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The Function of Singing in Female Black-Headed Grosbeaks (Pheucticus Melanocephalus): Family-Group Maintenance

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The Function of Singing in Female Black-Headed Grosbeaks (Pheucticus Melanocephalus): Family-Group Maintenance THE FUNCTION OF SINGING IN FEMALE BLACK-HEADED GROSBEAKS (PHE UCTICU$ MELANOCEPHAL US): FAMILY-GROUP MAINTENANCE GARY RITCHISON 1 Departmentof Biology,UMC 53, Utah StateUniversity, Logan, Utah 84322USA ABSTRACT.--Theresponses of young Black-headedGrosbeaks (Pheucticus melanocephalus) to playbackof the songsof parentsand strangerswere examined.I found that young gros- beaks moved about more, called more, and were more often oriented toward speakers in responseto parental song(both male and female) than in responseto the songsof strangers. I suggest that such a responseis part of a system used by Black-headedGrosbeaks to maintain family groups after the young fledge. In this system, an adult with food, but unawareof the locationof its young, singsto elicit beggingfrom its young. The parent bird is then able to locateand feed its young. Received31 August1981, accepted26 May 1982. AMONG passerines, song is typically the and Hooker 1969, Bertram 1970, Payne 1971). function of the male. There is also much evi- Concerninginstances of female singing other dence,however, of regular, occasional,and ar- than antiphonalsinging or duetting, Kern and tificially inducedsong by females.It has long King (1972)have suggesteda number of func- been recognized,for instance,that femalesof tions, including stimulating the breeding ac- many speciescan be induced to sing by injec- tivities of the male. Nottebohm (1975) sug- tion of testosterone (Baldwin et al. 1940, Kern gestedthat singingby femalesmay influence and King 1972). There are also many reports of a bird's socializationand choiceof partner and, speciesin which the female sings only in ex- in some cases, aid in territorial defense. ceptionalcases, e.g. the WesternMeadowlark Among the speciesof birds in which singing (Sturnellaneglecta; Lanyon 1957), Indigo Bun- by females has been reported is the Black- ting (Passerinacyanea; Nolan 1958), Song Spar- headed Grosbeak (Pheucticusmelanocephalus; row (Melospizamelodia; Van Tyne and Berger see Fig. 1). Weston (1947), in a general study 1976), EasternPhoebe (Sayornisphoebe; Smith of the breeding behavior of the grosbeak,re- 1969), Eastern Bluebird (Sialia sialis; Morton et ported that femalessang "while incubatingor al. 1978), and White-crowned Sparrow (Zono- brooding, usuallyas the male comesto take his trichialeucophrys; Kern and King 1972).In oth- placeon the eggsor young. Severaltimes dur- er species,singing appears to be a regularfea- ing nest-building, the female uttered songsin ture of femalebehavior, i.e. many specieshave the vicinity of the nest and always in the pres- beenreported in which femalescommonly sing ence of the male. The female will also occa- in a variety of situations (Table 1). sionally sing while foraging in the peripheral Despite these many observations, the sig- foliageof trees,but only when the male is close nificance of singing in females remains ob- by." Armstrong(1963) statedthat singing by scure. Possible functions, however, have been females tends to be characteristic of cardueline postulated. Armstrong (1963) suggestedthat finches.Van Tyne and Berger(1976: 249), in a antiphonal singing is a "means whereby con- general discussionof female song, suggested tact, rapport, and the social bond are main- that the songs of the female Black-headed tained." Duetting is believed to be important Grosbeakare nearly as elaborateas thoseof the in the synchronizationof breeding behavior male. and in the reinforcementof the pair bond (e.g. Despite these observations, the function of Thorpe and North 1965, Thorpe 1966, Hooker female songin the Black-headedGrosbeak, as in other species,is unclear. The present study 1 Present address: Department of Biological Sci- is an attempt to ascertainthe function(s) of this ences, Eastern Kentucky University, Richmond, song. The initial year of this 2-yr study was Kentucky 40475 USA. devoted to extensive field observation and re- 105 The Auk 100: 105-116. January 1983 106 GARY RITCHISON [Auk, Vol. 100 TABLE1. Reportsof singing in femalesamong various arian species. Situations in which Suggested female singsa function(s) a Species NB NR I B FY TD PB-M C F-GM Source Greenshank Simms 1958 Tringa nebularia European Nightjar X Selous 1905 Caprimulguseuropaeus Gray-capped Flycatcher X Skutch 1953 Myiozetetesgranadensis LoggerheadShrike X Armstrong 1963 Lanius ludovicianus European Dipper X X Van Tyne and Berger Cinclus cinclus 1976 American Dipper X X Bakus 1959a, b Cinclus mexicanus House Wren X X X Armstrong 1955 Troglodytesaedon Wrentit X Armstrong 1963 Chamaeafasciata Blue Wren X Robinson 1949 Malurus cyaneus ParadiseFlycatcher X Moreau 1949 