Sangster et al 1999

Transcription

Sangster et al 1999

139
DUTCH AVIFAUNAL LIST: SPECIES CONCEPTS, TAXONOMIC
INSTABILITY, AND TAXONOMIC CHANGES IN 1977-1998
GEORGE SANGSTER 1, CORNELIS
J.
HAZEVOET2,3, ARNOUD
C.S. (KEES) ROSELAAR2
&
B. VAN DEN BERG4,
RONALD SLUYS2
Sangster G., c.J. Hazevoet, A.B.Van den Berg, C.S. Roselaar & R. Sluys 1999. Ardea 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 contrast to the 'Biological Species Concept', this approach is compatible with the reconstruction 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 monophyletic 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: 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 Laborat6rio
Zool6gico e Antropol6gico (Museu Bocage), Rua da Escola Politecnica 58, 1250
Lisboa, Portugal; 4Duinlustparkweg 98, 2082 EG Santpoort-Zuid, Netherlands
e9
INTRODUCTION
This report includes taxonomic and nomenclatural changes adopted by the Dutch committee for
avian 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 the
Dutch Birding Association (DBA) support the
CSNA and its reports are published in both Ardea
and Dutch Birding. The present report is essentially similar to two separate publications in
Dutch Birding (Sangster et al. 1997; 1998). The
committee currently consists of five members
(year of election between parentheses): Arnoud
B. van den Berg (1995), Cornelis J. Hazevoet
(1996), C.S. (Kees) Roselaar (1995), George
Sangster, Secretary (1996) and Ronald Sluys
(1998). Due to limitations of space, the present
report only includes a brief summary of the rationale for each decision. More extensive and explicit motivations for the recognition of some of
the species-level and higher taxa listed here will
be published elsewhere.
Choice of species concept and its application
Over the past few years the fields of systematic and evolutionary biology have been invigorated by a scientific discussion on the concept of
species (for reviews, see Sluys 1991; Mayden
1997; Zink 1997). Although this discussion continues, it is also true that new and useful insights
have been gained already that can be incorporated
fruitfully into an up to date and modem description of avian diversity. One of the insights that has
surfaced in the literature is that different species
concepts may be best for different purposes
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ARDEA 87(1),1999
(Endler 1989) and it seems increasingly likely that
no single species concept will satisfy the multiple
purposes of 'species' in the biological sciences
(Sluys 1991; Hull 1997), However, a theme common to all biological sciences is that all taxa are
historically connected through their pattern of ancestry and descent. All living taxa are the product
of history, and we can understand little about their
diversity 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 and
among 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-level
taxa is that they are compatible with the reconstruction of evolutionary relationships. Species
concepts which group taxa that are not closely related in a single species misrepresent evolutionary history. Second, species-level taxa should be
delimited on the basis of historical subdivisions
(i.e. historical patterns), rather than present-day
or possible future interactions and processes
(Liden & Oxelman 1989), such as hybridisation
and gene flow. Species concepts which are prospective and which require speculations about the
future are not helpful in biology; since all of our
data are of the present and past, the units by
which we interpret these data must also be strictly
historical (Maddison 1997). Third, species should
be 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 distinct
taxa in a single (polytypic) 'species' because such
'species' actually are higher, more inclusive taxa.
Species concepts which recognise not only single
units (monotypic species) but also polytypic assemblages (polytypic species) as 'species' run
counter to the fundamental need for species-level
taxa to be basal and comparable.
The decision by the CSNA to abandon the traditional Isolation Species Concept (ISC) in favour
of a Phylogenetic Species Concept (PSC) was lar-
gely based on these views. The Isolation Species
Concept (popularly known as the 'Biological
Species Concept') is rejected because its properties violate all three aforementioned principles.
First, interbreeding taxa are not necessarily more
closely related to each other than they are to taxa
from 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 are
not closely related in a single species and, hence,
the misrepresentation of evolutionary history, is
inherent to the ISC and does not simply result
from errors in application. Phylogenetic analyses
indicate that in various groups of birds 'polytypic' species recognised by the ISC do not represent natural (monophyletic) groups. Evidence
comes from phylogenies based on both morphological (Livezey 1995a; Chu 1998; Veron 1999) and
molecular 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 the
sense 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 fuse
into each other. This poses both theoretical and
practical problems. It makes little sense to try to
interpret 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 in
WOOs or even millions of years (Zink & McKitrick 1995). Third, many 'species' recognised by
the ISC contain more than one taxon. The ISC recognises monotypic species but may also unite up
to 10 or more diagnosable taxa and still recognise
the resulting unit as a single 'species'. This not
only underestimates and misrepresents biodiversity but also compromises interspecific comparisons (Prum 1994; Hazevoet 1996; Cracraft 1997).
Sangster et at.: TAXONOMIC CHANGES IN 1977-1998
Two distinct Phylogenetic Species Concepts
have been advocated. These versions agree in
viewing species as products of evolution, not as
players in evolution, and support the notion of
species as basal taxa. The original version, proposed by Cracraft (1983) and further developed by
Nixon & Wheeler (1990) and Davis & Nixon
(1992), considers a phylogenetic species to be an
irreducible cluster of organisms possessing at
least one diagnostic character state. This version
thus focuses on the diagnosability of species. Diagnostic character states are discrete character
states which are fixed within the species and are
absent from close relatives. Diagnosability of
species may be based on any intrinsic attribute,
either morphological, molecular, ethological or a
combination of these. It should be emphasised
that this concept is populational (Cracraft 1997);
the criterion of diagnosability applies to (groups
of) populations, and not to, e.g. family groups or
individuals. An alternative approach was developed by Donoghue (1985) and De Queiroz &
Donoghue (1988; 1990) and considers phylogenetic species to be the smallest monophyletic
groups of organisms supported by autapomorphies
(unique derived character states).
The diagnosability and monophyly versions
of the PSC are significantly different; both in theory and application (Baum 1992; Davis 1997). A
major difference is that the monophyly version
requires phylogenetic analysis prior to delimiting
species, whereas the diagnosability version does
not. Implementation of the monophyly version
will, therefore, result in a significant shift in taxonomic practice, whereas the diagnosability version will not (Baum 1992). In addition to this
practical difference, the two versions also differ
with regard to which basal taxa are called 'species'. When a small group of related individuals
leaves a species, evolves diagnostic character
states and thus forms a descendant species, the
ancestral 'species' will cease to be monophyletic.
Because both the ancestral and descendant 'species' are characterised by diagnostic character
states, the diagnosability version would recognise
both as phylogenetic species; the monophyly ver-
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sion, however, would recognise the descendant
species, but not the ancestral 'species', as a phylogenetic species because only the former is
monophyletic. Proponents of the monophyly version propose that such ancestral 'species' are recognised as a separate class of species, for which
they propose the term 'metaspecies' (Donoghue
1985; De Queiroz & Donoghue 1988).
