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Variations in the Weight of Birds • 416 BALDWINANDI•ENDEIGH, Variations in Bird Weights [July[Auk VARIATIONS IN THE WEIGHT OF BIRDS • BY S. PRENTISS BALDWIN AND S. CHARLES KENDEIGH INTRODUCTION I• spiteof the fact that the weight of birds may be easilyobtained, there is very little informationavailable for wild species. Very few collectorsin the past have taken the trouble to weigh the birds that they obtain for taxonomicpurposes. One reasonfor this may lle in the lack of a suitable portableweighing instrument. A moreimportant reason why the weighing of birds has not been more commonlydone may lie in the recognizedvari- ability of bird weights and the consequentlack of appreciationof their importance. Yet the weight of birds and the variationsand fluctuationsof theseweights furnish criteria of considerableimportance in the understand- ing of the physiologicaland ecologicalreactions of the bird as a living organism. The physiologyof the bird, its behavior, and the influenceof environmentare interacting factors, no one of which can be understood without a knowledgeof the two others. The weighing of a]] birds collected might well be made a fundamental policy in all museums. There is a de- velopingtendency to do so amongsome of the youngerornithologists. A good accuratebalance may be obtained at a very reasonableprice. Bird- banding operators have an especiallygood opportunity for obtaining weightsof living birds of many speciesand alsoof the sameindividual bird at frequent intervals. At the Baldwin Bird ResearchLaboratory, Gates Mills, near Cleveland, Ohio, the first weights of birds were obtained in 1925, and in 1926 the weighingof all birds taken from the bandingtraps becamean established practice in the laboratory routine. The presentpaper is an analytical summaryof the data obtainedduring the nine yearsup to January 1, 1934, in the courseof which 13,546 weights were obtained of 5812 individuals representingeighty-five species. More than ten individualswere weighed in eachof thirty-threespecies, and thesereceive major considerationin this report. Acknowledgmentis made of aid from th e fo]]owlngpersons who, as as- sistantsin the laboratory during various periods,have been of much help in gatheringthese records: Rudyerd Bou]ton,W. W. Bowen,Leonard G. Wor]ey, Delos E. Johnson,Carl Johnson,James Stevenson, Theodore C. Kramer, and RoscoeW. Franks. We wishto make specialacknowledgment to Dr. Worley who was actlve]y interestedin this line of work while at the laboratoryand madea usefulpre]Jminary summary of the first four years' data. Contribution No. 32 from the Baldwin Bird Research Laboratory, Gates Mills, Ohio. Vol.1038 55]J BALDWINANDKENDEIGH, Variations in Bird Weights 417 LITERATURE No attempt is here made to review in an exhaustivemanner all the litera- ture on bird weights. Attention is centerednot on the averageand extreme weightsof individual species,but on analysisof the variationsin the weight of individuals and speciesunder different circumstances. Data on the weight of variousspecies have beenrecently obtained or compiledfrom the literatureby Krohn (1915), Bergtold(1917, 1926),Hesse (1921), Heinroth (1922), Przibram (1922), Weigold (1926), Whittle and Whittle (1926), Esten (1931), Roberts(1932), Groebbels(1932), Fiora (1933-34), Wether- bee(1934), Marpies (1935), Mountfort (1935),Stewart (1937), lmler (1937) and Nice (1938). Severalof theseauthors, as well as I(nappen (1928),give usefulbibliographies relative to the weightsof birds. Someauthors are content with giving singleor averageweights, with or without indication of the limits of variation. In some cases, distinctions in weight betweenthe two sexesor betweenadult and immature birds are made, but seldomis any extensiveanalysis given. Thosein the abovelist who have obtainedweights of living birds taken from banding traps are Whittle and Whittle (1926), Wetherbee(1934), Marpies (1935), Mount- fort (1935), Stewart (1937), lmler (1937) and Nice (1938). Most students, especiallythose who have weighedliving birds and often the sameindividual at differenttimes, are impressedwith the great variability in their weights. Groebbels(1932) finds this variability relatively greaterin small than in large birds. Linsdale and Sumner (1934) adequately emphasizethis variability in their statement that the "weight of a bird is not a static quality but is one of continuousand ordered change." Other literature dealingwith specificproblems will be consideredin the varioussections that follow. METHODS Practicallyall the weightsentering into the presentstudy were obtained from living birds; a very few were obtainedfrom birds collectedfor other investigations. All the living birds were gathered,at intervals of two or three hours, from the banding traps scatteredover the fifteen or more acresimmediately surrounding the laboratory. They were broughtto the laboratory in small gathering cagesand weighedimmediately. All the birds had thus been recently feedingand had a varying amount of food undergoingdigestion in their alimentarytract. Differencesin activity and diet previousto enteringthe trapsmay haveinfluenced the weightsobtained, but thesewere beyondcontrol. Accordingto Stevenson(1933), the amount of food in the stomachsof twenty-fiveadult SongSparrows, I English Sparrows,Starlings, and White- Scientific names of species are given in Table 5, p•ge 436. 