Molt Limits in North American

Peter Pyle Point Reyes Observatory 4990 Shoreline Highway Stinson Beach, CA 94970

ABSTRACT American passerines are complete (Pyle et al. 1987), the presenceof moltlimits indicates HY/SY "Molt limits," or the boundaries between replaced and (first-year), at leastuntil the prealternatemolt, retained wing feathers and rectrices during partial or and often untilthe secondprebasic molt (Mulvihill incompletemolts, were investigatedin 288 speciesof North American passerines through the examination of over 1993, Jenni and Winkler 1994). Thus, molt limits 16,000 specimens. Thirty-sixspecies showed evidence of can be especiallyuseful for ageing North American completefirst prebasic molts, including27 species that did passerines in winter and spring, after first-year not have extensive prealternate molts (those includingat birds have typicallycompleted skull pneumatiza- least some greater coverts) and nine species that did have tion. Patterns of replacement among the wing extensive prealternate molts. Of the remaining252 species, 183 did not have extensive prealternate molts and 69 feathers vary substantiallyboth among species species did. Fifty-fourspecies showed evidence of partialor and among individualsof the same species. This incomplete replacement of primaries, during the first variationis very poorlydocumented for most North prebasic molt, the prealternate molt, or both, in at least a American species (see Mulvihill 1993). Addition- proportionof the populations.The replacement pattern of primarieswas either"eccentric" (proceeding distally from the ally, some species undergo partial or incomplete center of the primaries;46 species), or "typical"(proceeding prealternate molts in both HY/SY and AHY/ASY distallyfrom the innermostprimary; 8 species).In six species (adult)birds (Pyle et al. 1987, Mulvihill1993). To use that showed an eccentric replacement pattern, a small molt limits effectively,therefore, variation in the proportion of individuals also showed the typical replace- extentof replacementduring the firstprebasic molt, ment pattern. Data on variation in the extent of greater- covert, secondary, rectrix, primary, and primary-covert and the occurrenceand extent of prealternate molts replacement for each partial or incomplete molt in each (especially in AHY/ASYs), must be known. To species, referencesto previousdetailed studies on molt, and assess variation in the location of molt limits notes on geographicvariation, discrepanciesbetween the resulting from partial and incomplete, results of this study and that of previous work, and other interestingcases, are presented. presupplemental,first prebasic, first prealternate, and adult prealternate molts of North American INTRODUCTION passerines, I examined over 16,000 specimens of 288 species. The results of this examination are In most North American passerines, the first presented here. prebasicmolt is "partial"or "incomplete,"some but not all feathers being replaced(Pyle et al. 1987, METHODS Mulvihill1993). Recently,Jenni and Winkler(1994) have illustrated the utility of "molt limits," the Specimensexamined for this study were locatedat boundaries between replaced and retained the California Academy of Sciences (CAS), feathersthat resultfrom partial molts,in ageing Museumof VertebrateZoology (MVZ), PointReyes such passerines.Retained juvenal wing coverts Bird Observatory (PRBO), Natural History and flight feathers are relativelyworn and often Museum of Los Angeles County (LACM), San showmore subduedcolor patterns than adjacent, Diego Natural HistoryMuseum (SDNHM), Moore supplementalor first-basicfeathers. Because adult Laboratory of Zoology (MLZ), and Western (definitive)prebasic molts in virtually all North Foundationof Vertebrate Zoology (WFVZ). On Apr.- Jun, 1997 NorthAmerican Bird Bander Page 49 each specimenthe wingcoverts and flightfeathers An attempt was made to sample at least 15-20 were studiedcarefully for evidenceof partial or specimensfor each moltwithin a species,from as incompletemolts. The numberof replacedwing wide a geographicrange (withinNorth America) as coverts and flight feathers were recorded on all specimenmaterial allowed. Although all collections birds showingevidence of incompletefeather were locatedin California,a significantproportion of replacement,after active molting had ceased. Both specimens were collected from other localities wingswere examinedon each specimento ensure throughout . Larger samples of that resultswere basedon incompletemolts rather specimens were examined for species showing than adventitiousreplacement; specific data were wide geographicvariation or complex incomplete taken fromthe rightwing. molts,and smallersamples indicate fewer available specimens. Ranges in the number of feathers The age of each bird when it was collected was replaced,within each tract, are presentedas mean determinedby informationon the specimenlabels, _+twice the standarddeviation. These ranges esti- the presenceof molt limits(assuming that adult