Strategies of space use and foraging effort in two Neotropical flocking warbler species (Parulidae) during their nonbreeding period
DOI:
https://doi.org/10.22201/ib.20078706e.2026.97.5753Keywords:
Competition, Golden-cheeked warbler, Mixed-species bird flocks, NeotropicsAbstract
We explored habitat use and foraging effort in 2 migratory species, the Golden-cheeked warbler and the Townsend’s warbler (Parulidae) to identify wintering habitats and foraging costs associated to group behavior. We compared densities, sex/age ratios and use of foraging substrates within and between species as a function of habitat conditions in 2 landscapes of varying forest cover. Then, we estimated home range and proxies of foraging effort to identify potential fitness costs of group behavior. Densities along with sex and age ratios of the 2 focal species differed between higher and lower quality sites, with adult males of both species being more abundant in flocks. Higher frequency in use of inner branches and trunks and lower stratums such as the understory at the micro-site level was observed in the Townsend’s warbler, whereas use of outermost leaves, twigs and upper stratums such as canopy were observed in the Golden-heeked warbler. Home ranges of individual Townsend’s warblers overlapped areas used by flocks suggesting potential benefits of retaining a territory in areas where flocks occur. Traveled distances, daily used areas and searching times were higher in lower-quality sites for both species with increased group size as an underlying variable influencing cumulative searching times.
References
Brown, D. R., & Sherry, T. W. (2008). Alternative strategies of space use and response to resource change in a wintering migrant songbird. Behavioral Ecology, 19, 1314 –1325. https://doi.org/10.1093/beheco/arn073
BirdLife International. (2020). Species factsheet: Golden-cheeked warbler Setophaga chrysoparia. Downloaded July 28, 2025, from https://datazone.birdlife.org/species/factsheet/golden–cheeked–warbler–setophaga–chrysoparia
Brown, J. L. (1969). Territorial behavior and population regulation in birds: a review & re-evaluation. The Wilson Bulletin, 81, 293–329.
Brown, D. R., & Sherry, T. W. (2008). Alternative strategies of space use and response to resource change in a wintering migrant songbird. Behavioral Ecology, 19, 1314–1325. https://doi.org/10.1093/beheco/arn073
Darrah, A., & Smith, K. (2014). Ecological and behavioral correlates of individual flocking propensity of a tropical bird. Behavioral Ecology, 25, 1064–1072. https://doi.org/10.1093/beheco/aru086
Dolby, A. S., & Grubb, T. C. (1998). Benefits to satellite members in mixed species foraging groups: an experimental analysis. Animal Behaviour, 56, 501–509. https://doi.org/10.1006/anbe.1998.0808
Elgar, M. A. (1989). Predator vigilance & group size in mammals and birds: a critical review of the empirical evidence. Biological Reviews, 64, 13–33. https://doi.org/10.1111/j.1469–185X.1989.tb00636.x
Gentry, R. K. E., Roche, D. P., Mugel, S. G., Lancaster, N. D., Sieving, K. E., Freeberg, T. M. et al. (2022). Social dynamics of core members in mixed-species bird flocks change across a gradient of foraging habitat quality. Plos One, 17, e0262385. https://doi.org/10.1371/journal.pone.0262385
Gil, M. A. Z., Emberts, H. J., & St. Mary, C. M. (2017). Social information on fear and food drives animal grouping & fitness. American Naturalist, 189, 227–241. https://doi.org/10.1086/690055
Goodale, E., Beauchamp, G., Magrath, R. D., Nieh, J. C., & Ruxton, G. D. (2010). Interspecific information transfer influences animal community structure. Trends in Ecology & Evolution, 25, 354–361. https://doi.org/10.1016/j.tree.2010.01.002
Goodale, E., Sridhar, H., Sieving, K. E., Bangal, P., Colorado, Z., & Farine, D. R. (2020). Mixed company: a framework for understanding the composition and organization of mixed-species animal groups. Biological Reviews, 95, 889–910. https://doi.org/10.1111/brv.12591
Greenberg, R., Gonzáles, C. E., Bichier, P., & Reitsma, R. (2001). Nonbreeding habitat selection and foraging behavior of the Black-throated green warbler complex in southeastern Mexico. Condor, 103, 31–37. https://doi.org/10.1093/condor/103.1.31
Greenberg, R., & Salewski, V. (2005). Ecological correlates of wintering social systems in New World and Old-World migratory passerines. In R. Greenberg, & P. P. Marra (Eds.), Birds of two worlds: the ecology & evolution of migration (pp. 336–358). Baltimore, MD: John Hopkins University Press.
