Menu

Volume 54, No. 1

Search by author or title:

Comparative diet analysis of three larid species nesting at Isla Rasa, Gulf of California, Mexico.


Authors

ENRIQUETA VELARDE1 & LLOYD T. FINDLEY2
1Instituto de Ciencias Marinas y Pesquerías, Universidad Veracruzana, Boca del Río, Veracruz, Mexico (enriqueta_velarde@yahoo.com.mx)
2Centro de Investigación en Alimentación y Desarrollo, A. C.—Unidad Guaymas, Guaymas, Sonora, Mexico

Citation

Velarde, E., & Findley, L. T. (2026). Comparative diet analysis of three larid species nesting at Isla Rasa, Gulf of California, Mexico. Marine Ornithology, 54(1), 87-97.
http://doi.org/10.5038/2074-1235.54.1.1688

Received 09 June 2025, accepted 03 September 2025

Date Published: 2026/04/15
Date Online: 2026/04/15
Key words: comparative seabird diets, fishery impacts, Gulf of California, Isla Rasa, Larus heermanni, Thalasseus elegans, Thalasseus maximus

Abstract

Isla Rasa, situated in highly productive waters of the Gulf of California, has the largest nesting colony of Royal Terns Thalasseus maximus (as many as 17,000 individuals) in the Mexican Pacific, sharing this site with 300,000 Elegant Terns T. elegans and 260,000 Heermann's Gulls Larus heermanni. During eight breeding seasons between 1996 and 2011, we found that the Royal Tern diet contained over twice the number of prey species as the other two seabirds' diets: the Royal Tern's diet included 36 species of fishes, in 20 families, and pelagic squat lobster Grimothea planipes, whereas the Elegant Tern and Heermann's Gull preyed upon 17 taxa (15 fishes and two pelagic crustaceans), in nine families. Most fishes taken by Elegant Terns and Heermann's Gulls (93.8%) represented only three pelagic species (Pacific Sardine Sardinops sagax, Northern Anchovy Engraulis mordax, and Pacific Chub Mackerel Scomber japonicus), whereas these three species made up only 73.3% of the Royal Tern diet—a high proportion, nonetheless. The Royal Tern diet also reflected more diverse prey habitats. Only ~50% of Royal Tern prey species were pelagic, far fewer than for the other two seabirds (99%). The dietary differences among these three species appear to be key to their sharing of a crowded nesting site. The continuing regional growth of the commercial fishery for pelagic forage fish represents a threat to these seabirds and other components of the Gulf of California food web.

References


Ainley, D. (2019, June 14). How people saved the seabirds of the California Current. Bay Nature. https://baynature.org/2019/06/14/how-people-saved-the-seabirds-of-the-california-current/

Ainley, D. G., & Boekelheide, R. J. (Eds.). (1990). Seabirds of the Farallon Islands: Ecology, dynamics, and structure of an upwelling-system community. Stanford University Press.

Alvarez-Borrego, S. (1983). Gulf of California. In B. H. Ketchum (Ed.), Estuaries and enclosed seas (pp. 427-449). Elsevier.

American Ornithologists' Union. (1998). Check-list of North American birds (7th ed.).

Anderson, D. W., & Gress, F. (1984). Brown Pelicans and the anchovy fishery off Southern California. In D. N. Nettleship, G. A. Sanger, & P. F. Springer (Eds.), Marine birds: Their feeding ecology and commercial fisheries relationships (pp. 128-135). Pacific Seabird Group; Canadian Wildlife Service.

Anderson, D. W., Gress, F., & Mais, K. F. (1982). Brown pelicans: Influence of food supply on reproduction. Oikos, 39(1), 23-31. http://dx.doi.org/10.2307/3544527

Anderson, D. W., & Keith, J. O. (1980). The human influence on seabird nesting success: Conservation implications. Biological Conservation, 18(1), 65-80. https://doi.org/10.1016/0006-3207(80)90067-1

Angulo-Gastélum, U. T., Castillo-Guerrero, J. A., & Mellink, E. (2011). Breeding ecology of the Royal Tern (Thalasseus maximus) at Isla El Rancho, Mexico: Colony size and nest location affect predation. Ornitología Neotropical, 22, 131-142.

