VARIATION IN GROWTH OF NESTLING TREE SWALLOWS ACROSS MULTIPLE TEMPORAL AND SPATIAL SCALES Author(s): John P. McCarty Source: The Auk, 118(1):176-190. 2001. Published By: The American Ornithologists' Union DOI: 10.1642/0004-8038(2001)118[0176:VIGONT]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1642/0004- 8038%282001%29118%5B0176%3AVIGONT%5D2.0.CO%3B2 BioOne (www.bioone.org) is an electronic aggregator of bioscience research content, and the online home to over 160 journals and books published by not-for-profit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. The Auk 118(1):176±190, 2001 VARIATION IN GROWTH OF NESTLING TREE SWALLOWS ACROSS MULTIPLE TEMPORAL AND SPATIAL SCALES JOHN P. M CCARTY1 Department of Ecology and Systematics, Cornell University, Ithaca, New York 14853, USA ABSTRACT.ÐDifferences within a species in rates of growth of nestlings can be used as indicators of the quality of parental care, environmental conditions, and future success of offspring, whereas comparisons among different species may re¯ect a history of different ecological conditions or life-history strategies. The presesnt study examines the patterns of variation in growth in nestling Tree Swallows (Tachycineta bicolor) from across the species' range and compares Tree Swallows to other species. Growth of Tree Swallows was typical of other species in the family Hirundinidae. As a family, the Hirundinidae have slower growth than typical for passerines. Growth rate of species of Hirundinidae was not corre- lated with adult body mass or average brood size. Contrary to predictions, species that are double-brooded did not have higher growth rates, but swallow species living at higher lat- itudes did have higher growth rates than tropical species. Substantial variation in growth rates was observed among populations of Tree Swallows, yet the amount of variation ob- served between breeding colonies only a few kilometers apart, or from the same colony in different years, was as great as that seen in populations separated by hundreds of kilometers. Within a population, differences in growth among years were correlated with temperature and food supply when nestlings were being raised. No correlation between climate and growth was seen when comparing different populations. Differences between populations were not explained by local habitat, nor were large-scale geographic patterns evident. I used both experimental and observational evidence to evaluate the implications of short-term re- duction in growth for subsequent growth and survival. Nestlings were slow to recover from even very short periods of delayed growth that occur early in the nestling phase. Return of nestlings with experimentally or naturally induced delayed growth was reduced, which sug- gests that short interruptions in growth may have long term effects on post¯edging survival, even though mass at ¯edging is not affected. Received 9 August 1999, accepted 16 September 2000. NESTLINGS OF ALTRICIAL BIRDS exhibit sub- growth. In a broader context, those energy de- stantial variation in growth rates both within mands, coupled with physiological constraints and among species. Variation within a species on growth, accentuate the trade-offs that exist is often used as an indicator of variability in pa- between maximizing growth and attempting to rental care, environmental conditions, or nes- optimize other aspects of life histories (Lack tling quality. Growth rates of nestlings in¯u- 1968, Ricklefs 1984, Ricklefs and Starck 1998, ence length of time offspring are dependent on Starck and Ricklefs 1998b). their parents, their energy requirements and Within a species, nestlings with below-aver- rate of food delivery required of the parents, age growth or size at ¯edging generally suffer and length of time they are exposed to nest from reduced post¯edging survival (Gebhardt- predators (Lack 1968, Bosque and Bosque 1995, Henrich and Richner 1998). Reduced growth Halupka 1998). Variation among species is typ- may also have a long-term effect on ®tness, ically seen as a result of variation in life-history even when it does not appear to result in lower strategies. Altricial birds are among the fastest growing vertebrates (Case 1978), with most post¯edging survival, by decreasing the ability small passerines attaining full adult mass with- to obtain a breeding territory or mate or by in 10 to 20 days of hatching. High rates of en- lowering subsequent fecundity (Gustafsson ergy intake are necessary to sustain that rapid and Sutherland 1988, Richner 1992, Richner et al. 1989, Lozano 1994). Although it is clear that 1 Present address: Department of Biology, Univer- growth is a good indicator of future success, sity of Maryland, College Park, Maryland 20742, factors that determine variation in size and USA. E-mail: [email protected] mass are not as well understood. 