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. Ar­dea 87: 139-165.

lNieuwe Rijn 27, 2312 JD Leiden, Netherlands; E-mail: g.sangster@tele2.nl; 2Insti­tute of Systematics and Population Biology, Zoological Museum, University of Am­sterdam, P.O. Box 94766, 1090 GT Amsterdam, Netherlands; 3Museu e Laborat6rioZool6gico e Antropol6gico (Museu Bocage), Rua da Escola Politecnica 58, 1250Lisboa, Portugal; 4Duinlustparkweg 98, 2082 EG Santpoort-Zuid, Netherlands

Key words: systematics - taxonomy - phylogeny - species concepts - species - highertaxa

e9

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

INTRODUCTION

This report includes taxonomic and nomenclatu­ral changes adopted by the Dutch committee foravian systematics (Commissie Systematiek Ne­derlandse Avifauna, CSNA) since Voous (1977).Proposals affecting the Dutch list that were re­jected by the CSNA are also discussed. The Neth­erlands Ornithologists' Union (NOU) and theDutch Birding Association (DBA) support theCSNA and its reports are published in both Ardeaand Dutch Birding. The present report is essen­tially similar to two separate publications inDutch Birding (Sangster et al. 1997; 1998). Thecommittee currently consists of five members(year of election between parentheses): ArnoudB. van den Berg (1995), Cornelis J. Hazevoet(1996), C.S. (Kees) Roselaar (1995), GeorgeSangster, Secretary (1996) and Ronald Sluys

(1998). Due to limitations of space, the presentreport only includes a brief summary of the ratio­nale for each decision. More extensive and ex­plicit motivations for the recognition of some ofthe species-level and higher taxa listed here willbe published elsewhere.

Choice of species concept and its applicationOver the past few years the fields of system­

atic and evolutionary biology have been invigo­rated by a scientific discussion on the concept ofspecies (for reviews, see Sluys 1991; Mayden1997; Zink 1997). Although this discussion contin­ues, it is also true that new and useful insightshave been gained already that can be incorporatedfruitfully into an up to date and modem descrip­tion of avian diversity. One of the insights that hassurfaced in the literature is that different speciesconcepts may be best for different purposes

140 ARDEA 87(1),1999

(Endler 1989) and it seems increasingly likely thatno single species concept will satisfy the multiplepurposes of 'species' in the biological sciences(Sluys 1991; Hull 1997), However, a theme com­mon to all biological sciences is that all taxa arehistorically connected through their pattern of an­cestry and descent. All living taxa are the productof history, and we can understand little about theirdiversity without knowledge of their history, i.e.the phylogenetic knowledge provided by system­atics (O'Hara et al. 1988). Virtually all compara­tive studies of biological variation within andamong taxa depend on such phylogenetic knowl­edge for interpretation (Felsenstein 1985; Brooks& McLennan 1991; Harvey & Pagel 1991). There­fore, in biodiversity studies and comparative biol­ogy a fundamental requirement of species-leveltaxa is that they are compatible with the recon­struction of evolutionary relationships. Speciesconcepts which group taxa that are not closely re­lated in a single species misrepresent evolution­ary history. Second, species-level taxa should bedelimited on the basis of historical subdivisions(i.e. historical patterns), rather than present-dayor possible future interactions and processes(Liden & Oxelman 1989), such as hybridisationand gene flow. Species concepts which are pro­spective and which require speculations about thefuture are not helpful in biology; since all of ourdata are of the present and past, the units bywhich we interpret these data must also be strictlyhistorical (Maddison 1997). Third, species shouldbe basal, taxonomically comparable units (Cra­craft 1987; 1989): species should be basal taxa,that is, taxa that contain no included taxa. A spe­cies concept should not combine several distincttaxa in a single (polytypic) 'species' because such'species' actually are higher, more inclusive taxa.Species concepts which recognise not only singleunits (monotypic species) but also polytypic as­semblages (polytypic species) as 'species' runcounter to the fundamental need for species-leveltaxa to be basal and comparable.

The decision by the CSNA to abandon the tra­ditional Isolation Species Concept (ISC) in favourof a Phylogenetic Species Concept (PSC) was lar-

gely based on these views. The Isolation SpeciesConcept (popularly known as the 'BiologicalSpecies Concept') is rejected because its proper­ties violate all three aforementioned principles.First, interbreeding taxa are not necessarily moreclosely related to each other than they are to taxafrom which they are reproductively isolated. Be­cause interbreeding is the prime criterion for con­specificity under the ISC, the ISC could still re­gard such interbreeding taxa as conspecifics.Therefore, the problem of lumping taxa which arenot closely related in a single species and, hence,the misrepresentation of evolutionary history, isinherent to the ISC and does not simply resultfrom errors in application. Phylogenetic analysesindicate that in various groups of birds 'polyty­pic' species recognised by the ISC do not repre­sent natural (monophyletic) groups. Evidencecomes from phylogenies based on both morpho­logical (Livezey 1995a; Chu 1998; Veron 1999) andmolecular data sets (Zink 1988; Friesen et al. 1996;

Leisler et al. 1997; Roy et al. 1997; Trewick 1997).Second, under the ISC taxa are recognised as spe­cies if they remain 'reproductively isolated' , in thesense that they do not fuse into a single population(Mayr 1982; 1996). The ISC, therefore, is prospec­tive (O'Hara 1993; 1994; Maddison 1997); only fu­ture events will show whether currently recog­nised taxa remain reproductively isolated or fuseinto each other. This poses both theoretical andpractical problems. It makes little sense to try tointerpret the past and present diversity of organ­isms with a taxonomy that is based on expecta­tions ('dreams', Maddison 1997) about the future.A practical problem is that, except for rare cases,the process of fusion transcends observable time.The likely time-to-fusion may be measured inWOOs or even millions of years (Zink & McKi­trick 1995). Third, many 'species' recognised bythe ISC contain more than one taxon. The ISC rec­ognises monotypic species but may also unite upto 10 or more diagnosable taxa and still recognisethe resulting unit as a single 'species'. This notonly underestimates and misrepresents biodiver­sity but also compromises interspecific compari­sons (Prum 1994; Hazevoet 1996; Cracraft 1997).

Sangster et at.: TAXONOMIC CHANGES IN 1977-1998 141

Two distinct Phylogenetic Species Conceptshave been advocated. These versions agree inviewing species as products of evolution, not asplayers in evolution, and support the notion ofspecies as basal taxa. The original version, pro­posed by Cracraft (1983) and further developed byNixon & Wheeler (1990) and Davis & Nixon(1992), considers a phylogenetic species to be anirreducible cluster of organisms possessing atleast one diagnostic character state. This versionthus focuses on the diagnosability of species. Dia­gnostic character states are discrete characterstates which are fixed within the species and areabsent from close relatives. Diagnosability ofspecies may be based on any intrinsic attribute,either morphological, molecular, ethological or acombination of these. It should be emphasisedthat this concept is populational (Cracraft 1997);the criterion of diagnosability applies to (groupsof) populations, and not to, e.g. family groups orindividuals. An alternative approach was devel­oped by Donoghue (1985) and De Queiroz &Donoghue (1988; 1990) and considers phylogen­etic species to be the smallest monophyleticgroups of organisms supported by autapomorphies(unique derived character states).

The diagnosability and monophyly versionsof the PSC are significantly different; both in the­ory and application (Baum 1992; Davis 1997). Amajor difference is that the monophyly versionrequires phylogenetic analysis prior to delimitingspecies, whereas the diagnosability version doesnot. Implementation of the monophyly versionwill, therefore, result in a significant shift in taxo­nomic practice, whereas the diagnosability ver­sion will not (Baum 1992). In addition to thispractical difference, the two versions also differwith regard to which basal taxa are called 'spe­cies'. When a small group of related individualsleaves a species, evolves diagnostic characterstates and thus forms a descendant species, theancestral 'species' will cease to be monophyletic.Because both the ancestral and descendant 'spe­cies' are characterised by diagnostic characterstates, the diagnosability version would recogniseboth as phylogenetic species; the monophyly ver-

sion, however, would recognise the descendantspecies, but not the ancestral 'species', as a phy­logenetic species because only the former ismonophyletic. Proponents of the monophyly ver­sion propose that such ancestral 'species' are rec­ognised as a separate class of species, for whichthey propose the term 'metaspecies' (Donoghue1985; De Queiroz & Donoghue 1988).

The CSNA has adopted the diagnosabilityversion of the PSC as its operational species con­cept because no phylogenetic analysis is requiredprior to delimiting species, its implementation in­volves little modification of existing taxonomicpractices and does not require the recognition ofadditional classes of species. It is believed thatthese are advantages over the monophyly version,and similar versions (Baum & Shaw 1995), andthat these outweigh any disadvantages. Althoughboth aspects (diagnosability and monophyly) maybe meaningful at the level of species (Baum1992), the monophyly version is not sufficientlypractical as an operational species concept.

