Portal de Periódicos da CAPES

Taxonomic recommendations for British birds: eighth report

2012; Wiley; Volume: 154; Issue: 4 Linguagem: Inglês

10.1111/j.1474-919x.2012.01273.x

1474-919X

George Sangster, J. Martin Collinson, Pierre‐André Crochet, Alan G. Knox, David T. Parkin, Stephen C. Votier,

Genetic diversity and population structure

This paper is the eighth report of the Taxonomic Sub-Committee of the BOU Records Committee relating to the British List. Species-level decisions are based on criteria outlined by Helbig et al. (2002). Taxonomic Sub-Committee membership is George Sangster (Secretary), Martin Collinson (Convenor), Pierre-André Crochet and Stephen C. Votier. The seventh report of the Sub-Committee was published by Sangster et al. (2011). Grouse (Tetraonidae, sensu Voous 1977) are nested within Phasianidae (sensu Voous 1977), as demonstrated by a number of molecular phylogenetic studies (Kimball et al. 1999, Dimcheff et al. 2000, 2002, Crowe et al. 2006, Pereira & Baker 2006, Kaiser et al. 2007, Kriegs et al. 2007, Kimball & Braun 2008, Meng et al. 2008, Shen et al. 2010). The grouse are therefore best included in Phasianidae (Crowe et al. 2006). New World quails, which were included in Phasianidae by Voous (1977), form a distinct clade sister to Phasianidae (sensu Crowe et al. 2006) and are better included in a separate taxon, Odontophoridae (Crowe et al. 2006, Cox et al. 2007, Kriegs et al. 2007). Most studies of the grouse support non-monophyly of the genus Bonasa (Ellsworth et al. 1996, Dimcheff et al. 2000, 2002, Gutiérrez et al. 2000, Crowe et al. 2006, but see Lucchini et al. 2001, Drovetski 2002). Hazel Grouse B. bonasia and Severtsov's Grouse B. sewerzowi are therefore best placed in Tetrastes. The black grouse (Tetrao tetrix and T. mlokosiewiczi) have been placed in the genus Lyrurus (Peters 1934) but were lumped in Tetrao by Voous (1977). More recently, some authors have reinstated Lyrurus (e.g. Gutiérrez et al. 2000, Dickinson 2003, Svensson et al. 2009). Molecular phylogenetic studies support a sister relationship between the black grouse and capercaillies (Ellsworth et al. 1996, Dimcheff et al. 2000, Crowe et al. 2006) and studies that included all four species found that the two black grouse are sister to the two capercaillies (Gutiérrez et al. 2000, Lucchini et al. 2001, Dimcheff et al. 2002, Drovetski 2002). Thus, both arrangements (one genus vs. two genera) are consistent with current knowledge of the phylogeny of grouse. Other evidence is not helpful in deciding between the two alternative arrangements: the two black grouse are more similar to each other in morphology (plumage, size) and habits (ecotonal/open habitats vs. forested habitats) than they are to the capercaillies, but on the other hand there are clear morphological (very dark/black plumage) and behavioural similarities (e.g. polygyny, lekking behaviour). Recognition of Lyrurus would be consistent with all molecular studies but it is not warranted and would create two genera consisting of only two species. We therefore retain the black grouse in Tetrao. Phylogenetic analysis of molecular and morpho-behavioural characters suggests that the genus Francolinus is not monophyletic (Crowe et al. 1992, 2006, Bloomer & Crowe 1998). A re-arrangement of four (Crowe et al. 1992) or five genera (Crowe et al. 2006) has been proposed, of which two occur in the Western Palearctic: Pternistis (P. bicalcaratus and P. erckelii) and Francolinus (F. francolinus). The classification suggested here is based on the phylogeny of Crowe et al. (2006, their fig. 4) except for the placement of Perdix which is poorly supported in the latter study and is therefore based on the results of other studies (Kriegs et al. 2007, Kimball & Braun 2008, Shen et al. 2010). The position of Coturnix relative to junglefowls (Gallus) and peafowls (Pavo) is unclear and generally poorly supported; some studies support a closer relationship to Gallus than to Pavo (Kriegs et al. 2007, Bonilla et al. 2010, Shen et al. 2010) whereas others place Coturnix outside the Gallus–Pavo clade (Crowe et al. 2006, Kimball & Braun 2008). We have tentatively followed Crowe et al. (2006) but stress the need for further study. The phylogeny of Crowe et al. (2006) lacked c. 8 small genera of Phasianidae (one to three