The ‘soft-plumaged petrel’ complex: a review of the literature on taxonomy, identification and distribution Andrew H. J. Harrop

ABSTRACT This paper reviews the literature concerning the taxonomy, identification and distribution of the ‘soft-plumaged petrel’ Pterodroma feae/madeira/mollis complex in the Western Palearctic.There are no known consistent plumage differences between feae and madeira and, at present, only bill structure can be regarded as diagnostic.A number of consistent differences exist between mollis and feae/madeira which make field identification of mollis possible.To date, all well-documented records of ‘soft-plumaged petrels’ away from breeding grounds in the North Atlantic are compatible with feae; there are no safe records of mollis in the northern hemisphere, or of madeira away from its breeding grounds.

wo species of petrel of the genus Ptero- History and taxonomy droma breed in the Western Palearctic: Archaeological remains found in two cave sites TFea’s Petrel P. feae, with the nominate in Gibraltar are clearly identifiable as those of form on Cape Verde and the subspecies P. f. Pterodroma petrels (Cooper 1999). The approx- deserta occurring on Bugio, in the Madeiran imate age range of the majority of these speci- archipelago; and Zino’s Petrel P. madeira, in the mens is between 60,000 and 25,000 years BP, mountains of Madeira (table 1). In addition, although none of the specimens has been dated Soft-plumaged Petrel P. mollis may occur as a precisely. The remains fall into two distinct size vagrant from the South Atlantic or Indian groupings: a small type, similar in size to Ocean. These three closely similar species, here- madeira, and the more abundant of the two; after referred to simply as feae, madeira and and a larger but less numerous type. This evi- mollis, have often been described collectively as dence suggests the presence of two species, but ‘soft-plumaged petrels’. Rare and genuinely might possibly represent the size range of one enigmatic, the Pterodroma petrels of the eastern species (Jo Cooper in litt.). These remains are North Atlantic have both excited birdwatchers believed to be the oldest examples of the genus and generated a substantial body of literature, in the Western Palearctic, and are unusual in especially in the past 20 years. Nonetheless, that they originate from continental sites, some aspects of their taxonomy, identification although it is uncertain whether they represent and distribution remain controversial. This the site of a former breeding colony, or are the paper provides a critical review of the literature result of a seabird wreck. They do suggest, to date, in particular that relating to field identi- however, that members of the genus were for- fication. merly more widespread in the region than they

6© British Birds 97 • January 2004 • 6-15 The ‘soft-plumaged petrel’ complex

Table 1. Scientific and common names, breeding distributions and population sizes of the two Pterodroma petrels which breed in the Western Palearctic. Population sizes are from Snow & Perrins (1998) for P. feae and from Jorge Garzon (in litt.) for P. madeira.

