Taxonomic recommendations for British birds:eighth report

GEORGE SANGSTER,1,2* J. MARTIN COLLINSON,3 PIERRE-ANDRÉ CROCHET,4 ALAN G. KNOX,5

DAVID T. PARKIN6 & STEPHEN C. VOTIER7

1Department of Vertebrate Zoology, Swedish Museum of Natural History, PO Box 50007, SE–104 05, Stockholm,Sweden

2Department of Zoology, Stockholm University, SE-10691, Stockholm, Sweden3School of Medical Sciences, Institute of Medical Sciences, University of Aberdeen, Aberdeen AB25 2ZD, UK

4CNRS-UMR 5175 Centre d’Ecologie Fonctionnelle et Evolutive, 1919 route de Mende, 34293, Montpellier Cedex 5,France

5University Museums, King’s College, University of Aberdeen, Aberdeen AB24 3SW, UK6Institute of Genetics, University of Nottingham, Queen’s Medical Centre, Nottingham NG7 2UH, UK

7Marine Biology & Ecology Research Centre, University of Plymouth, Plymouth PL4 8AA, UK

This paper is the eighth report of the TaxonomicSub-Committee of the BOU Records Committeerelating to the British List. Species-level decisionsare based on criteria outlined by Helbig et al.(2002). Taxonomic Sub-Committee membershipis George Sangster (Secretary), Martin Collinson(Convenor), Pierre-André Crochet and Stephen C.Votier. The seventh report of the Sub-Committeewas published by Sangster et al. (2011).

Family and generic limits of WesternPalearctic Galliformes

Grouse (Tetraonidae, sensu Voous 1977) arenested within Phasianidae (sensu Voous 1977), asdemonstrated by a number of molecular phylo-genetic studies (Kimball et al. 1999, Dimcheffet 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, Shenet al. 2010). The grouse are therefore bestincluded in Phasianidae (Crowe et al. 2006).

New World quails, which were included inPhasianidae by Voous (1977), form a distinct cladesister to Phasianidae (sensu Crowe et al. 2006) andare better included in a separate taxon, Odonto-phoridae (Crowe et al. 2006, Cox et al. 2007,Kriegs et al. 2007).

Most studies of the grouse support non-mono-phyly 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 andSevertsov’s Grouse B. sewerzowi are therefore bestplaced in Tetrastes.

The black grouse (Tetrao tetrix and T. mlokosie-wiczi) have been placed in the genus Lyrurus (Peters1934) but were lumped in Tetrao by Voous (1977).More recently, some authors have reinstated Lyru-rus (e.g. Gutiérrez et al. 2000, Dickinson 2003,Svensson et al. 2009). Molecular phylogenetic stud-ies support a sister relationship between the blackgrouse and capercaillies (Ellsworth et al. 1996,Dimcheff et al. 2000, Crowe et al. 2006) and stud-ies that included all four species found that the twoblack grouse are sister to the two capercaillies(Gutiérrez et al. 2000, Lucchini et al. 2001, Dim-cheff et al. 2002, Drovetski 2002). Thus, botharrangements (one genus vs. two genera) are consis-tent with current knowledge of the phylogeny ofgrouse. Other evidence is not helpful in decidingbetween the two alternative arrangements: the twoblack grouse are more similar to each other in mor-phology (plumage, size) and habits (ecotonal/openhabitats vs. forested habitats) than they are to thecapercaillies, but on the other hand there are clearmorphological (very dark/black plumage) andbehavioural similarities (e.g. polygyny, lekkingbehaviour). Recognition of Lyrurus would be consis-tent with all molecular studies but it is not war-ranted and would create two genera consisting ofonly two species. We therefore retain the blackgrouse in Tetrao.