species) which would not affect the taxonomic sequence derived from it. The taxonomic sequence recommended here for Western Palearctic taxa is based on the principles that, for each branching point in the phylogeny, the less-speciose group should be listed first, and equally speciose groups are listed alphabetically. Species sequence within each genus follows Voous (1977) except for Alectoris which is based on Randi (1996) and Randi and Lucchini (1998), Pternistis which is based on Bloomer and Crowe (1998), and Tetrao which is based on Drovetski (2002). Phylogenetic analyses of mitochondrial DNA sequences indicate that the three subspecies of Cory's Shearwater form reciprocally monophyletic groups and suggest that C. d. diomedea and C. d. edwardsii are more closely related to each other than either is to C. d. borealis (Gómez-Díaz et al. 2006, 2009). C. d. edwardsii differs from both C. d. borealis and C. d. diomedea in size, and in the pattern and coloration of the upperparts, cheeks, scapulars, underparts and bill (Hazevoet 1995, Hillcoat et al. 1997). C. d. borealis and C. d. diomedea differ in mean size and in the typical extent of white on the inner webs of the primaries (Thibault et al. 1997). Multivariate analysis of morphometric data identified three groups corresponding to C. d. borealis, C. d. diomedea and C. d. edwardsii (Granadeiro 1993, Gómez-Díaz & González-Solís 2007, see also Gómez-Díaz et al. 2006, 2009). The three taxa further differ in vocalizations. Duet calls of C. d. borealis have three brief syllables, whereas most (97–99%) C. d. diomedea have two longer syllables (Bretagnolle & Lequette 1990, Thibault & Bretagnolle 1998, see also Robb et al. 2008). Of c. 400 male calls of C. d. borealis and C. d. diomedea examined, none was detected from one taxon showing simultaneously two call character states of the other (Thibault & Bretagnolle 1998). Playback studies have documented differential responses to recordings of C. d. borealis and C. d. diomedea (Bretagnolle & Lequette 1990). Duet calls of C. d. edwardsii have two syllables like those of C. d. diomedea but differ from both C. d. borealis and C. d. diomedea in being noticeably higher pitched (Robb et al. 2008). There are several reports of C. d. borealis within breeding colonies of C. d. diomedea but most of these involved non-breeding individuals or birds of unknown breeding status (Lo Valvo & Massa 1988, Thibault & Bretagnolle 1998, Martinez-Abrain et al. 2002). Despite intensive monitoring of Atlantic and Mediterranean breeding colonies, reports of interbreeding by C. d. borealis and C. d. diomedea are limited to one record of a mixed pair raising a young in the Collumbretes Islands in 2001 (Martinez-Abrain et al. 2002), two birds showing phenotypic characters of one taxon and a genotype of the other taxon (Gómez-Díaz et al. 2009), and one record of a male C. d. borealis or hybrid breeding with a female with characters intermediate or atypical for C. d. diomedea on Giraglia Island in 1993, 1994 and 1995 (Thibault & Bretagnolle 1998). C. d. borealis breeds in at least two colonies in the Mediterranean Sea (Almeria: Gómez-Díaz et al. 2006, Chafarinas Islands: Navarro et al. 2009) and a few pairs of C. d. diomedea have bred along the Bay of Biscay coast in western France (Mays et al. 2006). The Almeria colony apparently consists of C. d. borealis only (Gómez-Díaz et al. 2006). The Chafarinas Islands hold 10 000 breeding pairs; based on morphometric data from 82 individuals, it has been estimated that 78% are C. d. diomedea and 22% are C. d. borealis (Gómez-Díaz et al. 2009, Navarro et al. 2009). This shows that the breeding grounds of these two forms are not fully allopatric and that some mechanisms of reproductive isolation are likely to be involved in maintaining their differences. C. d. borealis and C. d. diomedea breeding in sympatry in the Chafarinas Islands differ in their feeding ecology and foraging areas during both chick-rearing and wintering periods (Navarro et al. 2009). Cory's and Scopoli's Shearwaters are in Category A of the British List. Birds breeding in the cool season in Madeira and the Selvagens (and also in the cool season in the Azores and islands off Portugal) are sympatric with hot-season breeders in Madeira (Nunes 2000) and the Selvagens (Faria 1998), but differ from