Abstract
Parental investment in offspring provides fitness benefits for progeny but incurs costs for parents. However, in birds there is a paucity of information on the foraging behaviors of altricial young once they leave the nest. Here we report a novel form of food acquisition in an altricial bird that is driven by a fledgling itself rather than its parents. In a population of densely nesting Red-winged Blackbirds (Agelaius phoeniceus) in Eastern Illinois, USA, we encountered multiple instances of a free-moving fledgling present in a focal nest that also had chicks too young to fly. We observed this behavior six different times in the field and obtained detailed video footage of six total instances at three independent nests, with two of these resulting in an apparent acceptance (i.e., feeding) of the focal fledgling by the attending parental adults. In one of the two nests where the fledgling(s) were accepted, feeding adults provisioned the fledgling as much as the average chick in the nest while at the other nest attending adults fed the fledgling above random expectations. Nest integration appears to be a novel form of food acquisition that is distinct from other fledgling self-maintenance behaviors.
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Data availability
All data is available in the manuscript. Supplementary material is available on Figshare.com (https://figshare.com/s/1dd6c20a769b785a73fd, https://figshare.com/s/bada505195147e412943, https://figshare.com/s/fa372e336170ec68bb7c).
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References
Antonson ND, Rubenstein DR, Hauber ME, Botero CA (2020) Ecological uncertainty favours the diversification of host use in avian brood parasites. Nat Commun 11:4185
Beletsky LD, Orians GH (1996) Red-winged blackbirds decision-making and reproductive success. University of Chicago Press
Brooke MDL, Davies NB (1989) Provisioning of nestling Cuckoos Cuculus canorus by Reed Warbler Acrocephalus scirpaceus hosts. Ibis 131:250–256
Brown CR, Brown MB (1988) A new form of reproductive parasitism in cliff swallows. Nature 331:66–68
Canestrari D, Bolopo D, Turlings TCJ et al (2014) From parasitism to mutualism : unexpected interactions between a cuckoo and its host. Science 80(343):1350–1352
Clutton-Brock TH (1991) The evolution of parental care. Princeton University Press, Princeton
Davies NB (2011) Cuckoo adaptations: trickery and tuning. J Zool 284:1–14
Davies NB, Welbergen JA (2008) Cuckoo-hawk mimicry? An experimental test. Proc R Soc B Biol Sci 275:1817–1822
De Mársico MC, Gantchoff MG, Reboreda JC (2012) Host-parasite coevolution beyond the nestling stage? Mimicry of host fledglings by the specialist screaming cowbird. Proc R Soc B Biol Sci 279:3401–3408
Delmore KE, Van Doren BM, Conway GJ et al (2020) Individual variability and versatility in an eco-evolutionary model of avian migration: migratory strategies of blackcaps. Proc R Soc B Biol Sci 287:20201339
Edwards S, Messenger E, Yasukawa K (1999) Do red-winged blackbird parents and their nestlings recognize each other? J Field Ornithol 70:297–309
Franchini P, Irisarri I, Fudickar A et al (2017) Animal tracking meets migration genomics: transcriptomic analysis of a partially migratory bird species. Mol Ecol 26:3204–3216
Gittleman JL (1988) Energy allocation in mammalian reproduction. Am Zool 28:863–875
Gloag R, Fiorini VD, Reboreda JC, Kacelnik A (2013) The wages of violence: mobbing by mockingbirds as a frontline defence against brood-parasitic cowbirds. Anim Behav 86:1023–1029
Gloneková M, Brandlová K, Pluháček J (2021) Further behavioural parameters support reciprocity and milk theft as explanations for giraffe allonursing. Sci Rep 11:1–9
Grim T (2007) Experimental evidence for chick discrimination without recognition in a brood parasite host. Proc R Soc B Biol Sci 274:373–381
Hauber ME (2006) A future cost of misdirected parental care for brood parasitic young? Folia Zool 55:367–374
Heinsohn RG (1991) Kidnapping and reciprocity in cooperatively breeding white-winged choughs. Anim Behav 41:1097–1100
Honza M, Šicha V, Procházka P, Ležalová R (2006) Host nest defense against a color-dimorphic brood parasite: great reed warblers (Acrocephalus arundinaceus) versus common cuckoos (Cuculus canorus). J Ornithol 147:629–637
Jackson MV, Kyne PM (2010) Potential host species fatally attack female eastern koel Eudynamys orientalis, a brood-parasite. Aust F Ornithol 27:133–136
Jamieson IG (1989) Behavioral heterochrony and the evolution of birds’ helping at the nest: an unselected consequences of communal breeding? Am Nat 133:394–406
Jamieson IG, Craig JL (1987) Critique of helping behavior in birds: a departure from functional explanations. In: Perspectives in ethology P. Bateson and P. Klopfer(eds). New York Plenum Press.