The CSNA has adopted the diagnosability
version of the PSC as its operational species concept because no phylogenetic analysis is required
prior to delimiting species, its implementation involves little modification of existing taxonomic
practices and does not require the recognition of
additional classes of species. It is believed that
these are advantages over the monophyly version,
and similar versions (Baum & Shaw 1995), and
that these outweigh any disadvantages. Although
both aspects (diagnosability and monophyly) may
be meaningful at the level of species (Baum
1992), the monophyly version is not sufficiently
practical 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 from
a narrow view of species, based on the ISC,
which restricts the term 'species' to entities which
are involved in the process of evolutionary
change. According to this view, species are real
only 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 organisms
living at different times, it does not follow that
other species concepts should also be applicable
in one time-slice only. In any case, the PSC does
not define species in terms of reproductive isolation (which, indeed, can only be assessed at a
given moment in time) but in terms of diagnostic
character states. The presence of diagnostic character states in a population may extend over time
and, therefore, phylogenetic species-level taxa
will have reality over time. However, this does
not mean that the PSC defines the evolutionary
lineage, as claimed by Szalay & Bock (1991) and
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ARDEA 87(1), 1999
Bock (1994). Since diagnostic character states
may become fixed in a population after a lineage
has split, speciation may be completed after the
origin of a lineage and, therefore, phylogenetic
species are not synonymous with evolutionary
lineages.
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 and
interpreted with reference to the diagnosability
version of the PSc. This does not imply that the
authors of the original publications necessarily
proposed the relevant taxa to be considered as
species-level taxa or explicitly endorse a PSC. In
fact, many of the authors concerned - implicitly
or explicitly - worked under the umbrella of the
ISC. The present interpretation of the data is fully
the responsibility of the CSNA. In the absence of
a fully documented system of phylogenetically
defined species level taxa, the CSNA continues to
use subspecific names to denote the likely geographic origin of the populations occurring in the
Netherlands.
Phylogeny and higher taxa
Although 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, development
and evolution (e.g. Dobzhansky 1937; Mayr
1942). With the 'Evolutionary Synthesis' of the
1930s and 1940s a dichotomy developed between
the study of species and the study of phylogeny
and higher taxa. Studies at the level of populations and species proceeded with great vigour, but
the study of phylogeny suffered from a lack of
conceptual and methodological clarity. For instance, it was not clear whether the term 'phylogeny' should only denote the branching pattern of
evolutionary 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 consistent
method 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 and
logic of phylogenetic systematics was greeted
with enthusiasm by systematists (e.g. Nelson
1970; Brundin 1972; Cracraft 1972) and is now almost universally accepted (e.g. Wiley 1981; Hillis
et al. 1996; Dingus & Rowe 1998). One reason for
its success is that units of study in biology (from
genes through organisms to higher taxa) do not
represent statistically independent observations
but rather are interrelated through their historical
connections. Therefore, almost any comparative
study requires information on phylogeny (Felsenstein 1985; Brooks & McLennan 1991; Harvey &
Pagel 1991). As a result, phylogenetic analyses are
now firmly entrenched in contemporary biology
and new applications continue to appear (e.g.
Harvey et al. 1996). Being the most central and
unifying concept in biology, phylogeny should
also 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 the
recognition of higher taxa (i.e. taxa above the level
of species): (1) higher taxa are named clades and,
therefore, represent monophyletic groups of species or less inclusive clades; hence, higher taxa
are delimited on the basis of common ancestry,
rather than a shared set of character states; (2)
phylogenetic knowledge should be expressed as
accurately as possible in nomenclature. As a consequence of the first rule, para- and polyphyletic
taxa should be abandoned. This has resulted in the
recognition of Casmerodius for Great White Egret C. aibus and Mergellus for Smew M. aibellus
and in the transfer of some species of Hippolais to
Acrocephalus.
The second rule implies that adjustments to
the present system should be enacted when im-
Sangster et al.: TAXONOMIC CHANGES IN 1977-1998
proved phylogenetic knowledge becomes available. For instance, in the past 15 years, evidence
for a sister-group relationship of Anseriformes
and Galliformes has accumulated and now greatly
outweighs all other hypotheses of relationships of
these orders. This has resulted in the recognition
of the taxon Galloanserae. More detailed knowledge of the relationships among cormorants and
shags, gannets and boobies, dabbling ducks, and
cardueline finches has resulted in the recognition
of four additional genera, Stictocarbo, Morus,
Mareca and Chloris. Generally, established usage
is maintained unless alternative hypotheses are
better supported. As a consequence, Wilson's
Phalarope Phalaropus tricolor is not placed in
Steganopus and Gelochelidon is not included in
Sterna. It has been attempted to not recognise
higher taxa that are not or only weakly supported
by phylogenetic analyses. This has resulted in the
inclusion of Chettusia in Vanellus, Catharacta in
Stercorarius and Tachymarptis in Apus.
Systematics and taxonomic (in)stability
There is a large hiatus between our knowledge
of phylogenetic relationships, which has greatly
increased since the late 1970s, and the avian taxonomic system (classification), which is basically
still the same as the one proposed by Alexander
Wetmore in 1930, which in tum was largely based
on the work of Max Ftirbringer and Hans Gadow
in the late 19th century. Some have suggested that
the goal of systematics is to produce a 'stable
standard sequence' and that the Wetmore classification and sequence is now so well entrenched
that it should continue to serve as the standard sequence (Mayr 1989; Mayr & Bock 1994). The
'stability' of the Wetmore sequence, especially in
the light of much systematic research, was viewed
by Mayr & Bock (1994) as 'of major advantage to
all avian biologists' because, in the Wetmore sequence, biologists can easily locate a particular
group or species. However, this 'stability' is not
to be regarded as a proof of its correctness. In
fact, the 'stability' of the Wetmore sequence
throughout this century is entirely due to failure
to incorporate new ideas about relationships (Si-
143
bley 1989; 1994). Fortunately, in recent years taxonomic systems have become more consistent
with current knowledge of phylogenetic relationships. Closely associated with this development is
a reappraisal of systematics as a historical science
(Gould 1986), in which phylogenies are viewed as
hypotheses of relationships and taxonomies based
on them as dynamic systems, subject to further
modification as our knowledge of relationships
grows.
Systematics is not an exercise in producing
classifications and sequences convenient for humans, but an attempt to discover an underlying
real structure in nature (Griffiths 1994). That real
structure, or natural system, is the pattern of historical (phylogenetic) relationships. This natural
system 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 of
these relationships. With its focus on past events
(i.e. historical subdivisions), systematics is a historical science.
