Translator Disclaimer
1 March 2006 Historic Populations of the Double-crested Cormorant (Phalacrocorax auritus): Implications for Conservation and Management in the 21st Century
Author Affiliations +
Abstract

In North America, the Double-crested Cormorant (Phalacrocorax auritus) is currently abundant, widely distributed across five broad geographic regions, and often perceived as overabundant. In many U.S. states and Canadian provinces, policy makers are pressured to significantly reduce cormorant numbers, primarily to minimize conflicts between cormorants and fish resources. Concurrently, large-scale conservation plans recently developed for birds in the Americas depart from the traditional narrow focus on threatened and endangered species to encompass broader and more representative goals (e.g., Partners in Flight’s objective to “keep common birds common”). In recent waterbird conservation initiatives, historic distribution and abundance provide the basis for conservation focus; these initiatives advocate conservation of birds in natural numbers and natural habitats. To provide a context in which current populations of Double-crested Cormorants can be understood, we reviewed historic and current breeding and wintering records to determine historic distribution (pre-1900), current distribution (1970-1999), and extent of range expansion across North America. Early records suggest Double-crested Cormorants were present in large numbers throughout much of their current range; colonies and flocks much larger than any known in the 1990s are well documented. However, numbers sharply declined through the late 1800s as cormorants were greatly reduced and/or extirpated in many areas. The population partially recovered through at least the mid-1900s, but experienced a second major decline during the 1950s-1970s. In the late 1970s, a second rebound began across much of the continent; the largest breeding populations (Canadian/U.S. interior, Atlantic Coast >80% of total) increased from approximately 32,000 pairs in the early 1970s to >226,000 pairs in the late 1990s. Comparison of historic and current records challenges the opinion that cormorants are currently overabundant, and suggests that perception of overabundance rests on socio-political rather than biological or ecological factors. For this species, and others that are seen as competitors with humans, limits of human tolerance (i.e. “social carrying capacity”) are far narrower than those of biological carrying capacity. Because large numbers have been typical for cormorants historically, population targets based on fishery or other objectives derived from human values will likely be readily surpassed, require intensive management, and significantly depart from the concept of conserving birds in natural numbers and natural habitats. Although managing fish-eating birds to benefit fishery yields may increase some fish populations, this approach does not resolve or address the underlying problems causing current fish population declines across the continent, and is in direct conflict with current broad scale conservation initiatives. To ensure inclusion of cormorants and other fish-eating birds in these conservation plans, the avian conservation community must continue to press for programs based on ecosystem health and process that recognize humans, fish and cormorants as three components of a complex system driven by many species and dynamic interactions.

Introduction

In May of 1604, Samuel de Champlain sailed along the southwest coast of Nova Scotia and visited several islands. One, “The Isle of Cormorants”, west of Cape Sable, was “so named because of the infinite number of these birds of whose eggs we took a barrel full” (Champlain 1922). On his visit to the nearby Seal Island group, Champlain wrote “the abundance of birds of different kinds is so great that no one would believe it possible unless he had seen it-such as cormorants . . .” Though Champlain does not identify the species of cormorant he observed, Lewis (1929) concluded some of the birds were likely Double-crested Cormorants (Phalacrocorax auritus) (DCCO); more than 200 years after Champlain’s visit, Audubon (1840-1844) reported Double-crested Cormorants breeding on the Seal Islands.

By the late 19th and early 20th centuries, however, the abundance of cormorants Champlain and others observed across much of North America had greatly diminished. As early as 1634, cormorants were reported “to destroy abundance of small fish” (Wood 1634); since this time, the perception of cormorant diet and foraging behavior has changed little and has been a significant factor affecting the distribution and abundance of the DCCO in North America. Human efforts to reduce numbers and eliminate populations have been documented throughout European settlement, and species history during the latter part of the 19th and the first quarter of the 20th centuries has been described as “a history of persecution and gradual abandonment of one breeding place after another” (Lewis 1929).

Between the 1920s-1940s, DCCOs began a period of population recovery and expansion in several areas. This period was relatively short-lived, however, as widespread use of DDT beginning in the 1940s, combined with legal and illegal control activities and habitat change, led to a second major period of DCCO population declines (e.g., Carter et al. 1995; Hatch 1995; Krohn et al. 1995; Ludwig et al. 1995; Weseloh et al. 1995). Between the late 1940s-1970s, numbers plummeted sharply across much of the species range.

Conversely, history of the species during the last third of the 20th century can be described as one of protection and conservation efforts. In 1972 the DCCO was Blue Listed by National Audubon Society (Tate and Tate 1982) and added to the U.S. Migratory Bird Treaty Act protected bird list (23 U.S.T. 260 (1972)). In the same year, DDT was also banned (U.S. EPA 1972). These actions, along with changes in the prey base (e.g., increases in forage fish in natural waters and development of large-scale aquaculture facilities) contributed to the recent period of spectacular growth in numbers and return of DCCOs to many portions of the historic range from which they had long been absent. Presently, the species is widespread and abundantly distributed with five major breeding regions described: Alaska, the Pacific Coast, Canadian and U.S. Interior, Gulf Coast, and Atlantic Coast (Hatch and Weseloh 1999; Wires et al. 2001).

In this paper, we identify conflicts between federal policy and continental/regional conservation plans that are closely linked to interpretations of recent population growth. Current DCCO numbers are reported to be at “all time” or “historic” highs (USDA/WS Activity/Fact sheet; USFWS News Release for DCCO Proposed Rule, Mar. 2003; Wywialowski 1999), and to have “irrupted” during the last 20 years (Glahn et al. 2000). Though historical information is limited (Hatch 1995; Wires et al. 2001) the U.S. Fish and Wildlife Service (USFWS) Proposed Rule for DCCO (USDI/FWS 2003a) stated, “population levels are greater now than in the past.” Additionally, the species is frequently characterized as “overabundant” (e.g., Farquhar 2001; USDA/APHIS/WS 2003; USFWS Waterbird Fact Sheet, Jan 2002). Growing concern over population changes and possible impacts to natural resources led USFWS, in conjunction with U.S. Dept. of Agriculture/Wildlife Services (USDA/WS), to prepare an Environmental Impact Statement (USDI/FWS 2003b), and publish a Final Rule which establishes a Public Resource Depredation Order for DCCOs effective in 24 states (geared towards Interior, Southeastern and Atlantic coast populations), and revises the 1998 DCCO Aquaculture Depredation Order to include lethal control at winter roost sites (USDI/FWS 2003c). At the same time, large-scale conservation plans for waterbirds in North America and Canada are based on sustaining and/or restoring waterbird populations throughout their historical range (Kushlan et al. 2002; Milko et al. 2003).

To provide a context for the abundance of current (1970-2000) populations we review historic (circa 1900 and earlier) populations of Double-crested Cormorants. Though human modification of the environment was well under way in the 19th century, human induced environmental change greatly accelerated in the 20th century and severely impacted many wildlife species (Askins 1999), including DCCOs (Hatch and Weseloh 1999). Therefore, we examine occurrence during the historic period because populations at this time were more representative of the numbers that existed under relatively “natural” conditions. We explore the concept of “overabundance” as it relates to distribution and abundance of cormorants, perceptions about cormorant population growth and recovery, and biological carrying capacity vs. “wildlife acceptance capacity”. Because establishing population objectives is a goal for major bird conservation plans (e.g., Partners in Flight (Bonney et al. 1999), North American Waterbird Plan (Kushlan et al. 2002), North American Waterfowl Management Plan (NAWMP 1998), U.S. Shorebird Conservation Plan (Brown et al. 2001)), we also discuss factors that have influenced population objectives for cormorants.

Methods

To describe historic (pre-1900) breeding distribution of DCCOs in North America, we reviewed early ornithological and archaeological records reported in journals, state breeding bird atlases, books and maps. All records that reported observations of nests, chicks and/or eggs were considered breeding records. Records that contained circumstantial or anecdotal evidence indicating cormorants may have been breeding were explored and considered probable breeding records based on factors such as long-term occurrence in area, season record was obtained, age of specimen seen, and author’s comments and apparent level of knowledge. Records of breeding obtained post-1900 were included if they indicated the colony site was historic or had likely been occupied prior to 1900. Because most early records did not provide numerical estimates we were unable to estimate historic number of breeding birds on a regional level. However, early records often included qualitative descriptions of abundance and sometimes estimates of numbers of pairs or nests. We carefully examined these descriptions to obtain information on and assess historic abundance.

To describe current (1970-2000) breeding distribution and abundance, we obtained census and survey data from university, state and provincial biologists monitoring DCCOs across the continent. Most of these individuals and agencies regularly census DCCO colonies and they provided us with estimates of breeding pairs.

In an effort to present biologically relevant information for current and historic populations, we report population data in the context of distinct breeding zones. Hatch and Weseloh (1999) described five main breeding zones for DCCOs in North America: Alaska, Pacific Coast, Canadian and U.S. Interior, Florida and the western Caribbean, and the Atlantic Coast, which largely correspond to distribution of the five subspecies (Palmer 1962; Johnsgard 1993). Though thought to reflect fairly distinct breeding populations, recent expansion and re-colonization has blurred boundaries between and among these zones (Hatch and Weseloh 1999; Wires et al. 2001). Border states between Interior and Florida/Caribbean populations, Interior and Pacific Coast populations, and Florida/Caribbean and Atlantic Coast populations (e.g., Texas, New Mexico, Idaho, inland southern states, the Carolinas), qualify as “gray areas” or regions where it is not possible to identify precisely what zone breeding cormorants belong to without banding and molecular studies. To determine the most likely population zone for each colony documented prior to 1900 and between 1970-2000, we relied on subspecies distribution information and consideration of logical geographic units. Latitude/longitude coordinates were obtained for historic and current breeding sites and mapped within each breeding zone. For some historic sites, only general locations (e.g., county, central portion of state) were documented; for these sites, best approximate locations were mapped. Changes in abundance and distribution were examined by comparing current and historic records within each of the breeding zones.

Results

Distribution

Archaeological records.--We did not conduct an exhaustive search for archeological records. Those we report were obtained while searching for early records and are based on skeletal remains retrieved from Native American middens. Though only four records were found, they indicate DCCOs were present in three of five breeding zones (Fig. 1) between 500-5000 years ago. Bones of young birds collected from middens on the Pacific and North Atlantic coasts indicate DCCOs were breeding on both coasts. Skeletal material retrieved from middens on Amchitka, Aleutian Islands, documents presence but does not verify breeding because no bones of young birds were found.

Early ornithological records (1500-1900).--Prior to 1900, cormorants were widely distributed across North America, and occurred in all five of the hypothesized breeding zones (Fig. 1). Approximately 80% of the locations shown are documented breeding sites (i.e., have records of nests, eggs, or young); we designated the remaining 20% as probable or possible breeding locations based on circumstantial or anecdotal information (e.g., time of year observed, author’s comment). Systematic surveys for birds were not formalized at most locations prior to the twentieth century; thus, the number of records within each breeding zone does not reflect abundance of breeding birds or extent of distribution. In some areas (e.g., Pacific and Atlantic coasts), multiple records prior to 1900 are available; in others (e.g. some parts of the Interior and Gulf Coast), little effort was made to survey avifauna prior to the twentieth century (Ridgway 1874).