mate what wouldbe expectedfor 95% of the popu- prebasicmolts are complete),and the color and lation, assuming a normal distributionto replace- amountof wear to the primarycoverts (Figure 1), ment patterns (Pyle 1997). whichtypically are retained,at leastin part, by birds undergoingincomplete molts (see below). Other When the results of this examination contradicted plumagecriteria (Pyle et al. 1987) were usedwhere thoseof otherpublished or unpublishedinformation appropriate.Age terminology follows that ofthe Bird (see the Tables for other references discussing BandingLaboratory (Canadian Wildlife Service and molt in North American passerines),specimens U.S. Fishand Wildlife Service 1991). Terminology were reexamined to either confirm or correct the of molt,plumages, and feather generations follows original data of this study, before they were Humphreyand Parkes(1959; see alsoThompson tabulated. and Leu 1994). Plumagecharacters, along with date and location of collection, were used to RESULTS AND DISCUSSION determine whether observed molt limits resulted from the presupplementalmolt "PS" (Thompson Molt patterns in the 288 North American species and Leu 1994), the prebasic molt "PB", or the examined could be categorized into several prealternatemolt "PA". groups. In 27 species (9.4%), data indicatedthat replacement of wing coverts and flight feathers Fig 1. Shapeand relativecondition of the primarycoverts in duringboth the first and adult prebasicmolts was HY/SY and AHY/ASY passerines,in fresh (fall) and worn (spring)condition. The contrastbetween these feathereand typicallycomplete, and that the prealternatemolts replaced greater coverts is very useful in ageing many were either absent or limited,including no greater species. Note that the edging on these feathere is often coverts or flight feathers. These species were: present but thinnerin HY birdsthan in AHY birds in the fall, Northern Beardless-Tyrannulet (Camptostoma and it is oftenabsent in SY birdsbut still present in ASY birds imberbe),the two speciesof wood-pewees,Alder in the spring. Flycatcher (Empidonax alnorum), Horned Lark (Eremophilaalpestris), the eightspecies of martins and swallows, Bushtit (Psaltriparus minimus), Wrentit (Chamaea fasciata), GrasshopperSpar- row (Ammodramussavannarum), nine species of blackbirds, grackles, and cowbirds (all North Fall American species except Yellow-headed Black- bird), and the two speciesof meadowlarks.These species are not consideredfurther in this paper. The other 261 species are listed in Table 1, along with data on the replacementof greater coverts, .•HY/SY CSprinKAHY/ASY • ,D tertials/secondaries, and rectrices.

Page 50 NorthAmerican Bird Bander Vol.22 No.2 In ninespecies of passerines(the four speciesof Of these 252 species, data indicated that 183 Myiarchusflycatchers; Sulphur-bellied Flycatcher; species (72.6%) either lack a prealternatemolt or Eastern Kingbird; and Bachman's, Botteri's and have a limited prealtemate molt that does not Cassin'ssparrows), the first prebasicmolt was include greater coverts or flight feathers in any complete or nearly so, and the first and adult birds.Just the extent of the first prebasicmolt is prealternatemolts includedsome flight feathers summarizedfor these 183 species (Table 1). The and/or greatercoverts in at least some birds.In four remaining69 species(27.4%) showedevidence of of these species (the kingbird and the three prealternatemolts that includedone or more greater sparrows), age of the bird subsequent to covertsor flightfeathers in at least some birds.For completion of the prebasic molt could not be these species,the extent of the first prebasicmolt determined,so data on the extent of the prealternate (1st PB), the firstprealternate molt (1 st PA), and the molt(in both age groupspooled) is given in Table 1. adult prealternatemolt (adult PA) are summarized In Sulphur-belliedFlycatcher and the fourspecies (Table 1). of Myiarchus,most or all juvenalprimary coverts were retainedthrough the secondprebasic molt, Finally, the extent of the presupplementalmolts allowingageing of birdscollected in springand (PS) of six speciescould be determined,those in summer. For these, the extents of both the first and which this molt does not overlap in timing or the adultprealternate molts are given(Table 1). location, at the populationslevel, with the first prebasic molt (Thompsonand Leu 1994). In four The remaining 252 species have partial or other species that have presupplementalmolts incompletefirst prebasic molts. The extentof these (NorthernCardinal, Pyrrhuloxia,Yellow-breasted varied from no greater coverts or flight feathers Chat,and LarkSparrow) (Thompson and Leu 1994, replaced(17 species)to mostor allgreater coverts, Pyleunpublished data), the periodin whichgreater a variable number of secondaries and rectrices, covertsor flightfeathers were replacedcould not be andone or moreprimaries replaced in at leastsome determined, and these two molts are combined in birds (51 species;Table 2). Table 1 (as "PS/PB").

