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HISTORICAL BREEDING BY PELAGIC, DOUBLE-CRESTED, AND BRANDT'S CORMORANTS IN THE SALISH SEA, 1891-1955.

Historical breeding by Pelagic Cormorant (Phalacrocorax pelagicus), Double-crested Cormorant (P. auritus) and Brandt's Cormorant (P. penicillatus) in the Salish Sea region of British Columbia and Washington in the late 19th and early 20th centuries has been summarized to some degree but never fully examined (Drent and Guiguet 1961; Speich and Wahl 1989; Campbell and others 1990; Carter and others 1995; Carter 2016). I present published and unpublished information to identify when breeding was first documented at Salish Sea colonies for all 3 species of cormorant prior to 1956, when surveys became more common. This information will provide a better background for interpreting population trends of cormorants in the Strait of Georgia (Carter, Chatwin, and Drever, submitted.) and other regions of the Salish Sea after 1955.

METHODS

The Salish Sea encompasses the southwestern portion of the Canadian province of British Columbia and the northwestern portion of the state of Washington in the United States. The southern and central sections of the Salish Sea include the southern Gulf Islands, the San Juan Islands, and the Juan de Fuca Strait (Fig. 1), whereas the northern section of the Salish Sea includes the Strait of Georgia northward from Vancouver (Fig. 2). To gather data on historical cormorant breeding in the Salish Sea, I conducted: (1) a literature review; (2) a search for specimens in museums, using VertNet (vertebrate distributed database network), in-person museum visits, and correspondence with museum staff; and (3) an examination of miscellaneous unpublished field notes and correspondence of early British Columbia naturalists in the British Columbia Archives (BCA), Canadian Museum of Nature Archives (CMNAC) and Museum of Zoology at Cambridge University (UMZC). Drent and Guiguet (1961) provided the most extensive summary of pre-1956 nesting by Double-crested Cormorants in the Salish Sea, based on literature, specimens, and correspondence; they also summarized nesting by Pelagic Cormorants in the British Columbia portion of the Salish Sea. Speich and Wahl (1989) later conducted a fairly exhaustive search of the literature, specimens, and field notes in U.S. museums and institutions, which turned up more information for Washington colonies than found by Drent and Guiguet (1961). But Speich and Wahl (1989) did not interpret this additional information and did not attempt to summarize unpublished Canadian or English information. I did not attempt to collate every breeding record at each colony because information is very limited at most colonies and much of it is already provided in Drent and Guiguet (1961) and Speich and Wahl (1989). Instead, I focused on: (1) examining when breeding was first noted at colonies in the Salish Sea, which served to highlight early historical records not included in past summaries; and (2) integrating published and unpublished information to outline the historical status of breeding cormorants in the Salish Sea.

RESULTS

Lack of Cormorant Breeding (1792-1890)

Baird and others (1884) did not report any known breeding by Pelagic, Double-crested or Brandt's cormorants in the Salish Sea, yet the central portion of this region (southern Gulf Islands, San Juan Islands and inner Strait of Juan de Fuca) had received considerable attention from early ornithologists and naturalists since the late 18th century. First reports of many waterbird species exist from this period. In 1792, Black Oystercatchers (Haematopus bachmani) were first reported breeding at Smith Island in the central Salish Sea, as well as Pigeon Guillemots (Cepphus columba) at Cutts Island in the southern Salish Sea by A Menzies, during the voyage of HMS Discovery (Captain G Vancouver; (Speich and Wahl 1989; Carter and others in press). In contrast, no specimens of cormorants were deposited in the Natural History Museum (NHM; previously British Museum). Cooper and Suckley (1859) reported breeding by Pigeon Guillemots at the entrance to Hood Canal and Whidbey Island in 1855 (Carter and others in press) and breeding by Rhinoceros Auklets (Cerorhinca monocerata) at Protection Island, as reported by indigenous people (Carter and Sealy 2011a), during surveys related to the Pacific Railroad in 1853-1857. They did not report breeding cormorants, although most of their time in the southern Salish Sea (southern Puget Sound) was spent in winter, with only March 1855 spent at Whidbey Island in the central Salish Sea. However, Baird (1858) reported a specimen of a Pelagic Cormorant collected by CBR Kennerly at Orcas Island, San Juan Islands, in February 1858. Cormorants were not collected and no specimens deposited in the NHM in 1857-1860 during the voyage of the HMS Plumper (Captain GH Richards) when Black Oystercatchers, Pigeon Guillemots, Rhinoceros Auklets, and Tufted Puffins (Fratercula cirrhata) were reported breeding in the central Salish Sea at Waldron Island, Mandarte Island, Bird Rocks, Williamson Rocks, and White Rock by D Lyall and possibly CB Wood (Mayne 1862; Carter and others 2011a,b, in press). At this time, Pelagic Cormorants were noted as "extremely common" by Wood, while Double-crested and Brandt's cormorants were not mentioned (Mayne 1862: page 418). However, Lyall did collect or obtain one adult Double-crested Cormorant on the northwest coast of Vancouver Island at Quatsino Sound in September 1860 (NHM 61.8.12.20).

