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Factors Affecting Feeding and Brooding of Gray Catbird Nestlings

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Factors Affecting Feeding and Brooding of Gray Catbird Nestlings FACTORS AFFECTING FEEDING AND BROODING OF GRAY CATBIRD NESTLINGS ELLEN J. JOHNSONAND LOUIS B. BEST Departmentof Animal Ecology,Iowa State University,Ames, Iowa 50011 USA ABSTRACT.--GrayCatbirds (Dumetellacarolinensis) were studied to documentpatterns of feeding and brooding nestlingsand to relate theseto nestlingage, brood size, time of day, time of season,and weather. Major factors affecting feeding were nestling age and brood size. The averagenumber of food deliveries.brood-•. h-• increasedlinearly with the log of nestlingage for both male and femaleparents. Also, volumeof food delivered/feedingtrip increasedlinearly with nestling age. As brood size increased,feeding frequencies/brood increasedbut food deliveries/nestlingdecreased. Brooding and shadingwere affectedmainly by nestlingage and ambient temperature.Time spent brooding decreasedsubstantially as nestlingsgrew older. Shading was important when the sun shone directly on the nest. Femalesspent significantlymore time brooding nestlingswhen rain increased.As ambient temperature increased, time spent shading increased and time spent brooding declined. Received7 May 1980, accepted27 April 1981. ONE of the greateststresses on birds during STUDY AREA AND METHODS the breeding seasonis caring for their nest- lings. Frequent feeding trips are required to This study was conductedin a shrubby and partly satisfythe food demandsof the rapidly devel- wooded pasture near Ames, Iowa during the springs and summers of 1977 and 1978. The area covered 16 oping young, and nestling body temperature ha of which approximately50% containedshrubs, must be maintained at a relatively constant primarily gooseberry(Ribes spp.) and multiflora rose level. Ratesof feeding and brooding by parent (Rosa multiflora), suitable as nesting sites for cat- birds can be related either to requirements of birds. A stream meandered through the study site. the nestlings or to environmental factors af- During 1976-1977, a drought occurred in central fecting the adults. Knowledgeof these rela- Iowa. Rainfall in Ames was 52 cm less than the nor- tionshipsis importantto understandbetter the mal of 116 cm during the 15-month period, May nestlingperiod of altricialbirds, but few work- 1976-July 1977 (Natl. Ocean. Atmos. Admin. 1976, ers have addressedthese topics. 1977, 1978). The drought ended in August 1977, and Although the Gray Catbird (Dumetellacaro- during April-July 1978,precipitation was 9 cm above the normal of 43 cm. linensis)is a commonspecies throughout much To locate all active nests, intensive nest searches of the United Statesand its nestingbiology has were conductedweekly over the entire study area. been extensivelydocumented (for a review see The status of active nests was checked at least once Johnsonand Best 1980), factors affecting feed- daily. Nearly all adult catbirdson the studyarea were ing and brooding frequencieshave not been capturedby using mist nets and marked with colored reported. Slack (1973) documented effects of leg bands.Additionally, a spot of paint was placed temperature,time of day, and day of incuba- on the head of one member of each pair. Nestlings tion on female attentivenessduring incubation were marked with a spot of paint on the top of the but reportedvery little concerningbrooding of beak. nestlings.Zimmerman (1963)collected limited Catbird nests were observed from blinds during information on attentive periodsof female cat- both 1977 and 1978. A mirror was positioned per- manently over each nest so that its contentscould be birds relative to nestling age. viewed. This was done prior to hatching; thus, birds The objectivesof this study were (1) to doc- became accustomed to the mirror before behavioral ument patterns of feeding and brooding nest- observationswere begun. Mirrors were positioned ling Gray Catbirds and (2) to relate these to so that they did not shade the nest. Feeding fre- nestlingage, brood size, time of day, time of quenciesand time spentbrooding and shadingnest- season, and weather. lings by each parent were recorded.Brooding was 148 The Auk 99: 148-156. January 1982 January1982] Feedingand Brooding Catbird Nestlings 149 distinguished from shading becauseeach served a TABLE1. Total volume delivered/feeding trip and different function. Brooding occurredwhen the fe- estimated volume delivered.nestling-•.h -• for Gray Catbird young of different ages.a male settled down on the nestlings and provided heat to them. During shading, the bird stood over Volume the nestlings,with wings slightly spread,usually to (cm3) protect them from sunlight (sometimes birds as- Number Volume (cma) delivered. sumed this position when no direct sunlight was •e (x) of trips