Current records (1970-2000).--In general, distribution of the current population is similar to that of the historic population (Fig. 1). All records shown are documented nest sites, and, with the exception of states and provinces for which data were not available, represent the continental breeding distribution. Important areas that appear different from historic distribution (Fig. 1) include the Great Lakes and western portion of the Interior range; interior portion of southeastern U.S.; and southeastern Alaska. Comparison of historic and current distribution maps suggests the species currently has a wider distribution in these zones.

Abundance

Historic abundance.--While precise counts are not available for most colonies prior to the twentieth century, records located for each population zone suggest historic populations of DCCO were very large. To indicate relative abundance within each population zone we summarized colony observations and qualitative descriptions for each breeding region (see Zone Summaries below).

Current abundance.--Census efforts among states and provinces are rarely coordinated across breeding zones due to several factors (e.g., limited funding, political boundaries). Therefore, colony size typically is not estimated during the same year or using the same census technique across a population. Additionally, some important areas that provide breeding habitat for large numbers of cormorants (e.g., Mexico, Manitoba) are not regularly censused and little is known about cormorants in major portions of these areas. Thus, accurate estimates for regional (zone-wide) populations are not available. However, most areas within the five breeding zones are regularly censused and estimates of breeding pairs are available for many colony sites. To indicate relative abundance of DCCOs in each population zone we report (Table 2) population estimates for each breeding region based on the most recent and complete census efforts.

Alaska Summary

Birds in this zone comprise the subspecies P.a. cincinatus, the most restricted (Palmer 1962) and smallest of the five populations. Between 1970 and 2000, DCCOs were confirmed nesting at 126 colonies, mostly along the southern coast and on the Aleutian Islands (Fig. 1). Alaska has not conducted statewide censuses of all colonies in the same year and the number of breeding birds is not known. However, 106 sites were censused at least once between 1970 and 2000; most colonies (93%) were small (<100 pairs), and a total of 3,029 pairs was reported (Table 2). Because so few data are available over such a broad time period it is not possible to comment on how closely this estimate approximates actual numbers, but the abundance of small colonies suggests the population is small.

Determining historic distribution and abundance in this zone is complicated by the large-scale introduction of foxes (Alopex lagopus; Vulpes vulpes) to Alaska that occurred between 1750 and the early 1900s (Bailey 1993). Because introduced foxes had major impacts on nesting seabirds (Bailey 1993; Carter 1995), records collected during the historic period may reflect a recently modified distribution and abundance. Turner (1885) described DCCOs as abundant residents and breeders in the Near Islands in the western Aleutians in the late 1800s, and Clark (1911) reported they were still present on these islands as breeders in the early 1900s. However, the peak in fur farming did not occur until the early 1900s, with the most rapid growth occurring in the 1920s (Bailey 1993); by the mid-1930s, DCCOs were no longer breeding in the Near Islands (Murie 1959). Today the western-most breeding point is Chuginidak, Aleutian Islands (52°51’04”, 169°49’69”).

Gabrielson and Lincoln (1959) note that reports of DCCOs breeding throughout southeastern Alaska could not be confirmed with available information. They reviewed early records up to 1945, and defined the breeding distribution “from Kodiak westward at suitable places along the Alaska Peninsula far out into the Aleutian Islands.” All observed colonies were small (<100 pairs).

Based on available information, current DCCO distribution appears more restricted than in the past. In this portion of the range, DCCOs utilize level ground for nesting, which makes colonies vulnerable to terrestrial predators (Siegel-Causey et al. 1991). Introduction of foxes may have limited cormorant distribution, and if large colonies existed, they may have been substantially reduced before they were documented, particularly in the Aleutian Islands and Gulf of Alaska (Siegel-Causey et al. 1991; Carter et al. 1995). Murie (1959) noted a “drastic change” in the Alaska distribution since about 1906. Disappearance from portions of the range combined with increased predation pressure suggests DCCOs may have been more abundant historically, at least in parts of this region, than they are today. Additional causes that may have contributed to declines include other introduced predators and rabbits, and human disturbance (Siegel-Causey et al. 1991; Bailey 1993; Carter et al. 1995).

Pacific Coast Summary

Birds in this zone comprise the subspecies P.a. albociliatus. Between 1970 and 2000, nesting was documented at 248 sites (Table 2). The current breeding population occurs mostly along the coast from southern British Columbia to at least Bird Island, Sinaloa, Mexico (Fig. 2), and possibly further south (Carter et al. 1995). Significant colonies also occur inland. Summing available estimates for the late 1980s and 1990s, approximately 33,000 nesting pairs were documented in the region primarily during the mid-to-late 1990s. However, estimates from important portions of this region are dated, and large-scale movement among breeding birds occurs. In the Salton Sea, dramatic increases in breeding birds occurred during the mid-to-late 1990s, and cannot be attributed to productivity rates alone (D. Shuford, Point Reyes Bird Observatory (PRBO), pers. comm.); most birds were believed to be immigrants from other locations, possibly Mexico (D. Shuford, PRBO, pers. comm.). Because of large shifts and intercolony movement, frequent monitoring efforts over wide areas are required to estimate regional population size (Carter et al. 1995). Although data collection and census efforts have not been coordinated across the region, available information indicates this population is the third largest, but relatively small in comparison to the North Atlantic Coast and Interior populations.

With the exception of British Columbia, the DCCO was recorded as a breeding species throughout most of the Pacific Coast prior to 1900. The earliest breeding records we located were bones of young cormorants retrieved from a shell mound in Emeryville, CA. Samples from the mound indicate that aboriginal people utilized the area as long as 2,550 years ago and hunted a great diversity of waterbirds, including DCCOs (Howard 1929; Sher 1994; Broughton 2004). Although bones of DCCOs are abundant at archaeological sites throughout the Straits of Georgia, British Columbia, indicating the species occupied this area for the past 5,000 years, no skeletal remains of young birds were identified in excavations (Hobson and Driver 1989). Nesting was not documented in the province until 1927 (Munro 1928). However, based on archaeological evidence, Hobson and Driver (1989) suggest that the first documented nesting for this species may actually represent a re-colonization of the area.

Multiple records indicate breeding DCCOs were historically abundant across much of their Pacific Coast range (Squires 1917; Ray 1915; Chamberlin 1895; Willett 1910, 1933; Howell 1917; Grinnell 1908; Finley 1907, 1915; Linton 1907; Lamb and Howell 1913; Goldman 1908; Salvadori 1865; Lamb 1927; Bryant 1889; Bancroft 1927b; Bendire 1877; Dawson 1908, 1911). The largest cormorant colony ever recorded on the continent existed at San Martin Island, Baja California, MX; Wright (1913) estimated 348,840 nests at this site. The estimated size of this colony alone is larger than each of the five current regional populations and may be comparable to current continental population size. Although Wright’s estimate was later considered an overestimate and revised by J. R. Jehl to 213,500 pairs (Hatch 1995) based on the area of the island, the revised estimate still dwarfs the present size of the entire Pacific Coast and much of the North American population. Even if Jehl’s estimate is off by an order of magnitude, no colony approaches this size anywhere in North America.

By the late 1800s and early 1900s, the DCCO had experienced substantial decline and loss of breeding colonies along several portions of its Pacific Coast range. The species was heavily persecuted by humans, and breeding birds were shot at colonies and nests destroyed. In addition, habitat was lost due to agricultural and water developments (Carter et al. 1995). Review of early records suggests that the DCCO was far more abundant in this zone historically than it is today.

Interior Summary

Birds in this zone represent a substantial portion of P.a. auritus’ distribution. Between 1970 and 2000, nesting was reported at 704-754 sites (Table 2). This is the largest, most widespread breeding population; it spans the prairie provinces of Canada, the Canadian and U.S. Great Lakes, and southwestern Quebec. The distribution extends west of Minnesota to southwestern Idaho, and as far south as central Utah and central Colorado. Local breeding also occurs in central Kansas and possibly northern New Mexico. Breeding birds in Idaho were included in this zone based on Burleigh (1972), who reported that 11 specimens collected from different locations in Idaho were intermediate in their characters between P.a. auritus and P.a. albociliatus, but closer to auritus. Birds breeding in the Upper Rio Grande drainage and Middle Pecos River, New Mexico, were also included in this zone, based on plumage characters of P.a. auritus (S. Williams, NM Dept. Game and Fish, pers. comm.). Genetic analyses need to be undertaken to confirm these subspecies designations.

Recent estimates based on complete counts are available for about half the states and provinces where cormorants breed. With the exception of Manitoba, complete counts are available for most of the areas where cormorants nest in large numbers. The largest numbers occur in the Great Lakes and Prairie Provinces. Summing the most recent estimates available for each state and province gives a very rough estimate of 170,000 pairs for the region in the 1990s (Table 2). However, the breeding population is significantly larger than this. Recent estimates are not available for some areas, and several estimates for other areas were based on partial counts. With the exception of Manitoba, DCCOs are not known to breed in very large numbers (several thousands) in most portions of the Interior range where estimates were missing or incomplete (Hatch 1995, Wires et al. 2001). In Manitoba, however, DCCOs do breed in large numbers, particularly on Lake Winnipegosis. The last province-wide survey was conducted in 1979, at which time 22,642 active nests were documented; 9,053 nests were on Lake Winnipegosis, an extremely important area for waterbird nesting (Koonz and Rakowski 1985). The last census of Lake Winnipegosis in 1999 indicated pairs of nesting cormorants had quadrupled since the previous survey 20 years earlier (Table 2). Assuming cormorant numbers across Manitoba increased at the same rate, there may be as many as 90,000 pairs in Manitoba province-wide (W. Koonz, Manitoba Dept. Nat. Res., pers. comm.). Given these gaps, the Interior population in the late 1990s may have been closer to 270,000 pairs but the accuracy of this estimate is unknown.

In many parts of the Interior region, the early breeding history of the DCCO is well known. Pre-1900 records document nesting by the species across most of the region and suggest it had been a long-time and abundant breeder in several areas, particularly in the Prairie Provinces and the mid-western states, where many large colonies were documented. In Minnesota, Hatch (1892) reported DCCOs bred in nearly all parts of the state, and was “occasional to innumerable” depending on how close one was to breeding colonies. At the Minnesota-Iowa border, observers reported “the air is jist black with em’ an they’re nestin’ on the island so yer can’t see it for eggs” (Whitehead 1887). In Missouri, Widmann (1907) reported cormorants breeding in “considerable numbers.” In Ohio, “boatloads” were killed at St. Mary’s Reservoir (Langdon 1878). At Lake of the Woods, Ontario, “great numbers” of young were killed in the late 1700s (Tanner 1994). Lakes Winnipeg and Winnipegosis, Manitoba, have long been recognized for their very large colonies (Seton 1886; Bent 1922). Lake Isle a la Crosse, Saskatchewan, also had a “large” regular breeding colony (Seton 1908). For additional records see Barnes 1890; Cooke 1888; Roberts 1932; Agersborg 1885; Stansbury 1853.