It has been argued that the PSC actually de­fines the evolutionary lineage and does not repre­sent a species concept at all (Bock 1979; 1994;Szalay & Bock 1991). This complaint stems froma narrow view of species, based on the ISC,which restricts the term 'species' to entities whichare involved in the process of evolutionarychange. According to this view, species are realonly at a single point in time because repro­ductive isolation and processes such as natural se­lection operate only in a single time-slice. Al­though the ISC can not be applied to organismsliving at different times, it does not follow thatother species concepts should also be applicablein one time-slice only. In any case, the PSC doesnot define species in terms of reproductive isola­tion (which, indeed, can only be assessed at agiven moment in time) but in terms of diagnosticcharacter states. The presence of diagnostic char­acter states in a population may extend over timeand, therefore, phylogenetic species-level taxawill have reality over time. However, this doesnot mean that the PSC defines the evolutionarylineage, as claimed by Szalay & Bock (1991) and

142 ARDEA 87(1), 1999

Bock (1994). Since diagnostic character statesmay become fixed in a population after a lineagehas split, speciation may be completed after theorigin of a lineage and, therefore, phylogeneticspecies are not synonymous with evolutionarylineages.

Throughout this paper, 'qualitative differ­ences' are differences that can be coded as dis­crete character states. Published data on diagnos­tic character states of taxa were evaluated andinterpreted with reference to the diagnosabilityversion of the PSc. This does not imply that theauthors of the original publications necessarilyproposed the relevant taxa to be considered asspecies-level taxa or explicitly endorse a PSC. Infact, many of the authors concerned - implicitlyor explicitly - worked under the umbrella of theISC. The present interpretation of the data is fullythe responsibility of the CSNA. In the absence ofa fully documented system of phylogeneticallydefined species level taxa, the CSNA continues touse subspecific names to denote the likely geo­graphic origin of the populations occurring in theNetherlands.

Phylogeny and higher taxaAlthough the study of phylogeny assumed an

important place in biology in the decades follow­ing Darwin's Origin of species (Darwin 1859),biologists' interest in phylogeny was sub­sequently replaced by new emphases on the pro­cesses and mechanisms of genetics, developmentand evolution (e.g. Dobzhansky 1937; Mayr1942). With the 'Evolutionary Synthesis' of the1930s and 1940s a dichotomy developed betweenthe study of species and the study of phylogenyand higher taxa. Studies at the level of popula­tions and species proceeded with great vigour, butthe study of phylogeny suffered from a lack ofconceptual and methodological clarity. For in­stance, it was not clear whether the term 'phylo­geny' should only denote the branching pattern ofevolutionary history or whether it should also in­clude reference to the level of divergence subse­quent to cladogenesis. Also, it was commonly be­lieved that, in contrast to species, higher taxa are

artificial and without biological relevance (e.g.Voous 1964). Above all, there was no consistentmethod to reconstruct phylogeny.

Hennig (1966) restricted the term 'phylogeny'to the branching pattern of evolutionary history,argued convincingly that higher taxa (i.e. mono­phyletic groups) are real, and that these can be re­constructed by phylogenetic analysis (sub­sequently termed cladistics). The consistency andlogic of phylogenetic systematics was greetedwith enthusiasm by systematists (e.g. Nelson1970; Brundin 1972; Cracraft 1972) and is now al­most universally accepted (e.g. Wiley 1981; Hilliset al. 1996; Dingus & Rowe 1998). One reason forits success is that units of study in biology (fromgenes through organisms to higher taxa) do notrepresent statistically independent observationsbut rather are interrelated through their historicalconnections. Therefore, almost any comparativestudy requires information on phylogeny (Felsen­stein 1985; Brooks & McLennan 1991; Harvey &Pagel 1991). As a result, phylogenetic analyses arenow firmly entrenched in contemporary biologyand new applications continue to appear (e.g.Harvey et al. 1996). Being the most central andunifying concept in biology, phylogeny shouldalso be a central principle in taxonomy.

In line with phylogenetic theory and method­ology (Hennig 1966; Wiley 1981; de Queiroz &Gauthier 1992), two basic rules are applied for therecognition of higher taxa (i.e. taxa above the levelof species): (1) higher taxa are named clades and,therefore, represent monophyletic groups of spe­cies or less inclusive clades; hence, higher taxaare delimited on the basis of common ancestry,rather than a shared set of character states; (2)phylogenetic knowledge should be expressed asaccurately as possible in nomenclature. As a con­sequence of the first rule, para- and polyphyletictaxa should be abandoned. This has resulted in therecognition of Casmerodius for Great White Eg­ret C. aibus and Mergellus for Smew M. aibellusand in the transfer of some species of Hippolais toAcrocephalus.

The second rule implies that adjustments tothe present system should be enacted when im-

Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 143

proved phylogenetic knowledge becomes avail­able. For instance, in the past 15 years, evidencefor a sister-group relationship of Anseriformesand Galliformes has accumulated and now greatlyoutweighs all other hypotheses of relationships ofthese orders. This has resulted in the recognitionof the taxon Galloanserae. More detailed knowl­edge of the relationships among cormorants andshags, gannets and boobies, dabbling ducks, andcardueline finches has resulted in the recognitionof four additional genera, Stictocarbo, Morus,Mareca and Chloris. Generally, established usageis maintained unless alternative hypotheses arebetter supported. As a consequence, Wilson'sPhalarope Phalaropus tricolor is not placed inSteganopus and Gelochelidon is not included inSterna. It has been attempted to not recognisehigher taxa that are not or only weakly supportedby phylogenetic analyses. This has resulted in theinclusion of Chettusia in Vanellus, Catharacta inStercorarius and Tachymarptis in Apus.

Systematics and taxonomic (in)stabilityThere is a large hiatus between our knowledge

of phylogenetic relationships, which has greatlyincreased since the late 1970s, and the avian taxo­nomic system (classification), which is basicallystill the same as the one proposed by AlexanderWetmore in 1930, which in tum was largely basedon the work of Max Ftirbringer and Hans Gadowin the late 19th century. Some have suggested thatthe goal of systematics is to produce a 'stablestandard sequence' and that the Wetmore classifi­cation and sequence is now so well entrenchedthat it should continue to serve as the standard se­quence (Mayr 1989; Mayr & Bock 1994). The'stability' of the Wetmore sequence, especially inthe light of much systematic research, was viewedby Mayr & Bock (1994) as 'of major advantage toall avian biologists' because, in the Wetmore se­quence, biologists can easily locate a particulargroup or species. However, this 'stability' is notto be regarded as a proof of its correctness. Infact, the 'stability' of the Wetmore sequencethroughout this century is entirely due to failureto incorporate new ideas about relationships (Si-

bley 1989; 1994). Fortunately, in recent years tax­onomic systems have become more consistentwith current knowledge of phylogenetic relation­ships. Closely associated with this development isa reappraisal of systematics as a historical science(Gould 1986), in which phylogenies are viewed ashypotheses of relationships and taxonomies basedon them as dynamic systems, subject to furthermodification as our knowledge of relationshipsgrows.

Systematics is not an exercise in producingclassifications and sequences convenient for hu­mans, but an attempt to discover an underlyingreal structure in nature (Griffiths 1994). That realstructure, or natural system, is the pattern of his­torical (phylogenetic) relationships. This naturalsystem is something we discover (i.e. recon­struct), not something we create (Ghiselin 1987).

The objective of systematics, therefore, is the re­construction of phylogenetic relationships; tax­onomy is concerned with the representation ofthese relationships. With its focus on past events(i.e. historical subdivisions), systematics is a his­torical science.

It needs to be emphasised that reconstructedphylogenies (cladograms) are hypotheses. Thetrue phylogeny is buried in history and is un­known and probably unknowable. Therefore,proof, in a literal sense, of phylogenetic relation­ships may never be obtained. Because the truephylogeny is unknown, there is no a priori basisfor accepting or rejecting a given phylogeny; hy­potheses must be tested in the light of additionaldata. A phylogenetic hypothesis is open to test bythe addition of new characters, the addition ofnew taxa and the re-evaluation of other charac­ters. Thus, a phylogeny can be corroborated or re­jected and replaced by another phylogeny. Cor­roboration may come in the form of congruence.Phylogenies are congruent if they show the samebranching pattern. If congruence exists betweenphylogenies based on different sets of data, thismay indicate a strong historical signal; congru­ence may be regarded as evidence that the rele­vant phylogenies identify the true organismalphylogeny. Rejection of a phylogenetic hypoth-

144 ARDEA 87(1), 1999

esis requires that an alternative hypothesis bettersummarises all available evidence.

Because the purpose of taxonomy is commu­nication of information about phylogenetic rela­tionships, taxonomic systems should reflect thecurrent best-supported hypothesis of relation­ships. Taxonomic systems should be adjustedonly if it is believed that an alternative hypothesisis better supported by the available evidence. Be­cause phylogenies are hypotheses which are sub­ject to further testing, taxonomies based on themare provisional and may later be replaced or modi­fied. Therefore, taxonomies are dynamic systems;a definitive taxonomic system is probably unat­tainable in practice.

