Body Condition and Survival of Vagrant Long-billed Murrelets, Brachyramphus perdix, in North America
SPENCER G. SEALY 1 and HARRY R. CARTER 2
1Department of Biological Sciences, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 2Carter Biological Consulting, 1015 Hampshire Road, Victoria, British Columbia V8S 4S8 Canada
Sealy, Spencer G., and Harry R. Carter. 2012. Body condition and survival of vagrant Long-billed Murrelets, Brachyram - phus perdix, in North America. Canadian Field-Naturalist 126(1): 6–14. From 1979 through 2009, 81 records of long-distance vagrancy in the Long-billed Murrelet ( Brachyramphus perdix ) in North America south of Alaska were examined to assess body condition and survival after first observation. Sixty-one records were of live birds, of which 38 (62.3%) were discovered at sea along the west coast of North America, 18 (29.5%) were encountered inland, and 5 (8.2%) were encountered along the Atlantic coast. Fifteen of the 20 individuals salvaged (19 adults, 1 juvenile) were discovered on lake shores (75.0%) and the other 5 (25.0%) on marine coasts; 85.0% were dead when initially found (15 dead, 2 shot), and 3 (15.0%) were moribund (2 died within one day, 1 later released). Of 10 sexed individuals, 5 were adult males, 4 were adult females, and 1 was a juvenile female. Eight of 10 murrelets observed foraging were diving on lakes, but 2 others surfaced with fish; two species of common freshwater fish were removed from stomachs of 2 birds shot by hunters. Most birds (72.1%, n = 61) disappeared after one observation, which suggests survival and mov - ing on; one bird stayed at the same location for at least 25 days before disappearing. Dead or dying Long-billed Murrelets found on shorelines of fresh water may have been too emaciated to regain lost mass after arrival—they weighed less than those shot, presumably because they were not able to locate prey or too weak to capture it. Survival for weeks or longer on freshwater stopover sites better explains how Long-billed Murrelets move across North America, with some reaching the Atlantic Ocean. Long-surviving vagrants may establish a new breeding population of Long-billed Murrelet on the west coast of North America. Key Words: Brachyramphus perdix , Long-billed Murrelet, body condition, freshwater habitat, foraging, length of stay, marine habitat, prey, stopover sites, survival, vagrancy. What are the fates of vagrant birds? Almost a cen - relet beyond its normal range in western North Amer - tury ago, Grinnell (1922) pondered this question, won - ica, before or since 1979, despite careful scrutiny of dering whether all vagrants (or “accidentals”, as they all out-of-range murrelets encountered by birders and were usually called then) perished or some individuals other field workers (see Svingen 2009). The recentness founded new populations. Data on survival of vagrants of the vagrancy of the Long-billed Murrelet appears and return to populations of origin have been gathered, to reflect in part enhanced prevailing westerly winds mainly through banding of land birds, at widespread across the North Pacific Ocean during warm phases localities in Europe (Newton 2008) and at Southeast of the Pacific Decadal Oscillation (Sealy et al. 1991; Farallon Island, about 43 km west of San Francisco, Mlodinow 1997; Carter et al. 2011). California (DeSante 1973; DeSante and Ainley 1980). Long-billed Murrelets breed in eastern Russia from Some authors have considered the implications of the northern Sea of Okhotsk and Karaginsky Gulf in vag rants in the dynamics of bird populations (e.g., the southwestern Bering Sea (~60–63° N; Kamchatka DeSante 1983; Patten and Marantz 1996; Fraser 1997; Krai [territory]) south to Vladivostok (~43° N; Pri - Veit 1997; Newton 2008), but short-term survival of morsky Krai) (Gaston and Jones 1998; Kondratyev et obligate marine birds inland with possible eventual al. 2000), but individuals have been reported in coastal return of individuals to the sea, and possibly home, waters in winter