139

DUTCH AVIFAUNAL LIST: SPECIES CONCEPTS, TAXONOMIC INSTABILITY, AND TAXONOMIC CHANGES IN 1977-1998

GEORGE SANGSTER1, CORNELIS J. HAZEVOET2,3, ARNOUD B. VAN DEN BERG4, C.S. (KEES) ROSELAAR2 & RONALD SLUYS2

Sangster G., c.J. Hazevoet, A.B.Van den Berg, C.S. Roselaar & R. Sluys 1999. Ar­ dea 87: 139-165.

The Dutch avifaunal list is revised based on the principles of phylogenetic theory and methodology. A phylogenetic approach to species-level taxa is adopted. In con­ trast to the 'Biological Species Concept', this approach is compatible with the recon­ struction of evolutionary relationships, recognises species on the basis of historical patterns and views species as products of history. Two basic rules are applied for the recognition of higher taxa: (1) higher taxa are named clades and represent monophy- letic groups of species or less inclusive clades, and (2) phylogenetic knowledge should be expressed as accurately as possible by taxonomy. Systematics is viewed as a historical science in which phylogenies are hypotheses of historical relationships. Taxonomies should reflect the best supported hypotheses of relationships and are subject to further modification as knowledge of relationships grows.

Key words: systematics - taxonomy - phylogeny - species concepts - species - higher taxa

lNieuwe Rijn 27, 2312 JD Leiden, Netherlands; E-mail: [email protected]; 2Insti­ tute of Systematics and Population Biology, Zoological Museum, University ofAm­ sterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands; 3Museu e Laborat6rio Zool6gico e Antropol6gico (Museu Bocage), Rua da Escola Politecnica 58, 1250 e9 Lisboa, Portugal; 4Duinlustparkweg 98, 2082 EG Santpoort-Zuid, Netherlands

INTRODUCTION (1998). Due to limitations of space, the present report only includes a brief summary of the ratio­ This report includes taxonomic and nomenclatu­ nale for each decision. More extensive and ex­ ral changes adopted by the Dutch committee for plicit motivations for the recognition of some of avian systematics (Commissie Systematiek Ne­ the species-level and higher taxa listed here will derlandse Avifauna, CSNA) since Voous (1977). be published elsewhere. Proposals affecting the Dutch list that were re­ jected by the CSNA are also discussed. The Neth­ Choice of species concept and its application erlands Ornithologists' Union (NOU) and the Over the past few years the fields of system­ Dutch Birding Association (DBA) support the atic and evolutionary biology have been invigo­ CSNA and its reports are published in both Ardea rated by a scientific discussion on the concept of and Dutch Birding. The present report is essen­ species (for reviews, see Sluys 1991; Mayden tially similar to two separate publications in 1997; Zink 1997). Although this discussion contin­ Dutch Birding (Sangster et al. 1997; 1998). The ues, it is also true that new and useful insights committee currently consists of five members have been gained already that can be incorporated (year of election between parentheses): Arnoud fruitfully into an up to date and modem descrip­ B. van den Berg (1995), Cornelis J. Hazevoet tion of avian diversity. One of the insights that has (1996), C.S. (Kees) Roselaar (1995), George surfaced in the literature is that different species Sangster, Secretary (1996) and Ronald Sluys concepts may be best for different purposes 140 ARDEA 87(1),1999

(Endler 1989) and it seems increasingly likely that gely based on these views. The Isolation Species no single species concept will satisfy the multiple Concept (popularly known as the 'Biological purposes of 'species' in the biological sciences Species Concept') is rejected because its proper­ (Sluys 1991; Hull 1997), However, a theme com­ ties violate all three aforementioned principles. mon to all biological sciences is that all taxa are First, interbreeding taxa are not necessarily more historically connected through their pattern of an­ closely related to each other than they are to taxa cestry and descent. All living taxa are the product from which they are reproductively isolated. Be­ ofhistory, and we can understand little about their cause interbreeding is the prime criterion for con­ diversity without knowledge of their history, i.e. specificity under the ISC, the ISC could still re­ the phylogenetic knowledge provided by system­ gard such interbreeding taxa as conspecifics. atics (O'Hara et al. 1988). Virtually all compara­ Therefore, the problem of lumping taxa which are tive studies of biological variation within and not closely related in a single species and, hence, among taxa depend on such phylogenetic knowl­ the misrepresentation of evolutionary history, is edge for interpretation (Felsenstein 1985; Brooks inherent to the ISC and does not simply result & McLennan 1991; Harvey & Pagel 1991). There­ from errors in application. Phylogenetic analyses fore, in biodiversity studies and comparative biol­ indicate that in various groups of birds 'polyty­ ogy a fundamental requirement of species-level pic' species recognised by the ISC do not repre­ taxa is that they are compatible with the recon­ sent natural (monophyletic) groups. Evidence struction of evolutionary relationships. Species comes from phylogenies based on both morpho­ concepts which group taxa that are not closely re­ logical (Livezey 1995a; Chu 1998; Veron 1999) and lated in a single species misrepresent evolution­ molecular data sets (Zink 1988; Friesen et al. 1996; ary history. Second, species-level taxa should be Leisler et al. 1997; Roy et al. 1997; Trewick 1997). delimited on the basis of historical subdivisions Second, under the ISC taxa are recognised as spe­ (i.e. historical patterns), rather than present-day cies ifthey remain 'reproductively isolated', in the or possible future interactions and processes sense that they do not fuse into a single population (Liden & Oxelman 1989), such as hybridisation (Mayr 1982; 1996). The ISC, therefore, is prospec­ and gene flow. Species concepts which are pro­ tive (O'Hara 1993; 1994; Maddison 1997); only fu­ spective and which require speculations about the ture events will show whether currently recog­ future are not helpful in biology; since all of our nised taxa remain reproductively isolated or fuse data are of the present and past, the units by into each other. This poses both theoretical and which we interpret these data must also be strictly practical problems. It makes little sense to try to historical (Maddison 1997). Third, species should interpret the past and present diversity of organ­ be basal, taxonomically comparable units (Cra­ isms with a taxonomy that is based on expecta­ craft 1987; 1989): species should be basal taxa, tions ('dreams', Maddison 1997) about the future. that is, taxa that contain no included taxa. A spe­ A practical problem is that, except for rare cases, cies concept should not combine several distinct the process of fusion transcends observable time. taxa in a single (polytypic) 'species' because such The likely time-to-fusion may be measured in 'species' actually are higher, more inclusive taxa. WOOs or even millions of years (Zink & McKi­ Species concepts which recognise not only single trick 1995). Third, many 'species' recognised by units (monotypic species) but also polytypic as­ the ISC contain more than one taxon. The ISC rec­ semblages (polytypic species) as 'species' run ognises monotypic species but may also unite up counter to the fundamental need for species-level to 10 or more diagnosable taxa and still recognise taxa to be basal and comparable. the resulting unit as a single 'species'. This not The decision by the CSNA to abandon the tra­ only underestimates and misrepresents biodiver­ ditional Isolation Species Concept (ISC) in favour sity but also compromises interspecific compari­ of a Phylogenetic Species Concept (PSC) was lar- sons (Prum 1994; Hazevoet 1996; Cracraft 1997). Sangster et at.: TAXONOMIC CHANGES IN 1977-1998 141

Two distinct Phylogenetic Species Concepts sion, however, would recognise the descendant have been advocated. These versions agree in species, but not the ancestral 'species', as a phy­ viewing species as products of evolution, not as logenetic species because only the former is players in evolution, and support the notion of monophyletic. Proponents of the monophyly ver­ species as basal taxa. The original version, pro­ sion propose that such ancestral 'species' are rec­ posed by Cracraft (1983) and further developed by ognised as a separate class of species, for which Nixon & Wheeler (1990) and Davis & Nixon they propose the term 'metaspecies' (Donoghue (1992), considers a phylogenetic species to be an 1985; De Queiroz & Donoghue 1988). irreducible cluster of organisms possessing at The CSNA has adopted the diagnosability least one diagnostic character state. This version version of the PSC as its operational species con­ thus focuses on the diagnosability of species. Dia­ cept because no phylogenetic analysis is required gnostic character states are discrete character prior to delimiting species, its implementation in­ states which are fixed within the species and are volves little modification of existing taxonomic absent from close relatives. Diagnosability of practices and does not require the recognition of species may be based on any intrinsic attribute, additional classes of species. It is believed that either morphological, molecular, ethological or a these are advantages over the monophyly version, combination of these. It should be emphasised and similar versions (Baum & Shaw 1995), and that this concept is populational (Cracraft 1997); that these outweigh any disadvantages. Although the criterion of diagnosability applies to (groups both aspects (diagnosability and monophyly) may of) populations, and not to, e.g. family groups or be meaningful at the level of species (Baum individuals. An alternative approach was devel­ 1992), the monophyly version is not sufficiently oped by Donoghue (1985) and De Queiroz & practical as an operational species concept. Donoghue (1988; 1990) and considers phylogen­ It has been argued that the PSC actually de­ etic species to be the smallest monophyletic fines the evolutionary lineage and does not repre­ groups oforganisms supported by autapomorphies sent a species concept at all (Bock 1979; 1994; (unique derived character states). Szalay & Bock 1991). This complaint stems from The diagnosability and monophyly versions a narrow view of species, based on the ISC, of the PSC are significantly different; both in the­ which restricts the term 'species' to entities which ory and application (Baum 1992; Davis 1997). A are involved in the process of evolutionary major difference is that the monophyly version change. According to this view, species are real requires phylogenetic analysis prior to delimiting only at a single point in time because repro­ species, whereas the diagnosability version does ductive isolation and processes such as natural se­ not. Implementation of the monophyly version lection operate only in a single time-slice. Al­ will, therefore, result in a significant shift in taxo­ though the ISC can not be applied to organisms nomic practice, whereas the diagnosability ver­ living at different times, it does not follow that sion will not (Baum 1992). In addition to this other species concepts should also be applicable practical difference, the two versions also differ in one time-slice only. In any case, the PSC does with regard to which basal taxa are called 'spe­ not define species in terms of reproductive isola­ cies'. When a small group of related individuals tion (which, indeed, can only be assessed at a leaves a species, evolves diagnostic character given moment in time) but in terms of diagnostic states and thus forms a descendant species, the character states. The presence of diagnostic char­ ancestral 'species' will cease to be monophyletic. acter states in a population may extend over time Because both the ancestral and descendant 'spe­ and, therefore, phylogenetic species-level taxa cies' are characterised by diagnostic character will have reality over time. However, this does states, the diagnosability version would recognise not mean that the PSC defines the evolutionary both as phylogenetic species; the monophyly ver- lineage, as claimed by Szalay & Bock (1991) and 142 ARDEA 87(1), 1999