Ashmole, N. P. (1963). The regulation of numbers of tropical oceanic birds. Ibis, 103b(3), 458-473. http://dx.doi.org/10.1111/j.1474-919X.1963.tb06766.x

Ashmole, N. P. (1971). Seabird ecology and the marine environment. In D. S. Farner, & J. R. King (Eds.), Avian biology (Vol. I, pp. 223-286). Academic Press.

Ashmole, N. P., & Ashmole, M. J. (1967). Comparative feeding ecology of sea birds of a tropical oceanic island (Bulletin 24). Peabody Museum of Natural History, Yale University. https://oceanrep.geomar.de/id/eprint/53591/1/4160.pdf

Ashmole, N. P., & Tovar S., H. (1968). Prolonged parental care in Royal Terns and other birds. The Auk, 85(1), 90-100. http://dx.doi.org/10.2307/4083627

Aygen, D., & Emslie, S. D. (2006). Royal Tern (Sterna maxima) chick diet at Fisherman Island National Wildlife Refuge, Virginia. Waterbirds, 29(3), 395-400. https://doi.org/10.1675/1524-4695(2006)29[395:RTSMCD]2.0.CO;2

Barbour, B. (1978). Species of special concern, Royal Tern, Thalasseus maximus. In H. W. Kale (Ed.), Rare and endangered biota of Florida: Vol. 2. Birds (pp. 91-92). University Presses of Florida.


Bearhop, S., Phillips, R. A., McGill, R., Cherel, Y., Dawson, D. A., & Croxall, J. P. (2006). Stable isotopes indicate sex-specific and long-term individual foraging specialisation in diving seabirds. Marine Ecology Progress Series, 311, 157-164. https://www.int-res.com/articles/meps2006/311/m311p157.pdf

Bent, A. C. (1921). Life histories of North American gulls and terns: Order Longipennes (Bulletin of the United States National Museum No. 113). Smithsonian Institution; U.S. Government Printing Office. https://doi.org/10.5479/si.03629236.113.i

BirdLife International. (2018). Species factsheet: Royal Tern Thalasseus maximus. Retrieved May 13, 2025, from https://datazone.birdlife.org/species/factsheet/royal-tern-thalasseus-maximus

Boersma, P. D. (1978). Breeding patterns of Galápagos penguins as an indicator of oceanographic conditions. Science, 200(4349), 1481-1483. http://dx.doi.org/10.1126/science.200.4349.1481

Brusca, R. C., Álvarez-Borrego, S., Hastings, P. A., & Findley, L. T. (2017). Colorado River flow and biological productivity in the Northern Gulf of California, Mexico. Earth-Science Reviews, 164, 1-30. https://doi.org/10.1016/j.earscirev.2016.10.012

Cherel, Y., Hobson, K. A., & Hassani, S. (2005). Isotopic discrimination between food and blood and feathers of captive penguins: Implications for dietary studies in the wild. Physiological and Biochemical Zoology, 78(1), 106-115. http://dx.doi.org/10.1086/425202

Clapp, R. B., & Buckley, P. A. (1984). Status and conservation of seabirds in the southeastern United States. In J. P. Croxall, P. G. H. Evans, & R. W. Schreiber (Eds.), Status and conservation of the world's seabirds (ICBP Technical Publication No. 2, pp. 135-155). International Council of Bird Preservation.

