Specimen Record of a Long-Billed Murrelet from Eastern Washington, with Notes on Plumage and Morphometric Differences Between Long-Billed and Marbled Murrelets

Home , Ancient murrelet, Brachyramphus, Perdix

SPECIMEN RECORD OF A LONG-BILLED MURRELET FROM EASTERN WASHINGTON, WITH NOTES ON PLUMAGE AND MORPHOMETRIC DIFFERENCES BETWEEN LONG-BILLED AND MARBLED MURRELETS

CHRISTOPER W. THOMPSON, WashingtonDepartment of Fish and Wildlife, 16018 Mill Creek Blvd., Mill Creek, Washington98012, and Burke Museum,Box 353100, Universityof Washington,Seattle, Washington 98195 KEVIN J. PULLEN, ConnerMuseum, Washington State University, Pullman, Wash- ington 99164 RICHARD E. JOHNSON, Conner Museumand School of BiologicalSciences, WashingtonState University,Pullman, Washington 99164 ERICB. CUMMINS, WashingtonDepartment of Fishand Wildlife, 600 CapitolWay North, Olympia,Washington 98501

ABSTRACT:On 14 August2001, RobertDice found a Brachyramphusmurrelet approximately12 mileseast of Pomeroyin easternWashington state more than 200 milesfrom the nearestmarine waters. The bird died later that day. It had begun definitiveprebasic body molt, but not flightfeather molt. Necropsy indicated that the birdwas a female,probably in her secondcalendar year. Johnson and Thompson identifiedthe birdas a Long-billedMurrelet, Brachyramphus perdix, on the basisof plumageand measurements;it is the firstspecimen of thisspecies for Washington state. Contrary to many recent publicationsstating that Long-billedand Marbled Murreletshave white and brownunder wing coverts, respectively, we confirmedthat bothspecies typically have white under wing coverts prior to definitiveprebasic molt andbrown under wing coverts after this molt. Absence of anyextensive storm systems in the North Pacificin the dayspreceding 14 Augustsuggest that the specimen discussedhere became disoriented during postbreeding migration rather than having been blown inland by easterlywinds.

At approximately1100 PST on 14 August2001, RobertDice, an upland wildliferestoration biologist with WashingtonState Department of Fishand Wildlife,found a moribundrobin-sized brownish bird on a ruralcounty road approximately12 mileswest of Pomeroy,Garfield County, eastern Wash- ington.The locationis near the borderof Idaho, approximately210 and 270 milesfrom the nearestpoint in PugetSound and the PacificOcean, respectively.The bird was apparentlyheat-stressed because the ambient temperaturewas 90-95 ø F. Dice took the bird to the home of a localresident and tried to resuscitateit, but it died at about 1600 that afternoon. The bird was taken to WashingtonState Universityin Pullman,where Johnson tentativelyidentified it as a Long-billedMurrelet, Brachyramphus perdix. Pullenprepared the specimen(KJP 4328) as a roundstudy skin with an associatedextended wing (Spaw 1989, Winker 2000) and sent it to Thompsonto confirmits identity.

IDENTIFICATION OF THE SPECIMEN

The specimenwas a femalewith a matureovary (13 x 6 mm)and slightly enlargedova (2 mm),indicating the birdwas at leastone year old. The bird's

WesternBirds 34:157-168, 2003 157 LONG-BILLED MURRELET FROM EASTERN WASHINGTON plumage further corroboratesthis conclusion:its ventral feathers were broadlyedged with brown,giving the breastand bellya scallopedor mottled appearance, typical of adult Long-billed and Marbled Murrelets, Brachyramphusmarmoratus, in first and definitivealternate plumage (Thompsonunpubl. data). In contrast,the breastand bellyof both of these speciesare finely flecked with brown in juvenalplumage in theirfirst summer and fall, are completelywhite in first basicplumage except for a small percentageof birdsthat retain some juvenal breast and belly plumage in their first winter and subsequentspring, and are alwayscompletely white in definitivebasic plumage (Thompson unpubl. data). In addition,the birdwas in the earlystages of definitiveprebasic body molt (e.g., lesserand median wing coverts)but had not yet begunflight-feather molt. Long-billedMurrelets have billsabout 15% to 30% longer,on average, than thoseof MarbledMurrelets (Table 1); as a result,Sealy et al. (1991) suggestedthat birdswith billsexceeding 18 mm canreliably be identifiedas the Long-billed.Long-billed Murrelets also are generallylarger in overall bodysize than Marbled Murrelets, as reflectedby a varietyof standardbody measurements(Table 1).