It needs to be emphasised that reconstructed
phylogenies (cladograms) are hypotheses. The
true 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 true
phylogeny is unknown, there is no a priori basis
for accepting or rejecting a given phylogeny; hypotheses must be tested in the light of additional
data. A phylogenetic hypothesis is open to test by
the addition of new characters, the addition of
new 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 same
branching pattern. If congruence exists between
phylogenies based on different sets of data, this
may indicate a strong historical signal; congruence may be regarded as evidence that the relevant phylogenies identify the true organismal
phylogeny. Rejection of a phylogenetic hypoth-
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ARDEA 87(1), 1999
esis requires that an alternative hypothesis better
summarises all available evidence.
Because the purpose of taxonomy is communication of information about phylogenetic relationships, taxonomic systems should reflect the
current best-supported hypothesis of relationships. Taxonomic systems should be adjusted
only if it is believed that an alternative hypothesis
is better supported by the available evidence. Because phylogenies are hypotheses which are subject to further testing, taxonomies based on them
are provisional and may later be replaced or modified. Therefore, taxonomies are dynamic systems;
a definitive taxonomic system is probably unattainable in practice.
TAXONOMIC CHANGES
Galloanserae
A sister-group relationship of Anseriformes
and Galliformes is strongly supported by congruency of phylogenetic analyses of several independent data sets. These include morphological
characters (Cracraft 1986; 1988; Cracraft & Mindell 1989; Andors 1991; 1992; Kurochkin 1995;
Livezey 1997a), DNA-DNA hybridisation (Sibley
et al. 1988; Sibley & Ahlquist 1990; Harshman
1994; Bleiweiss et al. 1995), 12S and 16S ribosomal RNA sequences (Hedges et al. 1995), (Xcrystallin sequences (Hedges et al. 1995; Caspers
et al. 1997) and mitochondrial DNA sequences
(Mindell et al. 1997). The clade formed by Anseriformes and Galliformes was named Galloanserae
by Sibley et al. (1988). Most of these analyses
also indicate that Galloanserae is the sister taxon
of all extant birds except Paleognathae (Cracraft
1986; 1988; Cracraft & Mindell 1989; Sibley &
Ahlquist 1990; Hedges et ai. 1995; Kurochkin
1995; 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 list
Anseriformes before Galliformes and to place
these taxa before the remaining taxa on the Dutch
list.
Cygnus columbianus Whistling Swan
Fluitzwaan
Cygnus 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 differences
in morphology (Evans & Sladen 1980; Livezey
1996).
Anser fabalis Taiga Bean Goose Taigarietgans
Anser serrirostris Thndra Bean Goose
Toendrarietgans
Taiga Bean Goose and Tundra Bean Goose are
specifically distinct (Sangster & Oree11996; King
1997; Persson 1997; Wells 1998) based on differences in proportions, vocalisations, feeding habitat and diet, photosensitivity, activity pattern,
behaviour, phenology and responses to periods
of extreme cold (Berry 1938; Coombes 1951;
Mathiasson 1963; Huyskens 1977; 1986; Van Impe
1980; Kurechi et al. 1983; Van den Bergh 1985;
Barthel 1989; 1995; Burgers et al. 1991; Miyabayashi et al. 1994). Ringing recoveries suggest that
Taiga Bean Goose and Tundra Bean Goose have
allopatric breeding distributions (Burgers et al.
1991) and there is no evidence for 'intergradation'
(Sangster & Oreel 1996). In both Taiga Bean
Goose 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 Goose
Kleine Canadese Gans
Branta canadensis Greater Canada Goose
Grote Canadese Gans
Lesser Canada Goose and Greater Canada
Goose are specifically distinct (Sibley 1996) based
on congruence of phylogeographic analyses of
mitochondrial DNA restriction fragments
(Shields & Wilson 1987; Shields 1988; Van Wagner & Baker 1990; Quinn et al. 1991), mitochondrial DNA sequences (Quinn et al. 1991; Ba-
ker & Marshall 1997) and morphometry (Van
Wagner & 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 with
B. canadensis.
Branta bernicla Dark-bellied Brent Goose
Rotgans
Branta hrota Pale-bellied Brent Goose
Witbuikrotgans
Branta nigricans Black Brant Zwarte Rotgans
Dark-bellied Brent Goose, Pale-bellied Brent
Goose and Black Brant are specifically distinct
(Millington 1997) based on qualitative differences
in morphology (Delacour 1954; Johnsgard 1978;
Millington 1997), overlapping breeding ranges of
Pale-bellied Brent Goose and Black Brant in arctic Canada (Gavin 1947; Handley 1950) and segregation of Pale-bellied and Dark-bellied Brent
Goose in the Netherlands and Denmark (Lambeck 1981). The alleged hybrid origin of 'intermediate' populations in central arctic Canada, which
formed the basis for including Pale-bellied Brent
Goose and Black Brant, along with Dark-bellied
Brent Goose, in a single species (Delacour &
Zimmer 1952), has been falsified by genetic analysis (Shields 1990).
Mareca penelope Eurasian Wigeon Smient
Mareca americana American Wigeon
Amerikaanse Smient
Mareca falcata Falcated Duck Bronskopeend
Mareca strepera Gadwall Krakeend
Phylogenetic analyses based on mitochondrial
DNA and morphology (Kessler & Avise 1984;
Livezey 1991) provide strong support for the existence of two major clades within the dabbling
ducks traditionally included in Anas (Omland
1994): (1) a clade formed by Cape Teal M. capensis, the wigeons, Falcated Duck and Gadwall; and
(2) a clade formed by the remaining species. We
adopt the classification of Livezey (1991; 1997b)
in which the members of the former clade are
placed in Mareca and the remaining species in
145
Anas (King 1997). Current knowledge of the phylogenetic relationships of dabbling ducks is better
represented with the recognition of two genera
than with the placement of all species in Anas.
Anas crecca Common Teal Wintertaling
Anas carolinensis Green-winged Teal
Amerikaanse 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 Scoter
Zwarte Zee-eend
Melanitta americana Black Scoter
Amerikaanse 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-eend
Velvet Scoter and White-winged Scoter M.
deglandi are specifically distinct (Stepanyan
1990; Livezey 1995b; King 1997) based on qualitative differences in morphology (Johnsgard 1978;
Livezey 1995b).
Mergellus albellus Smew Nonnetje
Lophodytes cucullatus Hooded Merganser
Kokardezaagbek
A phylogenetic analysis based on morphology
(Livezey 1995b) indicates that Smew is more
closely related to the goldeneyes Bucephala than
to the mergansers Mergus. Because the inclusion
of 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). We
place Hooded Merganser in the monotypic genus
Lophodytes (AOU 1983; BOURC 1997; King
1997) to indicate its basal position among the mergansers (Livezey 1995b). The latter species is fre-
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ARDEA 87(1),
quently recorded in the Netherlands but records
are considered to refer to escapes from captivity.