Infrequent visitation to many parts of the central Salish Sea between 1792 and 1860 could have led to the lack of records if cormorants were nesting only sporadically or in small numbers. However, the most convincing data supporting a general lack of nesting by cormorants in that area was obtained in 1861-1867. During this period, JE Hepburn, a very knowledgeable English ornithologist living in Victoria and San Francisco, collected many seabirds and eggs from the central Salish Sea, and his collection is currently housed in the UMZC (Swarth 1926; Kinnear 1931; Jewett and others 1953). Hepburn obtained breeding records for other water species (Black Oystercatchers, Glaucous-winged Gulls [Larus glaucescens], Pigeon Guillemots, and Rhinoceros Auklets) on Smith Island, Bird Rocks, Williamson Rocks, Mandarte Island, Flattop Island, Bare Island, and San Juan Island, but he did not document breeding for cormorant species in this area (Baird and others 1884; Carter and others 2011a, b, in press; Hepburn, unpubl. notes [UMZC]). He may have missed a few nests at certain locations or years, since many of these islands later supported Pelagic Cormorant colonies, and he did collect 2 specimens of Pelagic Cormorants, indicating that at least some birds were present in the central Salish Sea at this time, although uncommon. On 31 May 1867, a male in adult plumage was collected at Saanich, Vancouver Island (UMZC 10/Phal/3/r/4; JH 1390). Hepburn (unpubl. notes; UMZC) noted that this individual was "immature", with "testes not enlarged." On 27 June 1867, a male in immature plumage was also collected at "Bare Island, Vancouver Island" (UMZC 10/Phal/3/r/9; JH 1393). It is unclear if this locality was Mandarte Island (also referred to as "Bare Island") or Bare Island east of Waldron Island; both of these islands occurred within the area referred to as 'Vancouver Island' by Hepburn. The next day (28 June 1867), he collected 2 Glaucous-winged Gull chicks from a nest at Flattop Island, located between these different Bare Islands (see Fig. 1). Hepburn did not collect any specimens of Double-crested or Brandt's cormorants, suggesting that they were absent or very uncommon in the central Salish Sea in 1861-1867. Lord (1866) stated that Pelagic and Double-crested Cormorants were abundant at "Fort Rupert" off northeastern Vancouver Island in 1862, likely referring in part to 2 specimens of Pelagic Cormorants obtained in this area in November 1862 (Carter and McClaren 2016). Brown (1868), who prepared a first list of known bird species for Vancouver Island assisted by Hepburn, noted Double-crested Cormorant and Pelagic Cormorant but not Brandt's Cormorant.

Pelagic Cormorant (1891-1955)

Fannin (1891) first reported breeding Pelagic Cormorants in the central Salish Sea at Mandarte Island (Table 1; see Figs. 1 and 2 for colony locations). Definite evidence of breeding was secured when 4 eggs (Western Foundation of Vertebrate Zoology [WFVZ] 81,319; set mark 1) were collected on 1 July 1892 at Mandarte Island (referred to as "Bare Isle, Sts. of Georgia") by W Burton, a well-known naturalist and collector from Victoria. On the egg card, Burton described the nest as "grasses + their own doing [feces] matted together on the side of cliff. A filthy nest." Between 1900 and 1940, 14 more nesting colonies were noted in the San Juan Islands (n = 8), inner Strait of Juan de Fuca (n = 3), Gulf Islands (n = 2), and northern Strait of Georgia (n = 1). During the 1941-1955 period, 5 other new colonies were first noted in the Gulf Islands (n = 2), San Juan Islands (n = 2), and inner Strait of Juan de Fuca (n = 1).

Greater detail is provided below for 3 other breeding records of Pelagic Cormorants not previously published: (1) Skipjack Island ("Skip Jack Rock Harro [sic] Straits", likely the bare rock about 135 m east of Skipjack Island)--on 10 July 1922, Burton collected 4 eggs (WFVZ 157,884; set mark 10) and the nest was "in a colony on face of cliff"; (2) Christie Islet--on 11 June 1924, RA Cumming, a well-known naturalist and collector from Vancouver, collected 2 eggs (Royal British Columbia Museum [RBCM] E1388); and (3) Jones Island--on 15 June 1936, WA Newcombe (unpubl. notes; BCA MS-1077) found 12 nests (2 with 4 eggs, 3 with 1 egg, and 7 either empty or could not be viewed) on a low cliff on the south side of the island. Newcombe had earlier worked as assistant biologist at the British Columbia Provincial Museum (BCPM; now known as the RBCM) in 1928-1933 (see Carter 2016).

