Prob ble Yellow-belliedS ,psuc,cer (Sphyr icusv flus)x Red-breasted Sapsucker(S. ru r) bri frome st- ern Kansas,with commentson th field identifiction of adultsapsuckers
Abstract 1995),both before and after the recognition of Mark B. Robbins A sapsucker(Sphyrapicus) found 15 Decem- thesesapsuckers as separatespecies (AOU ber 2001 near Lawrence,Kansas was collected 1985), Robbinscollected the bird on 20 De- NaturalHistory Museum andstudied through molecular methods (mi- cember2001 (KUNHM 92332; Figure 3) and tochondrialDNA sequencingand allozyme preservedtissue for analysisby Cicero. UniversityofKansas analysis),consultation with researchers work- ingon hybridization ofsapsuckers, and analy- Age,Sex, Plumage, and 1345Jayhawk Boulevard Geneticsof the Lawrencebird sisof 201 adultmale specimens of Red-naped Lawrence,Kansas 66045 (S. m•chalis)and Yellow-bellied(S. vatit,s) Upon dissection,the bird surprisingly Sapsuckers.We concludethat the individual provedto be an adult female with a granular (eraall:[email protected]) sapsuckerin question is nota Red-napedSap- ovarymass 3.5 x 2 min.Internally, the size suckeras originally identified in thefield but andshape of thegonads coupled with the ab- rathera hybrid,most likely the result of a pair- sence of the bursa of Fabricius indicated that ingof a femaleYellow-bellied with a male Red- thebird was at leasta yearand a halfold. The David E. $eibel breastedSapsucker (S. tuber), thus potentially uniformlydark and freshprimary coverts, thefirst specimen of thishybrid combination theuniformly adult secondaries, and the uni- JohnsonCounty Community College awayfrom the very limited contact zone of formlyadult flight feathers indicate that the ScienceDepartment thesespecies in BritishColumbia. We offer birdwas at leastfour calendar years old (Pyle commentson pitfallsin theidentification of 1997).In additionto havinga typicalYellow- OverlandPark, Kansas 66210 sapsuckersbyplumage, as well as an overview belliedback and ventral pattern (i.e., inten- of reportsof similarsapsuckers in eastern siveyellow underparts), it alsohad the out- (eraall:[email protected]) NorthAmerica. especially the Great Plains re- ermosttwo pairs of rectrices(numbers 4 and gion,where unusually plumaged sapsuckers 5) heavilymottled and tipped with white (in- have occasioneddebates for over a decade. termediatebetween Figure 129A and Figure Carla Cicero 129Bin Pyle[1997]). Adult male Red-naped Field Encounter and Red-breastedSapsuckers (S. ruber)have MuseumofVertebrate Zoology On 15 December 2001, Seibel discovereda the outerrectrix mostly black, and adult fe- sapsuckerwith unusualplumage on the maleRed-naped and Red-breastedhave less UniversityofCalifornia Lawrence,Kansas Christmas Bird Count near mottlingthan femaleYellow-bellied. espe- Lawrence.This individual was initially identi- ciallyon rectrixfour. Berkeley,California 94720 fied as an adultmale Red-naped Sapsucker Mitochondrial(mtDNA) sequencing of a (Sphyrapicusnuthalls) because ofthe presence diagnostic300 bp fragmentof the mtDNA (eraall:ccicero@berkeley. edu) of anextensive red nape and the complete in- cytochrome-b(Cicero and Johnson1995) cursionof redin theblack malar (Figure 1). identifiedthe bird as a Yellow-belliedSap- However,Seibel and othersrecogmzed that sucker.This resultwas confirmed by addi- quencydifferences at purine nucleoside thebird was atypical for an adultmale Red- tionalsequencing of theentire cytochrome- phosphorylase(NP) and aminopeptidase napedbecause of extensivered suffusion on b genefor the Lawrencebird and known leucyl-glycyl-glycine(LGG). Ciceroana- the broad black chest band. When Robbins individuals of S. m•chalis(MVZ 168594) and lyzedLGG for the Kansassample, using the wasshown photographs of the bird, he recog- S. varms (MVZ 168650). However,because sametwo MVZ specimensfor standards, and nizedthat the backpattern (Figure 2) was mtDNA is maternally inherited, the se- foundthe Kansas bird to behomozygous