Introduction

Ends and beginnings: the taxonomic cycle Truly integrative taxonomy:

With this companion volume to del Hoyo & Collar (2014) the HBW-Bird-
building the broadest evidence base
Life International project to review and revise the taxonomy of the birds
of the world reaches its first goal. A multiplicity of concerns and consid- There is an appealing and increasing recognition in modern avian taxon­
erations has driven the endeavour, including the geographical patchiness omy of the value of what has been termed an “integrative” approach to
of taxonomic research, the consequent neglect of many pressing and de- revisions, one which seeks to include all varieties of evidence in order
serving cases (often highlighted by the species profiles in the Handbook of to test and improve the robustness of the results obtained by one partic­
the Birds of the World [HBW] itself), the sometimes glacial speeds at which ular—almost invariably genetic—line of inquiry. Plumage pattern and
appointed regional committees take decisions, the potentially dramatic structure, bare part colours and formations, morphometrics, vocalizations,
disparities of treatment resulting from choice and understanding of spe- ecological factors, behavioural traits and molecular findings all represent
cies concept, the poor standards of some published analyses, the often variables that can be assessed for information relevant to a taxonomic
uncritical adoption of proposed changes by existing world checklists, the identity. Truly “integrative” work will endeavour to cover as many of these
increasing availability and relevance of sound recordings, and BirdLife’s parameters as possible (see such exemplary studies as Payne & Sorenson
enduring obligation to IUCN (International Union for Conservation 2007, Davison et al. 2012, Sangster et al. 2013, 2016, Alström et al. 2016).
of Nature) to evaluate the conservation status of all species of bird. We This is the fundamental premise on which this checklist is built.
wanted to make fullest use of the results of recent studies, assembling and The more these factors line up together, the more confident we can
integrating modern assessments and evidence with the often detailed tax- be of the taxonomic conclusion to which their alignment points. (Thus
onomic notes already elaborated by the many and various HBW authors, the plumage differences and morphometric disjunctions—so lightly
paying particular attention to the findings of molecular research; but we and benightedly dismissed as “relatively minor” by the author of HBW’s
also wanted to maintain as much independence of judgement as circum- Turdidae—shown by the Eastern Slaty Thrush Turdus subalaris mark it
stances allowed, and to be proactive where nobody else was taking a lead. emphatically, with a score of 9, as a species distinct from Andean Slaty
Consequently this volume, like its predecessor, contains hundreds of Thrush T. nigriceps.) Moreover, it is also then more likely that other, as yet
original reworkings of long-established taxonomic arrangements, and untested parameters will be found to show a difference. (The vocalizations
hundreds more evaluations of recent work by others in the field of mod- of T. subalaris add a further score of 6 in their divergence from those of
ern avian systematics, all using a single simple system to quantify levels T. nigriceps.) The system of taxonomic evaluation that we use, the Tobias
of phenotypic differentiation while also being guided (and sometimes criteria (Tobias et al. 2010, del Hoyo & Collar 2014: 30–41), allows for the
goaded!) by the findings of myriad molecular studies. By this means, quantification of all the variables listed above, with the exception of genet-
although emphatically not pretending to provide a definitive solution to ics. The inability of the criteria to accommodate genetic evidence within
the seemingly intractable problem of species limits in birds, we hope to the current scoring system is, as explained at length in the Introduction
have achieved at least a moderate degree of consistency, even-handedness to Volume 1, a function of the inability of molecular science to establish
and transparency of treatment that will help to serve the contemporary a fixed level of genetic differentiation to serve as the species/subspecies
interests and needs of science, conservation and recreational experience boundary. However, the fact that genetics is not (or not yet) integrated
in equal and equitable measure. into these criteria does not mean it is not integral to the evaluation pro-
We say “contemporary” with no very clear idea of the time-span this cess. On the contrary, the results and insights of recent molecular studies
word might encompass. In the six-and-a-half years from the initiation saturate this checklist: their evidence is constantly accounted for and
of our project in late February 2010 to 31 August 2016, our last date for incorporated. Indeed, the single greatest driver of our use of the Tobias
the consideration of new published evidence, at least 1,242 papers and criteria has been the findings of DNA studies.
articles appeared on the taxonomy of the passerines alone, a rate—0.5 So we take the view that the Tobias criteria have established the basis
per day—surely unprecedented in the history of avian systematics. (Simply for a process of taxonomic revision which, while not fitting every circum-
keeping track of this material is a discipline in itself; assessing its import stance and never claiming to be fault-free, is at once integrative, com-
and incorporating its insights into a global framework is something else prehensive, practical, robust, rapid, open and consistent. Consistency in
again.) It is not yet clear whether this extraordinary surge of information particular is a vital aspiration—too easily lost sight of—in a global check-
might be waning, although at some stage it surely must. Meanwhile, new list. If species limits in allopatric taxa are, as we are commonly reminded,
connections, new interpretations, new insights—inflections of the verb arbitrary constructs for human convenience, the Tobias criteria at least
“reveal” are ubiquitous in the titles of modern molecular papers on sys- serve as one way to standardise arbitrariness. Whatever the judgement of the
tematics—will constantly require new adjustments to be made to the world system or the reception of its results—and published criticism (Remsen
list of birds. Checklists are part of an enduring taxonomic cycle. What you 2015) has indicated potential areas for improvement but identified no
have on your lap or table as you read these words is as much a beginning irremediable weaknesses (Collar et al. 2016, Remsen 2016)—nothing is
as an end. hidden from plain view: throughout this checklist the evidence we as-
Even so, the completion of this checklist marks the fulfilment of a spe- semble in support of our decisions is laid out for scrutiny, contemplation,
cific ambition, fuelled over two decades with each new volume of HBW rejection, acceptance and/or adjustment. “Improved standardization and
(and indeed over three-and-a-half decades of modern Red List assess- transparency of species taxonomy”, Gill (2014) creditably asserted, “is
ments by BirdLife), to make a serious attempt at addressing the mani- essential”. This is precisely what this checklist, autonomous in its explora-
fest, manifold inconsistencies, inequalities and inadequacies that have tions of fact and argument, and holistic in its use and open in its presenta-
so long beset world lists of birds, and to establish a firm and unbiased tion of evidence, is attempting to provide.
foundation for the conservation of bird species across the planet. So
many taxonomic notes in HBW contain comments about one subspecies
or another that merits consideration for species rank; and so many of Taxonomic democracy:
its plates and descriptions point to the potential and need for a fresh in- taking account of all characters
vestigation of taxonomic status. Within the all-too-narrow constraints of
time and resources we have sought to address as many of these issues as
possible (although some, as we acknowledge further below, have proved Until relatively recently, morphology, internal as well as external, was the
beyond reach, but already with encouraging results from Volume 1, as only means of assessing a taxon’s relationships and distinctiveness. Now,
with the rediscovery of the Blue-bearded Helmetcrest Oxypogon cyanolae­ however, two parameters have come to be regarded as fundamental to the
mus: Rojas & Vasquez 2015). Of course the extensive body of recent work robustness of taxonomic revisions: voice and DNA. These two variables
in avian systematics, running in parallel with our venture, has played by have achieved a primacy of value which is, of course, entirely understand-
far the greater part; but through the combination of these endeavours able: songs and calls are a principal way, along with outward appearance,
we do not anticipate another undertaking on such a scale, at least for by which bird species identify themselves, while DNA provides the best
the foreseeable future. evidence of a population’s internal coherence and composition. Even

