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Effects of Band Color on Survivorship, Body Condition and Reproductive Effort of Free-Living Australian Zebra Finches

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Effects of Band Color on Survivorship, Body Condition and Reproductive Effort of Free-Living Australian Zebra Finches The Auk 111(1):131-142, 1994 EFFECTS OF BAND COLOR ON SURVIVORSHIP, BODY CONDITION AND REPRODUCTIVE EFFORT OF FREE-LIVING AUSTRALIAN ZEBRA FINCHES RICHARD ZANN Departmentof Zoology,La TrobeUniversity, Bundoora, Victoria, 3083 Australia ABSTRACT.--Indomesticated laboratory Zebra Finches (Taeniopygiaguttata), leg bands of certaincolors affect mate selection,parental effort, reproductivesuccess, sex ratio of offspring, and survivorship.I tested the effectsof these same colorson survivorship,body condition and reproductiveeffort of free-living Zebra Finchesover severalbreeding seasonsat two coloniesin northern Victoria,Australia. At the "Cloverlea" colony,adults of eachsex received either "attractive"or "unattractive"color bands.Nevertheless, survivorship to 12 months was not affectedby band color nor was there compellingevidence (mass, bill color and molt) for deterioratingbody conditiondue to reproductivecosts; there were no sexdifferences in survivorship.Reproductive effort eachbreeding season in thesefree-living birds may not be as great as that in some laboratory populationsbecause in nature the seasonis shorter and the intervalfor recoverycorrespondingly longer. In contrast,at the "Danaher"colony, where birds had unique color combinationsof three bands(plus one metal band), a retrospective analysisof four years of data showed that there were significanteffects of band color on reproductive effort of females;they laid more eggs if mated to males having a single red band (attractivecolor) in their combinationthan if mated to maleshaving a single light- green band (unattractive color) or bands of other colors (neutral attractiveness).They laid an extra clutch, but this did not result in significantly more fledglings and independent young being raisedby the pair. There were no band-coloror sex effectson survivorshipof adults at this colony. There were no obvious biases in the allocation of band colors or identificationof breedingpairs that might explainthe extra-clutcheffect. The finding on red- bandedmales is largely consistentwith that describedfor domesticatedZebra Finches, where femalesmake a differential reproductiveallocation in responseto their partner'sband color. No similar effectswere found for female band color where, contrary to expectation,females with black (attractive)bands had significantly fewer fledglings than did femaleswith color bandsof neutral attractiveness.The overall conclusionis that colorbands do not significantly affect the behavior of wild Zebra Finches, but it would be wise to omit red bands on males where possible.Received 18 August1992, accepted 8 December1992. COLOREDLEG BANDS affect mate selection (Bur- cluded that the potential of leg-band colors to ley 1981, 1986a, Burley et al. 1982), parental create biases in behavioral studies of natural investment (Burley 1988a), reproductive suc- bird populationsmay be overestimateddue to cess(Burley 1986b),sex ratio of offspring(Bur- many confounding effects. ley 1986c)and longevity (Burley 1985) in lab- A number of studies have investigated the oratory stocksof domesticatedAustralian Zebra effects of leg band color on behavior of wild Finches (Taeniopygiaguttata castanotis).The fact birds. Some a posteriorianalyses of long-term that wild-caught female Zebra Fincheshave the banding data have demonstrated negative ef- same band-color preferences as domesticated fectsof band color on reproductive success(Ha- ones(Burley 1988b)strengthens Burley's (1985) gan and Reed 1988), whereas others have found warning about possibleartifacts and biasescre- no effect (Beletsky and Orians 1991, Weather- ated by color marking of individuals in wild head et al. 1991). Similarly, findings are mixed populationsof Zebra Finchesand, by implica- with a priori designs:Metz and Weatherhead tion, other avian species.Conversely, Ratcliffe (1991, 1993) found negative effects, whereas and Boag(1987) found that, under experimental Cristol et al. (1992) found no effect. The influ- conditions promoting intermale competition, ence of leg-band colorson behavior appears to leg-bandcolors of their domesticatedlaboratory vary accordingto species,in particular whether Zebra Finchesdid not affect a male's ability to the color matchesone found on the bird's plum- compete for nest sites and females; they con- age or soft parts; if so, it may modify visual 131 132 RICHARDZANI• [Auk, Vol. 111 communication processeswith other members out the year, and continued to do so during the of the species. breeding season(Zann 1985). Consequently, if To investigate effects of band color on be- birds undergoreproductive stress resulting from havior of