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'Soft-Plumaged Petrel' Complex

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'Soft-Plumaged Petrel' Complex 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.
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