Scientific name Pterodroma feae Pterodroma madeira

Common names Fea’s Petrel Zino’s Petrel Cape Verde Petrel Madeira Petrel Gon-gon Freira Breeding distribution Desertas, Madeira Islands Madeira highlands Cape Verde Population size Desertas: 150-200 pairs 45 pairs Cape Verde: 500-1,000 pairs have been in the recent past. plumaged Petrel, showing as the latter does an As noted by Bourne (1983a), the first ‘soft- overlap in its variation in colour’. This treat- plumaged petrel’ recorded in historical times ment was followed by Imber (1985), though he appears to have been collected off the coast of did not provide any additional information in West Africa in October 1768, during Captain support of this split, other than noting that the Cook’s first voyage. No description was pub- species of Halipeurus louse supported by feae lished at the time, but an excellent drawing of differed from that found on mollis. Zino & Zino the specimen by Sydney Parkinson (reproduced (1986) provided a thorough account of the bio- in Lysaght 1959) is recognisable as the form metric differences of the two forms within the breeding locally in the Cape Verde Islands. This Madeiran archipelago. Despite this, and without was described subsequently as a distinct species, giving any reasons, Enticott (1991) continued to Oestrelata feae, by Salvadori (1899), who later treat all ‘soft-plumaged petrels’ as a single published comparisons between feae and mollis, species. Zonfrillo (1993), who studied the noting that the southern birds (mollis) are feather lice of the North Atlantic Pterodroma smaller, have a grey pectoral band, and have the species, inferred sister-taxon relationships sides of the body less freckled with grey (Sal- between Jamaican Petrel P. hasitata caribbaea vadori 1900). and feae, and between Bermuda Petrel P. cahow In 1934, Mathews described two new forms and madeira. which he labelled ‘soft-plumaged petrel’: P. m. Bretagnolle (1995) further complicated madeira from Madeira and P. m. deserta from matters by suggesting, on the basis of multi- the Desertas Islands in the Madeiran archi- variate statistical analysis of morphological pelago (Mathews 1934a,b). Subsequently, Ban- (biometrics and coloration) and behavioural nerman & Bannerman (1965) described characters (vocalisations) of the six populations madeira in greater detail, and made a plea for within the ‘soft-plumaged petrel’ grouping, that the continued recognition of deserta as a valid the complex should be split into two species, subspecies, despite acknowledging that it was one breeding in each hemisphere. His interpre- difficult to separate from feae. Other authors, tation of the data was challenged by Knox including Bourne (1957), Jouanin et al. (1969), (1995), who argued that there were significant and Cramp & Simmons (1977), considered differences between the calls of madeira and deserta inseparable from feae. deserta. Meanwhile, Hazevoet (1995) estab- Bourne (1957) noted that two forms of ‘soft- lished that feae breeds exclusively in the higher plumaged petrel’ bred within the Madeiran parts of four of the main islands in the Cape archipelago at different seasons, but felt that ‘it Verde group, while those from the Desertas would defeat the whole purpose of classification breed on small, low islets, close to sea level. to regard these closely related forms as distinct Hazevoet (1997), citing Nunn & Zino (in species’. Later, however, he reversed his earlier prep.), stated that phylogenetic studies of Ptero- view and advocated treating mollis, feae and droma petrels using mitochondrial cytochrome madeira binomially, as distinct species (Bourne b gene sequences showed that it is no longer 1983a), since ‘it is difficult to say which of the tenable to assign feae and madeira to the two [feae and madeira] is closer to the Soft- ‘P. mollis species group’. Unfortunately, in their

British Birds 97 • January 2004 • 6-15 7 The ‘soft-plumaged petrel’ complex

P. m. mollis

P. madeira

P. m. mollis

P. feae

P. madeira P. feae Ian Lewington Fig. 1. Bill structure of members of the ‘soft-plumaged petrel’ Pterodroma feae/madeira/mollis complex. Note that most mollis are like the uppermost example.

P. m. mollisP. madeira P. feae P. m. mollis (dark morph) Ian Lewington Fig. 2. Head pattern of members of the ‘soft-plumaged petrel’ Pterodroma feae/madeira/mollis complex.

study, Nunn & Zino used a sample of deserta is under review. Ratcliffe et al. (2000) discussed but did not include feae from Cape Verde, so the the relationship between nominate feae from taxonomic status and phylogenetic relationship Cape Verde and deserta. Based upon differences between birds from the Desertas and Cape in their morphometrics, their different breeding Verde remains unresolved. phenology (laying in December-January on Nunn & Stanley (1998) analysed cytochrome Cape Verde, July-August on Desertas), and the b sequences from 85 species of Procellari- distance between breeding sites, they suggested iformes, including mollis and feae, and found that the two taxa are effectively reproductively that these two taxa do not even share a sister- isolated through philopatry and ‘are probably’ taxon relationship. The following year, the ABA cryptic species. (A ‘cryptic species’ is one that is Checklist Committee (1999) accepted Fea’s/ genetically distinct but which cannot be distin- Zino’s Petrel as new for their recording area. At guished readily from congeners using tradi- present, the BOU regards all British records as tional morphological characters alone.) referable to ‘Soft-plumaged Petrel’ Pterodroma Sangster et al. (2002) recommended that mollis/madeira/feae (BOU 2000), although this feae, madeira and mollis are best treated as

8 British Birds 97 • January 2004 • 6-15 The ‘soft-plumaged petrel’ complex

Pterodromas from above illustrating P. m. mollis true tonal values in flat light (dark morph) mollis feae

madeira

Although the upperparts of some dark mollis are similar to those of feae, the under- parts are strikingly different

mollis in flat light

mollis madeira

feae feae in strong light

feae in strong madeira in strong light light with apparent breast-band

Pale mollis with faint breast-band

Tw o feae/madeira in calm conditions Five feae/madeira showing less angled wings illustrating angled wings with blunter tips with pointed tips in windy conditions Ian Lewington Fig. 3. Members of the ‘soft-plumaged petrel’ Pterodroma feae/madeira/mollis complex in flight. Note that true tonal values may be difficult to judge in field conditions.

