Seabird breeding behavior
The term seabird is used for many families of birds in several orders that spend the majority of their lives at sea. Seabirds make up some, if not all, of the families in the following orders: Procellariiformes, Sphenisciformes, Pelecaniformes, and Charadriiformes. Many seabirds remain at sea for several consecutive years at a time, without ever seeing land. Breeding is the central purpose for seabirds to visit land. The breeding period (courtship, copulation, and chick-rearing) is usually extremely protracted in many seabirds and may last over a year in some of the larger albatrosses; this is in stark contrast with passerine birds. Seabirds nest in single or mixed-species colonies of varying densities, mainly on offshore islands devoid of terrestrial predators. However, seabirds exhibit many unusual breeding behaviors during all stages of the reproductive cycle that are not extensively reported outside of the primary scientific literature.
The courtship stage of breeding is when pair bonds are formed and occurs before copulation and occasionally continues through the copulatory and chick-rearing stages of the breeding phenology. The sequence and variety of courting behaviors vary widely among species, but they typically begin with territorial defense, followed by mate-attraction displays, and selection of a nest site. Seabirds are long-lived, socially monogamous, birds that usually mate for life. This makes selecting a mate extremely important with lifelong implications for the reproductive success of both individuals in the pair.
Seabirds are one of the only avian families that include ritualized dances in their courtship. These dances are complex and can include displays and vocalizations that vary greatly between families and orders. Albatrosses are well known for their intricate mating dances. All species of albatross have some form of ritualized dance, with many species displaying very similar forms. Albatrosses’ complex visual and vocal dances are considered some of the most developed mating displays in any long-lived animal. Both members of the pair use these dances as a proxy for mate quality and it is believed to be a very important aspect of mate choice in this family. For Black-footed (Phoebastria nigripes) and Laysan Albatrosses (P. immutabilis) there are ten described parts to their mating dance which can be given in various sequences. Several parts include “billing” where one individual gently touches the others bill and “sky pointing” where the bird rises on the tips of its toes, stretches its neck and points its bill upward. In the Wandering Albatross (Diomedea exulans), sky pointing is accompanied with “sky calling” where the displaying individual spreads its wings, revealing his massive 12 foot wingspan while pointing and vocalizing skyward. The mating dance may last for several minutes. It has been noted that many albatross species dance upon reuniting with their partner every year; however, for Waved Albatross (P. irrorata), the dance is longer and more involved in new pairs, or in pairs that failed to breed the previous season.
Boobies are another group of seabirds known for their mating displays. Brown (Sula leucogaster), Red-footed (S. sula) and Blue-footed Boobies (S. nebouxii) have at least nine described parts to their mating display. Sky pointing in boobies is similar to albatrosses; in the Brown Booby, sky pointing is described as a display where the male throws his head backwards, stretches his neck out, and usually gives a whistling vocalization. Parading is a well-known display in boobies as well; in this display, one individual in the pair - usually the male - walks upright, with his tail erect, swaying in an exaggerated manner from side to side while taking small steps. In Blue-footed and Red-footed Boobies, parading also includes lifting and flaunting their brightly colored feet at their prospective partner.
Frigatebirds are known for their unusual displays and breeding system. Unlike other seabirds, frigatebirds have a lek-breeding system where displaying males aggregate in groups of up to 30 individuals with prospecting females flying overhead. However, unlike classic leks, the pair then builds a nest on the male’s display site. The male then participates fully in nest defense, incubation, and chick-rearing. The main display that male frigatebirds use to attract females is a “gular presentation” where the male inflates his bright red throat pouch, points his head upwards and opens his wings. Interestingly, it has been shown experimentally that there is no correlation between energy expended by males during courtship display and mate selection by females.
Once the pair bond is formed, courtship feeding occurs in some species. Courtship feeding is when one member of the pair presents the other with food in a ritualized way. Often the male feeds the female, but in certain species where the sex roles are reversed, the female may feed the male. Several reasons proposed as to why courtship feeding occurs is: 1) to help strengthen the pair bond 2) to reduce aggression between males and females and 3) to provide additional nutrition to the females during the egg-laying stage.
Courtship feeding is seen in many gull and tern species. In Common Terns (Sterna hirundo), courtship feeding begins right at the start of pair formation with male terns carrying a fish around the breeding colony, displaying it to prospective mates. The direct benefits hypothesis (where the female obtains some immediate benefit for copulating with the male, food in this case) may explain why courtship feeding has evolved; however, this theory has recently been disputed with the suggestion that the rate of courtship feeding is a way for females to determine the quality of their mate through the handicap principle.
Homosexual behavior has been well documented in over 500 species of non-human animals ranging from insects to lizards to mammals (reviewed in:). In birds, same-sex pairing has been shown in many families of non-passerines including vultures, ducks, and pigeons. There is also a remarkably high incidence of homosexual behavior in seabirds. Here, homosexual behavior refers to same-sex pair-formation and chick-rearing, not to same-sex copulation, for which there are very few documented examples. Almost all the examples of same-sex pairing in seabirds are of female-female pairs. Furthermore, this phenomenon doesn’t seem to be phylogenetically constrained to any specific order or family of seabirds.
There are many examples of homosexual behavior in wild gulls. In Herring Gull (Larus argentatus) populations nesting on the Great Lakes, Fitch (1980)  reported a low, yet consistent prevalence of female-female pairs. It appears that female-female Herring Gull pairs are more common in colonies with a female-biased operational sex ratio (OSR) and occasionally these homosexual pairs will remain stable for several breeding seasons. In Western Gulls (Larus occidentalis), female-female pairs are often associated with supernormal clutches (clutches of 4-6 eggs; a normal clutch for Larus gull species is 2-3 eggs) and these clutches are usually infertile. Female-female pairs have also been widely reported in wild populations of Ring-billed Gulls (Larus delawarensis). Studies of Ring-billed Gulls has shown that same sex pairs are rare (<1% of pairs in a colony) but consistent interannually and that they also lay supernormal clutches at a significantly higher rate than do heterosexual pairs. It has also been shown that these clutches of female-female pairs have significantly lower hatching and fledging success than heterosexual pairs. There is even one example of an unusual mixed female-female pair of two gull species, the Caspian (Larus cachinnans) and Yellow-legged Gull (Larus michahellis).
Female-female pairing has also been documented and studied in several tern species including Whiskered (Chlidonias hybyida), Roseate (Sterna dougalii) and Caspian Terns (Hydroprogne caspia) with similar attributes (supernormal clutches and reduced hatching/fledging success as compared to heterosexual pairs) to same-sex gull pairs. Also in the order Charadriiformes (family Chionididae), there has been one reported occurrence of a female-female pair in the Black-faced Sheathbill (Chionis minor), but eggs in the clutch proved to be inviable.
Same-sex pairing has also been shown in several families of true seabirds including the petrels and shearwaters. Antarctic Petrels (Thalassoica antarctica) have been shown to form female-female pairs in colonies where there is a surplus of females; it is hypothesized that “pairing” with another female may be a favorable strategy for some females because it allows them to become established in the colony. The experience with a site gained through forming a female-female pair may greatly improve the chances of future successful breeding for the non-genetic parent, which explains why it might be worth the short-term cost of raising another bird’s offspring. In another member of this family, the Cory’s Shearwater (Calonectris diomedea), same-sex pairing was recently discovered for the first time in a burrow-nesting seabird. This study proposed that similar factors cause female-female pairs to form in burrow-nesting seabirds as in surface-nesting seabirds (a female-biased OSR), and that female-female pairing in burrow-nesting seabirds might have remained undetected for so long due to the secretive nature of these animals.
In albatrosses, female-female pairing has recently received major press coverage. Last year, when a Southern Royal Albatross (D. epomophora) couple hatched a chick in New Zealand, it represented the first record of a successful same-sex pair in this species. In a landmark study by Young et al. (2008), she reported over 30% of Laysan Albatrosses in a colony in Oahu, Hawaii were same sex pairs. Even though these female-female pairs had less reproductive success than heterosexual pairs, it was better than not breeding at all. Young et al. (2008) also cited a female-biased OSR as the primary reason for such a high proportion of same-sex pairs. Additionally, an unsuccessful female-female pair of highly endangered of Short-tailed Albatrosses (P. albatrus) has been documented on Kure Atoll, Hawaii.
Penguins represent the only known examples of male-male pairings in seabirds. On the Otago Peninsula of New Zealand, two-male Yellow-eyed Penguins (Megadyptes antipodes) were reported incubating an egg in 2009 In captivity, Chinstrap (Pygoscelis antarcticus), Humboldt (Spheniscus humboldti), Magellanic (S. magellanicus), and African Black-footed Penguins (S. demersus) have all been documented to form male-male pairs.
In seabirds, the copulatory stage usually occurs after, and occasionally concurrently, with the formation of the pair bond. Copulation occurs mainly on land at the breeding colony. Usually the pair copulates several times, even in orders that lay only one egg per-clutch. These additional copulations are thought of as a mechanism to strengthen the pair bond. This is important for strongly monogamous, long-lived organisms and is especially important in seabirds that spend most of the non-breeding season apart on the open ocean.
Birds are one of the only major taxa where monogamy is the dominant mating system. Prior to the advent of genetic techniques, it was assumed that the majority of monogamous birds remained faithful to their partners. However, it is now known that extra-pair copulations (EPCs), extra-pair fertilizations (EPFs), and extra-pair paternity (the raising of another’s offspring, EPP) are actually quite common in a variety of avian orders and families. Roughly 70% of birds that used to be considered genetically monogamous actually engage in EPCs and raise extra-pair young (reviewed by:). Furthermore, it has been proposed that birds that nest in high densities, as seabirds do in breeding colonies, have higher rates of EPCs and EPFs than birds that do not nest colonially. Despite this, Westneat and Sherman (1997)  found no significant correlation between nesting density and EPFs in a meta-analysis. Many seabird species raise only one chick per breeding season, which would make the prevalence of EPFs and EPP in seabirds surprising due to the fact that the male’s entire breeding success for a year is dependent on the lone egg/chick he is raising to be his genetic offspring. Moreover, all seabirds have obligate biparental care, so it would be evolutionarily costly for the male to spend months of effort raising a chick that is not his genetic offspring.
In line with this prediction, many studies of seabirds have revealed no EPCs or EPFs. Several genetic studies of Storm-Petrels, show no evidence of EPCs or EPFs, which is not surprising considering these are burrow-nesting seabirds that lay only one egg per year. Dovekie (Alle alle), a surface nesting alcid that raises one chick per year, has shown no EPFs or EPP. Nazca Boobies (Sula granti) have been well studied at breeding colonies in the Galapagos for decades and also show no evidence of EPCs or EPFs; also not a surprising result since they only have one surviving offspring per year. Chinstrap Penguins, which raise two chicks annually, have also shown no EPCs or EPFs.
Contrary to these empirical results, there has been a multitude of studies where EPCs or EPFs have been found in seabirds. Perhaps the most surprising EPFs have been found in the Procellariiforms because all members of this order only lay one egg per year and some do not even breed every year. Waved Albatross show high rates (up to 25% of offspring are extra-pair young) of both EPCs and EPFs and behavioral observations have shown that many of the EPCs are forced by the extra-pair male. Studies of Wandering Albatross have shown over 10% of chicks are extra-pair young; an extremely surprising result since adult Wandering Albatrosses only breed once every other year at most. Similar results were seen in Black-browed (Thalassarche melanophris) and Grey-headed Albatrosses (T. chrysostoma) nesting on South Georgia Island. EPFs have also been shown in Antarctic Petrels. Perhaps the most unexpected result was when EPCs and EPFs were documented in a burrow-nesting Procellariid, the Short-tailed Shearwater (Puffinus tenuirostris), because the female would have to leave her burrow to solicit an EPC.
EPCs and EPFs have also been demonstrated to occur in other families of seabirds. In contrast to the results found in genetic studies of Dovekie, EPP has been shown in several species of alcid including Common Murre (Uria aalge) and Razorbill (Alca torda), both of which raise only one chick per year. Interestingly, it has been shown that female Razorbills can determine whether or not an EPC leads to an EPF and only accept extra-pair sperm when it gives them a fitness advantage over their current mate. A low-rate of EPP has also been shown in the sexually dimorphic Great Frigatebird (Fregata minor).
Mating with related individuals is rare in naturally occurring populations of birds due to the production of lower quality offspring suffering from the genetic effects of inbreeding depression. Seabirds have an inherently high risk of inbreeding because most are natally philopatric, and many are highly endangered with some species’ entire populations breeding on one small island. Despite this, inbreeding was observed no more than expected by random chance in the Wandering Albatross and the critically endangered Amsterdam Albatross (D. amsterdamensis). In contrast, some studies of seabirds have shown evidence of inbreeding. Huyvaert and Parker (2010) detected low frequencies of inbreeding in Waved Albatrosses and genetic similarity was negatively related to EPFs, which is an unusual result that does not support the inbreeding avoidance hypothesis. Close inbreeding was observed at low frequencies in the Cory’s Shearwater where two mother-son pairs were reported.
Chick-rearing is the most crucial stage of the reproductive cycle in determining final reproductive success during a breeding season. Chick-rearing includes brooding, feeding, defending, and in some cases, teaching the chick skills it will need to know to survive independently. Chick-rearing can be totally absent in some birds (the Brush-Turkeys of southeast Asia), to a couple weeks long in many passerines, to several months long in larger birds. Seabirds, along with some Australian and Southern African landbirds such as the Southern Ground Hornbill or White-winged Chough, have the longest chick-rearing stage of any bird on earth. It is not unusual for many seabirds to spend 3–4 months raising their chicks until they are able to fledge and forage independently. In the great albatrosses, chick-rearing can take over 9 months. It is because of this extremely protracted chick-rearing stage that many of the larger procellariiform seabirds can breed only once every other year.
Siblicide, the death of an individual due to the actions of members of its own clutch, is seen in several avian orders including egrets and kingfishers, some raptors, and grackles. In most of these examples, siblicide is facultative (i.e. not obligate) and only occurs when there is a shortage of food. However in some seabirds, siblicide proves to be obligate and occurs no matter how productive the breeding season is. The Nazca Booby is one species that practices obligate siblicide. The parents lay two eggs, several days apart. The second egg laid is seen as fertility insurance if the first egg is inviable. If both eggs hatch, the elder chick will push its sibling out of the nest area, leaving it to die of thirst or cold. The parent booby will not intervene and the younger chick will inevitably die. Research has shown that high hormone levels in Nazca Booby chicks are responsible for inciting their murderous behavior. Facultative siblicide is seen in the closely related Blue-footed Booby. Unlike the Nazca Booby, Blue-footed Boobies chicks only perform siblicide when food is scarce. Furthermore, the parents actually try to suppress the siblicidal behavior, rather than ignoring or encouraging it.
For most seabirds if breeding is successful they will continue breeding with the same partner year after year until one member of the pair dies or doesn’t return to the breeding colony. However, these pair bonds occasionally dissolve or are forced apart while both members of the pair are still alive, a process known as divorce. Reasons for divorce in seabirds are wide ranging and include asynchronous arrival of mates to the breeding colony, declining reproductive success of the pair, and competition for mates. Coercive divorce is seen in the Nazca Booby and Common Murre where one member of the pair actively deserts the other or where an intruder enters and forcibly splits the breeding pair to form a new pair.
Divorce is relatively common in gulls and their relatives (reviewed by:); in one study, Black-legged Kittiwakes (Rissa tridactyla) proved to be more faithful to their nesting site than their partner. Divorce rates are surprisingly high (>80% of pairs annually) in King (Aptenodytes patagonicus) and Emperor Penguins (A. forsteri). Asynchronous arrival of mates at the breeding colony is cited as the main reason for this because these penguins have extreme time constraints on their breeding. In Great Skuas (Stercorarius skua) divorce occurs annually, but at low frequencies (6-7% of pairs annually) and death is responsible for approximately three times more pair interruptions than divorce. Divorce is uncommon in procellariiforms and usually only occurs after several years of breeding failure. However, one study of Short-tailed Shearwaters observed the divorce rate in a colony to be as high as 16% annually.
- Carboneras, C. 1992. "Family Diomedeidae (Albatross)" in Handbook of Birds of the World Vol. 1. Barcelona: Lynx Editions.
- Brooke, M. 2004. Albatrosses And Petrels Across The World. Oxford University Press, Oxford, UK.
- Schreiber, Elizabeth A. and Burger, Joanne. 2001. Biology of Marine Birds. CRC Press, Boca Raton, Florida.
- Alcock, J. 2004. Understanding bird behavior, p. 6-1–6-98. In S. Podulka, R. W. Rohrbaugh, and R. Bonney [Eds.], Handbook of Bird Biology. Princeton University Press, Princeton, NJ.
- Warham, J. 1996. The behaviour, population biology, and physiology of the petrels. Academic Press, London, UK.
- Meseth, E. H. 1975. Dance of Laysan Albatross, Diomedea immutablis. Behaviour 54:217-257.
- Awkerman, Jill A., David J. Anderson and G. Causey Whittow. 2008. Black-footed Albatross (Phoebastria nigripes), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/065
- Awkerman, Jill, David Anderson and G. Causey Whittow. 2009. Laysan Albatross (Phoebastria immutabilis), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/066
- Pickering, S. P. C., S. D. Berrow. 2001. Courtship behaviour of the Wandering Albatross Diomedea exulans at Bird Island, South Georgia. Marine Ornithology 29: 29-37.
- Rothman, R. 1998. Waved Albatross. Seabirds of the Galapagos. Retrieved 24 Jan. 2011 http://people.rit.edu/rhrsbi/GalapagosPages/Albatross.html
- Schreiber, E. A., R. W. Schreiber and G. A. Schenk. 1996. Red-footed Booby (Sula sula), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/241
- Schreiber, E. A. and R. L. Norton. 2002. Brown Booby (Sula leucogaster), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/649
- Diamond, A. W. 1973. Notes on breeding biology and behavior of Magnificent Frigatebird. Condor 75:200-209.
- Diamond, Antony W. and Elizabeth A. Schreiber. 2002. Magnificent Frigatebird (Fregata magnificens), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online: http://bna.birds.cornell.edu/bna/species/601
- Dearborn, D. C., A. D. Anders, and J. B. Williams. 2005. Courtship display by great frigatebirds, Fregata minor: an energetically costly handicap signal? Behavioral Ecology and Sociobiology 58:397-406.
- Ehrlich, P.R., D.S. Dobkin and Wheye, D. 1988. Courtship feeding. <http://www.stanford.edu/group/stanfordbirds/text/essays/Courtship_Feeding.html>
- Gwynne, D.T. 1991. Sexual competition among females: What causes courtship-role reversal? Trends in Ecology & Evolution 6(4):118-121.
- Blanchard, L. and R. D. Morris. 1998. Another look at courtship feeding and copulation behavior in the Common Tern. Colonial Waterbirds 21:251-255.
- Velando, A. 2004. Female control in yellow-legged gulls: trading paternity assurance for food. Animal Behaviour 67(5):899-907.
- Kempenaers, B., R. B. Lanctot, V. A. Gill, S. A. Hatch, and M. Valcu. 2007. Do females trade copulations for food? An experimental study on kittiwakes (Rissa tridactyla). Behavioral Ecology 18:345-353.
- Bagemihl, B. 1999. Biological Exuberance: Animal Homosexuality and Natural Diversity. St. Martin's Press, New York, New York.
- Zuk, M. and N. W. Bailey. 2008. Birds gone wild: same-sex parenting in albatross. Trends in Ecology & Evolution 23:658-660.
- Bailey, N. W. and M. Zuk. 2009. Same-sex sexual behavior and evolution. Trends in Ecology & Evolution 24:439-446.
- MacFarlane, G. R., S. P. Blomberg, and P. L. Vasey. 2010. Homosexual behaviour in birds: frequency of expression is related to parental care disparity between the sexes. Animal Behaviour 80:375-390.
- Fitch, M. 1980. Monogamy, polgamy, and female-female pairs in Herring Gulls. Proceedings of the Colonial Waterbird Group 3: 44-48.
- Shugart, G. W., M. A. Fitch, and G. A. Fox. 1988. Female pairing – a reproductive strategy for Herring Gulls. Condor 90:933-935.
- Hunt, G. L. and M. W. Hunt. 1977. Female-female pairing in Western Gulls (Larus occidentalis) in southern California. Science 196:1466-1467.
- Conover, M. R., D. E. Miller, and G. L. Hunt. 1979. Female-female pairs and other unusual reproductive associations in Ring-billed and California Gulls. Auk 96:6-9.
- Kovacs, K. M. and J. P. Ryder. 1983. Reproductive performance of female-female pairs and polygynous trios of Ring-billed Gulls. Auk 100:658-669.
- Conover, M. R. and G. L. Hunt. 1984. Female-female pairing and sex ratios in gulls – an historical perspective. Wilson Bulletin 96:619-625.
- Evolution 24:439-446. Betleja, J., P. Skorka, and M. Zielinska. 2007. Super-normal clutches and female-female pairs in Gulls and Terns breeding in Poland. Waterbirds 30:624-629.
- Conover, M. R. 1983. Female-female pairing in Caspian Terns. Condor 85:346-349.
- Nisbet, I. C. T. and J. J. Hatch. 1999. Consequences of a female-biased sex-ratio in a socially monogamous bird: female-female pairs in the Roseate Tern Sterna dougallii. Ibis 141:307-320.
- Bried, J., O. Duriez, and G. Juin. 1999a. A first case of female-female pairing in the Black-faced Sheathbill Chionis minor. Emu 99:292-295.
- Lorentsen, S. H., T. Amundsen, K. Anthonisen, and J. T. Lifjeld. 2000. Molecular evidence for extrapair paternity and female-female pairs in Antarctic Petrels. Auk 117:1042-1047.
- Bried, J., M. P. Dubois, and P. Jouventin. 2009. First Case of Female-female Pairing in a Burrow-nesting Seabird. Waterbirds 32:590-596.
- Tedmanson, S. 2 Feb. 2010. Lesbian albatrosses become proud parents. The Times online. Retrieved 24 Jan. 2011 <http://www.timesonline.co.uk/tol/news/environment/article7011851.ece>
- Leach, B. 3 Feb. 2010. Lesbian albatrosses to raise chick. The Telegraph online. Retrieved 22 Jan. 2011 <http://www.telegraph.co.uk/earth/wildlife/7144393/Lesbian-albatrosses-to-raise-chick.html>
- Young, L. C., B. J. Zaun, and E. A. VanderWerf. 2008. Successful same-sex pairing in Laysan albatross. Biology Letters 4:323-325.
- Young, L. 8 Jan. 2011. The Short-tailed Albatross nest fails on Kure Atoll, Hawaii. Agreement on the conservation of albatrosses and petrels online. Retrieved 24 Jan. 2011. <http://www.acap.aq/latest-news/the-short-tailed-albatross-nest-fails-on-kure-atoll-hawaii>
- "Name search for NZ same sex couple's chick >". New Zealand. 2010-01-05. Retrieved 2012-05-15.
- Cardoze, C. 10 June 2002. They're in love. They're gay. They're penguins… And they're not alone. Columbia University News. Retrieved 25 Jan. 2011 <http://www.timelessspirit.com/SEPT04/cristina.shtml>
- Smith, D. 7 Feb. 2004. Central Park Zoo's gay penguins ignite debate. San Francisco Chronicle online. Retrieved 22 Jan 2011 <http://www.sfgate.com/cgibin/article.cgi?f=/c/a/2004/02/07/MNG3N4RAV41.DTL>
- May, M. 14 July 2009. Widow a wedge between zoo's male penguin pair. San Francisco Chronicle online. Retrieved 22 Jan. 2011 <http://www.sfgate.com/cgi-bin/article.cgi?f=/c/a/2009/07/14/BAUS18NTE7.DTL>
- Wagner, R. H. 2003. Social uses of copulation in socially monogamous Razorbills, p. 95-108. In U. Reichard and C. Boesche [Eds.], Monogamy: mating strategies and partnerships in birds, humans and other mammals. Cambridge University Press, Cambridge, UK.
- Moller A. P. and T. R. Birkhead. 1992. A pairwise comparative method as illustrated by copulation frequency in birds. American Naturalist 139: 644-656.
- Westneat, D. F. and I. R. K. Stewart. 2003. Extra-pair paternity in birds: Causes, correlates, and conflict. Annual Review of Ecology Evolution and Systematics 34:365-396.
- Grifﬁth S. C., Owens I. P. F. and K. A. Thuman. 2002. Extra pair paternity in birds: a review of interspeciﬁc variation and adaptive function. Molecular Ecology. 11:2195–212.
- Westneat, D. F. and P. W. Sherman. 1997. Density and extra-pair fertilizations in birds: a comparative analysis. Behavioral Ecology and Sociobiology 41:205-215.
- Mauck, R. A., T. A. Waite, and P. G. Parker. 1995. Monogamy in Leach's Storm-Petrel: DNA-fingerprinting evidence. Auk 112:473-482.
- Quillfeldt, P., T. Schmoll, H. U. Peter, J. T. Epplen, and T. Lubjuhn. 2001. Genetic monogamy in Wilson's Storm-Petrel. Auk 118:242-248.
- Lifjeld, J. T., A. M. A. Harding, F. Mehlum, and T. Oigarden. 2005. No evidence of extra-pair paternity in the Little Auk Alle alle. Journal of Avian Biology 36:484-487.
- Anderson, D. J. and P. T. Boag. 2006. No extra-pair fertilization observed in Nazca Booby (Sula granti) broods. Wilson Journal of Ornithology 118:244-247.
- Moreno, J., L. Boto, J. A. Fargallo, A. de Leon, and J. Potti. 2000. Absence of extra-pair fertilisations in the Chinstrap Penguin Pygoscelis antarctica. Journal of Avian Biology 31:580-583.
- Huyvaert, K. P., D. J. Anderson, T. C. Jones, W. R. Duan, and P. G. Parker. 2000. Extra-pair paternity in waved albatrosses. Molecular Ecology 9:1415-1419.
- Jouventin, P., A. Charmantier, M. P. Dubois, P. Jarne, and J. Bried. 2007. Extra-pair paternity in the strongly monogamous Wandering Albatross Diomedea exulans has no apparent benefits for females. Ibis 149:67-78.
- Burg, T. M. and J. P. Croxall. 2006. Extrapair paternities in black-browed Thalassarche melanophris, grey-headed T-chrysostoma and wandering albatrosses Diomedea exulans at South Georgia. Journal of Avian Biology 37:331-338.
- Austin, J. J., R. E. Carter, and D. T. Parkin. 1993. Genetic evidence for extra-pair fertilizations in socially monogamous Short-tailed Shearwaters, Puffinus tenuirostris (Procellariiformes, Procellariidae), using DNA fingerprinting. Australian Journal of Zoology 41:1-11.
- Austin, J. J. and D.T. Parkin. 1996. Low frequency of extra-pair paternity in two colonies of the socially monogamous short-tailed shearwater Puffinus tenuirostris. Molecular Ecology. 5(1):145-150.
- Birkhead, T. R., S. D. Johnson, and D. N. Nettleship. 1985. Extra-pait matings and mate-guarding in the Common Murre Uria aalge. Animal Behaviour 33:608-619.
- Birkhead, T. R., B. J. Hatchwell, R. Lindner, D. Blomqvist, E. J. Pellatt, R. Griffiths, and J. T. Lifjeld. 2001. Extra-pair paternity in the Common Murre. Condor 103:158-162.
- Wagner, R. H. 1992. Behavioral and breeding-habitat related aspects of sperm competition in Razorbills. Behaviour 123:1-26.
- Wagner, R. H. 1991. Evidence that female Razorbills control extra-pair copulations. Behaviour 118:157-169.
- Dearborn, D. C., A. D. Anders, and P. G. Parker. 2001. Sexual dimorphism, extrapair fertilizations, and operational sex ratio in great frigatebirds (Fregata minor). Behavioral Ecology 12:746-752.
