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Lerner et. al.; 2005

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http://heatherlerner.com/pdfs/Lerner.Mindell.MPE.2005.pdf 
Phylogeny of eagles, Old World vultures, and other
Accipitridae based on nuclear and mitochondrial DNA
Heather R.L. Lerner *
, David P. Mindell
Department of Ecology and Evolutionary Biology, University of Michigan Museum of Zoology, 1109 Geddes Ave., Ann Arbor, MI 48109-1079, USA
Received 16 November 2004; revised 31 March 2005
Available online 31 May 2005
Abstract
[1] We assessed phylogenetic relationships for birds of prey in the family Accipitridae using molecular sequence from two mitochondrial genes (1047 bases ND2 and 1041 bases cyt-b) and one nuclear intron (1074 bases b-fibrinogen intron 7). We sampled representatives of all 14 Accipitridae subfamilies, focusing on four subfamilies of eagles (booted eagles, sea eagles, harpy eagles, and
snake eagles) and two subfamilies of Old World vultures (Gypaetinae and Aegypiinae) with nearly all known species represented.
Multiple well-supported relationships among accipitrids identified with DNA differ from those traditionally recognized based on
morphology or life history traits. Monophyly of sea eagles (Haliaeetinae) and booted eagles (Aquilinae) was supported; however,
harpy eagles (Harpiinae), snake eagles (Circaetinae), and Old World vultures were found to be non-monophyletic. The Gymnogene
(Polyboroides typus) and the Crane Hawk (Geranospiza caerulescens) were not found to be close relatives, presenting an example of
convergent evolution for specialized limb morphology enabling predation on cavity nesting species. Investigation of named subspecies within Hieraaetus fasciatus and H. morphnoides revealed significant genetic differentiation or non-monophyly supporting recognition of H. spilogaster and H. weiskei as distinctive species.
2005 Elsevier Inc. All rights reserved.

Keywords: Accipitridae; Avian systematics; Phylogenetics; Molecular evolution; Cytochrome-b; ND2; b-Fibrinogen intron 7
1. Introduction
Accipitridae is a diverse avian family, comprising up
to 14 subfamilies, 65 genera, and 231 species (see Table
1, Dickinson, 2003; Stresemann and Amadon, 1979). [2] Of
the Accipitridae species,
some of the largest and most
threatened by anthropogenic factors belong to four eagle subfamilies
(Circaetinae, Haliaeetinae, Aquilinae,
and Harpiinae) and two Old World vulture subfamilies
(Gypaetinae and Aegypiinae). All Accipitridae species
are protected under the Convention on International
Trade in Endangered Species (CITES) and four eagles
are listed as top priority species (CITES I, CITES-Secretariat, 2003). As ecologically sensitive predators, birds of
prey are valuable indicators of habitat quality. The
Accipitridae are found in a variety of habitats from primary rainforest to arctic tundra throughout the world.
Some taxa are restricted in distribution such as the
snake eagles (Circaetinae) which are found only in the
Old World, while others, such as the sea eagles (Haliaeetinae), are global in distribution. Thorough phylogenetic analyses are needed to delineate the genetic and
overall biological diversity of this family, and to inform
conservation programs which aim to preserve genetic
diversity of distinguishable taxonomic units.
Phylogeny for Accipitridae based on morphological
traits has been difficult to resolve (e.g., Brown and Amadon, 1968; Jollie, 1976, 1977a,b). [3] The few published
molecular studies have been limited in sampling and
have proposed some previously unrecognized relationships (see below). The goal of the present study is to
identify phylogenetic relationships within and among
the six subfamilies of eagles and Old World vultures in the context of the other primary accipitrid groups using
molecular data.