Given infrequent colony visitations, many of these colonies may not have been noted until years after they had first formed. For example, in 1905, nesting was described by JM Edson and WL Dawson at 2 colonies in the San Juan Islands (Flattop Island and Waldron Island; Table 1), but was not noted at 3 other locations (Williamson Rocks, Colville Island, and Bare Island) where breeding subsequently occurred in 1937-1949 (Edson 1929; Speich and Wahl 1989; Table 1). Dawson and Bowles (1909) did not clearly state that any Pelagic Cormorants bred in the San Juan Islands, although they did include a photograph taken by Dawson of a clutch of 3 eggs in a nest on Flattop Island, presumably from 1905. Three other colonies in the northern Strait of Georgia (Mitlenatch Island, Ballenas Islands, and Hornby Island) also did not appear to form until the 1944-1955 period, given that nesting did not occur at Mitlenatch Island in 1922-1943 or at Hornby Island in 1938 (Pearse 1956).

Double-crested Cormorant (1896-1955)

Breeding by Double-crested Cormorants was first reported in the central Salish Sea at Mandarte Island in 1927, followed by Ballingall Islets in 1934 and Christie Islet in 1941 (Munro 1928,1937; Drent and Guiguet 1961; Carter 2016; Table 1). Regular breeding occurred afterwards at these locations (Drent and Guiguet 1961). Three colonies in the San Juan Islands (Waldron Island, Colville Island and Viti Rocks) were first noted in 1947-1949, followed by the Chain Islands in the Gulf Islands in 1951-1952 (Drent and Guiguet 1961).

Two intriguing earlier breeding records of Double-crested Cormorants in the Salish Sea have not been previously mentioned in the literature and appear to indicate sporadic breeding in the Salish Sea prior to 1927. The 1st record occurred on 18 June 1896, when an unknown collector obtained a set of 4 eggs on a cliff "south of Comox [British Columbia] for J. Fannin" (Sam Noble, Oklahoma Museum of Natural History [OMNH] Egg #95). This specimen was originally in the collection of Walter Raine, a well-known naturalist from Toronto, ON (see Raine 1892). I suspect strongly that the collector was CF Newcombe, a well-known naturalist working for the BCPM. In a letter to the Provincial Secretary, dated 24 June 1896 (BCA MS-1077), Newcombe stated that:
"I have also the honour to report that I left Victoria for Denman
Island on Tuesday the 16th inst. on a collecting expedition amongst the
islands near Cape Mudge, the Rev. H. C. Nixon [Reverend Harpur Colville
John Nixon] having placed at the disposal of a representative of the
Prov. Museum for a few days his steamer the Antolycus. We obtained
several specimens of sea bird nest & eggs... [which] I considered would
be of assistance to Dr. Fannin in preparing groups of birds in the
Museum. ... I made my way over the coast on foot + by stage to Nanaimo
and returned to Victoria by train yesterday the 23rd inst. [this month]"


Newcombe apparently collected eggs of several seabird species on this trip. In pencil, additional notes on the egg card for OMNH Egg #95 are: "sets 1/4 to 9/4 same data", suggesting that 9 clutches were collected, although only this one (set 5/4) could be located in a museum using VertNet. The only other egg specimen of a seabird that I could locate was one Glaucous-winged Gull (Larus glaucescens) egg collected on 20 June 1896 by Newcombe at the "West coast of Vancouver Island" (MCZ 359116). The original egg card was missing, and the locality may have been mis-copied from the original, which may have been written as or intended to be "west coast [of Canada], Vancouver Island." Few cliffs occur in this region and these cormorant eggs most likely were collected on the cliffs west of St. John's Point, Hornby Island, about 29 km SE of Comox. The nest was described as "seaweed on ledge of cliff.... several sets taken same data." The nest material and cliff location described for the 1896 egg record best match Pelagic Cormorant or Brandt's Cormorant, both of which frequently nest on cliffs and whose nests are usually made of seaweed. Double-crested Cormorant nests are typically built out of sticks and located on flat surfaces, although cliff ledges and seaweed construction also are used on occasion. However, the dimensions of the 4 eggs measured by T Yuri (OMNH) were 61.4 X 37.3 mm, 67.3 X 39.3 mm, 60.2 x 39.5 mm, and 64.0 X 39.4 mm. The size of these eggs fell within the range of Double-crested or Brandt's cormorants, and outside the range of Pelagic Cormorant (Davie 1898). Although Newcombe listed the species identity as "correct" on the egg card, Fannin (BCPM curator) may not have accepted the species identification for this specimen, possibly because breeding had not yet been reported in British Columbia. He did not report it in his update of the Checklist of the Birds of British Columbia (Fannin 1898), and the record was not reported in any subsequent summary of the birds of British Columbia. The specimen somehow was excluded from the BCPM egg collection and likely was sold or traded to Raine. With knowledge of later nesting by Double-crested Cormorants at Mandarte Island in 1927 (and possibly at Ballingall Islets even before 1927; see Munro 1937, Drent and Guiguet 1961), I cannot find any convincing reason to reject this breeding record and have treated it as valid for this paper, although genetic confirmation of this specimen is desirable in the future.