Hooded Merganser, therefore, is not formally admitted to the Dutch list.
Soft-plumaged petrel complex donsstormvogels
Fea's Petrel Pterodromafeae, Zino's Petrel P.
madeira and Soft-plumaged Petrel P. mollis are
specifically distinct (Bourne 1983; Collar & Stuart
1985; Zino & Zino 1986; Sibley & Monroe 1990;
Beaman 1994; Hazevoet 1995; 1997; Sibley 1996;
Snow & Pemns 1998) based on phylogeographic
analysis of mitochondrial DNA sequences (Nunn
& Zino in press) and concordance of differences in
morphology (Zino & Zino 1986), vocalisations
(Bretagnolle 1995) and reproductive behaviour
(Zino & Zino 1986). Analysis of mitochondrial
DNA sequences suggests that the divergence of P.
feae and P. madeira occurred 840 000 years ago
and that P. mollis is not closely related to P. feae
and P. madeira (Nunn & Zino in press). Populations of Fea's Petrel breeding on the Deserta
Islands, Madeira Cdeserta'), are provisionally retained conspecific with P. feae (Snow & Pemns
1998). 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 et
al. 1995), was accepted as P.feaelmadeiralmollis.]
Puffinus mauretanicus Balearic Shearwater
Vale Pijlstormvogel
Puffinus puffinus Manx Shearwater
Noordse Pijlstormvogel
Balearic Shearwater is specifically distinct
from Manx Shearwater and Yelkouan Shearwater
P. yelkouan (McMinn et al. 1990; Walker et al.
1990; Altaba 1995; Sibley 1996; King 1997; Heidrich et al. 1998; Snow & Perrins 1998; Wells
1998) based on analyses of qualitative morphological characters (Walker et al. 1990; Altaba
1995) and phylogenetic analysis of mitochondrial
DNA (Austin 1996; Heidrich et al. 1998). The recent discovery and subsequent description of two
1999
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, casts
doubt on the alleged sister relationship (and conspecificity) of Balearic Shearwater and Yelkouan
Shearwater (Altaba 1995). It has been suggested
that Balearic Shearwater may actually be more
closely related to Hole's Shearwater than to Yelkouan Shearwater (Walker et al. 1990).
Calonectris borealis Cory's Shearwater
Kuhls Pijlstormvogel
Cory's Shearwater and Scopoli's Shearwater
C. diomedea are specifically distinct based on
phylogeographic analysis of allozymes (Randi et
al. 1989) and mitochondrial DNA (Heidrich et al.
1996; 1998), qualitative differences in vocalisations (Bretagnolle & Lequette 1990) and analysis
of 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)
and 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 of
sight records of C. borealislC. diomedea in the
Netherlands is currently being investigated by the
Dutch rarities committee (CDNA).]
Morus bassanus Northern Gannet Jan van Gent
Phylogenetic relationships among Sulidae
(Warheit 1992; Friesen & Anderson 1997) are best
represented with the recognition of Morus for the
gannets and Sula for the boobies, with the exception of Abbott's Booby Papasula abbotti (Olson
1985; Olson & Warheit 1988; Van Tets et al. 1988;
AOU 1989; Sibley & Monroe 1990; Warheit 1992;
Friesen & Anderson 1997).
Stictocarbo aristotelis European Shag
Kuifaalscholver
A phylogenetic analysis of morphological
characters (Siegel-Causey 1988) identified nine
major 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 of
all species in a single genus Phalacrocorax
(Bourne & Casement 1996).
Casmerodius albus Great White Egret
Grote Zilverreiger
Phylogenetic analyses based on morphology
and DNA-DNA hybridisation (Payne & Risley
1976; Sheldon 1987; Sheldon et al. 1995) indicate
that Great White Egret is not closely related to the
Egretta clade and instead suggest a closer relationship with Bubulcus and Ardea. However,
given the unresolved relationships between Great
White Egret, Bubulcus and Ardea, inclusion of
Great White Egret in Ardea (AOU 1995; BOURC
1997) is premature. Until the relationships of
Great White Egret are better understood it is best
placed in a monotypic genus Casmerodius (AOU
1983; Eck 1996; Inskipp et al. 1996; King 1997;
Wells 1998).
Phoenicopterus roseus Greater Flamingo
Flamingo
Greater Flamingo and Caribbean Flamingo P
ruber are specifically distinct (Allen 1956; Morony et al. 1975; Hazevoet 1995; Sibley 1996;
King 1997; Sangster 1997a) based on qualitative
differences in plumage and bill pattern (Van den
Berg 1987; Sangster 1997a) and display behaviour
and vocalisations (Studer-Thiersch 1964; 1974;
1975).
Aquila nipalensis Steppe Eagle Steppearend
Steppe Eagle and Tawny Eagle A. rapax are
specifically distinct (Brooke et al. 1972; Clark
1992; Olson 1994; King 1997; Wells 1998) based
on qualitative morphological differences (Brooke
et al. 1972; Clark 1992; Olson 1994).
147
Porphyrio madagascariensis African Swamp-hen
Smaragdpurperkoet
Porphyrio poliocephalus Grey-headed Swamp-hen
Grijskoppurperkoet
Western Swamp-hen P porphyrio, African
Swamp-hen, Grey-headed Swamp-hen, Philippine Swamp-hen P pulverulentus, Black-backed
Swamp-hen P indicus and Australian Swamp-hen
P melanotus are specifically distinct (Sangster
1998) based on qualitative differences in morphology (Ripley 1977; Cramp & Simmons 1980; Del
Hoyo et al. 1996). Analyses of mitochondrial
DNA suggest that forms previously included under the name 'Purple Swamp-hen P porphyrio'
(Von Boetticher 1935; Ripley 1977; Del Hoyo et
al. 1996) are paraphyletic with respect to two
large flightless New Zealand taxa, South Island
Takahe P hochstetteri and extinct North Island
Takahe P mantelli (Trewick 1997). These results
argue against continued inclusion of swamp-hen
forms in a single polytypic species. The six
groups here treated as species (P porphyrio, P
madagascariensis, P poliocephalus, P pulverulentus, P indicus and P melanotus) are similar to
those recognised by Roselaar (in Cramp & Simmons 1980) as subspecies groups. Pending further
analysis, caspius and seistanicus are tentatively
included in P poliocephalus; viridis is tentatively
included in P indicus; and bellus, chathamensis,
melanopterus, pelewensis and samoensis are tentatively included in P melanotus. [The inclusion
of African Swamp-hen and Grey-headed Swamphen on the Dutch list is currently under review by
the Dutch rarities committee (CDNA).]
