Taxonomic Recommendations for British Birds: Fourth Report

Ibis (2007), 149, 853–857

Blackwell Publishing Ltd Taxonomic recommendations for British birds: Fourth report

GEORGE SANGSTER,1* J. MARTIN COLLINSON,2 ALAN G. KNOX,3 DAVID T. PARKIN4 & LARS SVENSSON5 1Stevenshof 17, 2312 GM Leiden, The Netherlands 2Biomedical Sciences, Institute of Medical Sciences, University of Aberdeen, Aberdeen AB25 2ZD, UK 3Historic Collections, King’s College, University of Aberdeen, Aberdeen AB24 3SW, UK 4Institute of Genetics, University of Nottingham, Queen’s Medical Centre, Nottingham NG7 2UH, UK 5S:ta Toras väg 28, S-260 93 Torekov, Sweden

This paper is the fourth report of the Taxonomic 149: 1–95) supported a sister relationship between Sub-Committee of the BOU Records Committee divers and grebes. This latter paper contained several relating to the British List. Species-level decisions hypotheses about higher order avian relationships are based on criteria outlined by Helbig et al. that are not supported by other highly congruent (2002, Guidelines for assigning species rank; Ibis studies, and it seems likely that any apparent close 144: 518–525). The third report of the Sub-Committee relationship between grebes and divers is due to con- was published in Ibis 147: 821–826. vergence. Recognition of the flamingo-grebe clade is therefore recommended. This clade was recently named Mirandornithes (Sangster 2005. Ibis 147: 612–615) and Greater Scaup Aythya marila is placed between Ciconiiformes and Falconiformes, In our previous report we referred to the name with the flamingos listed before the grebes. mariloides as being unavailable (Ibis 147: 821–826). This is incorrect; the name is inapplicable. American Bittern Botaurus lentiginosus The author of the scientific name for the American The position of grebes and flamingos Bittern was changed without explanation in the 6th Phylogenetic analyses based on DNA-DNA hybrid- Checklist (BOU 1992. Checklist of the Birds of Britain ization data (van Tuinen et al. 2001. Proc. R. Soc. Lond. and Ireland, 6th edn.) from Montagu to Rackett. For B 268: 1345–1350), mitochondrial and nuclear DNA clarification, both descriptions were based on the sequences (van Tuinen et al. 2001; Chubb 2004. same specimen, but Rackett’s publication pre-dated Mol. Phylogen. Evol. 30: 140–151; Cracraft et al. 2004. that of Montagu by several weeks and thereby takes In: Cracraft & Donoghue, Reconstructing the Tree of precedence (Macdonald & Grant 1951. Bull. Brit. Orn. Life, pp. 468–489; Ericson et al. 2006. Biol. Lett. 2: Club 71: 30). 543–547) and morphology (Mayr & Clarke 2003. Cladistics 19: 527–553; Mayr 2004. Zool. J. Linn. Soc. Spanish Imperial Eagle Aquila adalberti 140: 157–169; Manegold 2006. Acta Ornithol. 41: 79–82) provide congruent support for a sister-group The scientific name of this species is as given above relationship of flamingos Phoenicopteriformes and (cf. Ibis 147: 821–826). grebes Podicipediformes. Storer (2006. Auk 123: 1183– 1184) challenged some of the morphological evi- Allen’s Gallinule Porphyrula alleni dence for the flamingo-grebe grouping, and a recent Purple Gallinule Porphyrula martinica comprehensive phylogeny based on morphological characters (Livezey & Zusi 2007. Zool. J. Linn. Soc. The purple gallinules Porphyrula, Porphyrio, and Notornis constitute a monophyletic group. Porphyrula differs from Porphyrio in its smaller size, less massive *Corresponding author. bill and more oval nostril but the two genera share a Email: [email protected] number of characters, particularly of the hindlimb

