Rainy cycles in South America as a driver for the breeding of the Black Skimmer (Rynchops niger) and the Large-billed Tern (Phaetusa simplex) (Aves, Charadriiformes)





Biogeography, Waterbirds, South American rivers, Migration, Rynchops, Phaetusa


The Black Skimmer (Rynchops niger) and the Large-billed Tern (Phaetusa simplex) are two migratory waterbirds that breed simultaneously on many river beaches in South America. Both are polytypic taxa with little information about the distribution and nonbreeding (“wintering”) areas. Based on data from the literature, citizen science websites, fieldwork, and specimens housed in natural history museums, we revised the distribution of some of the main breeding colonies in South America, comparing it with continental rainy cycles to identify generalities about the role of precipitation seasonality on the defining intratropical migration routes of these species. Our data suggest that the seasonal precipitation cycle of South America directly influences the reproductive timing and distribution of both species, which is largely circumscribed by South America’s rivers. After breeding on sandy beaches during the dry season, both species disperse in small groups or even individually – not in large flocks as seen in breeding areas – making it difficult to find general migration patterns during the rainy season. Nonetheless, individuals of both species tend to follow the course of the largest rivers of the continent and even alternative routes to disperse into several areas throughout South America during the nonbreeding season.


Download data is not yet available.


Antas, P.D.T.Z. 1994. Migration and other movements among the lower Paraná River valley wetlands, Argentina, and the south Brazil/Pantanal wetlands. Bird Conservation International, 4(2-3): 181-190.

Antas, P.D.T.Z.; Carrara, L.A.; Ubaid, F.K.; Oliveira-Junior, S.B. & Ferreira L.P. 2016. As Aves Coloniais da Reserva Particular do Patrimônio Natural SESC Pantanal. Rio de Janeiro. 236p.

Barbieri, E. 2007. Variação sazonal e abundância de Rynchops niger no estuário de Cananéia-Iguape-Ilha Comprida, São Paulo. Biota Neotropica, 7(2): 21-26.

Belton, W. 1994. Aves do Rio Grande do Sul. Distribuição e biologia. São Leopoldo, Editora Unisinos. 584p.

Bisbal, G.A. 1995. The Southeast South American shelf large marine ecosystem. Evolution and components. Marine Policy, 19(1): 21-38.

Burger, J. & Gochfeld, M. 1994. Predation and effects of human on island-nesting seabirds. In: Nettleship, D.N.; Burger, J. & Gochfeld, M. (Eds.). Seabirds on islands: threats, case studies and action plans. XX Proceedings of the Seabird Specialist Group. New Zealand. p. 39-67. (Birdlife International Series, 1).

Burger, J. & Gochfeld, M. 1996. Family Laridae (Gulls). In: Del Hoyo, J.; Elliott, A. & Sargatal, J. (Eds.). Handbook of the Birds of the World. Barcelona, Lynx Edicion, v. 3, p. 572-623.

Chesser, R.T. 2010. Migration in South America: An overview of the austral system. Bird Conservation International, 4(2-3): 91-107.

Coburn, L.M.; Cobb, D.T. & Gore, J.A. 2001. Management opportunities and techniques for roof- and ground-nesting black skimmers. Wildlife Society Bulletin, 29(1): 342-348.

Das, D.K. 2015. Breeding status of Indian Skimmer Rynchops albicollis in the National Chambal Sanctuary, India. Indian Birds, 10(2): 53-54.

Davenport, L.C.; Goodenough, K.S. & Haugaasen, T. 2016. Birds of two oceans? Trans-Andean and divergent migration of black skimmers (Rynchops niger cinerascens) from the Peruvian Amazon. PLoS One, 11(1): 1-14.

Davenport, L.C.; Nole-Bazán, I. & Carlos-Erazo, N. 2012. East with the Night: Longitudinal Migration of the Orinoco Goose (Neochen jubata) Between Manu National Park, Perú and the Llanos de Moxos, Bolivia. PLoS One, 7: 1-7.

Demey, R.; Lack, P. & Webb, R. 2007. Africa Round-up: Protection for African migrants. Bulletin African Bird Club, 14(1): 10.