Table 1 Measurementsof an Adult Female Long-billedMurrelet from EasternWashington in Relation to Those of Adult Female Marbled and Long-billedMurrelets a

Specimenfrom Long-billed Measurement MarbledMurrelet easternWashington Murrelet

Exposedculmen 17.4 + 0.9, 108b 19.6 21.2 + 0.7, 5c Billdepth 5.7 + 0.3, 107b 6.2 6.6 + 0.1, 5c Tarsuslength 16.9 + 0.6, 107• 19.1 18.3 + 0.3, 5c Wing (flattened) Adults" 124.1 + 3.5, 45 a.e 142.9 + 4.7, 5c Juvenalf 115.6 + 4.4, 10a 132 131.0 + 2.8, 2• Tail length "Adults" 31.7 + 1.6, 7f. • 35.9 + 1.5, 5h Juvenal• 30.9 + 0.7, 5• 31 31,--, 1•

øAllmeasurements in millimeters.Data presentedas mean + standarddeviation, n. bDatafrom "adultfemales" in BritishColumbia (Sealy 1975). ½Data from Califomia Academy of Sciences68017, Hoffman and Woolfenden (1988), and Sealyet al. (1991). a Data from specimensin Universityof WashingtonBurke Museum, University of California Museumof VertebrateZoology and CalifomiaAcademy of Sciences. e Data from probableadult females, i.e., specimensexhibiting evidence of breeding(brood patch, enlargedovaries and/or oviduct)and/or with no bursaof Fabricius;these birds have likely replacedtheir juvenalprimaries at leastonce. fDatafrom females with discemible retained juvenal plumage. These birds have not replaced their juvenalprimaries. • Data from specimensin Universityof WashingtonBurke Museum. hDatafrom Stejneger (1886), Hoffman and Woolfenden (1988), Sealy et al. (1991:U.S. National Museum599498), and Universityof WashingtonBurke Museum 44434 and 44435.

158 LONG-BILLED MURRELET FROM EASTERN WASHINGTON

Similarly,many plumagecharacters also have been argued to be reliable indicatorsof one speciesor the other in alternateplumage. It has been claimedthat in alternate plumage Long-billedMurrelets have (1) more pronouncedwhite crescentsor arcs immediatelyabove, and especially below,the eye than do MarbledMurrelets (Stejneger 1886, Ridgway1919, Jehl and Jehl 1981, Harrison 1983, Sibley 1993, Piatt et al. 1994, Konyukhovand Kitaysky 1995), (2) entirelydark plumage above their gape, whereasMarbled Murrelets often have a palestripe above their gape (Sibley 1993, Konyukhovand Kitaysky1995), and (3) a completelydark maxilla, whereas the maxilla of Marbled Murrelets is always partially white (Konyukhovand Kitaysky 1995). Lethaby's(2000) subsequentevaluation of theseplumage differences, however, found them to be unreliable. Two additionalcharacters, wing liningand outer rectrixcolor, also have been suggestedas consistentdifferences between the Long-billedand MarbledMurrelets. The winglining (underwing coverts) of MarbledMurrelets usuallyhas been described as dark, e.g., "duskybrown" to "smokybrownish black"(Coues 1868), "clearash" (Barrows 1877), "smokygray" (Baird et al. 1884), "grayishbrown" to "duskygray" (Stejneger1886), and "uniform fuscous"(Ridgway 1919). In contrast,National Geographic's Field Guide to the Birds of North America (4th ed., 2002), Gastonand Jones (1998: plate 8), and Sibley (2000) illustratethe under wing covertsof Long-billed Murreletsas beingextensively white. Similarly,Lethaby (2000) notedthat most(13 of 15) Long-billedMurrelets in alternateplumage that he examined had "obviouspale feathering on the underwing coverts" and concluded that "presenceof extensivepale in the underwing coverts is strongly supportive, but not diagnostic (italics his), for alternate-plumagedLong-billed Murrelets." However,Lethaby's tentative conclusion is incorrect.Indeed, many au- thorshave previously noted that Long-billed Murrelets have wing linings that are brownishor vary in colorfrom whiteto brown.Specifically, Dement'ev andGladkov (1951) statedthat the winglining of adultLong-bfiled Murrelets is "grayish"or "gray-brown."Sealy et al. (1991) publisheda ventralphoto of a Long-billedMurrelet specimen with its wingspartially outstretched that clearlyshows dark underwing coverts.Oka (1999) describedtwo of three Long-billedMurrelets that he assumedwere adultsas having"brown" and "darkbrown" under wing coverts.In contrastto theseprevious descriptions of underwing-covertcolor in the Long-billedMurrelet, Shibaev (1990) noted that thesecoverts may varyfrom "solidlybrownish-gray" to almostwhite. Lethaby(2000) alsonoted that 2 of 15 Long-billedMurrelets he examined had "all-dark"under wing coverts. This apparent discrepancyhas arisen becausein both the Marbledand Long-billedMurrelets the color of the underwingcoverts is an indicatorof age. In the Long-billedMurrelet this variationwas recognizedmore than a centuryago by Taczanowski(1893), who statedthat the underwing coverts of adultsare "gray-brown,"whereas those of second-yearbirds are "white with a little brown";Dement'ev and Gladkov(1951) subsequentlynoted Taczanowski'sfindings. Kozlova (1957) alsostated that underwingcoverts of adultsare "brownish-gray"whereas those of birdsin juvenalplumage are "brownishgray with somewhite."