The 2nd record occurred on 10 July 1900, when EM Anderson obtained 2 Double-crested Cormorant eggs (RBCM E0038) at "Sidney Island", most likely referring to Mandarte Island, adjacent to Sidney, British Columbia. Anderson later worked as assistant curator of natural history at the BCPM in 1903-1916. On the original egg card, the nest was described as "grass, weeds and guano on steep cliff but the species identity was recorded as "unsure." Fannin or F Kermode (who succeeded Fannin as BCPM curator in 1904) likely also rejected this record as being of suitable quality to represent the first breeding record for the province but the specimen remained in the BCPM collection. It is not mentioned in an update of the checklist (Kermode 1904) or subsequent avifaunal summaries for the province. I also treat this breeding record as valid, prior to future genetic confirmation of species identity.

Brandt's Cormorant (7928-1953)

Brandt's Cormorants were first reported breeding in the central Salish Sea at Flattop Island in 1928 (Speich and Wahl 1989; Table 1). Breeding also was noted at Matia Island in 1940 and at Lopez Island (possibly the Davis Bay cliffs also used for nesting by Pelagic Cormorants) pre-1953 (Miller and others 1935; Jewett and others 1953). Regular breeding did not occur after these sporadic nesting events. Breeding was thought to occur at Colville Island in 1948, but empty stick nests were later clarified as those of Double-crested Cormorants (Goodge 1950; Drent and Guiguet 1961; Speich and Wahl 1989). By 1955, breeding had never been documented in the British Columbia portion of the Salish Sea, although partial nest building without egg laying was noted shortly after at Mandarte Island in 1957-1962 (Drent and others 1964). Breeding was later documented in the British Columbia portion of the Salish Sea at Race Rocks in 1987 (Campbell and others 1990) and at Mandarte Island in 2013-2015 (Carter and others 2014, unpubl. data).

DISCUSSION

Origin of Salish Sea Colonists

For Pelagic and Double-crested cormorants, Salish Sea colonists appeared to originate from long-established colonies on the outer west coast of Washington. Drent and Guiguet (1961) first identified this area as a likely source for Double-crested Cormorants that bred at Mandarte Island in 1927, but they did not make this same connection for Pelagic Cormorants that bred at Mandarte Island in 1891, possibly because they did not know if breeding records existed prior to 1891. On the outer coast of Washington, nesting Pelagic Cormorants were first documented at Tatoosh Island in 1882 (Goss 1884); a total of 21 colonies was recorded on the outer coast in 1905-1907 (Dawson 1908a,b; Speich and Wahl 1989). Double-crested and Brandt's cormorants also were found breeding at 12 and 4 colonies, respectively, in the same area in 1905-1907 (Dawson 1908a,b). All 3 species of cormorant appeared to be well-established breeders in this region by the late 19th century. In contrast, cormorants did not appear to breed on most or all of the west coast of Vancouver Island at this time (Drent and Guiguet 1961). Pelagic Cormorants were first reported breeding at Triangle Island in 1909 and may have long bred in the nearby Scott Islands (Carter and Sealy 2011c). However, the earliest reported breeding on the southwest coast of the island was in a cave near Cape Beale, Barkley Sound, in 1947 (Drent and Guiguet 1961; Carter and others 2007). Off northeast Vancouver Island, the first colony known in British Columbia had been found at Deep Sea Bluff in Queen Charlotte Strait in 1792 (Drent and Guiguet 1961), but no other colonies were reported in this strait prior to 1968 (Carter and McClaren 2016). Pelagic Cormorants also were reported breeding farther north in Haida Gwaii (Skedans Islands and Kunghit Island) in 1900 (Osgood 1901). In addition, CF Newcombe noted "glaucous wing gulls and cormorants [probably Pelagic Cormorant]" nesting on the north mainland coast at "Bird Rock" near Calvert Island (probably Upward Rock) in 1913 (Kermode 1914). From this limited information, Pelagic Cormorants appeared to breed at only a few widely spaced colonies on the outer coasts of British Columbia (outside the Salish Sea) in the late 19th and early 20th centuries, whereas Double-crested Cormorants did not appear to breed at all in these locations at this time and still do not breed outside of the Salish Sea (Rodway 1991). Thus these outer areas were unlikely to have been the source of birds colonizing the Salish Sea in the late 19th and early 20th centuries.