for typicalof an adultYellow-bellied (S. varius). quencedata only confirmed that the mater- the allelethat occurs in higherfrequency in Furthermore,his pastexamination of adult nalparent was varius. S. nuthallsthan S. varius.Attempts to score malesapsucker specimens (see below) had In an attemptto identifyboth parents, Ci- thesame samples for theNP locuswere am- shownthe presenceof extensivered on the ceroalso used allozymes following the work biguous.A reviewof theraw allozyme data napeto be an unreliable character for separat- ofJohnson and Zink (1983).Their study did usedin Johnsonand Zink (1983) revealed ingthese two sapsucker species. Knowing that not find tixed differencesbetween any that two of the eightS. varit•sthat theyex- geneticstudies had been done on sapsuckers speciesof Sphyrapicusother than S. thy- aminedlikewise were homozygous for this (Johnsonand Zink 1983,Cicero and Johnson wideus,although there were notablefre- allele; the other six birds were either het-
360 NORTH AMERICAN BIRD rubel daggetti)and Red- While reviewinga controversialMissour• re- naped Sapsucker--also port of a Red-napedin 1993 (seebelow), showsa brightyellow belly. Robbins examined a total of 120 adult male From a distributional stand- Yellow-belliedSapsucker and 81 adultmale point, a hybrid event be- Red-napedSapsucker museum specimens. tween Yellow-bellied and Of the adult male Yellow-belliedsexamined, nominate Red-breasted is 23 (19%)had red in thenape ranging from a far more probablethan a few red feathers that would not be visible in Yellow-bellied x S. ruber thefield to twoindividuals taken during mi- daggettievent (Campbell et gration near Philadelphia,Pennsylvania al. 1990, A.O.U. 1998). (ANSP 173632 and 173633) that had as Because of the wide much red in the nape as averagefresh- rangein variationin hybrid plumaged,adult male Red-napeds.Except phenotypesamong all three for thered nape, the plumage of bothof the of thesesapsucker species lattertwo individualsappeared to be classic (Howell 1952,Johnson and Yellow-bellieds.In Howell (1952), E. God- Johnson 1985, J. Hudon, frey relatedthat a specimenfrom Megantic M. Gossefin,pers. comm.), County,QuEbec, Canada had an extensive it is uncertain if the amountof red on the napebut showedno Lawrence individual is an otherRed-naped characters. A morerecent F1 or theresult of a back- Quebecrecord of a breedingadult male Yel- cross.Interestingly, except low-belliedwith an extensivered napewas for the Yellow-bellied back documentedwith a color photographat pattern of the Lawrence Saint-Jacquesde Leedsin 1988 (American bird, it is very similar to Birds42: 1225). specimensof Red-naped Sapsuckerx Red-breasted Malar Figure1. Adult female probable Yellow-bellied Sapsucker x Red-breasted Sapsucker, 16 Sapsuckerhybrids of both The coloration of the dark malar in adult December2001, near Lawrence, Kansas. Note that the dark malar is mostly red and the redof the throat masks most of the black chest band. The prominent red nape is visible sexes,e.g., Canadian Muse- malesapsuckers is a valuablecharacter for evenin this profile. Photograph byl(yleGetstrier. um of Nature,Ottawa spec- identification.Adult male Red-napeds usual- imens 47951 and 52385 ly havesome encroachment of thered throat erozygotesor homozygousfor a secondal- (Devillers 1970,Johnson and Johnson 1985, onto the malar.However, this can vary con- lele. Therefore,the allozymedata for the M. Gossdin,pers. comm.). sidembly,ranging from birds that have an all- Lawrencebird are not unexpectedbut un- fortunatelydonot provide evidence toiden- DISCUSSION tifythe paternal parent of thisindividual. Althoughthe genetic data indicate that the Plumagecharacters femaleparent was a Yellow-bellied,we are and their relevance for identi- limitedto an analysisof plumageto inferthe fication identityof the male parent.There are two plausible hypothesesfor the male-like Nape plumagein thisindividual. It is conceivable • mentionedabove, the