Chlamydotis macqueenii Macqueen's Bustard
Oostelijke Kraagtrap
Macqueen's Bustard and Houbara Bustard C.
undulata are specifically distinct (Gaucher et al.
1996; Sangster 1996; King 1997; Wells 1998) based
on qualitative differences in courtship behaviour
and genetic analysis (Granjon et al. 1994;
Gaucher et al. 1996).
148
ARDEA 87(1),
Pluvialis dominicus American Golden Plover
Amerikaanse Goudplevier
Pluvialis fulva Pacific Golden Plover
Aziatische Goudplevier
American Golden Plover and Pacific Golden
Plover are specifically distinct (Knox 1987; AOU
1993; King 1997; Wells 1998) based on differences
in plumage, morphology, moult, vocalisations,
ecology and overlap of breeding ranges (Connors
1983; Connors et af. 1993). The correct name of
American Golden Plover is P. dominicus, not P.
dominica (AOU 1995).
Vanellus gregarius Sociable Lapwing
Steppekievit
Vanellus leucurus White-tailed Lapwing
Witstaartkievit
Phylogenetic analyses of behavioural characters (Ward 1992) have been unable to resolve relationships among lapwings. Given the doubtful
monophyly of Chettusia (and Hoplopterus), the
recognition of Chettusia (and Hoplopterus) is not
justified. Therefore, all lapwings are placed in
Vanellus (BOURC 1997; King 1997; Wells 1998).
Gallinago gallinago Common Snipe Watersnip
Common Snipe and Wilson's Snipe G. delicata are specifically distinct (Olsson 1987; Gantlett et af. 1996; King 1997) based on qualitative
differences in morphology, vocalisations and
drumming 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, Paraguayan
Snipe G. paraguaiae, Magellan Snipe G. magellanica and Puna Snipe G. andina are specifically
distinct from Common Snipe based on qualitative
differences in morphology, vocalisations and
drumming display (Tuck 1972; Sutton 1981; Hayman et af. 1986; Fjeldsa & Krabbe 1990; Del Hoyo
et af. 1996).
1999
Phalaropus tricolor Wilson's Phalarope
Grote Franjepoot
Results of phylogenetic analyses based on allozymes (Dittmann et al. 1989), mitochondrial
DNA (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; Del
Hoyo et al. 1996; Higgins & Davies 1996) is unjustified and, therefore, we retain Wilson's Phalarope in Phalaropus.
Phalaropus fulicaria Grey Phalarope
Rosse Franjepoot
The correct name of Grey Phalarope is P. fulicaria, not P. fulicarius (Parkes 1982).
Stercorarius skua Great Skua Grote Jager
Recent phylogenetic analyses of allozymes
and mitochondrial DNA sequences (Cohen et af.
1997; Braun & Brumfield 1998) confirm and extend
previous suggestions based on short mitochondrial
DNA sequences (Blechschmidt et al. 1993) and
ecology (Andersson 1973) that Pomarine Skua S.
pomarinus is more closely related to species placed
in Catharacta than to Arctic Skua S. parasiticus
and 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 a
paraphyletic taxon, all skuas are placed in Stercorarius.
Larus graellsii Lesser Black-backed Gull
Kleine Mantelmeeuw
Larus fuscus Baltic Gull
Baltische Mantelmeeuw
Lesser Black-backed Gull and Baltic Gull are
specifically distinct based on qualitative differences in morphology and differences in moult and
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
form 'intermedius' is diagnosably distinct from
graellsii; 'intermedius' is, therefore, considered
conspecific 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 and
Vega Gull L. vegae based on qualitative differences in morphology and behaviour, and differences in ecology (Grant 1986; Filchagov et aL
1992; Hario 1992; Kennerley et aL 1995; Yesou &
Hirschfeld 1997), The breeding range of Heuglin's Gull overlaps with that of Herring Gull and
Baltic Gull, with evidence for reproductive isolation (Filchagov & Semashko 1987; Filchagov et
aL 1992; Filchagov 1994). Pending further analysis, taimyrensis and heuglini are provisionally retained as conspecific (Kennerley et al. 1995).
Larus argentatus Herring Gull Zilvermeeuw
Larus michahellis Yellow-legged Gull
Geelpootmeeuw
Larus 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; Kennerley
et aL 1995; Dubois 1997; Chu 1998). There is no
evidence that the form 'argenteus' is diagnosably
distinct from argentatus. Current evidence indicates a clinal pattern of variation (Barth 1968;
Cramp & Simmons 1983); the design of studies
which have suggested clear differences between
populations of 'argenteus' and argentatus (Monaghan et aL 1983; Golley 1993) was inadequate to
substantiate such claims (Chylarecki 1993). The
form '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) based
on qualitative differences in adult and immature
plumages, bare parts, behaviour and vocalisations
and overlap of breeding ranges (Nicolau-Guillaumet 1977; Glutz von Blotzheim & Bauer 1982;
149
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 in
morphology and vocalisations, and differences in
behaviour 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 is
provisionally retained as conspecific with L.
michahellis; barabensis and mongolicus are provisionally retained as conspecific with L. cachinnans.
Armenian Gull L. armenicus is specifically
distinct from Pontic Gull, Yellow-legged Gull and
Heuglin's Gull L. heuglini based on qualitative
differences in morphology and vocalisations
(Geroudet 1982; Hume 1983; Dubois 1985; Grant
1986; 1987; Satat & Laird 1992; Buzun 1993; Filchagov 1993; Frede & Langbehn 1997; Yesou &
Hirschfeld 1997).
Gelochelidon nilotica Gull-billed Tern
Lachstern
Although one phylogenetic study based on allozymes suggests that Gull-billed Tern originates
within the Sterna clade (Randi & Spina 1987),
others, based on allozymes (Hackett 1989), hindlimb musculature (McKitrick 1991) and osteology
(Chu 1995), suggest a more distant relationship
and do not support the inclusion of Gull-billed
Tern 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 among
terns, as indicated by the incongruence of available analyses, we retain Gelochelidon for Gullbilled Tern.
Apus melba Alpine Swift Alpengierzwaluw
Evidence is lacking for a sister relationship
between Alpine Swift and Mottled Swift A. aequatorialis and monophyly of the other species
traditionally placed in Apus. Therefore, the recog-
150
ARDEA 87(1), 1999
mtlOn of Tachymarptis for Alpine Swift and
Mottled Swift and a more restricted Apus for the
remaining species (Brooke 1972; Fry et al. 1988;
Short et al. 1990; Chantler & Driessens 1995; Eck
1996) may render Tachymarptis and/or Apus paraphyletic. In the absence of a relevant phylogenetic
analysis of the swifts, we retain Alpine Swift in
Apus.