Bock (1994). Since diagnostic character states artificial and without biological relevance (e.g. may become fixed in a population after a lineage Voous 1964). Above all, there was no consistent has split, speciation may be completed after the method to reconstruct phylogeny. origin of a lineage and, therefore, phylogenetic Hennig (1966) restricted the term 'phylogeny' species are not synonymous with evolutionary to the branching pattern of evolutionary history, lineages. argued convincingly that higher taxa (i.e. mono­ Throughout this paper, 'qualitative differ­ phyletic groups) are real, and that these can be re­ ences' are differences that can be coded as dis­ constructed by phylogenetic analysis (sub­ crete character states. Published data on diagnos­ sequently termed cladistics). The consistency and tic character states of taxa were evaluated and logic of phylogenetic systematics was greeted interpreted with reference to the diagnosability with enthusiasm by systematists (e.g. Nelson version of the PSc. This does not imply that the 1970; Brundin 1972; Cracraft 1972) and is now al­ authors of the original publications necessarily most universally accepted (e.g. Wiley 1981; Hillis proposed the relevant taxa to be considered as et al. 1996; Dingus & Rowe 1998). One reason for species-level taxa or explicitly endorse a PSC. In its success is that units of study in biology (from fact, many of the authors concerned - implicitly genes through organisms to higher taxa) do not or explicitly - worked under the umbrella of the represent statistically independent observations ISC. The present interpretation of the data is fully but rather are interrelated through their historical the responsibility of the CSNA. In the absence of connections. Therefore, almost any comparative a fully documented system of phylogenetically study requires information on phylogeny (Felsen­ defined species level taxa, the CSNA continues to stein 1985; Brooks & McLennan 1991; Harvey & use subspecific names to denote the likely geo­ Pagel 1991). As a result, phylogenetic analyses are graphic origin of the populations occurring in the now firmly entrenched in contemporary biology Netherlands. and new applications continue to appear (e.g. Harvey et al. 1996). Being the most central and Phylogeny and higher taxa unifying concept in biology, phylogeny should Although the study of phylogeny assumed an also be a central principle in taxonomy. important place in biology in the decades follow­ In line with phylogenetic theory and method­ ing Darwin's Origin of species (Darwin 1859), ology (Hennig 1966; Wiley 1981; de Queiroz & biologists' interest in phylogeny was sub­ Gauthier 1992), two basic rules are applied for the sequently replaced by new emphases on the pro­ recognition ofhigher taxa (i.e. taxa above the level cesses and mechanisms of genetics, development of species): (1) higher taxa are named clades and, and evolution (e.g. Dobzhansky 1937; Mayr therefore, represent monophyletic groups of spe­ 1942). With the 'Evolutionary Synthesis' of the cies or less inclusive clades; hence, higher taxa 1930s and 1940s a dichotomy developed between are delimited on the basis of common ancestry, the study of species and the study of phylogeny rather than a shared set of character states; (2) and higher taxa. Studies at the level of popula­ phylogenetic knowledge should be expressed as tions and species proceeded with great vigour, but accurately as possible in nomenclature. As a con­ the study of phylogeny suffered from a lack of sequence of the first rule, para- and polyphyletic conceptual and methodological clarity. For in­ taxa should be abandoned. This has resulted in the stance, it was not clear whether the term 'phylo­ recognition of Casmerodius for Great White Eg­ geny' should only denote the branching pattern of ret C. aibus and Mergellus for Smew M. aibellus evolutionary history or whether it should also in­ and in the transfer of some species ofHippolais to clude reference to the level of divergence subse­ Acrocephalus. quent to cladogenesis. Also, it was commonly be­ The second rule implies that adjustments to lieved that, in contrast to species, higher taxa are the present system should be enacted when im- Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 143

