- ing is to a very great extent practised by individuals cally and politically influential large-scale whaling living in small communities where whaling has been industries. However, today, those industries no longer an honoured profession for generations. It is in such exist, and there is no unmet consumer demand to jus- communities that whaling traditions and identity re- tify replacing them. In addition, there exists wide- main matters of intense personal and collective pride, spread public and government concern about whales and consequent social sanctions remain effective in and whaling, and many well-funded non-government ensuring compliance with best practices (Kalland & organizations are committed to keeping governments Moeran 1992, Aron et al. 2000). vigilant.

DISCUSSION AND CONCLUSIONS SUSTAINABILITY OF WHALE FISHING Many of the scientific uncertainties that justified a Continuation of community-based whale fisheries pause in commercial whaling in the 1980s have, for the that have persisted over centuries suggests that sus- species of interest to commercial whalers, been tainability of whaling can be attained and maintained, removed due to increased attention given to whale Freeman: Cetacean population status and recovery 181

research by the whaling nations and the Scientific 2007, Armstrong et al. 2008), but the appropriate fore- Committee of the IWC. Scientists have developed rig- casting methodology is rarely (if ever) applied by biol- orous sighting-survey methods, robust statistically reli- ogists who nevertheless make confident and dire pre- able methods for obtaining abundance estimates of dictions about the effect of environmental changes on whale stocks, and a safe and simple method of setting species’ survival prospects (Ehrlich & Ehrlich 1975, catch limits; these limits allow stable and continuing Myers 1983, Amstrup et al. 2007). yields, while ensuring that depleted stocks continue to Some biologists express the view that climate recover in the face of changing environmental condi- changes in the 21st century will, due to the rapidity of tions (Gambell 2003). the predicted change, be quite unlike warming It appears that the resilience of many whale stocks episodes in the past, while others note that the pre- and the time elapsed since the collapse of the global dicted changes are no greater nor rapid than most market for whale products have together removed extant species have experienced in the past (Botkin et what was earlier considered the major threat to al. 2007). Whales are highly mobile oceanic animals, whales. The IWC has endorsed its Scientific Commit- without the geographic barriers or habitat fragmenta- tee’s risk-averse and highly conservative procedure for tion or loss commonly faced by many terrestrial spe- setting sustainable catch levels, and these measures, cies. The history of whaling suggests that major com- together with an ongoing decline in market demand, positional changes to whale populations and their allow a very high level of assurance that fear of a associated biotic communities have occurred (Gambell return to unsustainable whaling is unjustified. Never- 2003) and very rapidly (over mere decades), suggest- theless, the major fears that prompted a pause in com- ing that whale species’ resilience has allowed severely mercial whaling 20 yr ago linger (see Table 4), and in depleted populations to nevertheless recover in a rela- regard to listing of whales ‘the difficulty of balancing tively short time (in relation to whale longevity). In the reality of a decline from previous global abun- those cases where recovery has not appeared to have dance with present low probability of extinction is a occurred, it is possible that recovery is occurring very problem that the IUCN criteria have not yet solved’ slowly (e.g. Hay et al. 2000, Heide-Jorgensen et al. (Ross 2000, p. 28; see also Webb & Carrillo 2000, 2007) or that extirpation or displacement of that popu- Mrosovsky 2004). Although the 1994 changes to the lation will prove to be long term or permanent for envi- Red List criteria refined the assessment process, the ronmental or biological reasons (Clapham et al. 2008). current detailed criteria likely cannot overcome prob- Arguably, population recovery or persistence has been lems originating not