854 G. Sangster et al.

(Olson 1973. Wilson Bull. 85: 381–416). Porphyrula • Common Greenshank Tringa nebularia (i.e. P. alleni, P. martinica, P. flavirostris) forms a • Lesser Yellowlegs Tringa flavipes monophyletic group which is sister to Porphyrio/ • Marsh Sandpiper Tringa stagnatilis Notornis, which also forms a monophyletic group • Wood Sandpiper Tringa glareola (Livezey 1998. Philos. Tr. R. Soc. Lond. B 353: 2077– • Common Redshank Tringa totanus 2151). Both the continued recognition of Porphyrula and Porphyrio as separate genera and the inclusion of Herring Gull Larus argentatus, Porphyrula in Porphyrio are potentially consistent Yellow-legged Gull L. michahellis, with their evolutionary relationships. However in Armenian Gull L. armenicus and light of the striking morphological similarities Lesser Black-backed Gull L. fuscus between the species in these genera there is growing international support for the inclusion of Porphyrula The recommendation that Yellow-legged Gull L. in Porphyrio. This treatment has been adopted in michahellis and Armenian Gull L. armenicus should major checklists and handbooks (e.g. Urban et al. be treated as separate species from Herring Gull 1986. The Birds of Africa 2; del Hoyo et al. 1996. L. argentatus was communicated previously (Sangster Handbook of the Birds of the World 3; Taylor & van et al. 2005. Ibis 147: 821–826). Phylogenetic evidence Perlo 1998. Rails; Banks et al. 2002. Auk 119: based on analysis of mitochondrial DNA (mtDNA) 897–906; Dickinson 2003. The Howard and Moore sequences indicates that the large white-headed gull Complete Checklist of the Birds of the World). It is complex consists of two main clades: (1) an ‘Atlantic’ recommended that Porphyrula be treated as a sub- clade consisting of Yellow-legged Gull, Armenian Gull, genus within Porphyrio. most individuals of Herring Gull, and including Allen’s Gallinule and Purple Gallinule are on the Great Black-backed Gull L. marinus and Palearctic British List and become Porphyrio alleni and Porphyrio individuals of Glaucous Gull L. hyperboreus; (2) an martinica, respectively. ‘Aralo-Caspian’ clade consisting of L. a. cachinnans, L. a. barabensis, L. a. heuglini, L. a. taimyrensis, Lesser Black-backed Gull L. fuscus, Kelp Gull L. dominicanus, Grey-tailed Tattler Heteroscelus brevipes some individuals of L. a. argentatus, and including Molecular phylogenetic analysis (Pereira & Baker 2005. an ‘Arctic/Pacific’ grouping of L. a. vegae, L. a. Condor 107: 514–526) groups two species traditionally smithsonianus, L. a. mongolicus, Slaty-backed Gull placed in the genus Heteroscelus (i.e. Grey-tailed Tattler L. schistisagus, Iceland Gull L. glaucoides, Glaucous- H. brevipes and Wandering Tattler H. incanus) and Willet winged Gull L. glaucescens and Nearctic individuals Catoptrophorus semipalmatus in the Tringa clade. These of Glaucous Gull (Crochet et al. 2002. Auk 119: data suggest that the tattlers and Willet should be 603–620; Crochet et al. 2003. Evolution 57: 2865– reclassified in the genus Tringa. Grey-tailed Tattler is 2878; Liebers et al. 2004. Proc. R. Soc. Lond. B 271: on the British List and becomes Tringa brevipes. 893–901). Western Gull L. occidentalis is not included in these clades and forms an outgroup. Genetic structure between the taxa is generally Taxonomic sequence of shanks well-defined, except within the Arctic/Pacific group- Phylogenetic analyses of mitochondrial and nuclear ing of the Aralo-Caspian clade, for which the taxa DNA sequences and morphology (Pereira & Baker are poorly separated with some shared haplotypes 2005. Condor 107: 514–526) have clarified the (Gay et al. 2005. Auk 122: 684–688). evolutionary relationships among the shanks. The species Morphological and genetic variation in Lesser in the genera Xenus, Actitis and Tringa should be listed Black-backed Gulls L. fuscus fuscus, L. f. intermedius in the following sequence: and L. f. graellsii is clinal (Liebers & Helbig 2002. • Terek Sandpiper Xenus cinereus J. Evol. Biol. 15: 1021–1033). These taxa are closely • Common Sandpiper Actitis hypoleucos related to the West Siberian taxa L. a. heuglini, L. a. • Spotted Sandpiper Actitis macularius taimyrensis and L. a. barabensis with evidence of • Green Sandpiper Tringa ochropus continuing low levels of gene flow in spite of apparent • Solitary Sandpiper Tringa solitaria ecological separation of L. f. fuscus and L. a. heuglini • Grey-tailed Tattler Tringa brevipes in parapatry (Filchagov et al. 1992a. Zool. Zh. 71: • Spotted Redshank Tringa erythropus 148–152; Rauste 1999. Limicola 13: 105–128; 153– • Greater Yellowlegs Tringa melanoleuca 188; Liebers & Helbig 2002; Liebers et al. 2004).