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 16 Introduction

so, the tendency to consider these parameters decisive to the point of judge that the molecular evidence must for the present prevail, include
veto—thereby rendering some characters “more equal than others”—is Olivaceous Mourner Schiffornis olivacea split from Northern Mourner
not wholly welcome or wise: voices, especially among the oscine passer- S. veraepacis, Winter Wren Troglodytes hiemalis and Pacific Wren T. pacifi­
ines, vary with circumstance, and DNA has some intriguing, instructive cus from Northern Wren T. troglodytes, Southern Variable Pitohui Pitohui
mysteries, as when two clear species—or as many as five, in the case of the uropygialis from Northern Variable Pitohui P. kirhocephalus and Highland
Tawny-bellied Seedeater Sporophila hypoxantha, Dark-throated Seedeater Rush-warbler Bradypterus centralis from Little Rush-warbler B. baboecala.
S. ruficollis, Pearly-bellied Seedeater S. pileata, Marsh Seedeater S. palustris Perhaps the most baffling issue we have encountered in preparing this
and Black-bellied Seedeater S. melanogaster (Benites et al. 2014, Campagna second volume concerns the identity and rank of a cluster of Pomatorhinus
et al. 2015)—prove genetically inseparable while other species possess scimitar-babblers (White-browed P. schisticeps, Chestnut-backed P. montanus
startling genetic divides that have no effect on their behaviour or status and Streak-breasted P. ruficollis), for which the genetic and morphological
(see, e.g., del Hoyo & Collar 2014: 28–29). Comments by certain molecu- data are so wholly at variance that we feel obliged to stick with the status
lar taxonomists—“genetic estimates of lineage divergence may not be… quo until renewed—and integrative—analyses can shed stronger light on
reliable indicators of species limits” (Humphries & Winker 2011); “levels an extraordinarily convoluted circumstance.
of sequence divergence alone should not be used for describing species” Overall, however, we see the frequent complementarity of phenotypic
(Schultz & Burns 2013)—reveal an awareness of this limitation. Indeed, and genetic distinctiveness as evidence that the Tobias criteria are well
the very techniques for producing evidence of molecular structure are rooted in established standards of taxonomic discrimination. This view
still open for debate, as in the case of the California Gnatcatcher Polioptila seems to be slowly gaining traction with practitioners of molecular studies,
californica (Zink et al. 2013, 2016, McCormack & Maley 2015, Anon. 2016). albeit with reservations and conditions: Lifjeld et al. (2016) give a score
The Tobias criteria spread the power to drive taxonomic change fairly of 11 to Gran Canaria Blue Chaffinch Fringilla polatzeki, while Campbell
evenly across all phenotypic characters (plumage, size, voice), allowing et al. (2016) “also consider a phenotypic score of 7, conservatively, to be a
them to stand alone or to operate in (sometimes unexpected) combina- conceptual threshold for identifying what we term phenotypically highly
tion to achieve species status for their bearers. A case in point is the Short- divergent taxa”. Such confluence and complementarity are encourag-
tailed Antbird Poliocrania maculifer, which we split here from Chestnut- ing. Molecular taxonomy thus need not be characterized either as a
backed Antbird P. exsul on the evidence of four colour characters and a Cinderella, left out of consideration by a self-serving (and some might say
markedly shorter tail yet with seemingly no difference in voice, despite self-sealing) system of evaluation, or as a Cyclops, dominating the modern
song being the almost exclusive means by which changes to thamnophilid taxonomic landscape with its unsparing, all-seeing eye. The combina-
taxonomy have been effected over the past two decades. Equally, cued tion of phenotypic and genetic analysis is clearly key to advances in avian
by the cautionary comments above about the acceptability of genetic systematics for the foreseeable future.
evidence on its own or in large measure, the Tobias criteria have the
capacity to reverse splits such as that of Highland Hepatic-tanager Piranga
lutea from what we now simply call Hepatic Tanager P. hepatica, which was Representative sampling: the limits of confidence
made in HBW on molecular grounds, given the very slight morphological
and vocal differences these two taxa show. The system thus possesses the Nevertheless, both phenotypic and genetic analysis depend on the
flexibility to liberate assessments, with no loss of analytical rigour, from the number and distribution of the samples involved, and this issue crops up
prevailing (if often seemingly unconscious) orthodoxy that places voice sufficiently often to be worth a brief commentary here. Statistically robust
and DNA on a higher tier of taxonomic relevance. evidence of a consistent morphometric disjunction demands an adequate
In many cases the Tobias criteria may even constitute a useful inde- sample size. The magnitude of this depends on the scale of natural vari-
pendent test of the results of genetic analyses. If, as we suggest above, the ation for each case, but we used a default minimum of 10 specimens of
number of variables that show differences predicts differences in other one sex for each test (Tobias et al. 2010), and our pursuit of this sample
as-yet unmeasured variables (albeit not in the voice of the Short-tailed size has accounted for many months of work in various museums. (Here
Antbird), it is plausible to assume that multiple phenotypic dissimilarities we should just repeat and refine information given in the Introduction
predict significant genetic distinctiveness. In this regard, an encouraging to Volume 1: when an effect size is given with no indication of sample
finding in this volume is the rather high degree of concordance between size or sex, this means that the sample was at least 10 and the sex of the
phenotypic and genetic evidence. Among the non-passerines there were sample was male.) The same requirements are of course highly desirable
several cases where simultaneous molecular and morphological analyses for the confident discrimination of constant vocal characters and when
converged on the same answers (e.g. Hainan Peacock-pheasant Polyplectron considering plumage differences in closely similar taxa. Inevitably there
katsumatae, Black-browed Barbet Psilopogon oorti), and more recently our are shortfalls: many taxa are known from fewer than ten specimens, while
split of Lesser Sri Lanka Flameback Dinopium psarodes has been supported many that are known by more have large proportions of them unsexed or
by genetic evidence (Fernando et al. 2016). Among the passerines the list scattered uneconomically among several distant museums. Similar prob-
of species where genetic and phenotypic evidence coincides is notable: lems attend the recordings of vocalizations.
examples are to be found in virtually every family consisting of more The uneven spatial sampling of taxa is perhaps a less recognized prob-
than just a few species. We particularly acknowledge a good number of lem. When evidence of clinality, involving voice, plumage and genetics,
cases which, owing to the similarity of appearance of the taxa in question, was first documented in a suboscine antbird, the Variable Antshrike Tham­
were not ones we originally selected for further study but which, after nophilus caerulescens (Isler et al. 2005), Brumfield (2005) was prompted to
being identified by genetic evidence, have proved also to meet the Tobias stress the cardinal importance of sampling over a wide area:
criteria, often because of the role played by voice: for example, Campina A more restricted sampling design in the present study might have
Thrush Turdus arthuri and Floodplain Thrush T. debilis from Black-billed led to the erroneous conclusion that T. c. aspersiventer, T. c. dinellii, and
Thrush T. ignobilis, Pacific Cacique Cacicus pacificus from Scarlet-rumped T. c. paraguayensis have reciprocally monophyletic mitochondrial lineages,
Cacique C. microrhynchus, and Amazonian Grosbeak Cyanoloxia rothschildii making them full species according to some species concepts.
from Blue-black Grosbeak C. cyanoides. The warning continues to be missed, however. The elevation to species
This is not to pretend that phenotypic and genetic evidence is not status of two clades in the Australian Raven Corvus coronoides (Jønsson
sometimes in conflict. Rarely—or at least we hope this is so, but ­inevitably et al. 2012) was declined by Dickinson & Christidis (2014) for lack of
detection rates are low—the confusion is attributable to clerical error sampling intergradient material, a position we fully support ­irrespective
(mislabelled samples, misattributed specimens): we know of several of our agreement that levels of sequence divergence alone are not ap-
unpublished cases that have led to published mistakes, and others have propriate arbiters of species limits. We likewise resist a claim of species
been openly acknowledged (Zou et al. 2008) or suspected (Davies & rank for northern Venezuela’s Green-and-black Fruiteater Pipreola riefferii
Peacock 2014). Meanwhile, the commonest problems involve paraphyly melanolaema on the basis of its 4.6% sequence divergence from subspe-
and polyphyly, and can often only be dealt with by calls for more re- cies confusa in distant Peru (Berv & Prum 2014), because south-western
search and denser sampling—a case in point is the seemingly intractable Venezuelan (i.e. geographically intermediate) specimens of nominate
conundrum of the Great Grey Shrike Lanius excubitor complex, where the riefferii not only remain genetically unsampled but also “clearly present
most recent molecular studies (Olsson et al. 2010) overturn some previ- plumage transitional to P. r. melanolaema” (Kirwan & Green 2012). We
ous notions of relationship but also fail to provide an arrangement that do the same with the Chinese form mutica of Scaly-breasted Cupwing
can be reconciled with any species concept or with complete diagnos- Pnoepyga albiventer, catapulted from synonym to species on the basis of “so
ability. Other such cases where the Tobias criteria fail to vindicate claims pronounced” differences in molecular and bioacoustic markers compared
based on the molecular evidence, but where circumstances compel us to to those in Nepal (Päckert et al. 2013), because our own sampling of songs

