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Eccentric First-Year Molt Patterns in Certain Tyrannid Flycatchers

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Eccentric First-Year Molt Patterns in Certain Tyrannid Flycatchers ECCENTRIC FIRST-YEAR MOLT PATrERNS IN CERTAIN TYRANNID FLYCATCHE!.S PETER PYLE, Point ReyesBird Observatory,4990 ShorelineHwy., StinsonBeach, California 94970 Most passerinesfollow a similarsequence of remex molt, replacementof the primariescommencing with the innermostand proceedingdistally, while that of the secondaries(except for the tertials, S7-S9) beginswith the outermostand proceeds proximally (Ginn and Melville 1983). Duringthe first- year molts(here defined as all periodsof moltingbefore the secondprebasic molt),many passerines replace no remigeswhereas others replace some or all remiges(Pyle et al. 1987, Jenni and Winkler 1994). "Incomplete"first-year remexmolts occasionally result when the typicalmolting sequence is arrested, the distaljuvenal primaries and proximalsecondaries (excluding the tertials) being retained.This pattern of incompletemolt is found in severalNorth Americanpasserines, notably, the Phainopepla(Phainopepla nitens), North- ern Cardinal (Cardinalis cardinalis), and Lark Sparrow (Chondestes grammacus)(Pyle et al. 1987, Thompsonand Leu 1994). In a few passerinespecies, incomplete first-year primary molts (and in some casessecondary molts) that do not follow the typical sequencebut involvereplacement of centralor distalprimaries in varyingpatterns have been documented(Jenni and Winkler 1994, Thompson and Leu 1994). These replacementpatterns, termed "eccentric"by Jenni and Winkler (1994), are poorly understood.A common eccentricreplacement pattern involvesrenewal of the outer four to six primariesand inner three to five secondaries,feather replacementproceeding distally in the primariesand proximallyin the secondaries,as in the typicalsequence, but commencingat differentpoints along the wing (Jenni and Winkler 1994; Figure 1). This pattern has been documentedin severalNorth American passerines,nota- bly, the LoggerheadShrike (Lanius ludovicianus),Yellow-breasted Chat (Icteria virens),and Passerina buntings (Miller 1928, Phillips1974, Thomp- son and Leu 1994). While examiningspecimens of North Americantyrannid flycatchers for molt-relatedage criteria, I found that first-yearbirds of severalspecies of differentgenera display eccentric remex-replacement patterns. Depending on the species,replacement of remigesoccurs during fall molting periods, spring molting periods, or both. Here I summarizethese patterns, and suggestseveral hypotheses for their occurrence. MATERIALS AND METHODS This studywas limited to birdscollected during their firstyear. Results were basedon datafrom 589 specimensof 22 north-temperatespecies examined at the CaliforniaAcademy of Sciences(CAS), Moore Laboratoryof Zoology (MLZ), Museumof VertebrateZoology (MVZ), and WesternFoundation of VertebrateZoology (WFVZ). On each specimenI carefullyexamined the primaries,secondaries, and primary covertsfor feather-retentionpatterns WesternBirds 29:29-35, 1998 29 ECCENTRIC FIRST-YEAR MOLT PATTERNS IN CERTAIN TYRANNID FLYCATCHERS •..... ""'• 4 7 7 6 5 432 12 primaries secondaries Figure1. Variationin the eccentricfirst prebasicmolt of the VermilionFlycatcher, fromnine remiges and no primarycoverts replaced (A) to allremiges and four primary covertsreplaced (C). Shadingdenotes replaced feathers. Note the differencein shape and wear betweenretained juvenal primary coverts(inner six feathersin C) and replacedadult primary coverts(outer four feathersin C). In many other tyrannid flycatchers,adults have primary covertsuniformly broad and fresh and are thus separatedfrom first-yearbirds, with entirelyor mostlynarrow, pointed, and worn coverts.See alsoPyle (1997). reflectingthe extent of previousmolts. Birdsthat had been undergoing activemolt when collected(as evidenced by growingor sheathedremiges) were excludedfrom analysesof extent,as extentcould not be determined from thesespecimens; however, I usedspecimens in molt for informationon the timingand sequence of featherreplacement. I examinedboth wings to assessmolt patterns,while all specificdata were recordedfrom the right wings. During this studyI discovered(as confirmed by replacementpatterns on specimenscollected during active primary molt) that most North American 3O ECCENTRIC FIRST-YEAR MOLT PA•FrERNS IN CERTAIN TYRANNID FLYCATCHERS tyrannidflycatchers retain mostor all of their juvenalprimary coverts until their secondprebasi½ molt and that juvenaland adult (definitive)primary covertscould be identifiedby differencesin shape,color pattern, and relative wear (Figure 1C; see also Pyle 1997). These differences,along with differencesin outerprimary shape, rectrix shape, feather wear, color of the wing bars, and color of the plumage (Pyle et al. 1987), allowedreliable separationof most first-yearfrom