TAXONOMIC CHANGES

GalloanseraeA sister-group relationship of Anseriformes

and Galliformes is strongly supported by con­gruency of phylogenetic analyses of several inde­pendent data sets. These include morphologicalcharacters (Cracraft 1986; 1988; Cracraft & Min­dell 1989; Andors 1991; 1992; Kurochkin 1995;Livezey 1997a), DNA-DNA hybridisation (Sibleyet al. 1988; Sibley & Ahlquist 1990; Harshman1994; Bleiweiss et al. 1995), 12S and 16S ribo­somal RNA sequences (Hedges et al. 1995), (X­

crystallin sequences (Hedges et al. 1995; Casperset al. 1997) and mitochondrial DNA sequences(Mindell et al. 1997). The clade formed by Anseri­formes and Galliformes was named Galloanseraeby Sibley et al. (1988). Most of these analysesalso indicate that Galloanserae is the sister taxonof all extant birds except Paleognathae (Cracraft1986; 1988; Cracraft & Mindell 1989; Sibley &Ahlquist 1990; Hedges et ai. 1995; Kurochkin1995; Caspers et al. 1997; Livezey 1997a). In con­formity with the suggestion of De Queiroz &Gauthier (1992) to list, of each pair of sister taxa,the less speciose groups first, we propose to listAnseriformes before Galliformes and to placethese taxa before the remaining taxa on the Dutchlist.

Cygnus columbianus Whistling SwanFluitzwaanCygnus bewickii Bewick's Swan Kleine Zwaan

Whistling Swan and Bewick's Swan are spe­cifically distinct (Stepanyan 1990; Gantlett et al.1996; King 1997) based on qualitative differencesin morphology (Evans & Sladen 1980; Livezey1996).

Anserfabalis Taiga Bean Goose TaigarietgansAnser serrirostris Thndra Bean GooseToendrarietgans

Taiga Bean Goose and Tundra Bean Goose arespecifically distinct (Sangster & Oree11996; King1997; Persson 1997; Wells 1998) based on differ­ences in proportions, vocalisations, feeding habi­tat and diet, photosensitivity, activity pattern,behaviour, phenology and responses to periodsof extreme cold (Berry 1938; Coombes 1951;Mathiasson 1963; Huyskens 1977; 1986; Van Impe1980; Kurechi et al. 1983; Van den Bergh 1985;Barthel 1989; 1995; Burgers et al. 1991; Miyabay­ashi et al. 1994). Ringing recoveries suggest thatTaiga Bean Goose and Tundra Bean Goose haveallopatric breeding distributions (Burgers et al.1991) and there is no evidence for 'intergradation'(Sangster & Oreel 1996). In both Taiga BeanGoose and Tundra Bean Goose, geographic varia­tion in size (Johansen 1945; Delacour 1951;Cramp & Simmons 1977; Roselaar 1977) is con­sistent with a clinal variation pattern (Sangster &Oreel 1996); therefore, johanseni and midden­dorffi are included in A. fabaiis; rossicus is in­cluded in A. serrirostris (Sangster & OreeI1996).

Branta hutchinsii Lesser Canada GooseKleine Canadese GansBranta canadensis Greater Canada GooseGrote Canadese Gans

Lesser Canada Goose and Greater CanadaGoose are specifically distinct (Sibley 1996) basedon congruence of phylogeographic analyses ofmitochondrial DNA restriction fragments(Shields & Wilson 1987; Shields 1988; Van Wag­ner & Baker 1990; Quinn et al. 1991), mito­chondrial DNA sequences (Quinn et al. 1991; Ba-

Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 145

ker & Marshall 1997) and morphometry (VanWagner & Baker 1990). Pending further analysis,leucopareia, minima and taverneri are provision­ally retained conspecific with B. hutchinsii; fulva,interior, maxima, moffitti, occidentalis and par­vipes are provisionally retained conspecific withB. canadensis.

Branta bernicla Dark-bellied Brent GooseRotgansBranta hrota Pale-bellied Brent GooseWitbuikrotgansBranta nigricans Black Brant Zwarte Rotgans

Dark-bellied Brent Goose, Pale-bellied BrentGoose and Black Brant are specifically distinct(Millington 1997) based on qualitative differencesin morphology (Delacour 1954; Johnsgard 1978;Millington 1997), overlapping breeding ranges ofPale-bellied Brent Goose and Black Brant in arc­tic Canada (Gavin 1947; Handley 1950) and segre­gation of Pale-bellied and Dark-bellied BrentGoose in the Netherlands and Denmark (Lam­beck 1981). The alleged hybrid origin of 'interme­diate' populations in central arctic Canada, whichformed the basis for including Pale-bellied BrentGoose and Black Brant, along with Dark-belliedBrent Goose, in a single species (Delacour &Zimmer 1952), has been falsified by genetic anal­ysis (Shields 1990).

Mareca penelope Eurasian Wigeon SmientMareca americana American WigeonAmerikaanse SmientMareca falcata Falcated Duck BronskopeendMareca strepera Gadwall Krakeend

Phylogenetic analyses based on mitochondrialDNA and morphology (Kessler & Avise 1984;Livezey 1991) provide strong support for the exis­tence of two major clades within the dabblingducks traditionally included in Anas (Omland1994): (1) a clade formed by Cape Teal M. capen­sis, the wigeons, Falcated Duck and Gadwall; and(2) a clade formed by the remaining species. Weadopt the classification of Livezey (1991; 1997b)in which the members of the former clade areplaced in Mareca and the remaining species in

Anas (King 1997). Current knowledge of the phylo­genetic relationships of dabbling ducks is betterrepresented with the recognition of two generathan with the placement of all species in Anas.

Anas crecca Common Teal WintertalingAnas carolinensis Green-winged TealAmerikaanse Wintertaling

Common Teal and Green-winged Teal are spe­cifically distinct (Stepanyan 1990; Livezey 1991;Gantlett et al. 1996; Johnson & Sorenson 1998)based on qualitative differences in morphology(Delacour 1956; Johnsgard 1978; Livezey 1991).

Melanitta nigra Common ScoterZwarte Zee-eendMelanitta americana Black ScoterAmerikaanse Zee-eend

Common Scoter and Black Scoter are specifi­cally distinct (Stepanyan 1990; Livezey 1995b;Gantlett et al. 1996; King 1997) based on quali­tative differences in morphology (Johnsgard 1978;Livezey 1995b).

Melanittafusca Velvet Scoter Grote Zee-eendVelvet Scoter and White-winged Scoter M.

deglandi are specifically distinct (Stepanyan1990; Livezey 1995b; King 1997) based on quali­tative differences in morphology (Johnsgard 1978;Livezey 1995b).

Mergellus albellus Smew NonnetjeLophodytes cucullatus Hooded MerganserKokardezaagbek

A phylogenetic analysis based on morphology(Livezey 1995b) indicates that Smew is moreclosely related to the goldeneyes Bucephala thanto the mergansers Mergus. Because the inclusionof Smew in Mergus would render Mergus poly­phyletic, we adopt Livezey's (1995b) classifica­tion and place Smew in a monotypic genus Mer­gellus (AOU 1983; BOURC 1997; King 1997). Weplace Hooded Merganser in the monotypic genusLophodytes (AOU 1983; BOURC 1997; King1997) to indicate its basal position among the mer­gansers (Livezey 1995b). The latter species is fre-

146 ARDEA 87(1), 1999

quently recorded in the Netherlands but recordsare considered to refer to escapes from captivity.Hooded Merganser, therefore, is not formally ad­mitted to the Dutch list.

Soft-plumaged petrel complex donsstormvogelsFea's Petrel Pterodromafeae, Zino's Petrel P.

madeira and Soft-plumaged Petrel P. mollis arespecifically distinct (Bourne 1983; Collar & Stuart1985; Zino & Zino 1986; Sibley & Monroe 1990;Beaman 1994; Hazevoet 1995; 1997; Sibley 1996;Snow & Pemns 1998) based on phylogeographicanalysis of mitochondrial DNA sequences (Nunn& Zino in press) and concordance of differences inmorphology (Zino & Zino 1986), vocalisations(Bretagnolle 1995) and reproductive behaviour(Zino & Zino 1986). Analysis of mitochondrialDNA sequences suggests that the divergence of P.feae and P. madeira occurred 840 000 years agoand that P. mollis is not closely related to P. feaeand P. madeira (Nunn & Zino in press). Popula­tions of Fea's Petrel breeding on the DesertaIslands, Madeira Cdeserta'), are provisionally re­tained conspecific with P. feae (Snow & Pemns1998). Non-monophyly of the soft-plumaged pet­rel complex precludes the recognition of a 'super­species' taxon for P.feae, P. madeira andP. mollis.[There are no accepted records of P. feae, P. ma­deira or P. mollis in the Netherlands, although a re­cord at Camperduin, Noord-Holland (Stegeman etal. 1995), was accepted as P.feaelmadeiralmollis.]