*Corresponding author.Email: george.sangster@nrm.se

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Ibis (2012), 154, 874–883

Phylogenetic analysis of molecular and morpho-behavioural characters suggests that the genus Fran-colinus is not monophyletic (Crowe et al. 1992,2006, Bloomer & Crowe 1998). A re-arrangementof four (Crowe et al. 1992) or five genera (Croweet al. 2006) has been proposed, of which twooccur in the Western Palearctic: Pternistis (P. bi-calcaratus and P. erckelii) and Francolinus (F. fran-colinus).

The classification suggested here is based onthe phylogeny of Crowe et al. (2006, theirfig. 4) except for the placement of Perdix whichis poorly supported in the latter study and istherefore based on the results of other studies(Kriegs et al. 2007, Kimball & Braun 2008, Shenet al. 2010). The position of Coturnix relative tojunglefowls (Gallus) and peafowls (Pavo) isunclear and generally poorly supported; somestudies support a closer relationship to Gallusthan to Pavo (Kriegs et al. 2007, Bonilla et al.2010, Shen et al. 2010) whereas others placeCoturnix outside the Gallus–Pavo clade (Croweet al. 2006, Kimball & Braun 2008). We havetentatively followed Crowe et al. (2006) butstress the need for further study.

The phylogeny of Crowe et al. (2006) lacked c.8 small genera of Phasianidae (one to three spe-cies) which would not affect the taxonomicsequence derived from it. The taxonomic sequencerecommended here for Western Palearctic taxa isbased on the principles that, for each branchingpoint in the phylogeny, the less-speciose groupshould be listed first, and equally speciose groupsare listed alphabetically. Species sequence withineach genus follows Voous (1977) except for Alec-toris which is based on Randi (1996) and Randiand Lucchini (1998), Pternistis which is based onBloomer and Crowe (1998), and Tetrao which isbased on Drovetski (2002).

NumidaeNumida meleagris

OdontophoridaeCallipepla californicaColinus virginianus

PhasianidaeCoturnix coturnixTetraogallus caucasicusTetraogallus caspiusAlectoris barbaraAlectoris rufaAlectoris graeca

Alectoris chukarAmmoperdix griseogularisAmmoperdix heyiPternistis erckeliiPternistis bicalcaratusFrancolinus francolinusTetrastes bonasiaLagopus lagopusLagopus mutaTetrao mlokosiewicziTetrao tetrixTetrao urogallusPerdix perdixSyrmaticus reevesiiPhasianus colchicusChrysolophus amherstiaeChrysolophus pictus

Cory’s Shearwater Calonectrisdiomedea

Phylogenetic analyses of mitochondrial DNAsequences indicate that the three subspecies ofCory’s Shearwater form reciprocally monophyleticgroups and suggest that C. d. diomedea and C.d. edwardsii are more closely related to each otherthan either is to C. d. borealis (Gómez-Díaz et al.2006, 2009).

C. d. edwardsii differs from both C. d. borealisand C. d. diomedea in size, and in the patternand coloration of the upperparts, cheeks, scapu-lars, underparts and bill (Hazevoet 1995, Hillcoatet al. 1997). C. d. borealis and C. d. diomedea dif-fer in mean size and in the typical extent ofwhite on the inner webs of the primaries(Thibault et al. 1997). Multivariate analysis ofmorphometric data identified three groups corre-sponding 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 sylla-bles, whereas most (97–99%) C. d. diomedea havetwo longer syllables (Bretagnolle & Lequette 1990,Thibault & Bretagnolle 1998, see also Robb et al.2008). Of c. 400 male calls of C. d. borealis andC. d. diomedea examined, none was detected fromone taxon showing simultaneously two call charac-ter states of the other (Thibault & Bretagnolle1998). Playback studies have documented differen-

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tial responses to recordings of C. d. borealis and C.d. diomedea (Bretagnolle & Lequette 1990). Duetcalls of C. d. edwardsii have two syllables likethose of C. d. diomedea but differ from both C.d. borealis and C. d. diomedea in being noticeablyhigher pitched (Robb et al. 2008).