Terpsiphoneviridis Elepaio X X X X Conant 1977 Chasiempissandwichensis Willie Wagtail X Robinson 1949 Rhipiduraleucophrys White-crowned Sparrow X X Blanchard 1941, Kern Zonotrichialeucophrys and King 1972 Grasshopper Sparrow X X X Smith 1959 Ammodramus savannarum Variable Seedeater X Gross 1952 Sporaphilaaurita Cuban Grassquit X Baptista 1978 Tiaris canora Gray Catbird X Palmer 1949 Dumetella carolinensis Mockingbird X Armstrong 1963 Mimus polyglottos Brown Thrasher X X X Thomas 1952 Toxostomarufum European Robin X Lack 1939, 1943 Erithacus rubecula Eastem Bluebird X Thomas 1946 Sialia sialis RussetNightingale Thrush X X X Skutch 1958 Catharus occidentalis Gray-cheekedThrush X X Van Tyne and Berger Catharus minimus 1976 European Blackbird Messmer and Messmer Turdus merula 1956 American Robin X Armstrong 1963 Turdusmigratorius Brown Towhee X X Marshall 1960 Pipilofuscus January 1983] Singingin FemaleGrosbeaks 107 TABLE 1. Continued. Situations in which Suggested female sings• function(s)a Species NB NR I B FY TD PB-M C F-GM Source Abert's Towhee X X Marshall 1960 Pipilo aberti Orange-billedSparrow X Skutch1954 Arremon aurantiirostris Rose-breasted Grosbeak X X X X Ivor 1944, Dunham Pheucticus ludovicianus 1964 Northern Cardinal X X X X X Laskey 1944 Cardinalis cardinalis Common Grackle X Wiley 1976 Quiscalusquiscula Greenfinch X Ferguson-Lees1943 Carduelis chloris American Goldfinch X Berger 1953 Carduelis tristis Lawrence's Goldfinch X Linsdale 1950 Carduelis lawrencei Red Crossbill X Lawrence 1949 Loxia curvirostra White-winged Crossbill X Bent 1968 Loxialeucoptera Magpie Lark X X Robinson 1949 Grallina cyanoleuca Butcherbird X Robinson 1949 Cracticussp. Black-backedMagpie X Robinson 1949 Gymnorhinatibicen • NB - nest-building; NR = nest-relief; I - incubation; B - brooding; FY - feeding young; TD - territorial defense; PB-M - pair-bond maintenance;C - courtship;F-GM - family-groupmaintenance. cording. During this period the situations in singing. Such singing apparently serves a ter- which females sang were noted, and hypoth- ritorial functionand, in addition, probably en- eses concerningthe function(s) of this song ables the female to maintain contact with the were derived. During the secondyear of the male. Females infrequently sang while forag- study, these hypotheses were tested experi- ing near the male (Weston 1947, pers. obs.). mentally. Singing, alone, is not sufficient to maintain a territory, at least early in the breeding sea- SYNOPSIS OF THE BLACK-HEADED son, and agonistic encountersinvolving chas- GROSBEAK BREEDING CYCLE ing and even physical contact occurred. Nearly all chasesinvolved males, although several fe- The first birds arrived in the study area (Mal- male-female chases were observed, and, on ibu-Guinavah Campground, in Cache Nation- three occasions, females were observed chas- al Forest 10 km east of Logan, Cache County, ing males. On one of these occasions,a female Utah) about the first week in May. Observa- was observedchasing a male, and, upon land- tions indicated that some birds were paired ing, she sang one loud song. On another oc- upon arrival. Such pairs may have been formed casion a female appeared to engagein a brief on the wintering grounds or during migration. singing duel with a neighboring male. During the early part of the breeding season, Following territory establishment, Black- paired birds foraged together within their ter- headed Grosbeaksbecome progressivelyless ritories. The female usually followed the male aggressive.This change in behavior was quan- as he moved through the territory feeding and tified in two ways: (1) male singing rates de- 108 GARY Rr?cmso2½ [Auk, Vol. 100 :] TIME (•) TIME(•) TIME(•) TIME(•) Fig. 1. Representativesonograms of the songsof male (above)and female(below) Black-headed Gros- beaks. clined as the seasonprogressed, and (2) the Incubation.--Both sexesare surprisingly vo- frequencyof intraspecificagonistic encounters cal on and around the nest. Males frequently (i.e. chasesand/or actualphysical encounters) sangwhile incubating.At timesthis songap- decreased(Fig. 2). peared to be in responseto the singing of Nest building.--Thefemale normallybuilds neighboringmales, i.e. a male would be qui- the nest, and I observed three instances in etly incubatingwhen, upon hearinga neigh- which females sang while involved in such boring male singing,the incubatingbird be- construction.Twice, females sang while gath- gan to sing. At other timesthe male'ssinging ering nestmaterials, and, on anotheroccasion, appearedto be a signal to the female that he a female was observedsinging while sitting in was about to leave the nest. a partially constructednest. On eachof these Femalesrarely sing while incubating.On two occasions a male was within a few meters of occasionsincubating females sang in apparent the singing female. responseto the singingof neighboringmales. January1983] Singingin FemaleGrosbeaks
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