Dilawar, M. & Sharma, V. 2016. A new breeding location of Indian Skimmer Rynchops albicollis, and notes on other birds in Son Gharial Wildlife Sanctuary, Madhya Pradesh, India. Indian Birds, 11(2): 35-38.

Duffy, D.C.; McKnight, A.L.Y. & David, B.I. 2013. Trans-andean passage of migrating arctic terns over Patagonia. Marine Ornithology, 41(2): 155-159.

Elphick, J. 2007. The Atlas of bird migration: tracing the great journeys of the world’s birds. London, Natural History Museum. 180p.

Faaborg, J.; Holmes, R.T.; Anders, A.D.; Bildstein, K.L.; Dugger, K.J. & Gauthreaux, S.A. 2010. Recent advances in understanding migration systems of New World land birds. Ecological Monographs, 80(1): 3-48.

Favero, M.; Bachmann, S.; Copello, S.; Mariano-Jelicich, R.; Silva, M.P.; Ghys, M.; Khatchikian, C. & Mauco, L. 2001. Aves marinas del Sudeste Bonaerense. In: Iribarne, O. (Ed.). Reserva de Biósfera Mar Chiquita: características físicas, biológicas y ecológicas. Universidad Nacional de Mar del Plata, Editorial Martin. p. 251-267.

Ffrench, R. 1991. A guide to the Birds of Trinidad and Tobago. Ithaca, Cornell University Press. 426p.

Gochfeld, M. 1979. Prevalence of oiled plumage of terns and skimmers on western Long Island, New York: baseline data prior to petroleum exploration. Environmental Pollution, 20(2): 123-130.

Gochfeld, M. & Burger, J. 1996. Family Sternidae (Terns). In: Del Hoyo, J.; Elliott, A. & Sargatal, J. (Eds.). Handbook of the Birds of the World. Barcelona, Lynx Edicion, v. 3, p. 624-667.

Gochfeld, M.; Burger, J.; Kirwan, G.M. & Garcia, E.F.J. 2018. Large-billed Tern (Phaetusa simplex). In: Del Hoyo, J.; Elliott, A.; Sargatal, J., Christie, D. & Juana, E. (Eds.). Handbook of the Birds of the World Alive. Barcelona, Lynx Edicions,. Available: www.hbw.com/node/54048. Access: 18/07/2018.

Gopi, G.V.; Jena, A.K. & Pandav, B. 2006. Bhitarkanika Wildlife Sanctuary (Orissa), a key congregation area for Indian Skimmer Rynchops albicollis. Birding Asia, 5: 78.

Gouvêa, A.C. 2018. Taxonomy and biogeography of Rynchops niger (Rynchopinae) and Phaetusa simplex (Sterninae) (Aves, Charadriiformes): using morphology and molecular markers to investigate the population structure and the role of the rivers in the evolution and migration of waterbirds. Tese de doutorado, Universidade de São Paulo, São Paulo.

Gudmundsson, A.G.; Benvenuti, S.; Alerstam, T.; Papi, F.; Lilliendahl, K. & Akesson, S. 1995. Examining the limits of flight and orientation performance: satellite tracking of brent Geese migrating across the Greenland Ice-Cap. Proceedings of The Royal Society, 261: 73-79.

Harrison, P. 1985. Seabirds: an identification guide. Boston, Houghton Mifflin Company. 448p.

Hawkes, L.A.; Balachandran, S.; Batbayar, N.; Butler, P.J.; Chua, B.; Douglas, D.C.; Frappell, P.B.; Hou, Y.; Milsom, W.K.; Newman, S.H.; Prosser, D.J.; Sathiyaselvam, P.; Scott, G.R.; Takekawa, J.Y.; Natsagdorj, T.; Wikelski, M.; Witt, M.J.; Yan, B. & Bishop, C.M. 2013. The paradox of extreme high-altitude migration in bar-headed geese Anser indicus. Proceedings of Biological sciences, 280(1750): 2012-2114.