pres- that the abnormal amount of red on the ence or absence of red in the throatand upper breast are the result of a ge- napeis an unreliablecharacter neticabnormality or an unknownaging phe- for separatingYellow-bellied nomenon,perhaps related to reducedhor• and Red-napedSapsuckers. monelevels (e.g., estrogen). Older females in Kaufman (1990) correctly somespecies may acquire male-like plumage, pointed out that some Red- e.g.,Summer Tanager (Pyle 1997, Owens and napeds,especially worn indi- Short1995). However,our discussionswith J. vidualsin late summer,lack red Hudonand M. Gosselin,who examined pho- on the nape.In mid-July2004, tographsof theLawrence specimen, and our Robbins conducted field work examinationof hybridshave led us to believe m the SouthDakota/Wyoming that the mostlikely explanation for the un- Black Hills and observed 14 usualdegree of redin theLawrence bird is a adultRed-naped Sapsuckers (in hybridizationevent between a femaleYellow- which nape coloration was belliedand a maleRed-breasted Sapsucker. clearlyvisible) with thefollow- As mentioned above, the Lawrencebird ing distribution:one female hasback, rectr•x, and facial patterns of a Yel- with no red, two females and low-bellied,whereas the extensionof the red two maleswith onlya hint of of thethroat masking the black breast band is red, and five birds that had exhibitedonly in Red-breasted.Although the moderate to extensive red Figure2. Dorsalview of the same Lawrence female depicted inFigure 1.The red brightyellow underparts may simply be the napes. napeis conspicuous andthe malar, which ist•pically black in both sexes ofYellow- resultof Yellow-belliedgenes, we notethat Kaufman (1990) also noted bellledand in fernale Red-naped, isheavily suffused with red. However, the t•pical the northern(nominate) subspecies of Red- that some adult male Yellow- Yellow-belliedbackpattern--i.e., the extensive light-colored plumage inproportion breastedunlike the southern subspecies (S. belliedshave red in the nape. toblack, isapparent. Photographbyl(yleGerstner.
OLUME 59 (2005) NUMBER 2 361 Figure3. Ventral view of Lawrence female (KUMNH 92332). Note that the red throat masks most of the black chest band. This feature isunknown inpure Yellow-bellied andRed- napedbut is characteristic ofRed-breasted. Thebright yellow underparts aretypical for Yellow-bellied andthe nominate subspecies ofRed-breasted. Photo9ral•h t•)'ldar•Robbins.
black malar to more extreme cases in which oredarea in the back,especially in mid- to Hybridization the black malar is nonexistent--i.e., the late summer,when birdsare heavilyworn It has been known for some time that the malarregion is entirely red. Robbins observed andmay appear washed out (Robbins,pers. identificationof a sinallpercentage of adult five of sevenadult male Red-napcds during obs.).Nonetheless, female Red-napeds still Yellow-bellied,Red-nape& and Red-breasted mid-Julyin the BlackHills that appeared to possessnoticeabk more black in the back Sapsuckersis problematic,both because of haveall-black malars. Given that red pigmen- thaneither sex of Yellow-bellied(Figure 3). overlapin plumagecharacters among these tationin sapsuckersispresent only in thetips Xdditionally,the reducedpale area in adult speciesand because of hybridizationamong of feathers,this can be easilyworn off, re- Rcd-napedsis typicallywhiter than that of them. Kaufman (1990) underscored that vealingthe all-blackremainder of the feath- Yellow-bellicds;in the latter, this pale region someadult individuals of thiscomplex cannot ers. However,this should not be an identifi- is oftenwashed with a light buffy-yellow.It be identified under field conditions. Until re- cationcouccrn outside the periodfrom late shouldbe notedthat the paledorsal region cently,hybridization between Yellow-bellied June throughAugust. Female Rcd-napeds of nominateRed-breasted also is yellowish. andRed-naped Sapsuckers was thought to be typicallydo not haveany red in the black "limited and localized" (Howell 1952, AOU malar,and we are unax• are of an,•Ycllow-bel- Throat 1998); however,unpublished details from lieds.x• ith theexception of possiblehybrids. In addition to unusual individuals like the western Alberta indicate that there havingred in theblack malar. Thus, if redis kawrencesapsucker, sexing of adult Red- is "extensive"hybridization (l. Hudon,pers. presentin the malarand thereis no unusual napcdSapsuckers can be problematicbe- comm.;
