Molt
Limits
North
in
American
Passerines
Peter Pyle
Point Reyes Bird Observatory
4990 Shoreline Highway
Stinson Beach, CA 94970
ABSTRACT
"Molt limits," or the boundaries between replaced and
retained wing feathers and rectrices during partial or
incompletemolts,were investigatedin 288 speciesof North
American passerines through the examination of over
16,000 specimens. Thirty-sixspecies showed evidence of
completefirst prebasic molts, including27 species that did
not have extensive prealternate molts (those includingat
least some greater coverts) and nine species that did have
extensive prealternate molts. Of the remaining252 species,
183 did not have extensive prealternate molts and 69
speciesdid. Fifty-fourspeciesshowedevidenceof partialor
incomplete replacement of primaries, during the first
prebasic molt, the prealternate molt, or both, in at least a
proportionof the populations.The replacementpattern of
primarieswas either"eccentric"
(proceedingdistallyfromthe
center of the primaries;46 species), or "typical"(proceeding
distallyfromthe innermostprimary;8 species).In sixspecies
that showed an eccentric replacement pattern, a small
proportionof individuals also showed the typical replacement pattern. Data on variation in the extent of greatercovert, secondary, rectrix, primary, and primary-covert
replacement for each partial or incomplete molt in each
species, referencesto previousdetailedstudies on molt, and
notes on geographicvariation, discrepanciesbetween the
results of this study and that of previous work, and other
interestingcases, are presented.
INTRODUCTION
In most North American passerines, the first
prebasicmoltis "partial"or "incomplete,"
some but
not all feathers being replaced(Pyle et al. 1987,
Mulvihill1993). Recently,JenniandWinkler(1994)
have illustrated the utility of "molt limits," the
boundaries between replaced and retained
feathersthat resultfrom partial molts,in ageing
such passerines.Retainedjuvenal wing coverts
and flight feathers are relativelyworn and often
showmore subduedcolorpatternsthan adjacent,
supplementalor first-basicfeathers. Becauseadult
(definitive)prebasic molts in virtuallyall North
Apr.- Jun, 1997
American passerines are complete (Pyle et al.
1987), the presenceof moltlimitsindicatesHY/SY
(first-year)birds,at leastuntilthe prealternatemolt,
and often untilthe secondprebasicmolt (Mulvihill
1993, Jenni and Winkler 1994). Thus, molt limits
canbe especiallyusefulforageingNorthAmerican
passerinesin winter and spring, after first-year
birds have typicallycompletedskull pneumatization. Patterns of replacement among the wing
feathers vary substantiallyboth among species
and among individualsof the same species. This
variationis very poorlydocumentedfor most North
American species (see Mulvihill 1993). Additionally, some species undergo partial or incomplete
prealternate molts in both HY/SY and AHY/ASY
(adult)birds(Pyleet al. 1987, Mulvihill1993).To use
molt limits effectively,therefore, variation in the
extentof replacementduringthe firstprebasicmolt,
and the occurrenceand extentof prealternatemolts
(especially in AHY/ASYs), must be known. To
assess
variation
in the
location
of molt limits
resulting from partial and incomplete,
presupplemental,first prebasic, first prealternate,
and adult prealternate molts of North American
passerines, I examined over 16,000 specimens of
288 species. The results of this examinationare
presented here.
METHODS
Specimensexaminedforthisstudywere locatedat
the California Academy of Sciences (CAS),
MuseumofVertebrateZoology(MVZ), PointReyes
Bird Observatory (PRBO), Natural History
Museum of Los Angeles County (LACM), San
Diego Natural HistoryMuseum (SDNHM), Moore
Laboratory of Zoology (MLZ), and Western
Foundationof Vertebrate Zoology (WFVZ). On
NorthAmericanBirdBander
Page 49
each specimenthe wingcovertsandflightfeathers
were studiedcarefullyfor evidenceof partial or
incompletemolts.The numberof replacedwing
coverts and flight feathers were recorded on all
birds showingevidence of incompletefeather
replacement,afteractivemoltinghadceased. Both
wingswere examinedon each specimento ensure
that resultswere basedon incompletemoltsrather
than adventitiousreplacement;specificdata were
taken fromthe rightwing.