Hawkes, L.A.; Balachandran, S.; Batbayar, N.; Butler, P.J.; Frappell, P.B.; Milsom, W.K.; Seveenmyadag, N.; Newman, S.H.; Scott, G.R.; Sathiyaselvam, P.; Takekawa, J.Y.; Wikelski, M. & Bishop, C.M. 2011. The trans-Himalayan flights of bar-headed geese (Anser indicus). Proceedings of the National Academy of Sciences, 108(23): 9516-9519.

Hughes, R.A. 1970. Notes on the birds of the mollendo district, southwest Peru. Ibis, 112(2): 229-241. International Union for Conservation of Nature and Natural Resources (IUCN). 2018. The IUCN Red List of Threatened Species. Version 2018-1. Available: www.iucnredlist.org. Access: 01/04/2018.

Jahn, A.E.; Cueto, V.R.; Fontana, C.S.; Guaraldo, A.C.; Levey, D.J.; Marra, P.P. & Ryder, T.B. 2020. Bird migration within the Neotropics. The Auk: Ornithological Advances, 137: 1-23.

Lambertucci, S.A.; Alarcón; P.A.E.; Hiraldo, F.; Anchez-Zapata, J.A.; Blanco, G. & Donázar, J.A. 2014. Apex scavenger movements call for transboundary conservation policies. Biological Conservation, 170: 156-150.

Livezey, B.C. & Zusi, R.L. 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society, 149: 1-95.

Mallach, T.J. & Leberg, P.L. 1999. Use of dredged materials as substrates by nesting terns and Black Skimmers. The Journal of Wildlife Management, 63(1): 137-146.

Mariano-Jelicich, R.; Favero, M. & Silva, M.P. 2003. Fish prey of the Black Skimmer Rynchops niger at Mar Chiquita, Buenos Aires Province, Argentina. Marine Ornithology, 31(2): 199-202.

Matsuura, K. 2017. The Climate Data Guide: Global (land) precipitation and temperature: Willmott and Matsuura. University of Delaware. National Center for Atmospheric Research. Available: www.climatedataguide.ucar.edu/climate-data/global-land-precipitation-and-temperature-willmott-matsuura-university-delaware. Access: 02/01/2017.

Miloslavich, P.; Klein, E.; Díaz, J.M.; Hernández, C.E.; Bigatti, G.; Campos, L.; Artigas, F.; Castillo, J.; Penchaszadeh, P.E.; Neill, P.E.; Carranza, A.; Retana, M.V.; Astarloa, J.M.D.; Lewis, M.; Yorio, P.; Piriz, M.L.; Rodríguez, D.; Yoneshigue-Valentin, Y.; Gamboa, L. & Martín, A. 2011. Marine Biodiversity in the Atlantic and Pacific Coasts of South America: Knowledge and Gaps. PLoS One, 6(1): 1-46.

Morris-Pacock, J.A.; Steeves, T.E.; Estela, F.A.; Anderson, D.J. & Friesen, V.L. 2010. Comparative phylogeography of brown (Sula leucogaster) and red-footed boobies (S. sula): the influence of physical barriers and habitat preference on gene flow in pelagic seabirds. Molecular Phylogenetics and Evolution, 54(3): 883-896.

Myers, J.P.; Morrison, R.I.G.; Antas, P.Z.; Harrington, B.A.; Lovejoy, T.E.; Sallaberry, M.; Senner, S.E. & Tarak, A. 1987. Conservation strategy for migratory species. American Scientist, 75(1): 19-26.

Nascimento, J.L.X.; Antas, P.T.Z.; Silva, F.M.B.V. & Scherer, S.B. 2000. Migração e dados demográficos do marrecão Netta peposaca (Anseriformes, Anatidae) no sul do Brasil, Uruguai, Paraguai e norte da Argentina. Melopsittacus, 3(4): 143-158.

Nores, M. & Yzurieta, D. 1980. Aves de ambientes acuáticos de Córdoba y centro de Argentina. Córdoba, Secretaria de Estado de Agricultura y Ganaderia. 236p.