There are several reports of C. d. borealis withinbreeding colonies of C. d. diomedea but most ofthese involved non-breeding individuals or birds ofunknown breeding status (Lo Valvo & Massa1988, Thibault & Bretagnolle 1998, Martinez-Abrain et al. 2002). Despite intensive monitoringof Atlantic and Mediterranean breeding colonies,reports of interbreeding by C. d. borealis and C.d. diomedea are limited to one record of a mixedpair raising a young in the Collumbretes Islands in2001 (Martinez-Abrain et al. 2002), two birdsshowing phenotypic characters of one taxon and agenotype of the other taxon (Gómez-Díaz et al.2009), and one record of a male C. d. borealis orhybrid breeding with a female with charactersintermediate or atypical for C. d. diomedea onGiraglia Island in 1993, 1994 and 1995 (Thibault& Bretagnolle 1998).

C. d. borealis breeds in at least two coloniesin the Mediterranean Sea (Almeria: Gómez-Díazet al. 2006, Chafarinas Islands: Navarro et al. 2009)and a few pairs of C. d. diomedea have bred alongthe Bay of Biscay coast in western France (Mayset al. 2006). The Almeria colony apparently con-sists of C. d. borealis only (Gómez-Díaz et al.2006). The Chafarinas Islands hold 10 000 breed-ing pairs; based on morphometric data from 82individuals, 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 showsthat the breeding grounds of these two forms arenot fully allopatric and that some mechanisms ofreproductive isolation are likely to be involved inmaintaining their differences. C. d. borealis and C.d. diomedea breeding in sympatry in the Chafari-nas Islands differ in their feeding ecology and for-aging areas during both chick-rearing andwintering periods (Navarro et al. 2009).

The Cory’s Shearwater complex is best treatedas three full species, which should be listed in thefollowing sequence:

• Cory’s Shearwater Calonectris borealis(monotypic)

• Scopoli’s Shearwater Calonectris diomedea(monotypic)

• Cape Verde Shearwater Calonectris edwardsii(monotypic)

Cory’s and Scopoli’s Shearwaters are in Cate-gory A of the British List.

Madeiran Storm Petrel Oceanodromacastro

Recent studies based on morphology, vocalizationsand molecular phylogenetics suggest that MadeiranStorm Petrel represents a complex of species. Pop-ulations in the Azores which breed sympatricallybut in different seasons differ from each other inmorphology, have different calls, discriminateamong these calls, and further differ in mitochon-drial DNA (mtDNA) and microsatellite loci(Monteiro & Furness 1998, Bolton 2007, Friesenet al. 2007, Smith et al. 2007, Bolton et al. 2008,Robb et al. 2008). In addition, there are differ-ences in ecology and moult (Monteiro & Furness1998, Bolton et al. 2008). The concordance ofthese patterns of variation strongly indicates thatthese two populations have unique evolutionaryhistories, and are genetically and demographicallydistinct. The hot season breeding birds haverecently been named O. monteiroi (Bolton et al.2008). Storm petrels breeding on the Cape VerdeIslands have very different calls from those in theAzores, and are highly divergent in mtDNA (moreso than the sympatric O. monteiroi and O. castro,or indeed any other population in this complex).The evidence indicates that there are at least threepopulation lineages in the Western Palearcticwhich are best treated as species.

• Cape Verde Storm Petrel Oceanodromajabejabe (monotypic) (endemic to the CapeVerde Islands)

• Madeiran Storm Petrel Oceanodroma castro(monotypic) (hot-season breeders in Madeiraand the Selvagens; also provisionally includescool-season breeders in the Azores, Madeira,the Selvagens, and islands off Portugal)

• Monteiro’s Storm Petrel Oceanodromamonteiroi (monotypic) (hot-season breedersin the Azores)

Birds breeding in the cool season in Madeira andthe Selvagens (and also in the cool season in theAzores and islands off Portugal) are sympatric withhot-season breeders in Madeira (Nunes 2000) andthe Selvagens (Faria 1998), but differ from these

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in vocalizations (Robb et al. 2008) and morphol-ogy (Faria 1998, Nunes 2000). However, DNAdifferences are less clear than those among Azoreanpopulations (Friesen et al. 2007, Smith et al.2007). As a consequence, although potentially afourth species, this unnamed taxon is provisionallyincluded in O. castro pending further study andtaxonomic description.

