Sea Otter Population Declines in the Aleutian Archipelago
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Journal of Mammalogy, 84(1):55±64, 2003 SEA OTTER POPULATION DECLINES IN THE ALEUTIAN ARCHIPELAGO ANGELA M. DOROFF,* JAMES A. ESTES,M.TIM TINKER,DOUGLAS M. BURN, AND THOMAS J. EVANS United States Fish and Wildlife Service, Marine Mammals Management, 1011 East Tudor Road, Anchorage, AK 99503, USA (AMD, DMB, TJE) Long Marine Laboratory, Biology Department and Institute of Marine Sciences, United States Geological Survey, University of California, Santa Cruz, CA 95004, USA (JAE, MTT) Sea otter (Enhydra lutris) populations were exploited to near extinction and began to re- cover after the cessation of commercial hunting in 1911. Remnant colonies of sea otters in the Aleutian archipelago were among the ®rst to recover; they continued to increase through the 1980s but declined abruptly during the 1990s. We conducted an aerial survey of the Aleutian archipelago in 2000 and compared results with similar surveys conducted in 1965 and 1992. The number of sea otters counted decreased by 75% between 1965 and 2000; 88% for islands at equilibrial density in 1965. The population decline likely began in the mid-1980s and declined at a rate of 17.5%/year in the 1990s. The minimal population estimate was 8,742 sea otters in 2000. The population declined to a uniformly low density in the archipelago, suggesting a common and geographically widespread cause. These data are in general agreement with the hypothesis of increased predation on sea otters. These data chronicle one of the most widespread and precipitous population declines for a mam- malian carnivore in recorded history. Key words: aerial surveys, Aleutian Islands, Enhydra lutris, population declines, population trends, sea otters, skiff-based surveys Early European explorers reported vast of sea otters after the fur trade occurred in numbers of sea otters (Enhydra lutris)in the Aleutian archipelago. Remnant popula- coastal waters of the Aleutian archipelago tions, located in the central Aleutian Is- and mainland Alaska (Bancroft 1959; Len- lands, were monitored infrequently through sink 1962). Extensive hunting of these an- the 1950s (Kenyon 1969; Lensink 1962; imals for their valuable pelts eliminated the Murie 1959). Systematic aerial surveys of species from most of its historic range, sea otters in the Aleutian Islands were ini- which extended across the Paci®c Rim from tiated in 1957 (Kenyon 1969) in conjunc- Japan to the Paci®c coast of Mexico (Ken- tion with site-speci®c surveys that em- yon 1969; Lensink 1962; Riedman and Es- ployed a variety of techniques (Estes et al. tes 1990; L. Rotterman and T. Simon-Jack- 1978; Kenyon 1969; Lensink 1962). Col- son, in litt.). The International Fur Seal lectively, these efforts chronicle a pattern of Treaty protected the surviving remnant col- population recovery that began in the cen- onies from further harvest beginning in tral Aleutians and gradually spread through- 1911. These colonies subsequently in- out the archipelago. By the 1950s, sea otter creased to repopulate much of the sea ot- numbers apparently had recovered to pre- ters' former range. commercial harvest levels at some islands The earliest and most extensive recovery in the central Aleutians, although the ma- jority of the archipelago was yet to be re- * Correspondent: [email protected] colonized (Estes 1990; Kenyon 1969). By 55 56 JOURNAL OF MAMMALOGY Vol. 84, No. 1 FIG. 1.ÐMap of the study area denoting 6 major island groups (Near, Rat, Delarof, Andreanof, Four Mountains, and Fox islands) in the Aleutian archipelago. the 1980s, an estimated 55,000±74,000 an- surveys conducted at several islands during imals inhabited the archipelago, and contin- the 1990s are used to de®ne decline trajec- ued population growth was expected (D. tories more precisely and provide a minimal Calkins and K. Schneider, in litt.; Estes estimate of bias in the aerial counts. 1990; L. Rotterman and T. Simon-Jackson, in litt.; K. Schneider, in litt.). Sea otters had MATERIALS AND METHODS spread to all island groups by 1992, but the The Aleutian archipelago extends westward overall count at that time had declined by from Unimak Pass to Attu Island (Fig. 1). The approximately 50% in the central Aleutians islands are volcanic in origin, forming a bound- since the 1965 survey (T. Evans et al., in ary between the Bering Sea and the North Pa- litt.). Population trends from a time series ci®c Ocean. There are 6 major island groups of skiff-based counts documented a precip- (Near, Rat, Delarof, Andreanof, Four Mountains, itous decline in sea otter numbers at Adak and Fox) that collectively contain 78 major is- Island during the early to mid-1990s, and lands with .5,000 km of coastline. subsequent surveys at Little Kiska, Am- Aerial surveys.