Merops persicus Blue-cheeked Bee-eater
Groene 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 Pieper
Richard's Pipit is specifically distinct from
Grassland A. cinnamomeus, Paddyfield A. rufuIus, Australian A. australis and New Zealand Pipit
A. novaeseelandiae (Glutz von Blotzheim &
Bauer 1985; Sibley 1996; Wells 1998) based on
qualitative differences in plumage and vocalisations (Glutz von Blotzheim & Bauer 1985 and references cited therein).
Anthus spinoletta Water Pipit Waterpieper
Anthus petrosus Rock Pipit Oeverpieper
Water Pipit, Rock Pipit and Buff-bellied Pipit
A. rubescens are specifically distinct (Oreel 1980;
AOU 1989; King 1997) based on qualitative differences in plumage, vocalisations and ecology
(Bijlsma 1977; Alstrom & Olsson 1987; Knox
1988a).
Motacillaflavissima Yellow Wagtail
Engelse Kwikstaart
Motacillaflava Blue-headed Wagtail
Gele Kwikstaart
Motacilla thunbergi Grey-headed Wagtail
Noordse Kwikstaart
Motacillafeldegg Black-headed Wagtail
Balkankwikstaart
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, Kamtchatka
Wagtail 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 von
Blotzheim & Bauer 1985; Cramp 1988; Leader
1996). For several named populations there is no
evidence or insufficient evidence that these are diagnosably distinct. Therefore, 'beema' is included
in M. jlava, 'angarensis', 'macronyx' and 'plexa'
in M. thunbergi, 'aralensis', 'kaleniczenkoi' and
'melanogrisea' in M. feldegg and 'pygmaea' in M.
cinereocapilla. The form 'dombrowskii' probably
refers 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 Kwikstaart
Motacilla 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 Baikal
Wagtail M. baicalensis are specifically distinct
(Glutz von Blotzheim & Bauer 1985; Cramp
1988). There is no evidence that populations in
western Asia (,dukhunensis') are diagnosably distinct from alba. Therefore, 'dukhunensis' is included in M. alba. The form 'persica' probably
represents a variable hybrid population of alba
and personata (Vaurie 1959; Cramp 1988) and is
not recognised.
Saxicola rubicola European Stonechat
Roodborsttapuit
Saxicola maura Siberian Stonechat
Aziatische Roodborsttapuit
European Stonechat, Siberian Stonechat and
African 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 that
populations 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). Pending
further analysis, variegata, armenica, indica and
przewalskii are provisionally retained as conspecific with S. maUTa.
Oenanthe hispanica Western Black-eared
Wheatear Westelijke Blonde Tapuit
Oenanthe melanoleuca Eastern Black-eared
Wheatear Oostelijke Blonde Tapuit
Western Black-eared Wheatear and Eastern
Black-eared Wheatear are specifically distinct
(Clement & Harris 1987; Cramp 1988).
Zoothera aurea White's Thrush Goudlijster
White's Thrush and Scaly Thrush Z. dauma
are specifically distinct (Eck 1996) based on qualitative differences in morphology and vocalisations (Seebohm & Sharpe 1902; Ali & Ripley
1973; Cramp 1988; Glutz von Blotzheim & Bauer
1988; Martens & Eck 1995). There is no evidence
that populations in south-eastern Siberia, Russia
and 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 Thrush
Z. machiki, New Britain Thrush Z. talaseae, San
Cristobal Thrush Z. margaretae, Guadalcanal
Thrush Z. turipavae, Bassian Thrush Z. lunulata
and Russet-tailed Thrush Z. heinei are specifically
distinct (Mayr 1955; Deignan et al. 1964; Ford
151
1983; Ishihara 1986; White & Bruce 1986; Christidis & Boles 1994; Gibbs 1996; Inskipp et al. 1996;
Sibley 1996; King 1997; Wells 1998) based on
qualitative differences in morphology and vocalisations (Seebohm & Sharpe 1902; Jahn 1942;
Mayr 1955; Ali & Ripley 1973; Ford 1983; Ishihara 1986).
Acrocephalus agricola Paddyfield Warbler
Veldrietzanger
Paddyfield Warbler and Manchurian Warbler
A. tangorum are specifically distinct (Round 1994;
King 1997) based on qualitative differences in
plumage (Alstrbm et al. 1991; Round 1994). Phylogenetic analysis of mitochondrial DNA sequences
indicates 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 Reed
Warbler A. avicenniae, African Reed Warbler A.
baeticatus and Caspian Reed Warbler A. fuscus
are specifically distinct (Leisler et al. 1997; Sangster 1997b) based on qualitative differences in
morphology (Pearson 1981; Ash et al. 1989; Harris
et al. 1995). Phylogenetic analysis of mitochondrial DNA sequences indicates that Mangrove
Reed Warbler, which is currently regarded as a
subspecies of A. baeticatus, is actually more
closely related to European Reed Warbler, and
that European and Caspian Reed Warbler, until
recently regarded as subspecies of A. scirpaceus,
are not sister-taxa (Leisler et al. 1997).
Acrocephalus caligatus Booted Warbler
Kleine Spotvogel
Phylogenetic analyses of mitochondrial DNA
sequences indicate that Booted Warbler and Olivaceous Warbler A. pallidus are more closely related to species traditionally included in Acrocephalus clade rather than to Icterine Warbler
Hippolais icterina (Leisler et al. 1997; Sangster
1997b). Therefore, Booted Warbler and Oliv-
152
ARDEA 87(1),
aceous Warbler are placed in Acrocephalus.
Booted Warbler and Sykes's Warbler A. rama
are specifically distinct (Stepanyan 1990; Glutz
von Blotzheim & Bauer 1991; Sibley & Monroe
1993; 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, its
placement in Acrocephalus is tentative.
Phylloscopus proregulus Pallas's Leaf Warbler
Pallas' Boszanger
Pallas's Leaf Warbler and Lemon-rumped
Warbler P. chloronotus are specifically distinct
(Inskipp et al. 1996; King 1997; Wells 1998) based
on qualitative differences in plumage and vocalisations (Martens 1985; Alstrom & Olsson 1990).
Phylloscopus inornatus Yellow-browed Warbler
Bladkoning
Phylloscopus humei Hume's Warbler
Humes Bladkoning
Yellow-browed Warbler and Hume's Warbler
are specifically distinct (Svensson 1992; Gantlett
et al. 1996; BOURC 1997; King 1997; Wells 1998)
based on qualitative differences in vocalisations
and plumage and overlap of breeding ranges
(Mild 1987; Aistrom & Olsson 1988; Dathe &
Loskot 1989).