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 Introduction 17

from geographically intermediate areas (Bhutan, north-east India) reveals starlings, thrushes, warblers—are highly imitative. As a consequence of all
a cline that reduces mutica to subspecific status. Certain recent proposed these things, a great deal of effort can be spent on analyses that ultimately
revisions by Pratt & Mittermeier (2016), who remark (without discernible fail to be convincingly robust.
supporting evidence) that the Tobias criteria “in many cases fail to include These drawbacks were borne in mind during the process of evaluat-
relevant behavioral and ecological data”, fail to achieve an appropriate ing vocal differences for use in this volume. Not all cases produced an
level of comprehensiveness, with Samoan Robin Petroica pusilla (nominate outcome, owing to the scarcity or quality of the material. Occasionally we
of a fourteen-race species) split on a vocal comparison with Fijian birds are able to point to previously unreported levels of distinctiveness which
only, and Manu’a Starling Aplonis manuae given species rank on the basis are highly suggestive but which require much closer investigation (e.g.
of vocal evidence which our own review of all available sound recordings Short-tailed Antthrush Chamaeza campanisona, Common Miner Geositta
simply does not confirm. cunicularia, Wedge-billed Woodcreeper Glyphorynchus spirurus, White-eyed
But we have no intention of disguising or diminishing the fact that Tody-tyrant Hemitriccus zosterops, Mountain Elaenia Elaenia frantzii). How-
a significant proportion of the taxonomic judgements we make here is ever, in many dozens of cases the scoring for voice was decisive in elevating
based on small sample sizes and assumptions. A striking case by way of taxa to species rank. The findings are of necessity highly condensed in the
illustration is that of the Slender-billed Cicadabird Edolisoma tenuirostre, a taxonomic notes sections where phenotypic characters are enumerated
single species in HBW which in this checklist we break into eleven. This and scored, but in the interests of transparency the full texts are to be
is based on as many specimens as we could access but which commonly published on the HBW Alive website very shortly.
did not reach the double figures we require; and in many cases also on
just one or two sound recordings. In such circumstances, where morphol-
ogy and voice are combining to point, with considerable strength, to a Our null hypothesis
change in taxonomic ranking, the choice then arises whether to allow the
weakness of the sample size to override the assumption that the material In an elegant, erudite and impassioned essay that appeared as Volume 1
we are reviewing is representative. Our inclination has almost always been was in press, Gill (2014) strove to make the case for a “reverse list” taxon­
to opt for what we hope is a very small risk in accepting the evidence. omy of birds, arguing that the new null hypothesis in a polytypic species
However, wherever sample sizes have been too small to make a score with concept should be that: “distinct and reciprocally monophyletic sister
confidence we insert the word “allow” before the (typically in these cases populations of birds exhibit essential reproductive isolation and would
conservatively) estimated number to acknowledge the increased degree of not inter­breed freely if they were to occur in sympatry”. By this formula-
uncertainty in the overall assessment. tion Gill effectively presses the basic premises of the phylogenetic species
concept (diagnosability and reciprocal monophyly) into the service of
the biological species concept. Rather than treating isolated taxa on
Quantifying voice: the new taxonomic dimension mountains and islands as conspecific with each other or with continental
taxa, as under current arrangements, the working assumption would be
The vocalizations of birds are often, as we note above, of crucial impor- that, unless and until shown otherwise, such taxa are presumed to be
tance in determining taxonomic status. Among the non-passerines evalu- reciprocally monophyletic and therefore each a species. In a reversal of
ated for species rank in Volume 1 of this checklist, at least 34 were given the decades-long tradition in which ornithologists of various backgrounds
scores under the Tobias criteria for “different voice”, comprising seven have slowly and laboriously, species by species, unpicked the overlumped
pigeons, seven owls, five woodpeckers, two guineafowl, two cuckoos, two global avifauna, the burden of proof would fall on the shoulders of those
kingfishers, two barbets and one penguin, snipe, rail, hornbill, trogon, who would keep particular taxa lumped, not on those who want to see
motmot and parrot. The evidence for these differences was in the pub- them split.
lished literature, and the sources were cited; the scores themselves were That this proposition has gained some traction is apparent in a recent
based on a subjective assessment of the degree of distinctiveness conveyed molecular study of the Rufous-tailed Tailorbird Orthotomus sericeus. Lim et
by the descriptions. al. (2014) reasonably interpreted Gill’s species-rank hypothesis to apply
Not all these taxa needed the awarded scores to achieve species rank, to all “physically separated, moderately divergent populations” of birds,
but the majority did. While we remain reasonably comfortable with and ­argued that the taxonomically undifferentiated population of the
the outcomes in these cases, it is obviously far from satisfactory in such tailorbird on Palawan (Philippines) is in line for elevation because of its
important matters to depend so completely on the statements of others, clear but moderate genetic distinctiveness from populations on Borneo:
which are themselves liable to subjective bias. This danger was underlined “If diagnosable differences in song or behavior are discovered…, it should
during work on the present volume when evaluating the distinctiveness of then be named a different species”.
the race longicaudata, from Anjouan, of Madagascar Brush-warbler Nesillas This exercise of Gill’s argument exposes its rather deep implications.
typica. This form was split in a recent field guide (Sinclair & Langrand How many “physically separated, moderately divergent populations” of
2013) with the remark that its notes are more rapid and higher-pitched birds are there in the world? In this case the moderate divergence is evident
than those of the nominate, and in consequence N. longicaudata has at the genetic level, so all that is needed is an additional phenotypic differ-
been accepted by at least two world checklists. However, our own analysis ence, no matter how small, to take Palawan birds from population to spe-
of the voice of this form, using publicly available recordings, reveals no cies in one bound. With sufficient sampling this may well happen (although
quantitative differences. We therefore retain longicaudata as a subspecies our own comparisons have revealed nothing clear), as differences in voice
of N. typica. A similar case is that of the Golden-bellied Gerygone Gerygone in insular forms, at least of oscine passerines, are commonly not difficult
sulphurea, where supposed differences in voice between populations prove to find, may simply relate to habitat, and can evolve at great speed (Baker
to be no stronger than those within populations. 2006, Baker et al. 2006). Thus if the criteria for bestowing species rank
This analysis was just one product of an initiative never previously at- can apply to any populations exhibiting physical separation and modest
tempted in a world checklist, to incorporate quantitative analysis of vocal­ divergence in voice, even recently translocated North Island Saddlebacks
izations as a central component of the taxonomic decision-making pro- Philesturnus rufusater appear to be on a path to eligibility (Parker et al. 2010).
cess. Resources—including time—did not permit a truly comprehensive Gill does, however, specify reciprocal monophyly, which (a) ought to
review of recorded material, but the decisive importance of vocalizations prevent any absurdities resulting from voice shifts on newly colonized
in the species-specific signals of almost all the passerines, coupled with islands but (b) appears to lock non-molecular taxonomy out of all future
our continuing adherence to the Tobias criteria (for acoustic para­meters consideration. Without genetic study, how will reciprocal monophyly or its
see Tobias et al. 2010: 734–735), required us at least to investigate as many absence ever be demonstrated? What criteria will govern the assumptions
cases as possible—more than eight hundred—where vocal evidence behind the new ranking of taxa as species? Or will the burden of proof
seemed likely to prove indispensable. simply require evidence of the absence of reciprocal monophyly? (Anyway,
Quantifying vocal differences in birds is, of course, challenging. Vocal­ is there anything close to consensus that reciprocal monophyly is the new
izations can vary within and between individuals, ages and sexes, and yardstick for implementing species-rank revisions? The five “capuchino”
with time of day, time of year, circumstance and—manifestly—location. seedeaters mentioned above all behave as species by recognising each
Sample size (see above!) is therefore a major issue. Moreover, determin- other by song and plumage, but do not exhibit reciprocal monophyly.)
ing homologous calls for comparison can be problematic. Quality of A further question arises of how to judge the situation—what crite-
recordings matters. Recordists’ competence in field identification matters ria to use—when the burden of proof is duly shouldered and evidence
more. Perhaps most taxing are the facts that oscine passerines learn their ­furnished to challenge the results of adopting the new null hypothesis.
songs, and that many species and families—drongos, larks, shrikes, chats, Ire­stedt et al. (2013) split Red-bellied Pitta Erythropitta erythrogaster into