British Birds 97 • January 2004 • 6-15 9 The ‘soft-plumaged petrel’ complex distinct species, and noted that analysis of mito- Peter Hayman. Gantlett suggested that madeira chondrial-DNA sequences suggests that the has a relatively short ‘hand’ and broader, divergence of feae and madeira occurred blunter wing-tip than feae (with mollis interme- approximately 840,000 years ago. They also diate between the two), and that these differ- ‘provisionally’ retained deserta as conspecific ences should be apparent in the field. with P. feae. This cautious approach towards the Nonetheless, he failed to draw attention to the position of deserta is prudent, given that its bill fact that the photographs claimed as madeira in structure does not differ from that of feae (see his article had previously been published as feae below). Furthermore, differences in morpho- (in Fisher 1989). Howell (1996) referred to the metrics are small, and there are comparable dif- same photographs and drew attention to a ferences in breeding phenology among number of apparent differences in wing and tail populations of both Bulwer’s Petrel Bulweria patterns which he suggested could be used to bulwerii and Madeiran Storm-petrel Ocean- separate feae and madeira, but Zino & Biscoito odroma castro (Snow & Perrins 1998). (2001) were unable to establish any reliable plumage differences. Until such proposed dif- Identification ferences are proven, on the basis of photographs Given their rarity and taxonomic complexity, of birds of known identity, they remain conjec- and the effort required to obtain good views, it tural. Tove (1997a,b) developed his earlier posi- is not surprising that field identification criteria tion regarding the identification of Pterodroma for members of this group have been slow to petrels in the North Atlantic, and supported evolve. Most early accounts focused on distin- Gantlett’s suggested differences in wing shape as guishing Pterodroma petrels from other a useful feature. He also documented 17 indi- seabirds. Even the standard text (Harrison viduals recorded from North American waters, 1983) offered little help in separating members and included nine photographs of five individ- of the complex in the northern hemisphere. It uals. was only following publication of the first pho- Brinkley & Patteson (1998) considered sepa- tographs of what were claimed as feae at sea ration of feae and madeira at sea to be feasible (Fisher 1989) that real progress began, although on the basis of overall size and, especially, the as recently as 1998, Anthony McGeehan sug- depth of the bill, although they implied that gested that the dark morph of Fulmar Fulmarus other potential characters remained to be glacialis (‘Blue Fulmar’) may be an identifica- tested. But, using measurements and wing out- tion pitfall (McGeehan 1998). lines drawn from photographs, Tove (2001) Fisher (1989) concluded that the most useful claimed to have verified the proposed differ- field characters for distinguishing feae from ences in wing shape, and even went so far as to madeira are likely to be the size of the head and make an analogy with Laughing Gull Larus atri- of the bill. In contrast, Carter (1989) noted that cilla and Franklin’s Gull L. pipixcan.He com- the most striking features of feae in comparison mented on the bird reported off the Isles of with mollis seemed to be its contrastingly pale Scilly in July 2001 (Fisher & Flood 2001), rump and tail, and dark crown. Madge (1990) stating that it was an ‘obvious and unam- failed to notice the pale tail on many feae seen biguous Fea’s’. around Cape Verde, however, and this feature Examination of published photographs of was considered to be poorly defined by Harrop birds of known identity does not, however, (2001). inspire confidence in the validity of differences By the early 1990s, there had been several in wing shape as a field character. Directly com- extralimital records of ‘soft-plumaged petrels’ parable photographs of hand-held outstretched from both sides of the North Atlantic (Enticott underwings of deserta (Jouanin et al. 1969, plate 1999), although their specific identification 4) and madeira (Wingate et al. 1998, fig. 11) do remained controversial. Using photographic not reveal a difference in shape (contra Tove and other evidence, Tove (1993) concluded that 2001), while the shape of an unheld, out- those occurring off North Carolina, USA, were stretched upperwing of deserta (Jouanin et al. feae. Gantlett (1995) discussed the field identifi- 1969, plate 5) is similar to the wing shape of the cation of the complex based on photographs bird in plate 7 of Gantlett (1995), which is taken off the Desertas, in the Madeiran archi- claimed to be madeira. pelago, and measured drawings of specimens by Tove’s methods, though not necessarily his

