Anseriformes comprises about 150 species, divided over three families: Anhimidae (screamers), Anseranatidae (Magpie Goose) and Anatidae (ducks, geese and swans). Overviews of the incidence of hybridization among Anseriformes have been published (Johnsgard, 1960; Randler, 2000; Randler, 2008; Scherer & Hilsberg, 1982). Apart from a phylogenetic analysis, Gonzalez et al. (2009) also discuss patterns of hybridization. In waterfowl, the occurrence of hybridization can be explained by interspecific brood amalgamation and (to a lesser extent) forced extrapair copulations (Randler, 2005).
Geese (Anser, Branta & Chen)
Hybridization is common among geese, but few studies have compiled all occurrences (Delany, 1992; Kampe-Persson & Lerner, 2007; Randler, 2000; Randler, 2008). The mitochondrial DNA of Anser species is closely related, but this could also be due to incomplete lineage sorting (Ruokonen, Kvist & Lumme, 2000). Several species combinations have been studied. Hybrids between Snow Goose (C. caerulescens) and Ross’ Goose (C. rossii) have been reported (Hatch & Shortt, 1976; Trauger, Dzubin & Ryder, 1971) and experimental (Macinnes & Kerbes, 1987) and genetic (Weckstein et al., 2002) studies are known. Canada Goose (B. canadensis) hybrids with Snow Goose (Nelson, 1952; Prevett & Macinnes, 1973) and Greater White-fronted Goose (Craven & Westemeier, 1979) have only been documented, with some estimates of frequency. The occurrence of wild hybrids between Greater and Lesser White-fronted Goose (A. erythropus) is certain (Nijman, Aliabadian & Roselaar, 2010), while the genetic purity of captive birds that were destined for a reintroduction programme remains a matter of debate (Ruokonen, Andersson & Tegelstrom, 2007). And a hybrid zone between Canada Goose and Cackling Goose (B. hutchinsii) is located in Northern Canada (Leafloor, Moore & Scribner, 2013). Hybrids between Greylag Goose and domestic breeds have been confirmed by mtDNA analyses (Heikkinen et al., 2015).
The morphology of the following hybrids have been described in more detail: Bar-headed Goose x Snow Goose (Lehmhus & Gustavsson, 2014), Greylag Goose x Canada Goose (Gustavsson, 2011) and several hybrids between Barnacle Goose and Anser species (Gustavsson, 2009). In addition, some hybrids have coloured tail-coverts although all goose species have white tail-coverts (Gustavsson, 2010).
The behaviour of hybrids between Swan Goose (A. cygnoides) and Greylag Goose (A. anser), such as vigilance (Randler, 2003) and aggression (Randler, 2004), is no different from pure species. Also, the breeding times of Snow Goose x Greylag Goose hybrids are intermediate in a captive setting (Davies, Fischer & Gwinner, 1969).
Next to these morphological, ecological and genetic studies, goose hybrids have also been extensively studied with regard to the meat industry (Kowalczyk, Adamski & Lukaszewicz, 2013; Kowalczyk & Lukaszewicz, 2012; Mazanowski & Bernacki, 2006).
The duck genus Anas has been studied extensively. The key species in this group is the Mallard (A. platyrhynchos), which hybridizes with the majority of other duck species. Phylogeographic analysis of the Mallard revealed two mitochondrial haplotype clades, A and B (Avise, Ankney & Nelson, 1990; Kraus et al., 2011; Kulikova et al., 2005; Kulikova, Poysa & Zhuravlev, 2012). Worldwide, several monochromatic duck species evolved, with which the Mallard hybridizes extensively:
- Black Duck (A. superciliosa) in Australia and New Zealand (Braithwaite & Miller, 1975; Gillespie, 1985; Haddon, 1984; Rhymer, Williams & Braun, 1994; Taysom, Johnson & Guay, 2014)
- Hawaii Duck (A. wylvilliana) on the Hawaiian islands (Browne et al., 1993; Fowler, Eadie & Engilis, 2009)
- American Black Duck (A. rubripes) in North America (Johnsgard, 1967; Kirby, Sargeant & Shutler, 2004; Mank, Carlson & Brittingham, 2004)
- Spot-billed Duck (A. zonorhyncha) in the Russian Far East (Kulikova, Zhuravlev & McCracken, 2004)
- Mottled Duck (A. fulvigula) in North America (Bielefeld et al., 2016; McCracken, Johnson & Sheldon, 2001; Peters et al., 2016; Peters et al., 2014a; Seyoum et al., 2012; Williams et al., 2005).