It is recommended that these taxa be treated as a 2004, Gay et al. 2005). In addition, European Herring single species L. fuscus. Variation in heuglini is slight Gulls are reported to respond poorly to calls of smith- (Buzun 2002. Br. Birds 95: 216–232) and the validity sonianus (Frings et al. 1958. Ecology 39: 126–131). of taimyrensis has been questioned (Filchagov et al. Based on current evidence, American Herring Gull 1992b. L’Oiseau 62: 128–148; Yésou 2002. Dutch L. smithsonianus is best treated as a separate species. Birding 64: 271–298). Given their lack of diagnostic differences, vegae Caspian Gull L. a. cachinnans is diagnosably and mongolicus are considered conspecific with L. distinct from all other taxa on the basis of plumage smithsonianus. and vocalisations (Panov et al. 1991a. Zool. Zh. 70/1: We recommend recognition of the following species 76–90; Panov et al. 1991b. Zool. Zh. 70/3: 73–89; to better reflect recent advances in knowledge of the Garner & Quinn 1997. Br. Birds 90: 25–62; Klein & evolution and systematics of large gulls: Gruber 1997. Limicola 11: 49-75; Liebers & Dierschke • Caspian Gull Larus cachinnans (monotypic) 1997. Dutch Birding 19: 277–280; Jonsson 1998. Alula • Lesser Black-backed Gull Larus fuscus (polytypic, 3: 74–100; Yésou 2002). It forms a discrete genetic including fuscus, intermedius, graellsii, heuglini, grouping and is probably most closely related to the taimyrensis, barabensis) West Siberian gulls heuglini and barabensis. How- • American Herring Gull Larus smithsonianus (poly- ever, introgression between cachinnans and the West typic, including smithsonianus, vegae, mongolicus) Siberian taxa is restricted and probably unidirectional • Yellow-legged Gull Larus michahellis (polytypic, (Panov & Monzikov 2000. Br. Birds 93: 227–241; including michahellis, atlantis) Liebers et al. 2004). Recent range expansion has led • Armenian Gull Larus armenicus (monotypic) to widespread hybridization with argentatus Herring • Herring Gull Larus argentatus (polytypic, including Gulls in Central and Eastern Europe which may argentatus, argenteus) partly underlie the apparently large intrataxon A manuscript explaining these decisions has been variation in cachinnans (Klein & Gruber 1997. Limicola submitted to British Birds. 11: 49–75; Liebers & Dierschke 1997; Panov & Monzikov 1999. Russ. J. Zool. 3: 129–141). However Generic arrangement of gulls the hybrid zone is reported to be narrow in relation to the potential dispersal distance of the parental Two research programmes, one based on morphology taxa (Neubauer et al. 2006. Vogelwelt 127: 11–22; (Chu 1998. Cladistics 14: 1–43) and another based Yakovets 2006. Vogelwelt 127: 23–30). Based on primarily on mitochondrial DNA sequences (Crochet this evidence, it is recommended that Caspian Gull et al. 2000. J. Evol. Biol. 13: 47–57; Pons et al. 2005. be treated as a monotypic species L. cachinnans. Mol. Phylogen. Evol. 37: 686–699) have examined Genetic evidence suggests that East Siberian and phylogenetic relationships of the entire group of American Herring Gulls, L. a. vegae, L. a. mongolicus gulls. Both studies indicate that the genus Larus, as and L. a. smithsonianus fall within the Arctic/Pacific currently defined [e.g. Voous 1977. List of Recent species group of the Aralo-Caspian clade (Liebers Holarctic Bird Species. Br. Ornithol. Union, London; et al. 2004; de Knijff et al. 2005. Birding 37: 402– Cramp & Simmons 1983. The Birds of the Western 411; Gay et al. 2005). There are sharp genetic and Palearctic. Vol. 3. Oxford UP, Oxford; Burger & morphological boundaries between vegae and the Gochfield 1996. Gulls. In: del Hoyo, J., Elliot, A. & parapatric or partially sympatric West Siberian taxon Sargatal, J. (eds.) Handbook of the Birds of the World. heuglini (Yésou 1994. Alauda 62: 247-252; Liebers Vol. 3. Hoatzin to Auks. Lynx Edicions, Barcelona], et al. 2004; Yésou 2002). Many individuals of vegae, is not monophyletic. These studies indicate that the mongolicus and smithsonianus can be identified to generic limits of the gulls need revision. subspecies (Lonergan & Mullarney 2004. Dutch Bird- Both studies indicated a separate position of Creagrus, ing 26: 1–35), but diagnosability with respect to each Rissa, Xema, Pagophila from all other gulls, supporting other has not been demonstrated. Although many the continued recognition of these genera, but also individuals can be identified, smithsonianus is not supported another well-defined clade (which includes fully diagnosable from European Herring Gulls on Slender-billed Gull Larus genei, Bonaparte’s Gull the basis of plumage alone (Lonergan & Mullarney L. philadelphia and Black-headed Gull L. ridibundus) 2004; Adriaens & Mactavish 2004. Dutch Birding 26: which is not the sister group to other gulls in Larus 151–179) but is clearly differentiated on the basis of as currently defined. Both studies also resolved a mtDNA (Crochet et al. 2002, 2003, Liebers et al. sister-group relationship of Ross’s Gull and Little