10 British Birds 97 • January 2004 • 6-15 The ‘soft-plumaged petrel’ complex Tony Marr Tony

1. Fea’s Petrel Pterodroma feae deserta, Desertas, Marr Tony Madeira Islands, August 1990.The heavy bill is 2. Fea’s Petrel Pterodroma feae deserta, Desertas, apparent. Note how bright sunshine makes Madeira Islands, August 1990.The bill structure the crown and tail appear paler than their is just visible in this photograph. Note the true colour tones. appearance of a partial dark breast-band. Tony Marr Tony

Tony Marr Tony 4. Presumed Fea’s Petrel Pterodroma feae, 3. Fea’s Petrel Pterodroma feae deserta, Desertas, Desertas, Madeira Islands, August 1990.Typical Madeira Islands, August 1990 (same individual as in heavy-billed and hooded appearance, with paler plate 2). Note how the grey tail appears only a tail catching the light. Note how the wing-tips shade or two paler than the mantle in flat light. appear quite rounded in this photograph.

British Birds 97 • January 2004 • 6-15 11 The ‘soft-plumaged petrel’ complex conclusion that the identity of all the well-doc- feae may appear to show a breast-band in the umented North Atlantic records is consistent field. Dark morphs are relatively rare, and diffi- with feae, were contested by both Marr (2001) cult to distinguish from some other dark petrels and Harrop (2001). Harrop pointed out that which are outside the scope of this paper. there are differences in the structure of the bills of feae and madeira. On feae, the distance Extralimital records between the tip of the nostril (naricorn) and Until there is a pattern of records which have the back of the hook at the tip of the upper been formally accepted at species level, assess- mandible (maxillary unguis) is very short, ment of the distribution of these three species forming a small ‘notch’ in the contour of the bill away from the breeding areas will remain highly in profile. On madeira, this distance is longer speculative. In his discussion of the first record and forms the impression of a ‘wedge’. This of a ‘soft-plumaged petrel’ off Ireland, in Sep- feature had not been noted before, presumably tember 1974, Enticott (1999) made a good case because it does not constitute one of the stan- for associating the record with the path of a dard bill measurements (cf. Zino & Zino 1986, dying hurricane. Since the early 1980s, and in Wingate et al. 1998). Further research has con- particular since the early 1990s, there has been a firmed that, if visible, this is a reliable feature phenomenal increase of records in British for the separation of feae and madeira, but there waters, many of which cannot be related to hur- is no apparent difference in bill structure ricanes. between nominate feae and deserta. The emerging pattern of records, mainly in To summarise, there are no known consis- late spring and summer in the western North tent plumage differences between feae and Atlantic, and in late summer and early autumn madeira, and the proposed differences in wing in the eastern North Atlantic, is likely to reflect structure remain of unproven validity in the a combination of factors including observer field. Size may be helpful in direct comparison, activity and improved optical instruments. It but is unlikely to be much help when identi- may also, however, be partly attributable to a fying lone vagrants away from the breeding real rise in numbers, and it would not be sur- grounds. On current knowledge, only bill struc- prising if global warming brings increasing ture can be regarded as diagnostic, and this will numbers of tropical species into temperate require excellent views at close range. If, at some waters. Since all the well-documented records to point in the future, deserta were to be split from date are compatible with feae, the suggestion feae, field identification would almost certainly that birds from Cape Verde disperse rapidly be impossible. after breeding (from May onwards) to the Study of skins of mollis, together with pho- western North Atlantic, then follow a clockwise tographs of birds of known identity, has shown route which brings them back into eastern that a number of consistent differences exist North Atlantic waters (Enticott 1999), seems between mollis and feae/madeira which, given the best working hypothesis. good views, should make field identification Leaving aside the records from both sides of possible. The bill of mollis tends to be relatively the North Atlantic discussed above, the only light and more similar in structure to that of extralimital records to date involving birds madeira than feae, although a few individuals unequivocally identified to the species level have heavier bills more like feae (fig. 1). More involve single examples of feae trapped on the importantly for field identification, on pale Azores in June 1990 (Bibby & del Nevo 1991); morphs of mollis the crown is typically paler again on the Azores in September 1993, grey than on feae/madeira and therefore con- retrapped in August 1994 (Monteiro & Furness trasts more strongly with the dark facial ‘mask’ 1995); and a specimen of feae found in Feb- (fig. 2), while the mantle and greater coverts are ruary 1963 on the western shore of the Dead also paler grey. It must be emphasized, however, Sea, Israel (Bourne 1983b; Shirihai 1999 – note that wind and light make a significant differ- that the date was given incorrectly as November ence to the appearance of birds in the field (fig. 1968 in the latter reference). The identity of the 3), so these factors need to be taken into Israeli specimen was confirmed by Killian account before attempting identification. Pale Mullarney (in litt. and from photographs), morphs also have a variable, but typically bold, though he had not examined the actual speci- grey breast-band, though, as noted below, some men as implied by Shirihai (1999).

