J Zool Syst Evol Res. 2019;00:1–8. wileyonlinelibrary.com/journal/jzs
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1 © 2019 Blackwell Verlag GmbH
1 | INTRODUCTION
The family Anhingidae contains the birds known colloquially as
snakebirds (due to their appearance when swimming—with their
body submerged all that is visible is a snake‐like elongated neck,
non‐differentiated elongated head and slender beak), anhingas
(“devil bird” or “snake bird” in Brazilian Tupian language), and dart‐
ers (due to their unique foraging behavior, spearing fish and other
aquatic prey using their beaks). These active fish hunters are found
in freshwater habitats in most parts of the tropics and subtropics.
Like their close relatives the cormorants (family Phalacrocoracidae)
and the gannets and boobies (Sulidae), with which they form the
Suliformes, they exhibit a remarkable degree of morphological con‐
servatism, to the point where the number of species recognized has
varied among different authorities. The New World species, the
Anhinga or American Darter Anhinga anhinga (Linnaeus, 1766), is
clearly separable from the remaining Old World forms. The latter
are variously treated as a single species, the Darter Anhinga mela‐
nogaster (Pennant, 1769), with three subspecies (Christidis & Boles,
1994; Clements, 2007; Cramp & Simmons, 1977; Dickinson, 2003;
Harrison, 1978; Johnsgard, 1993; Nelson, 2005; O'Brien, 1990;
Orta, 1992; Worthy, 2012), or, perhaps less often, as three separate
species, the Oriental or Indian Darter ( A. melanogaster), the African
Darter Anhinga rufa (Daudin, 1802), and the Australasian Darter
Anhinga novaehollandiae (Gould, 1847) (Clements et al., 2017; del
Hoyo, Collar, Christie, Elliott, & Fishpool, 2014; Ogilvie‐Grant, 1898;
Peters, 1931; Schodde, Kirwan, & Porter, 2012; Sibley & Monroe,
1990; Vaurie, 1965). The more recent treatments by some of these
Received: 17 December 2018
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Revised: 2 May 2019
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Accepted: 5 May 2019
DOI: 10.1111/jzs.12299
ORIGINAL ARTICLE
Sorting out the Snakebirds: The species status, phylogeny, and
biogeography of the Darters (Aves: Anhingidae)
Martyn Kennedy
1
| Sampath S. Seneviratne
2
| Ushawarni K. Mendis
2
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Hamish G. Spencer
1
1
Department of Zoology, University of
Otago, Dunedin, New Zealand
2
Avian Evolution Node, Department of
Zoology & Environment Sciences, University
of Colombo, Colombo, Sri Lanka
Correspondence
Martyn Kennedy, Department of Zoology,
University of Otago, Dunedin, New Zealand.
Email: martyn.kennedy@otago.ac.nz
Funding information
American Museum of Natural History,
Grant/Award Number: 3, 2, 41 and 14;
University of Otago; Allan Wilson Centre
Contributing authors: Sampath S.
Seneviratne (sam@sci.cmb.ac.lk), Ushawarni
K. Mendis (mushawarni@gmail.com), Hamish
G. Spencer (hamish.spencer@otago.ac.nz)
Abstract
Although the avian family Anhingidae is unequivocally monophyletic, the number
and relationships of the component species within the single genus ( Anhinga) have
long remained unclear. Here, we use extensive mitochondrial and nuclear DNA se‐
quence data (8,878 bp) to show that four species should be recognized. Our fully
resolved and well‐supported tree shows that the American Anhinga ( Anhinga an‐
hinga) is sister to the three Old World species, with the Oriental ( A. melanogaster) and
African ( A. rufa) Darters sister within the Old World clade, which also includes the
Australian Darter ( A. novaehollandiae). We estimate that the divergence between the
New World and Old World branches occurred 19–22 mya, with the Australian Darter
separating from its Old World congeners 14–16 mya and the Oriental and African
species splitting ~10 mya. The genus is yet another example of osteological conserva‐
tism in the Suliformes, which is comparable to that shown by the cormorants and
shags. Nevertheless, the relationships we infer are congruent with recent plumage
studies and are biogeographically plausible. We suggest that further investigation of
the variation within the African and Australian Darters would be of interest.
KEYWORDS
Anhinga, Anhingidae, Darters, phylogeny, taxonomy