AFROAVES II

ACCIPITRIMORPHAE: Cathartiformes, Accipitriformes

Afroaves

Coliiformes

Coliidae: Mousebirds

Accipitrimorphae

Strigiformes

Picimorphae

Leptosomiformes

Leptosomidae: Cuckoo Roller

Trogoniformes

Trogonidae: Trogons

Bucerotiformes

Coraciiformes

Piciformes

ACCIPITRIMORPHAEVieillot, 1816

CATHARTIFORMESCoues, 1884

Although New World vultures are rather similar to Old World Vultures (part ofAccipitridae), they are not closely related. This was already recognized byHuxley in 1876 based on morphological evidence. Nonetheless, they weretraditionally placed in Falconiformes, as where the Accipitridae and Falconidae.There are also some superficial similarities to storks, and when Sibley and Ahlquistfound support for this using DNA hybridization techniques, they were removedfrom Falconiformes and placed next to the storks. More recent analysis haveshown this is incorrect, and have also broken up Falconiformes. As mentioned above,more recent research (e.g., Hackett et al., 2008; Han et al., 2011;McCormack et al., 2013; Yuri et al., 2013; Jarvis et al., 2014) puts the New World vultures close to the Accipitriformes.

Cathartidae tree

The phylogeny of Cathartidae is based on Johnson et al. (2016).

Cathartidae: New World Vulturesde Lafresnaye, 1839

5 genera, 7 speciesHBW-2

Andean Condor,Vultur gryphus
California Condor,Gymnogyps californianus
King Vulture,Sarcoramphus papa
Black Vulture,Coragyps atratus
Turkey Vulture,Cathartes aura
Lesser Yellow-headed Vulture,Cathartes burrovianus
Greater Yellow-headed Vulture,Cathartes melambrotus

ACCIPITRIFORMESVieillot, 1816

Those interested in ancient raptors should readDarren Naish's post on Titan-Hawks.

The Secretarybird is the first branch of Accipitriformes, followed bythe Osprey. After that, come the Accipitridae (hawks, kites, andeagles).

Sagittariidae: SecretarybirdFinsch & Hartlaub, 1870 (1825)

1 genus, 1 speciesHBW-2

Secretarybird,Sagittarius serpentarius

Pandionidae: OspreyBonaparte, 1854

1 genus, 1 speciesHBW-2

It is not clear whether the Osprey is a single species. The IOC recognizestwo Osprey species, Eastern (Indo-Australasiancristatus) and Western(everything else). Some have suggested that the Caribbean population(ridgwayi) should also be separated.

The TiF list does not currently accept either of these splits. Monti et al.(2015) sampled over 200 individuals world-wide. They found 4 clades: New world(carolinensis, includingridgwayi), Old World (haliaetus),Indo-Australian (cristatus), and North-East Asia (name uncertain). The North-East Asian clade included samples from Japan and the Pacific coast ofSiberia, as well as a few from Indonesia/New Guinea and one from East Asia.Also, acristatus haplotype was found in India. At first glance, itseemsfriedmanni may apply to the NE Asian clade, but Wolfe's descriptionoffriedmanni (Auk, Oct., 1946) states that Ospreys from Japan are notfriedmani, making me wonder which clade the type offriedmanniactually belongs to.

Monti et al. suggest that the New World clade is basal, as suggested by thecyt-b analysis. However, combining ND2 with cyt-b yielded a different topology,with the Old World clade basal. In any event, the separation of all 4 cladesappears to have taken place over a short time interval. The cyt-b analysissuggested the New World clade is a million or so years old. A few of the samples showed a mismatch between geography and clade.

At present, it remains unclear whether or not the Osprey should be considered aconsidered a single species, or 3-4.

Osprey,Pandion haliaetus

Accipitridae: Hawks, Kites, EaglesVigors, 1824

73 genera, 255 speciesHBW-2

At the genus level, the order presented here attempts to synthesizethe papers by Amaral et al. (2006, 2009), Barrowclough et al. (2014),Breman et al. (2013), Griffiths et al. (2007), Haring et al. (2007b),Helbig et al. (2005), Kocum (2006), Lerner and Mindell (2005), Lerner etal. (2008), Nagy and Tökölyi (2014), Oatley et al. (2015),Ong et al. (2011), and Wink and Sauer-Gürth (2004). Some of the generawere restructured based on these and other papers.