12 British Birds 97 • January 2004 • 6-15 The ‘soft-plumaged petrel’ complex Tony Marr Tony Tony Marr Tony 5. Fea’s Petrel Pterodroma feae deserta, Desertas, 6. Fea’s/Zino’s Petrel Pterodroma feae/madeira, Madeira Islands, August 1990.The typical heavy- Desertas, Madeira Islands, August 1990. Although the billed and hooded appearance is again apparent. bird is close, at this angle it is impossible to assess bill structure. On current knowledge, none of the visible characters is sufficient for specific diagnosis. Tony Marr Tony Marr Tony 7. Soft-plumaged Petrel Pterodroma mollis mollis,at 8. Soft-plumaged Petrel Pterodroma mollis mollis,at sea between Argentina and the Falklands, December sea between Argentina and the Falklands, December 1992. Note the quite pale grey breast-band and the 1992. Even in a poor view, the pale grey crown and head pattern of this typical bird in the South Atlantic. mantle, as well as the concolorous tail are visible.

British Birds 97 • January 2004 • 6-15 13 The ‘soft-plumaged petrel’ complex

Although Jepson & Zonfrillo (1988) Atlantic Islands,Vol. 2. A history of the birds of Madeira, reported two possible mollis between Madeira the Desertas, and the Porto Santo Islands. Oliver & Boyd, Edinburgh. and Deserta Grande in 1986, the claim was Bibby, C., & del Nevo, A. 1991. A first record of regarded as dubious by Zino et al. (1995) on the Pterodroma feae from the Azores. Bull. Brit. Orn. Club basis that some feae have extensive breast-bands 111: 183-186. Bourne,W. R. P.1957. Additional notes on the birds of the (with a gap of as little as about 10 mm) which Cape Verde Islands, with particular reference to may appear complete in the field (cf. fig. 3). Bulweria mollis and Fregata magnificens. Ibis 99: 182-190. Zino et al. required an unambiguous photo- — 1983a.The Soft-plumaged Petrel, the Gon-gon and the graphic record or specimen for a record to be Freira, Pterodroma mollis, P. feae and P. madeira. Bull. Brit. Orn. Club 103: 52-58. acceptable. There is one accepted sight record of — 1983b. A Gon-gon Pterodroma (mollis) feae in Israel. mollis in the Western Palearctic, at Eilat, Israel, Bull. Brit. Orn. Club 103: 110-111. in March 1997 (Shirihai 1999). In the absence Bretagnolle,V. 1995. Systematics of the Soft-plumaged Petrel Pterodroma mollis (Procellariidae): new insights of any other confirmed records from anywhere from the study of vocalizations. Ibis 137: 207-218. in the northern hemisphere, however, this Brinkley, E. S., & Patteson, J. B. 1998. Gadfly petrels in the record is open to question. According to the western North Atlantic. Birding World 11: 341-354. BOU. 2000. The British List. British Ornithologists’ Union, published description, the bird had a dark Tring. crown, an incomplete breast-band, and the dark Carter, M. 1989. Identification of Soft-plumaged Petrels. bill was prominent. Its identification as mollis Birding World 2: 405. was based on concolorous dark upperparts, Clancey, P.A., Brooke, R. K., & Sinclair, J. C. 1981.Variation in the current nominate subspecies of Pterodroma rump and uppertail, yet these areas can also mollis (Gould) (Aves: Procellariidae). Durban Mus. Novit. appear concolorous on feae. The combination 12 (18): 203-213. of characters shown by this individual is more Cooper, J. 1999. The Late Pleistocene avifaunas of Gibraltar and their palaeoenvironmental significance. Unpublished