The Great Lakes is the one area within the region where status (e.g., breeder or migrant) and early history of the cormorant are not clear. For example, investigators reported that DCCOs colonized and invaded the Great Lakes in the early 20th century, moving in an easterly direction after the first nesting was reported on western Lake Superior in 1913 (Postupalsky 1978; Weseloh and Collier 1995). However, records and other evidence indicate DCCOs were present in the area prior to the 20th century, suggesting this portion of the cormorant’s North American range was occupied earlier than 1913, and possibly breeding. These records and evidence, described below, fall into the following categories: anecdotal, geographic place names, and general breeding distribution.

In the first category, three anecdotal records are important in presumed colonization history. The earliest reported observation of cormorants breeding at a specific location on the Great Lakes proper is an anecdotal record; Postupalsky (1978) cites a personal communication from Frank Novy that cormorants were breeding in the Great Lakes on western Lake Superior in 1913. No additional information is available about this record. However, because no other records were known at the time of Postupalsky’s publication, this 1913 personal communication is regarded as the first nesting of cormorants in the Great Lakes. Additionally, the order in which the Great Lakes were hypothetically colonized is traced to it (Postupalsky 1978; Weseloh and Collier 1995). The first documented account of cormorants nesting on the Great Lakes proper is reported in Fargo and Van Tyne (1927). This record reports discovery of a small colony (10 nests) in 1926 at Agawa Rocks, Agawa Bay, on the eastern shore of Lake Superior. However, Fargo and Van Tyne noted that local residents told them “the cormorants had nested there for years.” No additional information is available about this colony, but this anecdotal information suggests cormorants had a history of breeding in the area prior to discovery of the colony. A third record (Baillie 1947) from Georgian Bay, Lake Huron, reports that cormorants were nesting in the general vicinity of the Mink Islands in 1919; though this record was never verified, based on other nesting records, Baillie (1947) believed the record was probably accurate. This record places breeding cormorants much farther east into the Great Lakes more than a decade earlier than was hypothesized by Postupalsky (1978) and Weseloh and Collier (1995).

In the second category, geographic place names suggest cormorants were present in the area and recognized by both Native Americans and settlers, but their status as breeders or migrants is not reported. The earliest of these place names is found on a General Surveyors Office Plat Map of Forsyth Township, Michigan. On 4 Aug, 1854, this map was signed and filed in the General Land Office in Detroit, and documents the existence of two lakes, Shag Lake and Little Shag Lake. Shag, a common vernacular name for cormorant, suggests cormorants were present in the area when the map was prepared. These lakes still exist and are located in the upper peninsula in Marquette Co, about 8-16 km from Lake Superior and within 64 km of Lake Michigan, Bay de Noc area, where cormorants are currently abundant. The map survey work was done during the third quarter (July-Sept) of 1844, and further subdivisions of the township were completed during the third and fourth (Oct-Dec) quarters of 1852. Thus cormorants occurred in the area during the summer and or fall months but their status (breeder, migrant) is unknown. The second geographic place name is from Lake Onaping, Ontario, about 64 km north of the North Channel, Georgian Bay area (Bell 1891). At this location a rock forming part of the cliffs on the west side of the lake was named by the Ojibwa “Kakakeshiwishtagwaning”, which means “the cormorant’s head”, and suggests that the Ojibwa were familiar with cormorants in this landscape.

In the third category, the first four editions of the Checklist of North American Birds (1886, 1895, 1910 and 1931) all report the Great Lakes as part of the breeding range of the DCCO but give no specific locations within the lakes. These records indicate cormorants were breeding in the Great Lakes region earlier than 1913, but because specific locales are not provided it is not possible to determine if cormorants nested on the Great Lakes proper, or on nearby inland lakes. An additional record reports capture of a cormorant in June, 1877, at Sandusky Bay, OH (Wheaton 1882). No other information is available about this record, but the presence of a cormorant at this time of year suggests the bird may have been breeding.

As occurred over much of the continental range, DCCOs in the interior underwent substantial declines in the late 1800s and early 1900s. Persistent human persecution and habitat degradation resulted in colony declines, abandonment and local extirpation from several states (e.g., South Dakota, Indiana, Ohio, Missouri, Iowa, Illinois, Arkansas) (Lewis 1929). Because so few estimates are available pre-1900 and data vary greatly among states and provinces, it is not possible to determine if cormorants were more abundant historically in this zone than they are today. For some areas (e.g., Indiana, Ohio, Iowa, Arkansas, Illinois, Minnesota, Missouri) available data suggest breeding cormorants were more abundant historically. Additionally, very large migrant flocks were observed that were much larger than those currently known to pass through the region. In Minnesota, a flock of migrating cormorants was described that was “four miles long and one-and-a-half miles wide” (Sennett 1891). Though numbers declined throughout much of the Interior by the turn of the century, large numbers were still occasionally observed; in 1926, a flock estimated at 100,000 to one million individuals was described migrating up the Mississippi River past LaCrosse, and was so large “that at times it was impossible to see the sunset sky through the mass” (Grassett 1926). During recent years (1970-2000), no flocks anywhere near this size have been reported.

In other areas of this zone (e.g., the Great Lakes), current numbers are much greater than they were in the late 1800s-early 1900s, and are the highest recorded in the history of the region. In the Prairie Provinces, cormorants were historically and are currently abundant, but it is not possible to determine if abundance has greatly changed.

Southeast U.S. and Caribbean

Birds breeding along the Gulf Coast, the south Atlantic Coast, and in the Caribbean comprise the subspecies P.a. floridanus, while birds breeding in the Bahamas and Cuba are classified as P.a. heuretus (Hatch and Weseloh 1999). This breeding zone extends from southern and central Texas to Florida, and along the Atlantic Coast through Florida, Georgia, and the Carolinas. Birds breeding in southern Texas and southwestern Arkansas were included in this zone based on subspecies distribution maps (Palmer 1962; Johnsgard 1993). Because North Carolina has been reported as the northern breeding limit for P.a. floridanus (Audubon 1840-1844; Bent 1922; Johnsgard 1993), and other authors have suggested that Carolina birds may comprise this subspecies (Palmer 1962; Clapp and Buckley 1984; Post and Post 1988), we included birds breeding in the Carolinas as part of the Southeastern U.S./Caribbean zone. Recent estimates based on complete counts are available for most of the known colonies, with the exception of southern Texas, the Yucatan and the Caribbean. The largest known numbers occur in Florida. Summing estimates available for the region in the 1990s gives an approximate estimate of 9,400 pairs at >108 sites (Table 2).

Prior to the 20th century, the DCCO was “constantly resident in the Floridas and their Keys, and along the coast to Texas” (Audubon 1840-1844). Breeding was recorded in North Carolina as early as the 18th century; at that time cormorants were reported to “lay their eggs in . . . the Islands, in the Sound and near the Sea Shoar in the Banks, and sometimes on high trees, as the Shags do” (Brickell 1737). Several records document breeding along both the Gulf and Atlantic coasts (Table 1).

Although no quantitative data were located to provide a comparison between current and historic abundance, descriptions suggest breeding cormorants were more abundant in this zone prior to the 20th century than they are today. Presently, breeding cormorants are only abundant in Florida (Wires et al. 2001), while historically they were very numerous in multiple areas of the region. Audubon (1843) reported “many thousands” breeding in the Florida Keys in 1832; Brickell (1737) stated that in North Carolina, they were “as numerous all over these Parts of America as in any part of the World”; Audubon (1843) and Beyer et al. (1907) described cormorants as “abundant” in the Louisiana interior and on the coast; and Howell (1911) reported this species as “formerly abundant in the rivers and swamps of eastern Arkansas”. Additionally, based on Audubon’s (1840-1844) description, DCCOs may have been present as breeders along the Alabama, Mississippi and Texas coasts.

Northeast Atlantic Coast

Birds in this zone are also placed in the subspecies P.a. auritus, and breed along the Atlantic Coast from southern Newfoundland, the northern shore of the Gulf and estuary of the St. Lawrence River, Anticosti Island, Magdalen Island, south along the coast to New York City and Long Island (Hatch and Weseloh 1999). The range is expanding south with recent breeding and re-colonization in New Jersey, Delaware, Maryland and Virginia, as far south as Hopewell. Recent estimates based on complete counts are available for nearly all colonies known to be active in this region, with the exception of New Brunswick (Table 2). The largest numbers occur in Quebec in the St. Lawrence River, Estuary and Gulf, along the coast of Maine, and in the Maritime Provinces. Summing estimates available for the region in the 1990s gives an approximate estimate of 87,000 breeding pairs at 381-382 sites. This number should be interpreted cautiously because intercolony movement is common and the size of individual colonies may change dramatically from year to year in substantial portions of the region (J. F. Rail, Canadian Wildlife Service (CWS), pers. comm.). Additionally, a culling program in the St. Lawrence River in 1989 significantly reduced numbers in Quebec in the 1990s.

Archaeological records document cormorants as a breeding species in this zone from the late prehistoric period (about 1500 AD) (Luedtke 1980). Several historic records report breeding from the Straits of Belle Isle, northwest coast of the Island of Newfoundland, south to as far as Boston Harbor, Massachusettes (Table 1). The historic breeding distribution may have extended further south; Mendall (1936) notes that Williams’ (1643) observations of Native American use of cormorants for food referred either to Massachusetts or Rhode Island or both. Mackay (1894) noted that in the late 1800s, hundreds were observed every spring around Newport County, Rhode Island, at a rock outcrop near Sachuest Point called Cormorant Rocks, which was named on a map as early as 1776. Cormorants were also present historically in Virginia but seasonal presence and distribution is not clear (Lewis 1929).

As in the other breeding zones, sometime during European settlement DCCOs began declining in most parts of the Atlantic range due to heavy human persecution (Lewis 1929; Mendall 1936; Gross 1944; Krohn et al. 1995). By the early 19th century, they were extirpated in New England. By the late 19th and early 20th centuries, breeding cormorants had disappeared from Nova Scotia and were greatly reduced in the other Maritime Provinces and Quebec (Lewis 1929; Erskine 1992). Although we found no early estimates or archaeological evidence to which historic and current abundance can quantitatively be compared, early comments and records suggest cormorants were very abundant in this breeding zone prior to European settlement (Williams 1643; Audubon 1843; Reeks 1869; Champlain 1922; Goss 1889; Townsend 1917). Mendall (1936) cites Morton (1637), who wrote of cormorants in New England, “There are greate store of Pilchers: at Michelmas, in many places, I have seene the Cormorants in length 3.0 Miles feedinge upon the Sent.” Lewis (1929) reported that several 17th century references mention the abundance of cormorants in Virginia, and that in 1610, one anonymous author reported cormorants among the birds in Virginia rivers “in such abundance as are not in all the world to be equaled.”