Studies of vocalizations and phylogeographysuggest that there are multiple lineages outside theWestern Palearctic which should also be treated asspecies (Bolton 2007, Friesen et al. 2007, Smithet al. 2007).

There is a single British record of ‘MadeiranStorm Petrel’ sensu lato (off Isles of Scilly, 28July 2007; British Ornithologists’ Union (BOU)2012).

European Storm Petrel Hydrobatespelagicus

European Storm Petrel has been treated as amonotypic species. However, Mediterranean andAtlantic populations show mean morphometricdifferences and differences in vocalizations (Hém-ery & D’Elbée 1985, Bretagnolle 1992, Bret-agnolle & Zotier 1998, Lalanne et al. 2001, Robbet al. 2008). Phylogenetic analysis of mtDNAsequences has suggested that birds from Mediter-ranean and Atlantic populations are reciprocallymonophyletic with an estimated separation of lin-eages at least 350 000 years ago (Cagnon et al.2004). These differences strongly suggest thatMediterranean breeders should be recognizedtaxonomically. Pending ongoing analyses of vocali-zations, Mediterranean breeding birds are recogni-zed at subspecific level as H. p. melitensis.

Nominate H. p. pelagicus is in Category A ofthe British List.

Generic limits of Egretta

A study using an array of osteological characters(McCracken & Sheldon 1998) found Little BlueHeron Hydranassa caerulea and Tricoloured HeronH. tricolor to form a clade with Snowy EgretEgretta thula, Little Egret E. garzetta, White-facedHeron E. novaehollandiae and Pacific Reef HeronE. sacra. These results are supported by an analysisof mtDNA sequences (Sheldon et al. 2000). Con-sequently, no solid evidence exists for reciprocalmonophyly of ‘Hydranassa’ and ‘Egretta’ (sensu

Voous 1977). The genus ‘Hydranassa’ is thereforemerged with Egretta.

A revised taxonomic sequence ofCharadriiformes

In the early 1990s, molecular studies of relation-ships among Charadriiformes identified threemajor clades: plover-like birds (Charadrii), sand-piper-like birds (Scolopaci), and skuas, gulls, ternsand auks (Lari) (Sibley & Ahlquist 1990, Christianet al. 1992). A series of comprehensive molecularstudies published between 2003 and 2008 havecorroborated the existence of the three majorgroups identified by Sibley and Ahlquist (1990)and Christian et al. (1992) but also provide strongsupport for the relationships within and amongthese three groups (Ericson et al. 2003, Patonet al. 2003, Thomas et al. 2004a, Paton & Baker2006, 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 congru-ent support for the placement of Charadrii as thesister to (Scolopaci and Lari) (Ericson et al. 2003,Paton et al. 2003, Thomas et al. 2004a, Paton &Baker 2006, Baker et al. 2007, Fain & Houde2007), whereas a morphological study recoveredthese clades but placed Scolopaci as the sister-group of (Charadrii and Lari) (Mayr 2011, hisfig. 3C) and another morphological study did notrecover any of the three clades (Livezey 2010).

Some molecular studies suggested that the tun-dra plovers (Pluvialis) are sister to (Haematopodi-dae and Recurvirostridae) (Fain & Houde 2007) orto (Haematopodidae, Recurvirostridae and Chara-driidae) (Ericson et al. 2003, Baker et al. 2007).However, a more detailed analysis indicates thatthe tundra plovers are closely related to Charadrii-dae (Baker et al. 2012).