159 LONG-BILLED MURRELET FROM EASTERN WASHINGTON

Regardingthe MarbledMurrelet, Coues(1868:64) notedthat the underwingcoverts of birdsin juvenaland first basic plumage vary from being dark as in adults(e.g., USNM 46547) to being"dusky along the edgeof the wing, but...elsewherevariegated with dull whitish"(e.g., USNM 46542). Nelson (1997) also noted that underwingcoverts of birdsin juvenalplumage are "brownishgray with somewhite," whereas those of birdsin definitivebasic and definitivealternate plumage are "uniformfuscous" and "uniformgray brown,"respectively. Lethaby (2000) subsequentlynoted that "virtuallyall" MarbledMurrelets (n: 34) in juvenal,first basic, and first alternate plumage "showedat leastsome and often extensive white in the underwing coverts," whereasall birdsin subsequentdefinitive basic and alternateplumages (n -- 33) "showedcompletely dark under wings." Thompsonexamined additional Long-billed and Marbledmurrelet specimens (see acknowledgmentsfor museums)and corroboratedthat wing- liningcolor in thesespecies is an indicationof age and not a species-specific character;nine of nine hatch-yearLong-billed Murrelets in juvenalor first basicplumage (aged by retentionof juvenalplumage and/or bursasize) had whitewing linings (underwing coverts), whereas four of fivebirds in alternate plumagehad brownwing linings.Similarly, for the MarbledMurrelet, 89% (64 of 72) of hatch-yearbirds in juvenalor firstbasic plumage had white wing linings,whereas 91% (160 of 176) of birdsin alternateplumage or definitive basic plumagehad brown wing linings.In addition,Marbled Murrelets bandedas juveniles and recapturedas second-yearbirds typically have white wing linings(E Cooke, N. Parker,L. McFarlaneTranquilla, unpubl. data). This suggeststhat in mostcases specimens in alternateplumage that have white wing liningsprobably are second-yearbirds. The specimenfound in easternWashington has a whitewing lining (Figure 1), suggestingthat it was a second-yearbird, thoughreproductively mature (discussed above). This is unusual,however, because Marbled Murrelets, and presumablyLong-billed Murrelets,are generallybelieved to reproducefor the firsttime in the third, rather than their second,calendar year of life (Beissinger1995, DeSanto and Nelson 1995). Second,although this character was not mentionedby Mlodinow(1997) or Lethaby(2000), Dement'evand Gladkov (1951), Konyukhovand Kitaysky (1995) and Shibaev (1990) stated that the outer vane of the outermost rectrices(rectrix 6) of Long-billedMurrelets has "more or lesswell-developed white marbling,""a narrow,white marginalstripe," and "narrowfringes or mottles(white with reddish),"respectively, whereas Marbled Murrelets do not exhibitthis character.Kozlova (1957) statedthat the outer rectricesof adultLong-billed Murrelets "occasionally" have "narrowwhite marginsand brownishdots," whereas birds in juvenalplumage always have "whitebars" on their outer rectrices.However, only 4 of 16 Long-billedMurrelets that Thompsonexamined possessed whitish or buffon their outermostrectrices, so absenceof this characterdoes not identify specimensas Marbled Murrelets.Conversely, none of the 464 MarbledMurrelet specimens we examinedhad any white in the outerrectrices, so sucha patternappears to be a reliablecriterion for identifyingthe Long-billedMurrelet in the hand. In alternateplumage, the mostreliable field character "for separatingthe two speciesis the palethroat of the Long-billedMurrelet: The chinand sides