Discussion

The current breeding range of the DCCO is similar to pre-1900, although some changes have occurred. The range has retracted in Alaska and in southeastern U.S, but expanded along the Atlantic Coast and in the Interior. In the latter regions, some important changes may represent re-colonization events. DCCO historic abundance and widespread distribution has several implications for current conservation and management efforts, discussed below.

The Great Lakes Population

The Great Lakes region is one of the most important areas in the human-cormorant conflict, and concerns from this region have strongly influenced federal policy decisions for cormorants. Most of the conflict centers on perceived impacts to fisheries from cormorant predation. Currently many factors affect the health of the Great Lakes ecosystem; one of the most significant is establishment of >160 exotic invasive species (Mills et al. 1993; Ricciardi 2001). Of these, several (e.g., zebra mussel (Dreissena polymorpha) alewife (Alosa pseudoharengus), sea lamprey (Petromyzon marinus) directly and indirectly impact valuable fisheries (e.g., Mills et al. 1993; Belyea et al. 1999). Additionally, many other variables (e.g., water temperature variation, declines in benthos abundance) also impact fisheries (Hoyle et al. 1998, Pothoven et al. 2001). Nevertheless, in areas where declines in sport fish species and catch have occurred, many anglers and fisheries biologists attribute declines to recent increases in DCCO numbers (e.g., Fielder 2004). Additionally, some anglers believe cormorants are non-native to the region, and report that the species was intentionally introduced by the Japanese (F. Cuthbert, pers. obs.). The media has also published articles stating the species is an exotic (Sharp 2004). Finally, large cormorant colonies in this region have once again become the focus of vandalism and destruction as well as legal control activities. For example, in the late 1990s-2000, about 1,350 DCCO adults and chicks were illegally shot at Little Galloo Island, Lake Ontario, New York, and at Little Charity Island, Saginaw Bay, Michigan, by anglers who believed cormorants were responsible for poor catches (Kloor 1999; Lounsbury 2000). In summer 2004, all colonies in the Les Cheneaux Islands area, Michigan, were targeted by USDA/WS for control (e.g., harassment, oiling eggs, shooting adults) to aid the ailing yellow perch fishery (USDA/APHIS/WS 2004), despite an earlier rigorous study of cormorant predation in this area that concluded cormorants were not a substantial factor in yellow perch mortality (Belyea et al. 1999).

One critical component that shapes perception of a species’ role within an ecosystem is knowledge of species history. Previously, occurrence and movement of cormorants through the Great Lakes in the early 1900s was described as an “invasion” (Postupalsky 1978; Weseloh and Collier 1995). Perception of this species as an invader will strongly influence its management because invasive species, variously described as “exotics”, “nonindigenous” or “aliens,” typically spread and cause net harm to a system (Lodge and Shrader-Frechette 2003). Review of all available records to assess the history of the Double-crested Cormorant in the Great Lakes suggests characterization as an “exotic” or “invasive” is inappropriate, and history in the Great Lakes needs further exploration. Considering records documenting nesting prior to 1900 are available from the western and southern border states of the Great Lakes, and abundant nesting habitat would have been available, apparent species absence is puzzling.

Historic Populations and Current “Overabundance”

Most historic records cited in this work were obtained during the period of European settlement, particularly in the 19th century (Table 1). We assume abundances that existed during this period represent populations under relatively natural conditions (i.e., abundance attained when relatively free from artificial/human modified elements that could lead to inflated or diminished numbers). However, it is possible that in some locations settlers were encountering spectacularly abundant concentrations of cormorants and other wildlife that were responding to release from Native American harvest. Broughton (2004) suggests that the abundant wildlife populations observed in pre-1900 California resulted after Native Americans experienced dramatic disease-based population declines through coastal contact with European explorers in the 16th century. Thus, for some species, assuming that the pre-1900 time period represents the baseline or benchmark of past populations, may be a questionable assumption (Broughton 2004). Nevertheless, although the impact of Native American harvest on cormorants at a large scale (e.g., state, regional, continental) is not known, the breadth of cormorant records spanning hundreds of years and locations across the continent (Table 1) suggests the Double-crested Cormorant had been an abundant species for a substantial time. Additionally, the clearly documented record of human persecution throughout the European settlement period marks a significant era in the known history of the cormorant, to which massive population declines and local extirpations can be traced.

On a continental level, comparisons of historic and current records indicate cormorants were likely more abundant during initial European settlement than they are today. Therefore, describing the current continental population at an “all time” or “historic” high is misleading. Additionally, in several areas across the continent, portions of the public and some natural resource biologists view the DCCO as a recent addition to the regional avifauna despite historic records that demonstrate new nest records actually represent re-colonization of former breeding areas (Wires et al. 2001). Lack of knowledge about historical distribution and abundance has led to significant misconceptions about numbers of cormorants the environment may support. For example, some portions of the public in conflict with this species (e.g., anglers, aquaculturalists) consider mid-century (pre-1980) numbers to be the “norm” for DCCOs; however, numbers and distribution during this period were greatly diminished due to contaminants and persecution (e.g., Gress et al. 1973; Vermeer and Rankin 1984; Weseloh et al. 1995). Similarly, in the Great Lakes, where cormorants are currently at recorded highs, growth during much of the 20th century was suppressed through legal and illegal control activities and contaminants (Ludwig and Summer 1995; Weseloh et al. 1995). Thus population size prior to the 1970s (when national legislation protecting cormorants and reducing contaminants was introduced) was smaller due to human intervention and environmental degradation, rather than to natural regulation and biological carrying capacity. Misconceptions about historic numbers and what is “normal” in current ecosystems, rather than science, has fueled much of the debate about need to control DCCOs in North America.

The term “overabundant” is a judgment that needs qualification. Kushlan et al. (2002) imply one way to judge overabundance is to determine if a species has exceeded historical norms. This criterion is useful because comparison of current to historic numbers may help determine biologically if there are “too many” individuals of a species. Based on available records, the continental DCCO population has not exceeded historical norms. Additionally, no study has demonstrated DCCOs exceed biological carrying capacity by surpassing food supplies or habitat. Although concern about potential impacts to various resources has been reported in several areas (USDI/FWS 2003b), negative changes in natural resources directly linked to DCCO population growth via rigorous scientific study have been documented at only a few locations (Wires et al. 2001; Cuthbert et al. 2002).

In biological terms, we suggest characterization of the continental DCCO population as overabundant is inappropriate. This characterization is applicable, at best, only on a local and site-specific level where numbers can be demonstrated to outstrip resources. Applied to the eastern U.S. population, use of the term “overabundant” is meaningful only in terms of “wildlife acceptance capacity” (Decker and Purdy 1988). This term was created to highlight that many wildlife or social acceptance capacity levels may exist for a particular wildlife population at any point in time; this is unlike the concept of biological carrying capacity. Usually, wildlife acceptance capacity is far smaller and more variable than that determined by biological carrying capacity.

Conservation vs. Management Goals

Over the last decade and a half, large-scale conservation plans for birds in the Americas have developed that depart from the traditional narrow focus on threatened and endangered species, and now encompass broader and more representative goals, such as that defined by the vision of Partner’s in Flight’s (PIF) “keep common birds common” (Fitzpatrick 2002). Waterbird conservation plans at national and continental scales directly link species and population goals to historic distribution and abundance (Milko et al. 2003; Kushlan et al. 2002). Fitzpatrick (2002) takes current bird conservation initiatives one step further and offers the following as a robust mission statement: “Ensure persistence of all American bird populations in their natural numbers, natural habitats, and natural geographic ranges . . .” In these more recent initiatives and perspectives, historic distribution and abundance provide the basis for conservation focus. Thus, when designing management and conservation plans, and developing biologically meaningful population objectives, it is important to recognize where and in what numbers and habitats DCCOs formerly and naturally occurred. Because very large numbers have been typical for this species historically, population targets based on fishery objectives or other human imposed values may be readily surpassed and likely will require intensive management.

In some areas, return of DCCOs after a long period of absence or local/regional rarity has created conflicts over resources because humans are not accustomed to dense concentrations of DCCOs and now more intensively use areas that were formerly occupied by cormorants (Wires et al. 2001; Wires and Cuthbert 2003). These conflicts have led to policy decisions and management plans (Glahn et al. 2000; USDI/FWS 2003c; USDA/APHIS-WS 2004) that are incompatible with current conservation goals (Wires and Cuthbert 2001; Fitzpatrick 2002; Kushlan et al. 2002; Milko et al. 2003). For example, the management plan prepared for DCCOs to minimize damage to southern aquaculture (Glahn et al. 2000) proposes that southern breeding colonies be managed at levels compatible with aquaculture to forestall future depredation problems. Cormorants have recolonized portions of Arkansas and Mississippi after several decades of absence from these states; current colonies are small and few in number (Table 2). Despite no record of cormorant breeding in Arkansas since 1951, most (85%) of the birds at the only known nesting site were collected for research by USDA/APHIS-WS when they resumed breeding in 1999 (Hutchinson 1999). The Arkansas Wildlife Services state director commented that he hoped “we can keep this population down and maybe eliminate it . . . if we let it get out of control we could have cormorants throughout the year” (The Catfish Journal 1999; Hutchinson 1999). Though year-round residence in Arkansas by DCCOs may result in more conflicts with the aquaculture industry, year-round residence was documented historically (Howell 1911; James and Neal 1986).

In spring 2004, several islands in the Great Lakes were selected for large-scale cormorant control but these islands have also been identified as providing significant habitat for birds. For example, the islands in northern Lake Huron, MI, which include the Les Cheneaux Islands, have been proposed as an Important Bird Area (J. P. Cecil, Audubon, pers. comm.), and several of these islands with nesting cormorants were previously prioritized for conservation because of their regional importance to breeding colonial waterbirds (Wires and Cuthbert 2001). At Presqu’ile Provincial Park in Ontario, a formally designated Important Bird Area, over 6,000 breeding cormorants were killed in 2004, allegedly to protect an old-growth forest (Ontario Parks 2005).

Conservation of DCCOs and other abundant waterbird species cannot be effective without a unified conservation philosophy based on knowledge of historical populations and policy decisions founded on sound science. Public dissatisfaction with waterbirds is one of the greatest threats affecting long-term survival of waterbird populations (Kushlan et al. 2002). Much of this negative attitude is held because waterbirds eat fish and are perceived as competitors. In the case of the DCCO, its image is especially negative; the human-cormorant conflict is exacerbated because the species was not present or existed in very low numbers for all or most of current human memory in North America.

The fact that very large numbers of DCCOs existed prior to European settlement has fundamental implications for multiple natural resource management issues. Historically, vigorous fish populations sustained large numbers of fish-eating birds; in relatively natural systems, this is still possible (e.g., Davoren and Montevecchi 2003). In highly modified systems (e.g., Great Lakes), controlling native predators does not address the diverse problems that affect this ecosystem and its ailing fisheries. However, in many areas of the Great Lakes, DCCOs have been identified as the major factor limiting fish populations. This, in turn, has led to the development of population objectives for cormorants based largely on fishery objectives. For example, agencies managing cormorants in the U.S. eastern basin of Lake Ontario and on Long Island (Oneida Lake) (New York Department EC), and Canadian colonies on lakes Huron and Ontario (Ontario Ministry of Natural Resources) have established cormorant population sizes hypothesized to increase local fisheries or to achieve local fishery yields (Farquhar et al. 2001, 2004; Council of Lake Committees 2003; NYSDEC Newsletter 2004).