proved phylogenetic knowledge becomes avail­ bley 1989; 1994). Fortunately, in recent years tax­ able. For instance, in the past 15 years, evidence onomic systems have become more consistent for a sister-group relationship of Anseriformes with current knowledge of phylogenetic relation­ and Galliformes has accumulated and now greatly ships. Closely associated with this development is outweighs all other hypotheses of relationships of a reappraisal of systematics as a historical science these orders. This has resulted in the recognition (Gould 1986), in which phylogenies are viewed as of the taxon Galloanserae. More detailed knowl­ hypotheses of relationships and taxonomies based edge of the relationships among cormorants and on them as dynamic systems, subject to further shags, gannets and boobies, dabbling ducks, and modification as our knowledge of relationships cardueline finches has resulted in the recognition grows. of four additional genera, Stictocarbo, Morus, Systematics is not an exercise in producing Mareca and Chloris. Generally, established usage classifications and sequences convenient for hu­ is maintained unless alternative hypotheses are mans, but an attempt to discover an underlying better supported. As a consequence, Wilson's real structure in nature (Griffiths 1994). That real Phalarope Phalaropus tricolor is not placed in structure, or natural system, is the pattern of his­ Steganopus and Gelochelidon is not included in torical (phylogenetic) relationships. This natural Sterna. It has been attempted to not recognise system is something we discover (i.e. recon­ higher taxa that are not or only weakly supported struct), not something we create (Ghiselin 1987). by phylogenetic analyses. This has resulted in the The objective of systematics, therefore, is the re­ inclusion of Chettusia in Vanellus, Catharacta in construction of phylogenetic relationships; tax­ Stercorarius and Tachymarptis in Apus. onomy is concerned with the representation of these relationships. With its focus on past events Systematics and taxonomic (in)stability (i.e. historical subdivisions), systematics is a his­ There is a large hiatus between our knowledge torical science. of phylogenetic relationships, which has greatly It needs to be emphasised that reconstructed increased since the late 1970s, and the avian taxo­ phylogenies (cladograms) are hypotheses. The nomic system (classification), which is basically true phylogeny is buried in history and is un­ still the same as the one proposed by Alexander known and probably unknowable. Therefore, Wetmore in 1930, which in tum was largely based proof, in a literal sense, of phylogenetic relation­ on the work of Max Ftirbringer and Hans Gadow ships may never be obtained. Because the true in the late 19th century. Some have suggested that phylogeny is unknown, there is no a priori basis the goal of systematics is to produce a 'stable for accepting or rejecting a given phylogeny; hy­ standard sequence' and that the Wetmore classifi­ potheses must be tested in the light of additional cation and sequence is now so well entrenched data. A phylogenetic hypothesis is open to test by that it should continue to serve as the standard se­ the addition of new characters, the addition of quence (Mayr 1989; Mayr & Bock 1994). The new taxa and the re-evaluation of other charac­ 'stability' of the Wetmore sequence, especially in ters. Thus, a phylogeny can be corroborated or re­ the light of much systematic research, was viewed jected and replaced by another phylogeny. Cor­ by Mayr & Bock (1994) as 'of major advantage to roboration may come in the form of congruence. all avian biologists' because, in the Wetmore se­ Phylogenies are congruent if they show the same quence, biologists can easily locate a particular branching pattern. If congruence exists between group or species. However, this 'stability' is not phylogenies based on different sets of data, this to be regarded as a proof of its correctness. In may indicate a strong historical signal; congru­ fact, the 'stability' of the Wetmore sequence ence may be regarded as evidence that the rele­ throughout this century is entirely due to failure vant phylogenies identify the true organismal to incorporate new ideas about relationships (Si- phylogeny. Rejection of a phylogenetic hypoth- 144 ARDEA 87(1), 1999 esis requires that an alternative hypothesis better Cygnus columbianus Whistling Swan summarises all available evidence. Fluitzwaan Because the purpose of taxonomy is commu­ Cygnus bewickii Bewick's Swan Kleine Zwaan nication of information about phylogenetic rela­ Whistling Swan and Bewick's Swan are spe­ tionships, taxonomic systems should reflect the cifically distinct (Stepanyan 1990; Gantlett et al. current best-supported hypothesis of relation­ 1996; King 1997) based on qualitative differences ships. Taxonomic systems should be adjusted in morphology (Evans & Sladen 1980; Livezey only ifit is believed that an alternative hypothesis 1996). is better supported by the available evidence. Be­ cause phylogenies are hypotheses which are sub­ Anserfabalis Taiga Bean Goose Taigarietgans ject to further testing, taxonomies based on them Anser serrirostris Thndra Bean Goose are provisional and may later be replaced or modi­ Toendrarietgans fied. Therefore, taxonomies are dynamic systems; Taiga Bean Goose and Tundra Bean Goose are a definitive taxonomic system is probably unat­ specifically distinct (Sangster & Oree11996; King tainable in practice. 1997; Persson 1997; Wells 1998) based on differ­ ences in proportions, vocalisations, feeding habi­ tat and diet, photosensitivity, activity pattern, TAXONOMIC CHANGES behaviour, phenology and responses to periods of extreme cold (Berry 1938; Coombes 1951; Galloanserae Mathiasson 1963; Huyskens 1977; 1986; Van Impe A sister-group relationship of Anseriformes 1980; Kurechi et al. 1983; Van den Bergh 1985; and Galliformes is strongly supported by con­ Barthel 1989; 1995; Burgers et al. 1991; Miyabay­ gruency of phylogenetic analyses of several inde­ ashi et al. 1994). Ringing recoveries suggest that pendent data sets. These include morphological Taiga Bean Goose and Tundra Bean Goose have characters (Cracraft 1986; 1988; Cracraft & Min­ allopatric breeding distributions (Burgers et al. dell 1989; Andors 1991; 1992; Kurochkin 1995; 1991) and there is no evidence for 'intergradation' Livezey 1997a), DNA-DNA hybridisation (Sibley (Sangster & Oreel 1996). In both Taiga Bean et al. 1988; Sibley & Ahlquist 1990; Harshman Goose and Tundra Bean Goose, geographic varia­ 1994; Bleiweiss et al. 1995), 12S and 16S ribo­ tion in size (Johansen 1945; Delacour 1951; somal RNA sequences (Hedges et al. 1995), (X­ Cramp & Simmons 1977; Roselaar 1977) is con­ crystallin sequences (Hedges et al. 1995; Caspers sistent with a clinal variation pattern (Sangster & et al. 1997) and mitochondrial DNA sequences Oreel 1996); therefore, johanseni and midden­ (Mindell et al. 1997). The clade formed by Anseri­ dorffi are included in A. fabaiis; rossicus is in­ formes and Galliformes was named Galloanserae cluded in A. serrirostris (Sangster & OreeI1996). by Sibley et al. (1988). Most of these analyses also indicate that Galloanserae is the sister taxon Branta hutchinsii Lesser Canada Goose of all extant birds except Paleognathae (Cracraft Kleine Canadese Gans 1986; 1988; Cracraft & Mindell 1989; Sibley & Branta canadensis Greater Canada Goose Ahlquist 1990; Hedges et ai. 1995; Kurochkin Grote Canadese Gans 1995; Caspers et al. 1997; Livezey 1997a). In con­ Lesser Canada Goose and Greater Canada formity with the suggestion of De Queiroz & Goose are specifically distinct (Sibley 1996) based Gauthier (1992) to list, of each pair of sister taxa, on congruence of phylogeographic analyses of the less speciose groups first, we propose to list mitochondrial DNA restriction fragments Anseriformes before Galliformes and to place (Shields & Wilson 1987; Shields 1988; Van Wag­ these taxa before the remaining taxa on the Dutch ner & Baker 1990; Quinn et al. 1991), mito­ list. chondrial DNA sequences (Quinn et al. 1991; Ba- Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 145 ker & Marshall 1997) and morphometry (Van Anas (King 1997). Current knowledge ofthe phylo­ Wagner & Baker 1990). Pending further analysis, genetic relationships of dabbling ducks is better leucopareia, minima and taverneri are provision­ represented with the recognition of two genera ally retained conspecific with B. hutchinsii; fulva, than with the placement of all species in Anas. interior, maxima, moffitti, occidentalis and par­ vipes are provisionally retained conspecific with Anas crecca Common Teal Wintertaling B. canadensis. Anas carolinensis Green-winged Teal Amerikaanse Wintertaling Branta bernicla Dark-bellied Brent Goose Common Teal and Green-winged Teal are spe­ Rotgans cifically distinct (Stepanyan 1990; Livezey 1991; Branta hrota Pale-bellied Brent Goose Gantlett et al. 1996; Johnson & Sorenson 1998) Witbuikrotgans based on qualitative differences in morphology Branta nigricans Black Brant Zwarte Rotgans (Delacour 1956; Johnsgard 1978; Livezey 1991). Dark-bellied Brent Goose, Pale-bellied Brent Goose and Black Brant are specifically distinct Melanitta nigra Common Scoter (Millington 1997) based on qualitative differences Zwarte Zee-eend in morphology (Delacour 1954; Johnsgard 1978; Melanitta americana Black Scoter Millington 1997), overlapping breeding ranges of Amerikaanse Zee-eend Pale-bellied Brent Goose and Black Brant in arc­ Common Scoter and Black Scoter are specifi­ tic Canada (Gavin 1947; Handley 1950) and segre­ cally distinct (Stepanyan 1990; Livezey 1995b; gation of Pale-bellied and Dark-bellied Brent Gantlett et al. 1996; King 1997) based on quali­ Goose in the Netherlands and Denmark (Lam­ tative differences in morphology (Johnsgard 1978; beck 1981). The alleged hybrid origin of 'interme­ Livezey 1995b). diate' populations in central arctic Canada, which formed the basis for including Pale-bellied Brent Melanittafusca Velvet Scoter Grote Zee-eend Goose and Black Brant, along with Dark-bellied Velvet Scoter and White-winged Scoter M. Brent Goose, in a single species (Delacour & deglandi are specifically distinct (Stepanyan Zimmer 1952), has been falsified by genetic anal­ 1990; Livezey 1995b; King 1997) based on quali­ ysis (Shields 1990). tative differences in morphology (Johnsgard 1978; Livezey 1995b). Mareca penelope Eurasian Wigeon Smient Mareca americana American Wigeon Mergellus albellus Smew Nonnetje Amerikaanse Smient Lophodytes cucullatus Hooded Merganser Mareca falcata Falcated Duck Bronskopeend Kokardezaagbek Mareca strepera Gadwall Krakeend A phylogenetic analysis based on morphology Phylogenetic analyses based on mitochondrial (Livezey 1995b) indicates that Smew is more DNA and morphology (Kessler & Avise 1984; closely related to the goldeneyes Bucephala than Livezey 1991) provide strong support for the exis­ to the mergansers Mergus. Because the inclusion tence of two major clades within the dabbling of Smew in Mergus would render Mergus poly­ ducks traditionally included in Anas (Omland phyletic, we adopt Livezey's (1995b) classifica­ 1994): (1) a clade formed by Cape Teal M. capen­ tion and place Smew in a monotypic genus Mer­ sis, the wigeons, Falcated Duck and Gadwall; and gellus (AOU 1983; BOURC 1997; King 1997). We (2) a clade formed by the remaining species. We place Hooded Merganser in the monotypic genus adopt the classification of Livezey (1991; 1997b) Lophodytes (AOU 1983; BOURC 1997; King in which the members of the former clade are 1997) to indicate its basal position among the mer­ placed in Mareca and the remaining species in gansers (Livezey 1995b). The latter species is fre- 146 ARDEA 87(1), 1999 quently recorded in the Netherlands but records extinct shearwaters, Hole's Shearwater P. holeae are considered to refer to escapes from captivity. (Walker et al. 1990) and Olson's Shearwater P. 01­ Hooded Merganser, therefore, is not formally ad­ soni (McMinn et al. 1990), which occurred sym­ mitted to the Dutch list. patrically in the eastern Canary Islands, casts doubt on the alleged sister relationship (and con­ Soft-plumaged petrel complex donsstormvogels specificity) of Balearic Shearwater and Yelkouan Fea's Petrel Pterodromafeae, Zino's Petrel P. Shearwater (Altaba 1995). It has been suggested madeira and Soft-plumaged Petrel P. mollis are that Balearic Shearwater may actually be more specifically distinct (Bourne 1983; Collar & Stuart closely related to Hole's Shearwater than to Yel­ 1985; Zino & Zino 1986; Sibley & Monroe 1990; kouan Shearwater (Walker et al. 1990). Beaman 1994; Hazevoet 1995; 1997; Sibley 1996; Snow & Pemns 1998) based on phylogeographic Calonectris borealis Cory's Shearwater analysis of mitochondrial DNA sequences (Nunn Kuhls Pijlstormvogel & Zino in press) and concordance ofdifferences in Cory's Shearwater and Scopoli's Shearwater morphology (Zino & Zino 1986), vocalisations C. diomedea are specifically distinct based on (Bretagnolle 1995) and reproductive behaviour phylogeographic analysis of allozymes (Randi et (Zino & Zino 1986). Analysis of mitochondrial al. 1989) and mitochondrial DNA (Heidrich et al. DNA sequences suggests that the divergence of P. 1996; 1998), qualitative differences in vocalisa­ feae and P. madeira occurred 840 000 years ago tions (Bretagnolle & Lequette 1990) and analysis and that P. mollis is not closely related to P. feae of morphological characters (Granadeiro 1993; and P. madeira (Nunn & Zino in press). Popula­ Gutierrez 1998). Cape Verde Shearwater C. ed­ tions of Fea's Petrel breeding on the Deserta wardsii is specifically distinct from Cory's Shear­ Islands, Madeira Cdeserta'), are provisionally re­ water and Scopoli's Shearwater (Bannerman & tained conspecific with P. feae (Snow & Pemns Bannerman 1968; Norrevang & den Hartog 1984; 1998). Non-monophyly of the soft-plumaged pet­ Hazevoet 1995; 1997; Sibley 1996; Hillcoat et al. rel complex precludes the recognition of a 'super­ 1997; Porter et al. 1997; Snow & Pemns 1998) species' taxon for P.feae, P. madeira andP. mollis. based on qualitative differences in morphology [There are no accepted records of P. feae, P. ma­ and vocalisations (Alexander 1898; Murphy 1924; deira or P. mollis in the Netherlands, although a re­ Bourne 1955; Bannerman & Bannerman 1968; cord at Camperduin, Noord-Holland (Stegeman et Hazevoet 1995; 1997; Porter et al. 1997; Snow & al. 1995), was accepted as P.feaelmadeiralmollis.] Pemns 1998). [In the Netherlands, all specimen re­ cords of Calonectris were identified as C. borealis Puffinus mauretanicus Balearic Shearwater (Van den Berg & Bosman 1999). The identity of Vale Pijlstormvogel sight records of C. borealislC. diomedea in the Puffinus puffinus Manx Shearwater Netherlands is currently being investigated by the Noordse Pijlstormvogel Dutch rarities committee (CDNA).] Balearic Shearwater is specifically distinct from Manx Shearwater and Yelkouan Shearwater Morus bassanus Northern Gannet Jan van Gent P. yelkouan (McMinn et al. 1990; Walker et al. Phylogenetic relationships among Sulidae 1990; Altaba 1995; Sibley 1996; King 1997; Hei­ (Warheit 1992; Friesen & Anderson 1997) are best drich et al. 1998; Snow & Perrins 1998; Wells represented with the recognition of Morus for the 1998) based on analyses of qualitative morpho­ gannets and Sula for the boobies, with the excep­ logical characters (Walker et al. 1990; Altaba tion of Abbott's Booby Papasula abbotti (Olson 1995) and phylogenetic analysis of mitochondrial 1985; Olson & Warheit 1988; Van Tets et al. 1988; DNA (Austin 1996; Heidrich et al. 1998). The re­ AOU 1989; Sibley & Monroe 1990; Warheit 1992; cent discovery and subsequent description of two Friesen & Anderson 1997). Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 147

