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Artificial Incubation of Eggs of Various Bird Species And Vol. XXXII 1961 WETHERBEE,D. K. AND N. S.,•4rtificial Egg lncubatio• [141 HOLLOM, P. A.D., and H. G. BROWNLOW.1955. Trapping methods for bird ringers. (Rev. edition) British Trust/or Ornithology,Field Guide No. 1, 56 pp. HUSSELL,D. J. T., and JAMESWOODFORD. 1961. The use of ,a Heligoland trap and mist-nets at Long Poin,t, Ontario. (in •his issue) LINCOLN,F. C., and S. P. BALDWIN. 1929. ,Mant•al for bird banders.U.S. Dept. of Agriculture, Misc. Publ. No. 58; 1-112. LO.CKLEY,R. M., and R. RUSSELL. 1953. Bird-Ringing. Crosby, Lockwood and son. M[ACKINTOSH,J.R. 1956. Planting your garden for wild hirds. Audubon Society of Canada, Toronto. _MATHER,J. R. 1960. An alternative to the sleeve nsed on catching boxes. The Ringers' Bulletin, 1 (8): [10-11, unpaged]. SPENCER,ROBERT. 1959. Progressand prospectsin ringing. Ibis, 101: 416-424. TEBBES,JOHN K. 1953. Songbirds in your garden. Cromwell. WEIGOLD,HUGO. 1956. How we began ringing on Heligoland. The Ring, 8: 159-162. [_WHITTLE,CHARLE• L.] C.L.W. 1930. I"Der Vogelzug," a new ornkhological journal.] Review in Bird-Banding, 1: 83. WILLIAMSON,KENNETH. 1957. Mist-nets versus,Heligoland traps. B[rd-Band•'ng, 28: 213-222. WOODFORD,J. 1959. The use of mist-netsand a Heligol,and trap at Point Pelee. Bird-Banding, 30: 38-46. Royal Ontario Museum, 100 Qaeen's Park, Toronto, Ontario, Canada: 17 Farns- worth Drive, Weston, Ontario, Ca,ada. Received February, 1961 ARTIFICIAL INCUBATION OF E'GGS OF VARIOUS BIRD SPECIES AND SOME ATTRIBUTES OF NEONATES By DAVIDKENNETH WETHERBEE and NANCYS. WETHERBEE Introduction. In connectionwith a study of the morphology of just-hatchedbirds we have accumulatedcollateral data that may be of somegeneral value to ornithologists.More than 2,000 specimensof 83 specieswere hatched in incubators. No speciestried in adequate samplesfailed to be amenableto artificial incubation. Incubation data determinedin the laboratory where conditionscan be standardizedare basicto an understandingof the biologicalproblems of natural incuba- tion studiedby ecologists. Evans (1891 and 1892) in Scotlandand Heinroth (1908 and 1922) in Germanyfirst and mostextensively hatched out Old World species. Baldwinand Kendeigh(1932), Kendeigh(1940), and Graber i1955'• investigatedartificial incubationof wild birds' eggs in America. Presentmanuscript was completed in 1957. Acknowledgments. Grateful acknowledgmentis made to the many peoplewho helpedsearch for nests. Dr. Kay T. Rogersen- couragedthe useof artificial incubationin 1951 when sciencefounda- tionsspurned the practicality of arti'ficialincubat4on of wild birds'eggs. Bird-Banding 142] WETHERBEE,D. K. aN,• N. S..Arti/•cial Egg lacub,tio,, stay Dr. Richard Graber kindly contributedseveral speciinenshatched in his incubator. Incubators were made available by Dr. Rudolph F. Nunnemacherat Worcester, Massachusetts,Dr. Bradley M. Patten at Ann Arbor, Michigan,Dr. Hugh Clark at Storrs,Connecticut, and the late Dr. Charles S. Johnson at Nashville, Tennessee. Dr. Oliver L. Austin, Jr. and otherskindly mademany suggestionsthat were adopted in improving the manuscript; responsibilityfor the accuracy and organizationof the final draft howeveris entirely our own. Methods. All the eggsof each clutch found were collected,whether or not the clutch was complete. Date, time, locality, number of eggs in the clutch,degree of certaintyof speciesidentification, and data on the incubationattendance of the parent were recorded. A separatedata sheetfor eachegg was kept. At the incubatora clutch-and egg-number was brushedon each egg. The length of time and conditionsof trans- potration from the field were recorded. Each egg was measured or weighed,put in waterto determinebuoyancy, and candiedbefore being pu,t into the incubator. Efforts were made over the 6-year period to obtainas manyeggs as possible. Artificial incubationwas aimed at 37.8øC. and 64 percent relative humidi,ty.Most of the eggswere incubatedin forced draft, 600-chicken- egg capacity,American-Lincoln cabinets. The eggs,rolled twice daily, were held in open, partitioned petri dishes lined with gauze. To determinevolumes, most of the eggs were weigheddaily until they reachedspecific gravity of unity as determinedby buoyancyin waterat 37øC. At this point egg weightsin gramsequal their volumesin cubic centimeters.The comparisonof volumesis more significantthan com- parison of weights of fresh eggs (or at least is more amenable to standardization)because for practicalpurposes there is no suchthing as a "fresh egg" in nature. Balancescould be read to at least four digits and were sensitiveto at least ,threedecimal places. Thin-shelled eggs,such as those of swallows,were dried completely by blottersbefore being returned to the incubator. When volumeswere not determinedin