these in vocalizations (Robb et al. 2008) and morphology (Faria 1998, Nunes 2000). However, DNA differences are less clear than those among Azorean populations (Friesen et al. 2007, Smith et al. 2007). As a consequence, although potentially a fourth species, this unnamed taxon is provisionally included in O. castro pending further study and taxonomic description. Studies of vocalizations and phylogeography suggest that there are multiple lineages outside the Western Palearctic which should also be treated as species (Bolton 2007, Friesen et al. 2007, Smith et al. 2007). There is a single British record of 'Madeiran Storm Petrel' sensu lato (off Isles of Scilly, 28 July 2007; British Ornithologists' Union (BOU) 2012). European Storm Petrel has been treated as a monotypic species. However, Mediterranean and Atlantic populations show mean morphometric differences and differences in vocalizations (Hémery & D'Elbée 1985, Bretagnolle 1992, Bretagnolle & Zotier 1998, Lalanne et al. 2001, Robb et al. 2008). Phylogenetic analysis of mtDNA sequences has suggested that birds from Mediterranean and Atlantic populations are reciprocally monophyletic with an estimated separation of lineages at least 350 000 years ago (Cagnon et al. 2004). These differences strongly suggest that Mediterranean breeders should be recognized taxonomically. Pending ongoing analyses of vocalizations, Mediterranean breeding birds are recognized at subspecific level as H. p. melitensis. Nominate H. p. pelagicus is in Category A of the British List. A study using an array of osteological characters (McCracken & Sheldon 1998) found Little Blue Heron Hydranassa caerulea and Tricoloured Heron H. tricolor to form a clade with Snowy Egret Egretta thula, Little Egret E. garzetta, White-faced Heron E. novaehollandiae and Pacific Reef Heron E. sacra. These results are supported by an analysis of mtDNA sequences (Sheldon et al. 2000). Consequently, no solid evidence exists for reciprocal monophyly of 'Hydranassa' and 'Egretta' (sensu Voous 1977). The genus 'Hydranassa' is therefore merged with Egretta. In the early 1990s, molecular studies of relationships among Charadriiformes identified three major clades: plover-like birds (Charadrii), sandpiper-like birds (Scolopaci), and skuas, gulls, terns and auks (Lari) (Sibley & Ahlquist 1990, Christian et al. 1992). A series of comprehensive molecular studies published between 2003 and 2008 have corroborated the existence of the three major groups identified by Sibley and Ahlquist (1990) and Christian et al. (1992) but also provide strong support for the relationships within and among these three groups (Ericson et al. 2003, Paton et al. 2003, Thomas et al. 2004a, Paton & Baker 2006, Baker et al. 2007, Fain & Houde 2007, Hackett et al. 2008, see also Thomas et al. 2004b, van Tuinen et al. 2004, Baker & Pereira 2009). Molecular phylogenetic studies provide congruent support for the placement of Charadrii as the sister to (Scolopaci and Lari) (Ericson et al. 2003, Paton et al. 2003, Thomas et al. 2004a, Paton & Baker 2006, Baker et al. 2007, Fain & Houde 2007), whereas a morphological study recovered these clades but placed Scolopaci as the sister-group of (Charadrii and Lari) (Mayr 2011, his fig. 3C) and another morphological study did not recover any of the three clades (Livezey 2010). Some molecular studies suggested that the tundra plovers (Pluvialis) are sister to (Haematopodidae and Recurvirostridae) (Fain & Houde 2007) or to (Haematopodidae, Recurvirostridae and Charadriidae) (Ericson et al. 2003, Baker et al. 2007). However, a more detailed analysis indicates that the tundra plovers are closely related to Charadriidae (Baker et al. 2012). Within Scolopaci, (phalaropes and shanks), (turnstones and sandpipers), and (snipes, woodcocks and dowitchers) form three well-supported clades but the relationships among these are still poorly resolved (Ericson et al. 2003, Baker et al. 2007, Gibson & Baker 2012). We have tentatively followed Gibson and Baker (2012) because it is based on the most extensive data set, in terms of both sequence data and species included. The placement of the buttonquails (Turnicidae) in Charadriiformes, as the sister taxon of other Lari, is strongly supported (Paton et al. 2003, Paton & Baker 2006, Baker et al. 2007, Fain & Houde 