Gull and their separate position from the main clade The Chroicocephalus gull clade is strongly supported of gulls. by congruency between Pons et al. (2005) and the A taxonomic revision that would maintain mono- results of Chu (1998). The groups designated as phyly of gull genera with minimum change to the ‘hooded’ and ‘black-headed’ gulls by Pons et al. (2005) British List would be to include all gulls in Larus, and which may merit generic status were not found except those currently placed in Creagrus, Rissa, Xema by Chu (1998), so further division within Larus s.s. and Pagophila. This arrangement however does not is not recommended at this stage. reflect the taxonomic information derived from the cited studies. Pons et al. (2005) in contrast suggested Aleutian Tern Onychoprion aleutica recognition of ten genera, requiring adoption of five Sooty Tern O. fuscata new generic names: Chroicocephalus (which includes Slender-billed Gull, Bonaparte’s Gull and Black- The gender of the name Onychoprion is masculine headed Gull); Saundersilarus (Saunder’s Gull); and the correct names for these taxa are: Hydrocoloeus (Ross’s and Little Gull); Leucophaeus • Aleutian Tern Onychoprion aleuticus (some of the New World ‘hooded’ gulls); Ichthyaetus • Sooty Tern Onychoprion fuscatus (southern Palaearctic ‘black-headed gulls’). Four of these putative genera have been recorded in Marbled Murrelet Brachyramphus Britain. Retention of separate genera for Little and marmoratus Ross’s Gull would be justifiable on the basis of the long branch lengths separating the two, which are Brachyramphus m. marmoratus and B. m. perdix comparable with those separating Xema and show differences in plumage [Konyukhov & Kitaysky Pagophila. 1995. In: Ralph et al. (eds), USDA Forest Service For the purposes of the British List, the TSC General Technical Report PSW-GTR-152, pp. recommends recognition of an intermediate taxonomy 23–29; Gaston & Jones 1998. The Auks], mtDNA that adopts Chroicocephalus for the clade which restriction fragment lengths (Zink et al. 1995. includes Slender-billed Gull, Bonaparte’s Gull and Condor 97: 639–649) and allozymes and mtDNA Black-headed Gull and adopts Hydrocoloeus for sequences (Friesen et al. 1996. Condor 98: 681–690); Little Gull, but does not change the generic status of allozymes and mtDNA sequences suggest that other gull species, as follows: marmoratus is more closely related to Kittlitz’s Murrelet • Ivory Gull Pagophila eburnea B. brevirostris than to perdix. B. marmoratus is best • Sabine’s Gull Xema sabini treated as two species: • Black-legged Kittiwake Rissa tridactyla • Marbled Murrelet