Potential movements of Double-crested Cormorants to the Salish Sea region from colonies at lakes in interior Oregon and Washington also cannot be dismissed as potential sources. Some lake colonies were known in the mid-19th century (such as Malheur Lake, Oregon; Baird and others 1884) and early 20th century (Vancouver Lake, Washington; Jewett and others 1953). Birds from these possible sources may have bred in the Puget Sound area or San Juan Islands, areas closer to lake colonies, prior to first known breeding at St. John's Point, Mandarte Island, and Ballingall Islets along the east coast of Vancouver Island. It is doubtful that Pelagic Cormorants colonized the Strait of Georgia through Johnstone Strait because no colonies were known in Queen Charlotte Strait at this time (Drent and Guiguet 1961; Carter and McClaren 2016). If colonists had passed through Johnstone Strait, they also may have first formed colonies in the northern Strait of Georgia. Finally, no apparent breeding by any of the 3 cormorant species occurred in the southern Salish Sea (Puget Sound) prior to 1955, and none was recorded there by the early 1980s (Speich and Wahl 1989; Carter and others 1995), presumably due to little suitable nesting habitat and greater human disturbance.

Expansion of Cormorant Nesting in the Salish Sea (1891-1955)

Initial colonization of the Salish Sea by Pelagic Cormorants apparently occurred at Mandarte Island in about 1891, with the next recorded nesting occurring at Flattop Island and Davis Bay Cliffs (Lopez Island) in 1905 (Table 1). A gradual expansion of nesting by Pelagic Cormorants seemed to occur first in the central Salish Sea before extending to the northern Salish Sea (Christie Islet) by 1924. At Mandarte Island, the population also increased from 25 or 50 nests in 1915 (Kermode 1916) to 164 nests in 1923 (Munro 1937). However, numbers of nests did not change greatly at Ballingall Islets (with limited nesting habitats) from 1934 (16 nests; Carter 2016) to 1959 (11 nests; Drent and Guiguet 1961). Between 1925 and 1955, nesting continued to expand in the central and northern Salish Sea and population size increased or decreased at certain colonies. At Mitlenatch Island, rapid growth occurred between 1949 (18 nests) and 1959 (143 nests) (Drent and Guiguet 1961). At South Peapod Island, nesting was not noted after 1949 when first observed (Speich and Wahl 1989). However, at most colonies, data were insufficient for assessing apparent trends.

For Double-crested Cormorants, sporadic breeding was first noted in the northern Salish Sea in 1896 (Hornby Island) and the central Salish Sea in 1900 (Mandarte Island), prior to regular breeding in the central Salish Sea at Mandarte Island (1927) and Ballingall Islets (1934 or earlier). Colony size increased slowly at Mandarte Island from 1927 (3 nests) to 1936 (9 nests) to 1945 (23 nests) and then more rapidly to 1953 (145 nests) (Drent and Guiguet 1961). Colony growth also occurred at Ballingall Islets from 1934 (23 nest structures, 10 active; Carter 2016) to 1957 (74 nest structures, 8 active; Drent and Guiguet 1961). Breeding in the northern Salish Sea was first noted in 1941 at Christie Islet but further expansion in this area was not noted by 1955, even though other colonies formed in the central Salish Sea during this period. At most colonies, data were insufficient for generally assessing apparent trends.

Disturbance by indigenous or non-indigenous peoples or extensive predation likely led to abandonment of certain colonies, contributing to expansion of other colonies, as well as slowing population growth. Only a few instances were documented prior to 1955:

(1) Mandarte Island: in 1915-1916, a warden (WB Anderson) was placed on the island in June and July to prevent disturbance of nesting seabirds by indigenous peoples, especially during Glaucous-winged Gull egg harvesting (Kermode 1916, 1917). "Fifty" nesting Pelagic Cormorants (25 or 50 nests) had been noted by Anderson in 1915 (Kermode 1916). In 1916, Kermode (1917:Q12) reported that "violet-green cormorants (Phalacrocorax p. robustus) [Pelagic Cormorants] have increased in considerable numbers, as is to be noted by a visit to the high cliffs on the western side of the island, where these birds build their nests." However, in 1917, a warden was not assigned to the island, and LS Higgs noted that almost every Glaucous-winged Gull nest was empty prior to fledging (Kermode 1918); presumably Pelagic Cormorants also were disturbed during extensive gull egg harvesting by indigenous people and possibly cormorant eggs also were harvested. Disturbances after 1916 apparently were not frequent or severe enough to cause long-term abandonment by Pelagic Cormorants. Nesting was next noted at their "usual" colony in 1923 (Munro 1925);

(2) Third Sister Island: in 1945, a small colony of Pelagic Cormorants formed and bred annually through 1948; in 1948, "mink" (i.e., American Mink Neovison vison) were again noted on this island, after a period of absence due to heavy trapping, and cormorants did not breed there in 1949 or afterwards (Drent and Guiguet 1961);

(3) Colville Island: Double-crested Cormorants were severely persecuted, likely leading to empty nests in 1948-1949, although breeding was noted there later in 1957 (Goodge 1950; Drent and Guiguet 1961); and

(4) Christie Islet: after 1948, numbers of Pelagic Cormorants declined due to "ever-increasing disturbance" presumably from small boats. Only a few empty nests were found in 1960; adjacent Pam Rock was used for nesting from 1948 to 1956 but afterwards abandoned (Drent and Guiguet 1961).