likely to be shown by feae than mollis, so its PhD thesis, University of London. identification as mollis appears to be unsafe. Cramp, S., & Simmons, K. E. L. (eds.) 1977. The Birds of the Two specimen records from South Africa Western Palearctic.Vol. 1. OUP,Oxford. were formerly thought to be possible madeira Enticott, J.W. 1991. Identification of Soft-plumaged Petrel. Brit. Birds 84: 245-264. on the basis of morphology and biometrics — 1999. Britain and Ireland’s first ‘Soft-plumaged Petrel’ – (Clancey et al. 1981), but the only living author an historical and personal perspective. Brit. Birds 92: of this paper, Ian Sinclair, now believes that 504-518. Fisher, A., & Flood, B. 2001.The Fea’s Petrel off the Isles of both are probably southern mollis (per Jim Scilly. Birding World 14: 289-292. Enticott). There has, therefore, not been any Fisher, D. 1989. Pterodroma petrels in Madeira. Birding confirmed historical record of madeira away World 2: 283-287. Gantlett, S. 1995. Identification forum: field separation of from the breeding grounds. Fea’s, Zino’s and Soft-plumaged Petrels. Birding World 8: 256-260. Acknowledgments Harrison, P.1983. Seabirds: an identification guide. Croom Tony Marr was instrumental in drawing together all the Helm, Beckenham. people involved with this paper. He and Ian Lewington Harrop, A. 2001. Separation of Fea’s and Zino’s Petrels. contributed helpful advice and discussion, as well as, Birding World 14: 512-513. respectively, the excellent slides and artwork which Hazevoet, C. J. 1995. The Birds of the Cape Verde Islands. accompany this paper.Tim Inskipp, stimulated by his BOU Checklist No. 13,Tring. sighting of a Pterodroma off Dungeness, Kent, in October — 1997. Notes on distribution, conservation, and 1983, undertook a huge amount of the initial research on taxonomy of birds from the Cape Verde Islands, which this review is based, and made helpful comments including records of six species new to the archipelago. on the first draft. Jo Cooper provided information about Bull. Zool. Mus. Univ. Amsterdam 15: 89-100. the fossil record, while Robert Pry^s-Jones kindly provided Howell, S. 1996. Pterodroma identification revisited. Birding access to specimens at the NHM. Killian Mullarney World 9: 276-277. confirmed the identity of the Dead Sea specimen, and Jim Imber, M. J. 1985. Origins, phylogeny and taxonomy of the Enticott reported current thinking about the specimens gadfly petrels Pterodroma spp. Ibis 127: 197-229. from South Africa. Jimmy Steele provided useful discussion Jepson, P., & Zonfrillo, B. 1988. Bird notes from Madeira, in relation to his work on this group for BBRC. Linda summer 1986. Bocagiana 117: 1-9. Birch, at the Alexander Library in Oxford, answered Jouanin, C., Roux, F., & Zino, F. 1969.Visites aux lieux de queries about references patiently and often at short nidification de Pterodroma mollis ‘deserta’. L’Oiseau et notice. RFO 39: 161-175. Knox, A. 1995. Pterodroma taxonomy revisited. Birding World 8: 424. References Lysaght, A. M. 1959. Some eighteenth century bird ABA Checklist Committee. 1999. 1998-99 American paintings in the library of Sir Joseph Banks. Bull. Brit. Birding Association Checklist Report. Birding 31: 518- Mus. (Nat. Hist.) Hist. Ser. 1: 253-371. 524. Madge, S. 1990. Soft-plumaged Petrels at sea. Birding World Bannerman, D. A., & Bannerman,W. M. 1965. Birds of the 3: 138-139.