Puffinus mauretanicus Balearic ShearwaterVale PijlstormvogelPuffinus puffinus Manx ShearwaterNoordse Pijlstormvogel

Balearic Shearwater is specifically distinctfrom Manx Shearwater and Yelkouan ShearwaterP. yelkouan (McMinn et al. 1990; Walker et al.1990; Altaba 1995; Sibley 1996; King 1997; Hei­drich et al. 1998; Snow & Perrins 1998; Wells1998) based on analyses of qualitative morpho­logical characters (Walker et al. 1990; Altaba1995) and phylogenetic analysis of mitochondrialDNA (Austin 1996; Heidrich et al. 1998). The re­cent discovery and subsequent description of two

extinct shearwaters, Hole's Shearwater P. holeae(Walker et al. 1990) and Olson's Shearwater P. 01­soni (McMinn et al. 1990), which occurred sym­patrically in the eastern Canary Islands, castsdoubt on the alleged sister relationship (and con­specificity) of Balearic Shearwater and YelkouanShearwater (Altaba 1995). It has been suggestedthat Balearic Shearwater may actually be moreclosely related to Hole's Shearwater than to Yel­kouan Shearwater (Walker et al. 1990).

Calonectris borealis Cory's ShearwaterKuhls Pijlstormvogel

Cory's Shearwater and Scopoli's ShearwaterC. diomedea are specifically distinct based onphylogeographic analysis of allozymes (Randi etal. 1989) and mitochondrial DNA (Heidrich et al.1996; 1998), qualitative differences in vocalisa­tions (Bretagnolle & Lequette 1990) and analysisof morphological characters (Granadeiro 1993;Gutierrez 1998). Cape Verde Shearwater C. ed­wardsii is specifically distinct from Cory's Shear­water and Scopoli's Shearwater (Bannerman &Bannerman 1968; Norrevang & den Hartog 1984;Hazevoet 1995; 1997; Sibley 1996; Hillcoat et al.1997; Porter et al. 1997; Snow & Pemns 1998)based on qualitative differences in morphologyand vocalisations (Alexander 1898; Murphy 1924;Bourne 1955; Bannerman & Bannerman 1968;Hazevoet 1995; 1997; Porter et al. 1997; Snow &Pemns 1998). [In the Netherlands, all specimen re­cords of Calonectris were identified as C. borealis(Van den Berg & Bosman 1999). The identity ofsight records of C. borealislC. diomedea in theNetherlands is currently being investigated by theDutch rarities committee (CDNA).]

Morus bassanus Northern Gannet Jan van GentPhylogenetic relationships among Sulidae

(Warheit 1992; Friesen & Anderson 1997) are bestrepresented with the recognition of Morus for thegannets and Sula for the boobies, with the excep­tion of Abbott's Booby Papasula abbotti (Olson1985; Olson & Warheit 1988; Van Tets et al. 1988;AOU 1989; Sibley & Monroe 1990; Warheit 1992;Friesen & Anderson 1997).

Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 147

Stictocarbo aristotelis European ShagKuifaalscholver

A phylogenetic analysis of morphologicalcharacters (Siegel-Causey 1988) identified ninemajor clades among the connorants and shags.The classification proposed by Siegel-Causey(1988), which recognises nine genera, better rep­resents the phylogenetic relationships of the cor­morants and shags than the current inclusion ofall species in a single genus Phalacrocorax(Bourne & Casement 1996).

Casmerodius albus Great White EgretGrote Zilverreiger

Phylogenetic analyses based on morphologyand DNA-DNA hybridisation (Payne & Risley1976; Sheldon 1987; Sheldon et al. 1995) indicatethat Great White Egret is not closely related to theEgretta clade and instead suggest a closer rela­tionship with Bubulcus and Ardea. However,given the unresolved relationships between GreatWhite Egret, Bubulcus and Ardea, inclusion ofGreat White Egret in Ardea (AOU 1995; BOURC1997) is premature. Until the relationships ofGreat White Egret are better understood it is bestplaced in a monotypic genus Casmerodius (AOU1983; Eck 1996; Inskipp et al. 1996; King 1997;Wells 1998).

Phoenicopterus roseus Greater FlamingoFlamingo

Greater Flamingo and Caribbean Flamingo Pruber are specifically distinct (Allen 1956; Mo­rony et al. 1975; Hazevoet 1995; Sibley 1996;King 1997; Sangster 1997a) based on qualitativedifferences in plumage and bill pattern (Van denBerg 1987; Sangster 1997a) and display behaviourand vocalisations (Studer-Thiersch 1964; 1974;1975).

Aquila nipalensis Steppe Eagle SteppearendSteppe Eagle and Tawny Eagle A. rapax are

specifically distinct (Brooke et al. 1972; Clark1992; Olson 1994; King 1997; Wells 1998) basedon qualitative morphological differences (Brookeet al. 1972; Clark 1992; Olson 1994).

Porphyrio madagascariensis African Swamp-henSmaragdpurperkoetPorphyrio poliocephalus Grey-headed Swamp-henGrijskoppurperkoet

Western Swamp-hen P porphyrio, AfricanSwamp-hen, Grey-headed Swamp-hen, Philip­pine Swamp-hen P pulverulentus, Black-backedSwamp-hen P indicus and Australian Swamp-henP melanotus are specifically distinct (Sangster1998) based on qualitative differences in morphol­ogy (Ripley 1977; Cramp & Simmons 1980; DelHoyo et al. 1996). Analyses of mitochondrialDNA suggest that forms previously included un­der the name 'Purple Swamp-hen P porphyrio'(Von Boetticher 1935; Ripley 1977; Del Hoyo etal. 1996) are paraphyletic with respect to twolarge flightless New Zealand taxa, South IslandTakahe P hochstetteri and extinct North IslandTakahe P mantelli (Trewick 1997). These resultsargue against continued inclusion of swamp-henforms in a single polytypic species. The sixgroups here treated as species (P porphyrio, Pmadagascariensis, P poliocephalus, P pulverul­entus, P indicus and P melanotus) are similar tothose recognised by Roselaar (in Cramp & Sim­mons 1980) as subspecies groups. Pending furtheranalysis, caspius and seistanicus are tentativelyincluded in P poliocephalus; viridis is tentativelyincluded in P indicus; and bellus, chathamensis,melanopterus, pelewensis and samoensis are ten­tatively included in P melanotus. [The inclusionof African Swamp-hen and Grey-headed Swamp­hen on the Dutch list is currently under review bythe Dutch rarities committee (CDNA).]

Chlamydotis macqueenii Macqueen's BustardOostelijke Kraagtrap

Macqueen's Bustard and Houbara Bustard C.undulata are specifically distinct (Gaucher et al.1996; Sangster 1996; King 1997; Wells 1998) basedon qualitative differences in courtship behaviourand genetic analysis (Granjon et al. 1994;

Gaucher et al. 1996).

148 ARDEA 87(1), 1999

Pluvialis dominicus American Golden PloverAmerikaanse GoudplevierPluvialis fulva Pacific Golden PloverAziatische Goudplevier

American Golden Plover and Pacific GoldenPlover are specifically distinct (Knox 1987; AOU1993; King 1997; Wells 1998) based on differencesin plumage, morphology, moult, vocalisations,ecology and overlap of breeding ranges (Connors1983; Connors et af. 1993). The correct name ofAmerican Golden Plover is P. dominicus, not P.dominica (AOU 1995).

Vanellus gregarius Sociable LapwingSteppekievitVanellus leucurus White-tailed LapwingWitstaartkievit

Phylogenetic analyses of behavioural charac­ters (Ward 1992) have been unable to resolve rela­tionships among lapwings. Given the doubtfulmonophyly of Chettusia (and Hoplopterus), therecognition of Chettusia (and Hoplopterus) is notjustified. Therefore, all lapwings are placed inVanellus (BOURC 1997; King 1997; Wells 1998).

Gallinago gallinago Common Snipe WatersnipCommon Snipe and Wilson's Snipe G. deli­

cata are specifically distinct (Olsson 1987; Gan­tlett et af. 1996; King 1997) based on qualitativedifferences in morphology, vocalisations anddrumming display (Thonen 1969; Cramp & Sim­mons 1983; Olsson 1987; Carey & Olsson 1995;Miller 1996a; 1996b; Gibson & Kessel 1997).Pending further analysis, faeroeensis and galli­nago are provisionally retained as conspecific(Miller 1996b). African Snipe G. nigripennis,Madagascar Snipe G. macrodactyla, ParaguayanSnipe G. paraguaiae, Magellan Snipe G. magel­lanica and Puna Snipe G. andina are specificallydistinct from Common Snipe based on qualitativedifferences in morphology, vocalisations anddrumming display (Tuck 1972; Sutton 1981; Hay­man et af. 1986; Fjeldsa & Krabbe 1990; Del Hoyoet af. 1996).