362 NORTH AMERICAN BIRDS Lawrencebird (Howell1952, Campbell et al. Summary Picidae)•n Oregon.Bulletin of the Southern 1990,Walters et al. 2002,M. Gosselin,pers. Great care must be exercised in the identifi- CaliforniaAcademy of Sciences76: 38-41. comm.). As Howell (1952) noted,the identi- cationof sapsuckersbecause of overlapin Campbell,R. W.,N. K. Dawe,1. McTaggart- ficationof a variusx tuberhybrid collected plumagecharacters and becauseof hy- Cowan,J. M. Cooper,G. W. Kaiser, by H. S. Swarthat TelegraphCreek, north- bridization.The proportionof black vs. andM. C. E. McNall.1990. The Birds of western British Columbia (MVZ 39781) was whiteon the backand the presenceor ab- British Columbia. Vol. 2. Diurnal birds of based more on the fact that it was well north sence of red in the dark malar in adult males preythrough woodpeckers. University of of the nuchalis/ruber contact zone than on arethe most reliable plumage characters for British ColumbiaPress, Vancouver. plumagecharacters. Indeed, our examira- separatingRed-naped and Yellow-bellied Cicero,C., andN. K.Johnson. 1995. Specm- non of the specimenindicates there are no Sapsuckersunder field conditions. The pres- tion in sapsuckers(Sphyrapicus): II1 obvious variuscharacters, and if it were not enceof redin thenape should only be con- Mitochondrial-DNAsequence divergence for the localitywe would classifyit as a sideredsuggestive of Red-naped;the other at the cytochrome-blocus. nuchalisx tuberhybrid. Unlike the Lawrence twocharacters should be observedcarefully Auk 112: 547-563. presumedvanus x tuberhybrid, the Tele- in orderto distinguishthese species. Hy- Devillets, P 1970. Identification and distri- graphCreek bird hasa backpattern typical bridizationshould be consideredas a possi- butionin Californiaof theSphyrapicus of nuchalis and tuber. Mixed varius x ruber bilityfor anysapsucker that has a combina- variusgroup of sapsuckers.Califorma pairs have, however, been documented tion of the abovekey charactersand hasan Birds 1: 47-76. throughfieldwork in 1969by W.E.Godfrey unusualamount of red in otherregions of Grzybowski,J. A. 1993.The Winter Season: (summarizedin Campbellet al 1OO0)and in thehead and chest. The sapsucker from east- SouthernGreat Plains Region. American 1974by Scottet al. (1976). ern Kansasdocumented herein potentially Birds 47: 272-273. In additionto the examplesabove, we representsthe firstspecimen of Yellow-bel- Hamilton. R. A., andJ. L. Dunn. 2002. Fea- mentionthe followingrecords to underscore liedSapsucker x Red-breasted Sapsucker hy- turedPhoto. Red-raped and Red-breasted the extento[ unusual-appearingsapsuckers bridaway from the very limited contact zone Sapsucker.Western Birds 33: 128-130. in central and eastern North America. A bird in British Columbia. Howell,T. R. 1952.Natural history and dif- thatwas initially reported as an adultmale ferentiationin the Yellow-belliedSap- Red-napedin April1993 in JacksonCounty, Acknowledgments sucker. Condor 54: 237-282. Missouri was either a Yellow-bellied or a Yel- We especially want to thank Jocelyn Johnson,N. K.,and R. M. Zink.1983. Speci- low-belliedx Red-napedhybrid. Although Hudonand Michel Gossefinfor sharing ation in sapsuckers(Sphyrapicus): I. Ge- thisbird hadan extensivered nape,quite their unpublisheddata on sapsuckerhy- netic differentiation. Auk 100: 871-884. similar to the Lawrencebird, it too had the brids. Kyle Gerstner(Lawrence) and Dan Johnson,N. K., and C. B. Johnson.1985. classichack pattern of a