Although managing cormorants may benefit some fisheries and resolve human-cormorant conflicts, setting population objectives for cormorants based entirely on fishery or other objectives derived from human values significantly departs from the concept of conserving birds in natural numbers and natural habitats (Fitzpatrick 2002). Given the global state of ailing fisheries and increasing government sensitivity to wildlife acceptance capacity, self-regulating populations of cormorants may only be possible in cormorant “safe zones” where human interests are not allowed to influence cormorant numbers. Because few such “safe zones” are likely to exist, those committed to the conservation of fish-eating birds should vigorously oppose strategies that set population objectives based entirely on human interests rather than species biology and regional ecology. Finally, we urge the avian conservation community to support broad conservation strategies based on ecosystem health and process that recognize humans, fish and cormorants as three components of a complex system driven by many species and dynamic interactions.

Acknowledgments

The work of H. F. Lewis and H. L. Mendall brought together much of the historic literature on the distribution of Double-crested Cormorants in North America. Without their efforts our work may not have been possible. Many people, too numerous to mention individually, contributed unpublished data on current cormorant breeding numbers and provided key historic citations. We thank them and appreciate their willingness to help develop the continental picture of cormorant distribution and abundance. We are especially grateful to the following people: Dale Trexel helped find many obscure references to cormorants across the continent and Ron Dietz found and helped us obtain the historic maps showing Shag and Little Shag lakes in Michigan. D. V. “Chip” Weseloh provided substantial review of this manuscript. We also thank Douglas Causey for his comments on the manuscript. During discussion at an AOU banquet, David Blockstein and Robert Askins motivated this study by urging us to search the pre-1800 ornithological literature. Finally, we are indebted to the librarians in the Fisheries, Wildlife and Entomology Library at the University of Minnesota; they helped obtain most of the historic literature and were always helpful and generous with their time.