The age of each bird when it was collected was
An attempt was made to sample at least 15-20
specimensfor each moltwithina species,from as
wide a geographicrange (withinNorthAmerica) as
specimenmaterialallowed.Althoughallcollections
werelocatedinCalifornia,a significant
proportion
of
specimens were collected from other localities
throughout North America. Larger samples of
specimens were examined for species showing
wide geographicvariation or complex incomplete
molts,and smallersamplesindicatefewer available
specimens. Ranges in the number of feathers
replaced,withineach tract, are presentedas mean
_+twicethe standarddeviation.These rangesestimatewhatwouldbe expectedfor 95% of the population, assuming a normal distributionto replacement patterns(Pyle 1997).
determinedby information
onthe specimenlabels,
the presenceof molt limits(assumingthat adult
prebasicmoltsare complete),and the color and
amountof wear to the primarycoverts(Figure1),
whichtypicallyare retained,at leastinpart,bybirds
undergoingincompletemolts (see below). Other
When the results of this examination contradicted
plumagecriteria(Pyleet al. 1987) were usedwhere
thoseof otherpublishedor unpublishedinformation
appropriate.Ageterminologyfollowsthatofthe Bird
(see the Tables for other references discussing
BandingLaboratory(CanadianWildlifeServiceand
molt in North American passerines),specimens
U.S. FishandWildlifeService1991).Terminology were reexamined to either confirm or correct the
of molt,plumages,andfeathergenerations
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
determine
whether
observed
RESULTS
AND
DISCUSSION
molt limits resulted
from the presupplementalmolt "PS" (Thompson
and Leu 1994), the prebasic molt "PB", or the
prealternatemolt"PA".
Fig 1. Shapeand relativeconditionof the primarycovertsin
HY/SY and AHY/ASY passerines,in fresh (fall) and worn
(spring)condition.The contrastbetweenthesefeathereand
replaced greater coverts is very useful in ageing many
species. Note that the edging on these feathere is often
presentbut thinnerin HY birdsthan in AHY birds in the fall,
and it is oftenabsentin SY birdsbutstillpresentin ASY birds
in the spring.
Molt patterns in the 288 North Americanspecies
examined could be categorized into several
groups. In 27 species (9.4%), data indicatedthat
replacement of wing coverts and flight feathers
duringboththe firstand adultprebasicmoltswas
typicallycomplete,and that the prealternatemolts
were either absent or limited,includingno greater
coverts or flight feathers. These species were:
Northern Beardless-Tyrannulet (Camptostoma
imberbe),the two speciesof wood-pewees,Alder
Flycatcher (Empidonax alnorum), Horned Lark
(Eremophilaalpestris),the eightspeciesof martins
and swallows, Bushtit (Psaltriparus minimus),
Wrentit (Chamaea fasciata),GrasshopperSparrow (Ammodramussavannarum),nine speciesof
blackbirds, grackles, and cowbirds (all North
American species except Yellow-headed Blackbird), and the two speciesof meadowlarks.These
speciesare not consideredfurther in this paper.
The other 261 species are listed in Table 1, along
with data on the replacementof greater coverts,
Fall
HY/SY
C
AHY/ASY
.•
SprinK
• ,D
tertials/secondaries, and rectrices.
Page 50
NorthAmericanBirdBander
Vol.22 No.2
In ninespeciesof passerines(the fourspeciesof
Myiarchusflycatchers;
Sulphur-bellied
Flycatcher;
Eastern Kingbird;and Bachman's, Botteri's and
Cassin'ssparrows),the first prebasicmolt was
complete or nearly so, and the first and adult
prealternatemolts includedsome flight feathers
and/or greatercovertsinat leastsome birds.In four
of these species (the kingbird and the three
sparrows), age of the bird subsequent to
completion of the prebasic molt could not be
determined,
sodataontheextentoftheprealternate
molt(inbothage groupspooled)isgiveninTable1.