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17 species. Applying the Tobias criteria to the same complex, Collar et al. distinctiveness, whether as species or subspecies, will cause them to be
(2016) recognized 13. So by what standards are the rankings of the four ­accommodated into biodiversity conservation plans long into the future.
taxa that the latter authors retained as subspecies or synonymized to be In terms of the future of this endeavour after the publication of the
tested? It immediately becomes apparent that, rather than resolving this present volume, two points are worth conveying. First, the HBW/BirdLife
issue, the new null hypothesis has merely pushed it temporarily out of team will continue to review newly published material to produce at least
sight. But now, faced with evidence used to reject species rank for a taxon, annual updates to the Checklist in the future, which will be reflected in
a choice has to be made once again based on preference and belief. HBW Alive, in the BirdLife Data Zone and on the IUCN Red List. Anyone
Ultimately, therefore, the studies and debates generated by the adoption can contribute to this process by highlighting new information we may
of the new null hypothesis will be no shorter or less complex—or cost have overlooked, or taxonomic decisions that may require review, via the
less in terms of human resources—than those in play at the moment, and Comments boxes on the species accounts in HBW Alive. While we can-
they will still require taxonomists to choose between criteria for species not undertake to respond to individual comments, we will consider each
discrimination. We salute Gill’s intention with this initiative to seek to one carefully. Second, we hope to undertake further research to inform
establish a broader platform for the conservation of avian diversity, but we the Tobias criteria themselves, potentially exploring: expanding the
suspect that in reality it will serve no practical purpose. justification for the threshold score of 7 for species status, the feasibility
and impact of evaluating hybrid-zone width as a proportion of the entire
distribution width; broader issues relating to how to treat hybrid zones in
“A ‘rapid assessment program’ for assigning the criteria; differences between bird families in the challenges they pose
species rank?” Yes! for the application of the criteria; and the feasibility of integrating genetic
divergence into the criteria.

Remsen’s (2016) sympathy for the need for greater speed and efficacy in
evaluating species limits issues in birds gave him a good title, structured Technical aspects
as a question, and we readily answer him in the affirmative. We deem
it plausible that, in time, molecular studies might be able to resolve all However, in addition to reconsidering the taxonomic status of birds across
taxonomic issues, including at the species and subspecies levels. But that the whole class, this checklist offers a great deal more. For well over two
time is not yet, and it seems unlikely to come for many years. Meanwhile years our extended teams at both HBW/Lynx and BirdLife have been
we have to consider our options. fully occupied working on the present volume.
If Gill’s new null hypothesis is a non-starter for reasons given above, Given the many and sweeping changes to avian taxonomy to be found
the status quo he seeks to subvert is not acceptable either: it is, regrettably, in this volume, we have added extensively to the taxonomic texts imported
too little, too late. Despite the spreading compass of molecular studies, from HBW but have also extensively used explanatory notes to accompany
great swathes of taxa remain unsampled, part-sampled or in need of re- genus and family subheadings. These typically concentrate on indicating
sampling. Checklist committees serve an important purpose, but function how and why the group in question differs in treatment from what was
reactively, appear to abide by majority votes that are sometimes cast before presented in HBW.
all the arguments are heard and weighed, and understandably struggle Recovering and embellishing one of the features used throughout
with the issue of urgency (as at August 2016—this observation is neutral HBW, in the present volume we include summary totals for each taxon
and in no way a criticism—the South American Checklist Committee web- from genus upwards. Thus, a genus will have two figures in blue, which
site indicated 252 cases of “Proposal needed” and 72 cases of “Proposal represent species (in bold) and “taxa” (sensu HBW = total number of
badly needed”). Checklist compilers, meanwhile, rely on the published species-group taxa recognized, in other words all subspecies plus all
findings of others, in some cases often with little or no discrimination over monotypic species). Moving into family-group names, the numbers
the quality involved (although they make occasional unexplained choices continue to expand leftwards, with the inclusion of genera, and (when
between competing viewpoints, resulting in disconcerting levels of avoid- applicable, further left still) tribes and subfamilies. So we might have a
able contradiction and inconsistency). family with the figures 3sf·27·273·616, which would mean it contained three
All this underscores the necessity of our current exercise. We fully subfamilies, 27 genera, 273 species and 616 “taxa”. As in HBW, these fig-
acknowledge its multiple imperfections, and we have no doubt made er- ures always refer to extant forms. When extinct forms are also involved,
rors that will trouble our sleep for years to come. We admit, yet again, that the numbers of extinct taxa are displayed in a like fashion but in a sepa-
the Tobias criteria are open to refinement and improvement, perhaps on rate series of grey numbers, situated further to the right. These “extinct”
a scale and of a type which will reverse some of the taxonomic decisions figures continue leftwards until reaching the first “0”. Thus, if the above
they have produced to date. And we need to emphasise that constraints of family contained merely one extinct subspecies, the extinct figures would
time and resources have prevented us from being able to apply the Tobias be †0·1; if it were one species and three subspecies, it would appear as
criteria to every case, especially when the issue is one of lumping rather †0·1·3. The complete family entry in this case would be: 3sf·27·273·616 †0·1·3.
than splitting. Many taxa in the Troglodytidae merit evaluation in this re- Note once again that the extinct taxa are not included within the blue
gard, and a good number of cases are scattered throughout other families, figures for extant forms, so the total number of “recent” taxa in this fam-
among them (merely by way of illustration) White-tailed Swallow Hirundo ily would be 619.
megaensis and Pearl-breasted Swallow H. dimidiata, Hildebrandt’s Starling Our resident team of artists has been busy, and this volume contains a
Lamprotornis hildebrandti and Shelley’s Starling L. shelleyi, Japanese White- grand total of 12,629 bird illustrations, including 642 that are totally new,
eye Zosterops japonicus and Lowland White-eye Z. meyeni, Adelaide’s Warbler and a further 1,208 that have been improved from their original HBW
Setophaga adelaidae, Barbuda Warbler S. subita and St Lucia Warbler S. deli­ versions. A few figures published in HBW have been omitted as being not
cata, Scarlet-rumped Tanager Ramphocelus passerinii and Cherrie’s Tanager sufficiently relevant or in a few cases misleadingly different.
R. costaricensis, and Black-capped Hemispingus Kleinothraupis atropileus and Making the maps as accurate as possible involves extensive delving into
White-browed Hemispingus K. auricularis. an immense variety of sources, often to make very small changes that
Nevertheless, this volume, like its predecessor, makes the case for signifi- perhaps only a few will detect, and often struggling to make the best of
cant numbers of species-level changes, and we respectfully await the verdict what little is available to map some of the less well documented areas.
of history over their adoption. HBW treated 3,964 species of non-passerine Altogether 6,649 maps have been checked, rechecked and checked again
and 6,008 species of passerine; these figures rise in the present work to by our teams at HBW/Lynx and BirdLife to provide what we would like
4,372 and 6,592 respectively; thus the number of bird species in the world to hope are probably the most accurate set of maps for any complete class
rises in this exercise from 9,972 to 10,964, an increase of 11%. Somewhat of animals. A minor change on the maps this time is that arrows used to
over half this growth has been generated by the work of others; the remain- highlight small details of distribution now appear in red; this change has
der derives from our own endeavours after thousands of hours in museums been made for purely practical reasons, as the black arrows previously used
and hundreds of hours analysing recordings of vocalizations and studying sometimes proved difficult to see against coastal outlines. Also on the sub-
photographs and videos. We particularly hope that these “original” revi- ject of distributions, we ought perhaps to explain that we have attempted
sions will be subject to independent scrutiny, assessment and approval; in to respect local usage (as we understand it) in the positioning of the “R”
many cases fieldworkers may be motivated to collect the further evidence (for River) before or after the actual name so it normally comes before the
that will validate or refute them. But we feel confident, at least, that the name in most of the world (R Xingu, R Indigirka, R Zambezi, R Danube),
emphasis in this checklist on phenotypic differentiation has identified but after it in North America, Australia and New Guinea (Mississippi R,
numerous instances of previously neglected taxa whose newly recognized Murrumbidgee R, Fly R).