- Mexican Duck (A. diazi) in Mexico (Lavretsky et al., 2015a)
Besides the Mallard, other combinations of hybridizing duck species have been studied, such as Gadwalls (A. strepera) and Falcated ducks (A. falcata) in North America (Peters & Omland, 2007; Peters et al., 2007), Speckled Teal (A. flavirostris) and Yellow-billed Pintail (A. georgica) on the Falkland Islands (McCracken & Wilson, 2011), and Grey Teal (A. gracilis) and Chestnut Teal (A. castanea) in Australia (Joseph et al., 2009). Genetic analyses show that hybridization between Grey Teal and Pacific Black Duck occurs bidirectionally (Guay et al., 2015). Several subspecies of the Common Teal (A. crecca) also interbreed: there is continental gene flow between A. c. crecca and A. c. carolensis (Peters et al., 2012), and possible heteropatric speciation (Winker, 2006) between A. c. crecca and A. c. nimia (Winker et al., 2013). In Japan, the behaviour of a A. penelope x falcata hybrid was recorded (Chiba & Honma, 2010).
Hybridization between Speckled Teal and Yellow-billed Pintail might have led to the introgression of haemoglobin molecular adapted to high altitude, an possible example of adaptive introgression in birds (Natarajan et al., 2015)!
The Hawaiian duck may represent a young hybrid species. Genetic analyses show that the contemporary Hawaiian duck is descended from an ancient hybridization event (dated around the Pleistocene-Holocene boundary) between Mallard and Laysan duck (A. laysanensis) (Lavretsky et al., 2015).
Experimental work on Mallard x Black Duck hybrids showed that hybrids were more similar to Black Ducks in terms of salt toleration (Barnes & Nudds, 1991). Hybrids also tended to have more Sarcocystis parasites compared to “pure” species (Mason & Clark, 1990).
Finally, several studies provided clues for introgressive hybridization by studying several species (Kraus et al., 2012; Lavretsky, McCracken & Peters, 2014; Peters et al., 2005; Peters et al., 2014b).
One experimental study looked at sexual imprinting of a facultative brood parasite (A. americana) on his host (A. valisineria). They found that cross-fostered individuals approached and courted more towards their foster parents (Sorenson, Hauber & Derrickson, 2010). And Lavretsky et al. (2016) found evidence for gene flow from Lesser (A. affinis) into Greater Scaup (A. marila).
Bucephala (and Mergus)
Hybrids between species of this genus have only been described briefly, for instance Common Goldeneye (B. clangula) interbreeds with Barrow’s Goldeneye (B. islandica) (Martin & Dilabio, 1994), Smew (Mergus albellus) (Kovacovsky & Rychlik, 1999) and Bufflehead (B. albeola) (Finley & Huot, 2010).
Hybrids between Mute Swan (C. olor) and Whooper Swan (C. cygnus), and between Whooper Swan and Bewick’s Swan (C. columbianus bewickii) have been documented in the Baltic and are probably due to recent range expansions (Kampe-Persson & Boiko, 2011). Studies on hybrid swans are rare. One study compared the call characteristics of Trumpeter Swans (C. buccinators), Tundra Swans (C. columbianus) and their hybrids (Wood, Brooks & Sladen, 2002). Another study documented the behaviour of a hybrid between Whooper Swan and Mute Swan (Panov & Pavlova, 2010).
This genus is well known for the conservation issue of hybridization between the threatened White-headed Duck (O. leucocephala) and the invasive Ruddy Duck (O. jamaicensis) in Spain. The Ruddy Ducks are descendants of captive birds from the UK (Munoz-Fuentes et al., 2006), but a successful elimination program has prevented extensive introgression (Munoz-Fuentes et al., 2007). The situation is nicely summarized in Munoz-Fuentes et al. (2013). Both species and their hybrids use similar food sources (Sanchez, Green & Dolz, 2000).
One study describes a hybrid between Common Eider (S. mollissima) and King Eider (S. spectabilis) on the Kent Peninsula (Trefry, Dickson & Hoover, 2007).
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