with the stated criteria per se, but an outcome equal to or more prevalent than non- rather with the manner in which these objective crite- recovery among whale stocks. ria may be subjectively interpreted and applied. It may When considering the almost identical Red Listing of also be appropriate to ask whether the criteria used to the ubiquitous sperm whale and the localized Amazon list terrestrial species are, in every case appropriate river dolphin, one is reminded of the intense attraction when listing wide-ranging, long-lived and abundant that the general public and many biologists who study oceanic species. Just as small island endemic species whales feel toward whales and dolphins; in that are much more vulnerable to extinction than are wide- regard, the sperm whale is particularly iconic. Obtain- ranging species occupying continental land masses, so ing information needed to properly list whales — and too may oceanic species be that much, or more, secure. more importantly, to manage them — is at best an inex- Today, there is concern that whales, together with act science. It appears that for those whose passion for many other taxa worldwide, are beginning to face whales is great, the perception of an elevated risk grave threats that may be far greater than those posed associated with removing even one whale out of a by the current or expected future demand for their robust population may trigger an aversion that can be products. The new threats are associated with pre- just as intense as removing one whale out of a highly dicted effects of climate change on global biodiversity. depleted remnant population. For those individuals, Certainly, and plant distribution patterns and protection and population recovery may be more a population numbers will change in the future, as they moral than a technical and scientific matter. However, have in the past, but as others have noted: ‘recent eco- assessing candidate species for Red Listing does not logical and genetic research, along with specific prob- explicitly call for any moral assessment to be made in lems with present forecasting methods, lead us to the assessment process, although some biologists may believe that current projections of extinction rates are consider use of the ‘precautionary principle’ as justi- overestimates’ (Botkin et al. 2007, p. 228; see also fied moral (as opposed to evidence-based) grounds Berteaux et al. 2006). Science-based predictions when empirical data are lacking. require use of a rigorous methodology when carried The precautionary principle is an approach to out by forecasting specialists (Green & Armstrong responding to uncertainty in the face of risks. It 182 Endang Species Res: 6:173–184, 2008

involves ‘acting to avoid serious or irreversible poten- mammal management. Coast Manag 16:99–110 tial harm, despite lack of scientific certainty as to the Aron W, Burke W, Freeman MMR (2000) The whaling issue. likelihood, magnitude, or causation of that harm’ Mar Policy 24:179–191 Baker CS, Palumbi SR (1994) Which whales are hunted? a (www.pprinciple.net/the_precautionary_principle.html) molecular genetic approach to monitoring whaling. Sci- It appears that this principle can be applied in the ence 265:1538–1539 absence of sound scientific information to support a Beninca E, Huisman J, Heerkloss R, Jöhnk KD and others position regarding the degree of real or perceived (2008) Chaos in a long-term experiment with a plankton community. Nature 451:822–825 extinction threat a species faces. The dangers of such Berteaux D, Humphries MM, Krebs CJ, Lima M and others actions in international fora such as CITES, where del- (2006) Constraints to projecting the effects of climate egates may have little knowledge of the animals or change on mammals. Clim Res 32:151–158 plants being considered for listing, is that invoking the Botkin DB, Saxe H, Araúja MB, Betts R and others (2007) precautionary principle implies that a significant risk Forecasting the effects of global warming on biodiversity. BioScience 57:227–236 of endangerment exists, thus triggering actions that Boyce M (2000) Whaling models for cetacean conservation. overrule the potential benefits offered by another In: Ferson S, Burgmann M (eds) Quantitative methods for important conservation tool, namely sustainable