the mannerjust describedmost other eggswere measuredto greattest length and width by vernier calipers. Infertile and addled eggswere discardedas soon as detected. Dead embryoswere examined for anomalies.The timesof first peepingfrom the egg and of the first pipping of the shell were noted. Efforts were madeto preservethe hatchlingat, or as near the momentof, hatching as possible,as this was critical to the centralstudy (Wetherbee,1959). This time was recordedas plus or minus the nearesthour, one-half the interim betweenincubator tendings. The partitionsof the petri dishes isolated each "altricial" chick and its numbered shell from other hatchings;with "precocial"(in the strictornithological sense) species eachegg was individuallybagged. Each chick was weighed,described as to behavior,color of skin, mouth-liningand down,and often comparedto other specieshatched simultaneously.Each chick was labelled,tied in a gauzebag, and preservedin 10 percentneutral formalin or 75 percentethyl alcohol. Specimenswere la,ter transferredto 95 percentalcohol in individual snap-capvials. The shellswere collected and put into dry storagefor referencein caseof curatorialerrors. This work attemptsto establish Vol. 1961XXXlI WETHERBEE,D. K. AND N. S.,•4rtiy•cial Egg Inc,batior, [143 a standard for data on incubation under standardized conditions. Such measurementsmay allow for more accurateand more significan,tinter- specificcomparisons, for there should be (ideallyl very little intra- specificvariation. Embryologicaldevelopment and incubationperiods in the wild, however,vary greatly as parental attentiveness,environ- mental temperature,moisture, etc. are conditionswhich cannot be measuredaccurately. Definitions of terms. Incubation period is defined as the longest time it takesany egg of the givenspecies to hatch spontaneouslyunder artificial incubationin forced air incubatoraimed at 37.8øC., 64 percent relative humidity. Such standardizationis necessary,because, as every commercialhatcher knows, embryologicaldevelopment is sensitiveto slightdifferences in temperatureor humidity (Huggins,1941). Ideally, experimentalwork may some day establishthe optimum temperature for incubationof each species(Baldwin and Kendeigh, 1932, and Graber, 1955). Incubation periodsgiven here are the longestthat we recordedand are not necessarilyentire maximum potential incubation periods,though most are probably representative.Incubation period data were not usedwhen the chick required aid in emergingfrom the shell. Egg clutchesof larger specieshad greater variability in total emergencethan egg clutchesof smaller species. Mortality statistics for artificial incubationhave no bearing on incubationmortality rate in the wild. The expression of temperature in O.IøC. rather than 1.øF. is advocatedin the interestsof establishinga standard of high critical rigor sorelyneeded for comparativepurposes in this type of ornithologi- cal work. We have purposelyexpressed time duration in hours rather than days-and-hours. We believe that the sad state in which the literatureof incubationfinds itself can be rectifiedif ornithologists;viii use consistentlythe primary units of temporal embryology--hours. Even when we made a conscious effort to record to the neares,t hour we often came up with a hatchingtime "plus or minus 10 hr."; we can't help but wonder what the margins of error are in statementsin the literaturerecorded and expressedin days. What is worse,ornithologists often get confusedas to whetheror not to count.the first day; or they limit the meaningof day to calendar day; or they record incubation time in days and estimatedfractions, then later convert to hours. Interim betweenhatchi•gs. As eggsmay undergosome development in the nest due to variable, warm environmentaltemperature, even thoughthe eggsare not incubatedby the parent, the interim between hatching used as an indication of the relative time of the onset of parental incubationis of most value when expressedas the minimum interim observedin severalcomplete clutches all of which hatch, rather than as the maximum interim. Clutch weight. Clutch weights were reported only from actual clutchesknown to be completeby virtue of a hatchingtime in the incubator considerablyless .than known incubationperiods. These data should be distinguishedfrom clutch weights determinedfrom calculationsand extrapolations. Weights.Averages were determined from segregatedclutch averages, Bird-Banding ]44] W•:T•:I•:•:,D. K. a•I) N. S.,Arti]•cia[ Egg Incubation not arithmetic summation of individuals, unlessotherwise stated in the text. Neonatalweights were taken of specimensthat had just dried, after beingtaken directly from the constanthumidity of the incubator. Many chickswere observed,to defecateafter hatching: this probably accountsfor most disparities between
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