ÐWe conducted aerial surveys chitka, and Kagalaska islands indicated from 1±30 April 1992 and 11±29 April 2000. Viewing conditions were good to excellent dur- similar declines (Estes et al. 1998). These ing both surveys with approximately 90% of ®ndings prompted us to conduct another ae- sightings made in average wind speeds of #19 rial survey of the entire Aleutian archipel- km/h. We surveyed all major islands and off- ago in April 2000 to assess the magnitude shore rocks of the Aleutian archipelago except and geographic extent of the population de- for Chagulak Island in the Islands of Four cline. Mountains, which was omitted because of the We present general patterns of population high risk associated with ¯ying near its large change for sea otters in the Aleutian Islands seabird colony. through compilation of aerial survey data In 1992, the survey aircraft was a deHavilland from the late 1950s to 2000. We evaluated turbine Twin Otter, and in 2000 the survey air- the hypothesis that otter densities varied craft was a turbine Aero Commander 690A. among island groups over time. We analyze Both aircraft were equipped with large bubble windows and long-range fuel tanks. The survey trends in sea otter abundance by comparing crew consisted of a pilot, copilot, 2 observers, the number of animals counted over time and 1 data recorder. Observers sat aft of the pi- and by computing proportional changes be- lots on each side of the plane. The aircrafts were tween time periods. We evaluated the null ¯own approximately 0.23 km from the shoreline hypothesis that there was no difference be- at an altitude of 91 m and an average airspeed tween observed and expected distributions. of 185 km/h during the surveys, sampling an The more detailed results of skiff-based area from shoreline to approximately 0.7 km off- February 2003 DOROFF ET AL.ÐSEA OTTER POPULATION DECLINES 57 shore (T. Evans et al., in litt.). Location, altitude, licates. This analysis was done separately for ae- weather, visibility, and observer identi®cation rial survey data from 1965, 1992, and 2000. were recorded directly into an onboard computer We further evaluated trends in sea otter abun- interfaced with a global positioning system. dance by comparing the number of animals We surveyed the shoreline of each island in counted by island, island group, and across the its entirety. We sampled offshore habitat by sur- archipelago, and by computing proportional veying line transects that were perpendicular to changes between time periods (Nt2/Nt1). To avoid the shoreline and extended to the 50-fathom spurious results, we arbitrarily restricted the lat- (91.4 m) isobath (Burnham et al. 1980; T. Evans ter analyses to islands where 20 or more otters et al., in litt.). Transects were digitized for the were counted during at least 1 aerial survey. entire archipelago and of these, 61 were random- Proportional changes were then plotted as fre- ly sampled in 1992. In 2000, we resurveyed 35 quency distributions and contrasted with expect- transects in the western and central Aleutians. ed distributions for stable populations, assuming Mean encounter rates (otters/km) for the shore- sampling variation but with no prevailing ten- line and transects were compared between 1992 dency toward increase or decline. Expected dis- and 2000 surveys. tributions were computed from a lognormal den- Skiff-based surveys.ÐSkiff-based surveys sity function with a mean of 1 and the observed were conducted several times during the 1990s variance. The null hypothesis of no difference at Adak, Kagalaska, Kiska, Little Kiska, Am- between observed and expected distributions chitka, Shemya (Semichi Islands), and Attu is- was evaluated using a 1-sample K-S test. To lands (Fig. 1) when viewing conditions were avoid the unknown and potentially confounding good to excellent (Beaufort sea state of 1±2, and effects of range expansion and population .1 km of clear visibility at sea level). Two or growth between 1965 and 1992, we repeated the more observers counted sea otters from a 5.2-m analysis for those islands that Kenyon (1969) as- skiff as it was run parallel to shore along the sumed to be at or near equilibrial density in outer margins of kelp (Alaria ®stulosa) beds at 1965 (de®ned here as that which occurred when 15±22 km/h. We counted sea otters with the un- populations ceased growing because of resource aided eye, using binoculars to con®rm sightings limitation). The latter analysis included all the or to count animals in large groups. The shore- Rat and Delarof islands, and the Andreanof Is- line of each island was divided into contiguous lands from Great Sitkin westward (n 5 23 is- segments, each 3±10 km in length and separated lands; Table 1) in which $20 otters were count- by distinctive topographic features (e.g., promi- ed in at least 1 survey period. nent points of land). Counts were recorded sep- We computed independent rates of population arately for each section.