The Accipitridae are a complicated family. You can see from the diagram that Itreat them as consisting of ten subfamilies with large families Accipitrinae andButeoninae at the end. The latter two subfamilies are further divided intotribes. The timing estimates of Nagy and Tökölyi (2014) haveinfluenced what gets counted as a subfamily and what counts as a tribe. Afterreading Jarvis et al. (2014), I suspect Nagy and Tökölyi's divergencedates are a little short. Nagy and Tökölyi did not include thecrested goshawk subfamily, Lophospizinae, but indications are that it is nearAquilinae (Oatley et al., 2015).

The subfamily Accipitrinae is slightly smaller than Buteoninae, so I put itahead of Buteoninae. The crested goshawks seem to be basal to both of them. There's now some evidence they belong in the Harpiinae plus Aquilinae group,which is sister to the Accipitrinae + Buteoninae group. The grouping of thesefive families is supported by morphology as well as genetics. Mayr (2014b) foundthat four of them (with the fifth not tested) share a derived type of radialcarpal bone. The closest relatives of this big clade are Circaetinae andGypinae. All together are sister to Gypaetinae and Perninae, and the wholeshebang is sister to the basal group, Elaninae. This is all summed up in thediagram.

Click for genus-level tree of Accipitriformes

Elaninae: Elanine KitesBlyth, 1851

One thing that genetic results have made clear is that the kites are nota natural group. Rather, they are scattered in three subfamilie.The first group is the Elanine kites. Kites also occur in three clades inPerninae and four clades in Buteoninae.

Pearl Kite,Gampsonyx swainsonii
Click for Accipitridae
species tree
Scissor-tailed Kite,Chelictinia riocourii
Black-winged Kite,Elanus caeruleus
Black-shouldered Kite,Elanus axillaris
Letter-winged Kite,Elanus scriptus
White-tailed Kite,Elanus leucurus

Gypaetinae: African Vultures, Harrier-HawksBonaparte, 1831

Now that it is clear thatHenicopernis belongs within Perninae(Barrowclough et al., 2014), this small primarily African clade iscleanly separated from Perninae. The separation from Perninae seems tobe quite ancient (Nagy and Tökölyi, 2014), so I recognize bothgroups as subfamilies.

African Harrier-Hawk,Polyboroides typus
Madagascan Harrier-Hawk,Polyboroides radiatus
Palm-nut Vulture,Gypohierax angolensis
Egyptian Vulture,Neophron percnopterus
Bearded Vulture / Lammergeier,Gypaetus barbatus

Perninae: Bazas, Honey-BuzzardsBlyth, 1851

The Honey-Buzzards (Pernis) have been sorted out by Gamauf andHaring (2004), including the recently-split Philippine Honey-Buzzard. Barrowclough et al. (2014) provided additional information concerningthe Australo-Papuan species.

Three groups of kites are part of Perninae. The NeotropicalChondrohierax andLeptodon kites, the Swallow-tailedKite (Elanoides, sister to thePernis honey-buzzards),and the Square-tailed Kite (Lophoictinia, sister to theHenicopernis honey-buzzards). Note that the honey-buzzardsare not a natural group either.

Madagascan Serpent-Eagle,Eutriorchis astur
Hook-billed Kite,Chondrohierax uncinatus
Gray-headed Kite,Leptodon cayanensis
White-collared Kite,Leptodon forbesi
Swallow-tailed Kite,Elanoides forficatus
European Honey-Buzzard,Pernis apivorus
Philippine Honey-Buzzard,Pernis steerei
Oriental Honey-Buzzard / Crested Honey-Buzzard,Pernis ptilorhynchus
Barred Honey-Buzzard,Pernis celebensis
Black-breasted Buzzard,Hamirostra melanosternon
Square-tailed Kite,Lophoictinia isura
Long-tailed Honey-Buzzard,Henicopernis longicauda
Black Honey-Buzzard,Henicopernis infuscatus
Black Baza,Aviceda leuphotes
African Cuckoo-Hawk,Aviceda cuculoides
Madagascan Cuckoo-Hawk,Aviceda madagascariensis
Jerdon's Baza,Aviceda jerdoni
Pacific Baza,Aviceda subcristata

Gypinae: Old World VulturesBlyth, 1851

The name Gypinae (Blyth, 1851) has priority over Aegypiinae (W.P.Sclater, 1924). The Old World Vultures are gathered here and fall intotwo clades. The first four species (and genera!) are in one clade, withNecrosyrtes andGyps in the other. The arrangement ofspecies withinGyps has taken both Arshad et al. (2009) and Nagyand Tökölyi (2014) into account. I follow Lerner et al. (2005)and Nagy and Tökölyi (2014) for the genera. Arshad et al. isalmost the same, while Griffiths et al. (2009) is broadly consistent butless well-resolved.