Phylloscopus orientalis Eastern Bonelli's Warbler
Balkanbergfluiter
Phylloscopus bonelli Western Bonelli's Warbler
Bergfluiter
·Eastern Bonelli's Warbler and Western Bonelli's Warbler are specifically distinct (Gantlett et
al. 1996; BOURC 1997; King 1997; Wells 1998)
based on qualitative differences in vocalisations
and genetic analyses (Helb et al. 1982; Helbig et
al. 1995).
Phylloscopus collybita Common Chiffchaff
Tjiftjaf
1999
Phylloscopus brehmii Iberian Chiffchaff
Iberische Tjiftjaf
Common Chiffchaff, Iberian Chiffchaff and
Canarian 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 in
structure and plumage (Shirihai 1987; Cramp
1992), qualitative differences in vocalisations
(Martens & Hanel 1981; Martens 1982) and genetic analyses (Helbig et al. 1996). Pending further analysis, abietinus and tristis are tentatively
included in P. collybita.
Lanius phoenicuroides Thrkestan Shrike
Turkestaanse Klauwier
Lanius speculigerus Daurian Shrike
Daurische Klauwier
Turkestan Shrike and Chinese Shrike L. isabellinus are specifically distinct (Kryukov 1995;
Panov 1995; Panow 1996) based on qualitative
differences in morphology (Dean 1982; Cramp &
Pemns 1993; Panow 1996; Lefranc & Worfolk
1997) and analyses of their contact zone (Kryukov
1995).
Daurian Shrike is specifically distinct from
Turkestan Shrike and Chinese Shrike based on
qualitative differences in morphology (Dean
1982; Cramp & Pemns 1993; Panow 1996) and
vocalisations (Panov 1995). Pending further analysis, tsaidamensis is provisionally retained as
conspecific 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
being investigated by the Dutch rarities committee (CDNA).]
Lanius excubitor Great Grey Shrike Klapekster
Lanius pallidirostris Steppe Grey Shrike
Steppeklapekster
Great Grey Shrike and Southern Grey Shrike
L. 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 qualitative
differences in plumage, breeding ecology and behaviour (Isenmann & Bouchet 1993; Eck 1994;
Lefranc 1995a; 1995b; Panow 1996; Lefranc &
Worfolk 1997).
Steppe Grey Shrike is specifically distinct
from Great Grey Shrike and Southern Grey Shrike
(King 1997) based on qualitative differences in
plumage, breeding ecology, behaviour and overlap of breeding ranges (Panov 1995; Panow 1996;
Lefranc & Worfolk 1997).
Corvus corone Carrion Crow Zwarte Kraai
Corvus 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 in
plumage and analyses of their hybrid zone in Germany (Risch & Andersen 1998) and Italy (Saino
1990; Saino & Scatizzi 1991; Saino 1992; Saino &
Bolzern 1992; Saino & Villa 1992; Rolando 1993;
Rolando & Laiolo 1994; Rolando & Saino 1994).
Chloris chloris Common Greenfinch Groenling
Phylogenetic analyses of morphological characters (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 two
independent studies suggest that Chloris is not
part of the Carduelis clade and identify the same
sister-taxon (Raikow 1978; Fehrer 1996), whereas
inclusion of Chloris in Carduelis is supported by
only one study (Van den Elzen & Nemeschkal
1991), recognition of Chloris for the greenfinches
is justified.
153
Carduelis cannabina Linnet Kneu
Carduelis flavirostris Twite Frater
Carduelis cabaret Lesser Redpoll Kleine Barmsijs
Carduelisflammea Mealy Redpoll Grote Barmsijs
Carduelis hornemanni Arctic Redpoll
Witstuitbarmsij s
Published studies of phylogenetic relationships among cardueline finches (Marten & Johnson 1986; Van den Elzen & Nemeschkal 1991;
Fehrer 1996) are contradictory with regard to the
phylogenetic relationships of Acanthis and Carduelis. Because monophyly of Acanthis has not
been established and is contradicted by one study
(Van den Elzen & Nemeschkal 1991), recognition
of Acanthis may not significantly contribute to the
elimination of paraphyletic taxa. Although Carduelis as defined here (i.e. including Acanthis but
excluding Chloris) is still likely to be paraphyletic, other hypotheses of relationships, which
require changes in nomenclature, do not seem to
be better supported by available data. Therefore,
pending further phylogenetic analyses, Linnet,
Twite, Lesser Redpoll, Mealy Redpoll and Arctic
Redpoll are provisionally retained in Carduelis.
Lesser Redpoll and Mealy Redpoll are specifically distinct based on qualitative differences in
morphology (Knox 1988b; Herremans 1990) and
vocalisations (Herremans 1989), and overlap of
breeding ranges in south-eastern Norway without
hybridisation (Lifjeld & Bjerke 1996). Pending
further analysis, rostrata and exilipes are provisionally retained as conspecific with C. fiammea
and C. hornemanni, respectively.
Dendroica coronata Myrtle Warbler
Mirtezanger
Myrtle Warbler and Audubon's Warbler D.
auduboni are specifically distinct (Bermingham et
al. 1992) based on qualitative differences in plumage (Hubbard 1970; Kaufman 1979; Cramp & Perrins 1994; Dunn & Garrett 1997) and analysis of
their hybrid zone (Barrowclough 1980). It has
been calculated (Zink & McKitrick 1995) that it
would take more than 6 million years for these
taxa to completely fuse. Data in Bermingham et
al. (1992) show that many North American Den-
154
ARDEA 87(1), 1999
droica species are less than 3.5 million years old
which means that speciation in several species of
Dendroica took place during a much shorter time
than the period calculated as necessary for Myrtle
and Audubon's Warblers to fuse.
Icterus galbula Baltimore Oriole
B altimoretroepiaal
Baltimore Oriole, Bullock's Oriole I. bullockii
and Black-backed Oriole I. abeillei are specifically distinct (AOU 1995; Freeman & Zink 1995;
BOURC 1997) based on qualitative differences in
plumage and vocalisations, analyses of the hybrid
zone of Baltimore Oriole and Bullock's Oriole
(see AOU 1995 for references) and a phylogenetic
analysis (Freeman & Zink 1995) indicating that
Baltimore Oriole and Bullock's Oriole are not
eachother's closest relatives.
ACKNOWLEDGEMENTS
We thank Walter J. Bock and Ernst Mayr for providing
valuable and stimulating comments and criticisms. We
are 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 Netherlands
Ornithologists' Union (NOU) and the Dutch Birding
Association (DBA).
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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 een
aantal besluiten genomen ten aanzien van door haar
noodzakelijk geachte taxonornische en nomenclatorische wijzigingen. Dit artikel bevat een samenvatting
van de genomen besluiten en de eraan ten grondslag
liggende argumenten.