Stictocarbo aristotelis European Shag Kuifaalscholver Porphyrio madagascariensis African Swamp-hen A phylogenetic analysis of morphological Smaragdpurperkoet characters (Siegel-Causey 1988) identified nine Porphyrio poliocephalus Grey-headed Swamp-hen major clades among the connorants and shags. Grijskoppurperkoet The classification proposed by Siegel-Causey Western Swamp-hen P porphyrio, African (1988), which recognises nine genera, better rep­ Swamp-hen, Grey-headed Swamp-hen, Philip­ resents the phylogenetic relationships of the cor­ pine Swamp-hen P pulverulentus, Black-backed morants and shags than the current inclusion of Swamp-hen P indicus and Australian Swamp-hen all species in a single genus Phalacrocorax P melanotus are specifically distinct (Sangster (Bourne & Casement 1996). 1998) based on qualitative differences in morphol­ ogy (Ripley 1977; Cramp & Simmons 1980; Del Casmerodius albus Great White Egret Hoyo et al. 1996). Analyses of mitochondrial Grote Zilverreiger DNA suggest that forms previously included un­ Phylogenetic analyses based on morphology der the name 'Purple Swamp-hen P porphyrio' and DNA-DNA hybridisation (Payne & Risley (Von Boetticher 1935; Ripley 1977; Del Hoyo et 1976; Sheldon 1987; Sheldon et al. 1995) indicate al. 1996) are paraphyletic with respect to two that Great White Egret is not closely related to the large flightless New Zealand taxa, South Island Egretta clade and instead suggest a closer rela­ Takahe P hochstetteri and extinct North Island tionship with Bubulcus and Ardea. However, Takahe P mantelli (Trewick 1997). These results given the unresolved relationships between Great argue against continued inclusion of swamp-hen White Egret, Bubulcus and Ardea, inclusion of forms in a single polytypic species. The six Great White Egret in Ardea (AOU 1995; BOURC groups here treated as species (P porphyrio, P 1997) is premature. Until the relationships of madagascariensis, P poliocephalus, P pulverul­ Great White Egret are better understood it is best entus, P indicus and P melanotus) are similar to placed in a monotypic genus Casmerodius (AOU those recognised by Roselaar (in Cramp & Sim­ 1983; Eck 1996; Inskipp et al. 1996; King 1997; mons 1980) as subspecies groups. Pending further Wells 1998). analysis, caspius and seistanicus are tentatively included in P poliocephalus; viridis is tentatively Phoenicopterus roseus Greater Flamingo included in P indicus; and bellus, chathamensis, Flamingo melanopterus, pelewensis and samoensis are ten­ Greater Flamingo and Caribbean Flamingo P tatively included in P melanotus. [The inclusion ruber are specifically distinct (Allen 1956; Mo­ of African Swamp-hen and Grey-headed Swamp­ rony et al. 1975; Hazevoet 1995; Sibley 1996; hen on the Dutch list is currently under review by King 1997; Sangster 1997a) based on qualitative the Dutch rarities committee (CDNA).] differences in plumage and bill pattern (Van den Berg 1987; Sangster 1997a) and display behaviour Chlamydotis macqueenii Macqueen's Bustard and vocalisations (Studer-Thiersch 1964; 1974; Oostelijke Kraagtrap 1975). Macqueen's Bustard and Houbara Bustard C. undulata are specifically distinct (Gaucher et al. Aquila nipalensis Steppe Eagle Steppearend 1996; Sangster 1996; King 1997; Wells 1998) based Steppe Eagle and Tawny Eagle A. rapax are on qualitative differences in courtship behaviour specifically distinct (Brooke et al. 1972; Clark and genetic analysis (Granjon et al. 1994; 1992; Olson 1994; King 1997; Wells 1998) based Gaucher et al. 1996). on qualitative morphological differences (Brooke et al. 1972; Clark 1992; Olson 1994). 148 ARDEA 87(1), 1999

Pluvialis dominicus American Golden Plover Phalaropus tricolor Wilson's Phalarope Amerikaanse Goudplevier Grote Franjepoot Pluvialis fulva Pacific Golden Plover Results of phylogenetic analyses based on al­ Aziatische Goudplevier lozymes (Dittmann et al. 1989), mitochondrial American Golden Plover and Pacific Golden DNA (Dittmann & Zink 1991) and morphology Plover are specifically distinct (Knox 1987; AOU (Chu 1995) are contradictory with regard to the al­ 1993; King 1997; Wells 1998) based on differences leged polyphyletic origin of the phalaropes in plumage, morphology, moult, vocalisations, (Sibley & Monroe 1990). Because of this incon­ ecology and overlap of breeding ranges (Connors gruence, the recognition of Steganopus for Wils­ 1983; Connors et af. 1993). The correct name of on's Phalarope (Sibley & Monroe 1990; Dowsett American Golden Plover is P. dominicus, not P. & Dowsett-Lemaire 1993; Beaman 1994; Del dominica (AOU 1995). Hoyo et al. 1996; Higgins & Davies 1996) is un­ justified and, therefore, we retain Wilson's Phala­ Vanellus gregarius Sociable Lapwing rope in Phalaropus. Steppekievit Vanellus leucurus White-tailed Lapwing Phalaropus fulicaria Grey Phalarope Witstaartkievit Rosse Franjepoot Phylogenetic analyses of behavioural charac­ The correct name of Grey Phalarope is P. fuli­ ters (Ward 1992) have been unable to resolve rela­ caria, not P. fulicarius (Parkes 1982). tionships among lapwings. Given the doubtful monophyly of Chettusia (and Hoplopterus), the Stercorarius skua Great Skua Grote Jager recognition of Chettusia (and Hoplopterus) is not Recent phylogenetic analyses of allozymes justified. Therefore, all lapwings are placed in and mitochondrial DNA sequences (Cohen et af. Vanellus (BOURC 1997; King 1997; Wells 1998). 1997; Braun & Brumfield 1998) confirm and extend previous suggestions based on short mitochondrial Gallinago gallinago Common Snipe Watersnip DNA sequences (Blechschmidt et al. 1993) and Common Snipe and Wilson's Snipe G. deli­ ecology (Andersson 1973) that Pomarine Skua S. cata are specifically distinct (Olsson 1987; Gan­ pomarinus is more closely related to species placed tlett et af. 1996; King 1997) based on qualitative in Catharacta than to Arctic Skua S. parasiticus differences in morphology, vocalisations and and Long-tailed Skua S. longicaudus. Because in­ drumming display (Thonen 1969; Cramp & Sim­ dependent lines of evidence suggest that Sterco­ mons 1983; Olsson 1987; Carey & Olsson 1995; rarius, as currently defined (Furness 1987; Chris­ Miller 1996a; 1996b; Gibson & Kessel 1997). tidis & Boles 1994; BOURC 1997; King 1997), is a Pending further analysis, faeroeensis and galli­ paraphyletic taxon, all skuas are placed in Sterco­ nago are provisionally retained as conspecific rarius. (Miller 1996b). African Snipe G. nigripennis, Madagascar Snipe G. macrodactyla, Paraguayan Larus graellsii Lesser Black-backed Gull Snipe G. paraguaiae, Magellan Snipe G. magel­ Kleine Mantelmeeuw lanica and Puna Snipe G. andina are specifically Larusfuscus Baltic Gull distinct from Common Snipe based on qualitative Baltische Mantelmeeuw differences in morphology, vocalisations and Lesser Black-backed Gull and Baltic Gull are drumming display (Tuck 1972; Sutton 1981; Hay­ specifically distinct based on qualitative differ­ man et af. 1986; Fjeldsa & Krabbe 1990; Del Hoyo ences in morphology and differences in moult and et af. 1996). ecology (Barth 1968; Bergman 1982; Cramp & Simmons 1983; Hario 1992; Strann & Vader 1992; Jonsson 1998a). There is no evidence that the Sangster et at.: TAXONOMIC CHANGES IN 1977-1998 149 form 'intermedius' is diagnosably distinct from Teyssedre 1984; Marion et aL 1985; Yesou 1991). graellsii; 'intermedius' is, therefore, considered Pontic Gull and Yellow-legged Gull are specifi­ conspecific with L. graellsii. cally distinct (Klein & Buchheim 1997; Klein & Heuglin's Gull L. heuglini is specifically dis­ Gruber 1997) based on qualitative differences in tinct from Lesser Black-backed Gull, Baltic Gull, morphology and vocalisations, and differences in Armenian Gull L. armenicus, Pontic Gull L. ca­ behaviour and ecology (Klein 1994; Gruber 1995; chinnans, Yellow-legged Gull L. michahellis and Jonsson 1996; 1998b; Garner 1997; Gamer & Vega Gull L. vegae based on qualitative differ­ Quinn 1997; Garner et aL 1997; Klein & Buch­ ences in morphology and behaviour, and differ­ heim 1997; Klein & Gruber 1997; Larsson & Lo­ ences in ecology (Grant 1986; Filchagov et aL rentzon 1998). Pending further analysis, atlantis is 1992; Hario 1992; Kennerley et aL 1995; Yesou & provisionally retained as conspecific with L. Hirschfeld 1997), The breeding range of Heug­ michahellis; barabensis and mongolicus are pro­ lin's Gull overlaps with that of Herring Gull and visionally retained as conspecific with L. cachin­ Baltic Gull, with evidence for reproductive isola­ nans. tion (Filchagov & Semashko 1987; Filchagov et Armenian Gull L. armenicus is specifically aL 1992; Filchagov 1994). Pending further analy­ distinct from Pontic Gull, Yellow-legged Gull and sis, taimyrensis and heuglini are provisionally re­ Heuglin's Gull L. heuglini based on qualitative tained as conspecific (Kennerley et al. 1995). differences in morphology and vocalisations (Geroudet 1982; Hume 1983; Dubois 1985; Grant Larus argentatus Herring Gull Zilvermeeuw 1986; 1987; Satat & Laird 1992; Buzun 1993; Fil­ Larus michahellis Yellow-legged Gull chagov 1993; Frede & Langbehn 1997; Yesou & Geelpootmeeuw Hirschfeld 1997). Larus cachinnans Pontic Gull Pontische Meeuw Herring Gull, Vega Gull and American Her­ Gelochelidon nilotica Gull-billed Tern ring Gull L. smithsonianus are specifically dis­ Lachstern tinct based on qualitative differences in morphol­ Although one phylogenetic study based on al­ ogy and vocalisations (Frings et aL 1958; Hoff­ lozymes suggests that Gull-billed Tern originates man 1979; Grant 1986; Mullarney 1990; Kennerley within the Sterna clade (Randi & Spina 1987), et aL 1995; Dubois 1997; Chu 1998). There is no others, based on allozymes (Hackett 1989), hind­ evidence that the form 'argenteus' is diagnosably limb musculature (McKitrick 1991) and osteology distinct from argentatus. Current evidence indi­ (Chu 1995), suggest a more distant relationship cates a clinal pattern of variation (Barth 1968; and do not support the inclusion of Gull-billed Cramp & Simmons 1983); the design of studies Tern in Sterna. Therefore, the inclusion of Gelo­ which have suggested clear differences between chelidon in Sterna (AOD 1983; Eck 1996; populations of 'argenteus' and argentatus (Mon­ BODRC 1997) is not warranted by present knowl­ aghan et aL 1983; Golley 1993) was inadequate to edge of phylogenetic relationships. Given the un­ substantiate such claims (Chylarecki 1993). The certainty about phylogenetic relationships among form 'argenteus' is, therefore, considered con­ terns, as indicated by the incongruence of avail­ specific with L. argentatus. able analyses, we retain Gelochelidon for Gull­ Yellow-legged Gull and Herring Gull are spe­ billed Tern. cifically distinct (Oree! 1980; Marion et aL 1985; Stepanyan 1990; Beaman 1994; King 1997) based Apus melba Alpine Swift Alpengierzwaluw on qualitative differences in adult and immature Evidence is lacking for a sister relationship plumages, bare parts, behaviour and vocalisations between Alpine Swift and Mottled Swift A. ae­ and overlap of breeding ranges (Nicolau-Guil­ quatorialis and monophyly of the other species laumet 1977; Glutz von Blotzheim & Bauer 1982; traditionally placed in Apus. Therefore, the recog- 150 ARDEA 87(1), 1999