Phalaropus tricolor Wilson's PhalaropeGrote Franjepoot

Results of phylogenetic analyses based on al­lozymes (Dittmann et al. 1989), mitochondrialDNA (Dittmann & Zink 1991) and morphology(Chu 1995) are contradictory with regard to the al­leged polyphyletic origin of the phalaropes(Sibley & Monroe 1990). Because of this incon­gruence, the recognition of Steganopus for Wils­on's Phalarope (Sibley & Monroe 1990; Dowsett& Dowsett-Lemaire 1993; Beaman 1994; DelHoyo et al. 1996; Higgins & Davies 1996) is un­justified and, therefore, we retain Wilson's Phala­rope in Phalaropus.

Phalaropus fulicaria Grey PhalaropeRosse Franjepoot

The correct name of Grey Phalarope is P. fuli­caria, not P. fulicarius (Parkes 1982).

Stercorarius skua Great Skua Grote JagerRecent phylogenetic analyses of allozymes

and mitochondrial DNA sequences (Cohen et af.1997; Braun & Brumfield 1998) confirm and extendprevious suggestions based on short mitochondrialDNA sequences (Blechschmidt et al. 1993) andecology (Andersson 1973) that Pomarine Skua S.pomarinus is more closely related to species placedin Catharacta than to Arctic Skua S. parasiticusand Long-tailed Skua S. longicaudus. Because in­dependent lines of evidence suggest that Sterco­rarius, as currently defined (Furness 1987; Chris­tidis & Boles 1994; BOURC 1997; King 1997), is aparaphyletic taxon, all skuas are placed in Sterco­rarius.

Larus graellsii Lesser Black-backed GullKleine MantelmeeuwLarus fuscus Baltic GullBaltische Mantelmeeuw

Lesser Black-backed Gull and Baltic Gull arespecifically distinct based on qualitative differ­ences in morphology and differences in moult andecology (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 fromgraellsii; 'intermedius' is, therefore, consideredconspecific with L. graellsii.

Heuglin's Gull L. heuglini is specifically dis­tinct from Lesser Black-backed Gull, Baltic Gull,Armenian Gull L. armenicus, Pontic Gull L. ca­chinnans, Yellow-legged Gull L. michahellis andVega Gull L. vegae based on qualitative differ­ences in morphology and behaviour, and differ­ences in ecology (Grant 1986; Filchagov et aL1992; Hario 1992; Kennerley et aL 1995; Yesou &Hirschfeld 1997), The breeding range of Heug­lin's Gull overlaps with that of Herring Gull andBaltic Gull, with evidence for reproductive isola­tion (Filchagov & Semashko 1987; Filchagov etaL 1992; Filchagov 1994). Pending further analy­sis, taimyrensis and heuglini are provisionally re­tained as conspecific (Kennerley et al. 1995).

Larus argentatus Herring Gull ZilvermeeuwLarus michahellis Yellow-legged GullGeelpootmeeuwLarus cachinnans Pontic Gull Pontische Meeuw

Herring Gull, Vega Gull and American Her­ring Gull L. smithsonianus are specifically dis­tinct based on qualitative differences in morphol­ogy and vocalisations (Frings et aL 1958; Hoff­man 1979; Grant 1986; Mullarney 1990; Kennerleyet aL 1995; Dubois 1997; Chu 1998). There is noevidence that the form 'argenteus' is diagnosablydistinct from argentatus. Current evidence indi­cates a clinal pattern of variation (Barth 1968;Cramp & Simmons 1983); the design of studieswhich have suggested clear differences betweenpopulations of 'argenteus' and argentatus (Mon­aghan et aL 1983; Golley 1993) was inadequate tosubstantiate such claims (Chylarecki 1993). Theform 'argenteus' is, therefore, considered con­specific with L. argentatus.

Yellow-legged Gull and Herring Gull are spe­cifically distinct (Oree! 1980; Marion et aL 1985;Stepanyan 1990; Beaman 1994; King 1997) basedon qualitative differences in adult and immatureplumages, bare parts, behaviour and vocalisationsand overlap of breeding ranges (Nicolau-Guil­laumet 1977; Glutz von Blotzheim & Bauer 1982;

Teyssedre 1984; Marion et aL 1985; Yesou 1991).Pontic Gull and Yellow-legged Gull are specifi­cally distinct (Klein & Buchheim 1997; Klein &Gruber 1997) based on qualitative differences inmorphology and vocalisations, and differences inbehaviour and ecology (Klein 1994; Gruber 1995;Jonsson 1996; 1998b; Garner 1997; Gamer &Quinn 1997; Garner et aL 1997; Klein & Buch­heim 1997; Klein & Gruber 1997; Larsson & Lo­rentzon 1998). Pending further analysis, atlantis isprovisionally retained as conspecific with L.michahellis; barabensis and mongolicus are pro­visionally retained as conspecific with L. cachin­nans.

Armenian Gull L. armenicus is specificallydistinct from Pontic Gull, Yellow-legged Gull andHeuglin's Gull L. heuglini based on qualitativedifferences in morphology and vocalisations(Geroudet 1982; Hume 1983; Dubois 1985; Grant1986; 1987; Satat & Laird 1992; Buzun 1993; Fil­chagov 1993; Frede & Langbehn 1997; Yesou &Hirschfeld 1997).

Gelochelidon nilotica Gull-billed TernLachstern

Although one phylogenetic study based on al­lozymes suggests that Gull-billed Tern originateswithin the Sterna clade (Randi & Spina 1987),others, based on allozymes (Hackett 1989), hind­limb musculature (McKitrick 1991) and osteology(Chu 1995), suggest a more distant relationshipand do not support the inclusion of Gull-billedTern in Sterna. Therefore, the inclusion of Gelo­chelidon in Sterna (AOD 1983; Eck 1996;BODRC 1997) is not warranted by present knowl­edge of phylogenetic relationships. Given the un­certainty about phylogenetic relationships amongterns, as indicated by the incongruence of avail­able analyses, we retain Gelochelidon for Gull­billed Tern.

Apus melba Alpine Swift AlpengierzwaluwEvidence is lacking for a sister relationship

between Alpine Swift and Mottled Swift A. ae­quatorialis and monophyly of the other speciestraditionally placed in Apus. Therefore, the recog-

150 ARDEA 87(1), 1999

mtlOn of Tachymarptis for Alpine Swift andMottled Swift and a more restricted Apus for theremaining species (Brooke 1972; Fry et al. 1988;Short et al. 1990; Chantler & Driessens 1995; Eck1996) may render Tachymarptis and/or Apus para­phyletic. In the absence of a relevant phylogeneticanalysis of the swifts, we retain Alpine Swift inApus.

Merops persicus Blue-cheeked Bee-eaterGroene Bijeneter

Blue-cheeked Bee-eater and Madagascar Bee­eater M. superciliosus are specifically distinct(Glutz von Blotzheim & Bauer 1980; Eck 1996;King 1997; Wells 1998) based on qualitative dif­ferences in morphology (Fry 1984).

Anthus richardi Richard's Pipit Grote PieperRichard's Pipit is specifically distinct from

Grassland A. cinnamomeus, Paddyfield A. rufu­Ius, Australian A. australis and New Zealand PipitA. novaeseelandiae (Glutz von Blotzheim &Bauer 1985; Sibley 1996; Wells 1998) based onqualitative differences in plumage and vocalisa­tions (Glutz von Blotzheim & Bauer 1985 and ref­erences cited therein).

Anthus spinoletta Water Pipit WaterpieperAnthus petrosus Rock Pipit Oeverpieper

Water Pipit, Rock Pipit and Buff-bellied PipitA. rubescens are specifically distinct (Oreel 1980;AOU 1989; King 1997) based on qualitative dif­ferences in plumage, vocalisations and ecology(Bijlsma 1977; Alstrom & Olsson 1987; Knox1988a).

Motacillaflavissima Yellow WagtailEngelse KwikstaartMotacillaflava Blue-headed WagtailGele KwikstaartMotacilla thunbergi Grey-headed WagtailNoordse KwikstaartMotacillafeldegg Black-headed WagtailBalkankwikstaart

Yellow Wagtail, Blue-headed Wagtail, Grey­headed Wagtail, Black-headed Wagtail, Spanish

Wagtail M. iberiae, Ashy-headed Wagtail M. ci­nereocapilla, Yellow-headed Wagtail M. lutea,Green-headed Wagtail M. taivana, KamtchatkaWagtail M. simillima, Alaska Wagtail M. tschut­schensis and White-headed Wagtail M. leuco­cephala are specifically distinct based on qualita­tive differences in morphology (Sushkin 1925; Jo­hansen 1944; Voous 1950; Williamson 1955; Vau­rie 1957; Dittberner & Dittberner 1984; Glutz vonBlotzheim & Bauer 1985; Cramp 1988; Leader1996). For several named populations there is noevidence or insufficient evidence that these are di­agnosably distinct. Therefore, 'beema' is includedin M. jlava, 'angarensis', 'macronyx' and 'plexa'in M. thunbergi, 'aralensis', 'kaleniczenkoi' and'melanogrisea' in M. feldegg and 'pygmaea' in M.cinereocapilla. The form 'dombrowskii' probablyrefers to hybrids of M. jlava, M. thunbergi and M.feldegg (Vaurie 1957; Mayr & Greenway 1960).The form 'superciliaris' most likely refers to hy­brids of M. jlava and M. feldegg (Vaurie 1957;Mayr & Greenway 1960). The status of 'zaissa­nensis' remains unresolved.