Yellow-bellied,and Thalmann(Washington) provided splen- Speciationin sapsuckers(Sphyrapicus): 11. the malarstripes were entirely black (color did photographsof the Kansasbirds. Syrupairy,hybridization, and mate prefer- photographsin MissouriBird RecordsCom- JosephGrzybowski and Normand David encein S. tuberdaggetti and S. nuchalis. mitteefile #1993-33). A similarbird was pho- providedcolor photographsof the Okla- Auk 102: 1-15. tographed12-19 February 1993 at Tishomin- homa and Quebecbirds, respectively. Kaufman,K. 1990.A FieldGuide to Advanced go National Wildlife Refuge,Johnston CaseyTucker provided details of the Ohio Birding.Houghton Mifflin, Boston, Massa- County,Oklahoma (Grzybowski [1993] in- bird. John Gerwin shared his observation chusetts. cludesblack-and-white photograph; we also of the North Carolinasapsucker. Peter Lockwood,M. W., and C. E. Shackelford. examineda color image).Another similar Pyle, AlvaroJaramillo, Brian L. Sullivan, 1998. The occurrence of Red-breasted sapsuckerwas documented26 February TonyLeukering, and Steve Mlodinow made and suspectedhybrids with Red-raped 2002 at Washington,Washington County, helpful commentson the manuscript. Sapsuckerin Texas.Bulletin of theTexas Kansasby DanThaimann. Both of thesebirds Specimenswere examined at the following OrnithologicalSociety 31: 2-6. werepresumed to be males and had red napes institutions:Academy of NaturalSciences, Owens, I. P E, and R. V. Short. 1995. Hor- but otherwiseappeared as typical Yellow-bel- Philadelphia(ANSP), AmericanMuseum monal basisof sexualdimorphism in lied. An adult sapsucker,of unknownsex, of NaturalHistory (AMNH), Museumof birds:Trends in Ecologyand Evolution 10: that appearedin HolmesCounty, Ohio 3-8 VertebrateZoology, University of Califor- 44-47. April 2005 (underreview by the OhioBird nia-Berkeley(MVZ), and Universityof Phillips,A. R.,J. Marshall,and G. Mortson. RecordsCommittee; Casey Tucker, pers. Kansas Natural History Museum 1964.The Birds of Arizona. University of comm.)may havebeen a Yellow-belliedx (KUNHM). Possiblehybrids were loaned Arizona Press,Tucson, Arizona. Red-napedhybrid. The Ohio bird had a red to us by U.S. NationalMuseum of Natural Pyle,P 1997.Identification Guide to North napeand rotsflyred malars,but the ventral History,Smithsonian Institution (USNM), American Birds. Part 1. Columbidae to yellowwash was more typical of a Yellow-bel- andOccidental College (Moore Laboratory Ploceidae.Slate Creek Press, Bolinas, Cal- liedand the back pattern appeared intermedi- of Zoology,MLZ). ifornia. ate betweenthese two taxa.In Appalachian Scott,D. M., C. D. Ankney,and C. H.Jarosch. NorthCarolina, John Gerwin observed a pre- Literature cited 1976. Sapsuckerhybridization in British sumedadult maleYellow-bellied Sapsucker AmericanOrnithologists' Union IAOU]. Columbia. Condor 78:253-257 with a conspicuousred nape and inordinate 1985. Thirty-fifth Supplementto the Shunk,S. A. 2005. Sphyrapicusanxiety: "orange-yellow"patches, nearly forming a American identifyinghybrid sapsuckers. Birding 37: band,on theupper chest on 28 March2004 Ornithologists'Union Check-list of North 288-298. in MosesCone National Park, Watauga American Birds. Auk 102: 680-686. Walters, E. L., E. H. Miller, and P E. County.Except for the chestbeing orange- AmericanOrmthologists' Umon [AOU]. Lowther.2002. Red-breastedSapsucker yellowinstead of red,the North Carolinabird 1998. Check-listof NorthAmerican birds. (Sphyrapicustuber) and Red-napedSap- apparentlywas very similar to the Lawrence Seventhedition. American Ornithologists' sucker (Sphyrapicusnuchafis), in: A. sapsucker.The b•rd was not collected or pho- Union,Washington, D.C. Pooleand E Gill,eds. Birds of North Amer- tographed;if it wasindeed a male,one won- Browning,M. R. 1977.Interbreeding mem- ica, no. 663. Birdsof North America,Inc., derswhether it couldhave been a hybrid. bersof theSphyrapicus varius group (Aves: Philadelphia,Pennsylvania. •
VOLUME 59 (2005) NUMBER 2 363