Hamilton, G. (1971). Geometry of the selfish herd. Journal of Theoretical Biology, 31, 295–311. https://doi.org/10.1016/0022–5193(71)90189–5
Holmes, R. T., Sherry, T. W., & Reitsma, L. (1989). Population structure, territoriality, and overwinter survival of two migrant warbler species in Jamaica. Condor, 91, 545–561. https://doi.org/10.2307/1368105
Holt, R., & Lawton, J. (1993). The ecological consequences of shared natural enemies. Annual Review of Ecology & Systematics, 25, 495–520. https://doi.org/10.1146/annurev.ecolsys.25.1.495
Hutto, R. L. (1988). Foraging behavior patterns suggest a possible cost associated with participation in mixed-species bird flocks. Oikos, 51, 79–83. https://doi.org/10.2307/3565809
Janzen, D. H. (1980). Heterogeneity of potential food abundance for tropical small birds. In A. Keast, & E. S. Morton (Eds.), Migrant birds in the Neotropics: ecology, behavior, distribution, & conservation (pp. 545–556). Washington, D.C.: Smithsonian Institute Press.
Jullien, M., & Clobert, J. (2000). The survival value of flocking in neotropical birds: reality or fiction? Ecology, 81, 3416–3430. https://doi.org/10.1890/0012–9658(2000)081[3416:TSVOFI]2.0.CO;2
Kent, C. H., Huh, K., Chieko, S., Judson, K., Powell, L., & Sherry, T. (2022). High resource overlap and small dietary differences are widespread in food-limited warbler (Parulidae) communities. Ibis, 164, 44–59. https://doi.org/10.1111/ibi.13006
Kershaw, J. A. Jr., Ducey, M. J., Beers, T. W., & Husch, B. (2017). Forest mensuration, Fifth Edition. John Wiley & Sons. https://doi.org/10.1002/9781118902028
King, D. I., Chandler, C. C., Rappole, J. H., Chandler, R. B., & Mehlman, D. W. (2012). Establishing quantitative habitat targets for a "Critically endangered" neotropical migrant (Golden-cheeked warbler: Dendroica chrysoparia) during the nonbreeding season. Bird Conservation International, 22, 213–221. https://doi.org/10.1017/S095927091100027X
King, D., & Rappole, J. (2000). Winter flocking of insectivorous birds in montane pine-oak forests in middle America. Condor, 102, 664–672. https://doi.org/10.1093/condor/102.3.664
Krause, G. D. (1994). Differential fitness returns in relation to spatial position in groups. Biological Reviews, 69, 187–206. https://doi.org/10.1111/j.1469–185X.1994.tb01505.x
Krause, G. D., & Ruxton, J. (2002). Living in groups. Oxford, U.K.: Oxford University Press. https://doi.org/10.1093/oso/9780198508175.001.0001
Krebs, J. R. (1973). Social learning and the significance of mixed-species flocks of chickadees (Parus spp.). Canadian Journal of Zoology, 51, 1275–1288. https://doi.org/10.1139/z73–181
Komar, O., Smith, C., & Wolf, C. (2011). Winter ecology, relative abundance and population monitoring of Golden-cheeked warblers (Dendroica chrysoparia) throughout the known & potential winter range. Technical Report. Texas Parks & Wildlife Department. MOA E69 #172003.