Clapp, R. B., Klimkiewicz, M. K., & Kennard, J. H. (1982). Longevity records of North American birds: Gaviidae through Alcidae. Journal of Field Ornithology, 53(2), 81-124. https://digitalcommons.usf.edu/jfo/vol53/iss2/1

Croxall, J. P., & Prince, P. A. (1980). Food, feeding ecology and ecological segregation of seabirds at South Georgia. Biological Journal of the Linnean Society, 14(1), 103-131. http://dx.doi.org/10.1111/j.1095-8312.1980.tb00101.x

Decreto que declara Zona de Reserva Natural y Refugio de Aves a la Isla Rasa, Estado de Baja California (Secretaría de Agricultura y Ganadería), Diario Oficial de la Federación [DOF], CCLXIV(24), pp. 3-4, 30-05-1964 (Mex.). https://sidof.segob.gob.mx/notas/4787750

Diamond, A. W. (1975). The biology of tropicbirds at Aldabra Atoll, Indian Ocean. The Auk, 92(1), 16-39. http://dx.doi.org/10.2307/4084415

Diamond, A. W. (1983). Feeding overlap in some tropical and temperate seabird communities. In D. S. Farner & J. R. King (Eds.), Studies in avian biology (No. 8, pp. 24-46). Oxford University Press.

Dorward, D. F. (1962). Comparative biology of the White Booby and the Brown Booby Sula spp. at Ascension. Ibis, 103B(2), 174-220. https://doi.org/10.1111/j.1474-919X.1962.tb07244.x

Drinkwater, R., Goodwin, A., Cush, J., Korstian, J. M., Chumchal, M. M., Herrera M., L. G., Valdez, M., Otálora-Ardila, A., Flores-Martínez, J. J., & Clare, E. L. (2021). Molecular diet analysis of the marine fish-eating bat (Myotis vivesi) and potential mercury exposure. Canadian Journal of Zoology, 99, 752-759. https://doi.org/10.1139/cjz-2021-0018

Elias-Valdez, A., Velarde, E., Medina-Quej, A., Castro-Perez, J. M., Navarro, J., & Rosas-Luis, R. (2023). Feeding ecology of coexisting Heermann's gull (Larus heermanni) and elegant tern (Thalasseus elegans) chicks, based on stable isotope measurements. Marine Ecology Progress Series, 712, 101-111. https://doi.org/10.3354/meps14326

Erwin, R. M. (1977). Foraging and breeding adaptations to different food regimes in three seabirds: The Common Tern, Sterna hirundo, Royal Tern, Sterna maxima, and Black Skimmer, Rynchops niger. Ecology, 58, 389-397. https://doi.org/10.2307/1935613

Escalante, R. (1968). Notes on the Royal Tern in Uruguay. The Condor, 70(3), 243-247. http://dx.doi.org/10.2307/1366696

Eschmeyer, W. N., & Herald, E. S. (1983). A field guide to Pacific Coast fishes of North America. Houghton Mifflin.

Favero, M., Silva R., M. P., & Mauco, L. (2000). Diet of Royal (Thalasseus maximus) and Sandwich (T. sandvicensis) Terns during the austral winter in the Buenos Aires Province, Argentina. Ornitología Neotropical, 11(3), 259-262.

Flores-Martínez, J. J., Herrera, L. G., Arroyo-Cabrales, J., Alarcón, I., & Ruiz, E. A. (2015). Seasonal dietary differences of the Yellow-footed Gull (Charadriiformes: Laridae) in Isla Partida Norte, Gulf of California, Mexico. Revista Mexicana de Biodiversidad, 86(2), 412-418. https://doi.org/10.1016/j.rmb.2015.04.002

Froese, R., & Pauly, D. (Eds.). (2025). FishBase (Version 02/2025) [World Wide Web electronic publication]. https://www.fishbase.org

Furness, R. W., & Birkhead, T. R. (1984). Seabird colony distributions suggest competition for food supplies during the breeding season. Nature, 311, 655-656. https://doi.org/10.1038/311655a0

Furness, R. W., & Monaghan, P. (1987). Seabird Ecology. Blackie.