160 LONG-BILLED MURRELET FROM EASTERN WASHINGTON

Figure1. Ventralview of the wingof the Long-billedMurrelet specimen (Charles R. Conner MuseumIWashington State Universitylno. 01-37) found dead in eastern Washingtonstate on 14 August2001. of the throat all the way to the base of the neck are pale.... In contrast, MarbledMurrelets do not showpale sidesto the throat" (Lethaby2000). In the hand. but to a lesserdegree in the field, the most accurateplumage characterfor discriminatingbetween these species in alternateplumage is the presenceof "cinnamon-edged"or "rufous"feathers in the mantle and scapularsof the Marbledbut not the Long-billed(Ridgway 1919, Piattet al. 1994, Lethaby 2000). The specimenfound in eastern Washingtonis clearly a Long-billed Murreleton the basisof its pale throat, lack of any rufouson its mantle (Figure2; photosalso at http://www. ups.edu/biology/museum/LBMU01- 37.jpg [roundskin]), and slightwhitish edges on its outermostrectrices. Measurementsof the specimenalso support its being a Long-billed Murrelet;its exposed culmen of 19.6 mm andtarsus of 19.1 mm are too long for a MarbledMurrelet (Table 1). The relativelyshort wing (chord132 mm) and tail (31 mm) of the specimensupport our suggestionthat it is a second- year bird,i.e., a birdwith retainedjuvenal flight feathers shorter than thoseof subsequentgenerations of flightfeathers worn by olderbirds (Table 1).

POSSIBILITY OF HYBRIDIZATION

Although the Long-billed and Marbled Murrelets appear to be more similarto one anotherin plumagecolor and morphologythan to any other alcid species,they are not sistertaxa. Genetic evidenceindicates that the Kittlitz's Murrelet, Brachyrarnphus brevirostris, is the Marbled Murrelet's sisterspecies (Pitocchelli et al. 1995, Zink et al. 1995, Friesenet al. 1996a, b, 1997). Hybridization between Kittlitz's and Marbled murrelets is un- known. Thus it is unlikelythat Long-billedand Marbled Murreletswould hybridizeeven if their breedingranges were partiallysympatric, but their breedingranges are not known to overlap. However, identifyinga Long- billed X Marbled Murrelet hybrid may be impossibleexcept by genetic methods.Examination of the populationgenetics of MarbledMurrelets from Alaskaand the AleutianIslands, including specimens from Adak (n -- 5) and

161 LONG-BILLED MURRELET FROM EASTERN WASHINGTON

,•- :...•'"...... :-• .,.-o •..,,- ....-,,. :.:...... -• ';..•.• ,- ,.. -;: :•.•.., ..... , •" .. .:.. .• ..:; . • ...-:, ß':,: ..... 'c-:' ,...t,'.'-..¾:• ",.:.:•-.•,.'.,4:•,i•**--,?• .-"•,•.*'.-.*- '.;'•;•' *• •-.e-: '*,•

...... •:;:,c:;:'t• ...... :.....: ...... Figure2. Dorsal(top) and lateral(bottom) view of the Long-billedMurrelet specimen (CharlesR. ConnerMuseum ]Washington State University] no. 01-37) founddead in easternWashington s•te on 14 August2001.