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 Introduction 19

We have tried to be as informative as readability permits in backing up 2 began. For this privilege we particularly thank Robert Prŷs-Jones, whose
all new material or decisions, be they taxonomic, distributional or what- very positive support has been of the greatest value (and which we attempt
ever, with extensive reference to the literature we have been consulting, to acknowledge below: see page 21), Mark Adams, Hein van Grouw,
and this amounts to some 2,809 references. A great many more papers Alison Harding and other staff members, including those in security who
and books have been referred to, but in a work of this scope, regrettably it cheerfully supported our comings and goings at weekends and early
is not practical to include them all. and late in the working day. The American Museum of Natural History
The extensive internal rearrangements of numerous families in this (AMNH), New York, USA, has also allowed us to work there over several
volume have frequently had knock-on effects on the scientific names to be weeks in the past three years, and we gratefully acknowledge the kind help
used. Some of the issues are explained below, along with a few other mat- of Paul Sweet, Thomas Trombone, Lydia Garetano, Mary LeCroy and Joel
ters related to nomenclature (see page 21). Cracraft. Two other museums have hosted us several times for several days:
The English names of species have also come under new pressure as a the Smithsonian Institution (United States National Museum, USNM),
consequence of the unprecedented levels of change in the composition Washington DC, USA, where we thank Chris Milensky, Brian Schmidt and
of certain passerine families or groups, particularly in the New World (for Christina Gebhart, and the Naturkundemuseum (ZMB), Berlin, Germany,
example Fringillidae, Emberizidae, Cardinalidae and Thraupidae). Sum- where we thank Sylke Frahnert and Pascal Eckhoff. Other museums in
mer and Scarlet Tanagers turn out to be cardinals. The Paroaria cardinals which we undertook work on the project are Louisiana State University
prove to be tanagers. The Coal-crested Finch is a tanager. So are the two Museum of Natural Science (LSUMNS), Baton Rouge, USA, with thanks
“diuca-finches”, but they occupy different clades and arguably therefore to Fred Sheldon and Van Remsen; Naturalis (RMNH), Leiden, Nether-
should no longer be linked by a common name. The list of such misfits lands, with thanks to Steven van der Mije and Pepijn Kamminga; Muséum
is fairly long and, assuming that these new arrangements will not be over- National d’Histoire Naturelle, Paris, France (Patrick Boussès and Jérôme
turned by a further round of genetic analysis, it is obviously tempting to Fuchs); Royal Museum for Central Africa (RMCA), Tervuren, Belgium
consider changing the English names of species to reflect their true iden- (Alain Reygel and Michel Louette); and Royal Belgian Institute of Natural
tities. However, this is a sensitive issue and many birders and ornithologists Sciences (RBINS), Brussels, Belgium(Tom Geerinckx, René-Marie Lafon-
seem likely to prefer stability over accuracy, or at least to favour a more taine and Pierre Devillers). We also had help from John Bates at the Field
measured and consensual transition from one usage to another. It is also Museum of Natural History, Chicago, USA, and Markus Unsöld at the
potentially rather a complex issue: changing “White-winged Diuca-finch” Zoologische Staatssammlung München, Munich, Germany.
to “White-winged Tanager” immediately runs into trouble with Piranga The starting point for the texts of course was HBW, but the first stop
leucoptera, itself now a cardinal! Where opportunity arises and opposition when rechecking basic details for each species account and looking for
seems unlikely, we have made a very small number of changes (the Coal- potential conflicts was Dickinson & Christidis (2014) Howard and Moore
crested Finch becomes Coalcrest, for example); but otherwise we deem Checklist, 4th Edition, which rediscusses many of the issues of nomencla-
it more appropriate to assume a preference for a longer-term process of ture already tackled in HBW, as well as many new ones, and has made
adjustment by the global ornithological community. many advances in the field of establishing correct publication dates. Also
In order to avoid using up a great deal of space on what is a fairly minor most useful have been certain websites, notably Zoonomen (Peterson 2016),
feature, we have limited the “Other common names” to those used in Taxonomy in Flux (Boyd 2016) and the AOU’s South American Classifi­
recent world and regional checklists, and we do not list any that follow on cation Committee (Remsen et al. 2016), along with the world lists by
fairly obviously from what is already listed. ­Clements et al. (2016) and Gill & Donsker (2016).
As in Volume 1, we follow BirdLife International in indicating which We also offer a particularly hefty thank you with a warm recommen-
Critically Endangered species are likely to have gone extinct (or extinct in dation to Winkler et al. (2015) Bird Families of the World. This book has
the wild), but for which comprehensive searches are required to confirm provided the entire framework of families for our checklist, and is an
that the last individual has died. The application of the Possibly Extinct excellent source for gaining an idea of how the higher taxonomy of birds
and Possibly Extinct in the Wild tags to such taxa takes into account the has evolved in recent years.
time since the last record, the intensity, extensiveness and adequacy of We are most grateful to Normand David and Michel Gosselin for having
searches, the extent, intensity and timing of threats, and the likely suscep- checked through the entire list of scientific species-group names, and for
tibility of the taxon to these threats (Butchart et al. 2006, IUCN Standards having pointed out numerous errors in the drafts. For help and advice on
and Petitions Committee 2016). We have also continued the approach issues of taxonomy and nomenclature, we are particularly grateful to Per
in Volume 1 of identifying Extinct (“†”) avian subspecies, updated from Alström, Normand David, Edward Dickinson, Jon Fjeldså, Vítor Piacen-
Szabo et al. (2012). It should be noted that the species-level assessments in tini, Frank Steinheimer and Hans Winkler, as well as Walter Boles, Bob
particular need updating, but this has been postponed until publication Dowsett, John Fitzpatrick, Dick Schodde and Tom Schulenberg.
of an important new series of papers that describe and apply a new quan- We also wish to pay tribute to the website Xeno-canto, which has been
titative approach to assessing such information within a statistical frame- an extraordinary resource for our research into vocalizations, although
work. Once published, there are likely to be a number of changes to the we should also mention, with deep appreciation, AVoCet, the Macaulay
list of Extinct and Critically Endangered (Possibly Extinct) species. The Library at the Cornell Laboratory of Ornithology and Lynx’s own Internet
Red List categories assigned to all species in this volume—including all Bird Collection (IBC). Furthermore, the IBC, supplemented by a large
the taxonomic revisions—are those published in BirdLife’s 2016 update to number of other websites of different kinds, has been the main source of
the IUCN Red List. videos and photographs of live birds, which in this checklist have played a
For further information about the structure and technical workings of very important role for analysing morphological characteristics, comple-
the current checklist, we would point readers towards the Introduction to menting the traditional study of museum skins. Peter Boesman’s extensive
Volume 1 (pages 43–51). revision of vocalizations is reflected in there being a separate reference,
393 in total, for each different species (when appropriate). This has proved
essential, because he has amassed an enormous amount of m ­ aterial, all of
Acknowledgements which will be soon available as HBW Alive Ornithological Notes.
The teams that provide us with the French, German and Spanish names
Volume 1 of this checklist contains long lists of people and institutions to have had a busier time than ever, as they have had to revise all of the passer-
whom and to which we owe an enormous debt of gratitude, and we should ine names to fit with the new taxonomic ideas. We should like to acknowl-
at once stress that many of the contributions from those people and institu- edge our warm appreciation of Normand David and Michel Gosselin for the
tions have been to the entire project, not merely the first v­ olume. We hope French names; the “Standing Committee for German Names of the Birds of
that we may be forgiven for not repeating all these names here, where in- the World” of the Deutsche Ornithologen-Gesellschaft and IOU, consisting
stead we simply focus on those who have been particularly important to the of Peter H. Barthel, Christine Barthel, Einhard Bezzel, Pascal Eckhoff, Re-
second volume and to the completion of the project. We should, however, nate van den Elzen, Christoph Hinkelmann and Frank Steinheimer, for the
also acknowledge the immense contribution made by the authors, artists German names; and the committee of the Spanish Ornithological Society
and institutions involved with the production of the seventeen volumes of (SEO/BirdLife), comprising Eduardo de Juana, Josep del Hoyo, Manuel
HBW, the foundation on which this checklist builds. To all of them we take Fernández-Cruz, Xavier Ferrer and Jordi Sargatal, for the Spanish names.
this opportunity to register our sincere appreciation of their work. At BirdLife International we thank Gill Bunting, John Cornell, Mike
The Natural History Museum (NHMUK), Tring, UK, has been funda- Crosby, Mike Evans, Melanie Heath, Adrian Long, Ian May, Roger Safford,
mental to the fulfilment of this project, hosting us for several thousand Lizzie Sparrow and David Wege for their various contributions to the pro-
hours of research time over the two-and-a-half years since work on Volume ject. We also thank all those who posted comments on the BirdLife forums.