Literature Cited

  1. G. S. Agersborg 1885. The birds of southeastern Dakota. Auk 2:276–289. Google Scholar
  2. American Ornithologists’ Union 1886. The code of nomenclature and check-list of North American birds adopted by the American Ornithologists’ Union; being the report of the committee of the Union on classification and nomenclature. American Ornithologists’ Union, New York. Google Scholar
  3. American Ornithologists’ Union 1895. Check-list of North American birds. 2nd and rev. ed. American Ornithologists’ Union, New York. Google Scholar
  4. American Ornithologists’ Union 1910. Check-list of North American birds. Prepared by a committee of the American Ornithologists’ Union. 3rd ed. (rev.). American Ornithologists’ Union, New York. Google Scholar
  5. American Ornithologists’ Union 1931. Check-list of North American birds. Prepared by a committee of the American Ornithologists’ Union. 4th ed., constituting the “Systema avium” for North America north of Mexico. American Ornithologists’ Union, Lancaster, PA. Google Scholar
  6. J. J. Audubon 1840-1844. The Birds of America. Vol. 6. Dover Publications (Facsimile reprint, 1967), New York. Google Scholar
  7. E. P. Bailey 1993. Introduction of foxes to Alaskan Islands--history, effects on avifauna, and eradication. U.S. Department of the Interior, Fish and Wildlife Service, Resources Publication 193, Washington, DC. Google Scholar
  8. J. L. Baillie 1947. The Double-crested Cormorant nesting in Ontario. Canadian Field-Naturalist 61:119–126. Google Scholar
  9. S. F. Baird 1859. Notes on a collection of birds made by Mr. John Xantus, at Cape St. Lucas, Lower California, and now in the museum of the Smithsonian Institution. Proceedings of the Academy of Natural Sciences of Philadelphia. pp. 299-306. Google Scholar
  10. G. Bancroft 1927a. Breeding birds of Scammons Lagoon, Lower California. Condor 29:40. Google Scholar
  11. G. Bancroft 1927b. Notes on the breeding coastal and insular birds of central Lower California. Condor 29:193. Google Scholar
  12. R. M. Barnes 1890. List of birds breeding in Marshall County, IL. Ornithologist and Oologist 15:8113–116. Google Scholar
  13. R. B. Bell 1891. Report on the Sudbury mining district, to accompany sheet 130, series of geologically coloured maps of Ontario. S.E. Dawson, Ottawa. Google Scholar
  14. G. Y. Belyea, S. L. Maruca, J. S. Diana, P. J. Schneeberger, S. J. Scott, R. D. Clark, and J. P. Ludwig . 1999. Impact of Double-crested Cormorant predation on the yellow perch population in Les Cheneaux islands of Michigan. Pages 47-59 in M. E. Tobin [Ed.] Symposium on Double-crested Cormorants: population status and management issues in the Midwest. USDA/APHIS Technical Bulletin 1879. Google Scholar
  15. C. E. Bendire 1877. Notes on seventy-nine species of birds observed in the neighborhood of Camp Harney, Oregon, compiled from the correspondence of Capt. Charles Bendire, 1st Cavalry USA. Proceedings of the Boston Society of Natural History 18:153–168. Google Scholar
  16. C. E. Bendire 1878. Notes on some of the birds found in southeastern Oregon, particularly in the vicinity of Camp Harney, from November 1874, to January, 1877. Proceedings of the Boston Society of Natural History 19:109–149. Google Scholar
  17. A. C. Bent 1922. Life Histories of North American Petrels and Pelicans and their allies. Dover Publications, Inc., New York. Google Scholar
  18. G. E. Beyer, A. Allison, and H. H. Kopman . 1907. List of the birds of Louisiana. Auk 24:314–321. Google Scholar
  19. R. Bonney, D. N. Pashley, R. Cooper, and L. Niles . 1999. Strategies for bird conservation: the Partners in Flight planning process. Cornell Lab of Ornithology.  http://birds.cornell.edu/pifcapemayGoogle Scholar
  20. W. Brewster 1902. Birds of the Cape Region of Lower California. Bulletin of the Museum of Comparative Zoology 41:136. Google Scholar
  21. J. Brickell 1737. The Natural History of North Carolina. Printed by James Carson. Dublin. 408 pp. (pp. 211-212 of 1910 reprint. Also Plate facing p. 171 of reprint). Google Scholar
  22. J. M. Broughton 2004. Prehistoric human impacts on California birds: evidence from the Emeryville Shellmound avifauna. Ornithological Monographs No. 56. The American Ornithologists’ Union. Washington, DC. Google Scholar
  23. S. Brown, C. Hickey, B. Harrington, and R. Gill . 2001. The U.S. Shorebird Conservation Plan. 2nd ed. Manomet Center for Conservation Sciences, Manomet, MA. Google Scholar
  24. W. E. Bryant 1888. Birds and eggs from the Farallon Islands. Proceedings of the California Academy of Sciences 1:25–50. Google Scholar
  25. W. E. Bryant 1889. A catalogue of the birds of Lower California, Mexico. Proceedings of the California Academy of Sciences 2:237–320. Google Scholar
  26. BurleighT. D. 1972. Birds of Idaho. The Caxton Printers, Ltd., Caldwell, Idaho. Google Scholar
  27. C. F. Carr 1890. A list of the birds known to nest within the boundaries of Wisconsin, with a few notes thereon. Wisconsin Naturalist 1:28–31. Google Scholar
  28. H. R. Carter, A. L. Sowls, M. S. Rodway, U. W. Wilson, R. W. Lowe, G. J. McChesney, F. Gress, and D. W. Anderson . 1995. Population size, trends and conservation problems of the Double-crested Cormorant on the Pacific Coast of North America. Colonial Waterbirds 18:Special Publication 1189–215. Google Scholar
  29. H. R. Carter, G. J. McChesney, J. E. Takekawa, L. K. Ochikubo, D. L. Whitworth, T. W. Keeney, W. R. McIver, and C. S. Strong . 1996. Population monitoring of seabirds in California: 1993-1995 aerial photographic surveys of breeding colonies of Common Murres, Brandt’s Cormorants, and Double-crested Cormorants. Unpubl. Final Report, U.S. Geological Survey, California Science Center, Dixon, CA. Google Scholar
  30. H. R. Carter, P. J. Capitolo, G. J. McChesney, W. R. McIver, and J. E. Takekawa . 2000. Population monitoring of seabirds in California: colony and subcolony databases for 1985-1995 surveys of breeding colonies of Common Murres, Brandt’s Cormorants, and Double-crested Cormorants between Point Arena and Point Conception. Unpubl. Report, U.S. Geological Survey, Western Ecological Research Center, Dixon, California; Humboldt State University, Department of Wildlife, Arcata, California; and U.S. Fish and Wildlife Service, San Francisco Bay National Wildlife Refuge Complex, Newark, CA. Google Scholar
  31. C. Chamberlin 1895. An inland rookery of Phalacrocorax d. albociliatus. Nidologist 3:29–31. Google Scholar
  32. S. D. Champlain 1922. The works of Samuel de Champlain. H. P. Biggar, Ed. Toronto, 1922. pp. 240-241. Ibid., p. 243. Google Scholar
  33. G. Chapdelaine and J. Bédard . 1995. Recent changes in the abundance and distribution of Double-crested Cormorants in the St. Lawrence River, Estuary and Gulf, Québec. 1978-1990. Colonial Waterbirds 18:Special Publication 170–77. Google Scholar
  34. F. M. Chapman 1902. Double-crested Cormorants. Bird-Lore 4:146. [Photograph]. Google Scholar
  35. R. B. Clapp and P. A. Buckley . 1984. Status and conservation of marine birds in the southeastern United States. In Status and conservation of the seabirds of the world. J. P. Croxall, P. G. H. Evans and R. W. Schreiber, Eds. I.C.B.P. Publication No. 2. The University Press, Cambridge, UK. Google Scholar
  36. A. H. Clark 1911. The birds collected and observed during the cruise of the US Fisheries Steamer “Albatross” in the north Pacific Ocean, and in the Bering, Okhotsk, Japan, and Eastern seas, from April to December, 1906. Proceedings of the U.S. National Museum 38 25–74. Google Scholar
  37. W. W. Cooke 1888. Report on bird migration in the Mississippi Valley. U.S. Department of Agriculture, Division of Economic Ornithology, Bulletin 2, Washington. p. 59. Google Scholar
  38. E. Coues 1861. Notes on the ornithology of Labrador. Proceedings of the Academy of Natural Sciences of Philadelphia. pp. 215-257. Google Scholar
  39. Council of Lake Committees 2003. Summary notes: 17: Results of a 5-year Ontario study on cormorants. (J. Casselman, Ontario Ministry of Natural Resources). http://www.glfc.org/boardcomm/clc/2003Octminutes.pdf. Google Scholar
  40. F. J. Cuthbert, L. R. Wires, and J. E. McKearnan . 2002. Potential impacts of nesting Double-crested Cormorants on Great Blue Herons and Black-crowned Night-Herons in the U.S. Great Lakes region. Journal of Great Lakes Research 28:2145–154. Google Scholar
  41. F. J. Cuthbert, J. E. McKearnan, L. R. Wires, and A. R. Joshi . 2003. Distribution and abundance of colonial waterbirds in the U.S. Great Lakes: 1997-1999. Draft report to U.S. Fish and Wildlife Service. Fort Snelling, MN. Google Scholar
  42. G. K. Davoren and W. A. Montevecchi . 2003. Consequences of foraging trip duration on provisioning behaviour and fledging condition of Common Murres (Uria aalge). Journal of Avian Biology 34:44–53. Google Scholar
  43. W. L. Dawson 1908. The bird colonies of the Olympiades. Auk 25:153–166. Google Scholar
  44. W. L. Dawson 1911. Another fortnight on the Farallones. Condor 13:177. Google Scholar
  45. D. J. Decker and K. G. Purdy . 1988. Toward a concept of wildlife acceptance capacity in wildlife management. Wildlife Society Bulletin 16:53–57. Google Scholar
  46. J. E. Ducey 2000. Birds of the untamed West: the history of birdlife in Nebraska, 1750 to 1875. Making History, Omaha, NE. Google Scholar
  47. B. L. Ehresman 1996. Double-crested Cormorant. In L. S. Jackson, C. A. Thompson and J. J. Dinsmore. The Iowa Breeding Bird Atlas. 1996. University of Iowa Press, Iowa City. Google Scholar
  48. A. J. Erskine 1992. Atlas of breeding birds of the Maritime Provinces. Nimbus Publishing Limited and the Nova Scotia Museum, Halifax. Google Scholar
  49. W. G. Fargo and J. Van Tyne . 1927. Fall bird notes from the eastern shore of Lake Superior. Canadian Field-Naturalist 41:7–9. Google Scholar
  50. J. F. Farquhar 2001. Balancing act: managing cormorants in upstate New York. New York State Conservationist 56:126–28. Google Scholar
  51. J. F. Farquhar, R. D. McCullough, and I. M. Mazzochi . 2001. Cormorant management activities in Lake Ontario’s Eastern Basin. NYSDEC Special Report-March 1, 2001. https://www.dec.state.ny.us/website/dfwmr/cormorant/corm01sec2.pdf Last accessed 09/18/2004. Google Scholar
  52. J. F. Farquhar, I. M. Mazzochi, R. D. McCullough, and J. H. Johnson . 2004. Managing cormorant abundance to meet waterbird and fisheries objectives: an evaluation of a five-year program, Lake Ontario, NY: 1999-2003. Abstract from “Symposium on the biology and management of Double-crested Cormorants”. Quebec City, August 2004. Google Scholar
  53. D. G. Fielder 2004. Collapse of the yellow perch fishery in Les Cheneaux Islands, Lake Huron and possible causes. Pages 129-130 in Proceeding of Percis III: The Third International Percid Fish Symposium (T. P. Barry and J. A. Malison, Eds.). University of Wisconsin Sea Grant Institute, Madison, WI. Google Scholar
  54. W. L. Finley 1902. Among the seabirds of the Oregon coast. Condor 4:53–57. Google Scholar
  55. W. L. Finley 1905. Among the seabirds off the Oregon coast. Condor 7:119–127.161–169. Google Scholar
  56. W. L. Finley 1907. Among the Pelicans. Condor 9:36. Google Scholar
  57. W. L. Finley 1912. Reports of Field Agents. Report of William L. Finley, field agent for the Pacific Coast states. Bird-Lore 14:414–418. Google Scholar
  58. W. L. Finley 1915. Cruising the Klamath. Bird-Lore 17:485–491. Google Scholar
  59. J. W. Fitzpatrick 2002. The AOU and bird conservation: recommitment to the revolution. Auk 119–907–913. Google Scholar
  60. I. N. Gabrielson and F. C. Lincoln . 1959. The birds of Alaska. Stockpole Company, Harrisburg, Pennsylvania, and Wildlife Management Institute, Washington, DC. Google Scholar
  61. General Surveyors Office Plat Map 1854. Records of the Department of Natural Resources, Lands Division, RG87-157. State Archives of Michigan, Lansing. http://www.dnr.state.mi.us/spatialdatalibrary/pdf_maps/glo_plats/marquette/45n26w.pdf. Google Scholar
  62. J. F. Glahn, M. E. Tobin, and B. F. Blackwell . 2000. A science based initiative to manage Double-crested Cormorant damage to southern aquaculture. United States Department of Agriculture, Animal and Plant Health Inspection Service publication 11-55-010, 40 pp., Washington, DC. Google Scholar
  63. E. A. Goldman 1908. Summer birds of the Tulare Lake region. Condor 10:201. Google Scholar
  64. N. S. Goss 1889. The Double-crested Cormorant. Transactions of the Kansas Academy of Science 11:59–60. Google Scholar
  65. F. Grassett 1926. An unusual flight of cormorants. Wilson Bulletin 38:4234–235. Google Scholar
  66. F. Gress, R. W. Risebrough, D. W. Anderson, L. F. Kiff, and J. R. Jehl Jr. . 1973. Reproductive failures of Double-crested Cormorants in southern California and Baja California. Wilson Bulletin 85:197–208. Google Scholar
  67. J. Grinnell 1908. Birds of a voyage on Salton Sea. Condor 10:186. Google Scholar
  68. J. Grinnell and A. H. Miller . 1944. The distribution of the birds of California. Pacific Coast Avifauna 27:1–608. Google Scholar
  69. A. O. Gross 1951. The Herring Gull-Cormorant control project. Proceedings of the 10th International Ornithological Congress. pp. 532-536. Google Scholar
  70. R. Hakluyt 1904. The principal navigations, voyages, traffiques and discoveries of the English Nation. Vol. 8. J. MacLehose and Sons, Glasgow. pp. 154, 164. Google Scholar
  71. P. L. Hatch 1892. Notes on the birds of Minnesota, with specific characters. Harrison and Smith, Minneapolis, MN. Google Scholar
  72. J. J. Hatch 1995. Changing populations of Double-crested Cormorants. Colonial Waterbirds 18:Special Publication 18–24. Google Scholar
  73. J. J. Hatch and D. V. Weseloh . 1999. Double-crested Cormorant (Phalacrocorax auritus). In The Birds of North America, No. 441 (A. Poole and F. Gill, Eds.). The Birds of North America, Inc., Philadelphia, PA. Google Scholar
  74. C. L. Hayward, C. Cottam, A. M. Woodburry, and H. H. Frost . 1976. Great basin naturalist memoirs: birds of Utah. Brigham Young University Press, Provo, UT. Google Scholar
  75. C. Henderson 2001. Minnesota profile: Double-crested Cormorant (Phalacrocorax auritus). Minnesota Conservation Volunteer (May-June 2001). Google Scholar
  76. H. Herrick 1873. A partial catalogue of the birds of Grand Menan. N.B. Bulletin of the Essex Institute 5:28–41. Google Scholar
  77. K. A. Hobson and J. C. Driver . 1989. Archaeological evidence for the use of the Straight of Georgia by marine birds. Pages 168-173 in The ecology and status of marine and shoreline birds in the Straight of Georgia, British Columbia (K. Vermeer and R. W. Butler, Eds.). Canadian Wildlife Service Special Publication, Ottawa, Ontario. Google Scholar
  78. H. Howard 1929. The avifauna of Emeryville Shellmound. University of California Publications in Zoology 32:2301–394. Google Scholar
  79. A. H. Howell 1911. Birds of Arkansas. U.S. Department of Agriculture, Biological Survey-Bulletin 38. Government Printing Office, Washington. Google Scholar
  80. A. B. Howell 1917. Birds of the islands off the coast of southern California. Pacific Coast Avifauna 12:1–127. Google Scholar
  81. J. A. Hoyle, J. M. Casselman, and T. Schaner . 1999. Smallmouth bass (Micropterus dolomieu) population status in eastern Lake Ontario, 1978 to 1998. In Lake Ontario Fisheries Unit 1998. Ontario Ministry of Natural Resources. Picton, Ontario, Canada. Google Scholar
  82. L. M. Huey 1927. Northernmost breeding station of the Heerman Gull on the Pacific Ocean, and other notes from San Roque Island, Lower California. Condor 29:205–206. Google Scholar
  83. J. Hutchinson 1999. Large cormorant nesting site found in Arkansas. The Aquaculture News 7:91–2. Google Scholar
  84. D. A. James and J. C. Neal . 1986. Arkansas birds: their distribution and abundance. University of Arkansas Press, Fayetteville. Google Scholar
  85. Jesuit Relations 1896. The Jesuit Relations and allied documents: travels and explorations of the Jesuit missionaries in New France, 1610-1791. R. G. Thwaites, Ed. Burrows Bros., Cleveland 1: 153, 155. Google Scholar
  86. S. G. Jewett, W. P. Taylor, W. T. Shaw, and J. W. Aldrich . 1953. Birds of Washington State. University of Washington Press, Seattle. Google Scholar
  87. H. K. Job 1898. The enchanted isles. Osprey 3:37–41. Google Scholar
  88. P. A. Johnsgard 1993. Cormorants, darters and pelicans of the world. Smithsonian Institution Press, Washington, DC. Google Scholar
  89. H. E. Kingery 1998. Colorado Breeding Bird Atlas. Colorado Bird Atlas Partnership and Colorado Division of Wildlife. Google Scholar
  90. K. Kloor 1999. Killing all cormorants?. Audubon July-August 1999. p. 16. Google Scholar
  91. O. W. Knight 1908. The birds of Maine. Charles H. Glass and Company, Bangor, ME. Google Scholar
  92. W. H. Koonz and P. W. Rakowski . 1985. Status of colonial waterbirds nesting in southern Manitoba. Canadian Field-Naturalist 99:119–29. Google Scholar
  93. C. Korfanty, W. G. Miyasaki, and J. L. Harcus . 1997. Review of the population status and management of Double-crested Cormorants in Ontario. Ontario Ministry of Natural Resources, Fish and Wildlife Branch, Peterborough, Ontario. Google Scholar
  94. W. B. Krohn, R. B. Allen, J. R. Moring, and A. E. Hutchinson . 1995. Double-crested Cormorants in New England: Population and management histories. Colonial Waterbirds 18:Special Publication 199–109. Google Scholar