Within Scolopaci, (phalaropes and shanks),(turnstones and sandpipers), and (snipes, wood-cocks and dowitchers) form three well-supportedclades but the relationships among these are stillpoorly resolved (Ericson et al. 2003, Baker et al.2007, Gibson & Baker 2012). We have tentativelyfollowed Gibson and Baker (2012) because it isbased on the most extensive data set, in terms ofboth sequence data and species included.

The placement of the buttonquails (Turnicidae)in Charadriiformes, as the sister taxon of otherLari, is strongly supported (Paton et al. 2003,

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Paton & Baker 2006, Baker et al. 2007, Fain &Houde 2007, Hackett et al. 2008, but see Livezey2010). Most studies also support the placement ofGlareolidae 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 seeLivezey 2010). Molecular studies and one mor-phological study placed Crab Plover (Dromas) inLari (Hackett et al. 2008, Pereira & Baker 2010,Mayr 2011), whereas another morphological studyplaced it as the sister-group of the oystercatchers(Livezey 2010).

Most studies support a sister-relationship ofskuas and auks (Ericson et al. 2003, Paton et al.2003, Thomas et al. 2004a, Paton & Baker 2006,Baker et al. 2007, Fain & Houde 2007, Smith2011, but see Livezey 2010). A close relationshipbetween skimmers, terns and gulls is very stronglysupported, although exact relationships amongthese three groups are not yet clear (Ericson et al.2003, Paton et al. 2003, Paton & Baker 2006,Baker et al. 2007, Fain & Houde 2007, Livezey2010, Mayr 2011, Smith 2011). Several studiessupport a clade formed by Alca, Pinguinus, Alleand Uria, although except for a sister-group rela-tionship of Alca and Pinguinus, relationshipswithin this clade remain unresolved (Baker et al.2007, Pereira & Baker 2008, Smith 2011).

The sequence of taxa within Charadriiformes isbased on the principle that for each sister-groupthe less speciose taxon is listed first. The sequenceand generic limits of the shanks, terns and gullsare as previously defined (Sangster et al. 2005,2007). The taxonomic sequence of Alcidae isbased on phylogenetic studies by Baker et al.(2007) and Pereira and Baker (2008). The taxa onthe Western Palearctic list should be listed in thefollowing sequence:

CharadriiBurhinidae (Burhinus)Pluvianidae (Pluvianus)Recurvirostridae (Himantopus, Recurvirostra)Haematopodidae (Haematopus)Charadriidae (Pluvialis, Vanellus, Charadrius)

ScolopaciRostratulidae (Rostratula)Scolopacidae (Bartramia, Numenius, Limosa,Arenaria, Calidris, Phalaropus, Xenus, Actitis,Tringa, Lymnocryptes, Limnodromus, Scolopax,Gallinago)

LariTurnicidae (Turnix)Dromadidae (Dromas)Glareolidae (Glareola, Cursorius)Stercorariidae (Stercorarius)Alcidae (Fratercula, Aethia, Brachyramphus,Cepphus, Synthliboramphus, Alca, Pinguinus,Alle, Uria)Sternidae (Anous, Onychoprion, Sternula, Gelo-chelidon, Hydroprogne, Chlidonias, Sterna)Rynchopidae (Rynchops)Laridae (Pagophila, Xema, Rissa, Chroicocepha-lus, Hydrocoloeus, Rhodostethia, Larus)

Generic arrangement of calidrinesandpipers

A recent molecular phylogeny based on mito-chondrial and nuclear DNA sequences providesstrong evidence that the genus Calidris is para-phyletic, and that five monotypic genera are partof the Calidris clade (Aphriza, Philomachus,Limicola, Eurynorhynchus, Tryngites) (Gibson &Baker 2012).

We have considered two revisions of the cali-drine sandpipers: (1) include Aphriza, Philomachus,Limicola, Eurynorhynchus and Tryngites in Calidris,or, alternatively, (2) maintain some of these generaand split Calidris into several genera, e.g. Bahr(2011). Note that Bahr’s (2011) proposed generaEurynorhynchus and Tryngites cannot be main-tained as such in the light of the latest molecularresults since Eurynorhynchus was nested in Bahr’sLeimonites, and Tryngites was nested in his Ereune-tes (Gibson & Baker 2012).