mtlOn of Tachymarptis for Alpine Swift and Wagtail M. iberiae, Ashy-headed Wagtail M. ci­ Mottled Swift and a more restricted Apus for the nereocapilla, Yellow-headed Wagtail M. lutea, remaining species (Brooke 1972; Fry et al. 1988; Green-headed Wagtail M. taivana, Kamtchatka Short et al. 1990; Chantler & Driessens 1995; Eck Wagtail M. simillima, Alaska Wagtail M. tschut­ 1996) may render Tachymarptis and/or Apus para­ schensis and White-headed Wagtail M. leuco­ phyletic. In the absence ofa relevant phylogenetic cephala are specifically distinct based on qualita­ analysis of the swifts, we retain Alpine Swift in tive differences in morphology (Sushkin 1925; Jo­ Apus. hansen 1944; Voous 1950; Williamson 1955; Vau­ rie 1957; Dittberner & Dittberner 1984; Glutz von Merops persicus Blue-cheeked Bee-eater Blotzheim & Bauer 1985; Cramp 1988; Leader Groene Bijeneter 1996). For several named populations there is no Blue-cheeked Bee-eater and Madagascar Bee­ evidence or insufficient evidence that these are di­ eater M. superciliosus are specifically distinct agnosably distinct. Therefore, 'beema' is included (Glutz von Blotzheim & Bauer 1980; Eck 1996; in M. jlava, 'angarensis', 'macronyx' and 'plexa' King 1997; Wells 1998) based on qualitative dif­ in M. thunbergi, 'aralensis', 'kaleniczenkoi' and ferences in morphology (Fry 1984). 'melanogrisea' in M. feldegg and 'pygmaea' in M. cinereocapilla. The form 'dombrowskii' probably Anthus richardi Richard's Pipit Grote Pieper refers to hybrids ofM. jlava, M. thunbergi and M. Richard's Pipit is specifically distinct from feldegg (Vaurie 1957; Mayr & Greenway 1960). Grassland A. cinnamomeus, Paddyfield A. rufu­ The form 'superciliaris' most likely refers to hy­ Ius, Australian A. australis and New Zealand Pipit brids of M. jlava and M. feldegg (Vaurie 1957; A. novaeseelandiae (Glutz von Blotzheim & Mayr & Greenway 1960). The status of 'zaissa­ Bauer 1985; Sibley 1996; Wells 1998) based on nensis' remains unresolved. qualitative differences in plumage and vocalisa­ tions (Glutz von Blotzheim & Bauer 1985 and ref­ Motacilla alba White Wagtail Witte Kwikstaart erences cited therein). Motacilla yarrellii Pied Wagtail Rouwkwikstaart White Wagtail, Pied Wagtail, Moroccan Wag­ Anthus spinoletta Water Pipit Waterpieper tail M. subpersonata, Masked Wagtail M. person­ Anthus petrosus Rock Pipit Oeverpieper ata, Himalayan Wagtail M. alboides, Black-back­ Water Pipit, Rock Pipit and Buff-bellied Pipit ed Wagtail M. lugens, East Siberian Wagtail M. A. rubescens are specifically distinct (Oreel 1980; ocularis, Amur Wagtail M. leucopsis and Baikal AOU 1989; King 1997) based on qualitative dif­ Wagtail M. baicalensis are specifically distinct ferences in plumage, vocalisations and ecology based on qualitative differences in morphology (Bijlsma 1977; Alstrom & Olsson 1987; Knox (Glutz von Blotzheim & Bauer 1985; Cramp 1988a). 1988). There is no evidence that populations in western Asia (,dukhunensis') are diagnosably dis­ Motacillaflavissima Yellow Wagtail tinct from alba. Therefore, 'dukhunensis' is in­ Engelse Kwikstaart cluded in M. alba. The form 'persica' probably Motacillaflava Blue-headed Wagtail represents a variable hybrid population of alba Gele Kwikstaart and personata (Vaurie 1959; Cramp 1988) and is Motacilla thunbergi Grey-headed Wagtail not recognised. Noordse Kwikstaart Motacillafeldegg Black-headed Wagtail Saxicola rubicola European Stonechat Balkankwikstaart Roodborsttapuit Yellow Wagtail, Blue-headed Wagtail, Grey­ Saxicola maura Siberian Stonechat headed Wagtail, Black-headed Wagtail, Spanish Aziatische Roodborsttapuit Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 151