Kalmbach E (2006) Why do goose parents adopt unrelated goslings? A review of hypotheses and empirical evidence, and new research questions. Ibis 148:66–78
Katsis AC, Evans C, Common LK et al (2023) Intraspecific fledgling adoption by a pair of superb fairy-wrens Malurus cyaneus. Aust F Ornithol 40:49–55
Kilner RM, Langmore NE (2011) Cuckoos versus hosts in insects and birds: adaptations, counter-adaptations and outcomes. Biol Rev 86:836–852
Kilpatrick AM (2002) Variation in growth of brown-headed cowbird (Molothrus ater) nestlings and energetic impacts on their host parents. Can J Zool 80:145–153
Langmore NE, Hunt S, Kilner RM (2003) Escalation of a coevolutionary arms race through host rejection of brood parasitic young. Nature 422:157–160
Langmore NE, Cockburn A, Russell AF, Kilner RM (2009) Flexible cuckoo chick-rejection rules in the superb fairy-wren. Behav Ecol 20:978–984
Langmore NE, Stevens M, Maurer G et al (2011) Visual mimicry of host nestlings by cuckoos. Proc R Soc B Biol Sci 278:2455–2463
Latif QS, Grenier JL, Heath SK et al (2006) First evidence of conspecific brood parasitism and egg ejection in song sparrows, with comments on methods sufficient to document these behaviors. Condor 108:452–458
Lawson SL, Enos JK, Gill SA, Hauber ME (2021a) Eavesdropping on referential yellow warbler alarm calls by red-winged blackbirds is mediated by brood parasitism risk. Front Ecol Evol 9:1–10
Lawson SL, Enos JK, Wolf CS et al (2021b) Referential alarm calling elicits future vigilance in a host of an avian brood parasite. Biol Lett 17:5–8
Ligon JD, Stacey PB (1989) On the significance of helping behavior in birds. Auk 106:700–705
Lyon BE, Eadie JMA (2008) Conspecific brood parasitism in birds: a life-history perspective. Annu Rev Ecol Evol Syst 39:343–363
Medvin MB, Beecher MD (1986) Parent-offspring recognition in the barn swallow (Hirundo rustica). Anim Behav 34:161–163
Moskát C, Hauber ME (2007) Conflict between egg recognition and egg rejection decisions in common cuckoo (Cuculus canorus) hosts. Anim Cogn 10:377–386
Mota-Rojas D, Marcet-Rius M, Freitas-de-Melo A et al (2021) Allonursing in wild and farm animals: biological and physiological foundations and explanatory hypotheses. Animals 11:1–25
Numan M (2020) The parental brain: mechanisms, development, and evolution. Oxford University Press, Oxford
Packer C, Lewis S, Pusey A (1992) A comparative analysis of non-offspring nursing. Anim Behav 43:265–281
Pollock HS, Hoover JP, Uy FMK, Hauber ME (2021) Brood parasites are a heterogeneous and functionally distinct class of natural enemies. Trends Parasitol 37:588–596
R Development Core Team (2022) R: A language and environment for statistical computing (R Foundation for Statistical)
Ranjard L, Anderson MG, Rayner MJ et al (2010) Bioacoustic distances between the begging calls of brood parasites and their host species: a comparison of metrics and techniques. Behav Ecol Sociobiol 64:1915–1926
Ridley AR, Nelson-Flower MJ, Wiley EM et al (2022) Kidnapping intergroup young: an alternative strategy to maintain group size in the group-living pied babbler (Turdoides bicolor). Philos Trans R Soc B Biol Sci 377:20210153
Riedman ML (1982) The evolution of alloparental care and adoption in mammals and birds. Q Rev Biol 57:405–435
Riehl C (2011) Living with strangers: direct benefits favour non-kin cooperation in a communally nesting bird. Proc R Soc B Biol Sci 278:1728–1735
Riehl C (2013) Evolutionary routes to non-kin cooperative breeding in birds. Proc R Soc B 280:1–7