418 BALDWINANDKENDEIGH, Variatio• in Bird W•ights [•u•ky breastedNuthatches averaged about 1.5 per cent of the body weight. If as much is in the small intestineand a similar amountis in the large intes- tine and cloaca,about 4.5 per cent of the body weight may representun- assimilatedmaterial. Single excrementdroppings in these small birds averageless than 0.5 per cent of the body weight. A correctionof this amount(plus an allowanceof about0.5 per cent for weightof the band on the bird) may be made,if desired,to givethe approximatebasic weights of the birds. This correctionhas not beenmade in the presentstudy because of a desireto follow the natural weight fluctuationsof birds in the wild. The bird for weighingwas placedin a rectangularaluminum box (5 by 3 by 3.5 inches)with a looselyfitting cover. As the bird washere in darkness it generallyremained quiet, and therewas no dangerof suffocation.Weights were taken on a small Cenco balance. The box was accurately counter- balancedby meansof No. 12 shot,so that the weight of the bird couldbe read directly without subtractingthe weight of the box. The counter- balancingof the box wasperformed before each weighing because of gradual accumulationof excrementvoided by the birds while in the box. The bird was banded and examinedbefore the weighingwas done, so that by re- movingthe coverfrom the box, it couldat oncebe releasedwithout further handling. The bird was not in any way harmedby the procedureand often returnedto the traps severaltimes a day. The weightsof wet or obviously sick birds were not included with the other data. A largely negligibleloss in weight of the birds may have occurredfrom time of removalfrom the trapsuntil the weightwas taken. A carefulcalibration and checkingof the balanceand weightsindicated in weighingup to 20 grams,an accuracyof 4-0.1 gram, and between20 and 200 grams an accuracyof 4-0.2 to 0.3 gram. Sincemuch of the followingdiscussion is basedon weight averages, many of thesesmall plus and minusinaccuracies in individualweights are largely eliminated. Sinceweights of all birdsare includedin this study, althoughsome indi- vidualswere weighed only onceand othersmany times, a specialstudy was madeof the effectof the 'trap habit' on fluctuationsin weight. It might be expectedthat wherea bird comesrepeatedly to the traps and thus obtains muchof its food, there would be an interference in the weight physiology of the bird. Adult ChippingSparrows were usedto study this point because only in this specieswere there sufficientrecords for a singlemonth (May) betweenthe hoursof 8.30 a.m. and 5.30 p.m. free of the influenceof age and sex (seebeyond). Table 1 showsan increasedamount of fluctuation in the weight of an individual,as representedby the averagestandard deviation, with an in- creasednumber of timesit repeatsin the traps. This increase,at leastthe extremevalue, is of fairlyhigh statistical significance. However, this in- Vol.1938 5.51J BALDWINANDI•ENDEIGIt, Variations in Bird Weights 419 Variability in weightof adult ChippingSparrows correlated with numberof times capturedand weighed;there is no sex differencein weightof this species Number of times Number of Averageweight Averagestandard bird weighed birds in grams deviation, grams• 2- 6 30 12.4 4-0.40 7-10 24 12.4 0.50 11-20 15 12.2 0.54 21- 4 12.2 0.63 creasedfluctuation in weight seemsnot to be due to any influenceof the trap habit in itselfbut rather to the longerperiod of time necessarilycovered duringwhich the bird is subjectedto a largernumber of influencingfactors. Anotherreason for believingthat the trap habit has no intrinsiceffect is that the averageweight of the birds is nearly the sameregardless of the number of times they were caught. The differenceof 0.2 grams is not statisticallysignificant. In the easeof a very few individual birds of other species,notably the Cowbird,and occasionallya SongSparrow, a persistent trap habit did showa positiveeffect, in beingassociated with a considerable lossof weight. Suchrecords were eliminatedfrom the averagesgiven in this paper. In mostinstances the numberof recordsobtained from individ- uals that did not acquirethe trap habit greatlyoutnumbers those that did, and sono correctionor further considerationof
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