conservation . Springer, New York, p 109–126 resource use (IUCN 2000, Hutton & Leader-Williams Brownell RL Jr, Ralls K, Perrin WF (1989) The plight of the ‘forgotten’ whales. Oceanus 32:5–11 2003). Butterworth DS (1992) Science and sentimentality. Nature What, then, of the precautionary principle that 357:532–534 appears to have been invoked in to justify such Clapham PJ, Link J (2006) Whales, whaling and ecosystems assessments as may have occurred in the sperm in the North Atlantic. In: Estes J (ed) Whales, whaling and whale/Amazon river dolphin cases described earlier? ecosystems. University of California Press, Berkeley CA, p 241–250 Perhaps we should more fully recognize the subjectiv- Clapham PJ, Young SB, Brownell RL Jr (1999) Baleen whales: ity that enters the Red Listing process when reliable conservation issues and the status of the most endangered scientific data do not exist, especially in regard to populations. Mammal Rev 29:37–62 assessments involving charismatic or iconic taxa. Or Clapham PJ, Aguilar A, Hatch LT (2008) Determining spatial and temporal scales for management: lessons from whal- perhaps, heed the recent observation that ‘generally ing. Mar Mamm Sci 24:183–201 the precautionary principle is best left in interna- Clarke CW (1976) Mathematical bioeconomics: the optimal tional conventions’ (Milner-Gulland & Rowcliffe 2007, management of renewable resources. Wiley, New York quoted in Sharp 2007, p. 29) and that over-inflating the Clarke T (2003) Whale numbers disputed: pre-whaling pop- risk of extinction ‘should not be assigned to species ulation estimates upset conservationists. Nature. Avail- able online at: www.nature.com/news2003/030725/full/ that are abundant and widespread, with no known news030721-12.html scenarios that could lead to global extinction, no mat- Clover C (2007) Iceland ends whaling due to lack of demand. ter how compelling the case for public sympathy may Telegraph.co.uk, August 24 2007 posting available online be’ (Webb & Carillo 2000, p. 17). at: www.telegraph.co.uk/earth/main.jhtml?xml=/earth/ 2007/08/24/eawhale124.xml Conrad J, Bjørndal T (1993) On the resumption of commercial Acknowledgements. The author acknowledges helpful com- whaling: the case of the minke whale in the northeast ments provided by 2 anonymous reviewers, suggestions and Atlantic. Arctic 46:164–171 editorial guidance by Drs. Brendan Godley and Matthew Cosens S, Cleator H, Richard P (2006) Numbers of bowhead Godfrey, technical assistance from Ms. Cindy Mason and whales (Balaena mysticetus) in the eastern Canadian Arc- research funding from the National Centres of Excellence tic, based on aerial surveys in August 2002, 2003 and 2004. ArcticNet Program. Can Sci Advis Secretariat Res Doc 2006–052. Fisheries and Oceans Canada, Ottawa Dueck LP, Richard P (2006) Assessment of the recovery poten- LITERATURE CITED tial of the eastern arctic bowhead whale population by deterministic projections with a modified logistic growth Alter SE, Rynes E, Palumbi SR (2007) DNA evidence for his- model. Can Sci Advis Secretariat Res Doc 2006–053. Fish- toric population size and past ecosystem impacts on gray eries and Oceans Canada, Ottawa whales. Proc Natl Acad Sci USA 104:15162–15167 Edwards SR (1995) Conserving biodiversity: resources for our Amstrup SC, Marcot BG, Douglas DC (2007) Forecasting future. In: Bailey R (ed) The true state of the planet. Free the range-wide status of polar bears at selected times in Press, New York, p 211–265 the 21st century. USGS Science Strategy to Support US Ehrlich PH, Ehrlich A (1975) Extinction: the causes and conse- Fish and Wildlife Service Polar Bear Listing Decision. quences of the disappearance of species. Random House, Administrative Report, US Geological Survey, Reston, New York VA Ellis R (1991) Men and whales. Knopf, New York Armstrong JS, Green KC, Soon W (2008) Polar bear popula- Ford JKB, Reeves RR (2008) Fight or flight: antipredator tion forecasts: a public-policy forecasting audit. Working strategies of baleen whales. Mammal Rev 38:50–86 Paper Version 55: February 21, 2008. Available online at: Freeman MMR (2001) Is money the root of the problem? Cul- www.forecastingprinciples.com/Public_Policy/PolBears.pdf tural conflict in the IWC. In: Friedheim, RL (ed) Towards a Aron W (1988) The commons revisited: thoughts on marine sustainable whaling regime. University of Washington Freeman: Cetacean population status and recovery 183