In addition to instances of disturbance of cormorant colonies bv indigenous peoples noted above, Pearse (1923, 1929) noted extensive egging of Glaucous-winged Gulls at Mitlenatch Island in 1920-1928, prior to later nesting by Pelagic Cormorants in 1949 (Table 1) and Double-crested Cormorants in 1993 (Chatwin and others 2002). A warden was posted and caught several individuals. In a letter dated 14 June 1923 to PA Taverner (CMNAC 11996-021), HM Laing noted: "Martuboo [sic; possibly an indigenous warden] has just returned from 8 or 9 days on Mitlenatch. Caught a lot of egg poachers, got 7 convictions I think."

Seabird Hunting and Egg Harvesting by Indigenous Peoples

The chief reason for extension of breeding by Pelagic and Double-crested cormorants from the outer coast of Washington into the central Salish Sea in the late 19th and early 20th centuries appears to be the reduction of seabird hunting and egg harvesting by indigenous peoples in both regions. A major population decline among indigenous peoples of the Salish Sea occurred between about 1860 and 1890, related to a widespread smallpox epidemic in the 1860s and other diseases and factors related to non-indigenous people (Duff 1997). Although decline in indigenous populations also occurred on the outer coast, hunting and harvesting continued there. Dawson and Bowles (1909:851) noted that "The Indians used to be very fond of Shag's [Pelagic Cormorant] eggs, which are really very good eating, and they sacked the nesting rocks once or twice each season; but it has been found necessary to put a stop to this practice upon the Olympiades." Swan (1857:271) also described a bird hunting trip by indigenous people near Point Grenville, Washington, in July 1854:
"... taking the other two canoes and my gun, [they] started off for the
rocks. They were gone nearly an hour, and, when they returned, brought
with then thirty half-fledged loons--which were the size of ducks, and
very fat--and five pelicans [i.e., Brown Pelicans Pelecanus
occidentalis]."


These 30 "half-fledged loons" most likely were cormorant chicks, and all 3 species bred in the Point Grenville area in 1905-1907 (Dawson 1908a,b). Earlier regular breeding in the Salish Sea by Pelagic Cormorants than for Double-crested Cormorants may have reflected lower impacts on Pelagic Cormorants from indigenous peoples on the outer coast. Pelagic Cormorants nest primarily on sheer cliffs, often cementing seaweed to small ledges that were less accessible to indigenous peoples hunting seabirds. After efforts to stop such harvesting, Pelagic Cormorants may have reached relatively high population levels on the outer coast earlier than the Double-crested Cormorant, promoting an earlier extension to the Salish Sea. Double-crested Cormorants prefer to place their larger nests, usually composed of sticks, on flatter substrates often at the tops of steep slopes, more accessible to indigenous peoples until seabird harvesting ceased. The availability of these habitats increased greatly on the outer west coast once indigenous harvesting stopped, potentially allowing a population increase by Double-crested Cormorants prior to extending into the Salish Sea.

The lack of recorded nesting by cormorants in the Salish Sea prior to 1891 also may reflect earlier loss of breeding colonies in this region due to heavy harvesting by indigenous peoples within the Salish Sea, long before the arrival of Spanish, English, and American people. Bovy (2011) documented thousands of bones of Double-crested Cormorant chicks in a shell midden at Lopez Island, radiocarbon dated from 300-600 AD, possibly from a local colony. However, it is not possible to distinguish if these chick bones were related to periodic breeding at a nearby location, widespread local breeding, or if chicks were harvested on the outer coast of Washington and brought by canoe to Lopez Island by indigenous peoples before consumption. Although sampling was limited, Hobson and Driver (1989) found archeological evidence of all 3 species of cormorants at 6 locations in the Strait of Georgia, but chick bones were not found, suggesting that breeding may not have been widespread in the Strait of Georgia for any cormorant species between 3500 BC and 1800 AD.

Most coastal nesting habitats in the Salish Sea are easily accessible to indigenous hunters, except for certain sheer cliffs. Cliffs were first used for nesting efforts in the Salish Sea (at Hornby Island by Double-crested Cormorants in 1896; at Mandarte Island by Pelagic Cormorants in 1891 and Double-crested Cormorants in 1927), before birds spread out to use other more human-accessible nesting habitats that became suitable habitat after indigenous hunting reduced or ceased. These cliffs also may have provided similar habitat to outer coast nesting habitats and provided some protection to early colonizing birds while hunting by indigenous peoples was still occurring at reduced levels.