as far south as southern Japan (~30° N) has not been considered. (Nelson et al. 2002). The size of the global breeding More than 80 records of long-distance vagrancy in population is not accurately known but is likely less the Long-billed Murrelet ( Brachyramphus perdix ), than 10 000 breeding pairs (Kondratyev et al. 2000). which breeds in eastern Russia, have been reported An intriguing aspect of vagrancy by the Long-billed since the first specimen was taken in North America, in Murrelet is that some individuals are emaciated and 1979 (Sealy et al. 1982, 1991; Mlodinow 1997; Svingen apparently die soon after arriving in North America, 2009). At that time, and through the mid-1990s, this whereas others are in better condition and, if feeding form was recognized as a subspecies of the Marbled opportunities are located, they replenish their reserves Murrelet ( B. marmoratus ), that is, Brachyramphus and move on, possibly eastward as far as the Atlantic marmoratus perdix , until Friesen et al. (1996), using Ocean, or back to the Pacific Ocean. Long-billed Mur - molecular techniques, confirmed it to be a species, as relets are known to feed on fresh water year round originally described by Pallas in 1811 (Stejneger 1886). (Konyukhov and Kitaysky 1995; Oka and Hamosoto Vagrancy has not been reported in the Marbled Mur - 2001); therefore, the key to their survival inland is find -
6 2012 SEALY AND CARTER : C ONDITION AND SURVIVAL OF VAGRANT LONG -BILLED MURRELETS 7
ing a body of water with suitable prey on which to feed available, and therefore would weigh less. By contrast, while also avoiding predators. birds that were shot apparently were healthy enough to To gain a better understanding of survival for extend - feed after finding water and had thus regained lost mass; ed periods by vagrant Long-billed Murrelets, inland or these birds were expected to weigh more. Body mass at sea, we compiled (1) observations of foraging, prey (nearest 0.1 g in most cases) was usually measured capture, and prey taken by vagrants; (2) information on after birds had been frozen, in some cases for weeks or body condition of birds found dead or living when first more. We compared body mass of Long-billed Mur - encountered; and (3) data on length of stay and, hence, relet vagrants to Long-billed Murrelets weighed in survival of birds that were alive when initially encoun - Russia (Konyukhov and Kitaysky 1995). tered. We assessed the potential for survival and suc - The date of arrival of a Long-billed Murrelet at a cessive movements of vagrants that might explain oc - particular location was usually uncertain, as it is for cur rence of some individuals along the east coast of many vagrants (e.g., Armistead and Iliff 2003). Be - North America. We also consider the possibility of cause we did not know how long individuals had been long-term survival and establishment of a new breed - present at their original sites before disappearing, the ing population on the west coast of North America. length of stay was recorded as a minimum number of days. For example, we considered a Long-billed Mur - Methods relet first observed on day one but gone the next or, in Records of vagrants some cases, if no further observations were made, to In North America, observers now distinguish have spent a minimum of two days at that location (the between the Long-billed Murrelet and the resident bird likely was present for at least 12–24 hours before Marbled Murrelet of the west coast on the basis of the first observation and for at least another 12–24 hours differences in plumage (some differences are subtle, before departing, without being observed the next day, particularly in the fall and early winter) and the larg - and so on). Vagrants probably did not spend fewer than er size of the Long-billed Murrelet (e.g., Sibley 1993; 24 hours at one location, un less they were disturbed Erickson et al. 1995; Mlodinow 1997; Lethaby 2000; or they died. Maumary and Knaus 2000; Thompson et al. 2003; Foraging on fresh water and