Press, Seattle, WA and CCI Press, Edmonton, AB, IUCN (2007d) IUCN Red List of threatened species: p 123–146 Balaenoptera bonaerensis. Available online at: www. Freeman MMR, Kellert SR (1994) International attitudes to iucnredlist.org/search/details.php/2480/summ whales, whaling and the use of whale products: a six- IUCN (2007e) Extinction crisis escalates: Red List shows apes, country survey. In: Freeman MMR, Kreuter UP (eds) Ele- corals, vultures, dolphins all in danger. Available online at: phants and whales: resources for whom? Gordon and www.iucn.org/en/news/archive/2007/09/12_pr_redlist.htm Breach, Basel, p 293–315 Kalland A, Moeran B (1992) Japanese whaling: end of an era? Gambell R (1995) Management of whaling in coastal commu- Curzon Press, London nities. In: Blix AS, Walløe L, Ulltang O (eds) Whales, seals, Kenney RD, Scott GP, Thompson TJ, Winn HE (1997) Esti- fish and man. Elsevier, Amsterdam, p 699–708 mates of prey consumption and trophic impacts of Gambell R (2003) I am here, where should I be? In: Burns cetaceans in the USA northwest continental shelf ecosys- WCG, Gillespie A (eds) The future of cetaceans in a tem. J Northwest Atl Fish Sci 22:155–171 changing world. Transnational Publishers, Ardsley, NY Kinne O (2006) Protecting nature. Endang Species Res 1:1–2 Gerber L, DeMaster D, Roberts S (2000) Measuring success Klinowska M (1991) Dolphins, and whales of the in conservation. Assessing efforts to restore populations world. The IUCN Red Data book. International Union for of marine mammals is partly a matter of epistemology: Conservation of Nature and Natural Resources, Gland How do you know when enough is enough? Am Sci 88: Krebs CJ, Berteaux B (2006) Problems and pitfalls in relating 316–324 climate variability to population dynamics. Clim Res Gerber LR, Keller AC, DeMaster DP (2007) Ten thousand and 32:143–149 increasing: Is the western Arctic population of bowhead Lapointe E (1997) No red alert over conservation Red Listings. whale endangered? Biol Conserv 137:577–583 Nature 389:904 Green KC, Armstrong JS (2007) Global warming: forecasts by Lippsett L (2005) Diving into the right whale gene pool. scientists versus scientific forecasts. Energy Environ Oceanus 44:18–19 18:997–1021 Lubick N (2003) New count of old whales adds up to a big Greenpeace (2007) Whale meat: it’s what’s NOT for dinner. debate. Science 301:451 Whaling 101 webpage. Greenpeace, available online Marquardt O, Caulfield RA (1996) Development of west at: www.greenpeace.org/usa/campaigns/oceans/whale- Greenland markets for country food since the 18th defenders/whaling-101 century. Arctic 49:107–119 Hay K, Aglukark D, Igutsiaq J, Ikkidluak J, Mike M (2000) Martin AR (1990) Whales and dolphins. Salamander, London Final report of the Inuit bowhead knowledge study. Mills J, Ishihara A, Sakaguchi I, Kang S, Parry-Jones R, Nunavut Wildlife Management Board, Iqaluit, Nunavut. Phipps M (1997) Whale meat trade in east Asia: a review Available online at: www.nwmb.com/english/resources/ of the markets in 1997. TRAFFIC International, Cam- Bowheadreport.pdf bridge Heazle M (2004) Scientific uncertainty and the International Milner-Gulland EJ, Rowcliffe JM (2007) Conservation and Whaling Commission: an alternative perspective on sustainable use: a handbook of techniques. Oxford Uni- the use of science in policy making. Mar Policy 28: 361–374 versity Press, Oxford Heide-Jørgensen MP, Laidre K, Borchers D, Samarra F, Stern Mitchell ED (2004) Counting whales in the North Atlantic. H (2007) Increasing abundance of bowhead whales in Science 303:39 West Greenland. Biol Lett 3:577–580 Moore M (2004) Whither the North Atlantic right whale? Hilborn R (2006) Faith-based fisheries. Fisheries 31:554–555. Oceanus 43:1–5 Available online at: http://www.fisheries.org/afs/docs/ Mrosovsky N (2000) Sustainable use of hawksbill turtles: con- fisheries/fisheries_3111.pdf temporary issues in conservation. Key Centre for Tropical Hutton JM, Leader-Williams N (2003) Sustainable use and Wildlife Management, Darwin incentive-driven conservation: realigning human and con- Mrosovsky N (2004) Predicting extinction: fundamental flaws servation interests. Oryx 37:215–226 