European Stonechat, Siberian Stonechat and 1983; Ishihara 1986; White & Bruce 1986; Christi­ African Stonechat S. torquata are specifically dis­ dis & Boles 1994; Gibbs 1996; Inskipp et al. 1996; tinct (Sibley 1996; Wells 1998) based on qualita­ Sibley 1996; King 1997; Wells 1998) based on tive differences in morphology (Cramp 1988; qualitative differences in morphology and vocal­ Svensson 1992) and phylogeographic analysis isations (Seebohm & Sharpe 1902; Jahn 1942; (Wittmann et al. 1995). There is no evidence that Mayr 1955; Ali & Ripley 1973; Ford 1983; Ishi­ populations inhabiting western Europe are dia­ hara 1986). gnosably distinct from those in central and north­ ern Europe. Therefore, the form 'hibernans' re­ Acrocephalus agricola Paddyfield Warbler presents a synonym of S. rubicola. There is no ev­ Veldrietzanger idence that populations inhabiting eastern Siberia Paddyfield Warbler and Manchurian Warbler ('stejnegeri') are diagnosably distinct from west­ A. tangorum are specifically distinct (Round 1994; ern Siberian populations. Therefore, 'stejnegeri' King 1997) based on qualitative differences in is included in S. maUTa (Svensson 1992). Pending plumage (Alstrbm et al. 1991; Round 1994). Phylo­ further analysis, variegata, armenica, indica and genetic analysis of mitochondrial DNA sequences przewalskii are provisionally retained as conspe­ indicates that Paddyfield Warbler and Manchu­ cific with S. maUTa. rian Warbler are not sister-taxa (Leisler et al. 1997). Paddyfield Warbler is considered mono­ Oenanthe hispanica Western Black-eared typic (Williamson 1968). Wheatear Westelijke Blonde Tapuit Oenanthe melanoleuca Eastern Black-eared Acrocephalus scirpaceus European Reed War· Wheatear Oostelijke Blonde Tapuit bier Kleine Karekiet Western Black-eared Wheatear and Eastern European Reed Warbler, Mangrove Reed Black-eared Wheatear are specifically distinct Warbler A. avicenniae, African Reed Warbler A. based on qualitative differences in morphology baeticatus and Caspian Reed Warbler A. fuscus (Clement & Harris 1987; Cramp 1988). are specifically distinct (Leisler et al. 1997; Sang­ ster 1997b) based on qualitative differences in Zoothera aurea White's Thrush Goudlijster morphology (Pearson 1981; Ash et al. 1989; Harris White's Thrush and Scaly Thrush Z. dauma et al. 1995). Phylogenetic analysis of mitochon­ are specifically distinct (Eck 1996) based on qual­ drial DNA sequences indicates that Mangrove itative differences in morphology and vocalisa­ Reed Warbler, which is currently regarded as a tions (Seebohm & Sharpe 1902; Ali & Ripley subspecies of A. baeticatus, is actually more 1973; Cramp 1988; Glutz von Blotzheim & Bauer closely related to European Reed Warbler, and 1988; Martens & Eck 1995). There is no evidence that European and Caspian Reed Warbler, until that populations in south-eastern Siberia, Russia recently regarded as subspecies of A. scirpaceus, and southern Kuril Islands, Japan ('toratugumi') are not sister-taxa (Leisler et al. 1997). are diagnosably distinct from aurea. Therefore, 'toratugumi' is included in Z. aurea. Acrocephalus caligatus Booted Warbler Amami Thrush Z. major, Nilghiri Thrush Z. neil­ Kleine Spotvogel gherriensis, Sri Lanka Thrush Z. imbricata, Hors­ Phylogenetic analyses of mitochondrial DNA field's Thrush Z. horsfieldi, Fawn-breasted Thrush sequences indicate that Booted Warbler and Oliv­ Z. machiki, New Britain Thrush Z. talaseae, San aceous Warbler A. pallidus are more closely re­ Cristobal Thrush Z. margaretae, Guadalcanal lated to species traditionally included in Acro­ Thrush Z. turipavae, Bassian Thrush Z. lunulata cephalus clade rather than to Icterine Warbler and Russet-tailed Thrush Z. heinei are specifically Hippolais icterina (Leisler et al. 1997; Sangster distinct (Mayr 1955; Deignan et al. 1964; Ford 1997b). Therefore, Booted Warbler and Oliv- 152 ARDEA 87(1), 1999 aceous Warbler are placed in Acrocephalus. Phylloscopus brehmii Iberian Chiffchaff Booted Warbler and Sykes's Warbler A. rama Iberische Tjiftjaf are specifically distinct (Stepanyan 1990; Glutz Common Chiffchaff, Iberian Chiffchaff and von Blotzheim & Bauer 1991; Sibley & Monroe Canarian Chiffchaff P. canariensis are specifi­ 1993; Sibley 1996; Wells 1998) based on qualita­ cally distinct (Gantlett et al. 1996; Eck 1996; tive differences in morphology and vocalisations, Sibley 1996; King 1997; Wells 1998) based on dif­ and differences in ecology (Portenko et al. 1976; ferences in structure and plumage (Erard & Salo­ Glutz von Blotzheim & Bauer 1991; Cramp 1992; mon 1989; Salomon et al. 1997), qualitative differ­ Hirschfeld 1994). Pending information on phylo­ ences in vocalisations (Salomon 1989; Salomon & genetic relationships of Sykes's Warbler, its Hemim 1992) and genetic analyses (Helbig et al. placement in Acrocephalus is tentative. 1993; 1996). Pending further analysis, exsul is ten­ tatively included in P. canariensis. Phylloscopus proregulus Pallas's LeafWarbler Common Chiffchaff, Mountain Chiffchaff P. Pallas' Boszanger sindianus and Caucasian Mountain Chiffchaff P. Pallas's Leaf Warbler and Lemon-rumped lorenzii are specifically distinct (Martens 1982; Warbler P. chloronotus are specifically distinct Snow & Pemns 1998) based on differences in (Inskipp et al. 1996; King 1997; Wells 1998) based structure and plumage (Shirihai 1987; Cramp on qualitative differences in plumage and vocal­ 1992), qualitative differences in vocalisations isations (Martens 1985; Alstrom & Olsson 1990). (Martens & Hanel 1981; Martens 1982) and ge­ netic analyses (Helbig et al. 1996). Pending fur­ Phylloscopus inornatus Yellow-browed Warbler ther analysis, abietinus and tristis are tentatively Bladkoning included in P. collybita. Phylloscopus humei Hume's Warbler Humes Bladkoning Lanius phoenicuroides Thrkestan Shrike Yellow-browed Warbler and Hume's Warbler Turkestaanse Klauwier are specifically distinct (Svensson 1992; Gantlett Lanius speculigerus Daurian Shrike et al. 1996; BOURC 1997; King 1997; Wells 1998) Daurische Klauwier based on qualitative differences in vocalisations Turkestan Shrike and Chinese Shrike L. isa­ and plumage and overlap of breeding ranges bellinus are specifically distinct (Kryukov 1995; (Mild 1987; Aistrom & Olsson 1988; Dathe & Panov 1995; Panow 1996) based on qualitative Loskot 1989). differences in morphology (Dean 1982; Cramp & Pemns 1993; Panow 1996; Lefranc & Worfolk Phylloscopus orientalis Eastern Bonelli's Warbler 1997) and analyses oftheir contact zone (Kryukov Balkanbergfluiter 1995). Phylloscopus bonelli Western Bonelli's Warbler Daurian Shrike is specifically distinct from Bergfluiter Turkestan Shrike and Chinese Shrike based on ·Eastern Bonelli's Warbler and Western Bonel­ qualitative differences in morphology (Dean li's Warbler are specifically distinct (Gantlett et 1982; Cramp & Pemns 1993; Panow 1996) and al. 1996; BOURC 1997; King 1997; Wells 1998) vocalisations (Panov 1995). Pending further anal­ based on qualitative differences in vocalisations ysis, tsaidamensis is provisionally retained as and genetic analyses (Helb et al. 1982; Helbig et conspecific with L. isabellinus. [An accepted re­ al. 1995). cord of an 'isabelline shrike' on Texel, Noord­ Holland, in May 1995 (Wassink 1996), is now con­ Phylloscopus collybita Common Chiffchaff sidered to refer to Daurian Shrike (Van den Berg Tjiftjaf & Bosman 1999). The identity of all records of 'isabelline shrike' in the Netherlands is currently Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 153 being investigated by the Dutch rarities commit­ Carduelis cannabina Linnet Kneu tee (CDNA).] Carduelis flavirostris Twite Frater Carduelis cabaret Lesser Redpoll Kleine Barmsijs Lanius excubitor Great Grey Shrike Klapekster Carduelisflammea Mealy Redpoll Grote Barmsijs Lanius pallidirostris Steppe Grey Shrike Carduelis hornemanni Arctic Redpoll Steppeklapekster Witstuitbarmsijs Great Grey Shrike and Southern Grey Shrike Published studies of phylogenetic relation­ L. meridionalis are specifically distinct (Eck 1994; ships among cardueline finches (Marten & John­ 1996; Gantlett et al. 1996; Inskipp et al. 1996; son 1986; Van den Elzen & Nemeschkal 1991; Sibley 1996; BOURC 1997; King 1997; Lefranc & Fehrer 1996) are contradictory with regard to the Worfolk 1997; Wells 1998) based on qualitative phylogenetic relationships of Acanthis and Car­ differences in plumage, breeding ecology and be­ duelis. Because monophyly of Acanthis has not haviour (Isenmann & Bouchet 1993; Eck 1994; been established and is contradicted by one study Lefranc 1995a; 1995b; Panow 1996; Lefranc & (Van den Elzen & Nemeschkal 1991), recognition Worfolk 1997). ofAcanthis may not significantly contribute to the Steppe Grey Shrike is specifically distinct elimination of paraphyletic taxa. Although Car­ from Great Grey Shrike and Southern Grey Shrike duelis as defined here (i.e. including Acanthis but (King 1997) based on qualitative differences in excluding Chloris) is still likely to be para­ plumage, breeding ecology, behaviour and over­ phyletic, other hypotheses of relationships, which lap of breeding ranges (Panov 1995; Panow 1996; require changes in nomenclature, do not seem to Lefranc & Worfolk 1997). be better supported by available data. Therefore, pending further phylogenetic analyses, Linnet, Corvus corone Carrion Crow Zwarte Kraai Twite, Lesser Redpoll, Mealy Redpoll and Arctic Corvus cornix Hooded Crow Bonte Kraai Redpoll are provisionally retained in Carduelis. Carrion Crow and Hooded Crow are specifi­ Lesser Redpoll and Mealy Redpoll are specif­ cally distinct (Stepanyan 1990; Gantlett et al. 1996; ically distinct based on qualitative differences in King 1997) based on qualitative differences in morphology (Knox 1988b; Herremans 1990) and plumage and analyses of their hybrid zone in Ger­ vocalisations (Herremans 1989), and overlap of many (Risch & Andersen 1998) and Italy (Saino breeding ranges in south-eastern Norway without 1990; Saino & Scatizzi 1991; Saino 1992; Saino & hybridisation (Lifjeld & Bjerke 1996). Pending Bolzern 1992; Saino & Villa 1992; Rolando 1993; further analysis, rostrata and exilipes are provi­ Rolando & Laiolo 1994; Rolando & Saino 1994). sionally retained as conspecific with C. fiammea and C. hornemanni, respectively. Chloris chloris Common Greenfinch Groenling Phylogenetic analyses of morphological char­ Dendroica coronata Myrtle Warbler acters (Raikow 1978; 1985) and short mitochon­ Mirtezanger drial DNA sequences (Fehrer 1996) provide con­ Myrtle Warbler and Audubon's Warbler D. gruent evidence that Chloris is more closely re­ auduboni are specifically distinct (Bermingham et lated to Pyrrhula than to Carduelis. Because two al. 1992) based on qualitative differences in plum­ independent studies suggest that Chloris is not age (Hubbard 1970; Kaufman 1979; Cramp & Per­ part of the Carduelis clade and identify the same rins 1994; Dunn & Garrett 1997) and analysis of sister-taxon (Raikow 1978; Fehrer 1996), whereas their hybrid zone (Barrowclough 1980). It has inclusion of Chloris in Carduelis is supported by been calculated (Zink & McKitrick 1995) that it only one study (Van den Elzen & Nemeschkal would take more than 6 million years for these 1991), recognition of Chloris for the greenfinches taxa to completely fuse. Data in Bermingham et is justified. al. (1992) show that many North American Den- 154 ARDEA 87(1), 1999