B. marmoratus (monotypic) • Slender-billed Gull Chroicocephalus genei • Long-billed Murrelet B. perdix (monotypic) • Bonaparte’s Gull Chroicocephalus philadelphia • Black-headed Gull Chroicocephalus ridibundus Belted Kingfisher Ceryle alcyon • Little Gull Hydrocoloeus minutus • Ross’s Gull Rhodostethia rosea A recently-published phylogenetic analysis of the • Laughing Gull Larus atricilla kingfishers indicates that Ceryle rudis is the sister of • Franklin’s Gull Larus pipixcan Chloroceryle and is not closely related to Ceryle • Mediterranean Gull Larus melanocephalus alcyon (Moyle 2006. Auk 123: 487–499). This indi- • Audouin’s Gull Larus audouinii cates that the current treatment of Megaceryle as a • Pallas’s Gull Larus ichthyaetus subgenus of Ceryle does not accurately reflect their • Mew Gull Larus canus phylogenetic relationships. In view of their distinc- • Ring-billed Gull Larus delawarensis tive morphology and to avoid paraphyly of Ceryle, • Lesser Black-backed Gull Larus fuscus three genera of ceryline kingfishers are recognized: • Herring Gull Larus argentatus Megaceryle, Ceryle and Chloroceryle. Both Miller • Yellow-legged Gull Larus michahellis (1912. Bull. Am. Mus. Nat. Hist. 31: 239–311; 1920. • Caspian Gull Larus cachinnans Auk 37: 422–429) and Fry (1980. Living Bird 18: • American Herring Gull Larus smithsonianus 113–160) emphasized anatomical differences • Iceland Gull Larus glaucoides among the three groups in support for treatment as • Glaucous Gull Larus hyperboreus three genera. These data suggest that Belted King- • Great Black-backed Gull Larus marinus fisher should be reclassified in the genus Megaceryle.

Belted Kingﬁsher is on the British List and becomes includes Catharus, Hylocichla, Cichlopsis, Entomodestes Megaceryle alcyon. and Ridgwayia). On the basis of this evidence, Varied Thrush is reinstated in a monotypic genus and becomes Ixoreus naevius. Red-rumped Swallow Cecropis daurica The author of the scientiﬁc name is Laxmann Willow Tit Poecile montanus (Dickinson & Ericson 2002. Zool. Verh. 340: 205– 206). The gender of the name Poecile is feminine (not as given in Ibis 147: 821–826). Therefore, the correct name of Willow Tit is Poecile montana. Varied Thrush Zoothera naevia

Phylogenetic analyses of mitochondrial DNA sequences Acknowledgements (Klicka et al. 2005. Mol. Phylogen. Evol. 34: 486–500) indicate that Varied Thrush does not group with We thank Dick Banks, Edward Dickinson and Laurent Raty African and Asian Zoothera thrushes but is part of for their help and apologize to any others that we have a New World radiation of thrushes (which also omitted to mention.