prey consumed Svingen 2009). In one case, the identity of a Long- Using the literature and museum specimens, we billed Murrelet, salvaged in New Mexico, was con - summarized details of observations of vagrant Long- firmed by molecular analysis (Witt et al. 2010). For billed Murrelets foraging on fresh water and at sea and, this paper, we examined reports of Long-billed Mur - when available, included information on prey removed relets in North America through 2009 (Svingen 2 009) from digestive tracts (hereafter “stomachs”). We accept - from the west and east coasts and the interior of North ed authors’ descriptions of the following behaviour as America. evidence of foraging and/or feeding: (1) diving, often We compiled information on use of marine and repeatedly, which some authors interpreted as feed - freshwater habitats by vagrant Long-billed Murrelets ing (some also construed from this activity that the from (1) published accounts of occurrences in North bird was in good condition), and (2) birds surfacing America (see summaries and literature cited in Sealy with and swallowing a fish. Prey items removed from et al. 1982, 1991; Mlodinow 1997; Svingen 2009); (2) stomachs of birds found dead or shot by hunters pro - persons who originally observed and/or salvaged spec - vided further evidence of feeding. imens; and (3) managers of collections with specimens of Long-billed Murrelet. Data from records with insuf - Results ficient documentation and records from the Bering Sea, Records of vagrants the Aleutian Islands, and Europe were not included in Excluding two early coastal records from Alaska, our compilation (Mlodinow 1997; Maumary and Knaus in 1845 and 1897 (Mlodinow 1997; also see Svingen 2000; Hopkins et al. 2006; Svingen 2009). Museums 2009), the Long-billed Murrelet was not reported in from which information referred to in this paper was North America until 1979, when an adult was shot by obtained are listed in the acknowledgements and in a hunter in southern Quebec (Sealy et al. 1982, 1991). Tables 2 and 3. Minimum straight-line distances of Eighty-one records in North America (south of Alaska) travel were determined using Google Maps. were available from 1979 through 2009 (20 specimens Body condition and length of stay and 61 sightings; Table 1). Data on sex, age, mass, and body condition were Thirty-eight live individuals (62.3%) were discov - derived from information in published reports of occur - ered at sea along the west coast of North America, rences and from labels on specimens and information from Washington (~48° N) to California (~38° N); 18 in museum catalogues. Ages were reported as (1) “ad - (29.5%) were encountered inland and 5 (8.2%) were ults” (after-hatching-year (AHY) birds), or (2) “juve - found along the Atlantic coast from Newfoundland niles” (hatching-year (HY) birds). We hypothesized that and Labrador (~48° N) to Florida (~27° N). One of the individuals found dead were exhausted after a long 5 specimens salvaged at sea was from California, and flight and were too emaciated to exploit prey, even if the other 4 were from Florida (Tables 1 and 2). Fifteen 8 THE CANADIAN FIELD -N ATURALIST Vol. 126
TABLE 1. Habitat type of 81 vagrant Long-billed Murrelets salvaged or observed in North America south of Alaska. Records Habitat type Observation made Specimen collected Total Ocean 43 a 5 b 48 Freshwater lake 13 10 c 23 Reservoir/dam 10 1 River/creek 40 4 Street/road/railway tracks 04 4 Brine pool 01 1 Total 61 20 81 a Thirty-eight birds, including 36 individuals and one case with two birds together (treated as separate records) were observed within 1 km of shore along the coast of Washington, Oregon, and California, and single individuals were observed in each of Newfoundland and Labrador, Massachusetts, New Jersey, South Carolina, and Florida. b One bird was salvaged from the coast of California, one from the Atlantic coast of Florida, and three from the Gulf coast of Florida. c One bird was salvaged from a marsh near the ocean in Massachusetts.