delivered/trip nestling-•. falling on the nest). Observations were made at each ays) sampled (•)b (• + SD) h •e nest from the time of hatching until the nestlings 0 11 0.10 + 0.08 0.09 fledged. Nests were observed an average of 92 h 1 28 0.11 + 0.07 0.24 2 19 0.14 + 0.11 0.41 each. Ages of nestlings were divided into 24-h 3 12 0.24 + 0.15 0.60 classesbeginning from the time of hatching (0-24 4 31 0.26 + 0.14 0.80 h = 0 day old, 24--48h = 1 day old, etc.). Five nests 5 23 0.27 + 0.12 1.00 were observed in 1977, with brood sizes of 2• 2, 3, 6 23 0.41 + 0.27 1.22 7 46 0.38 + 0.25 1.44 4, and 5, and four in 1978, with sizes of 2, 3, 4, and 8 11 0.35 + 0.35 1.67 5. Nestling growth rates were determined by Rick- lefs' (1967) method. a For the size distributions of individual food items, see Johnson et al. 1980. In 1978, food samples were collectedfrom all cat- b Statisticsare: • = 0.075 + 0.047x, r2 = 0.96. bird nestlings on the area except those for which c Seetext for detailson how this was computed. feeding frequency was being observed. Wire liga- tures were placed around the nestlings' necks, pre- venting them from swallowing food delivered by the size. Because many of the independent variables parents (Johnsonet al. 1980). Nestlings were watched were intercorrelated (Appendix), all variables also from a blind, and, after each feeding trip made by were consideredsimultaneously in a multiple regres- parents, the food was collectedfrom the nestlings' sion procedureto determine the relative importance throats. Food was sampledfrom nestlings0 through of each. The level of significancefor statisticaltests 8 days old; after 8 days, disturbances would cause was set at P •< 0.05 unless noted otherwise. them to fledge prematurely (young normally fledge at about 11 days old). The volume of food items was RESULTS AND DISCUSSION determined by water displacement. Nearly all factorsconsidered had someeffect Each day was divided into five 3-h time periods: dawn-0900, 0900-1200, 1200-1500, 1500-1800, and on feeding or brooding frequencies,and inter- action effects were common. 1800--dusk. Observations from blinds and food sam- pling were conducted during alternate intervals, Nestlingage.---The average number of food with the schedulereversed every other day. In this deliveries-brood-•.h-•, (P), increasedlinearly way, all intervals were representedfor each part of with the logx0of nestlingage, (x), for both male the study during every 2-day period. Temperature (P = 4.14+ 3.74logIx + 1], r2 = 0.76)and fe- (øC),relative humidity (%), barometricpressure (mm male ('•= 1.55+3.43 logIx+ 1], r2= 0.83) Hg), cloud cover.(classes 1 through 5), wind speed parents. The logarithmic relationship shows (classes1 through 7), and light intensity (foot-can- that most of the increasein feeding frequency dies) were recordedhourly. occurredearly in the nestlingperiod. A similar The relationshipsbetween eachindependent vari- able and nestling care were evaluated using either relationshipbetween nestlingage and feeding linear regression analysis or analysis of variance frequencyhas been reported for other species (ANOVA). Nestling age had a noticeable and con- (Morton et al. 1972, Westerterp 1973). sistent influence on both feeding and nest atten- The volume of food delivered to nestlings/ dance. Thus, in ANOVA tests for the effects of brood unit time depends not only on frequency of size, time of season,and weather on the dependent feedingsbut alsoon the amountof food in each variables, a two-way analysiswas used in which one delivery. Both volume delivered/feedingtrip of the factorsalways was nestling age. Values used and the variability in food volume delivered/ in the analysiswere meansof the dependentvariable feeding trip increasedwith nestlingage (Table calculatedfor each age categorywithin each classof 1). These trends occurred through 8 days of the other independentvariable (unweighted analysis of cell means; Snedecor and Cochran 1967: 475). age, after which food could not be sampled. Regression equations were determined by using Most of the increasedenergy requirements of means of the dependent variable for each value of older catbird nestlingswere met by enlarging the independent variable. Using means in both of the amount of food delivered/feeding trip, the above proceduresadjusted for unequal sample rather than increasing the feeding frequency. 150 JOH•SO• •,•v BEST [Auk, Vol. 99 1•,25r ProportionofFeeding 1'5 •_• / Y= 0.272 + O.014x . i' ' ß ß , • O I 2 3NESTLING 4 5 AGE6 (DAYS)7 8 9 IO II Fig. 1. Nest attendanceand proportionof nestlingfeedings by femalecatbirds in relationto nestlingage. This was accomplishedby feeding largeritems lower than those of males (averaging about as the nestlingsincreased in sizeand were able one-half those of the males over the entire to consumethem (Johnsonet al. 1980). Volume nestling period), but females
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