Adler and Ritchison • PROVISIONING BY GRASSHOPPER SPARROWS 
519 
beneficial because smaller grasshoppers, relative 
to their mass, contain a greater percentage of 
chitin (Kaspari and Joem 1993) and provide less 
energy per unit of mass. Adult Grasshopper 
Sparrows may not prey on large grasshoppers 
l> 35 mm) because they may be too large tor 
nestlings to consume, especially during the tirst 
few days post-hatching, and may require longer 
handling times (i.e.. removing chitinous pans). 
ACKNOWLEDGMENTS 
We thank Ben Sutter and Chris Rutledge for assistance 
with the field work, C. E. Braun, Myles Falconer, and an 
anonymous reviewer for helpful comments, and the EKU 
research committee for financial support. 
LITERATURE CITED 
Ammer. F. K. 2003. Population level dynamics of 
Grasshopper Sparrow populations breeding on re¬ 
claimed mountainiop mines in West Virginia. Disser¬ 
tation. West Virginia University, Morguntown, USA, 
Barba, E„ F. Atienzar, M. Marin. J. S. Monros, and J. 
A. Gil-Delc,ado. 2009. Patterns of nestling provi¬ 
sioning by a single-prey loader bird. Great Tit Pams 
major. Bird Study 56:187-197. 
BCdard. J. and M. Meiinier. 1983. Parental care in the 
Savannah Sparrow. Canadian Journal of Zoology 
61:2836-2843. 
Bjornstad, G. and J. T. Liejeld. 1996. Male parental care 
promotes early fledging in an open-ncster, the Willow 
Warbler Phylloscopus tmchilus. Ibis 138:229-235. 
Bowman, K. M.. C. M. LESSELLS. and .1. Komdkuk. 2005. 
Male Reed Buntings do not adjust parental effort in 
relation to extrapair paternity. Behavioral Ecology 
16:499-506. 
Cockburn, A. 2006. Prevalence of different modes of 
parental care in birds. Proceedings of the Royal 
Society of London, Series B 273:1375-1383. 
Conrad, K, F. and R. J. Robertson. 1993. Patterns of 
parental provisioning hy Eastern Phocbes. Condor 
95:57-62. 
Crossman. T, I. 1989. Habitat use by Grasshopper and 
Savannah sparrows at Bradley International Airport 
and management recommendations. Thesis. University 
of Connecticut. Storrs. USA. 
East. M. 1981. Aspects of courtship and parental care of 
the European Robin Eiithacus mbecula. Ornis Scan- 
dinavica 12:230-239. 
Falconer. C. M.. M. L. Mallory, and E. Nol. 2008. 
Breeding biology and provisioning of nestling Snow 
Buntings in the Canadian High Arctic, Polar Biology 
31:483-489. 
FlLLlATER, T. S. and R. BRFurwisCH. 1997. Nestling 
provisioning by the extremely dichromatic Northern 
Cardinal. Wilson Bulletin 109:145-153. 
Goodbred, C. O. and R. T. Holmes. 1996. Factors 
affecting food provisioning of nestling Black-throated 
Blue Warblers. Wilson Bulletin 108:467-479. 
GRUNDEL, R. 1987. Determinants of nestling feeding rates 
and parental investment in the Mountain Chickadee. 
Condor 89:319-328. 
KaRASOV, W. H. 1990. Digestion ill birds: chemical and 
physiological determinants and ecological implica¬ 
tions. Studies in Avian Biology 13:319—415. 
Kaspari, M. 1990. Prey preparation and the determinants 
of handling time. Animal Behaviour 40:118-126. 
Kaspari, M. 1991. Central place foraging in Grasshopper 
Sparrows: opportunism or optimal foraging in a 
variable environment? Oikos 60:307-312. 
Kaspari. M. and A. Joern. 1993. Prey choice by three 
insectivorous grassland birds: reevaluating opportun¬ 
ism. Oikos 68:414—430. 
KLASING. K. C. 1998. Comparative avian nutrition. CAB 
International. Wallingford. United Kingdom. 
MacCoi.l. A. D. C. and B. J. HaTCHWELL.. 2003. Sharing 
of curing: nestling provisioning behaviour of Long¬ 
tailed Tit. Aegithalos caudatus . parents and helpers. 
Animal Behaviour 66:955-964. 
MaGRATH, R. 1991. Nestling weight and juvenile survival 
in the Blackbird (Tardus merula). Journal of Animal 
Ecology 60:335-351. 
Mitchell, D. p.. P- O. Dunn, l. a. Whittingham, and C. 
R. Freeman-Gallant. 2007. Attractive males provide 
less parental care in two populations of the Common 
Yellowthroal. Animal Behaviour 73:165-170. 
Nordlund. C. A. and C. A. Barber. 2005. Parental 
provisioning in Melospiza melodia (Song Sparrows). 
Northeastern Naturalist 12:425-432. 
Olson, V. A., A. Liker. R. p. Freckleton, and t. 
SZfcKELY. 2008. Parental conflict in birds: comparative 
analyses of offspring development, ecology and 
mating opportunities, Proceedings of the Royal 
Society of London. Series B 275:301-307. 
RiNGSBY, T. H.. B.-L. S/RTHER. AND E- J. SOLBERG. 1998. 
Factors affecting juvenile survival in House Sparrow 
Passer domestu us. Journal of Avian Biology 29:241-249. 
ROYAMA, T. 1966. Factors governing feeding rate, tood 
requirement and brood size of Great Tits Parus major. 
Ibis 108:313-347. 
RYTKONEN, S.. K. KOIVULA. AND M. ORELL. 1996. Patterns of 
per-brood and per-offspring provisioning efforts in the 
Willow Tit Parus mantanus. Journal of Avian Biology 
27:21-30. 
SAS Institute. 1999. SAS users guide: statistics. SAS 
Institute Inc., Cary. North Carolina, USA. 
Schadd. C. A. AND G. Ritchison. 1998. Provisioning ol 
nestlings by male and female Yellovv-brCasted Chats. 
Wilson Bulletin 110:398—402. 
SCHWAGMEYER, P, L. and D. W. MOCK. 2008. Parental 
provisioning and offspring fitness: size matters. 
Animal Behaviour 75:291-298. 
SMITH. R- L. 1963. Some ecological notes on the 
Grasshopper Sparrow. Wilson Bulletin 75:159-165. 
Stodola, K. W.. E. T. Linder, D. A. Buehler, K. E. 
FRANZREB, AND R. J. COOPER. 2009. Parental care in 
the multi-brooded Black-throated Blue Warbler. Con¬ 
dor 111:497-502. 
TrEmont. S. and H. A. Ford. 2000. Partitioning of 
parental care in the Leaden Rycatcher. Emu 100:1-11. 