García, G. O., Favero, M., & Vassallo, A. I. (2010). Factors affecting kleptoparasitism by gulls in a multi-species seabird colony. The Condor, 112(3), 521-529. https://doi.org/10.1525/cond.2010.090117

García-Rodríguez, F. J., & Aurioles-Gamboa, D. (2004). Spatial and temporal variation in the diet of the California sea lion (Zalophus californianus) in the Gulf of California, Mexico. Fishery Bulletin, 102(1), 47-62.

Gatto, A. J., & Yorio, P. (2009). Provisioning of mates and chicks by Cayenne and Royal Terns: Resource partitioning in northern Patagonia, Argentina. Emu - Austral Ornithology, 109(1), 49-55. https://doi.org/10.1071/MU08025

Gochfeld, M., & Burger, J. (1996). Family Sternidae (terns). In J. del Hoyo, A. Elliott, & J. Sargatal (Eds.), Handbook of the birds of the world: Vol. 3. Hoatzin to auks (pp. 624-667). Lynx Edicions.

Grant, G. S. (1981). Belly-soaking by incubating Common, Sandwich, and Royal Terns. Journal of Field Ornithology, 52(3), Article 19. https://digitalcommons.usf.edu/jfo/vol52/iss3/19

Harris, M. P. (1969). Food as a factor controlling the breeding of Puffinus lherminieri. Ibis, 111(2), 139-156. https://doi.org/10.1111/j.1474-919X.1969.tb02523.x

Hunt, G. L., Jr. (1972). Influence of food distribution and human disturbance on the reproductive success of Herring Gulls. Ecology, 53(6), 1051-1061. http://dx.doi.org/10.2307/1935417

Islam, K., & Velarde, E. (2020). Heermann's Gull (Larus heermanni), Version 2.0. In P. G. Rodewald & B. K. Keeney (Eds.), Birds of the world. Cornell Lab of Ornithology. https://doi.org/10.2173/bow.heegul.02

Kale, H. W., II, Sciple, G. W., & Tomkins, I. R. (1965). The Royal Tern colony of Little Egg Island, Georgia. Bird Banding, 36, 21-27. http://dx.doi.org/10.2307/4511131

Kilham, L. (1981). Courtship feeding and copulation of Royal Terns. The Wilson Bulletin, 93(3), 390-391. https://digitalcommons.usf.edu/wilson_bulletin/vol93/iss3/13

Liechty, J. S., Fontenot, Q. C., & Pierce, A. R. (2016). Diet Composition of Royal Tern (Thalasseus maximus) and Sandwich Tern (Thalasseus sandvicensis) at Isles Dernieres Barrier Island Refuge, Louisiana, USA. Waterbirds, 39(1), 58-68. https://doi.org/10.1675/063.039.0107

Loftin, R. W., & Sutton, S. (1979). Ruddy Turnstones destroy Royal Tern colony. The Wilson Bulletin, 91(1), 133-135. https://digitalcommons.usf.edu/wilson_bulletin/vol91/iss1/21

Markaida, U., & Sosa-Nishizaki, O. (2003). Food and feeding habits of jumbo squid Dosidicus gigas (Cephalopoda: Ommastrephidae) from the Gulf of California, Mexico. Journal of the Marine Biological Association of the United Kingdom, 83(3), 507-522. https://doi.org/10.1017/S0025315403007434h

Martin, G. R., & Prince, P. A. (2001). Visual fields and foraging in procellariiform seabirds: Sensory aspects of dietary segregation. Brain, Behavior and Evolution, 57(1), 33-38. http://dx.doi.org/10.1159/000047224

McGinnis, T. W., & Emslie, S. D. (2001). The foraging ecology of Royal and Sandwich Terns in North Carolina, USA. Waterbirds, 24(3), 361-370. https://doi.org/10.2307/1522066