Red-headed Vulture,Sarcogyps calvus
White-headed Vulture,Trigonoceps occipitalis
Cinereous Vulture,Aegypius monachus
Lappet-faced Vulture,Torgos tracheliotos
Hooded Vulture,Necrosyrtes monachus
White-backed Vulture,Gyps africanus
White-rumped Vulture,Gyps bengalensis
Himalayan Vulture,Gyps himalayensis
Griffon Vulture,Gyps fulvus
Rueppell's Vulture,Gyps rueppelli
Cape Vulture,Gyps coprotheres
Slender-billed Vulture,Gyps tenuirostris
Indian Vulture,Gyps indicus

Circaetinae: Serpent EaglesBlyth, 1851

In the Serpent Eagles,Dryotriorchis has been merged intoCircaetus. Exactly how manyspecies are inSpilornis remains an issue. There are six here, butFerguson-Lees and Christie (2001) list 13! I've not accepted the IOCsplit ofminimus because I don't see any evidence to split thisand not the others.

Andaman Serpent-Eagle,Spilornis elgini
Philippine Serpent-Eagle,Spilornis holospilus
Crested Serpent-Eagle,Spilornis cheela
Great Nicobar Serpent-Eagle,Spilornis klossi
Mountain Serpent-Eagle,Spilornis kinabaluensis
Sulawesi Serpent-Eagle,Spilornis rufipectus
Philippine Eagle,Pithecophaga jefferyi
Bateleur,Terathopius ecaudatus
Western Banded Snake-Eagle,Circaetus cinerascens
Southern Banded Snake-Eagle,Circaetus fasciolatus
Congo Serpent-Eagle,Circaetus spectabilis
Beaudouin's Snake-Eagle,Circaetus beaudouini
Black-chested Snake-Eagle,Circaetus pectoralis
Brown Snake-Eagle,Circaetus cinereus
Short-toed Snake-Eagle,Circaetus gallicus

Lophospizinae: Crested GoshawksInformal

What evidence there is suggests that the Crested Goshawk is notpart of Accipitrinae. Barcoding suggested it is basal to the Accipitrinae/Buteoninaeclade (Ong et al., 2011; Breman et al., 2013). Raty, in aBirdForum postgot similar results when adding a partial cytochrome-b gene.More recently, Oatley et al.'s (2015) 4-gene analysis found the Crested Goshawk near theAquilinae (Harpiinae was not in the analysis). I'm not sure I'm entirely convinced,but I've double-down on this and put it in a trichotomywith Harpiinae and Aquilinae. Besides, there's something that appeals to me inhaving the crested goshawks near the harpy eagles.

DNA from the Sulawesi (Crested) Goshawk has notbeen tested, but it is thought to be a very close relative of theCrested Goshawk, with the two forming a superspecies. The nameLophospiza (Kaup 1844, typetrivirgata) is available forthese two species.

Crested Goshawk,Lophospiza trivirgata
Sulawesi Goshawk,Lophospiza griseiceps

Harpiinae: Harpy EaglesLesson, 1828

The split between the harpy eagles and booted eagles seems to have occurredabout 20 million years ago (Nagy and Tökölyi, 2014), so I treat them as separate subfamilies.

Bat Hawk,Macheiramphus alcinus
Papuan Eagle,Harpyopsis novaeguineae
Harpy Eagle,Harpia harpyja
Crested Eagle,Morphnus guianensis

Aquilinae: Booted EaglesVigors, 1825

The key papers for the Booted Eagles (Aquilinae) are Bunce et al. (2005), Helbig et al. (2005), Lerner and Mindell (2005), Haring et al.(2007b), and Lerner et al. (2017). Aquilinae tree TheSpizaetus Hawk-Eagles belong in two different clades withinAquilinae. ThusSpizaetus is divided intoNisaetus andSpizaetus. The Black-and-chestnut Eagle (Oroaetus) mustbe merged into the remainingSpizaetus. The Rufous-bellied Hawk-Eagle(Lophotriorchis) is separated fromHieraaetus. whilethe spotted eagles are separated fromAquila asClanga.Hieraaetus loses a couple of species toAquila, which alsogains Cassin's Hawk-Eagle fromSpizaetus.