Attu (n = 9), found no evidenceof hybridizationwith the Long-billed Murrelet(Congdon et al. 2000). It is likelythat MarbledMurrelets breed as far westas Adak (Kesseland Gibson1978) and possiblyas far westas Attu (Sealy et al. 1982) in the Aleutian Islands,about 300 km east of the CommanderIslands, the easternmostpossible breeding locality of the Long- billed Murrelet.Thus, it is possible,though probablyunlikely, that these specieshybridize in the westernAleutian Islands.

BREEDING AND WINTERING RANGE OF THE LONG-BILLED MURRELET

The Long-billedMurrelet breedsin Russiaon the KamchatkaPeninsula, the KurilIslands, Sakhalin Island, and along the northernand western shores of the Seaof Okhotskand Sea of Japansouth to OlgaBay (Stejneger 1898, Labzyuk1987, Konyukhovand Kitaysky1995, Nelson 1997). There are alsosummer records from Japan on easternHokkaido. suggesting the birds may breedthere in smallnumbers, possibly only intermittently (Brazil 1991); recentevidence of breedingis lacking(Nelson et al. 1997). Summerrecords of Long-billedMurrelets from the Commander Islands(Hartert 1920, Kuzyakin1963), justwest of the Aleutians,also suggest that the speciesmay breedthere as well; however, Stejneger (1885) didnot reportseeing them in the vicinityof the CommanderIslands, and Kozlova (1957) statedthat Long- billedMurrelets "certainly do not breed"there. The Long-billedMurrelet is more migratorythan the Marbled,typically winteringabout 500 miles southof its breedinglocations, mainly in the

162 LONG-BILLED MURRELET FROM EASTERN WASHINGTON vicinityof SakhalinIsland, Hokkaido, Honshu (including inland lakes; Oka 1999), with smallernumbers as far south as coastalwaters of Kyushu, Amami-O-shima,and Kume-jima(Japan), and China, North Korea, and South Korea (Austin 1948, Vaurie 1959, Nechaev 1986, Shibaev 1990, Brazil1991, Nelson1997; seedistribution maps in Konyukhovand Kitaysky 1995, Mlodinow 1997).

POSSIBLE EXPLANATION FOR THIS VAGRANT RECORD

Vagrancyin seabirdsis common, especiallyin procellariids(Bourne 1967), but also in some alcids(Bent 1919, Grinnell 1938, Porsild1943, Salomonsen1944, Munyer1965, Nero 1968, Sealyet al. 1971, Roberson 1980, Pitmanet al. 1983). Amongalcids, however, some species are much more prone to vagrancythan others;for example,the AncientMurrelet, Synthliborarnphusantiquus (Munyer 1965, Smith 1966, Verbeek1966, Sealyand Carter 1980), Dovekie,Alle alle (Murphyand Vogt 1933, Sprunt 1938, Snyder1953), and ParakeetAuklet, Aethia psittacula(Jones et al. 2001) are frequentvagrants whereas most other alcidsare not. Becausethe Long-billedMurrelet's breeding range is muchfarther from the west coastof North Americathan the MarbledMurrelet's, one would expectmore vagrantsin North Americaof the Marbledthan of the Long- billed.However, the oppositeis true:there is not a singleinland record of a vagrantMarbled Murrelet for North America. In contrast,more than 50 vagrantLong-billed Murrelets have been recordedinland throughout the United Statesand Canada(Sealy et al. 1991, Langridge1994, DiLabio 1996, Mlodinow1997, Anon 1998, Grzybowski1998, Robersonet al. 1998, 1999, Ellisonand Martin 1999, Gilligan 1999, Lubahn 1999, Sundell1999, Tessen1999, Domagalski2000, Korducki2000, Martin 2000, Rottenbornand Morlan 2000, Ericksonand Hamilton 2001, Burgiel et al. 2002, Kratteret al. 2002), andeven Europe (Knaus and Balzari 1999, Maumaryand Knaus2000a, 2000b). This may be becausethe Long-billed Murrelet is more migratorythan the Marbled (Konyukhovand Kitaysky 1995). The currentrecord is the seventhdocumented record, first specimen record,and only inland record for Washingtonstate (Skriletz 1996, Aanerud and Mattocks1997, 2000, Mlodinow1997, Aanerud2002). Mlodinow(1997) suggestedthat the Long-billedMurrelet is morepredis- posedto vagrancybecause it is moremigratory than the MarbledMurrelet. Most vagrantLong-billed Murrelets have occurred from earlyJuly through lateAugust or fromlate October through early December (Mlodinow 1997). BecauseJuly and Augustcorrespond to the normalperiod of fiedgingand postbreedingdispersal/migration, Mlodinow (1997) suggestedthat vagrant recordsfrom this time periodmay representbirds that becamedisoriented duringnormal postbreeding dispersal/migration. Similarly, Mlodinow (1997) found a tendencyfor recordsfrom October through Decemberto be correlated with "storms that occurred off the east coast of Asia between Japan and the KamchatkaPeninsula within two to three days of each record."He alsofound records of theLong-billed Murrelet significantly more frequentwhen the mid-troposphericatmospheric circulation (about three