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 20 Introduction

Davison, G.W.H., Chang, J., Zhang, Z. & Chen, D. (2012). Full tree resolution of Polyplectron,
We should also like to acknowledge the important help received from Temminck 1813, confirms species status of Hainan P. katsumatae Rothschild, 1906, and
Desmond Allen, Johnnier Andres Arango, Norbert Bahr, Bas van Balen, Bornean Peacock-pheasants P. schleiermacheri Brüggemann, 1877. Bull. Brit. Orn. Club 132(4):
Jessie Barry, Bruce Beehler, Carlos Calimares, Les Christidis, Oswaldo 251–259.
Cortés, Paul Donald, Guy Dutson, James Eaton, Stephen Garnett, Guy Dickinson, E.C. & Christidis, L. (2014). The Howard and Moore Complete Checklist of the Birds of the
World. 4th edition, Vol. 2. Passerines. Aves Press, Eastbourne, UK.
Gibbon, Phil Gregory, Stephen Gregory, Steve Hilty, Steve Howell, Rob Fernando, S.P., Irwin, D.E. & Seneviratne, S.S. (2016). Phenotypic and genetic analysis support
Hutchinson, Michael Irwin, Allen Jeyarajasingam, Ron Johnstone, Leo distinct species status of the Red-backed Woodpecker (Lesser Sri Lanka Flameback: Dinopium
Joseph, Steve Kelling, Niels Krabbe, Thomas Kuenzel, Marek Kuziemko, psarodes) of Sri Lanka. Auk 133(3): 497–511.
Frank Lambert, Paul Leader, Miguel Lentino, Simon Mahood, Petter Gill, F.B. (2014). Species taxonomy of birds: which null hypothesis? Auk 131(2): 150–161.
Gill, F. & Donsker, D. (2016). IOC World Bird List. Version 6.4. doi:10.14344/IOC.ML.6.4.
Marki, Ben Marks, Willy Marthy, Rob Martin, Jay McEntee, Matthew Med­ del Hoyo, J. & Collar, N.J. (2014). HBW and BirdLife International Illustrated Checklist of the Birds of
ler, Michael Mills, Nial Moores, Pete Morris, David Moyer, Urban Olsson, the World. Volume 1: Non-passerines. Lynx Edicions, Barcelona.
Chikara Otani, Bob Payne, Alan Peterson, Praveen J., Josep Puig, Rich Humphries, E.M. & Winker, K. (2011). Discord reigns among nuclear, mitochondrial and
Pyle, Pam Rasmussen, Van Remsen, Robin Restall, Frank Rheindt, Bob phenotypic estimates of divergence in nine lineages of trans-Beringian birds. Mol. Ecol. 20(3):
573–583.
Ridgely, Craig Robson, Stephen Rumsey, George Sangster, Nat Seddon, Irestedt, M., Fabre, P.H., Batalha-Filho, H., Jønsson, K.A., Roselaar, C.S., Sangster, G. & Eric-
Claire Spottiswoode, Lars Svensson, Paul Sweet, Joe Taylor, Rob Tizard, son, P.G.P. (2013). The spatio-temporal colonization across the Indo-Pacific by a ‘great spe-
Joe Tobias, Colin Trainor, Jeremiah Trimble, Ben Wielstra and Roland ciator’ (Aves, Erythropitta erythrogaster). Proc. Roy. Soc. London (Ser. B Biol. Sci.) 280: 20130309.
Wirth. We extend our warmest thanks to them all. Isler, M.L., Isler, P.R. & Brumfield, R.T. (2005). Clinal variation in vocalizations of an antbird
(Thamnophilidae) and implications for defining species limits. Auk 122(2): 433–444.
And last, but by no means least, NJC’s position at BirdLife is funded IUCN Standards and Petitions Subcommittee (2016). Guidelines for using the IUCN Red List
by the A. G. Leventis Foundation, and we extend our gratitude for this categories and criteria. Version 12. Prepared by the Standards and Petitions Subcommittee.
invaluable support to Anastasios P. Leventis; moreover, the Leventis Foun- URL: http://www.iucnredlist.org/documents/RedListGuidelines.pdf.
dation also covered the participation of Lincoln Fishpool in this venture, Jønsson, K.A., Fabre, P.H. & Irestedt, M. (2012). Brains, tools, innovation and biogeography in
crows and ravens. BMC Evol. Biol. 12: 72. doi: 10.1186/1471-2148-12-72.
and our gratitude is accordingly the greater. We also particularly thank for Kirwan, G.M. & Green, G. (2011). Cotingas and Manakins. Christopher Helm, London.
their great generosity Stephen Eccles, who provided support that helped Lifjeld, J.T., Anmarkrud, J.A., Calabuig, P., Cooper, J.E.J., Johannessen, L.E., Johnsen, A.,
defray the costs of the museum work; Sarah Durant, Karen Hsu and Sally Kearns, A.M., Lachlan, R.F., Laskemoen, T., Marthinsen, G., Stensrud, E. & Garcia-del-
Davidson, whose homes were a frequent base for NJC in the course of Rey, E. (2016). Species-level divergences in multiple functional traits between the two
endemic subspecies of Blue Chaffinches Fringilla teydea in Canary Islands. BMC Zool. 1: 4.
the museum work in Tring, New York and Washington; Per Alström, who doi: 10.1186/s40850–016–0008–4.
reviewed all material relating to the leaf-warblers and advised on certain Lim, H.C., Chua, V.L., Benham, P.M., Oliveros, C.H., Rahman, M.A., Moyle, R.G. & Sheldon,
other taxa; John Croxall, who gave us his extensive notes on taxonomic F.H. (2014). Divergence history of the Rufous-tailed Tailorbird (Orthotomus sericeus) of
revisions over the past five years; Frank Lambert, who undertook an initial Sunda­­land: implications for the biogeography of Palawan and the taxonomy of island spe-
cies in general. Auk 131(4): 629–642.
set of vocal analyses and gave much advice on particular taxonomic cases McCormack, J.E. & Maley, J.M. (2015). Interpreting negative results with taxonomic and con-
to review; and Van Remsen, whose service to the ornithological commu- servation implications: another look at the distinctness of coastal California Gnatcatchers.
nity in distributing copies of newly published papers has been invaluable. Auk 132(2): 380–388.
And at Lynx, for help in a variety of ways, we express our personal Olsson, U., Alström, P., Svensson, L., Aliabadian, M. & Sundberg, P. (2010). The Lanius excubi­
tor (Aves, Passeriformes) conundrum—taxonomic dilemma when molecular and non-molec-
thanks to Dolors Buxó, Miguel Ángel Carbayo, Cam Christie, Heather ular data tell different stories. Mol. Phylogenet. Evol. 55(2): 347–357.
Elliott, Sandy Elliott, Fortunato Frías, Isabel Martínez, Olga Rius, Pilar Päckert, M., Martens, J., Liang, W., Hsu, W.-C. & Sun, Y.-H. (2013). Molecular genetic and bio-
Ruiz-Olalla and Iván Teba. acoustic differentiation of Pnoepyga wren-babblers. J. Orn. 154(2): 329–337.
Parker, K.A., Hauber, M.E. & Brunton, D.H. (2010). Contemporary cultural evolution of a
conspecific recognition signal following serial translocations. Evolution 64(8): 2431–2441.
Payne, R.B. & Sorenson, M.D. (2007). Integrative systematics at the species level: plumage,
References songs and molecular phylogeny of Quailfinches Ortygospiza. Bull. Brit. Orn. Club 127(1): 4–26.
Peterson, A.P. (2016). Zoonomen. Zoological nomenclature resource. URL: http://www.
Alström, P., Rasmussen, P.C., Zhao, C., Xu, J., Dalvi, S., Cai, T., Guan, Y., Zhang, R., Kalyakin, zoonomen.net.
M.V., Lei, F. & Olsson, U. (2016). Integrative taxonomy of the Plainbacked Thrush (Zoothera Pratt, H.D. & Mittermeier, J.C. (2016). Notes on the natural history, taxonomy, and conserva-
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Res. 7: 1. doi: 10.1186/s40657-016-0037-2. Remsen, J.V. (2015). Book reviews: HBW and BirdLife International Illustrated Checklist of the
Anon. (2016). 12-month finding on a petition to delist the coastal California Gnatcatcher. Carls- Birds of the World. Volume 1: Non-passerines. J. Field Orn. 86(2): 182–187.