We prefer to include Aphriza, Philomachus,Limicola, Eurynorhynchus and Tryngites in Calid-ris, rather than subdivide the last-named into anumber of genera, based on the following con-siderations. First, since some of the clades inGibson 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 mono-typic genera. Third, splitting Calidris wouldchange the names of many species, and only twospecies currently in Calidris would remain inthat genus (i.e. the two knots, since the typespecies of Calidris is C. canutus). In contrast, ifCalidris were expanded to include the five cur-rently recognized monotypic genera, only fivespecific names would be changed and all species

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878 G. Sangster et al.

currently in Calidris would remain in it. Finally,even if Calidris were subdivided into a numberof genera, the newly proposed polytypic generawould still be rather heterogeneous morphologi-cally whereas no major morphological or ecologi-cal differences are known to characterize thesegenera.

The names and taxonomic sequence of the cali-drine sandpipers become as follows (species on theBritish List are indicated by their category):

• Great Knot Calidris tenuirostris (Cat. A)• Red Knot Calidris canutus (Cat. A)

• Surfbird Calidris virgata• Ruff Calidris pugnax (Cat. A)

• Sharp-tailed Sandpiper Calidris acuminata(Cat. A)

• Broad-billed Sandpiper Calidris falcinellus(Cat. A)

• Curlew Sandpiper Calidris ferruginea (Cat. A)• Stilt Sandpiper Calidris himantopus (Cat. A)

• Spoon-billed Sandpiper Calidris pygmea• Red-necked Stint Calidris ruficollis (Cat. A)

• Long-toed Stint Calidris subminuta (Cat. A)

• Temminck’s Stint Calidris temminckii (Cat. A)

• Sanderling Calidris alba (Cat. A)

• Dunlin Calidris alpina (Cat. A)

• Purple Sandpiper Calidris maritima (Cat. A)• Rock Sandpiper Calidris ptilocnemis• Baird’s Sandpiper Calidris bairdii (Cat. A)

• Little Stint Calidris minuta (Cat. A)

• White-rumped Sandpiper Calidris fuscicollis(Cat. A)

• Least Sandpiper Calidris minutilla (Cat. A)

• Buff-breasted Sandpiper Calidris subruficollis(Cat. A)

• Pectoral Sandpiper Calidris melanotos (Cat. A)

• Western Sandpiper Calidris mauri (Cat. A)

• Semipalmated Sandpiper Calidris pusilla(Cat. A)

Cream-coloured Courser Cursoriuscursor

Populations of Cream-coloured Courser in Ethio-pia, Somalia and northern Kenya (littoralis and so-malensis) differ diagnosably from those elsewherein Africa and Asia in tail pattern and underwingcoloration in adults, in the pattern of the body andtail in juveniles, and in size and proportions(Pearson & Ash 1996). Based on this evidence,two species are recognized:

• Cream-coloured Courser Cursorius cursor(polytypic, with subspecies exsul, cursor andbogolubovi)

• Somali Courser Cursorius somalensis (poly-typic, with subspecies littoralis and somalensis)

Cream-coloured Courser is in Category A ofthe British List.

Arctic Warbler Phylloscopus borealis

Molecular studies have identified three stronglydivergent groups within the Arctic Warbler com-plex (Saitoh et al. 2010, see also Reeves et al.2008). These three groups also differ in morpho-metrics, territorial songs and calls (Saitoh et al.2008, Martens 2010, Alström et al. 2011), andare best treated as separate species. Severalnames could potentially be used within thisgroup. The type specimens of sylvicultrix Swin-hoe 1860 and hylebata Swinhoe 1861 could notbe located by Alström et al. (2011) and the iden-tity of these types is unclear, but the identitiesof the type specimens of flavescens Gray 1860,examinandus Stresemann 1913, and xanthodryasSwinhoe 1863 have been resolved, which hasenabled names to be applied to the three species(Alström et al. 2011). The taxon kennicotti is besttreated as a synonym of borealis (Alström et al.2011). Consequently, three monotypic speciesare recognized.