of 20 specimens (75.0%) were salvaged inland, most on The most frequent minimum length of stay was 2 lake shores (Table 1); most of these 20 (85.0%) were days (72.1%; n = 61 records), equivalent to a “one-day- dead when salvaged (15 found dead, 2 shot) and the bird” of Fraser (1997: 100). Most individuals were gone other 3 (15.0%) were moribund (2 died within one day, the day following first observation, having moved either 1 was later released). These 20 specimens were aged to another part of the same body of water or elsewhere. as follows: 95.0% were adults and 1 (5.0%) was a juve - Some individuals (9.8%) stayed for at least 3 or 4 days, nile (12 of these specimens are included in Table 2). and the longest minimum stay was 25 days. Some vag - Five of 10 individuals sexed were adult males, 4 were rants that moved on apparently were observed else - adult females, and 1 was a juvenile female (all sexed where, i.e., were observed more than once. For exam - specimens are included in Table 2). ple, a bird observed in coastal South Carolina on 19 November 1994 was gone the next day (Davis 1995), Body condition and length of stay Mass of 8 vagrants found dead ( = 7) or found but three weeks later (9–11 December) an individual n with similar plumage was photographed on a reservoir alive but died the next day ( n = 1) averaged 193.7 g (SD 20.6, range 154.2–217.8) (Table 2). Seven were about 250 km directly north, in North Carolina (Davis adults; the single juvenile weighed 207.2 g, near the and Carter 1996). The bird recovered at the edge of a upper end of the range of body mass for adult vagrants. marsh just inland from the coast of Massachusetts in All masses were below the mean of Long-billed Mur - September 1982 was molting and flightless (Table 2), relets reported from Russia—295.8 g (range 258–357) which suggests arrival in eastern North America at least (sample size and season unknown) (Konyukhov and 1–2 weeks earlier, before its wing molt rendered it Kitaysky 1995). Authors variously described vagrants flightless (see Carter and Stein 1995). found dead as emaciated with no subcutaneous fat, with Complicating our attempts to identify other possible empty and shrivelled (flaccid) digestive tracts, and with re-sightings, however, is the clustering of many inland relatively low body weights (Table 2). We assumed records in time and space. During intense El Niño con - dead birds found on lake shores were too emaciated ditions in July and August 1983, for example, three to be able to feed after arrival. Most grounded Long- birds were found dead at Mono Lake, California, and billed Murrelets also probably soon died, regardless of one at Railroad Lake, Alaska (Sealy et al. 1991). In condition; however, one grounded bird was rescued and March–December 1994 and July–November 1996, 10 released to fresh water. Two adults shot in November and 5 records, respectively, were scattered across North weighed 285.9 g with “… considerable subcutaneous America from the west coast to Florida (Mlodinow fat” (Canadian Museum of Nature 69845; Sealy et al. 1997). In any case, the short length of stay recorded 1982: 779) and 303.8 g (Smithsonian National Muse - for most individuals and the low number later found um of Natural History 582506; Mumford 1982: 191). dead suggest that a high proportion moved on after These values are 68.1 g and 86.0 g more than the heav - they were first observed. iest bird found dead (217.8 g; Table 2), respectively. Foraging on fresh water and prey consumed These individuals had fed before being shot (Table 2) Fish of freshwater origin removed from stomachs and may have survived inland for many days or weeks. and observations of ingestion of fish confirmed feed - Many individuals encountered inland apparently at - ing by Long-billed Murrelets at inland bodies of water tempted to forage before moving on after the obser - (Table 3). Both of the individuals shot by hunters had vation was made. consumed multiple individuals of at least two genera of 2012 SEALY AND CARTER : C ONDITION AND SURVIVAL OF VAGRANT LONG -BILLED MURRELETS 9
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) freshwater fish (Table 3), which could have been taken 1 6 1
1 where the birds were shot (southern Quebec (Sealy et 1
1
; : 1 0 . ) l 0 0 2
8 al. 1982) and northern Indiana (Mumford 1982)) or
a 1 2 y
5
y r t 0 d 1 e a e e i 2 - from previous foraging sites (Table 3). The two fish t
s n ( t 7
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5 l a o r J species identified—Emerald Shiner (
8 s a Notropis atheri - 1
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i i noides Dorosoma cepedianum m r t 0 M a a W t o u
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h —are common and widely distributed in freshwater o m m 2 S ( e T K A e W habitats in central and southeastern North America (Scott and Crossman 1973). Long-billed Murrelets surfacing with fish on the Moses Saunders Power Dam,
t St. Lawrence River (Di Labio 1996), and a reservoir
o ]
. n d
in Ohio (Fazio and Webb 1997), in addition to prey d e s t e a t o
a
identified above, lend credence to equating “repeated n
d i t
l ” e c s t o a y
a dives,” described by several observers, to foraging dives t o m m p w n
[ e
t (Table 3). The condition of the Long-billed Murrelet
y
m t u s d e a
a b f o
observed foraging at sea off Florida (Muschlitz 1995) s
b w d s a
t e i y d w apparently was adequate for survival after crossing
h
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e an ocean and a continent, possibly facilitated by suc -
, e
a t h d n ”
a w ’ , e o
m f t t
t cessive stopovers, regardless of the direction travelled i
t o t a h a o o t i i f g
s n n d c i from eastern Russia.