Fig. 2. Variationin the extentof wingcovert and tertial replacement during partial molts. HY/SYs of manyNorth American passerineswill show molt limits similar to thosein illustrationsA to E, althoughexceptions to thispattern of replacementare to be expected.Most AHY/ASY passerines show uniform replacement (F), at leastuntil the prealternatemolt, when some AHY/ ASYs undergoa partialmolt resulting in moltlimits as in A-E (see Table 1).

carpalcovert alulacovert

E Apr.-Jun. 1997 NorthAmerican Bird Bander Page51 Molt limits resulting from partial or incomplete, Note that the primarycoverts are retainedin all of firstprebasic molts- The sequenceand extentof these examplesof partialmolt (Figures 2 and 3). wing-covertand tertialreplacement generally follow similar patterns among North American passe- By comparingthe typicalreplacement sequences rines, althoughnumerous exceptions, both within andextents of Figure2 withinformation on variation and amongspecies, can be expected.Molt of the in the extent of the first prebasic molts of each wing coverts typically begins with the proximal speciesin Table1, moltlimits can be lookedfor and lesser coverts, and proceeds distallyand toward usedto age manyHY/SY birdsthrough at leastthe the greatercoverts (Jenni and Winkler1994, Figure prealternatemolt. Individuals of all NorthAmerican 2). Thus,it usuallycommences with the innerlesser passerinesin fall and winter (exceptfor a few and mediancoverts (Figure 2A). Often,when about specieswhich may suspend the adult prebasic molt halfof the lessercoverts have been replaced,molt formigration, such as Red-eyedVireo; see Mulvihill of the median coverts commences (Figure 2A); & Rimmer1997), notin activemolt, that show molt when about half of the median coverts have been limits(Figure 2A-E and Figure 3) are HY/SYs.AHY/ replaced, molt of the greater covertscommences ASYs typicallyshow wing coverts which are (Figure 2B); and when about half of the greater uniformin color,wear, and size (Figure2F), at least coverts have been replaced, molt of the tertials untilthe prealternatemolt. commences (Figure 2D); however, the relative timingof featherreplacement in thesefeather tracts Fig. 3. Many vireos,warblers, and sparrowsshow a slight can vary substantially. variationto the generalpattern of replacementshown in Fig. 2, replacingall wing coverts but no (sometimes1-2) alula feathem or flightfeathem (see Table 1). Replacement of the greater coverts usually proceeds proximally (Figure 2B-E), although irregularsequences and skipped feathers, particu- larly involvingthe innermostcovert (Figure2C) often are encountered (Jenni and Winkler 1994).Thealula covert is oftenreplaced when molt of the mediancoverts has been completed,and the carpal covert and alula feathers often are not replaceduntil molt of the greatercoverts has been completed(Figure 2C-E). Partialwing feather molts can suspendat any pointduring this replacement process,and variationin the pointof suspension, sometimessubstantial, occurs within each species (Table 1). In a few birdsof somespecies, s6 and In somespecies "pseudolimits" occur (See Table 1 occasionallys5 can be replacedafter all three and Notes2 and 3, followingthe tables).These are tertialShave been renewed(Figure 2E). naturalcontrasts in colorpattern between adjacent feathers, that can simulate molt limits. In Zonotrichia In many species,the central rectricescan be sparrows,for instance,the innermosttwo or three replacedif and when the tertialsare replaced.In a greatercoverts and the tertials are a darkeror richer few species,the centralrectrices are replacedbut brownthan adjacent,distal feathers, in both HY/ the tertialsare retained. In some species additional SYs and AHY/ASYs. With these species, care rectricescan be renewedduring incomplete molts. mustbe taken to distinguishbetween pseudolimits These often are replaced from the central pair andtrue molt limits; it is best to carefullyexamine the outwards, although in many individuals the extent of wear to the tips of these feathers to outermost pair may be replaced immediately determineif one or moregeneration of feathersis followingthe central pair. In many species of involved.Jenni and Winkler (1994) providemore passerines,particularly among the vireos, war- information,accompanied by numerousillustra- blers,and sparrows,all lesser, median,and greater tions,on pseudolimitsand the processof ageing coverts but no tertials, rectrices, or other flight passerinesusing molt limits. feathers are replaced (Figure 