Mellink, E., Palacios, E., & Amador, E. (2007). Colonies of four species of terns and the Black Skimmer in western Mexico. Waterbirds, 30(3), 358-366. https://doi.org/10.1675/1524-4695(2007)030[0358:COFSOT]2.0.CO;2

Miller, D. J., & Lea, R. N. (1972). Guide to the coastal marine fishes of California (Fish Bulletin 157). California Department of Fish and Game. https://escholarship.org/uc/item/6s04v367

Moser, H. G., & Ahlstrom, E. H. (1996). Myctophidae: Lanternfishes. In H. G. Moser (Ed.), The early stages of fishes in the California Current region (California Cooperative Oceanic Fisheries Investigation Atlas 33, pp. 387-475). National Marine Fisheries Service, Southwest Fisheries Science Center. https://calcofi.org/downloads/publications/atlases/CalCOFI_Atlas_33.pdf

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 (Secretaría de Medio Ambiente y Recursos Naturales), Diario Oficial de la Federación [DOF] 30-12-2010 (Mex.). https://www.gob.mx/profepa/documentos/norma-oficial-mexicana-nom-059-semarnat-2010

Petalas, C., Lazarus, T., Lavoie, R. A., Elliott, K. H., & Guigueno, M. F. (2021). Foraging niche partitioning in sympatric seabird populations. Scientific Reports, 11, Article 2493. https://doi.org/10.1038/s41598-021-81583-z

Portnoy, J. W. (1978). Colonial waterbird population status and management on the north Gulf of Mexico coast. Proceedings of the Colonial Waterbird Group, 1, 38-43. http://dx.doi.org/10.2307/1520899

Robison, B. H. (1972). Distribution of the midwater fishes of the Gulf of California. Copeia, 1972(3), 448-461. https://doi.org/10.2307/1442916

Roedel, P. M. (1948). Common marine fishes of California (Fish Bulletin 68). California Department of Fish and Game. https://doi.org/10.5962/bhl.title.6875

Santamaría-del-Ángel, E., Álvarez-Borrego, S., Müller-Karger, F. E. (1994). Gulf of California biogeographic regions based on coastal zone color scanner imagery. Journal of Geophysical Research: Oceans, 99(C4), 7411−7421. https://doi.org/10.1029/93JC02154

Schaffner, F. C. (1985). Royal Tern nesting attempts in California: Isolated or significant incidents? Western Birds, 16(2), 71-80. https://digitalcommons.usf.edu/western_birds/vol16/iss2/4

Tershy, B. R., Breese, D., Meyer, G. M. (1990). Kleptoparasitism of adult and immature Brown Pelicans by Heermann's Gulls. The Condor, 92, 1076-1077.

Thomson, D. A., & McKibbin, N. (1976). Gulf of California fishwatcher's guide. Golden Puffer Press.

Toland, B., & Gilbert, T. (1987). Roof nesting by Royal Terns in Vero Beach, Florida. Florida Field Naturalist, 15, 80-82.

Tomkins, I. R. (1963). Skimmer-like behavior in the Royal and Caspian terns. The Auk, 80, 549. http://dx.doi.org/10.2307/4082865

Van Velzen, W. T., & Benedict, R. D. (1972). Recoveries of Royal Terns banded in Virginia. Part I. The Caribbean. The Raven, 43, 39-41.

Veit, R. R., Velarde, E., Horn, M. H., & Manne, L. L. (2021). Population growth and long-distance vagrancy leads to colonization of Europe by Elegant Terns Thalasseus elegans. Frontiers in Ecology and Evolution, 9, Article 725614. https://doi.org/10.3389/fevo.2021.725614

Velarde, E. (1980). [Unpublished raw field data: Behavioral observations of Heermann's Gulls kleptoparasitizing Elegant and Royal Tern's prey during chick feedings].