e l a n l e ‘ t
o h t … … m s i a C l “ T “ e Discussion Long-billed Murrelets travelled at least 4700 km e
p from the nearest breeding area in Karaginsky Gulf y t
d D D D (northeast Kamchatka) to Cape Flattery, Washington, r F F F o
c and some individuals survived long enough to contin - e
R ue southeast across North America, reaching Florida, 7 . which would be another 4600 km (to Miami). Travel 0 3 o 0 0 - n 2 7
across North America along a roughly northwest to 1 9 1 n 0 2 e 4 # southeast pathway from Washington to Florida was
# 4
m # i M
c probably facilitated by a continuation of the winds and W F C e S p U R storms that initially carried individuals from eastern S M C Russia to the west coast of North America (from Wash - )
g ington to northern California), followed by successive 8 0 ( . .
D s 7 7
s stopovers once in North America (Sealy et al. 1991). 1 7 N a 2 1 Length of stopover likely depended on the degree of M * * emaciation, availability of prey at the site, and presence * * e Y g Y Y of predators, as various lengths of stay were reported. H A H H A A A However, three of four individuals salvaged in Florida were underweight (Table 2) and, no doubt, in poor con - x * D
e dition when recovered, regardless of whether they had F F N S fed along the way. Survival appeared greater for indi - viduals found farther north along the east coast, from Virginia to Newfoundland and Labrador, in the cooler 3 S 0 1
0 waters of the southern Labrador Current. These indi - G 0 9 2 0 S
0
r 2
0 viduals may have altered course along the Washington
y e t 2 e b
b s t
n y
a –Florida pathway, possibly assisted by changes in wind u d o l m i g e e D u t u n J v a direction or other conditions in certain years, and more
i o n A 2
i m 1 N
4 easily found prey along the way. a
m 1 x 2 a e 2 x We assumed that vagrants did not come to North
e h
p d America directly from Europe, although Maumary and a a r n
g Knaus (2000) hypothesized a direct route from Asia o o l g t
o ) o to Europe to explain the occurrence of a Long-billed y d o h b d e p
p a
a u Murrelet in Switzerland. We would have expected many d e r e n o s e i r o n
t
i ; ; n
. more Long-billed Murrelets to have been reported in - m r . n o r o o b n e i o c c
i f ( ; o C land in Europe or on the coast, and along the east coast
C . t m
x n .
i n o e g d o 2 d c r o l c n a of North America, if the direction of travel had been C e i
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Some vagrant Long-billed Murrelets certainly sur - assume that all perish, to have a significant impact on vive for weeks during long-distance travel, even though the breeding population. We also do not know, how - survival of vagrants for more than a few days has tra - ever, how many individuals become vagrants but are ditionally been assumed to be low or zero (but see not recorded. DeSante 1973; Newton 2008). Long-billed Murrelets A high level of vagrancy may reflect undetected may have stopped to feed along the west coast of the population-level impacts being experienced in east - United States, whether they arrived directly from east - ern Russia due to natural and anthropogenic factors, ern Russia or returned from inland sites, and they may such as climate change, changes in prey and distribu - have survived for some time before and/or after being tion of predators, and loss of coastal old-growth forests observed, as has been recorded for some other vagrant (Nelson et al. 2002). Ranges may change because alcids (Sealy and Carter 2012). Once they replenished pioneering individuals that were originally vagrants their reserves, some individuals on the west coast may survive and breed successfully in new areas. Perhaps have flown inland, continuing in the same (wrong) Long-billed Murrelets will breed in North America direction, that is, away from eastern Russia; some may some day or possibly a few already have, without our have returned to eastern Russia. “Refuelled” individ - knowledge, along the west coast where most birds have uals