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Baker, M.C. (2006). Differentiation of mating vocalizations in birds: acoustic features in main- Remsen, J.V., Areta, J.I., Cadena, C.D., Claramunt, S., Jaramillo, A., Pacheco, J.F., Pérez-Emán,
land and island populations and evidence of habitat-dependent selection on songs. Ethology J., Robbins, M.B., Stiles, F.G., Stotz, D.F. & Zimmer, K.J. (2016). A classification of the bird
112(8): 757–771. species of South America. American Ornithologists’ Union. Version August 2016. URL:
Baker, M.C., Baker, M.S.A. & Tilghman, L.M. (2006). Differing effects of isolation on evolution http://www.museum.lsu.edu/~Remsen/SACCBaseline.htm
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Neotropical radiation of grassland seedeaters. J. Avian Biol. 46(1): 55–62. of Old World chats and flycatchers reveals extensive paraphyly at family, subfamily and genus
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gas (Cotingidae, Aves) with a comparative evolutionary analysis of breeding system and plum- Sangster, G., King, B.F., Verbelen, P. & Trainor, C.R. (2013). A new owl species of the genus
age dimorphism and a revised phylogenetic classification. Mol. Phylogenet. Evol. 81: 120–136. Otus (Aves: Strigidae) from Lombok, Indonesia. PLoS ONE 8(2): e53712.
Boyd, J.H. (2016). Taxonomy in Flux Checklist 3.06. URL: http://jboyd.net/Taxo/List.html. Sangster, G., Rodríguez-Godoy, F., Roselaar, C.S., Robb, M.S. & Luksenburg, J.A. (2016). Inte-
Brumfield, R.T. (2005). Mitochondrial variation in Bolivian populations of the Variable Ant- grative taxonomy reveals Europe’s rarest songbird species, the Gran Canaria blue chaffinch
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Campagna, L., Gronau, I., Silveira, L.F., Siepel, A. & Lovette, I. (2015). Distinguishing noise Sinclair, I. & Langrand, O. (2013). Birds of the Indian Ocean Islands. 3rd edition. Struik Nature,
from signal in patterns of genomic divergence in a highly polymorphic avian radiation. Mol. Cape Town.
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Campbell, K.K., Braile, T. & Winker, K. (2016). Integration of genetic and phenotypic data in avian extinctions at the species and subspecies level. PLoS ONE 7(10): e47080.
48 lineages of Philippine birds shows heterogeneous divergence processes and numerous Tobias, J.A., Seddon, N., Spottiswoode, C.N., Pilgrim, J.D., Fishpool, L.D.C. & Collar, N.J.
cryptic species. PLoS ONE 11(7): e0159325. (2010). Quantitative criteria for species delimitation. Ibis 152(4): 724–746.
Clements, J.F., Schulenberg, T.S., Iliff, M.J., Roberson, D., Fredericks, T.A., Sullivan, B.L. & Winkler, D.W., Billerman, S.M. & Lovette, I.J. (2015). Bird Families of the World: An Invitation to
Wood, C.L. (2016). The eBird/Clements checklist of birds of the world. Version 2016. URL: the Spectacular Diversity of Birds. Lynx Edicions, Barcelona.
http://www.birds.cornell.edu/clementschecklist/download. Zink, R.M., Groth, J.G., Vázquez-Miranda, H. & Barrowclough, G.F. (2013). Phylogeography
Collar, N.J., del Hoyo, J. & Jutglar, F. (2015). The number of species and subspecies in the of the California Gnatcatcher (Polioptila californica) using multilocus DNA sequences and
Red-bellied Pitta Erythropitta erythrogaster complex: a quantitative analysis of morphological ecological niche modeling: implications for conservation. Auk 130(3): 449–458.
characters. Forktail 31: 13–23. Zink, R.M., Groth, J.G., Vázquez-Miranda, H. & Barrowclough, G.F. (2016). Geographic vari-
Collar, N.J., Fishpool, L.D.C., del Hoyo, J., Pilgrim, J.D., Seddon, N., Spottiswoode, C.N. & ation, null hypotheses, and subspecies limits in the California Gnatcatcher: a response to
Tobias, J.A. (2016). Towards a scoring system for species delimitation: a response to Remsen. McCormack and Maley. Auk 133(1): 59–68.
J. Field Orn. 87(1): 104–110. Zou, F., Lim, H.C., Marks, B.D., Moyle, R.G. & Sheldon, F.H. (2008). Corrigendum to “Mo-
Davies, G.B.P. & Peacock, D.S. (2014). Reassessment of plumage characters and morphomet- lecular phylogenetic analysis of the Grey-cheeked Fulvetta (Alcippe morrisonia) of China and
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 Introduction 21

Issues of nomenclature in the current volume

The relevant contents of papers referred to in this work is commonly commented briefly in the texts, but a notable exception is in alterations of the authors and dates of taxa. To avoid
tedious, long drawn-out explanations of very simple modifications, we have merely added the relevant reference number to the author(s) or year when these differ from the version used
in HBW.
The nomenclature of Heliobletus contaminatus (page 106) has recently suffered considerable upheaval, but the matter is now under control. Nevertheless, at the time of writing, the
northern race still lacks a valid name. Vítor de Q. Piacentini (in litt.) and Fernando Pacheco are in the process of remedying this situation, but meantime we are in the awkward position
of having a well-established and universally recognized subspecies temporarily without a valid name. In these exceptional circumstances, we decided that we had no option but to leave
this race in the text but for the present nameless; it is not a simple question of a replacement name, as the authors of the new name will explain in due course. This procedure should not
be confused with the numerous cases, some widely known over many years within the ornithological community, of taxa that are undoubtedly valid forms but have not yet been formally
described. Such forms have never been included formally in HBW, nor are they in this checklist.
Our treatment of a few names of authors of taxa has evolved with the current volume, in part due to new factors, and in part due to fuller information. For example, the inclusion of
an extinct species described by Julian Hume means that we have had to add Allan O. Hume’s initials to all of the many accepted taxa he described. Again, up to now the name Bartels
received the same treatment as Phelps and Gurney, involving father and son with the same initials. However, we now have a new Bartels (A.), and this means that the father and son now
also need their initial, M. Although Max, Sr was M. E. G. Bartels, to date we have not been able to verify the full initials of Max, Jr, but note that they were regularly referred to as Sr and Jr,
and so consider it anyway appropriate to maintain this, at any rate until proven wrong; see, e.g. J. H. Becking (2009) Bull. Brit. Orn. Club 129(1): 18–48. Some years ago we were advised to
style the name of a certain taxon describer as R. G. W. Ramsay. However, on revisiting this case recently, we established that the correct form is actually Wardlaw-Ramsay (with a hyphen).
It appears that his grandfather, Mr Wardlaw, married a Miss Ramsay, so, if any abbreviated form had been used, he would probably have been more likely to be called Mr Wardlaw than Mr
Ramsay. Nonetheless, due to our past treatment of this surname, for clarity we retain the initials for Mr E. P. Ramsay.