In Sulphur-bellied
Flycatcher
andthefourspecies
of Myiarchus,mostor all juvenalprimarycoverts
were retainedthroughthe secondprebasicmolt,
allowingageingof birdscollectedin springand
summer. For these, the extents of both the first and
theadultprealternatemoltsare given(Table1).
The remaining 252 species have partial or
incompletefirstprebasicmolts.The extentof these
varied from no greater covertsor flightfeathers
replaced(17 species)to mostor allgreatercoverts,
a variable number of secondaries and rectrices,
andoneor moreprimariesreplacedinat leastsome
birds(51 species;Table 2).
Of these 252 species, data indicated that 183
species(72.6%) either lack a prealternatemoltor
have a limited prealtemate molt that does not
include greater coverts or flight feathers in any
birds.Just the extent of the first prebasicmolt is
summarizedfor these 183 species(Table 1). The
remaining69 species(27.4%) showedevidenceof
prealternatemoltsthat includedone or moregreater
covertsor flightfeathersin at leastsome birds.For
these species,the extent of the first prebasicmolt
(1st PB),the firstprealternatemolt(1st PA),andthe
adult prealternatemolt (adult PA) are summarized
(Table 1).
Finally, the extent of the presupplementalmolts
(PS) of six speciescould be determined,those in
which this molt does not overlap in timing or
location, at the populationslevel, with the first
prebasicmolt (Thompsonand Leu 1994). In four
other species that have presupplementalmolts
(NorthernCardinal, Pyrrhuloxia,Yellow-breasted
Chat,and LarkSparrow)(Thompsonand Leu 1994,
Pyleunpublished
data),the periodinwhichgreater
covertsorflightfeatherswere replacedcouldnotbe
determined, and these two molts are combined in
Table 1 (as "PS/PB").
Fig. 2. Variationin the extentof wingcovertandtertialreplacement
duringpartialmolts.HY/SYsof manyNorthAmerican
passerines
willshowmoltlimitssimilarto thosein illustrations
A to E, althoughexceptions
to thispatternof replacement
are to
be expected.MostAHY/ASYpasserinesshowuniformreplacement(F), at leastuntilthe prealternatemolt,whensomeAHY/
ASYs undergoa partialmoltresultingin moltlimitsas in A-E (see Table 1).
carpalcovert alulacovert
E
Apr.-Jun.1997
NorthAmerican
BirdBander
Page51
Molt limits resulting from partial or incomplete,
firstprebasic molts- The sequenceand extentof
wing-covertandtertialreplacementgenerallyfollow
similar patterns among North American passerines, althoughnumerousexceptions,both within
and amongspecies,can be expected.Molt of the
wing coverts typically begins with the proximal
lesser coverts, and proceedsdistallyand toward
the greatercoverts(JenniandWinkler1994, Figure
2). Thus,it usuallycommenceswiththe innerlesser
and mediancoverts(Figure2A). Often,whenabout
halfof the lessercovertshave been replaced,molt
of the median coverts commences (Figure 2A);
when about half of the median coverts have been
replaced,molt of the greater covertscommences
(Figure 2B); and when about half of the greater
coverts have been replaced, molt of the tertials
commences (Figure 2D); however, the relative
timingof featherreplacementinthesefeathertracts
can vary substantially.
Replacement of the greater coverts usually
proceeds proximally (Figure 2B-E), although
irregularsequencesandskippedfeathers,particularly involvingthe innermostcovert (Figure2C)
often are encountered (Jenni and Winkler
1994).Thealulacovertisoftenreplacedwhenmolt
of the mediancovertshas been completed,and the
carpal covert and alula feathers often are not
replaceduntilmoltof the greatercovertshasbeen
completed(Figure2C-E). Partialwingfeathermolts
can suspendat any pointduringthisreplacement
process,andvariationin the pointof suspension,
sometimessubstantial,occurswithineach species
(Table 1). In a few birdsof somespecies,s6 and
occasionallys5 can be replacedafter all three
tertialShave been renewed(Figure2E).