may have survived better inland or developed suf - been observed and prey and forest habitat are similar ficient stores to attempt to return to Russia and possi - to those of eastern Russia. bly re-join Russian populations. On the other hand, those found dead or in poor condition inland may have Acknowledgements flown non-stop from Asia and possibly landed because We thank the many birders and other field workers they were near death. who carefully observed murrelets and recorded their By highlighting these survival scenarios for the behaviour, in some cases over several days, or sal - Long-billed Murrelet, we emphasize potential diffi - vaged dead and dying birds. Many individuals pub - culties with interpreting our data, despite the relative - lished their observations with detailed descriptions of ly high number of occurrences of this species across the plumage, often accompanied by photographs, and North America. We could not directly determine routes notes on the condition of specimens, or information on used by individuals before or after being recorded in habitat, weather conditions, and behaviour at the time North America, but many individuals on the west coast of observation. Many workers responded to requests apparently survived after observation, because only one for additional details on body condition and prey of dead Long-billed Murrelet was salvaged there (Table Long-billed Murrelets: R. Bowman, B. Di Labio, K. 2). In the absence of apparent direct lines of travel, Garrett, J. Hudon, F. Huettmann, J. Kendall, K. determining exact routes and time required for indi - McGowan, S. Mlodinow, B. Muschilitz, H. Nevins, viduals to fly from place to place will require use of W. Schmoker, P. Svingen, and G. van Vliet. Several satellite radios or GPS systems (e.g., Jouventin and museum curators and others provided additional infor - Weimerskirch 1990; Landers et al. 2011). General mation on specimens in their care: C. Angle (Smith - routes and patterns of movements across North Amer - sonian National Museum of Natural History), C. Cic - ica, however, can be surmised by examining samples ero and N. Johnson (Museum of Vertebrate Zoology, of records of vagrants (Sealy et al. 1991; Sealy and University of California, Berkeley); A. Kratter (Florida Carter 2012). Museum of Natural History, University of Florida, The proportion of individuals in a population that Gainesville); R. Johnson (Washington State University, become vagrants (see Fraser 1997), and whether all Pullman); and H. Nevins (Moss Landing Marine Lab - die or some establish new populations or even return oratory, Moss Landing, California). This work was to their original breeding populations, is a gap in our funded by grants to SGS from the Natural Sciences understanding of mortality and population changes in and Engineering Research Council of Canada. species. For the Long-billed Murrelet, the relatively Literature Cited high degree of long-distance vagrancy in North Ameri - Anderson, K. S. 1982. Marbled Murrelet: a first Massachu - ca, exhibited only since 1979, is remarkable in com - setts record. Bird Observer 10: 349-352. parison with other alcids and likely reflects changes Anderson, B. H. 1995. Eleventh report of the Florida Ornitho - in weather patterns that assist movements across the logical Society Records Committee. Florida Field Natu - North Pacific Ocean, as well as changes in breeding ralist 23: 38-43. habitats and populations in eastern Russia (see Carter Armistead, G. L., and M. J. Iliff. 2003. The vagrancy of Gray et al. 2011). In addition, the ability of vagrant Long- Kingbird in North America. North American Birds 57: billed Murrelets to survive for long periods enhances 148-161. the chances of discovering vagrants in North Ameri - Bailey, S. F., and D. Singer. 1996. Middle Pacific Coast. North American Birds 50: 218-220. ca and increases the likelihood of moving long dis - Barnes, S. 2009. New Jersey’s first record of Long-billed tances across the continent. Although this species is Murrelet. New Jersey Birds 35: 2-3. one of the rarest alcids, we do not consider the actual Carter, H. R., and J. L. Stein. 1995. Molts and plumages in numbers of vagrant individuals recorded, even if we the annual cycle of the Marbled Murrelet. Pages 99-109 2012 SEALY AND CARTER : C ONDITION AND SURVIVAL OF VAGRANT LONG -BILLED MURRELETS 13