The booted eagles (Aquilinae) are one of the largest
accipitrid groups containing 35–36 species in 8–9 genera
and are distributed worldwide. The majority of the species fall into three genera, Aquila, Hieraaetus, and Spizaetus, while the remaining five genera are all monotypic.
All species have ‘‘boots,’’ or feathered tarsi, a trait that
separates this group from most other accipitrid taxa.
The booted eagles have been considered to be monophyletic (Jollie, 1977b) or polyphyletic (Holdaway, 1994)
with morphological data, and only a few species in
one genus have been studied phylogenetically with
molecular data (cyt-b, Seibold et al., 1996; control region, Vali, 2002). Monophyly of the three Aquilinae
genera is not well supported with morphological characters, such that the Hieraaetus species and some Spizaetus
species have been placed in the genus Aquila by various
authors (described by Brown and Amadon, 1968; and
Thiollay, 1994). The two molecular studies included
about half of the species in the genus Aquila, and
both found that A. chrysaetos was genetically distant
from four other Aquila species. Sister relationships for
A. clanga and A. pomarina, A. nipalensis and A. heliaca
or A. heliaca and A. adalberti were also proposed.
The sea eagles (Haliaeetinae) are a much smaller and
more easily defined group of large eagles found in
coastal and riverine areas worldwide except South
America and Antarctica. The two sea eagle genera,
Haliaeetus and Ichthyophaga, share some morphological traits with two genera of kites (Milvus and Haliastur), suggesting a close relationship (Holdaway, 1994;
Jollie, 1977b; Thiollay, 1994). The sea eagles also share some traits with the palmnut vulture (Gypohierax angolensis), suggesting a relationship between them and Old
World vultures (Brown and Amadon, 1968). Using cytb sequence data, Seibold and Helbig (1996) studied
eight of the nine species of sea eagles in the genus
Haliaeetus. They supported a clear split between species
with temperate versus tropical distributions, and a close
relationship between the sea eagles and two Milvus
kites. The relationship between the two genera of sea
eagles has not been investigated with molecular sequence data and the possibility of paraphyly of the genera remains unresolved.
The four species and genera of harpy eagles (Harpiinae) are some of the largest raptors and are found in
tropical rain forests in the Americas, the Philippines
and New Guinea. This group is generally considered
monophyletic due to their large size, lack of feathers
on the tarsi, and similarities in behavior (Brown and
Amadon, 1968; Thiollay, 1994); however, [4] some have
suggested that the Old World species are not sister to
the New World species
(e.g., Jollie, 1977b). Holdaway
(1994) removed one Old World (Pithecophaga) and
one New World (Morphnus) species from the Harpiinae.
A close relationship between the booted eagles and the
harpy eagles has been proposed but not tested with
molecular data.
The 14 species of snake eagles (Circaetinae) in five
genera are found only in the Old World. Although usually considered monophyletic (Brown and Amadon,
1968; Friedmann, 1950), the possibility of polyphyletic
origins for snake eagles has been raised (Jollie, 1977b,
could not identify sister relationships for Eutriorchis and Dryotriorchis; Thiollay, 1994). The final group we focused on is the Old World vultures, a diverse mix of scavengers including at least one
species that uses tools (Egyptian vulture), and potentially including a frugivorous raptor (palmnut vulture).
One or two subfamilies have been proposed for the
Old World vultures. Three species are highly divergent
from the remaining 11 and have been placed by some
in a separate family called Gypaetinae (Mundy et al.,
1992). The core 11 species are called the Aegypiinae. Seibold and Helbig (1995) used cyt-b sequence from 11 Old
World vulture species and found evidence of polyphyly
for the Old World vultures.
There are no previously published molecular studies
that include representatives of all of the Accipitridae
subfamilies; however, several molecular studies have
used mitochondrial DNA to examine particular Accipitridae subgroups and have found evidence for polyphyly
of some traditionally recognized taxa (e.g., polyphyly of
the Perninae kites, Riesing et al., 2003; and the genus
Buteo, Gamauf and Haring, 2004). Relationships among
a small set of accipitrids based on mtDNA indicated a
closer relationship between a representative sea eagle
and kite in the genus Milvus, than between the sea eagle
and a snake eagle in the genus Circaetus. A representative Old World vulture was more closely related to the
snake eagle than other accipitrid taxa in the study,
including species of Buteo, Haliaeetus, Milvus, Circus,
Accipiter, and Pernis (Mindell et al., 1997). Increased
sampling of species and molecular characters are needed
to improve our understanding of phylogenetic relationships among the Accipitridae.
In this study, we focus on full or nearly complete taxonomic representation of five accipitrid subgroups (sea
and fish eagles, harpy eagles, booted eagles, snake eagles, and Old World vultures), corresponding to six potential subfamilies. We use both mitochondrial and
nuclear sequences for representatives of 51 out of 65
genera (78%) and just under half of the known Accipitridae species (n = 111). At least one representative of each
previously proposed subgroup/subfamily within the
Accipitridae have been included to help in phylogenetic placement of the focal taxa.
Circaetinae
a
Snake eagles Circaetus, Terathopius,
Dryotriorchis,
Eutriorchis, Spilornis
Old World species feeding mainly
on snakes, other reptiles and
small mammals, have a reticulate
pattern of heavy scales on the
tarsi and relatively short toes
Circaetus, Terathopius,
Dryotriorchis, Spilornis and
Pithecophaga
Circaetinae
Pithecophaga jefferyi Philippines TPEF, captive 227 AY987246 AY987064 AY987171
Pithecophaga jefferyi Philippines TPEF, captive 205 AY987245 AY987065 AY987172
Pithecophaga jefferyi Mindanao, Philippines AMNH 534856 n/0
with nearly all known species represented.
Multiple well-supported relationships among accipitrids identified with DNA differ from those traditionally recognized based on
morphology or life history traits. Monophyly of sea eagles (Haliaeetinae) and booted eagles (Aquilinae) was supported; however,
harpy eagles (Harpiinae), snake eagles (Circaetinae), and Old World vultures were found to be non-monophyletic. The Gymnogene
(Polyboroides typus) and the Crane Hawk (Geranospiza caerulescens) were not found to be close relatives, presenting an example of
convergent evolution for specialized limb morphology enabling predation on cavity nesting species. Investigation of named subspecies within Hieraaetus fasciatus and H. morphnoides revealed significant genetic differentiation or non-monophyly supporting recognition of H. spilogaster and H. weiskei as distinctive species.
Corresponding author. Fax: +1 734 763 4080.
E-mail address: hlerner@umich.edu (H.R.L. Lerner).
Molecular Phylogenetics and Evolution 37 (2005) 327–346
MOLECULAR
PHYLOGENETICS
AND
EVOLUTION
www.elsevier.com/locate/ympev
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