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Marr,T. 2001. Identification of Fea’s and Zino’s Petrels. Taxonomic recommendations for European birds. Ibis Birding World 14: 512. 144: 153-159. Mathews, G. M. 1934a. (Description of a new subspecies Shirihai, H. 1999. Fifty species new to Israel, 1979-1998: of the Soft-plumaged Petrel Pterodroma mollis.) Bull. Brit. their discovery and documentation, with tips on Orn. Club 54: 161. identification. Sandgrouse 21 (1): 45-105. — 1934b. (The Soft-plumaged Petrel, Pterodroma mollis, Snow, D.W., & Perrins, C. M. (eds.) 1998. Birds of the and its subspecies.) Bull. Brit. Orn. Club 54: 178-179. Western Palearctic. Concise edition. OUP,Oxford. McGeehan, A. 1998. Appearance of ‘Blue Fulmar’ and Tove, M. H. 1993. Soft-plumaged Petrels in the NW potential confusion with Soft-plumaged Petrels. Dutch Atlantic. Birding World 6: 414. Birding 20: 66-68. — 1997a. Fea’s Petrel in North America. Part 1. Birding Monteiro, L. R., & Furness, R.W. 1995. Fea’s Petrel 29: 206-214. Pterodroma feae in the Azores. Bull. Brit. Orn. Club 115: — 1997b. Fea’s Petrel in North America. Part 2. Birding 9-14. 29: 309-315. Nunn, G. B., & Stanley, S. E. 1998. Body size effects and — 2001.Verification of suspected field identification rates of cytochrome b evolution in tube-nosed differences in Fea’s and Zino’s Petrels. Birding World 14: seabirds. Mol. Biol. Ecol. 15: 1360-1371. 283-289. — & Zino, F. In press. A molecular genetic diagnosis of Wingate, D. B., Hass,T., Brinkley, E. S., & Patteson, J. B. 1998. Europe’s rarest seabirds: the endangered Pterodroma Identification of Bermuda Petrel. Birding 30: 19-36. gadfly petrels nesting in the Madeiran archipelago. Zino, F., & Biscoito, M. J. 2001. Comparative biometrics of Ratcliffe, N., Zino, F., Oliveira, P.,Vasconceles, A., Hazevoet, Pterodroma madeira and P. feae from the archipelago of C. J., Neves, H. C., Monteiro, L. R., & Zino, E. A. 2000. Madeira (NE Atlantic). Symposium Poster Presentation, The status and distribution of Fea’s Petrel Pterodroma Freira Conservation Project, Funchal, Madeira. feae in the Cape Verde Islands. Atlantic Seabirds 2: 73- —, — & Zino, P.A. 1995. Birds of the Archipelago of 86. Madeira and the Selvagens, new records and checklist. Salvadori,T. 1899. Collezioni ornitologische fatte nelle isole Bol. Mus. Mun. Funchal 47 (262): 63-100. del Capo Verde da Leonardo Fea. Ann. Mus. Civ. Genova Zino, P.A., & Zino, F. 1986. Contribution to the study of (2) 20: 283-312. the petrels of the genus Pterodroma in the archipelago — 1900. On Oestrelata mollis (Gould) and the allied of Madeira. Bol. Mus. Mun. Funchal 180: 141-165. species living at Madeira and the Cape Verde Islands. Zonfrillo, B. 1993. Relationships of the Pterodroma petrels Ibis (7) 6: 298-303. from the Madeira archipelago inferred from their Sangster, G., Knox, A. G., Helbig, A. J., & Parkin, D.T. 2002. feather lice. Bol. Mus. Mun. Funchal Suppl. 2: 325-331.

Andrew H. J. Harrop, 30 Dean Street, Oakham, Rutland LE15 6AF; e-mail: [email protected]

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