Motacilla alba White Wagtail Witte KwikstaartMotacilla yarrellii Pied Wagtail Rouwkwikstaart

White Wagtail, Pied Wagtail, Moroccan Wag­tail M. subpersonata, Masked Wagtail M. person­ata, Himalayan Wagtail M. alboides, Black-back­ed Wagtail M. lugens, East Siberian Wagtail M.ocularis, Amur Wagtail M. leucopsis and BaikalWagtail M. baicalensis are specifically distinctbased on qualitative differences in morphology(Glutz von Blotzheim & Bauer 1985; Cramp1988). There is no evidence that populations inwestern Asia (,dukhunensis') are diagnosably dis­tinct from alba. Therefore, 'dukhunensis' is in­cluded in M. alba. The form 'persica' probablyrepresents a variable hybrid population of albaand personata (Vaurie 1959; Cramp 1988) and isnot recognised.

Saxicola rubicola European StonechatRoodborsttapuitSaxicola maura Siberian StonechatAziatische Roodborsttapuit

Sangster et al.: TAXONOMIC CHANGES IN 1977-1998 151

European Stonechat, Siberian Stonechat andAfrican Stonechat S. torquata are specifically dis­tinct (Sibley 1996; Wells 1998) based on qualita­tive differences in morphology (Cramp 1988;Svensson 1992) and phylogeographic analysis(Wittmann et al. 1995). There is no evidence thatpopulations inhabiting western Europe are dia­gnosably distinct from those in central and north­ern Europe. Therefore, the form 'hibernans' re­presents a synonym of S. rubicola. There is no ev­idence that populations inhabiting eastern Siberia('stejnegeri') are diagnosably distinct from west­ern Siberian populations. Therefore, 'stejnegeri'is included in S. maUTa (Svensson 1992). Pendingfurther analysis, variegata, armenica, indica andprzewalskii are provisionally retained as conspe­cific with S. maUTa.

Oenanthe hispanica Western Black-earedWheatear Westelijke Blonde TapuitOenanthe melanoleuca Eastern Black-earedWheatear Oostelijke Blonde Tapuit

Western Black-eared Wheatear and EasternBlack-eared Wheatear are specifically distinctbased on qualitative differences in morphology(Clement & Harris 1987; Cramp 1988).

Zoothera aurea White's Thrush GoudlijsterWhite's Thrush and Scaly Thrush Z. dauma

are specifically distinct (Eck 1996) based on qual­itative differences in morphology and vocalisa­tions (Seebohm & Sharpe 1902; Ali & Ripley1973; Cramp 1988; Glutz von Blotzheim & Bauer1988; Martens & Eck 1995). There is no evidencethat populations in south-eastern Siberia, Russiaand southern Kuril Islands, Japan ('toratugumi')are diagnosably distinct from aurea. Therefore,'toratugumi' is included in Z. aurea.Amami Thrush Z. major, Nilghiri Thrush Z. neil­gherriensis, Sri Lanka Thrush Z. imbricata, Hors­field's Thrush Z. horsfieldi, Fawn-breasted ThrushZ. machiki, New Britain Thrush Z. talaseae, SanCristobal Thrush Z. margaretae, GuadalcanalThrush Z. turipavae, Bassian Thrush Z. lunulataand Russet-tailed Thrush Z. heinei are specificallydistinct (Mayr 1955; Deignan et al. 1964; Ford

1983; Ishihara 1986; White & Bruce 1986; Christi­dis & Boles 1994; Gibbs 1996; Inskipp et al. 1996;Sibley 1996; King 1997; Wells 1998) based onqualitative differences in morphology and vocal­isations (Seebohm & Sharpe 1902; Jahn 1942;Mayr 1955; Ali & Ripley 1973; Ford 1983; Ishi­hara 1986).

Acrocephalus agricola Paddyfield WarblerVeldrietzanger

Paddyfield Warbler and Manchurian WarblerA. tangorum are specifically distinct (Round 1994;King 1997) based on qualitative differences inplumage (Alstrbm et al. 1991; Round 1994). Phylo­genetic analysis of mitochondrial DNA sequencesindicates that Paddyfield Warbler and Manchu­rian Warbler are not sister-taxa (Leisler et al.1997). Paddyfield Warbler is considered mono­typic (Williamson 1968).

Acrocephalus scirpaceus European Reed War·bier Kleine Karekiet

European Reed Warbler, Mangrove ReedWarbler A. avicenniae, African Reed Warbler A.baeticatus and Caspian Reed Warbler A. fuscusare specifically distinct (Leisler et al. 1997; Sang­ster 1997b) based on qualitative differences inmorphology (Pearson 1981; Ash et al. 1989; Harriset al. 1995). Phylogenetic analysis of mitochon­drial DNA sequences indicates that MangroveReed Warbler, which is currently regarded as asubspecies of A. baeticatus, is actually moreclosely related to European Reed Warbler, andthat European and Caspian Reed Warbler, untilrecently regarded as subspecies of A. scirpaceus,are not sister-taxa (Leisler et al. 1997).

Acrocephalus caligatus Booted WarblerKleine Spotvogel

Phylogenetic analyses of mitochondrial DNAsequences indicate that Booted Warbler and Oliv­aceous Warbler A. pallidus are more closely re­lated to species traditionally included in Acro­cephalus clade rather than to Icterine WarblerHippolais icterina (Leisler et al. 1997; Sangster1997b). Therefore, Booted Warbler and Oliv-

152 ARDEA 87(1), 1999

aceous Warbler are placed in Acrocephalus.Booted Warbler and Sykes's Warbler A. rama

are specifically distinct (Stepanyan 1990; Glutzvon Blotzheim & Bauer 1991; Sibley & Monroe1993; Sibley 1996; Wells 1998) based on qualita­tive differences in morphology and vocalisations,and differences in ecology (Portenko et al. 1976;Glutz von Blotzheim & Bauer 1991; Cramp 1992;Hirschfeld 1994). Pending information on phylo­genetic relationships of Sykes's Warbler, itsplacement in Acrocephalus is tentative.

Phylloscopus proregulus Pallas's Leaf WarblerPallas' Boszanger

Pallas's Leaf Warbler and Lemon-rumpedWarbler P. chloronotus are specifically distinct(Inskipp et al. 1996; King 1997; Wells 1998) basedon qualitative differences in plumage and vocal­isations (Martens 1985; Alstrom & Olsson 1990).

Phylloscopus inornatus Yellow-browed WarblerBladkoningPhylloscopus humei Hume's WarblerHumes Bladkoning

Yellow-browed Warbler and Hume's Warblerare specifically distinct (Svensson 1992; Gantlettet al. 1996; BOURC 1997; King 1997; Wells 1998)based on qualitative differences in vocalisationsand plumage and overlap of breeding ranges(Mild 1987; Aistrom & Olsson 1988; Dathe &Loskot 1989).

Phylloscopus orientalis Eastern Bonelli's WarblerBalkanbergfluiterPhylloscopus bonelli Western Bonelli's WarblerBergfluiter

·Eastern Bonelli's Warbler and Western Bonel­li's Warbler are specifically distinct (Gantlett etal. 1996; BOURC 1997; King 1997; Wells 1998)based on qualitative differences in vocalisationsand genetic analyses (Helb et al. 1982; Helbig etal. 1995).

Phylloscopus collybita Common ChiffchaffTjiftjaf

Phylloscopus brehmii Iberian ChiffchaffIberische Tjiftjaf

Common Chiffchaff, Iberian Chiffchaff andCanarian Chiffchaff P. canariensis are specifi­cally distinct (Gantlett et al. 1996; Eck 1996;Sibley 1996; King 1997; Wells 1998) based on dif­ferences in structure and plumage (Erard & Salo­mon 1989; Salomon et al. 1997), qualitative differ­ences in vocalisations (Salomon 1989; Salomon &Hemim 1992) and genetic analyses (Helbig et al.1993; 1996). Pending further analysis, exsul is ten­tatively included in P. canariensis.

Common Chiffchaff, Mountain Chiffchaff P.sindianus and Caucasian Mountain Chiffchaff P.lorenzii are specifically distinct (Martens 1982;Snow & Pemns 1998) based on differences instructure and plumage (Shirihai 1987; Cramp1992), qualitative differences in vocalisations(Martens & Hanel 1981; Martens 1982) and ge­netic analyses (Helbig et al. 1996). Pending fur­ther analysis, abietinus and tristis are tentativelyincluded in P. collybita.