Sporadic Breeding by Brandt's, Cormorants

Sporadic nesting, possibly for only one year at a time, by Brandt's Cormorants in central Salish Sea in 1928, 1940, and <1953 may have represented temporary breeding by some birds that roosted there in the non-breeding season (Miller and others 1935), although most of these roosting birds bred on the outer coast of Washington or farther south. Reasons for sporadic nesting (at least prior to 2013) are not known, but this behavior may largely reflect less suitable conditions for regular breeding near the north end of the breeding range. Long-term regular breeding appears to occur on the west coast of the Olympic Peninsula but not on the west coast of Vancouver Island. Brandt's Cormorants bred on the southwest coast of Vancouver Island from 1965 to about 2007 (Stirling and Buffam 1966; Campbell and others 1990; T Chatwin, unpubl. data), but not earlier or more recently. Earlier sporadic breeding also may have occurred at Solander Island, outside of the Salish Sea. Brooks and Swarth (1925; also see Taverner 1926, 1928) noted "nesting in numbers" at Solander Island in 1904. Subsequently, Munro and Cowan (1947:52) stated that this record was based "on the presence of birds on Solander Island in summer", which may not have been nesting. It is unclear how this conclusion was reached, and Brooks died in January 1946 so he could not dispute an error. In a letter dated 14 February 1923 to PA Taverner (CMNAC 11996-021), A Brooks confirmed "Brandt's Cormorant I saw breeding at Solander Id. W. coast V. Id......". Drent and Guiguet (1961) did not include any mention of the 1904 Solander Island breeding record but may have deferred to Munro and Cowan (1947) or relied on 1954 data when only Pelagic Cormorants bred (Guiguet 1955). Later sporadic breeding was also recorded at Sartine Island, in the Scott Islands, British Columbia (20 nests noted on 2 August 1975 and 39 nests on 29 July 1989; Vermeer and others 1976, Campbell and others 1990), as well as much farther north at Seal Rocks, Prince William Sound, Alaska (11 nests on 22 July 1972) (Sowls and others 1978). Given much evidence of sporadic nesting north of the Olympic Peninsula, it is inappropriate to dismiss the 1904 Solander Island breeding record without a complete explanation, especially given that Brooks was a very competent observer. However, it seems quite reasonable to conclude that regular breeding by Brandt's Cormorants did not appear to occur on the west coast of Vancouver Island for at least a century prior to 1965. Future monitoring will determine if current nesting at Mandarte Island becomes regular nesting or another period of sporadic nesting.

ACKNOWLEDGMENTS

Information for museum specimens was obtained through in-person museum visits or on-line through VertNet, with assistance from museum staff to provide additional details, for the following museums: Museum of Zoology at Cambridge University, Cambridge, UK (M Lowe, A Charlton, M Brooke); Natural History Museum, Tring, UK (M Adams, D Russell); Sam Noble Oklahoma Museum of Natural History, University of Oklahoma, Norman, OK (T Yuri); Royal British Columbia Museum, Victoria, BC (L Kennes); and Western Foundation of Vertebrate Zoology, Camarillo, CA (R Corado). Access to correspondence and notes of CF and WA Newcombe, housed at the British Columbia Archives (Victoria, BC), was assisted by K Bridge, C Gilbert, K Hughes, A ten Cate, K-A Turkington, and F Verspoor. Access to correspondence and notes of A Brooks, H Laing, and P Taverner, housed at the Canadian Museum of Nature Archive Collections (Hull, QC), was assisted by M Gosselin and C Dussault. Access to notes of J Hepburn, housed at UMZC was assisted by A Charlton. Colony location maps were prepared by Mapmonsters (Victoria, BC; B Calder). The manuscript benefited from careful reviews and edits from S Thomas, M Drever, and an anonymous reviewer.

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Harry R Carter (1)

Carter Biological Consulting, 1015 Hampshire Road, Victoria, BC V8S 4S8 Canada

(1) See posthumous note at end of article.

Article contact: Mark Drever, mark.drever@ Canada.ca.

Submitted 28 January 2017, accepted 13 May 2017. Corresponding Editor: D Max Smith.

IN MEMORIAM

Sadly, Harry Carter (January 1956-April 2017) died during the final preparations of 2 historical manuscripts on cormorant populations in the Salish Sea. Typical of Harry's dedication to conservation of seabirds and ensuring that records be properly kept, he wanted to encourage his fellow collaborators to carry on his important legacy. Mark Drever addressed comments from reviewers on both manuscripts, and we have carried on with "cormorant counts" in the summer of 2017.

I have known Harry since 1977 when we were employed as summer students by the Province of British Columbia, Canada, to survey all the seabird colonies on Haida Gwaii. Harry ably captained his father's converted fish boat TedMac to the storm-swept islands on the west coast of Moresby Island. On that trip I remember Harry leaping down boulders of a particularly rugged islet yelling "HOPU, HOPU"! Harry had discovered the 1st record of Horned Puffins nesting in Canada! Today, those Horned Puffin colonies, along with many other colonies, are protected in Gwaii Haanas National Park Reserve. Harry had never-ending enthusiasm, courage, and dedication to both ensuring that every islet, ledge, cliff, and crack was checked for nesting birds. Equally, he was amazing at checking facts, recording details, and tracking down historical records, then publishing this work! Lack of funding didn't stop Harry from getting out in his zodiac to remote areas to collaborate on seabird projects. Harry was generous with his time and his knowledge. His spirit was infectious and the seabird community will dearly miss him. Harry will be honoured in February 2018 with the Pacific Seabird Group Lifetime Achievement Award.