Although it is fairly basal, there is still some uncertain concerningthe closest relatives of the Crowned Eagle,Stephanoaetus coronatus.

I've included the extinct Haast's Eagle of New Zealand on the speciestree since Bunce et al. (2005) analyzed DNA from it. However, it isn'tincluded on the main list because it likely became extinct prior to1500, which is the cutoff for inclusion.

Crowned Eagle,Stephanoaetus coronatus
Legge's Hawk-Eagle,Nisaetus kelaarti
Sulawesi Hawk-Eagle,Nisaetus lanceolatus
Pinsker's Hawk-Eagle,Nisaetus pinskeri
Changeable Hawk-Eagle,Nisaetus cirrhatus
Philippine Hawk-Eagle,Nisaetus philippensis
Wallace's Hawk-Eagle,Nisaetus nanus
Mountain Hawk-Eagle,Nisaetus nipalensis
Blyth's Hawk-Eagle,Nisaetus alboniger
Javan Hawk-Eagle,Nisaetus bartelsi
Black Hawk-Eagle,Spizaetus tyrannus
Black-and-white Hawk-Eagle,Spizaetus melanoleucus
Ornate Hawk-Eagle,Spizaetus ornatus
Black-and-chestnut Eagle,Spizaetus isidori
Rufous-bellied Hawk-Eagle,Lophotriorchis kienerii
Martial Eagle,Polemaetus bellicosus
Long-crested Eagle,Lophaetus occipitalis
Black Eagle,Ictinaetus malaiensis
Indian Spotted-Eagle,Clanga hastata
Lesser Spotted-Eagle,Clanga pomarina
Greater Spotted-Eagle,Clanga clanga
Wahlberg's Eagle,Hieraaetus wahlbergi
Ayres's Hawk-Eagle,Hieraaetus ayresii
Pygmy Eagle,Hieraaetus weiskei
Booted Eagle,Hieraaetus pennatus
Little Eagle,Hieraaetus morphnoides
Steppe Eagle,Aquila nipalensis
Tawny Eagle,Aquila rapax
Spanish Imperial-Eagle,Aquila adalberti
Eastern Imperial-Eagle,Aquila heliaca
Golden Eagle,Aquila chrysaetos
Cassin's Hawk-Eagle,Aquila africana
Gurney's Eagle,Aquila gurneyi
Wedge-tailed Eagle,Aquila audax
Verreaux's Eagle,Aquila verreauxii
Bonelli's Eagle,Aquila fasciata
African Hawk-Eagle,Aquila spilogaster

Accipitrinae: Accipiters, HarriersVigors, 1824

There is a deep division of the Accipitrinae into two groups which I rank as tribes: Melieraxini and Accipitrini.

Melieraxini: Chanting Goshawks and alliesInformal

The Tiny Hawk (and presumably Semicollared Hawk) do not seem closelyrelated toAccipiter (Kocum, 2006; Olson, 2006) and have beenremoved fromAccipiter.

New Name forHieraspiza:Sangster et al. (2021a) showed that the genus nameHieraspizadoes not apply to the Tiny Hawk. According, it and the Semicollared Hawkare transferred toMicrospizias, becomingTiny Hawk,Microspizias superciliosus andSemicollared Hawk,Microspizias collaris.

Kocum (2006) includes several species from this group in herdissertation. Her analysis places them in a weakly supported unresolvedbasal polytomy with Accipitrini, Harpagini, and the remainingButeoninae. Griffiths et al. (2007) grouped together the Lizard Buzzard, Gabar Goshawk, Long-tailed Hawk, and the chanting goshawks.The group was basal to Accipitrini and Buteoninae, but did notinclude the Tiny Hawk. The analysis by Nagy and Tökölyi(2014) bundles them together as shown here, but their evidence for thisseems weak.

The name Melieraxinae has appeared in the literature since at least Lernerand Mindell (2005), but does not seem to be formally established.