Lanius phoenicuroides Thrkestan ShrikeTurkestaanse KlauwierLanius speculigerus Daurian ShrikeDaurische Klauwier

Turkestan Shrike and Chinese Shrike L. isa­bellinus are specifically distinct (Kryukov 1995;Panov 1995; Panow 1996) based on qualitativedifferences in morphology (Dean 1982; Cramp &Pemns 1993; Panow 1996; Lefranc & Worfolk1997) and analyses of their contact zone (Kryukov1995).

Daurian Shrike is specifically distinct fromTurkestan Shrike and Chinese Shrike based onqualitative differences in morphology (Dean1982; Cramp & Pemns 1993; Panow 1996) andvocalisations (Panov 1995). Pending further anal­ysis, tsaidamensis is provisionally retained asconspecific with L. isabellinus. [An accepted re­cord of an 'isabelline shrike' on Texel, Noord­Holland, in May 1995 (Wassink 1996), is now con­sidered to refer to Daurian Shrike (Van den Berg& 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 commit­tee (CDNA).]

Lanius excubitor Great Grey Shrike KlapeksterLanius pallidirostris Steppe Grey ShrikeSteppeklapekster

Great Grey Shrike and Southern Grey ShrikeL. meridionalis are specifically distinct (Eck 1994;1996; Gantlett et al. 1996; Inskipp et al. 1996;Sibley 1996; BOURC 1997; King 1997; Lefranc &Worfolk 1997; Wells 1998) based on qualitativedifferences in plumage, breeding ecology and be­haviour (Isenmann & Bouchet 1993; Eck 1994;Lefranc 1995a; 1995b; Panow 1996; Lefranc &Worfolk 1997).

Steppe Grey Shrike is specifically distinctfrom Great Grey Shrike and Southern Grey Shrike(King 1997) based on qualitative differences inplumage, breeding ecology, behaviour and over­lap of breeding ranges (Panov 1995; Panow 1996;Lefranc & Worfolk 1997).

Corvus corone Carrion Crow Zwarte KraaiCorvus cornix Hooded Crow Bonte Kraai

Carrion Crow and Hooded Crow are specifi­cally distinct (Stepanyan 1990; Gantlett et al. 1996;King 1997) based on qualitative differences inplumage and analyses of their hybrid zone in Ger­many (Risch & Andersen 1998) and Italy (Saino1990; Saino & Scatizzi 1991; Saino 1992; Saino &Bolzern 1992; Saino & Villa 1992; Rolando 1993;Rolando & Laiolo 1994; Rolando & Saino 1994).

Chloris chloris Common Greenfinch GroenlingPhylogenetic analyses of morphological char­

acters (Raikow 1978; 1985) and short mitochon­drial DNA sequences (Fehrer 1996) provide con­gruent evidence that Chloris is more closely re­lated to Pyrrhula than to Carduelis. Because twoindependent studies suggest that Chloris is notpart of the Carduelis clade and identify the samesister-taxon (Raikow 1978; Fehrer 1996), whereasinclusion of Chloris in Carduelis is supported byonly one study (Van den Elzen & Nemeschkal1991), recognition of Chloris for the greenfinchesis justified.

Carduelis cannabina Linnet KneuCarduelis flavirostris Twite FraterCarduelis cabaret Lesser Redpoll Kleine BarmsijsCarduelisflammea Mealy Redpoll Grote BarmsijsCarduelis hornemanni Arctic RedpollWitstuitbarmsij s

Published studies of phylogenetic relation­ships among cardueline finches (Marten & John­son 1986; Van den Elzen & Nemeschkal 1991;Fehrer 1996) are contradictory with regard to thephylogenetic relationships of Acanthis and Car­duelis. Because monophyly of Acanthis has notbeen established and is contradicted by one study(Van den Elzen & Nemeschkal 1991), recognitionof Acanthis may not significantly contribute to theelimination of paraphyletic taxa. Although Car­duelis as defined here (i.e. including Acanthis butexcluding Chloris) is still likely to be para­phyletic, other hypotheses of relationships, whichrequire changes in nomenclature, do not seem tobe better supported by available data. Therefore,pending further phylogenetic analyses, Linnet,Twite, Lesser Redpoll, Mealy Redpoll and ArcticRedpoll are provisionally retained in Carduelis.

Lesser Redpoll and Mealy Redpoll are specif­ically distinct based on qualitative differences inmorphology (Knox 1988b; Herremans 1990) andvocalisations (Herremans 1989), and overlap ofbreeding ranges in south-eastern Norway withouthybridisation (Lifjeld & Bjerke 1996). Pendingfurther analysis, rostrata and exilipes are provi­sionally retained as conspecific with C. fiammeaand C. hornemanni, respectively.

Dendroica coronata Myrtle WarblerMirtezanger

Myrtle Warbler and Audubon's Warbler D.auduboni are specifically distinct (Bermingham etal. 1992) based on qualitative differences in plum­age (Hubbard 1970; Kaufman 1979; Cramp & Per­rins 1994; Dunn & Garrett 1997) and analysis oftheir hybrid zone (Barrowclough 1980). It hasbeen calculated (Zink & McKitrick 1995) that itwould take more than 6 million years for thesetaxa to completely fuse. Data in Bermingham etal. (1992) show that many North American Den-

154 ARDEA 87(1), 1999

droica species are less than 3.5 million years oldwhich means that speciation in several species ofDendroica took place during a much shorter timethan the period calculated as necessary for Myrtleand Audubon's Warblers to fuse.

Icterus galbula Baltimore OrioleBaltimoretroepiaal

Baltimore Oriole, Bullock's Oriole I. bullockiiand Black-backed Oriole I. abeillei are specifi­cally distinct (AOU 1995; Freeman & Zink 1995;BOURC 1997) based on qualitative differences inplumage and vocalisations, analyses of the hybridzone of Baltimore Oriole and Bullock's Oriole(see AOU 1995 for references) and a phylogeneticanalysis (Freeman & Zink 1995) indicating thatBaltimore Oriole and Bullock's Oriole are noteachother's closest relatives.

ACKNOWLEDGEMENTS

We thank Walter J. Bock and Ernst Mayr for providingvaluable and stimulating comments and criticisms. Weare grateful to Joel Cracraft for his support and encour­agement. As editor, Kees (C.J.) Camphuysen gener­ously shared his time and offered helpful suggestions.The work of the CSNA is supported by the NetherlandsOrnithologists' Union (NOU) and the Dutch BirdingAssociation (DBA).

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Alexander B. 1898. An ornithological expedition to theCape Verde Islands. Ibis (7) 4: 74-118.

Ali S. & S.D. Ripley 1973. Handbook of the birds ofIndia and Pakistan with those of Bangladesh, Ne­pal, Sikkim, Bhutan and Sri Lanka, 9. Oxford Uni­versity Press, Bombay.

Allen R.P. 1956. The flamingos: their life history andsurvival. Res. Rep. Nat. Audubon Soc. 5.

Alstrom P. & K. Mild 1987. Some notes on the tax­onomy of the Water Pipit complex. Proc. 4th Int.Identif. Meet. Eilat Nov. 1986: 47-48.

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Zink R.M. 1997. Species concepts. Bull. Br. OrnithoI.Club 117: 97-109.

Zink R.M. & M.C. McKitrick 1995. The debate overspecies concepts and its implications for ornithol­ogy. Auk 112: 701-719.

Zino P.A. & F. Zino 1986. Contribution to the study ofthe petrels of the genus Pterodroma in the archi­pelago of Madeira. BoI. Mus. Mun. Funchal 38:141-165.

SAMENVATTING

De Commissie Systematiek Nederlandse Avifauna(CSNA), een adviserende commissie van de Neder­landse Ornithologische Unie (NOU) en de Dutch Bir­ding Association (DBA), heeft zich gebogen over inter­nationale systematisch-biologische studies die betrek­king hebben op vogelsoorten die ook op de Neder­landse lijst voorkomen. Op grond van deze gepubli­ceerde studies en de evaluatie ervan heeft de CSNA eenaantal besluiten genomen ten aanzien van door haarnoodzakelijk geachte taxonornische en nomenclatori­sche wijzigingen. Dit artikel bevat een samenvattingvan de genomen besluiten en de eraan ten grondslagliggende argumenten.