Trudy Chatwin, September 2017
TABLE 1. First-reported breeding of Pelagic, Double-crested and
Brandt's cormorants at colonies in the Salish Sea, 1891-1955.

Year                      Colony (a)                        Area (b)

PELAGIC CORMORANT
<1891                     Mandarte Island, BC               GI
1892                      Mandarte Island, BC               GI
1905                      Flattop Island, WA                SJI
                          Davis Bay cliffs, WA (Lopez       SJI
                          Island)
1920-23                   Valdes Island, BC                 GI
1922                      Skipjack Island, WA               SJI
1924                      Chain Island and Islets, BC       GI
                          Christie Islet, BC                NSG
1927                      Viti Rocks, WA                    SJI
1933                      Waldron Island, WA                SJI
1934                      Ballingall Islets, BC             GI
1936                      Jones Island, WA                  SJI

1937                      Race Rocks, BC                    IJFS
                          Colville Island, WA               SJI
                          Williamson Rocks, WA              SJI
1939-42                   Protection Island, WA             IJFS
1945-48                   Third Sister Island, BC           GI
1948                      Bare Island, WA                   SJI
1949                      Mitlenatch Island, BC             NSG
                          South Peapod Island, WA           SJI
1951                      Gordon Head, BC                   IJFS
1952-56                   Ballenas Islands, BC              NSG
1952-59                   Bare Point, BC                    GI
1955                      St. John's Point, BC (Hornby      NSG
                          Island)
DOUBLE-CRESTED CORMORANT
1896                      "South of Comox", BC (likely St.  NSG
                          John's Point, Hornby Island)
1900                      "Sidney Island", BC (likely       GI
                          Mandarte Island)
1927                      Mandarte Island, BC               GI
1934                      Ballingall Islets, BC             GI
1941                      Christie Islet, BC                NSG
1947-49                   Waldron Island, WA (Point         SJI
                          Disney)
1948                      Colville Island, WA               SJI

1949                      Viti Rocks, WA                    SJI
1951-52                   Channel Islets, BC                GI
BRANDT'S CORMORANT
1928                      Flattop Island, WA                SJI
1940                      Matia Island, WA                  SJI
<1953                     Lopez Island, WA (possibly        SJI
                          Davis Bay Cliffs)

Year                      Observer             Source

PELAGIC CORMORANT
<1891                     Unknown              Fannin 1891
1892                      W Burton             WFVZ 81,319
1905                      JM Edson, WL Dawson  Edson 1929
                          JM Edson, WL Dawson  Edson 1929

1920-23                   EA Anderson          Drent & Guiguet 1961
1922                      W Burton             WFVZ 157,884
1924                      AL Meugens           Drent and Guiguet 1961
                          RA Cumming           RBCM El 388
1927                      EJ Booth             Speich and Wahl 1989
1933                      ED Lumley            Lumley 1934
1934                      WA Newcombe          Carter'2016
1936                      WA Newcombe          WA Newcombe, unpubl.
                                               notes
1937                      TT McCabe            Drent and Guiguet 1961
                          SG Jewett            Speich and Wahl 1989
                          LB Howslev           Speich and Wahl 1989
1939-42                   WQ Wick              Wick 1958
1945-48                   A Best               Drent and Guiguet 1961
1948                      McMannama            Speich and Wahl 1989
1949                      R Fryer              Pearse 1956
                          GE Hudson            Speich and Wahl 1989
1951                      AR Davidson          Drent and Guiguet 1961
1952-56                   AG Waldron           Drent and Guiguet 1961
1952-59                   J Stainer            Drent and Guiguet 1961
1955                      HM Laing             Pearse 1956

DOUBLE-CRESTED CORMORANT
1896                      Unknown              OMNH 95

1900                      EM Anderson          RBCM E0037

1927                      W Burton             Munro 1928
1934                      WA Newcombe          Carter 2016
1941                      WS Maguire           Drent and Guiguet 1961
1947-49                   ZM Shultz            Drent and Guiguet 1961

1948                      ZM Shultz            Drent and Guiguet 1961
                                               (see Good ge 1950)
1949                      GE Hudson            Speich and Wahl 1989
1951-52                   A Best               Drent and Guiguet 1961
BRANDT'S CORMORANT
1928                      FS Rathbun           Speich and Wahl 1989
1940                      SG Jewett            Speich and Wahl 1989
<1953                     GG Cantwell          Jewett and others 1953

(a) For locations, sec Figures 1 and 2.
(b) GI (Gulf Islands), IJFS (Inner Strait of Juan do Fuca), NSG
(Northern Strait of Georgia), and SJI (San Juan Islands).
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