Een belangrijk deel van de taxonornische veranderingen betreft de soortstatus van bepaalde taxa, die
vroeger veelal de status van ondersoort hadden. Dit is
een resultaat van een verandering in de wetenschappelijke inzichten in het soortsbegrip. Traditioneel wordt
er 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 tot
dezelfde soort. Het is echter allang bekend, dat er problemen aan dit, op het eerste gezicht zo logischlijkende, isolatie-soortsbegrip kleven. Zo blijkt het in de
natuur nogal eens voor te komen dat individuen uit
twee groepen weI met elkaar kunnen kruisen, terwijl zij
niet elkaars nauwste verwanten zijn (in de zin van
bloedverwantschap, genealogie, afstamming). Daarnaast gaat het isolatie-soortsbegrip er bij het afbakenen
van soorten vanuit dat deze soorten in de toekomst reproductief van elkaar gei"soleerd zullen blijven en niet
tot een populatie zullen fuseren. Dit betekent, dat in
feite toekomstige gebeurtenissen uitsluitsel moeten geven of we huidige groepen als aparte soorten of slechts
als 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 toch
tot slechts een soort gerekend (omdat de onderlinge
verschillen 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
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 het
zogenaamde 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 periode
van de systematische biologie. Als gevolg hiervan heeft
nu alom het idee postgevat dat kennis van en inzicht in
de afstammings- of fylogenetische relaties tussen groepen van doorslaggevend belang is in het vakgebied van
de systematische biologie en de evolutiebiologie. Dit
heeft als consequentie dat van een modem soortsbegrip
verwacht mag worden dat het in overeenstemming is
met de fylogenetische relaties tussen de onderscheiden
eenheden. Soortsbegrippen, zoals het isolatie-soortsbegrip, die groepen van individuen bij elkaar zetten die
niet elkaars nauwste verwanten zijn, geven dientengevolge een vertekend beeld van de evolutionaire geschiedenis. Fylogenetische soortsbegrippen voldoen in
dit opzicht beter.
Er zijn twee typen van fylogenetische soortsbegrippen, te weten de monofylie-versie en de diagnostische
versie. De monofylie-versie van het fylogenetische
soortsbegrip eist dat elke soort gekenmerkt wordt door
minimaal een uniek kenmerk dat niet bij andere soorten
voorkomt. De diagnostische versie is wat minder strikt
en eist slechts dat een soort gekarakteriseerd wordt
door een unieke combinatie van kenmerken; d.w.z. de
afzonderlijke kenmerken mogen weI bij andere soorten
voorkomen, maar hun combinatie moet uniek zijn. De
CSNA heeft de diagnostische versie van het fylogenetische soortsbegrip gehanteerd bij het samenstellen van
deze rapportage.
Een tweede aspect van de avifaunistiek en de taxonomie dat door de CSNA aan een nadere beschouwing
is onderworpen, betreft het groeperen van soorten in
hogere eenheden, zoals geslachten en families, en de
indeling van de lijst (dat wil zeggen welke groepen
staan bovenaan en welke onderaan de lijst). Ook deze
zaken dienen, naar huidige inzichten, een afgeleide te
zijn van de historische, fylogenetische relaties tussen
de onderscheiden groepen. Twee voorbeelden uit de
huidige 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-
165
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 hogere
orde plaatsen, de Galloanserae.
De eerder genoemde revolutionaire ontwikkeling
binnen de systematische biologie en de evolutiebiologie, en de moderne opvatting dat de nieuwe inzichten
en 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 voordeel
van de oude systemen en lijsten is dat ze stabiel zijn:
hun onveranderlijkheid geeft houvast bij het opzoeken
van informatie. Ret is echter een feit dat deze stabiliteit, zo niet starheid, tot gevolg heeft dat de oude avifaunistische en taxonomische lijsten geen afspiegeling
meer 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 die
lijsten en systemen dan weI vertegenwoordigen. Ret
klassieke antwoord dat zij ingeburgerde en gemakkelijke zoeksystemen vertegenwoordigen, is wetenschappelijk gezien niet acceptabel. Als wetenschappelijke
onderbouwing niet meer van belang wordt geacht, dan
zijn er praktischere zoeksystemen denkbaar, bijvoorbeeld alfabetische lijsten. Daarom is de CSNA van mening dat taxonomische systemen geregeld dienen te
worden aangepast om zo een afspiegeling te zijn van de
meest modeme hypothesen over de fylogenetische relaties tussen groepen van vogels. Ret is evident, dat bij
een voortschrijdende wetenschap ook taxonomische
systemen en lijsten niet stil kunnen blijven staan en altijd aan verandering onderhevig zullen zijn.
Received 11 September 1998, accepted 29 March 1999
Corresponding editor: Kees (C.J.) Camphuysen
Note from the editors of Ardea
The taxonomic and nomenclatural changes listed
above, as proposed by the Dutch committee for avian
systematics (CSNA), will be adopted by Ardea in this
and 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 clear
that there is still considerable opposition to the principles underlying the present changes. Some referees
were not convinced of the advantages of the Phylogenetic Species Concept (PSC). Not surprisingly, these
referees could not recommend the publication of this
paper in Ardea. Other referees were completely in
favour of the principles of a PSC and strongly recommended the present publication. Apparently, there was
little or no ground for compromise. In the sometimes
heated discussions generated by the proposals of the
CSNA (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 inspired
by a similar mix of reasons. We believe that the CSNA
has convincingly defended their proposal to abandon
the traditional Isolation Species Concept (ISC) in favour of a Phylogenetic Species Concept, both in the introductory statements in the present publication and at
the NOV symposium in Leiden in 1999. Yet, it is clear
that we deal with hypotheses, and CSNA has confirmed
that changes in specific status, sequences or positions
on the list are and will be proposed only if the evidence
is 'overwhelming'. Ever since the earlier publications
of CSNA in Dutch Birding, several editors of ornitho-
logical journals and books in The Netherlands have
adopted the changes while others decided to continue
to use more traditional lists. The present situation, a
split into two ('followers' and 'non-followers'), is very
inconvenient. Both the NOV and the editorial team of
Ardea are convinced that a decision has to be made, so
that the CSNA could either continue to function as an
advisory committee of NOVIDBA, or should operate
independently.
After the publication of this list, the editorial team
of Ardea will seek advice from CSNA regarding the
name 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 inform
the readers about the name/status of these species as
they were known prior to the present changes. Because
the CSNA is the only body in The Netherlands that is
specialised in the evaluation of the taxonomic status
and systematics of birds, we recommend that proposals
published in Ardea by this committee are followed. At
the same time, we challenge scientists to constructively
comment on CSNA decisions and proposals, so that, in
the process, we deepen our understanding of evolutionary relationships and patterns of ancestry and descent
of contemporary birds.

Sangster et al 1999

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