Een belangrijk deel van de taxonornische verande­ringen betreft de soortstatus van bepaalde taxa, dievroeger veelal de status van ondersoort hadden. Dit iseen resultaat van een verandering in de wetenschappe­lijke inzichten in het soortsbegrip. Traditioneel wordter in de ornithologie gebruikgemaakt van het isolatie­soortsbegrip, ook weI genoemd het biologische soorts­begrip. Dit soortsbegrip stelt dat het belangrijkste crite­rium bij het onderscheiden van soorten de kruisbaar­heid betreft: als individuen uit twee verschillende po­pulaties met elkaar kunnen kruisen, dan behoren zij totdezelfde soort. Het is echter allang bekend, dat er pro­blemen aan dit, op het eerste gezicht zo logischlij­kende, isolatie-soortsbegrip kleven. Zo blijkt het in denatuur nogal eens voor te komen dat individuen uittwee groepen weI met elkaar kunnen kruisen, terwijl zijniet elkaars nauwste verwanten zijn (in de zin vanbloedverwantschap, genealogie, afstamming). Daar­naast gaat het isolatie-soortsbegrip er bij het afbakenenvan soorten vanuit dat deze soorten in de toekomst re­productief van elkaar gei"soleerd zullen blijven en niettot een populatie zullen fuseren. Dit betekent, dat infeite toekomstige gebeurtenissen uitsluitsel moeten ge­ven of we huidige groepen als aparte soorten of slechtsals populaties van een soort moeten beschouwen. Ten­slotte heeft praktische toepassing van het isolatie­soortsbegrip geleid tot het onderscheiden van samenge­stelde, polytypische soorten. Hoewel bepaalde popula­ties weI degelijk onderscheidbaar zijn, worden zij tochtot slechts een soort gerekend (omdat de onderlingeverschillen als te gering worden beoordeeld); deze han­delswijze resulteert in een onderwaardering van de fei­telijke biodiversiteit.

Bovengenoemde en andere problemen met het iso­latie-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 bake­nen. En zo zijn er in de loop van de tijd vele altema­tieve soortsbegrippen geformuleerd. De CSNA acht hetzogenaamde fylogenetische soortsbegrip van groot be­lang voor avifaunistische lijsten, de systematische bio­logie en biodiversiteitsstudies in het algemeen. Ret fy­logenetische soortsbegrip is een uitvloeisel van een we­tenschappelijke revolutie in een iets eerdere periodevan de systematische biologie. Als gevolg hiervan heeftnu alom het idee postgevat dat kennis van en inzicht inde afstammings- of fylogenetische relaties tussen groe­pen van doorslaggevend belang is in het vakgebied vande systematische biologie en de evolutiebiologie. Ditheeft als consequentie dat van een modem soortsbegripverwacht mag worden dat het in overeenstemming ismet de fylogenetische relaties tussen de onderscheideneenheden. Soortsbegrippen, zoals het isolatie-soortsbe­grip, die groepen van individuen bij elkaar zetten dieniet elkaars nauwste verwanten zijn, geven dientenge­volge een vertekend beeld van de evolutionaire ge­schiedenis. Fylogenetische soortsbegrippen voldoen indit opzicht beter.

Er zijn twee typen van fylogenetische soortsbegrip­pen, te weten de monofylie-versie en de diagnostischeversie. De monofylie-versie van het fylogenetischesoortsbegrip eist dat elke soort gekenmerkt wordt doorminimaal een uniek kenmerk dat niet bij andere soortenvoorkomt. De diagnostische versie is wat minder strikten eist slechts dat een soort gekarakteriseerd wordtdoor een unieke combinatie van kenmerken; d.w.z. deafzonderlijke kenmerken mogen weI bij andere soortenvoorkomen, maar hun combinatie moet uniek zijn. DeCSNA heeft de diagnostische versie van het fylogeneti­sche soortsbegrip gehanteerd bij het samenstellen vandeze rapportage.

Een tweede aspect van de avifaunistiek en de taxo­nomie dat door de CSNA aan een nadere beschouwingis onderworpen, betreft het groeperen van soorten inhogere eenheden, zoals geslachten en families, en deindeling van de lijst (dat wil zeggen welke groepenstaan bovenaan en welke onderaan de lijst). Ook dezezaken dienen, naar huidige inzichten, een afgeleide tezijn van de historische, fylogenetische relaties tussende onderscheiden groepen. Twee voorbeelden uit dehuidige lijst kunnen dit duidelijk maken. Fylogeneti­sche studies hebben aangetoond dat de Grote Zilverrei­ger niet nauw verwant is aan de andere zilverreigers.Dat betekent dat we de Grote Zilverreiger niet in het­zelfde geslacht (Egretta) kunnen handhaven als de an­dere zilverreigers en zij in een ander geslacht (Casme-

rodius) wordt geplaatst. Veel studies hebben aange­toond dat de eendachtigen (Anseriformes) en hoender­achtigen (Galliformes) elkaars nauwste verwanten zijn.Daarom kunnen we deze twee in een groep van hogereorde plaatsen, de Galloanserae.

De eerder genoemde revolutionaire ontwikkelingbinnen de systematische biologie en de evolutiebiolo­gie, en de moderne opvatting dat de nieuwe inzichtenen resultaten hun reflectie dienen te hebben op de be­staande taxonomische systemen, hebben directe prakti­sche consequenties. Ret betekent dat de oude taxono­mische systemen en avifaunistische lijsten gaan veran­deren. Er is weI beargumenteerd dat een groot voordeelvan de oude systemen en lijsten is dat ze stabiel zijn:hun onveranderlijkheid geeft houvast bij het opzoekenvan informatie. Ret is echter een feit dat deze stabili­teit, zo niet starheid, tot gevolg heeft dat de oude avi­faunistische en taxonomische lijsten geen afspiegelingmeer vormen van verkregen wetenschappelijke inzich­ten en hun relatie met de huidige stand van de systema­tische biologie en de evolutiebiologie deels hebben ver­loren. Als dergelijke lijsten geen afspiegeling meer vor­men van modeme inzichten in de fylogenie en taxono­mie van vogels, dan rijst onmiddellijk de vraag wat dielijsten en systemen dan weI vertegenwoordigen. Retklassieke antwoord dat zij ingeburgerde en gemakke­lijke zoeksystemen vertegenwoordigen, is wetenschap­pelijk gezien niet acceptabel. Als wetenschappelijkeonderbouwing niet meer van belang wordt geacht, danzijn er praktischere zoeksystemen denkbaar, bijvoor­beeld alfabetische lijsten. Daarom is de CSNA van me­ning dat taxonomische systemen geregeld dienen teworden aangepast om zo een afspiegeling te zijn van demeest modeme hypothesen over de fylogenetische rela­ties tussen groepen van vogels. Ret is evident, dat bijeen voortschrijdende wetenschap ook taxonomischesystemen en lijsten niet stil kunnen blijven staan en al­tijd aan verandering onderhevig zullen zijn.

Received 11 September 1998, accepted 29 March 1999Corresponding editor: Kees (C.J.) Camphuysen

Note from the editors of ArdeaThe taxonomic and nomenclatural changes listed

above, as proposed by the Dutch committee for aviansystematics (CSNA), will be adopted by Ardea in thisand future issues. With this decision, we follow recom­mendations of the Netherlands Ornithologists' Union(NOU) and the Dutch Birding Association (DBA). In

166 ARDEA 87(1), 1999

the review process of this manuscript, it became clearthat there is still considerable opposition to the princi­ples underlying the present changes. Some refereeswere not convinced of the advantages of the Phylo­genetic Species Concept (PSC). Not surprisingly, thesereferees could not recommend the publication of thispaper in Ardea. Other referees were completely infavour of the principles of a PSC and strongly recom­mended the present publication. Apparently, there waslittle or no ground for compromise. In the sometimesheated discussions generated by the proposals of theCSNA (e.g. at the NOV symposium in Naturalis, Lei­den, 10 April 1999), it became clear that scientific argu­ments were mixed with practical reasons, such as a de­sire to have a 'stable' list rather than (frequent) changes.Our decision to publish the list in Ardea was inspiredby a similar mix of reasons. We believe that the CSNAhas convincingly defended their proposal to abandonthe traditional Isolation Species Concept (ISC) in fa­vour of a Phylogenetic Species Concept, both in the in­troductory statements in the present publication and atthe NOV symposium in Leiden in 1999. Yet, it is clearthat we deal with hypotheses, and CSNA has confirmedthat changes in specific status, sequences or positionson the list are and will be proposed only if the evidenceis 'overwhelming'. Ever since the earlier publicationsof CSNA in Dutch Birding, several editors of ornitho-

logical journals and books in The Netherlands haveadopted the changes while others decided to continueto use more traditional lists. The present situation, asplit into two ('followers' and 'non-followers'), is veryinconvenient. Both the NOV and the editorial team ofArdea are convinced that a decision has to be made, sothat the CSNA could either continue to function as anadvisory committee of NOVIDBA, or should operateindependently.

After the publication of this list, the editorial teamof Ardea will seek advice from CSNA regarding thename and taxonomic status of species of birds men­tioned in the articles published in this journal, includ­ing those not featuring on the Dutch list. For conve­nience, and to avoid misunderstanding, we will informthe readers about the name/status of these species asthey were known prior to the present changes. Becausethe CSNA is the only body in The Netherlands that isspecialised in the evaluation of the taxonomic statusand systematics of birds, we recommend that proposalspublished in Ardea by this committee are followed. Atthe same time, we challenge scientists to constructivelycomment on CSNA decisions and proposals, so that, inthe process, we deepen our understanding of evolution­ary relationships and patterns of ancestry and descentof contemporary birds.