163 LONG-BILLED MURRELET FROM EASTERN WASHINGTON miles above the Earth's surface)tracks from the Gulf of Alaska and North Pacific into the interior of Alaska. Robert Dice, who found the specimenin eastern Washington,told Thompsonthat a severestorm had occurredin the area the previousnight. However,our reviewof broaderregional climatic conditions in Washington stateand westacross the North Pacificbetween 11 and 14 Augustfound no large-scalesevere storm systems in the PacificNorthwest or North Pacific Oceanduring that time. As a result,we suspectthat thisspecimen became disorientedduring postbreeding migration as suggested by Mlodinow(1997) but wasgrounded locally by a storm.

ACKNOWLEDGMENTS

We thank Fred Cooke, NadineParker, and LauraTranquilla McFarlane for access to unpublisheddata on Marbled Murrelets.Russ Bradley, Janet Hinshaw, Falk Huettemann,Kim Nelson,Hal Opperman,Dennis Paulson, and Terry Wahl provided citationsand photocopiesof literatureregarding the Long-billedMurrelet. Janet Hinshaw, lan Jones,Alexander (Sasha) Kitaysky, Jevgeni Shergalin, and Francis Weiseprovided translations of Russianand French literature. George Divoky provided relevant climatologicaldata. Dennis Paulsonand Joe Morlan providedextensive commentson drafts of the manuscriptthat greatly improvedthe paper. Dennis Paulsonalso provided the photographs.For accessto their birdcollections, we thank DougLong (California Academy of Sciences),Leo Joseph(Philadelphia Academy of Natural Sciences),Ned Johnson,Carla Cicero, and Chris Conroy (Museumof Vertebrate Zoology, University of California, Berkeley), Dennis Paulson (Slater Museumof NaturalHistory, Universityof PugetSound), and Rob Faucett,Sievert Rohwer,and Chris Wood (Burke Museum, University of Washington).Dennis Paulson alsohelped verify the specificidentity of the specimendiscussed in thispaper.

LITERATURE CITED

Aanerud,K. 2002. Fifth report of the WashingtonBird RecordsCommittee. Wash. Birds 8:1-18. Aanerud,K. R., and Mattocks,P. W., Jr. 1997. Third report of the WashingtonBird Records Committee. Wash. Birds 6:7-31. Aanerud,K., and Mattocks,P. W., Jr. 2000. Fourthreport of the WashingtonBird Records Committee. Wash. Birds 7:7-24. Anonymous.1998. 1997 report of the KansasBird RecordsCommittee. Kans. Ornithol. Soc. Bull. 49:26-29. Austin,O. L, Jr. 1948. The birdsof Korea. Bull. Mus. Comp. Zool. 101. Baird,S. F., Brewer,T. M., andRidgway, R. 1884. The waterbirds of NorthAmerica, vol. 2. Mem. Mus. Comp. Zool. 13. Barrows,W. B. 1877. Catalogueof the Alcidae containedin the museumof the BostonSociety of NaturalHistory, with a reviewand proposedclassification of the family.Proc. BostonSoc. Nat. Hist. 19:150-165. Beissinger,S. R. 1995. Populationtrends of the MarbledMurrelet projected from demographicanalyses, in Ecologyand Conservationof the MarbledMurrelet (C. J. Ralph, G. L. Hunt, M. G. Raphael, and J. E Piatt, eds.), pp. 385-394. U.S.D.A. ForestServ. Gen. Tech. Rep. PSW-GTR-152, Pac. SouthwestRes. Sta., Albany,CA.

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Accepted24 September2003

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