3). Page 52 North American Bird Bander Vol. 22 No. 2 Molt limits resulting from partial or incomplete, Fig. 4. An exampleof an SY birdwith three generationsof prealternate molts - Most North American feathers,juvenal (white), first basic (lightly stippled), and first alternate(dark) feathers, after partialfirst prebasicand first passerines do not have prealternate molts that prealternatemolts. ASY birdsthat have partialprealternate includegreater covertsor tertials, but in those that moltswill showonly two generationsof feathers,as in Fig.2. do, the feather replacementsequence typicallyis similar to that of prebasic molts, as illustratedin Figure2. In mostspecies, partial prealternate molts occurin bothSYs and ASYs, althoughthe extentof this molt in ASYs usuallyis less than that of SYs (Table 1). In 14 of the 75 species listed in Table 1 with first and adult prealternate molts, no ASY specimens were found with replaced greater covertsor flightfeathers. These includedspecies with extensive first prealternate molts (several Species that replace at least some primaries flycatcherspecies), and species in whichthe first during incomplete molts - Fifty-fourspecies of prealternatemolt includedonly a few innergreater NorthAmerican passerines were foundin whichat coverts at most (several warbler species). One least a proportionof individualsregularly replaced species,the Bobolink,showed complete or nearly some butnot all primariesduring incomplete molts complete prealternate molts in both SYs and ASYs. (Table 2). Several replacement strategies were In some species, the tertials and/or central noted among these species. The majority (46 rectrices could be replaced during prealternate species, or 88.5%) showed "eccentric" replace- molts, that otherwise included few if any wing ment patterns (Figure 5), in which the outer coverts (see Table 1). primaries,inner secondariesand, sometimes,the outermostprimary coverts are replaced(Jenni and Care must be taken when ageing these species in Winkler 1994, Pyle in review). In 38 species, springand summer, as both SYs and ASYs can eccentric patterns were observed during the first show molt limits. Many SYs of certain species prebasic molt only. In two species (Yellow-bellied (those with more extensivefirst prebasicthan first and Willow flycatchers)it occurredonly duringthe prealternate molts) can show three generationsof first prealternate molt; and in four species of feathers in the wing or tail: juvenal feathers, first- kingbirds,replacement of primaries began during basic feathers, and first-alternatefeathers (Figure the first prebasicmolt, suspendedover winter,and 4). These individualscan be aged SY. Otherwise, resumed during the first prealternate molt (along the relativecontrast between retainedand replaced with a second replacementof body feathers; see feathers is the best means of distinguishingthe age Pyle in review). In one species (Nelson's Sharp- groups,this contrastbeing much greater between tailed Sparrow), eccentric replacement patterns juvenal and first-alternatefeathers than between were observed during both the first and the adult adult-basic and adult-alternate feathers (see prealternatemolts but not duringthe first prebasic Mulvihil11993,Jenni and Winkler 1994). Contrasts molt. Interestingly, no replaced primaries were involvingthe juvenal primary coverts, which are found in springor summer Saltmarsh Sharp-tailed retained completely or partially by most HY/SY Sparrows(Table 1), whichhave recentlybeen split North American passerines (see below), often from Nelson's (American Ornithologist'sUnion providethe best means of distinguishingSYs and 1995). Finally, in one species (Lesser Goldfinch; ASYs in the springand summer (Figure 1). see Notes following the tables), eccentric replacementpatterns were noted during the first prebasic molt of all forms, and duringthe first and the adult prealternatemolts of the "black-backed" form but not the "green-backed"form. Other examplesof geographicvariation in moltextent are discussedin the notes followingthe tables. If not specificallynoted, species did not show marked geographicvariation in molt extent. Apr. - Jun. 1997 NorthAmerican Bird Bander Page 53 Fig. 5. Eccentricmolt patterns in NorthAmerican passerines. Most species show a patternsimilar to that of illustrationA, althoughsome flycatchers can show more extensive eccentric replacement, as in illustrationsB andC. A fewspecies can show bothan eccentricand a typicalpattern, as in illustrationD (see Table 2).

Eight species showed primary and secondary Six species that showed eccentric molt patterns replacementin "typical"sequence (Figure 6), the also replacedup to three innerprimaries and three primaries commencingfrom the innermost and outersecondaries, in typical sequence (Figure 5D). proceedingdistally, and the secondaries (after In thesespecies (Table 2), onlysmall proportions of replacement of the tertials) commencingwith the birds(5-16%) showingeccentric replacement also outermost and proceeding proximally. In these had replacedfeathers in typicalsequence. Finally, cases,primary coverts typically were replacedwith one species (Green Jay) showed an irregular their correspondingprimaries, although one or two sequence, replacement of the secondaries coverts often were retained despite the replace- proceedingdistally from the tertials, followed by ment of the adjacent primary (Figure 6B). The replacementof the primaries,proceeding distally typical remex replacementsequence was ob- from the innermost feather. served duringthe first prebasicmolt only. Table 2 summarizes the of replacement Fig. 6. Examplesof replacementin typical pattern and extent of molts in species which sequence (as in completemolts), found during incomplete showed incomplete replacement of the primaries molts in eight speciesof North Americanpasserines (see Table 2). and primarycoverts. As withmolt limits among wing coverts,the limitsamong the flightfeathers of these species are helpfulin distinguishingHY/SYs from HY/ASYs (Uulvihill1993, Jenniand Winkler1994), in mostcases throughthe secondprebasic molt.

A call to banders: more study is needed - The informationpresented in Tables 1 and 2 shouldbe used as a startingpoint toward a more complete understandingof moltlimits and their use in ageing NorthAmerican passerines. Detection of moltlimits on specimensoften is difficult (see Note1 following the tables), in part becausethe wingscannot be examined freely without risking damage to the specimens.For instance,in severalspecies, the originalresults of this study contradictedthat of

Page54 NorthAmerican Bird Bander Vol.22 No.2 other detailed examinations based on either 2. Bancroft, G.T. and G.E. Woolfenden. 1982. The specimensor livebirds (see the notesfollowing the molt of the Scrub Jays and Blue Jays in tables). In a few of these examples,reexamination Florida. Ornith.Monogr. 29:1-51. indicatedthat the initialresults of thisstudy were in 3. Canadian Wildlife Service and U.S. Fish and error. Certainly, other errors exist within Tables 1 Wildlife Service. 1991. North American Bird and 2 whichwill need to be correctedby future Banding.Manual v. 1 and 2. U.S. Fish and workers.In addition,replacement patterns of the Wildl. Serv., , D.C. carpal covert, alula covert,and greater and lesser 4. Cannell, P.F., J.D. Cherry, and K.C. Parkes. alula feathers (see Figure 2), not covered 1983. Variationand migrationoverlap in specificallyby this study, should be examinedmore flightfeather molt of the Rose-breasted fully(Mulvihil11993). Grosbeak. Wilson Bull. 95:621-627. 5. Cherry,J.D. 1985. Earlyautumn movements and Molt limits are much easier to detect on live birds in prebasic molt of Swainson's Thrushes. the handthan they are on specimens.The abilityto Wilson Bull. 97:368-370. open a bird'swing to examinethe feathers,and the 6. Cherry, J.D. and P.F. Cannell. 1984. Rate and factthat the feathers are inbetter relative shape on timingof prebasicmolt of adult Boreal livebirds than on specimens,should allow banders Chickadees. J. Field Omith. 55:487-489. to readilydetect molt limits in mostspecies. (A few 7. Collier, B. and G.E. Wallace. 1989. Aging species, such as House Wren and Common Catharus thrushesby rectrixshape. J. Yellowthroat,will always present difficulties, even Field Omith. 60:230-240. on livebirds in the hand).I stronglyurge banders to 8. Cramp, S., ed. 1988. The birds of the western start lookingfor molt limits when ageing North Palearctic. v. 5. Oxford Univ. Press, American passerines and to publish their Oxford, U.K. information, whether it substantiates or contradicts 9. Cramp, S. and C.M. Perrins, eds. 1994a. The the resultsof this study. birds of the western Palearctic. v. 8. Oxford Univ. Press,Oxford, U.K. ACKNOWLEDGMENTS 10. Cramp, S. and C.M. Perrins, eds. 1994b. The birds of the western Palearctic. v. 9. Oxford Univ. Press, Oxford, U.K. Luis Baptista and Karen Cebra (CAS), Ned K. 11. deGraw, W.A. and M.D. Kern. 1990. Johnson,Carla Cicero,and BarbaraStein (MVZ), Postnuptualmolt in Harris'Sparrows. KimballL. Garrett (LACM), PhilipUnitt (SDNHM), Condor92:829-838. James Northern(MLZ), and Jon Fisherand Walter 12. Dixon, K.L. 1962. Notes on the molt schedule of Wehtje (WFVZ) gave me permissionto examine the Plain Titmouse. Condor64:134-139. specimens under their care. I thank Frank F. Pitelka 13. Emlen, J.T. Jr. 1936. Age determinationin the for pointingout to me the interestingmolt pattern of American Crow. Condor38:99-102. the Green Jay, and Robert S. Mulvihill,Ernest J. 14. Ewert, D.N. and W.E. Lanyon. 1970. The first Willoughby,David Cimprich,and Jon R. Kingfor prebasic molt of the Common Yellowthroat sharing with me their results on molts and molt (Parulidae).Auk87:362-363. limits. Steve N.G. Howell helped me in the 15. Foster, M.S. 1967. Molt cyclesof the Orange- collectionsand producedthe illustrations,and Ann crowned Warbler. Condor 69:169-200. Kiener and Ken Converyhelped me prepare and 16. George,W.G. 1973. Moltof juvenile White-eyed proof the tables. Critical reviews by Robert S. Vi•eos. Wilson Bull. 85:327-330. Mulvihilland ChristopherW. Thompsonimproved 17. Hubbard,J.P. 1980. The extent and sequence the manuscript.This is contribution741 of PRBO. of the moltsof the Yellow-rumpedWarbler. Nemouria 25:1-9. LITERATURE CITED AND REFERENCES 18. Humphrey, P.S. and K.C. Parkes. 1959. An approachto the studyof moltsand 1. AmericanOrnithologist's Union. 1995. 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Zool. 38:425-444. flycatchers.Pp. 147-154 in R.W. Dickerman 35. Morton,M.L., J.R. King,and D.S. Farner.1969. (comp.).The Era of Allan R. Phillips:A Postnuptualand postjuvenalmolt in White- Festschrift.R.W. Dickerman,AIbequerque, crowned Sparrows in central Alaska. NM. Condor71:376-385. Page 56 North Amedcan Bird Bander Vol. 22 No. 2 53. Pyle, P. In review.Eccentric first-year molt 68. Thompson,C.W. 1991. The sequenceof molts patternsin certaintyrannid flycatchers, and and plumagesin PaintedBuntings and relatedrefinements to moltterminology. W. implicationsfor theories of delayedplumage Birds. maturation. Condor93:209-235. 54. Pyle, P., S.N.G. Howell, R.P. Yunick,and D.F. 69.Thompson,C.W. and M. Leu. 1994. Determining DeSante.1987. Identificationguide to North homologyof moltsand plumagesto American passerines. Slate Creek Press, addressevolutionary questions: A rejoinder Bolinas, CA. regardingEmberizid . Condor96:769- 55. Pyle, P. and D.A. Sibley.1992. Juvenal- 782. plumagedLe Conte'sSparrows on migra- 70. Tordoff, H.B. and R.M. Mengel. 1951. The tion.Are they beingoverlooked? Birding occurrenceand possiblesignificance of a 24:70-76. spring molt in Le Conte's Sparrow. Auk 56. Pyle,P. andP. Unitt.In press.Molt and plumage 68:519-522. variationby age and sex in the California 71 .Traylor,M.A., Jr. 1968. Winter moltin the and Black-tailedgnatcatchers. Studies in Acadian Flycatcher, œmpidonaxvirens. Avian BioL Auk85:691. 57. Reese, J.G. 1975. Fall remix and rectrix molt in 72. Walters, P.M. and D.W. Lamm. 1980. A Hooded the Cardinal.Bird-Banding46:305-310. Warbler (Wi/soniacitrina) in south-east 58. Rohwer,S.A. 1986. A previouslyunknown Arizona. N. Am. Bird Bander5:15. plumageof first-yearIndigo Buntings and 73. Welter,W.A. 1936. Featherarrangement, theoriesof delayedplumage maturation. development,and moltof the Long-billed Auk 103:281-292. Marsh Wren. Wilson Bull. 48:256-269. 59. Rohwer, S.A., W.P. Klein Jr. and S. Heard. 74. Willoughby,E.J. 1986. An unusualsequence of 1983. Delayedplumage maturation and the moltsand plumagesin Cassin'sand presumed prealternate molt in American Bachman's sparrows. Condor88:461-472. Redstarts. Wilson Bull. 95:199-208. 75. Willoughby,E.J. 1989. The moltsof Chipping 60. Rohwer,S.A. and J. Manning.1990. Differ- Sparrowsand Field Sparrowsin Maryland. ences in timingand numberof moltsfor Maryland Birdlife45:127-134. Baltimoreand Bullock'sorioles: Implica- 76. Willoughby,E.J. 1991. Moltof the genus tions to hybrid fitness and theories of Spizella(Passeriformes, Emberizidae)in delayed plumagematuration. Condor relationto ecologicalfactors affecting 95:125-140. plumage wear. Proc. Western Found. Vert. 61. Scott, D.M. 1967. Postjuvenal molt and Zool. 4:247-286. determinationof age of the Cardinal.Bird- 77. Wiseman, A.J. 1977. Interrelationof variables in Banding38:37-51. postjuvenalmolt of Cardinals.Bird-Banding 62. Sealy, S.G. 1969. Prebasicmolt of the Northern 48:206-223. Oriole. Can. J. ZooL 57:1473-1478. 78. Wolf,L.L. 1977. Speciesrelationships in the 63. Selander,R.K. 1964. Speciationin wrensof the avian genusAimophila. Ornith. Monogr. genus Campylorhinchus.Univ. California 23:1-220. Pubs. ZooL 74:1-305. 79. Woolfenden,G.E. 1955. Springmolt of the 64. Stangel,P.W. 1985. Incompletefirst prebasic Harris Sparrow. WilsonBull. 67:212-213. molt of Massachusetts House Finches. J. 80. Young,B.E. 1991. Annualmolt and interruption Field Ornith. 56:1-8. of the fall migrationfor moltingin Lazuli 65. Sutton,G.M. 1935.The juvenalplumage and Buntings. Condor 93:236-250. postjuvenalmolt in several speciesof 81. Yunick,R.P. 1976. Incompleteprebasic molt in Michigan sparrows. CranbrookInst. Sci. a Dark-eyedJunco. Bird-Banding47:276- 3:1-36. 277. 66. Taylor,W.K. 1970. Moltsof the Verdin, 82.Yunick,R.P. 1992. A methodfor age deter- Auriparusflaviceps. Condor 72:493-496. minationof BlueJays in northeastern 67. Thompson,C.F. 1973. Postjuvenalmolt in the and southeastern Canada. White-eyedVireo. Bird-Banding 44:63-65. N. Am. Bird Bander 17:10-15.

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