Laundré, J. W., Hernandez, L., & Ripple, W. J. (2010). The landscape of fear: ecological implications of being afraid. Open Ecology Journal, 3, 1–7. https://doi.org/10.2174/1874213001003030001
Lima, S. L., & Dill, L. M. (1990). Behavioral decisions made under the risk of predation: a review and prospectus. Canadian Journal of Zoology, 68, 619–640. https://doi.org/10.1139/z90–092
Losin, N., Drury, J. P., Peiman, K. S., Storch, C., & Grether, G. F. (2016). The ecological and evolutionary stability of interspecific territoriality. Ecology Letters, 19, 260–267. https://doi.org/10.1111/ele.12561
MacArthur, R. H. (1958). Population ecology of some warblers of northeastern coniferous forests. Ecology, 39, 599– 619. https://doi.org/10.2307/1931600
Maldonado-Coelho, M., & Marini, M. A. (2004). Mixed-species bird flocks from Brazilian Atlantic forests: the effects of forest fragmentation and seasonality on their size, richness and stability. Biological Conservation, 116, 19–26. https://doi.org/10.1016/S0006–3207(03)00169–1
MacArthur, R. H. (1958). Population ecology of some warblers of northeastern coniferous forests. Ecology, 39, 599–619. https://doi.org/10.2307/1931600
Martínez, A., & Robinson, S. (2016). Using foraging ecology to elucidate the role of species interactions in two contrasting mixed-species flock systems in northeastern Peru. Wilson Journal of Ornithology, 128, 378–390. https://doi.org/10.1676/wils–128–02–378–390.1
Marra, P. (2000). The role of behavioral dominance in structuring patterns of habitat occupancy in a migrant bird during the nonbreeding period. Behavioral Ecology, 11, 299–308. https://doi.org/10.1093/beheco/11.3.299
Marra, P., Studds, C., Wilson, S., Sillet, T., Sherry, T., & Holmes, R. (2015). Nonbreeding period habitat quality mediates the strength of density-dependence for a migratory songbird. Proceedings of the Royal Society B, 282, 20150624. https://doi.org/10.1098/rspb.2015.0624
McNamara, J. M., & Houston, A. I. (1992). Evolutionarily stable levels of vigilance as a function of group size. Animal Behaviour, 43, 641–658. https://doi.org/10.1016/S0003–3472(05)81023–1
Mokross, K., Potts, J., Rutt, C., & Stouffer, P. (2018). What can mixed-species flock movement tell us about the value of Amazonian secondary forests? Insights from spatial behavior. Biotropica, 50, 664–663. https://doi.org/10.1111/btp.12557
Morris, D. W. (1988). Habitat-dependent population regulation and community structure. Evolutionary Ecology, 2, 253–269. https://doi.org/10.1007/BF02214286
Munn, C. A., & Terborgh, J. W. (1979). Multi-species territoriality in neotropical foraging flocks. Condor, 81, 338–347. https://doi.org/10.2307/1366956
Nadkarni, N., & Matelson, T. (1989). Bird use of epiphyte resources in Neotropical trees. Condor, 91, 891–907. https://doi.org/10.2307/1368074
Newell, F. L., Beachy, T. A., Rodewald, A. D., Rengifo, C., Ausprey, I., & Rodewald, P. (2014). Foraging behavior of migrant warblers in mixed-species flocks in Venezuelan shade coffee: interspecific differences, tree species selection, and effects of drought. Journal of Field Ornithology, 85, 134–151. https://doi.org/10.1111/jofo.12056
Norris, R. (2005). Carry-over effects and habitat quality in migratory populations. Oikos, 109, 178–186. https://doi.org/10.1111/j.0030–1299.2005.13671.x
Pomara, L., Cooper, R. J., & Petit, L. (2007). Modelling the flocking propensity of passerine birds in two Neotropical habitats. Oecologia, 153, 121–133. https://doi.org/10.1007/s00442–007–0701–7
Pollard, K. A., & Blumstein, D. T. (2008). Time allocation and the evolution of group size. Animal Behaviour, 76, 1683–1699. https://doi.org/10.1016/j.anbehav.2008.08.006
Powell, G. V. N. (1985). Sociobiology and adaptive significance of interspecific foraging flocks in the Neotropics. Ornithological Monographs, 36, 713–732. https://doi.org/10.2307/40168313
Pulliam, H. R., & Danielson, B. J. (1991). Sources, sinks, and habitat selection: a landscape perspective on population dynamics. American Naturalist, 137, S50–S66. https://doi.org/10.1086/285139
Ramírez-Marcial, N., González-Espinosa, M., & Williams-Linera, G. (2001). Anthropogenic disturbance and tree diversity in Montane Rain Forests in Chiapas, Mexico. Forest Ecology & Management, 154, 311–326. https://doi.org/10.1016/S0378–1127(00)00639–3
Rappole, J. H., King, D. I., & Leimgruber, P. (2000). Winter habitat and distribution of the endangered Golden-cheeked warbler (Dendroica chrysoparia). Animal Conservation, 2, 45–49. https://doi.org/10.1017/S1367943000000731
Semarnat (Secretaría del Medio Ambiente y Recursos Naturales). (2010). Norma Oficial Mexicana NOM-059-SEMARNAT-2010, Protección ambiental - Especies nativas de México de flora y fauna silvestres - Categorías de riesgo y especificaciones para su inclusión, exclusión o cambio - Lista de especies en riesgo. Diario Oficial de la Federación. 30 de diciembre de 2010, Segunda Sección, México.