Replacement names
As a result of recent studies, mainly genetic, the present volume contains many cases of altered generic limits. Sometimes the work is still in progress and exact divisions between redefined
genera have yet to be clarified, leading in a few instances to a provisional lumping together to form outsize genera in order to avoid paraphyly, but at the same time creating a few new
cases of homonymy.
In order to resolve these, the following new replacement names are hereby introduced. As all are new replacement names for species-group taxa, they are junior objective synonyms of
the taxa they replace. In each case, the type specimen is automatically designated as the same specimen that is the type of the taxon replaced (ICZN Code Art. 72.7). Similarly, in each case
the type locality is automatically that of the replaced taxon.
Phylloscopus intermedius zosterops nom. nov. A. Elliott & del Hoyo, 2016
New replacement name for Abrornis affinis F. Moore, 1854, in Phylloscopus preoccupied by Motacilla Offinis [sic] Tickell, 1833 [currently = Phylloscopus affinis].
Etymology. From the Greek zwsthρ, “girdle”, and wψ, “eye”, referring to the prominent white eyering of this Himalayan form, and recalling the white-eye genus of the same name. This
name is a noun in apposition and is thus invariable.
Phylloscopus montis barisanus nom. nov. Christie & A. Elliott, 2016
New replacement name for Cryptolopha montis inornata Robinson & Kloss, 1920, in Phylloscopus preoccupied by Regulus inornatus Blyth, 1842 [currently = Phylloscopus inornatus].
Etymology. An adjectival name based on Bukit Barisan, the mountain range that forms the spinal column of the island of Sumatra and encompasses the range of this taxon.
Tangara episcopus prysjonesi nom. nov. Collar & Kirwan, 2016
New replacement name for Tanagra Berlepschi Dalmas, 1900, in Tangara preoccupied by Calliste nigriviridis [sic] Berlepschi Taczanowski, 1884 [currently = Tangara nigroviridis berlepschi].
Etymology. Named in honour of Robert Prŷs-Jones, recently retired as Head of the Bird Group at the Natural History Museum (UK), for his unswerving championship of the importance
of museum-based ornithology over the past three decades, and his consistently generous assistance to visiting researchers, ourselves included. The name is a noun in the genitive singular,
and is invariable.
Tangara episcopus johntoddzimmeri nom. nov. A. Elliott, 2016
New replacement name for Thraupis episcopus urubambae J. T. Zimmer, 1944, in Tangara preoccupied by Tangara parzudakii urubambae J. T. Zimmer, 1943 [currently = Tangara parzudakii
urubambae].
Etymology. Named in honour of John Todd Zimmer (1889–1957), one of the most prolific contributors to our knowledge of Neotropical birds, late Curator of Birds at the American
Museum of Natural History, and describer of both the junior and senior homonyms in this case. The name is a noun in the genitive singular, and is invariable.
Tangara sayaca beniensis nom. nov. A. Elliott, 2016
New replacement name for Thraupis episcopus boliviana J. Bond & Meyer de Schauensee, 1941, in Tangara preoccupied by Callospiza boliviana Bonaparte, 1851 [currently = Tangara mexicana
boliviana].
Etymology. An adjectival name based on the department of Beni (northern Bolivia), which encompasses much of the range of this taxon, including its type locality of Chatarona.

First Reviser acts

Several cases have been encountered of simultaneously published names vying for precedence, and four of these appear to be unresolved. As no previous valid acts have been located that
clearly solve the following cases, A. Elliott hereby acts as First Reviser to establish the precedence of:
(a) Deconychura longicauda pallida J. T. Zimmer, 1929 over Deconychura longicauda connectens J. T. Zimmer, 1929;
(b) Anthreptes longmari [sic] angolensis Neumann, 1906 over Anthreptes longmari [sic] nyassae Neumann, 1906;
(c) Icterus graduacauda dickeyae van Rossem, 1938 over Icterus graduacauda nayaritensis van Rossem, 1938; and
(d) Melospiza heermanni S. F. Baird, 1858a over Melospiza gouldii S. F. Baird, 1858a.
Notes on (d)
Also included in this particular synonymy is Ammodromus samuelis S. F. Baird, 1858b, which was already placed in the synonymy of gouldii by Coues (1872). Subsequently, both Ridgway
(1901) and Hellmayr (1938) included gouldii in the synonymy of samuelis! In his original description of the taxa heermanni and gouldii, Baird (1858a) already suggested (page 477) that
gouldii might be merely a variation (“Var. gouldii”). In fact, Cooper & Baird (1870) appear maybe to include gouldii within heermanni, but the text is somewhat unclear, and perhaps not
wholly convincing as a First Reviser act, so it has been considered more appropriate to issue a new act here. The precise date of publication of samuelis is often reported to be August
1858, but internal evidence indicates that it was certainly no earlier than 25th October 1858; it has been deemed best to treat the two publications as appearing simultaneously, until
proven otherwise.
Many thanks to Frank Steinheimer and Normand David for their valuable advice. Any errors are, however, of course entirely mine (AE).

References
Baird, S.F. (1858a). Rep. Explor. Surv. Railroad Mississ. Pacific Ocean 9: 478–479.
Baird, S.F. (1858b). Proc. Boston Soc. Nat. Hist. 6: 379.
Blyth, E. (1842). J. Asiatic Soc. Bengal 11(1): 191.
Bonaparte, C.L. (1851). Compt. Rend. Acad. Sci. Paris 32: 80.
Bond, J. & Meyer de Schauensee, R. (1941). Notulae Naturae 93: 6.
Cooper, J.G. & Baird, S.F. (1870). Ornithology of California. Vol. 1. Land Birds. Geological Survey of California. Page 212.
Coues, E. (1872). Key North Amer. Birds. 1st edition. Page 139.
Dalmas, R. de (1900). Mém. Soc. Zool. France 13: 136.
Hellmayr, C.E. (1938). Catalogue of Birds of the Americas and the adjacent islands. Field Mus. Nat. Hist. (Zool. Ser.) 13(11): 603.
Moore, F. (1854). Cat. Birds Mus. Hon. East-India Co. 1: 341.
Neumann, O. (1906). Orn. Monatsber. 14(1): 6–7.
Ridgway, R. (1901). The Birds of North and Middle America. Bull. US Natl. Mus. 50(1): 370.
Robinson, H.C. & Kloss, C.B. (1920). J. Straits Branch Roy. Asiat. Soc. 81: 99.
van Rossem, A.J. (1938). Bull. Brit. Orn. Club 58(9): 137.
Taczanowski, L. (1884). Orn. Pérou 2: 469.
Tickell, S.R. (1833). J. Asiatic Soc. Bengal 2(23): 576.
Zimmer, J.T. (1929). Field Mus. Nat. Hist. (Zool. Ser.) 17: 6–8.
Zimmer, J.T. (1943). Amer. Mus. Novit. 1245: 10.
Zimmer, J.T. (1944). Amer. Mus. Novit. 1262: 13.

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