droica species are less than 3.5 million years old Alstrom P., U. Olsson & P.D. Round 1991. The taxo­ which means that speciation in several species of nomic status of Acrocephalus agricola tangorum. Forktail6: 3-13. Dendroica took place during a much shorter time Altaba c.R.. La sistematica i la conservaci6 de la biodi­ than the period calculated as necessary for Myrtle versitat: el cas de les baldritges (Puffinus). Anu. and Audubon's Warblers to fuse. Ornitol. Balears [1993] 8: 3-14. American Ornithologists' Union (AOU) 1983. Check­ Icterus galbula Baltimore Oriole list of North American Birds. Sixth edition. Baltimoretroepiaal American Ornithologists' Union, Washington, DC. American Ornithologists' Union (AOU) 1989. Thirty­ Baltimore Oriole, Bullock's Oriole I. bullockii seventh supplement to the American Ornithol­ and Black-backed Oriole I. abeillei are specifi­ ogists' Union Check-list of North American Birds. cally distinct (AOU 1995; Freeman & Zink 1995; Auk 106: 532-538. BOURC 1997) based on qualitative differences in American Ornithologists' Union (AOU) 1993. Thirty­ plumage and vocalisations, analyses of the hybrid ninth supplement to the American Ornithologists' Union Check-list of North American Birds. Auk zone of Baltimore Oriole and Bullock's Oriole 110: 675-682. (see AOU 1995 for references) and a phylogenetic American Ornithologists' Union (AOU) 1995. Fortieth analysis (Freeman & Zink 1995) indicating that supplement to the American Ornithologists' Union Baltimore Oriole and Bullock's Oriole are not Check-list of North American Birds. Auk 112: eachother's closest relatives. 819-830. Andersson M. 1973. Behaviour of the Pomarine Skua Stercorarius pomarinus Temm. with comparative remarks on Stercorariinae. Ornis Scand. 4: 1-16. ACKNOWLEDGEMENTS Andors AY. 1991. Paleobiology and relationships of the giant groundbird Diatryma (Aves: Gastornithi­ We thank Walter J. Bock and Ernst Mayr for providing formes). Acta. Congr. Int. Ornithol. 20: 563-571. valuable and stimulating comments and criticisms. We Andors AY. 1992. Reappraisal of the Eocene ground­ bird Diatryma (Aves: Anserimorphae). Nat. Hist. are grateful to Joel Cracraft for his support and encour­ Mus. Los Angeles County Sci. Ser. 36: 109-125. agement. As editor, Kees (C.J.) Camphuysen gener­ Ash J.S., D.J. Pearson, G. Nikolaus & P.R. Colston ously shared his time and offered helpful suggestions. 1989. The mangrove reed warblers of the Red Sea The work of the CSNA is supported by the Netherlands and Gulf ofAden coasts, with description of a new Ornithologists' Union (NOU) and the Dutch Birding subspecies ofthe African Reed Warbler Acroceph­ Association (DBA). alus baeticatus. Bull. Br. Ornithol. Club 109: 36­ 43. Austin U. 1996. Molecular phylogenetics of Puffinus shearwaters: preliminary evidence from mitochon­ REFERENCES drial cytochrome b gene sequences. Mol. Phylo­ genet. Evol. 6: 77-88. Alexander B. 1898. An ornithological expedition to the Baker AJ. & B.D. Marshall 1997. Mitochondrial con­ Cape Verde Islands. Ibis (7) 4: 74-118. trol region sequences as tools for understanding Ali S. & S.D. Ripley 1973. Handbook of the birds of evolution. In: Mindell D.P. (ed.) Avian molecular India and Pakistan with those of Bangladesh, Ne­ evolution and systematics: 51-82. Academic Press, pal, Sikkim, Bhutan and Sri Lanka, 9. Oxford Uni­ San Diego. versity Press, Bombay. Bannerman D.A & W.M. Bannerman 1968. History of Allen R.P. 1956. The flamingos: their life history and the birds of the Cape Verde Islands. Oliver & survival. Res. Rep. Nat. Audubon Soc. 5. Boyd, Edinburgh. Alstrom P. & K. Mild 1987. Some notes on the tax­ Barrowclough G.P. 1980. Genetic and phenotypic dif­ onomy of the Water Pipit complex. Proc. 4th Int. ferentiation in a wood warbler (genus Dendroica) Identif. Meet. Eilat Nov. 1986: 47-48. hybrid zone. Auk 97: 655-668. Alstrom P. & U. Olsson 1988. Taxonomy of Yellow­ Barth E.K. 1968. The circumpolar systematics ofLarus browed Warblers. Br. Birds 81: 656-657. argentatus and Larusfuscus with special reference A1strom P. & U. Olsson 1990. Taxonomy of the Phyl­ to the Norwegian populations. NyU Mag. Zool. 15 loscopus proregulus complex. Bull. Br. Ornithol. Suppl. 1: 1-50. Club 110: 38-43. Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 155

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pelago. Treubia 20: 647-656. SAMENVATTING Voous KH. 1964. Het genusbegrip in de zoOlogie in theorie en praktijk. Vakblad BioI. 44: 139-149. De Commissie Systematiek Nederlandse Avifauna Voous K.H. 1977. List of recent Holarctic bird species. (CSNA), een adviserende commissie van de Neder­ British Ornithologists' Union, London. landse Ornithologische Unie (NOU) en de Dutch Bir­ Walker c.A., G.M. Wragg & c.J.O. Harrison 1990. A new shearwater from the Pleistocene ofthe Canary ding Association (DBA), heeft zich gebogen over inter­ Islands and its bearing on the evolution of certain nationale systematisch-biologische studies die betrek­ Puffinus shearwaters. Hist. BioI. 3: 203-224. king hebben op vogelsoorten die ook op de Neder­ Ward D. 1992. The behavioural and morphological af­ landse lijst voorkomen. Op grond van deze gepubli­ finities of some vanelline plovers (Vanellinae: ceerde studies en de evaluatie ervan heeft de CSNA een Charadriiformes: Aves). J. ZooI. 228: 625-640. aantal besluiten genomen ten aanzien van door haar Warheit KI. 1992. The role ofmorphometries and clad­ noodzakelijk geachte taxonornische en nomenclatori­ istics in the taxonomy of fossils: a paleornithologi­ sche wijzigingen. Dit artikel bevat een samenvatting cal example. Syst. BioI. 41: 345-369. van de genomen besluiten en de eraan ten grondslag WassinkA. 1996. Izabelklauwier op Texel in mei 1995. Dutch Birding 18: 129-131. liggende argumenten. Wells M.G. 1998. World bird species checklist. World­ Een belangrijk deel van de taxonornische verande­ list, Bushey, Herts. ringen betreft de soortstatus van bepaalde taxa, die White C.M.N. & M.D. Bruce 1986. The birds ofWalla­ vroeger veelal de status van ondersoort hadden. Dit is cea. British Ornithologists' Union, London. een resultaat van een verandering in de wetenschappe­ Wiley E.O. 1981. Phylogenetics. Wiley, New York. lijke inzichten in het soortsbegrip. Traditioneel wordt Williamson K 1955. Migrational drift and the Yellow er in de ornithologie gebruikgemaakt van het isolatie­ Wagtail complex. Br. Birds 48: 382-403. soortsbegrip, ook weI genoemd het biologische soorts­ Williamson K 1968. Identification for ringers 1. The begrip. Dit soortsbegrip stelt dat het belangrijkste crite­ genera Cettia, Locustella, Acrocephalus and Hip­ rium bij het onderscheiden van soorten de kruisbaar­ polais. Third edition. British Trust for Ornithol­ ogy, Tring. heid betreft: als individuen uit twee verschillende po­ Wittmann u., P. Heidrich, M. Wink & E. Gwinner pulaties met elkaar kunnen kruisen, dan behoren zij tot 1995. Speciation in the Stonechat (Saxicola tor­ dezelfde soort. Het is echter allang bekend, dat er pro­ quata) inferred from nucleotide sequences of the blemen aan dit, op het eerste gezicht zo logischlij­ cytochrome-b gene. J. ZooI. Syst. EvoI. Research kende, isolatie-soortsbegrip kleven. Zo blijkt het in de 33: 116-122. natuur nogal eens voor te komen dat individuen uit Yesou P. 1991. The sympatric breeding of Larus fus­ twee groepen weI met elkaar kunnen kruisen, terwijl zij cus, L. cachinnans and L. argentatus in France. niet elkaars nauwste verwanten zijn (in de zin van Ibis 133: 256-263. bloedverwantschap, genealogie, afstamming). Daar­ Yesou P. & E. Hirschfeld 1997. Which large gulls from naast gaat het isolatie-soortsbegrip er bij het afbakenen the Larus fuscus-cachinnans-argentatus complex van soorten vanuit dat deze soorten in de toekomst re­ of (sub)species occur in Bahrain? Sandgrouse 19: 111-121. productief van elkaar gei"soleerd zullen blijven en niet Zink R.M. 1988. Evolution of Brown Towhees: allo­ tot een populatie zullen fuseren. Dit betekent, dat in zymes, morphometries and species limits. Condor feite toekomstige gebeurtenissen uitsluitsel moeten ge­ 90: 72-82. ven of we huidige groepen als aparte soorten of slechts Zink R.M. 1997. Species concepts. Bull. Br. OrnithoI. als populaties van een soort moeten beschouwen. Ten­ Club 117: 97-109. slotte heeft praktische toepassing van het isolatie­ Zink R.M. & M.C. McKitrick 1995. The debate over soortsbegrip geleid tot het onderscheiden van samenge­ species concepts and its implications for ornithol­ stelde, polytypische soorten. Hoewel bepaalde popula­ ogy. Auk 112: 701-719. ties weI degelijk onderscheidbaar zijn, worden zij toch Zino P.A. & F. Zino 1986. Contribution to the study of tot slechts een soort gerekend (omdat de onderlinge the petrels of the genus Pterodroma in the archi­ pelago of Madeira. BoI. Mus. Mun. Funchal 38: verschillen als te gering worden beoordeeld); deze han­ 141-165. delswijze resulteert in een onderwaardering van de fei­ telijke biodiversiteit. Bovengenoemde en andere problemen met het iso­ latie-soortsbegrip hebben veel biologen ertoe aangezet Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 165

