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The Sequence of Molts and Plumages in Painted Buntings and Implications for Theories of Delayed Plumage Maturation’

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The Sequence of Molts and Plumages in Painted Buntings and Implications for Theories of Delayed Plumage Maturation’ THE CONDOR AJOURNAL OF AVIAN BIOLOGY UWWN-Y OF IDAHOl_lBRARY Volume 93 Number 2 AUG !2 &# ay 1991 The Condor93:209-235 0 The Cooper Ornithological Society I99 1 THE SEQUENCE OF MOLTS AND PLUMAGES IN PAINTED BUNTINGS AND IMPLICATIONS FOR THEORIES OF DELAYED PLUMAGE MATURATION’ CHRISTOPHER W. THOMPSON Department of Zoology,Arizona State University,Tempe, AZ 85287-1501 Abstract. Within a week of fledging,juvenile Painted Buntings,Passerina ciris, undergo a previously unrecognizedfirst prebasicmolt that includes most to all body plumage except greater primary and greater secondarycoverts. This molt occurs from early June to early October, usually on the breeding ground. The resulting first basic plumage is similar to juvenal plumage, but is more adult female-like in color in both sexes. First-year, but not adult, Painted Buntings subsequentlyundergo a presupplemental molt during which the outer four or five primaries, inner four to six secondaries,all remiges, and all body plumage except some to all greaterprimary coverts typically are replaced.The resultingsupplemental plumage is identical in both sexesand is more adult female-like in color than the previous first basicplumage. Presupplementalmolt usuallyoccurs between early Septemberand early November. Most (90%) Painted Buntingsalong the easterncoast ofthe United Statesundergo this molt on the breeding ground prior to fall migration; whereas, in the western United States, most (60%) molt in exclusively migratory areas of the desert southwesternUnited Statesand northwestern Mexico. These desert locationsexhibit a large increasein plant and insect life in response to “monsoon” rains that occur predictably in these areas in late summer and fall. Selection may have favored evolution of molt-migration strategiesin Painted Buntings in the western United States as a mechanism to allow them to molt in areas with greater food resourcesthan exist at the same time of year on their breeding or wintering ground. Painted Buntingsare one of only six passerinesin which molt-migration has been documented. All age and sex classesundergo a partial prealtemate molt that was describedpreviously (Fisk 1974) but has been overlooked by all subseauentinvestigators. This molt usuallv occurson the wintering ground, and is typically limited to the her& breast, and belly. Most plumagegrown by subadultmales during first prealtemate molt is adult female-like in color. This is the only passerineknown in which sexuallymature subadultmales grow adult female- like rather than adult male-like plumage during prealtemate molt. Juvenal plumage has poor structural integrity compared to subsequentplumages. Its structure facilitates heat transfer to the young by brooding females, but is poorly adapted to protectingjuveniles from adverseeffects of abiotic factors. Therefore, selectionmay have favored replacement of juvenal plumage with a structurally strongerfirst basic plumage as soon as possible after fledging. First-year males and females may undergo presupplementalmolt to prevent being iden- tified as first-year birds by adult females and being dominated by them (and possibly by adult males as well if adult males do not usually dominate adult females), or to reduce predation on themselves.That no subadultmales acquire a winter plumage intermediate in color between that of adult males and adult females in winter is the first empirical support for Rohwer et al.‘s (1980) model which indicates that selectionshould favor subadultmales with plumagesthat are completely adult female-like or adult male-like more than plumages that are intermediate between adult males and adult females. These resultsalso support the winter female mimicry hypothesis(Brown and Brown 1988), are consistentwith the winter I Received 17 October 1990. Final acceptance3 January 1991. 12091 210 CHRISTOPHER W. THOMPSON cryptic hypothesis (Rohwer and Butcher 1988), and are inconsistentwith the winter status signalinghypothesis (Rohwer 1975, 1982). Becausesubadult males grow plumage during first prealtemate molt that is adult female- like in color, this strongly suggeststhat the resulting plumage functions during their first potential breeding season to mimic females. This supports the summer female mimicry hypothesis(Rohwer et al. 1980), but is inconsistentwith the summer cryptic (Sclander 1965, 1972), summer statussignaling (Lyon and Montgomerie 1986, Montgomerie and Lyon 1986) andjuvenile mimicry hypotheses(Lawton and Lawton 1986, Foster 1987).This also supports Rohwer et al.3 (1980) model discussedabove. Adult males exhibit a greater rate of feather replacement during prealtemate molt than any