Velarde, E. (1992). Predation of Heermann's Gull (Larus heermanni) chicks by Yellow-footed Gulls (Larus livens) in dense and scattered nesting sites. Colonial Waterbirds, 15(1), 8-13. https://doi.org/10.2307/1521349

Velarde, E. (1999). Breeding biology of Heermann's Gulls on Isla Rasa, Gulf of California, Mexico. The Auk, 116(2), 513-519. http://dx.doi.org/10.2307/4089383

Velarde, E. (2012). [Unpublished raw field data: Behavioral observations of Heermann's and Yellow-footed Gulls predating on Elegant and Royal Tern's eggs].

Velarde, E., & Anderson, D. W. (1994). Conservation and management of seabird islands in the Gulf of California: Setbacks and successes. In D. N. Nettleship, J. Burger, & M. Gochfeld (Eds.), Seabirds on islands: Threats, case studies and action plans (pp. 229-243). BirdLife International.

Velarde, E., Cartron, J.-L. E., Drummond, H., Anderson, D. W., Rebón-Gallardo, F., Palacios, E., & Rodríguez, C. (2005). Nesting seabirds of the Gulf of California's offshore islands: Diversity, ecology and conservation. In J.-L. E. Cartron, G. Ceballos, & R. S. Felger (Eds.), Biodiversity, ecosystems, and conservation in Northern Mexico (pp. 452-470). Oxford University Press.

Velarde, E., & Ezcurra, E. (2002). Breeding dynamics of Heermann's Gulls. In T. E. Case, M. L. Cody, & E. Ezcurra (Eds.), A new island biogeography of the Sea of Cortés (pp. 313-325). Oxford University Press.

Velarde, E., & Ezcurra, E. (2018). Are seabirds' life history traits maladaptive under present oceanographic variability? The case of Heermann's Gull (Larus heermanni). The Condor, 120(2), 388-401. https://doi.org/10.1650/CONDOR-17-5.1

Velarde, E., Ezcurra, E., & Anderson, D. W. (2013). Seabird diets provide early warning of sardine fishery declines in the Gulf of California. Scientific Reports, 3, Article 1332. http://www.nature.com/srep/2013/130225/srep01332/full/srep01332.html

Velarde, E., Ezcurra, E., & Anderson, D. W. (2015). Seabird diet predicts following-season commercial catch of Gulf of California Pacific Sardine and Northern Anchovy. Journal of Marine Systems, 146, 82-88. http://dx.doi.org/10.1016/j.jmarsys.2014.08.014

Velarde, E., Ezcurra, E., Cisneros-Mata, M. A., & Lavín, M. F. (2004). Seabird ecology, El Niño anomalies, and prediction of sardine fisheries in the Gulf of California. Ecological Applications, 14(2), 607-615. http://dx.doi.org/10.1890/02-5320

Velarde, E., Ezcurra, E., Horn, M. H., & Patton, R. T. (2015). Warm oceanographic anomalies and fishing pressure drive seabird nesting north. Science Advances, 1(5), Article e1400210. https://doi.org/10.1126/sciadv.1400210

Velarde, E., Tordesillas, M. S., Vieyra, L., & Esquivel, R. (1994). Seabirds as indicators of important fish populations in the Gulf of California. California Cooperative Oceanic Fisheries Investigations Reports, 35, 137-143. https://calcofi.org/downloads/publications/calcofireports/v35/Vol_35_Velarde_etal.pdf

Vieyra, L., Velarde, E., & Ezcurra, E. (2009). Effects of parental age and food availability on the reproductive success of Heermann's Gulls in the Gulf of California. Ecology, 90(4), 1084-1094. https://doi.org/10.1890/07-2009.1

Wambach, E. J., & Emslie, S. D. (2003). Seasonal and annual variation in the diet of breeding, known-age Royal Terns in North Carolina. The Wilson Bulletin, 115(4), 448-454.

Search by author or title:

Browse previous volumes: