This bird order contains small to medium birds. Most of the about 350 species occur near water. They were formerly divided into three suborders: the Charadrii (waders), Lari (gulls) and Alcae (auks). Hybridization occurs in many genera.



An apparent hybrid between Dunlin (C. alpina) and White-rumped Sandpiper (C. fuscicollis) was documented in Wisconsin (Anich, 2013).



There might be a hybrid zone between subspecies of the Common Ringed Plover (C. hiaticula), running from Northern Scandinavia to Belarus (Thies et al., 2018).

Several genetic studies uncovered gene flow between Kentish Plover (C. alexandrinus) and White-faced Plover (C. dealbatus), two species that recently established secondary contact (Wang et al., 2019a; Wang et al., 2019b).


A Kentish Plover (left) and White-faced Plover (right) at Tanjung Tokong in Malaysia. © D.N. Bakewell | PLoS One



In Norway, a hybrid between Great Snipe (G. media) and Common Snipe (G. gallinago) was described, based on behaviour, morphology and genetic data (Höglund et al., 2015).

Gallinago gallinago and G. media

Common Snipe (Gallinago gallinago) and Great Snipe (G. media)



Several hybrids in this genus have been documented, all based on morphological data:

  • Island Pied Oystercatcher (H. finschi) x Variable Oystercatcher (H. unicolor) (Baker, 1975Crocker, Petch & Sagar, 2010)
  • Magellanic Oystercatcher (H. leucopodus) x Blackish Oystercatcher (H. ater) (Jehl, 1978)
  • Black Oystercatcher (H. bachmani) x American Oystercatcher (H. palliatus) (Jehl, 1985)
Haematopus longirostris

Pied Oystercatcher (Haematopus longirostris)



Hybridization between the endemic Black Stilt (H. novaezelandiae) and the Pied Stilt (H. himantopus leucocephalus) is of major conservation concern (Wallis, 1999). The situation has been studied from a morphological (Pierce, 1984) and genetic (Greene, 1999; Steeves et al., 2010) perspective. The most recent study found evidence for extensive hybridization, but not for introgression (Steeves et al., 2010).

In addition, a hybrid between American Avocet (Recurvirostra americana) and Black-necked Stilt (H. mexicanus) is known (Principe, 1977).

Himantopus novaezelandiae and H. himantopus

Black Stilt (Himantopus novaezelandiae) and Pied Stilt (H. himantopus)



One study gives a morphological description of a Northern Jacana (J. spinosa) x Wattled Jacana (J. jacana) hybrid (Betts, 1973). A study of the Panamanian hybrid zone between both species revealed introgression from J. jacana into J. spinosa (Miller et al., 2014). Moreover, female body mass, a trait involved in competition between females, has introgressed from Northern into Wattled Jacana (Lipshutz et al., 2019).

Jacana jacana and J. spinosa

Wattled Jacana (Jacana jacana) and Northern Jacana (J. spinosa)



Hybridization is widespread among gulls (Lönnberg, 1919; Smith, 1966). Due to the high frequency of hybridization and the rapid speciation in this bird group, they are genetically closely related (Snell, 1991a), which hampers the construction of phylogenetic trees (Crochet et al., 2003; Crochet, Lebreton & Bonhomme, 2002; Sonsthagen et al., 2016) and leads to the lack of a genetic population structure (Sonsthagen et al., 2012). Gulls were once seen as a classical example of a ring species, but this has been proven incorrect (Liebers, de Knijff & Helbig, 2004).

The Herring Gull complex (Larus argentatus – cachinnans – fuscus) is circumpolar and has been studied extensively in terms of breeding area distribution (Adriaens, Vercruijsse & Stienen, 2012; Filchagov, 1994; Yesou, 1991) and genetic structure (Liebers, Helbig & De Knijff, 2001). Most studies have focused on certain species combinations, always involving the Herring Gull (L. argentatus). Gull species hybridizing with Herring Gull include Yellow-legged Gull (L. michahellis) in the Mediterranean (Pons et al., 2004), Great Black-backed Gull (L. marinus) in North America (Andrle, 1972; Jehl, 1960), Lesser Black-backed Gull (L. fuscus) in Europe (Brown, 1967; Harris, 1970; Harris, Morley & Green, 1978) and  North America (Ellis et al., 2014), and Glaucous-winged Gull (L. glaucescens) in North America (Mlodinow; Patten & Weisbrod, 1974; Williamson & Peyton, 1963).

The Iceland Gull complex (L. glaucoides – kumlieni – thayeri) consists of three (sub)species that all interbreed (Gaston & Decker, 1985; McGowan & Kitchener, 2001; Snell, 1989; Weir, Kitchener & McGowan, 2000). Other species combinations that have been studied were Red-billed Gull (L .novaehollandiae) x Black-billed Gull (L. bulleri) in New Zealand (Gurr, 1967) and American Herring Gull (L. smithsonianus) x Great Black-backed Gull (L. marinus) in North America (Pons et al., 2014). The former cross probably resulted in introgression of mtDNA (Mischler et al., 2018).

A few species combinations have received more attention.


Herring Gull (L. argentatus) x Caspian Gull (L. cachinnans)

There are hybrid zones between these species in Poland (Faber et al., 2001; Neubauer et al., 2006) and eastern Europe (Panov & Monzikov, 1999). Genetic studies revealed asymmetric introgression (Gay et al., 2007) and selection on plumage melanism and bare-parts coloration (Gay et al., 2009). The former can be explained by latitude (i.e., Gloger’s Rule), while the latter may be due to sexual selection. Behavioural observations revealed that there is premating isolation due to different breeding phenology and assortative mating according to bare-parts coloration (Neubauer et al., 2009). There is no strong postmating isolation: breeding performance did not differ between pure species and hybrids (Zagalska-Neubauer & Neubauer, 2012). However, female hybrids do seem to have a lower survival rate (Neubauer, Nowicki & Zagalska-Neubauer, 2014).

Larus argentatus and L. cachinnans

Larus argentatus and L. cachinnans


Herring Gull (L. argentatus) x Glaucous Gull (L. hyperboreus)

Numerous of these hybrids have been reported (Andrle, 1980; Jehl, 1971; Jehl & Frohling, 1965; Spear, 1987). Ingolfsson (1970; 1987) described the hybrid zone on Iceland. However, Snell (1991b) argued that the intermediate plumage was not due to hybridization, but a consequence of light-winged founders in the 1920s. Detailed genetic (and morphological) analyses settled the debate and indicated widespread hybridization (Palsson, Vigfusdottir & Ingolfsson, 2009; Sternkopf et al., 2010; Vigfusdottir, Palsson & Ingolfsson, 2008).

Larus hyperboreus

Glaucous Gull (Larus hyperboreus)


Western Gull (L. occidentalis) x Glaucous-winged Gull (L. glaucescens)

The hybrid zones between these species is located at the coast of Washington and Oregon (Scott, 1971) and has been described from a morphological and genetic perspective (Bell, 1996; Hoffman, Wiens & Scott, 1978). Cline analysis of the hybrid zone detected introgression (Gay et al., 2008), but also (sexual) selection on bare-parts coloration (Gay et al., 2009). The majority of literature on this hybrid zone focuses on breeding success. Bell (1997) found that L. occidentalis has a higher breeding success compared to L. glaucescens and mixed pairs, while Good et al. (2000) documented hybrid superiority. The breeding success of the hybrids was mainly influenced by nest predation, which decreased with increasing vegetation structure around the nest (Good, 2002). However, recent studies did not detect any differences in breeding performance (Megna et al., 2014; Moncrieff et al., 2013).

Larus occidentalis and L. glaucescens

Western Gull (Larus occidentalis) and Glaucous-winged Gull (L. glaucescens)



While studying the speciation process of American Golden-plover (P. dominica) and Pacific Golden-plover (P. fulva), Withrow and Winker (2014) detected one hybrid individual. Analyses showed that gene flow was close to zero.

Pluvialis dominica and P. fulva

American Golden-plover (Pluvialis dominica) and Pacific Golden-plover (P. fulva)


Tribe Sternini (Terns)

Numerous hybrids between various species have been recorded and described, but without any further studies.

Sterna hirundo

Common Tern (Sterna hirundo)



Several observations of hybrids (and backcrosses) between Common Murre (U. aalge) and Thick-billed Murre (U. lomvia) have been reported (Birkhead, Johnson & Nettleship, 1986; Cairns & deYoung, 1981). Reports of hybrids based on morphological characters are often questioned (Cairns, 1983; Sluys, 1983), but a backcross was successfully identified based on molecular data (Friesen et al., 1993). Furthermore, a detailed study detected cryptic introgression between the two species (Taylor et al., 2012).

A possible hybrid between Common Murre and Razorbill (Alca torda) has been reported (Wilhelm et al., 2001).

Uria aalge and U. lomvia

Common Murre (Uria aalge) and Thick-billed Murre (U. lomvia)



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Greene, B. (1999). Genetic variation and hybridisation of black stilts (Himantopus novaezelandiae) and pied stilts (H-h. leucocephalus), Order Charadriiformes. New Zealand Journal of Zoology 26, 271-277.

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Megna, L. C., Moncrieff, A. E., Hayward, J. L. & Henson, S. M. (2014). Equal reproductive success of phenotypes in the Larus glaucescens–occidentalis complex. Journal of Avian Biology 45, 410-416.

Miller, M. J., Aguilar, C., De León, L. F., Loaiza, J. R. & McMillan, W. O. (2014). Complete mitochondrial genomes of the New World jacanas: Jacana spinosa and Jacana jacana. Mitochondrial DNA, 1-2.

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Vigfusdottir, F., Palsson, S. & Ingolfsson, A. (2008). Hybridization of glaucous gull (Larus hyperboreus) and herring gull (Larus argentatus) in Iceland: mitochondrial and microsatellite data. Philosophical Transactions of the Royal Society B-Biological Sciences 363, 2851-2860.

Wallis, G. (1999). Genetic status of New Zealand black stilt (Himantopus novaezelandiae) and impact of hybridisation. Department of Conservation Wellington.

Wang, X., Que, P., Heckel, G., Hu, J., Zhang, X., Chiang, C. Y., et al. (2019). Genetic, phenotypic and ecological differentiation suggests incipient speciation in two Charadrius plovers along the Chinese coast. BMC Evolutionary Biology19(1), 135.

Wang, X., Maher, K. H., Zhang, N., Que, P., Zheng, C., Liu, S., et al. (2019). Demographic histories and genome-wide patterns of divergence in incipient species of shorebirds. Frontiers in Genetics10, 919.

Weir, D. N., Kitchener, A. C. & McGowan, R. Y. (2000). Hybridization and changes in the distribution of Iceland gulls (Larus glaucoides kumlieni/thayeri). Journal of Zoology 252, 517-530.

Whittam, R. M. (1998). Interbreeding of roseate and arctic terns. The Wilson Bulletin, 65-70.

Wilhelm, S., Walsh, C., Stenhouse, I. & Storey, A. (2001). A possible common guillemot (Uria aalge) X razorbill (Alca torda) hybrid. Atl. Seabirds 3, 85-88.

Williamson, F. S. & Peyton, L. J. (1963). Interbreeding of Glaucous-winged and Herring gulls in the Cook Inlet region, Alaska. Condor, 24-28.

Withrow, J. J. & Winker, K. (2014). Genetics of a High-Latitude Cryptic Speciation Event: American and Pacific Golden-Plovers. Wilson Journal of Ornithology 126, 429-442.

Yang, J., Chen, G., Yuan, L., Huang, Q., Fan, Z., Lu, Y., Liu, Y. & Chen, S. (2018) Genetic evidence of the world’s most endangered tern, the Chinese Crested Tern Thalasseus bernsteiniIbis

Yesou, P. (1991). The sympatric breeding of Larus fuscus, L. cachinnans and L. argentatus in western France. Ibis 133, 256-263.

Zagalska-Neubauer, M. & Neubauer, G. (2012). Reproductive performance and changes in relative species abundance in a mixed colony of Herring and Caspian Gulls, Larus argentatus and Larus cachinnans. Acta Ornithologica 47, 185-194.

Zingo, J., Church, C. & Spendelow, J. (1994). Two hybrid common x roseate terns fledge at Falkner Island, Connecticut. The Connecticut Warbler 14, 50-55.

* The Larus text has been reviewed by Grzegorz Neubauer (Polish Academy of Sciences, Warsaw)

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