• Arctic Warbler Phylloscopus borealis (mono-typic)

• Kamchatka Leaf Warbler Phylloscopus exam-inandus (monotypic)

• Japanese Leaf Warbler Phylloscopus xantho-dryas (monotypic)

Arctic Warbler is in Category A of the BritishList.

Marmora’s Warbler Sylvia sarda

Two studies based on mitochondrial gene sequencessuggest that Marmora’s Warbler Sylvia s. sarda andBalearic Warbler Sylvia s. balearica are not neces-sarily sister-taxa (Böhning-Gaese et al. 2003, Voel-ker & Light 2011). Instead, balearica is inferredeither as sister to Dartford Warbler S. undata, withsarda as their sister-taxon (Voelker & Light 2011),or both sarda and balearica are equally divergentfrom a clade comprising S. undata and Tristram’s

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Taxonomic recommendations for British birds: eighth report 879

Warbler S. deserticola (Böhning-Gaese et al. 2003).Precise relationships among these species are stillunresolved, but these studies, albeit inferred from asingle locus and very small samples, strongly sup-port a lengthy divergence of sarda and balearica(around 2.5 million years ago, Voelker & Light2011). There are minor but apparently consistentdifferences in bare-part and plumage colorationbetween balearica and sarda: breast and throat ofadult males are near-concolorous in sarda, whereasthe paler throat contrasts with the breast in baleari-ca; basal parts of the bill are pinkish in sarda, orangein balearica; and leg coloration is darker andbrowner in most first-winter and juvenile sarda,pale orange to orange-brown in balearica (Shirihaiet al. 2001). In addition, there are distinct differ-ences in contact calls and territorial song (Shirihaiet al. 2001). A fuller published description of thevocal differences would be valuable, but the com-bined morphological, vocal and molecular data sug-gest that Marmora’s Warbler is best treated as twomonotypic species.

• Balearic Warbler Sylvia balearica (mono-typic)

• Marmora’s Warbler Sylvia sarda (monotypic)

There are six accepted British records of Mar-mora’s Warbler sensu lato. None was formallyassigned to subspecies level, and all records shouldbe reviewed.

Eurasian Nuthatch Sitta europaea

Recent studies have shown that the northeastSiberian form arctica differs from other popula-tions of Sitta europaea in plumage, size andproportions (Red’kin & Konovalova 2006), vocali-zations (Leonovich et al. 1996) and mtDNAsequences (10% sequence divergence in ND2,Zink et al. 2006). The two groups have overlap-ping ranges in eastern Siberia, but only one puta-tive intermediate specimen has been reported inthe literature and no further intermediates werelocated in museums (Red’kin & Konovalova2006). The combined evidence indicates that Sittaarctica represents a distinct species (Red’kin &Konovalova 2006).

• Sitta arctica Siberian Nuthatch (monotypic)

• Sitta europaea Eurasian Nuthatch (polytypic)

Phylogeographical analyses (Zink et al. 2006,Hung et al. 2012) and reports of reduced intro-

gression in contact zones (Red’kin & Konoval-ova 2006) indicate that Eurasian Nuthatch mayinclude additional species but further study is nec-essary.

Eurasian Nuthatch is in Category A of the Brit-ish List.

We thank Vicki L. Friesen, P.J. Moore and MagnusRobb for their help, and Lars Svensson for extensivecomments on some of the taxa included in this report.

REFERENCES

Alström, P., Saitoh, T., Williams, D., Nishiumi, I., Shigeta,Y., Ueda, K., Irestedt, M., Björklund, M. & Olsson, U.2011. The Arctic Warbler Phylloscopus borealis—threeanciently separated cryptic species revealed. Ibis 153: 395–410.

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