om te trachten soorten op een andere manier af te bake­ rodius) wordt geplaatst. Veel studies hebben aange­ nen. En zo zijn er in de loop van de tijd vele altema­ toond dat de eendachtigen (Anseriformes) en hoender­ tieve soortsbegrippen geformuleerd. De CSNA acht het achtigen (Galliformes) elkaars nauwste verwanten zijn. zogenaamde fylogenetische soortsbegrip van groot be­ Daarom kunnen we deze twee in een groep van hogere lang voor avifaunistische lijsten, de systematische bio­ orde plaatsen, de Galloanserae. logie en biodiversiteitsstudies in het algemeen. Ret fy­ De eerder genoemde revolutionaire ontwikkeling logenetische soortsbegrip is een uitvloeisel van een we­ binnen de systematische biologie en de evolutiebiolo­ tenschappelijke revolutie in een iets eerdere periode gie, en de moderne opvatting dat de nieuwe inzichten van de systematische biologie. Als gevolg hiervan heeft en resultaten hun reflectie dienen te hebben op de be­ nu alom het idee postgevat dat kennis van en inzicht in staande taxonomische systemen, hebben directe prakti­ de afstammings- of fylogenetische relaties tussen groe­ sche consequenties. Ret betekent dat de oude taxono­ pen van doorslaggevend belang is in het vakgebied van mische systemen en avifaunistische lijsten gaan veran­ de systematische biologie en de evolutiebiologie. Dit deren. Er is weI beargumenteerd dat een groot voordeel heeft als consequentie dat van een modem soortsbegrip van de oude systemen en lijsten is dat ze stabiel zijn: verwacht mag worden dat het in overeenstemming is hun onveranderlijkheid geeft houvast bij het opzoeken met de fylogenetische relaties tussen de onderscheiden van informatie. Ret is echter een feit dat deze stabili­ eenheden. Soortsbegrippen, zoals het isolatie-soortsbe­ teit, zo niet starheid, tot gevolg heeft dat de oude avi­ grip, die groepen van individuen bij elkaar zetten die faunistische en taxonomische lijsten geen afspiegeling niet elkaars nauwste verwanten zijn, geven dientenge­ meer vormen van verkregen wetenschappelijke inzich­ volge een vertekend beeld van de evolutionaire ge­ ten en hun relatie met de huidige stand van de systema­ schiedenis. Fylogenetische soortsbegrippen voldoen in tische biologie en de evolutiebiologie deels hebben ver­ dit opzicht beter. loren. Als dergelijke lijsten geen afspiegeling meer vor­ Er zijn twee typen van fylogenetische soortsbegrip­ men van modeme inzichten in de fylogenie en taxono­ pen, te weten de monofylie-versie en de diagnostische mie van vogels, dan rijst onmiddellijk de vraag wat die versie. De monofylie-versie van het fylogenetische lijsten en systemen dan weI vertegenwoordigen. Ret soortsbegrip eist dat elke soort gekenmerkt wordt door klassieke antwoord dat zij ingeburgerde en gemakke­ minimaal een uniek kenmerk dat niet bij andere soorten lijke zoeksystemen vertegenwoordigen, is wetenschap­ voorkomt. De diagnostische versie is wat minder strikt pelijk gezien niet acceptabel. Als wetenschappelijke en eist slechts dat een soort gekarakteriseerd wordt onderbouwing niet meer van belang wordt geacht, dan door een unieke combinatie van kenmerken; d.w.z. de zijn er praktischere zoeksystemen denkbaar, bijvoor­ afzonderlijke kenmerken mogen weI bij andere soorten beeld alfabetische lijsten. Daarom is de CSNA van me­ voorkomen, maar hun combinatie moet uniek zijn. De ning dat taxonomische systemen geregeld dienen te CSNA heeft de diagnostische versie van het fylogeneti­ worden aangepast om zo een afspiegeling te zijn van de sche soortsbegrip gehanteerd bij het samenstellen van meest modeme hypothesen over de fylogenetische rela­ deze rapportage. ties tussen groepen van vogels. Ret is evident, dat bij Een tweede aspect van de avifaunistiek en de taxo­ een voortschrijdende wetenschap ook taxonomische nomie dat door de CSNA aan een nadere beschouwing systemen en lijsten niet stil kunnen blijven staan en al­ is onderworpen, betreft het groeperen van soorten in tijd aan verandering onderhevig zullen zijn. hogere eenheden, zoals geslachten en families, en de indeling van de lijst (dat wil zeggen welke groepen Received 11 September 1998, accepted 29 March 1999 staan bovenaan en welke onderaan de lijst). Ook deze Corresponding editor: Kees (C.J.) Camphuysen zaken dienen, naar huidige inzichten, een afgeleide te zijn van de historische, fylogenetische relaties tussen de onderscheiden groepen. Twee voorbeelden uit de Note from the editors ofArdea huidige lijst kunnen dit duidelijk maken. Fylogeneti­ The taxonomic and nomenclatural changes listed sche studies hebben aangetoond dat de Grote Zilverrei­ above, as proposed by the Dutch committee for avian ger niet nauw verwant is aan de andere zilverreigers. systematics (CSNA), will be adopted by Ardea in this Dat betekent dat we de Grote Zilverreiger niet in het­ and future issues. With this decision, we follow recom­ zelfde geslacht (Egretta) kunnen handhaven als de an­ mendations of the Netherlands Ornithologists' Union dere zilverreigers en zij in een ander geslacht (Casme- (NOU) and the Dutch Birding Association (DBA). In 166 ARDEA 87(1), 1999 the review process of this manuscript, it became clear logical journals and books in The Netherlands have that there is still considerable opposition to the princi­ adopted the changes while others decided to continue ples underlying the present changes. Some referees to use more traditional lists. The present situation, a were not convinced of the advantages of the Phylo­ split into two ('followers' and 'non-followers'), is very genetic Species Concept (PSC). Not surprisingly, these inconvenient. Both the NOV and the editorial team of referees could not recommend the publication of this Ardea are convinced that a decision has to be made, so paper in Ardea. Other referees were completely in that the CSNA could either continue to function as an favour of the principles of a PSC and strongly recom­ advisory committee of NOVIDBA, or should operate mended the present publication. Apparently, there was independently. little or no ground for compromise. In the sometimes After the publication of this list, the editorial team heated discussions generated by the proposals of the of Ardea will seek advice from CSNA regarding the CSNA (e.g. at the NOV symposium in Naturalis, Lei­ name and taxonomic status of species of birds men­ den, 10 April 1999), it became clear that scientific argu­ tioned in the articles published in this journal, includ­ ments were mixed with practical reasons, such as a de­ ing those not featuring on the Dutch list. For conve­ sire to have a 'stable' list rather than (frequent) changes. nience, and to avoid misunderstanding, we will inform Our decision to publish the list in Ardea was inspired the readers about the name/status of these species as by a similar mix of reasons. We believe that the CSNA they were known prior to the present changes. Because has convincingly defended their proposal to abandon the CSNA is the only body in The Netherlands that is the traditional Isolation Species Concept (ISC) in fa­ specialised in the evaluation of the taxonomic status vour of a Phylogenetic Species Concept, both in the in­ and systematics ofbirds, we recommend that proposals troductory statements in the present publication and at published in Ardea by this committee are followed. At the NOV symposium in Leiden in 1999. Yet, it is clear the same time, we challenge scientists to constructively that we deal with hypotheses, and CSNA has confirmed comment on CSNA decisions and proposals, so that, in that changes in specific status, sequences or positions the process, we deepen our understanding of evolution­ on the list are and will be proposed only ifthe evidence ary relationships and patterns of ancestry and descent is 'overwhelming'. Ever since the earlier publications of contemporary birds. of CSNA in Dutch Birding, several editors of ornitho-