Prance, G.T. & Goulding, M. 1981. The Fishes and the Fforest: explorations in Amazonian natural history. Brittonia, 33(2): 257.

Rappole, J. 2013. The Avian Migrant. New York, Columbia University Press. 464p.

Reboita, M.S.; Gan, M.A.; da Rocha, R.P. & Ambrizzi, T. 2010. Regimes de precipitação na América do Sul: uma revisão bibliográfica. Revista Brasileira de Meteorologia, 25(2): 185-204.

Reserva de Desenvolvimento Sustentável, Instituto Mamirauá (RDS). 2015. Aves recapturadas indicam que filhotes retornam adultos para praia onde nasceram. Available: www.mamiraua.org.br/pt-br/comunicacao/noticias/2013/12/23/aves-recapturadas-indicam-que-filhotes-retornam-adultos-para-praia-onde-nasceram. Access: 23/02/2019.

Sick, H. 1997. Ornitologia brasileira. 3 ed. Rio de Janeiro, Nova Fronteira. 862p.

Somenzari, M.; Amaral, P.P.; Cueto, V.R.; Guaraldo, A.C.; Jahn, A.E.; Lima, D.M.; Lima, P.C.; Lugarini, C.; Machado, C.C.; Martinez, J.; Nascimento, J.L.X.; Pacheco, J.F.; Paludo, D.; Prestes, N.P.; Serafini, P.P.; Silveira, L.F.; Sousa, A.E.B.A.; Sousa, N.A.; Souza, M.A.; Telino-Júnior, W.R. & Whitney, B.W. 2018. An overview of migratory birds in Brazil. Papéis Avulsos de Zoologia, 58(3): 1-66. http://doi.org/10.11606/1807-0205/2018.58.03.

Stotz, D.F.; Fitzpatrick, J.W.; Parker, T.A. & Moskovits, D.K. 1996. Neotropical birds: ecology and conservation. Chicago, University of Chicago Press. 502p.

Sullivan, B.L.; Wood, C.L.; Iliff, M.J.; Bonney, R.E.; Fink, D. & Kelling, S. 2009. Ebird: a citizen-based bird observation network in the biological sciences. Biological Conservation, 142(10): 2282-2292. Available: www.ebird.org. Access: 23/04/2018.

Taylor, S.A.; Zavalaga, C.B.; Luna-Jorquera, G.; Simeone, A.; Anderson, D.J. & Friesen, V.L. 2011. Panmixia and high genetic diversity in a Humboldt Current endemic, the Peruvian Booby (Sula variegata). Journal of Ornithology, 152(3): 623-630.

Tyler, S.J. 2004. The breeding and population status of the African Skimmer Rynchops flavirostris in Botswana. Ostrich, 75(4): 329-332.

Vieira, B.P.; Furness, R. & Nager, R.G. 2018. What Do we know about Black Skimmers? A review on its annual-cycle and life-history main events. Ardea, 106(2): 119-130.

Webster, M.S. & Marra, P.P. 2005. The importance of understanding migratory connectivity. In: Greenberg, R. & Marra, P.P. (Eds.). Birds of two worlds: the ecology and evolution of temperate-tropical migration systems. Baltimore, MD, Johns Hopkins University Press. p. 199-209.

WikiAves. 2017. Wiki Aves – A Enciclopédia das Aves do Brasil. Available: www.wikiaves.com. Access: 01/12/2017.

Zusi, R.L. 1996. Family Rhynchopidae (skimmers). In: Del Hoyo, J.; Elliott, A. & Sargatal, J. (Eds.). Handbook of the Birds of the World. Barcelona, Lynx Edicion, v. 3, p. 668-677.






Original Article

How to Cite

Gouvêa, A. C., Bravo, G. A., Antas, P. de T. Z., Schuchmann, K.-L., & Silveira, L. F. (2023). Rainy cycles in South America as a driver for the breeding of the Black Skimmer (Rynchops niger) and the Large-billed Tern (Phaetusa simplex) (Aves, Charadriiformes). Papéis Avulsos De Zoologia, 63, e202363028. https://doi.org/10.11606/1807-0205/2023.63.028

Funding data