  95. J. A. Kushlan, M. J. Steinkamp, K. C. Parsons, J. Capp, M. A. Cruz, M. Coulter, I. Davidson, L. Dickson, N. Edelson, R. Elliot, R. M. Erwin, S. Hatch, S. Kress, R. Milko, S. Miller, K. Mills, R. Paul, R. Phillips, R. E. Saliva, B. Sydeman, J. Trapp, J. Wheeler, and K. Wohl . 2002. Waterbird Conservation for the Americas: The North American Waterbird Conservation Plan, Version 1. Waterbird Conservation for the Americas, Washington, DC. 78 pp. Google Scholar
  96. C. C. Lamb 1927. The birds of Natividad Island, Lower California. Condor 29:69. Google Scholar
  97. C. Lamb and A. B. Howell . 1913. Notes from Buena Vista Lake and Fort Teton. Condor 15:115–118. Google Scholar
  98. F. W. Langdon 1878. Observations on Cincinnati Birds. The Journal of the Cincinnati Society of Natural History 1:110–118. Google Scholar
  99. H. F. Lewis 1929. The natural history of the Double-crested Cormorant (Phalacrocorax auritus auritus (Lesson)). Ph.D. Thesis, Cornell University, Ithaca, NY. Google Scholar
  100. W. Lewis and M. Clark . 1814. History of the expedition under the command of Captains Lewis and Clark to the sources of the Missouri, thence across the Rocky Mountains and down the River Columbia to the Pacific Ocean performed during the years 1804-5-6. By order of the Government of the United States of America: Bradford and Inskeep, Philadelphia; Abm. H. Inskeep, New York. Google Scholar
  101. C. B. Linton 1908. Notes from Buena Vista Lake, May 20 to June 16, 1907. Condor 10:196. Google Scholar
  102. A. R. Lock, R. G B. Brown, and S. H. Gerriets . 1994. Gazetteer of Marine Birds in Atlantic Canada. Canadian Wildlife Service, Environmental Conservation Branch, Environment Canada, Atlantic Region. Google Scholar
  103. D. M. Lodge and K. Shrader-Frechette . 2003. Nonindigenous species: ecological explanation, environmental ethics, and public policy. Conservation Biology 17:31. 37. Google Scholar
  104. W. E. Loucks 1893. Illinois River Valley from an ornithological standpoint. Oologist 10:4119–121. Google Scholar
  105. Louisiana Department of Conservation 1931. The birds of Louisiana. Published by the Department of Conservation, Baton Rouge, LA. Google Scholar
  106. H. Lounsbury 2000. No Sanctuary. The Bay Times Press, Vol. 127 (191): July 9, 2000. pp. 1A-4A. Google Scholar
  107. J. P. Ludwig and C. L. Summer . 1995. Final report of the SERE Group Ltd. study to: Model the population of Double-crested Cormorants in the Les Cheneaux region of Lake Huron, and to collect data relevant to fish species utilization by cormorants and other waterbirds from the northern Great Lakes. The SERE Group, Ltd., Eureka, MI. Google Scholar
  108. B. E. Luedtke 1980. The Calf Island site and the Late Prehistoric period in Boston Harbor. Man in the Northeast 20:25–76. Google Scholar
  109. G. H. Mackay 1894. Habits of the Double-crested Cormorant (Phalacrocorax dilophus) in Rhode Island. Auk 11:18–25. Google Scholar
  110. J. Macoun and J. M. Macoun . 1909. Catalogue of Canadian Birds. Government Printing Bureau, Ottawa. Google Scholar
  111. W. McInnes 1913. The basins of Nelson and Churchill rivers. Government printing bureau. Ottawa, Ontario. Google Scholar
  112. J. E. McKearnan 1997. Colonial waterbird population analysis project, phase 1. Final report. submitted to MN Department of Natural Resources, Natural Heritage and Non-Game Research Program, St. Paul, MN. Google Scholar
  113. H. L. Mendall 1936. The home-life and economic status of the Double-crested Cormorant (Phalacrocorax auritus auritus). The University Press, Orono, ME. Google Scholar
  114. J. C. Merrill 1888. Notes on the birds of Fort Klamath, Oregon, with remarks on certain species by William Brewster. Auk 5:139–146.251–262.357–366. Google Scholar
  115. R. Milko, D. Loney, R. Elliot, and G. Donaldson . 2003. Wings Over Water: Canada’s Waterbird Conservation Plan. Canadian Wildlife Service, Environment Canada. Ottawa, Ontario. Google Scholar
  116. E. L. Mills, J. H. Leach, J. T. Carlton, and C. L. Secor . 1993. Exotic species in the Great Lakes: a history of biotic crises and anthropogenic introductions. Journal of Great Lakes Research 19:11–54. Google Scholar
  117. G. R. Milton, P. J. Austin-Smith, and G. J. Farmer . 1995. Shouting at shags: A case study of cormorant management in Nova Scotia. Colonial Waterbirds 18:Special Publication 191–98. Google Scholar
  118. H. H. Mitchell 1925. Some more notes on Saskatchewan birds. Condor 27:94–97. Google Scholar
  119. MoffittJ. 1939. From field and study. Condor 41:33. Google Scholar
  120. T. Morton 1637. The new English Canaan. Amsterdam. Reprints: Peter Force, Washington, 1838; The Prince Society, Boston, 1883. Google Scholar
  121. R. E. Mumford and C. E. Keller . 1984. The Birds of Indiana. Indiana University Press, Bloomington. Google Scholar
  122. J. A. Munro 1928. Cormorants nesting on Bare Island, British Columbia. Condor 30:327–328. Google Scholar
  123. O. J. Murie 1959. Fauna of the Aleutian Islands and the Alaska Peninsula. North American Fauna 61:1–406. Google Scholar
  124. NAWMP 1998. U.S. Dept. of the Interior, Environment Canada, and Secretaría de Medio Ambiente y Recursos Naturales de Mexico. 1998 Update to the North American Waterfowl Management Plan: Expanding the Vision. Washington, DC. Google Scholar
  125. NYSDEC Newsletter 2004. DEC announces cormorant management plans. May 2004. http://www.dec. state.ny.us/website/environmentdec/2004a/cormorants04.html Last accessed 09/18/2004. Google Scholar
  126. H. C. Oberholser 1974. The bird life of Texas. University of Texas Press, Austin. Google Scholar
  127. Ontario Parks 2005. Annual report on the management of Double-crested Cormorants at Presqu’ile Provincial Park, 2004. 2005, Queens Printer of Ontario. 51860 ISSN 1710-3797 electronic. http://www.ontarioparks.com/english/pres_planning.html. Google Scholar
  128. R. S. Palmer 1962. Handbook of North American birds. Volume I. Yale University Press, New Haven, CT. Google Scholar
  129. J. G. Pearson 1899. Notes on some of the birds of eastern North Carolina. Auk 16:246–250. Google Scholar
  130. C. J. Pennock 1889. Nesting of the Florida Cormorant. Ornithologist and Oologist 14:154–155. Google Scholar
  131. B. G. Peterjohn and W. Zimmerman . 1989. The Birds of Ohio. Indiana University Press, Bloomington and Indianapolis. Google Scholar
  132. R. Peterson 1995. The South Dakota breeding bird atlas. South Dakota Ornithologists’ Union, Aberdeen. Google Scholar
  133. A. Phillips, J. Marshall, and G. Monson . 1964. The birds of Arizona–The University of Arizona Press, Tucson. Google Scholar
  134. L. O. Pindar 1925. Birds of Fulton County, Kentucky. Wilson Bulletin 37:77–88. Google Scholar
  135. W. Post and C. A. Post . 1988. Double-crested Cormorant nesting in South Carolina. The Chat 52:35. Google Scholar
  136. S. Postupalsky 1978. Toxic chemicals and cormorant populations in the Great Lakes. Canadian Wildlife Service, Wildlife Toxicology Division Manuscript Report 40:1–25. Google Scholar
  137. S. A. Pothoven, T. F. Nalepa, P. J. Schneeberger, and S. B. Brandt . 2001. Changes in diet and body condition of Lake Whitefish in southern Lake Michigan associated with changes in benthos. North American Journal of Fisheries Management 21:876–883. Google Scholar
  138. M. S. Ray 1915. Nesting of the American Osprey at Eagle Lake, California Condor 17 70–74. Google Scholar
  139. H. Reeks 1869. Notes on the zoology of Newfoundland. The Zoologist: a monthly journal of natural history 4:1848–1859. Google Scholar
  140. D. S. Reinhold, A. J. Mueller, and G. Ellis . 1998. Observations of nesting Double-crested Cormorants in the Delta Region of Mississippi. Colonial Waterbirds 21:3:466–467. Google Scholar
  141. A. C. Revkin 1998. A Slaughter of Cormorants in Angler Country. The New York Times, Aug 1, Section: Metropolitan Desk. Google Scholar
  142. S. N. Rhoads 1895. Contributions to the Zoology of Tennessee, No. 2. Birds. Proceedings of the Academy of Natural Sciences of Philadelphia pp. 463-501. Google Scholar
  143. A. Ricciardi 2001. Facilitative interactions among aquatic invaders: is an “invasional meltdown” occurring in the Great Lakes?. Canadian Journal of Fisheries and Aquatic Sciences 58:2513–2525. Google Scholar
  144. R. Ridgway 1874. The Lower Wabash Valley, Considered in its relation to the faunal districts of the eastern region of North America; with a synopsis of its avian fauna. Proceedings of the Boston Society of Natural History 16:304–332. Google Scholar
  145. T. S. Roberts 1932. The Birds of Minnesota. Vol. 1. University of Minnesota Press, Minneapolis. Google Scholar
  146. E. S. Rolfe 1898. Notes from the Devil’s Lake Region. Osprey 2:125–128. Google Scholar
  147. W. R. Salt and A. L. Wilk . 1958. The birds of Alberta. Government of Alberta. Department of Economic Affairs. The Queens Printer, Edmonton. Google Scholar
  148. T. Salvadori 1865. Descrizione di alter nuove species di uccelli esistenti nel Museo di Torina. Atti della societa di Scienze Naturali [Milan] 8:375–379. Google Scholar
  149. J. Schmitt 1902. A summer colony at Anticosti. Auk 19:181–183. Google Scholar
  150. G. B. Sennett 1891. Abstract of the Proceedings of the Linnaean Society of New York for the year ending Mar. 6, 1891. No. 3, p. 5. Google Scholar
  151. E. T. Seton 1886. The birds of western Manitoba. Auk 3:145–156. Google Scholar
  152. E. T. Seton 1908. Bird records from Great Slave Lake region. Auk 25:68–69. Google Scholar
  153. Eric Sharp 2004. Native cormorants? Not in Great Lakes. Detroit Free Press. Outdoors. Published April 8, 2004. http://www.freep.com/sports/outdoors/eric8_20040408.htm. Google Scholar
  154. S. Sher 1994. The native legacy of Emeryville, Part Two: The destruction of the Emeryville shellmound. Journal of the Emeryville Historical Society 5:33–10. Google Scholar
  155. D. Siegel-Causey, C. Lefevre, and A. B. Savinetskii . 1991. Historical diversity of cormorants and shags from Amchitka Island, Alaska. Condor 93:840–852. Google Scholar
  156. F. Smith 1911. Double-crested Cormorants breeding in Central Illinois. Auk 28:16–19. Google Scholar
  157. W. A. Squires 1917. Some Field notes for 1917. Condor 19:185–187. Google Scholar
  158. H. Stansbury 1853. Exploration and survey of the valley of the Great Salt Lake of Utah. Robert Armstrong, Public Printer, Washington. Google Scholar
  159. H. M. Stevenson and B. H. Anderson . 1994. The birdlife of Florida. University Press of Florida, Gainesville. Google Scholar
  160. R. E. Stewart 1975. Breeding birds of North Dakota. Tri-College Center for Environmental Studies, Fargo, ND. Google Scholar
  161. J. Tanner 1994. The falcon: a narrative of the captivity and adventures of John Tanner during thirty years residence among the Indians in the interior of North America. Penguin Books, New York. p. 62. Google Scholar
  162. J. Tate and D. J. Tate . 1982. The Blue List and the list of species with Special Concerns. American Birds 36:129–134. Google Scholar
  163. The Catfish Journal 1999. Cormorant nesting area located in southwest Arkansas. 8 (11): 14-15. Google Scholar
  164. The Jesuit Relations 1896. The Jesuit relations and allied documents: travels and explorations of the Jesuit missionaries in New France, 1610-1791. the original French, Latin, and Italian texts, with English translations and notes (R. G. Thwaites, Ed.). Vol. 1, pp. 155, 159. Burrows Bros., Cleveland, OH. Google Scholar
  165. W. E C. Todd 1963. Birds of the Labrador Peninsula and adjacent areas: a distributional list. Published in association with Carnegie Museum by University of Toronto Press. Toronto. 819 pp. Google Scholar
  166. C. W. Townsend 1917. In Audubon’s Labrador. Auk 34:133–146. Google Scholar
  167. C. W. Townsend and A. C. Bent . 1910. Additional notes on the birds of Labrador. Auk 27:1–18. Google Scholar
  168. M. B. Trautman 1940. The birds of Buckeye Lake, Ohio. University of Michigan Press, Ann Arbor. 466 pp. Google Scholar
  169. C. H. Trost 1994. Status and distribution of colonial nesting waterbirds in southern Idaho, 1993. Idaho Bureau of Land Management Technical Bulletin No. 94-6. July 1994. Google Scholar
  170. L. M. Turner 1885. Notes on the birds of the Nearer Islands, Alaska. Auk 2:154–159. Google Scholar
  171. USDA/WS 2004. Activity Sheet. Living with cormorants. http://migratorybirds.fws.gov/issues/cormorant/cormorant.html Last accessed 09/16/2004. Google Scholar
  172. USDA/APHIS/WS 2003. Wildlife Services: Managing bird damage to America’s resources. Program Aid No. 1750. Issued August 2003. http://www.aphis.usda.gov/oa/pubs/pub_wsbirddamage.pdf Last accessed 09/16/2004. Google Scholar
  173. USDA/APHIS/WS 2004. Environmental Assessment: Reducing Double-crested Cormorant damage through an integrated wildlife damage management program in the state of Michigan. http://www.usda.aphis.gov/ws/nepa/MIdoublecrest.pdf Last accessed 09/16/2004. Google Scholar
  174. U.S. EPA 1972. DDT ban takes effect. Press Release, December 31, 1972. http://www.epa.gov/history/topics/ddt/01.htm Last updated, 12/13/2004. Google Scholar
  175. USDI/FWS 2003a. Migratory bird permits; regulations for Double-crested Cormorant management, Proposed rule. Federal Register: March 17, 2003. 68(51): 12653-12660. Google Scholar
  176. USDI/FWS 2003b. Final environmental impact statement: Double-crested Cormorant management in the United States. Prepared by USDI/FWS, in cooperation with USDA/APHIS/WS, 2003. Google Scholar
  177. USDI/FWS 2003c. Migratory bird permits; regulations for Double-crested Cormorant management, Final rule. Federal Register: October 8, 2003. 68(195): 58022-58037. Google Scholar
  178. USFWS 2002. Colonial nesting waterbirds: a glorious and gregarious group. http://www.birds.fws.gov/Waterbird-Fact-Sheet.pdf Last accessed 09/16/2004. Google Scholar
  179. USFWS 2003. Service Releases Proposed Rule on Double-Crested Cormorant Management. News Release. Mar 2003. http://migratorybirds.fws.gov/issues/cor-morant/cormorant.html Last accessed 09/16/2004. Google Scholar
  180. K. Vermeer and L. Rankin . 1984. Population trends in nesting Double-crested and Pelagic Cormorants in Canada. Murrelet 65:1–9. Google Scholar
  181. A. T. Wayne 1910. Birds of South Carolina. Series: Contributions of the Charleston Museum, 1. The Daggett Printing Company, Charleston. Google Scholar
  182. D. V. Weseloh and B. Collier . 1995. The rise of the Double-crested Cormorant on the Great Lakes: winning the war against contaminants. Great Lakes Fact Sheet. Canadian Wildlife Service, Environment Canada and Long Point Observatory. http://www.on. ec.gc.ca/wildlife/factsheets/fs_cormorants-e.html. Google Scholar
  183. D. V. Weseloh, P. J. Ewins, J. Struger, P. Mineau, C. A. Bishop, S. Postupalsky, and J. P. Ludwig . 1995. Double-crested Cormorants of the Great Lakes: Changes in population size, breeding distribution and reproductive output between 1913 and 1991. Colonial Waterbirds 18:Special Publication 148–59. Google Scholar
  184. J. M. Wheaton 1882. Report on the birds of Ohio. Pages 187-628 in Report of the Geological Survey of Ohio. Vol. 4. Zoology and Botany. Nevins and Myers, State Printers, Columbus. Google Scholar
  185. C. S. Whitehead 1887. “Loon Lake”. Ornithologist and Oologist 12:221–22. Google Scholar
  186. O. Widmann 1907. A preliminary catalog of the birds of Missouri. Transactions of the Academy of Science. St. Louis 17:11–266. Google Scholar
  187. G. Willett 1910. A summer trip to the northern Santa Barbara Islands. Condor 12:170–174. Google Scholar
  188. G. Willett 1919. Bird notes from southeastern Oregon and northeastern California. Condor 21:194–207. Google Scholar
  189. G. Willett 1933.A revised list of the birds of southwestern California. Pacific Coast Avifauna 21. Google Scholar
  190. R. Williams 1643. A key into the language of America. London. Reprinted by Rhode Island Historical Society, 1827. Google Scholar
  191. L. R. Wires, F. J. Cuthbert, D. R. Trexel, and A. R. Joshi . 2001. Status of the Double-crested Cormorant (Phalacrocorax auritus) in North America. Final report to USFWS. Fort Snelling, Minnesota. Google Scholar
  192. L. R. Wires and F. J. Cuthbert . 2001. Prioritization of waterbird colony sites for conservation in the U.S. Great Lakes. Final report to U.S. Fish and Wildlife Service, Nov 2001. Fort Snelling, Minnesota. Google Scholar
  193. L. R. Wires and F. J. Cuthbert . 2003. Fish-eating bird predation at aquaculture facilities in Minnesota: a first step towards bridging the information gap. Final report to Minnesota Sea Grant. Duluth, Minnesota. Google Scholar
  194. W. Wood 1634. Nevv England’s prospect: a true, lively, and experimental description of that part of America, commonly called Nevv England. Tho. Cotes, for Iohn Bellamie, printed at London. Google Scholar
  195. H. W. Wright 1913. The birds of San Martin Island, Lower California. Condor 15:207–210. Google Scholar
  196. A. P. Wywialowski 1999. Wildlife-caused losses for producers of Channel Catfish Ictalurus punctatus in 1996. Journal of the World Aquaculture Society 30:4461–472. Google Scholar
  197. 23 U.S.T. 260 1972. United States Treaties. U.S. Department of State, Vol. 23, pp. 260-261. Google Scholar