in the IUCN’s Red List system exemplified by the case of ICR (Institute of Cetacean Research) (2007) Journalists sea turtles. Available online at: http://members.seaturtle. and editors misinforming the public — again. November org/mrosovsky/ 28, 2008 Press release, The Institute of Cetacean Re- Myers N (1979) The sinking ark: a new look at the problems search, Tokyo. Available online at: http://icrwhale.org/ of disappearing species. Pergamon, Oxford 0711128Release.htm Myers N (1983) A wealth of species. Westview Press, Boulder, Iino Y, Goodman D (2003) Japan’s position in the Interna- CO tional Whaling Commission. In: Burns WCG, Gillespie A Øen EO (2005) Electronic monitoring of Norwegian minke (eds) The future of cetaceans in a changing world. whaling. High North Alliance Fact Sheet, Reine. Transnational Publishers, Ardsley, NY, p 3–32 Available online at: www.highnorth.no/Library/Hunts/ IUCN (International Union for Conservation of Nature and Norway/el-mo-of.htm Natural Resources) (2000) IUCN policy statement on sus- Palsbøll PJ, Bérubé M Larsen, F (2007) Could genetic diver- tainable use of wild living resources. Available online at: sity in eastern North Pacific gray whales reflect global his- www.iucn.org/themes/ssc/susg/docs/valeurs_annex2.pdf toric abundance? Proc Natl Acad Sci 104(52) Available IUCN (2007a) The IUCN Red List of threatened species. online at: www.pnas.org/cgi/content/full/104/52/E2 Available online at: www.iucnredlist.org Perry S, DeMaster DP, Silber G (1999) The right whales. Mark IUCN (2007b) IUCN Red List of threatened species: Physeter Sci 61:7–23 macrocephalus. Available online at: www.iucnredlist.org/ Phipps M, Ishihara A, Kanda N, Suzuki H (1998) A prelimi- search/details.php/41755/summ nary report on DNA sequence analysis of whale meat and IUCN (2007c) Guidelines for using the IUCN Red List whale meat products collected in Japan. TRAFFIC Bul- categories and criteria. Version 6.2. Available online letin 17:91–94 at: http://intranet.iucn.org/webfiles/doc/SSC/RedList/ Pittman C (2001) Baby boom at sea. St. Petersburg Times, RedListGuidelines. pdf March 20, 2001. Available online at: www.sptimes. 184 Endang Species Res: 6:173–184, 2008

com/News/032001/State/Baby_boom_at_sea_.shtml 22:173–187 Roman J, Palumbi SR (2003) Whales before whaling in the Turvey ST, Pitman RL, Taylor BL, Barlow J and others (2007) North Atlantic. Science 301:508–510 First human-caused extinction of a cetacean species? Biol Ross JP (2000) Book reviews. Mar Turtle Newsl 88:28–29 Lett 3:537–540 Sharp R (2007) Book review: Conservation and sustainable Tyack PH (1989) Let’s have less public relations and more use: a handbook of techniques. Sustainable, December ecology. Oceanus 32:103–108 2007:28–29. Available online at: http://intranet.iucn.org/ Vikingsson G, Kapel FO (eds) (2000) Minke whales, harp and webfiles/doc\SSC\Gen_docs\SustainableDecember2007. hooded seals: major predators in the North Atlantic pdf ecosystem. Sci Publ 2, North Atlantic Marine Mammal Small GL (1971) The blue whale. Columbia University Press, Commission, Tromsø New York Ward E (1993) Indigenous peoples between human rights Smith TD, Reeves RR (2006) Pre-20th century whaling: impli- and environmental protection — based on an empirical cations for management in the 21st century. In: Ringstad, study of Greenland. Danish Centre for Human Rights, JE (ed) Whaling and history. II. Kommandør Chr. Chris- Copenhagen tensen Whaling Museum, Sandefjord, p 119–134 Webb JWW, Carrillo EC (2000) Risk of extinction and cate- Tillman MF, Donovan GP (eds) (1983) Special issue on histor- gories of endangerment: perspectives from long-lived rep- ical whaling records. IWC Special Issue 5, International tiles. Popul Ecol 42:11–17 Whaling Commission, Cambridge Young OR, Freeman MMR, Osherenko G, Andersen RR and Trites AW, Christensen V, Pauly D (1997) Competition be- others (1994) Subsistence, sustainability and sea mam- tween fisheries and marine mammals for prey and primary mals: reconstructing the international whaling regime. production in the Pacific Ocean. J Northwest Atl Fish Sci Ocean Coast Manag 23:117–127

Editorial responsibility: Matthew Godfrey, Submitted: December 11, 2007; Accepted: April 30, 2008 Beaufort, North Carolina, USA Proofs received from author(s): August 8, 2008