Royle NJ, Smiseth PT, Kölliker M (2012) The evolution of parental care. Oxford University Press
Sato NJ, Tokue K, Noske RA et al (2010) Evicting cuckoo nestlings from the nest: a new anti-parasitism behaviour. Biol Lett 6:67–69
Scharf HM, Hauber ME, Stenstrom KH, Schelsky WM (2022) Should i stay or should i go: the effect of avian brood parasitism on host fledging dynamics. Behav Ecol Sociobiol 76:64
Searcy WA (1986) Are female red-winged blackbirds territorial? Anim Behav 34:1381–1391
Searcy WA, Yasukawa K (1995) Polygyny and sexual selection in red-winged blackbirds. Princeton University Press, Princeton, New Jersey
Seddon LM, Nudds TD (1994) The costs of raising nidifugous offspring: brood rearing by giant Canada geese (Branta canadensis maxima). Can J Zool 72:533–540
Shah SS, Rubenstein DR (2017) Group augmentation underlies the evolution of complex sociality in the face of environmental. Proc Natl Acad Sci 120:e2212211120
Shizuka D, Lyon BE (2010) Coots use hatch order to learn to recognize and reject conspecific brood parasitic chicks. Nature 463:223–226
Shy MM (2010) Interspecific feeding among birds: a review. J F Ornithol 53:370–393
Spottiswoode CN, Kilner RM, Davies NB (2012) Chapter 13: Brood Parasitism. The evolution of parental care. Oxford University Press, Oxford, pp 226–243
Stoddard PK, Beecher MD (1983) Parental recognition of offspring in the cliff swallow. Auk 100:795–799
Stoddard MC, Hauber ME (2017) Colour, vision and coevolution in avian brood parasitism. Philos Trans R Soc B Biol Sci 372:1–11
Šulc M, Štětková G, Procházka P et al (2020) Caught on camera: circumstantial evidence for fatal mobbing of an avian brood parasite by a host. J Vertebr Biol 69:1–6
Welbergen JA, Davies NB (2011) A parasite in wolf’s clothing: hawk mimicry reduces mobbing of cuckoos by hosts. Behav Ecol 22:574–579
Acknowledgements
We are grateful for the permissions to work at our study site on vertebrate animals. We thank the University of Illinois Urbana-Champaign, Illinois Department of Natural Resources, the Champaign County Forest Preserve District, and the Vermilion County Conservation District for allowing us to conduct our research on their parks. We also thank the referees and the editor of the journal for providing valuable feedback on earlier drafts.
Funding
Funding was provided by internal SIB/PEEC grants (Harley J. Van Cleave Research Award, and the Program in Ecology, Evolution, and Conservation Biology Summer Research Award) from the University of Illinois Urbana-Champaign to FF-D. Additional support was provided by a National Science Foundation grant (#1953226# to MEH).
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FFD and SLL conducted the field work and recorded the field observations. FFD, EYH, and SC recorded the behavioral data from the videos. FFD and MEH analyzed the data. All authors contributed to writing the manuscript.
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All research was approved by the IACUC of the University of Illinois Urbana-Champaign and conducted under the Protocol #22007.
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Fernandez-Duque, F., Huerta, E.Y., Lawson, S.L. et al. Nest Integration: a novel form of food acquisition by altricial fledglings. Evol Ecol 37, 859–869 (2023). https://doi.org/10.1007/s10682-023-10250-8
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DOI: https://doi.org/10.1007/s10682-023-10250-8