- Pusey, A. and M. Wolf. 1996. Inbreeding avoidance in animals. Trends in Ecology & Evolution 11:201-206.
- Inchausti, P. and H. Weimerskirch. 2001. Risks of decline and extinction of the endangered Amsterdam Albatross and the projected impact of long-line fisheries. Biological Conservation 100:377-386.
- Bried, J., M. Nicolaus, P. Jarne, M. P. Dubois, and P. Jouventin. 2007. Population biology of the wandering albatross (Diomedea exulans) in the Crozet and Kerguelen archipelagos, southern Indian Ocean, approached through genetic and demographic methods. Journal of Zoology 272:20-29.
- Huyvaert, K. P. and P. G. Parker. 2010. Extra-pair paternity in waved albatrosses: genetic relationships among females, social mates and genetic sires. Behaviour 147:1591-1613.
- Rabouam, C., J. C. Thibault, and V. Bretagnolle. 1998. Natal philopatry and close inbreeding in Cory's shearwater (Calonectris diomedea). Auk 115:483-486.
- Goth, A. and C. S. Evans. 2004. Social responses without early experience: Australian brush-turkey chicks use specific visual cues to aggregate with conspecifics. Journal of Experimental Biology 207:2199-2208.
- Skutch; Alexander Frank (author) and Gardner, Dana (illustrator) Helpers at birds' nests : a worldwide survey of cooperative breeding and related behavior pp. 69–71. Published 1987 by University of Iowa Press. ISBN 0877451508
- Russell, Eleanor M.; “Avian Life Histories: Is Extended Parental Care the Southern Secret?”; in Emu; Vol. 100, 377-399 (2000)
- Creighton, J. C. and G. D. Schnell. 1996. Proximate control of siblicide in cattle egrets: A test of the food-amount hypothesis. Behavioral Ecology and Sociobiology 38:371-377.
- Watson, R. T., S. Razafindramanana, R. Thorstrom, and S. Rafanomezantsoa. 1999. Breeding biology, extra-pair birds, productivity, siblicide and conservation of the Madagascar Fish Eagle. Ostrich 70:105-111.
- Anderson, D. J. 1990. Evolution of obligate siblicide in boobies: a test of the insurance-egg hypothesis. American Naturalist 135:334-350.
- Tarlow, E. M., M. Wikelski, and D. J. Anderson. 2001. Hormonal correlates of siblicide in Galapagos Nazca Boobies. Hormones and Behavior 40:14-20.
- Muller, M.S., J.F. Brennecke, E.T. Porter, M.A. Ottinger and D.J. Anderson. 2008. Perinatal Androgens and Adult Behavior Vary with Nestling Social System in Siblicidal Boobies. PLoS One 3. <http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0002460>
- Lougheed, L. W. and D. J. Anderson. 1999. Parent blue-footed boobies suppress siblicidal behavior of offspring. Behavioral Ecology and Sociobiology 45:11-18.
- Johnstone, V. and J. P. Ryder. 1987. Divorce in larids: a review. Colonial Waterbirds 10(1):16-26.
- Jeschke, J. M., S. Wanless, M. P. Harris, and H. Kokko. 2007. How partnerships end in guillemots Uria aalge: chance events, adaptive change, or forced divorce? Behavioral Ecology 18:460-466.
- Maness, T. J. and D. J. Anderson. 2008. Mate rotation by female choice and coercive divorce in Nazca boobies, Sula granti. Animal Behaviour 76:1267-1277.
- Bried, J., F. Jiguet and P. Jouventin. 1999b.Why do Aptenodytes penguins have high divorce rates? Auk 116(2):504-512.
- Catry, P., N. Ratcliffe, and R. W. Furness. 1997. Partnerships and mechanisms of divorce in the great skua. Animal Behaviour 54:1475-1482.
- Bradley, J. S., R. D. Wooller, and D. L. Serventy. 1990. The influence of mate retention and divorce upon the reproductive success in Short-tailed Shearwaters Puffinus tenuirostris. Journal of Animal Ecology 59:487-496.