Note that the primarycovertsare retainedin all of
these examplesof partialmolt(Figures2 and 3).
By comparingthe typicalreplacementsequences
andextentsof Figure2 withinformation
onvariation
in the extent of the first prebasicmolts of each
speciesinTable1, moltlimitscanbe lookedforand
usedto age manyHY/SY birdsthroughat leastthe
prealternate
molt.Individuals
ofallNorthAmerican
passerinesin fall and winter (exceptfor a few
specieswhichmaysuspendtheadultprebasicmolt
formigration,
suchas Red-eyedVireo;see Mulvihill
& Rimmer1997), notin activemolt,thatshowmolt
limits(Figure2A-EandFigure3) areHY/SYs.AHY/
ASYs typicallyshow wing covertswhich are
uniformincolor,wear,and size (Figure2F), at least
untilthe prealternatemolt.
Fig. 3. Many vireos,warblers,and sparrowsshowa slight
variationto the generalpatternof replacementshownin Fig.
2, replacingall wing coverts but no (sometimes1-2) alula
feathem or flightfeathem (see Table 1).
In somespecies"pseudolimits"
occur(See Table 1
and Notes2 and 3, followingthe tables).These are
naturalcontrastsincolorpatternbetweenadjacent
feathers, that can simulate molt limits. In Zonotrichia
In many species,the central rectricescan be
replacedif and whenthe tertialsare replaced.In a
few species,the centralrectricesare replacedbut
the tertialsare retained. In some species additional
rectricescan be renewedduringincompletemolts.
These often are replaced from the central pair
outwards, although in many individualsthe
outermost pair may be replaced immediately
followingthe central pair. In many species of
passerines,particularlyamong the vireos, warblers,and sparrows,all lesser,median,and greater
coverts but no tertials, rectrices, or other flight
feathers are replaced (Figure 3).
Page 52
sparrows,for instance,the innermost
twoor three
greatercovertsandthetertialsarea darkerorricher
brownthan adjacent,distalfeathers,in both HY/
SYs and AHY/ASYs. With these species, care
mustbe taken to distinguishbetweenpseudolimits
andtruemoltlimits;itisbesttocarefullyexaminethe
extent of wear to the tips of these feathers to
determineif one or moregenerationof feathersis
involved.Jenni and Winkler (1994) providemore
information,accompaniedby numerousillustrations,on pseudolimits
and the processof ageing
passerinesusingmoltlimits.
North American Bird Bander
Vol. 22 No. 2
Molt limits resulting from partial or incomplete,
prealternate molts
- Most North American
passerines do not have prealternate molts that
includegreater covertsor tertials, but in those that
do, the feather replacementsequence typicallyis
similar to that of prebasic molts, as illustratedin
Figure2. In mostspecies,partialprealternatemolts
Fig. 4. An exampleof an SY birdwiththree generationsof
feathers,juvenal(white),firstbasic(lightlystippled),andfirst
alternate(dark)feathers,after partialfirstprebasicand first
prealternatemolts.ASY birdsthat have partialprealternate
moltswillshowonlytwo generationsof feathers,as in Fig.2.
occurinbothSYs 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
flycatcherspecies),and species in whichthe first
prealternatemoltincludedonlya few innergreater
coverts at most (several warbler species). One
species,the Bobolink,showedcompleteor nearly
complete prealternatemoltsin bothSYs and ASYs.
In some species, the tertials and/or central
rectrices could be replaced during prealternate
molts, that otherwise included few if any wing
coverts (see Table 1).
Care must be taken when ageingthese speciesin
springand summer,as both SYs and ASYs can
show molt limits. Many SYs of certain species
(those with more extensivefirst prebasicthan first
prealternatemolts)can show three generationsof
feathers in the wing or tail: juvenal feathers, firstbasic feathers, and first-alternatefeathers (Figure
4). These individualscan be aged SY. Otherwise,
the relativecontrastbetweenretainedand replaced
feathers is the best means of distinguishingthe age
groups,this contrastbeing muchgreater between
juvenal and first-alternatefeathers than between
adult-basic and adult-alternate feathers (see
Mulvihil11993,Jenniand Winkler 1994). Contrasts
involvingthe juvenal primary coverts, which are
retained completely or partially by most HY/SY
North American passerines (see below), often
providethe best means of distinguishing
SYs and
ASYs in the springand summer (Figure 1).
Species that replace at least some primaries
during incomplete molts - Fifty-fourspecies of
NorthAmericanpasserineswerefoundin whichat
least a proportionof individualsregularlyreplaced
somebutnotall primariesduringincompletemolts
(Table 2). Several replacement strategies were
noted among these species. The majority (46
species, or 88.5%) showed "eccentric" replacement patterns (Figure 5), in which the outer
primaries,innersecondariesand, sometimes,the
outermostprimarycovertsare replaced(Jenniand
Winkler 1994, Pyle in review). In 38 species,
eccentric patterns were observed during the first
prebasic molt only. In two species (Yellow-bellied
and Willow flycatchers)it occurredonly duringthe
first prealternate molt; and in four species of
kingbirds,replacementof primaries began during
the firstprebasicmolt,suspendedover winter,and
resumed during the first prealternate molt (along
with a second replacementof body feathers;see
Pyle in review). In one species (Nelson's Sharptailed Sparrow), eccentric replacement patterns
were observed during both the first and the adult
prealternatemoltsbut not duringthe firstprebasic
molt. Interestingly, no replaced primaries were
found in springor summer SaltmarshSharp-tailed
Sparrows(Table 1), whichhave recentlybeen split
from Nelson's (American Ornithologist'sUnion
1995). Finally, in one species (Lesser Goldfinch;
see Notes following the tables), eccentric
replacementpatternswere noted duringthe first
prebasicmolt of all forms, and duringthe first and
the adult prealternatemoltsof the "black-backed"
form but not the "green-backed"form. Other
examplesof geographicvariationin moltextentare
discussedin the notes followingthe tables. If not
specificallynoted, species did not show marked
geographicvariationin molt extent.
Apr.- Jun. 1997
NorthAmericanBirdBander
Page 53
Fig. 5. Eccentric
moltpatternsin NorthAmericanpasserines.
Mostspeciesshowa patternsimilarto thatof illustration
A,
although
someflycatchers
canshowmoreextensiveeccentricreplacement,
as in illustrations
B andC. A fewspeciescanshow
bothan eccentricand a typicalpattern,as in illustrationD (see Table 2).
Eight species showed primary and secondary
replacementin "typical"sequence (Figure 6), the
primaries commencingfrom the innermost and
proceedingdistally, and the secondaries (after
replacementof the tertials) commencingwith the
outermost and proceeding proximally. In these
cases,primarycovertstypicallywere replacedwith
their correspondingprimaries,althoughone or two
coverts often were retained despite the replacement of the adjacent primary (Figure 6B). The
typical remex replacementsequence was observed duringthe firstprebasicmoltonly.
Fig. 6. Examplesof flightfeather replacementin typical
sequence(as in completemolts),found during incomplete
moltsin eight speciesof North Americanpasserines(see
Table 2).
Six species that showed eccentric molt patterns
alsoreplacedupto three innerprimariesand three
outersecondaries,intypicalsequence(Figure5D).
Inthesespecies(Table2), onlysmallproportions
of
birds(5-16%) showingeccentricreplacementalso
had replacedfeathersin typicalsequence.Finally,
one species (Green Jay) showed an irregular
sequence, replacement of the secondaries
proceedingdistallyfrom the tertials, followed by
replacementof the primaries,proceedingdistally
from the innermost feather.
Table 2 summarizes the type of replacement
pattern and extent of molts in species which
showed incomplete replacement of the primaries
andprimarycoverts.Aswithmoltlimitsamongwing
coverts,the limitsamongthe flightfeathersof these
speciesare helpfulin distinguishingHY/SYs from
HY/ASYs (Uulvihill1993, Jenniand Winkler1994),
in mostcasesthroughthe secondprebasicmolt.
A call to banders: more study is needed - The
informationpresentedin Tables 1 and 2 shouldbe
used as a startingpointtoward a more complete
understandingof moltlimitsand theiruse in ageing
NorthAmericanpasserines.Detectionof moltlimits
onspecimensoftenisdifficult(see Note1 following
the tables), in part becausethe wingscannotbe
examined freely without riskingdamage to the
specimens.For instance,in severalspecies,the
originalresultsof this study contradictedthat of
Page54
NorthAmericanBirdBander
Vol.22 No.2
other
detailed
examinations
based
on
either
specimensor livebirds(see the notesfollowingthe
tables). In a few of these examples,reexamination
indicatedthatthe initialresultsof thisstudywere in
error. Certainly, other errors exist within Tables 1
and 2 whichwill need to be correctedby future
workers.In addition,replacementpatternsof the
carpal covert,alula covert,and greater and lesser
alula feathers (see Figure 2), not covered
specifically
bythisstudy,shouldbe examinedmore
fully(Mulvihil11993).
Molt limits are much easier to detect on live birds in
the handthantheyareonspecimens.
Theabilityto
opena bird'swingto examinethe feathers,and the
factthatthefeathersareinbetterrelativeshapeon
livebirdsthanon specimens,shouldallowbanders
to readilydetectmoltlimitsin mostspecies.(A few
species, such as House Wren and Common
Yellowthroat,willalwayspresentdifficulties,
even
on livebirdsinthe hand).I stronglyurgebandersto
start lookingfor molt limits when ageing North
American passerines and to
publish their
information, whether it substantiates or contradicts
the resultsof this study.
2. Bancroft, G.T. and G.E. Woolfenden. 1982. The
molt of the Scrub Jays and Blue Jays in
Florida.Ornith.Monogr.29:1-51.
3. Canadian
Wildlife Service and U.S. Fish and
Wildlife Service. 1991. North American Bird
Banding.Manual v. 1 and 2. U.S. Fish and
Wildl. Serv., Washington, D.C.
4. Cannell, P.F., J.D. Cherry, and K.C. Parkes.
1983. Variationand migrationoverlap in
flightfeather molt of the Rose-breasted
Grosbeak.
Wilson Bull. 95:621-627.
5. Cherry,J.D. 1985. Earlyautumnmovementsand
prebasic molt of Swainson's Thrushes.
Wilson Bull. 97:368-370.
6. Cherry, J.D. and P.F. Cannell. 1984. Rate and
timingof prebasicmoltof adult Boreal
Chickadees.
J. Field Omith. 55:487-489.
7. Collier, B. and G.E. Wallace. 1989. Aging
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10. Cramp, S. and C.M. Perrins,eds. 1994b. The
ACKNOWLEDGMENTS
birds of the western Palearctic. v. 9. Oxford
Luis Baptista and Karen Cebra (CAS), Ned K.
Johnson,Carla Cicero,and BarbaraStein (MVZ),
KimballL. Garrett(LACM), PhilipUnitt(SDNHM),
James Northern(MLZ), and Jon Fisherand Walter
Wehtje (WFVZ) gave me permissionto examine
specimens under their care. I thank Frank F. Pitelka
for pointingout to me the interestingmoltpatternof
the Green Jay, and Robert S. Mulvihill,Ernest J.
Willoughby,David Cimprich,and Jon R. Kingfor
sharing with me their results on molts and molt
limits. Steve N.G. Howell helped me in the
collectionsand producedthe illustrations,and Ann
Kienerand Ken Converyhelped me prepareand
proof the tables. Critical reviews by Robert S.
Mulvihilland ChristopherW. Thompsonimproved
the manuscript.This is contribution741 of PRBO.
Univ. Press, Oxford, U.K.
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