Appendices

Figure 1. Historic (1900 and earlier) and current (1970-2000) breeding locations of the Double-crested Cormorant in North America. 1) Alaska; 2) Pacific Coast; 3) Canadian and U.S. Interior; 4) Southeast U.S./Caribbean; 5) Northeast Atlantic Coast.

i1524-4695-29-1-9-f01.gif

Table 1. Early North American breeding records for Double-crested Cormorants (B = breeding; PB = possible/probable breeding; * = Best approximate location; NA = not available).

i1524-4695-29-1-9-t101.gif

Table 1. (Continued) Early North American breeding records for Double-crested Cormorants (B = breeding; PB = possible/probable breeding; * = Best approximate location; NA = not available).

i1524-4695-29-1-9-t102.gif

Table 1. (Continued) Early North American breeding records for Double-crested Cormorants (B = breeding; PB = possible/probable breeding; * = Best approximate location; NA = not available).

i1524-4695-29-1-9-t103.gif

Table 1. (Continued) Early North American breeding records for Double-crested Cormorants (B = breeding; PB = possible/probable breeding; * = Best approximate location; NA = not available).

i1524-4695-29-1-9-t104.gif

Table 1. (Continued) Early North American breeding records for Double-crested Cormorants (B = breeding; PB = possible/probable breeding; * = Best approximate location; NA = not available).

i1524-4695-29-1-9-t105.gif

Table 1. (Continued) Early North American breeding records for Double-crested Cormorants (B = breeding; PB = possible/probable breeding; * = Best approximate location; NA = not available).

i1524-4695-29-1-9-t106.gif

Table 2. Numbers of breeding pairs of Double-crested Cormorants in each breeding zone (A = complete count, estimate at all known colonies; B = partial estimate, does not include all known colonies; C = extrapolated older count or other informed estimate; D = estimate often based on knowledge of most colonies or combining counts of different years; E = only old, indirect or scattered recent knowledge available; NA = Not available).

i1524-4695-29-1-9-t201.gif

Table 2. (Continued) Numbers of breeding pairs of Double-crested Cormorants in each breeding zone (A = complete count, estimate at all known colonies; B = partial estimate, does not include all known colonies; C = extrapolated older count or other informed estimate; D = estimate often based on knowledge of most colonies or combining counts of different years; E = only old, indirect or scattered recent knowledge available; NA = Not available).

i1524-4695-29-1-9-t202.gif

Table 2. (Continued) Numbers of breeding pairs of Double-crested Cormorants in each breeding zone (A = complete count, estimate at all known colonies; B = partial estimate, does not include all known colonies; C = extrapolated older count or other informed estimate; D = estimate often based on knowledge of most colonies or combining counts of different years; E = only old, indirect or scattered recent knowledge available; NA = Not available).

i1524-4695-29-1-9-t203.gif

Table 2. (Continued) Numbers of breeding pairs of Double-crested Cormorants in each breeding zone (A = complete count, estimate at all known colonies; B = partial estimate, does not include all known colonies; C = extrapolated older count or other informed estimate; D = estimate often based on knowledge of most colonies or combining counts of different years; E = only old, indirect or scattered recent knowledge available; NA = Not available).

i1524-4695-29-1-9-t204.gif
Linda R. Wires and Francesca J. Cuthbert "Historic Populations of the Double-crested Cormorant (Phalacrocorax auritus): Implications for Conservation and Management in the 21st Century," Waterbirds 29(1), (1 March 2006). https://doi.org/10.1675/1524-4695(2006)29[9:HPOTDC]2.0.CO;2
Received: 11 July 2005; Accepted: 1 October 2005; Published: 1 March 2006
JOURNAL ARTICLE
29 PAGES


SHARE
ARTICLE IMPACT
Back to Top