Sih, A. (1980). Optimal behavior: can foragers balance two conflicting demands? Science, 210, 1041–1043. https://doi.org/10.1126/science.210.4473.1041
Smith, J. A., Reitsma, L. R., & Marra, P. P. (2010). Moisture as a determinant of habitat quality for a nonbreeding Neotropical migratory songbird. Ecology, 91, 2874–2882. https://doi.org/10.1890/09–2212.1
Smith, S. (1976). Ecological aspects of dominance hierarchies in black-capped chickadees. Auk, 93, 95–107.
Sodhi, N., & Paszkowski, A. (1995). Habitat use and foraging behavior of four parulid warblers in a secondary forest. Journal of Field Ornithology, 66, 277–288.
Steiniger, S., & Hunter, A. (2012). OpenJUMP HoRAE–A free GIS and toolbox for home-range analysis. Wildlife Society Bulletin, 36, 600–608. https://doi.org/10.1002/wsb.168
Steven, A., Wolfe, J., Kellerman, J., Sherry, T., Stutchbury, B., Bayly, N. et al. (2020). Habitat ecology of Nearctic-Neotropical migratory landbirds on the nonbreeding grounds. Condor, 122, 1–18. https://doi.org/10.1093/condor/duaa055
Stutchbury, B. J. (1994). Competition for winter territories in a Neotropical migrant: the role of sex and age. Auk, 111, 63–69. https://doi.org/10.2307/4088505
Sridhar, H., Beauchamp, G., & Shanker, K. (2009). Why do birds participate in mixed-species foraging flocks? A large-scale synthesis. Animal Behaviour, 78, 337–347. https://doi.org/10.1016/j.anbehav.2009.05.008
Stephens, D. W., & Krebs, J. R. (1986). Foraging theory. Princeton, N.J.: Princeton University Press.
Suhonen, J., Halonen, M., & Mappes, T. (1993). Predation risk and the organization of the Parus guild. Oikos, 66, 94–100. https://doi.org/10.2307/3545200
Telleria, J. L., Virgos, E., Carbonell, R., Pérez-Tris, J., & Santos, T. (2001). Behavioral responses to changing landscapes: flock structure and anti–predator strategies of tits wintering in fragmented forests. Oikos, 95, 253–264. https://doi.org/10.1034/j.1600–0706.2001.950207.x
Terborgh, J. (1990). Mixed-species flocks and polyspecific associations: costs and benefits of mixed groups to birds & monkeys. American Journal of Primatology, 21, 87–100.
https://doi.org/10.1002/ajp.1350210203
Thomson, R. L., Forsman, J. T., & Mönkkönen, M. (2003). Positive interactions between migrant and resident birds: testing the heterospecific attraction hypothesis. Oikos, 134, 431–438. https://doi.org/10.1007/s00442–002–1140–0
Valone, T. J. (2007). From eavesdropping on performance to copying the behavior of others: a review of public information use. Behavioral Ecology & Sociobiology, 62, 1–14. https://doi.org/10.1007/s00265–007–0439–6
Vidal, R. M., Macías-Caballero, C., & Duncan, C. (1994). The occurrence and ecology of the golden-cheeked warbler in the highlands of northern Chiapas, Mexico. Condor, 96, 684–691. https://doi.org/10.2307/1369471
Wahl, R., Diamond, D. D., & Shaw, D. (1990). The Golden-cheeked warbler: astatus review. Fort Worth, TX: Ecological Services, U.S. Fish & Wildlife Service.
Whittingham, M., & Evans, K. (2004). The effects of habitat structure on predation risk of birds in agricultural landscapes. Ibis, 146, 210–220. https://doi.org/10.1111/j.1474–919X.2004.00370.x
Winker, K. (1998). The concept of floater. Ornitologia Neotropical, 9, 111–119.
Worton, B. J. (1989). Kernel methods for estimating the utilization distribution in home-range studies. Ecology, 70, 164–8. https://doi.org/10.2307/1938423
Wright, A. L., Hayward, G. D., Matsuoka, S. M., & Hayward, P. H. (1998). Townsend's warbler (Setophaga townsendi). In P. G. Rodewald (Ed.), The birds of North America. Ithaca, N.Y.: Cornell Lab of Ornithology. https://doi.org/10.2173/tbna.333.p