older birds through the second or definitiveprebasic molt. Even first-yearbirds with completelyreplaced remigesretain five or more inner primary coverts,allowing their accurate separationfrom adults. First-yearflycatchers show a varietyof moltingstrategies (e.g., Johnson 1963, 1974; Pyleet al. 1987). Remigescan be replacedduring the fall(July to November),the spring(March to May), or both (Dickeyand van Rossem 1938, Johnson1963, this study);thus, fall and winterbirds can showup to two generationsof remigesand springand summerbirds can showup to three generationsof remiges.On first-yearspecimens, I recordedeach remex and primary covertas either juvenal,"first fall" (feathersreplaced duringthe fall),or "firstspring" (feathers replaced during the spring).After the springmolting period, I distinguishedjuvenal, first-fall, and first-spring feathersby their relativewear, first-fallfeathers being fresherthan juvenal feathersbut more worn than first-springfeathers. RESULTS Among first-yearflycatchers, eccentric replacement patterns were found in eight species(Table 1): the Olive-sidedFlycatcher (Contopus cooperi), Yellow-belliedFlycatcher (Empidonax fiaviventris), western Willow Fly- catcher(E. traillii brewsteri,adastus, and extirnus),Vermilion Flycatcher '!'aisle! Numbers(Mean and Range) of ConsecutiveOuter Primaries, Inner Secondaries,and Outer PrimaryCoverts Replaced on the RightWing During EccentricFirst-Year Molts in TyrannidFlycatchers Inner Outer Species n Primariesa Secondariesb PrimaryCoverts Olive-sidedFlycatcher 28 7.8 (4-10) 6.9 (3-9) 0.9 (0-4) Yellow-belliedFlycatcher 12 7.3 (6-10) 7.3 (6-9) 0.0 (--) WesternWillow Flycatcher 37 8.1 (5-10) 7.2 (3-9) 0.0 (--) VermilionFlycatcher 82 9.0 (5-10) 8.2 (3-9) 1.3 (0-5) WesternTropical Kingbird 18 5.3 (5-6) 3.9 (3-6) 0.0 Cassin'sKingbird 14 1.7 (0-5) 3.7 (3-5) 0.0 (--) WesternKingbird 55 6.0 (5-7) 3.8 (3-6) 0.0 (--) Scissor-tailedFlycatcher 16 6.5 (4-7) 3.5 (3-5) 0.0 (--) aStartingpoints (primary number) for primarymolts can be calculatedas ten minusthe numberof feathersreplaced. bNotethat one or moretertials can be replacedtwice during the firstyear in kingbirds(see Table 2), but the totalsfor these speciesinclude each tertial once. 31 ECCENTRIC FIRST-YEAR MOLT PAl-fERNS IN CERTAIN TYRANNID FLYCATCHERS (Pyrocephalusrubinus), western Tropical Kingbird (Tyrannus melancholi- cus occidentalis;other subspecieswere not examinedfor this study), Cassin's Kingbird (T. vociferans), Western Kingbird (T. verticalis), and Scissor-tailedFlycatcher (T. forficatus). Limitedspecimen evidence (n = 4) suggeststhat an eccentricreplacement pattern also occurs in first-yearGray Kingbirds(T. dorninicensis).Among other taxa of thesegenera, the West- ern (Contopussordidulus; n = 26) and Eastern(C. virens; n = 12) wood- pewees,eastern Willow (E. t. carnpestris/traillii;n = 6) and Alder (E. alno- rum; n = 22) flycatchers,and Eastern Kingbird (T. tyrannus; n = 12) replacedall remiges,whereas the GreaterPewee (C. pertinax; n = 15), the remainingErnpidonax flycatchers(virescens, n = 9; rninirnus, n = 18; harnrnondii, n = 28; oberholseri, n = 27; wrightii, n = 24; difficilis, n = 30; occidentalis, n = 16; and fulvifrons, n = 15), and the Thick-billed Kingbird(T. crassirostris;n = 16) replacedno remigesor primarycoverts, other than the tertials on some birds. The eccentricreplacement pattern is well illustratedin the Vermilion Flycatcher(Figure 1, Table1). The numberof remigesreplaced ranged from all secondariesand primaries(in 53.7% of 82 first-yearspecimens exam- ined, identifiedby the retentionof juvenalprimary coverts)to nine of 19 feathers(five outer primaries and four inner secondaries;CAS 39678). The numberof outer primarycoverts replaced varied from none (23.1%) to five (2.4%). In all cases,replacement proceeded consecutively from the starting pointsto the outermostprimary and primary covertand to the innermost secondaryother than the tertials.The sequencewas confirmedwith eight specimenscollected during active molt. This pattern was also typical of first- year remex molts in the other flycatcherspecies showing the eccentric pattern,except for mostCassin's Kingbirds (see below). All replacementof remigesby VermilionFlycatchers occurs in the late summerand fall; no springreplacement of flightfeathers was found. First-yearYellow-bellied and westernWillow flycatchersdiffer from the VermilionFlycatcher in that remexmolt occurs during the springrather than the fall (seealso Dickey and van Rossem1938, Mengel 1952, Johnson 1963); adultsof these forms replace flight feathersin the fall (Johnson 1963, P. Unitt pers. comm.). First-yearOlive-sided Flycatchers appear to have one protractedover-winter molt of flight feathers, commencingin September or October and concludingin March or April, althoughthis needsconfirmation as only three specimensI examinedwere collectedon the winter grounds.Among
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