in Ecology and conservation of the Marbled Murrelet. Konyukhov, N. B., and A. S. Kitaysky. 1995. The Asian Edited by C. J. Ralph, G. L. Hunt Jr., M. G. Raphael, and race of the Marbled Murrelet. Pages 23-29 in Ecology and J. F. Piatt. U.S. Forest Service, General Technical Report conservation of the Marbled Murrelet. Edited by C. J. PSW-GTR-152, Pacific Southwest Research Station, Ralph, G. L. Hunt Jr., M. G. Raphael, and J. F. Piatt. U.S. Albany, California. 420 pages. Forest Service, General Technical Report PSW-GTR-152, Carter, H. R., S. K. Nelson, S. G. Sealy, and G. B. van Pacific Southwest Research Station, Albany, California. Vliet. 2011. Occurrences of Kittlitz’s Murrelets south of 420 pages. the breeding range along the west coast of North America. Kratter, A. W., T. Webber, T. Taylor, and D. W. Steadman. Northwestern Naturalist 92: 186-199. 2002. New specimen-based records of Florida birds. Bul - Davis, R. 1995. Southern Atlantic Coast region. National letin of the Florida Museum of Natural History 43: 111- Audubon Society Field Notes 49: 135-139. 161. Davis, R., and D. Carter. 1996. First record of Marbled Mur - Landers, T. J., M. J. Rayner , R. A. Phillips , and M. E. relet in North Carolina. Chat 60: 27-28. Hauber . 2011. Dynamics of seasonal movements by a DeSante, D. F. 1973. An analysis of the fall occurrences and trans-Pacific migrant, the Westland Petrel. Condor 113: nocturnal orientation of vagrant wood warblers (Parulidae) 71-79. in California. Ph.D. dissertation, Stanford University, Palo Lethaby, N. 2000. The identification of Long-billed Mur - Alto, California. relet in alternate plumage. Birding 32: 438-444. DeSante, D. F. 1983. Annual variability in the abundance of Mactavish, B. 1989. Atlantic Provinces region. American migrant landbirds on Southeast Farallon Island. Auk 100: Birds 43: 1285-1288. 826-852. Maumary, L., and P. Knaus. 2000. Marbled Murrelet in DeSante, D. F., and D. G. Ainley. 1980. 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Ohio Cardinal 20: 1-6. conservation of the Long-billed Murrelet in Japan. Journal Fraser, P. A. 1997. How many rarities are we missing? Week - of the Yamashina Institute for Ornithology 33: 88-106. end bias and length of stay revisited. British Birds 90: 94- Newton, I. 2008. The migration of birds. Academic Press, 101. New York. 976 pages. Friesen, V. L., J. F. Piatt, and A. J. Baker. 1996. Evidence Oka, N., and H. Hamasoto. 2001. Nutritional condition and from cytochrome B sequences and allozymes for a ‘new’ muscular features of wintering Long-billed Murrelets in species of alcid: the Long-billed Murrelet ( Brachyram - Japan. Condor 103: 874-878. phus perdix ). Condor 98: 681-690. Patten M. A., and C. A. Marantz. 1996. Implications of Gaston, A. J., and I. L. Jones. 1998. The Auks: Alcidae. vagrant southeastern vireos and warblers in California. Oxford University Press, Oxford, U.K. 349 pages. Auk 113: 911-923. Grinnell, J. 1922. The role of the “accidental.” Auk 39: 373- Scott, W. 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Cummins. 2003. Specimen record of a Long-billed Mur - 141 pages. relet from eastern Washington, with notes on plumage 14 THE CANADIAN FIELD -N ATURALIST Vol. 126
and morphometric differences between Long-billed and Witt, W. C., M. S. Graus, and H. A. Walker. 2010. Molec - Marbled murrelets. Western Birds 34: 157-168. ular data confirm the first record of the Long-billed Mur - Veit, R. R. 1997. Long-distance dispersal and population relet for New Mexico. Western Birds 41: 160-167. growth of the Yellow-headed Blackbird Xanthocephalus xanthocephalus . Ardea 85: 135-143. Received 29 November 2011 Accepted 9 May 2012