wild Australian Zebra Finches, I stud- reproductiveeffort it is feasiblethat feather re- ied the effectsof leg-bandcolors in two adjacent placementmay be slowed or suspendedduring breeding colonies("Cloverlea" and "Danaher") the period of stress.Reproductive stress might in the southeasternpart of the Australianwide also affect bill color, which becomes less red distribution of the species.In the color-band over the breeding seasonin wild Zebra Finches experiment at Cloverlea I tested specificallythe (Burley et al. 1992); in domesticated Zebra effect of band colors on body condition and Finches bill color declines during a single survivorship. All males received either red breeding cycle, and there is an overall decline bandsor light-green bands,and all femaleswere with prolonged breeding (Burley 1985, Burley given black or light-blue bands. The bands et al. 1992).Therefore, to detectchanges in body (supplied by A. C. Hughes, Middlesex, En- conditiondue to reproductivestress, I measured gland) were the same size, shade and material differences in mass, molt and bill color before (celluloidplastic) as those used by Burley(1981, and after breeding in birds of each color-band 1985, 1986a, b, c, 1988a) in her studies of do- type trapped at Cloverlea. mesticatedpopulations, where she found that The secondcolony in which the effectsof leg- red bands (attractive to females relative to un- band colorswere investigatedwas the Danaher banded males) conferred significantly greater colony. The study was not designedwith this survivorship(89 vs. 40%) on male wearersthan specific purpose in mind. Rather, it was de- did light-green bands (unattractive to females); signed to investigategeneral aspectsof behav- similarly, females with black bands (attractive ior and population dynamics;here birds were to males) lived significantly longer than fe- given unique combinations of color bands for males with light-blue bands (unattractive to recognition of individuals. Survivorship and males). Birds with bands of other colors were reproductive performance of many pairs was indistinguishable in attractiveness from un- monitored each breeding season. These data bandedbirds. Consequently,these colors were were analyzed,a posteriori,according to whether assumed to be neutral in effect. In order to test the partnershad attractivebands (red and black), this finding in a natural population I predicted unattractive bands (light-green), or neutral col- that, all things being equal, the red-banded or bands; however, no unattractive light-blue males and black-banded female Zebra Finches bands were placed on females in this colony. would havea higher survivorshipat the Clover- lea colony than their counterparts with light- METHODS green and light-blue bands. Survivorship was estimatedfrom the number of birds recaptured Color-bandexperiment.--Members of this colony 12 months or more after banding. nested and roosted in a tall windbreak on a wheat/ Burley (1985) postulatedthat an asymmetry sheep property (Cloverlea; 36ø09'S,145ø18'E) in the northern portion of the stateof Victoria and had oc- in reproductive effort and its associatedcosts cupiedthe site sinceat least1981 when I first discov- between the partners of the breeding pair ac- ered it. Zebra Finches were present each visit and counts for the differential effects of band color breeding normally ran for eight months, from Sep- on survivorshipof domesticatedZebra Finches. tember(austral spring) to April (australautumn; Zann Differential parental expenditure arises when and Straw 1984). The banding experiment ran from the parent with the attractive band color has a January 1987 through September 1989, and covered lower per-offspringinvestment than its lessat- two and one-half breedingseasons--the second half tractive partner (the differential-allocationhy- of the 1986/1987 season, and the whole of the 1987/ pothesis;Burley 1986b);the different costsdi- 1988 and 1988/1989 seasons. Birds were captured one day each month (except rectly affect their respectivelife spans.If free- for the months of May and August 1989 when no living Zebra Finches also incur reproductive birds were trapped) at a walk-in trap baited with a costsduring breeding, these may be reflected weekly ration of commercialseed. The assignmentof in massloss and slow feather replacement over band colors was random. Unbanded birds were this period. Zebra Finchesat a neighboring col- grabbed"blind" from the holding box and allocated ony ("Padgett";3 km eastof Cloverlea)replaced red or light-green bandsif they were males,but black remiges in a slow, continuousmanner through- or light-blue bandsif they were females.A few im- January1994] Bandcolor Effects in WildZebra Finches 133 matures were banded if their sex could be positively her"; 36ø09'S, 145ø26'E) 6.7 km east of Cloverlea. All determined; otherwise, they were released with a birds were given unique combinationsof three
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