176 January 2001] Variation in Nestling Growth 177 Previous studies have generally not differ- into the degree of plasticity possible on an evo- entiated between growth reductions that are lutionary time scale, whereas differences in life due to chronic food shortages or that are history among related species may indicate caused by poor parental care and those caused how growth responds to changes in the relative by short-term ¯uctuations in food supply with- demands placed on developing nestlings. Sec- in a season. Several groups of birds with food ond, I examined the signi®cance of short-term supplies that are subject to short-term ¯uctua- periods of reduced feeding on growth and the tions, such as seabirds (Hawksley 1957, Dunn possible long-term effects of those reductions. 1975, Konarzewski and Taylor 1989) and aerial Although the importance of chronic food short- insectivores (Koskimies 1950, Lack and Lack ages is well known, relatively little attention 1951, Bryant 1978, Wrege and Emlen 1991), are has been paid to short-term ¯uctuations in re- observed to undergo periods of interrupted sources and their effect on subsequent aspects growth and development under adverse con- of an individual's biology. I present observa- ditions, resuming normal growth once condi- tional and experimental evidence to examine tions improve. The long-term effects of those the importance of short-term reductions in temporary growth reductions have seldom growth. been explored, but if starvation interferes with critical developmental stages, permanent METHODS changes could result. Previous studies have found that nestlings subject to short-term Tree Swallows breeding in nest boxes were studied shortages of food eventually attain full body at the Cornell University Experimental Ponds Facil- ity (428309N768279W), near Ithaca, New York. This mass (Wiggins 1990b, Negro et al. 1994), mak- facility consists of two breeding sites located ap- ing it unclear whether such reductions in proximately 2 km apart. Unit One supported ap- growth have a long-term effect on post¯edging proximately 55 to 75 pairs of breeding Tree Swal- survival if survival depends solely on body lows, and Unit Two had between 10 and 23 pairs. size. Insect abundance was measured daily using suction In contrast to the view that intraspeci®c var- traps running during daylight hours, and high tem- iation is due to effects of the environment, var- perature was recorded daily. These sites and meth- iation among species is often viewed as the ods for sampling insects are described in detail in adaptive outcome of different selection pres- McCarty and Winkler (1999a, b). All values are re- ported as means 6 SE. sures. Predation rates, food availability, num- Nestlings of all ages were weighed to the nearest ber of breeding attempts per year, and level of 0.1 g during the 1990±1993 breeding seasons using competition among siblings are all thought to either Pesola spring scales or a portable Ohaus elec- in¯uence interspeci®c differences in growth tronic balance. In addition, nestlings were measured (Lack 1968, Bosque and Bosque 1995, Halupka on day 10 only in 1989, and those data are included 1998). Even though the sources of intraspeci®c in analyses of return rates. Swallows breeding at and interspeci®c variation in growth are ulti- those sites are monitored closely for the exact date of mately the same, the two types of variation hatching to determine nestling age. All nestling ages have seldom been considered together. are given as hatch day equal to nestling day 1. Lengths of ¯attened and straightened wing chord The present study employed two approaches (hereafter ``wing length''), the 9th (outermost) pri- to address the question of the ecological sig- mary, and 6th (outermost) rectrix feather were mea- ni®cance of variation in growth rates of Tree sured to the nearest 0.5 mm using a ruler with a wing Swallows (Tachycineta bicolor) and variation and feather stop. Length of the manus was calculated among the species in the Hirundinidae. First, I from difference between wing and 9th primary described both the inter- and intraspeci®c var- length. Length of the tarsometatarsus (henceforth iation in growth. Variation in growth among ``tarsus'') was measured to the nearest 0.1 mm using different species and among different groups of dial calipers. Nestling Tree Swallows typically Tree Swallows was compared to factors that ¯edge on day 21; disturbing nestlings after day 15 may cause premature ¯edging, so sample sizes for might contribute to variation, such as climate, older nestlings are small and come primarily from life history (clutch size and number of broods nestlings removed for other studies.