other age or sex class. This suggeststhat the extent of this molt is less constrained in adult males than other age and sex classes,thus weakly supporting the molt constraints hypothesis (Rohwer and Butcher 1988). Becausesubadult males exhibit only a limited prealtemate molt, their summer plumage may not be functionally independent of their winter plumage. Thus, it is not possible to conclusively determine whether subadult male plumagesare adaptive during winter, sum- mer, or both. Becausepresupplemental body molt is essentially complete, this clearly in- dicates that failure of subadult males to obtain fully adult male-like supplementalplumage can not be due to energeticconstraints. Key words: Molt migration: delayedplumage maturation: Passerina;museum specimens; color; energetics;constraints. INTRODUCTION ually monochromatic and dichromatic, that have In nearly all speciesof birds, females reach sexual a variety of mating systemsand migratory strat- and somatic maturity at the same time (but see egies, and that occur throughout the world in Ligon 197 1, Hussell 1983, Stutchbury and Rob- diverse habitats ranging from arctic tundra at ertson 1987, Mountjoy and Robertson 1988). high latitudes to tropical rain forests at low lat- However, males of many speciesdo not attain itudes. somatic maturity in plumage or bare part color All hypotheses for the evolution of delayed for months to years after reaching sexual matu- plumage maturation propose that the relatively rity. In sexually mature but somatically imma- cryptic plumage of subadult males is adaptive ture (subadult sensuRohwer et al. 1980, Rohwer either during their first non-breeding season and Butcher 1988) males, these characters are (winter hypotheses)or their first potential breed- more cryptic than those in adult males (Rohwer ing season (summer hypotheses). The function et al. 1980, Lawton and Lawton 1986, Butcher of plumage color during breeding season(usually and Rohwer 1989) and usually are very similar summer) may be considered independent of to those of adult females. In many of these spe- plumage color during nonbreeding season (usu- cies subadult males successfullybreed occasion- ally winter) only when a complete body molt ally to regularly (e.g., Diamond 1972, Forshaw occurs between these seasons (Rohwer 1986, and Cooper 198 1, Proctor-Gray and Holmes Rohwer and Butcher 1988, Butcher and Rohwer 1981,Steenhofetal. 1983,Goodwin 1986,Grant 1989). 1986, Lawton and Lawton 1986, Rohwer and Although most studies of delayed plumage Butcher 1988). Such delayed acquisition of adult maturation have argued that subadult plumages plumage by sexually mature birds is termed de- evolved as adaptations for breeding, Rohwer and layed plumage maturation. Delayed plumage Butcher (1988) elegantly and unambiguously maturation has been studied almost exclusively demonstrated that subadult plumages often are in North American passerines (but see Foster adaptive only in winter. Males of 16 specieswear 1987, Jar-vi et al. 1987) but its occurrence is subadult plumages only during their first winter. geographically and taxonomically more wide- In all of these species,subadult males undergo a spread than is generally recognized. It occurs in complete body molt in spring which causesthem at least seven orders and 11 families of non- to lose their adult female-like winter plumage passerinesand at least 21 families of passerines and acquire a summer plumage that is very sim- (e.g., Mayr 1933, 1934; Lack 1968; Diamond ilar or identical to that of adult males. This in- 1972; Forshaw and Cooper 198 1; Schodde 1982; dicates that subadult plumage in these species Grant 1986; Lawton and Lawton 1986; Rohwer must be an adaptation to winter. and Butcher 1988) including speciesthat are sex- Males of 21 species wear subadult plumage DELAYED PLUMAGE MATURATION 211 during their first winter and subsequentsummer. tings are an ideal choice of speciesin which to In all of these species,subadult males undergo a look for a spring molt. In addition, three winter limited springbody molt thereby acquiring a more and five summer hypothesesimply testable pre- adult male-like plumage. Because this molt is dictions regarding subadult male molts and incomplete, summer plumage may not be func- plumages. These predictions were tested by de- tionally independent of winter plumage. There- termining the sequenceof molts and plumages, fore, this finding does not unequivocally support and resulting changesin plumage color, exhibited either winter or summer hypotheses.All plum- by male Painted Buntings from hatching to so- age replaced during spring molt in these 37 spe- matic maturity. The relevant winter and summer cies is always adult male-like and never adult hypothesesare summarized below. The predic- female-like or cryptic (Rohwer and Niles 1979, tions of these hypothesesare summarized in Ta- Rohwer et al. 1983, Rohwer 1986, Rohwer and ble 1 and
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