2007, Hackett et al. 2008, but see Livezey 2010). Most studies also support the placement of Glareolidae in the Lari clade (Ericson et al. 2003, Paton et al. 2003, Paton & Baker 2006, Baker et al. 2007, Fain & Houde 2007, Mayr 2011, but see Livezey 2010). Molecular studies and one morphological study placed Crab Plover (Dromas) in Lari (Hackett et al. 2008, Pereira & Baker 2010, Mayr 2011), whereas another morphological study placed it as the sister-group of the oystercatchers (Livezey 2010). Most studies support a sister-relationship of skuas and auks (Ericson et al. 2003, Paton et al. 2003, Thomas et al. 2004a, Paton & Baker 2006, Baker et al. 2007, Fain & Houde 2007, Smith 2011, but see Livezey 2010). A close relationship between skimmers, terns and gulls is very strongly supported, although exact relationships among these three groups are not yet clear (Ericson et al. 2003, Paton et al. 2003, Paton & Baker 2006, Baker et al. 2007, Fain & Houde 2007, Livezey 2010, Mayr 2011, Smith 2011). Several studies support a clade formed by Alca, Pinguinus, Alle and Uria, although except for a sister-group relationship of Alca and Pinguinus, relationships within this clade remain unresolved (Baker et al. 2007, Pereira & Baker 2008, Smith 2011). The sequence of taxa within Charadriiformes is based on the principle that for each sister-group the less speciose taxon is listed first. The sequence and generic limits of the shanks, terns and gulls are as previously defined (Sangster et al. 2005, 2007). The taxonomic sequence of Alcidae is based on phylogenetic studies by Baker et al. (2007) and Pereira and Baker (2008). The taxa on the Western Palearctic list should be listed in the following sequence: A recent molecular phylogeny based on mitochondrial and nuclear DNA sequences provides strong evidence that the genus Calidris is paraphyletic, and that five monotypic genera are part of the Calidris clade (Aphriza, Philomachus, Limicola, Eurynorhynchus, Tryngites) (Gibson & Baker 2012). We have considered two revisions of the calidrine sandpipers: (1) include Aphriza, Philomachus, Limicola, Eurynorhynchus and Tryngites in Calidris, or, alternatively, (2) maintain some of these genera and split Calidris into several genera, e.g. Bahr (2011). Note that Bahr's (2011) proposed genera Eurynorhynchus and Tryngites cannot be maintained as such in the light of the latest molecular results since Eurynorhynchus was nested in Bahr's Leimonites, and Tryngites was nested in his Ereunetes (Gibson & Baker 2012). We prefer to include Aphriza, Philomachus, Limicola, Eurynorhynchus and Tryngites in Calidris, rather than subdivide the last-named into a number of genera, based on the following considerations. First, since some of the clades in Gibson and Baker (2012) are poorly supported, there is a risk that these are non-monophyletic, whereas an expanded Calidris is well-supported. Second, an expanded Calidris removes all monotypic genera. Third, splitting Calidris would change the names of many species, and only two species currently in Calidris would remain in that genus (i.e. the two knots, since the type species of Calidris is C. canutus). In contrast, if Calidris were expanded to include the five currently recognized monotypic genera, only five specific names would be changed and all species currently in Calidris would remain in it. Finally, even if Calidris were subdivided into a number of genera, the newly proposed polytypic genera would still be rather heterogeneous morphologically whereas no major morphological or ecological differences are known to characterize these genera. Cream-coloured Courser is in Category A of the British List. Arctic Warbler is in Category A of the British List. There are six accepted British records of Marmora's Warbler sensu lato. None was formally assigned to subspecies level, and all records should be reviewed. Phylogeographical analyses (Zink et al. 2006, Hung et al. 2012) and reports of reduced introgression in contact zones (Red'kin & Konovalova 2006) indicate that Eurasian Nuthatch may include additional species but further study is necessary. Eurasian Nuthatch is in Category A of the British List. We thank Vicki L. Friesen, P.J. Moore and Magnus Robb for their help, and Lars Svensson for extensive comments on some of the taxa included in this report.