Lizard Buzzard,Kaupifalco monogrammicus
Gabar Goshawk,Micronisus gabar
Long-tailed Hawk,Urotriorchis macrourus
Tiny Hawk,Microspizias superciliosus
Semicollared Hawk,Microspizias collaris
Dark Chanting-Goshawk,Melierax metabates
Eastern Chanting-Goshawk,Melierax poliopterus
Pale Chanting-Goshawk,Melierax canorus

Notes on Accipitrini: Accipiters & Harriers

The papers by Wink and Sauer-Gürth (2004), Breman et al. (2013),Barrowclough et al. (2014), Nagy and Tökölyi (2014) andKocum's dissertation (2006) allow us to make a rough draft of aspecies-level tree for the accipitters. Doubtless it will need furtheradjustment.

The result is that the genusAccipiter has been divided into6 pieces. Two of those,Hieraspiza andLophospizahave been moved outside the Accipitrini. Kocum (2006), Griffiths et al.(2007), Lerner et al. (2008), and Nagy and Tökölyi (2014)found that theCircus harriers are embedded within thetraditionalAccipiter. The options are to lumpCircus intoAccipiter or divideAccipiter into at least three parts. I've chosen to take the latter course, dividingAccipter intofour parts.

The Cooper's Hawk, Northern Goshawk, and closely related taxa are theclosest relatives of the harriers. They go in genusAstur(Lacépède 1799, typegentilis). These are sister toa clade that includes the Eurasian Sparrowhawk and the Sharp-shinnedHawk complex. It retains the nameAccipiter (Brisson 1760, typenisus). I have included the Gray-bellied Hawk with this groupbased on Kocum (2006), although the support for this is not strong.

The remaining accipiters andErythrotriorchis form a cladesister to the combinedAccipiter-Astur-Circusclade. There is a deep division (ca. 15 million years) between a small groupof African accipiters and the remaining accipiters, so they are dividedinto two genera:Aerospiza (Roberts 1922, typetachiro)andTachyspiza.

The two oldest available names for theTachyspiza clade seem tobeTachyspiza (typesoloensis) andLeucospiza (typenovaehollandiae). Both are from the same 1844 monograph by Kaup,published in early 1844, no later than March. Another available name forthe clade is Blyth's (1844)Nisastur, typebadius, but thatseems to date from July (fide Richmond index).

NeitherTachyspiza norLeucospiza seems to have obtainedany priority over the other, and I'm somewhat arbitrarily usingTachyspiza here. However, someone will need to formally pick one inthe literature and give it priority, hopefullyTachyspiza. The nameLeucospiza really only fits the white morph ofnovaehollandiae. At present, it is unclear where exactlyErythrotriorchis fits in. It may be sister toAerospiza, ortoTachyspiza or even embedded within one of them (Barrowclough etal, 2014 is not conclusive).

WithinTachyspiza, the position of the Spot-tailed Sparrowhawk,Tachyspiza trinotata, is quite uncertain. I also have doubts aboutwhere thehenicogramma-novaehollandiae clade goes. Nagy andTökölyi (2014), apparently using the RAG-1 gene from subspeciesrufoschistaceus (hiogaster, notnovaehollandiae) putit nearimitator, but Breman et al. (2013), using the barcodingregion, put another member of the clade,fasciata, nearervirgata. There is also some uncertainty about exactly wheresoloensis goes. I've followed Kocum's (2006) multigene analysis, butNagy and Tökölyi (2014) have it in a rather differentposition.

This list includes three relatives of the Sharp-shinned Hawk thatSACC has not split (although they are split in the Sibley-Monore list).They could be allopatric forms best treated as one species. However,although the combined analysis of Kocum (2006) placedventralisanderythronemius in a cladestriatus, other arrangementswere seen for some of the other genes (chionogaster was notincluded). Breman et al. (2013) also found a separation betweenstriatus anderythronemius. This hints, but does notprove, that a species-level treatment is appropriate, as has beenfollowed by various authors.

Sharp-shinned Hawk complex:Catanach et al. (2021) found that the Sharpies of theGreater Antilles are related to the a clade consisting of North and SouthAmerican Sharp-shinned Hawks. The distances suggest species-level taxa, soI've added the three island species to the complex, bringing it to sevenspecies.

Rufous-thighed Hawk,Accipiter erythronemius
Plain-breasted Hawk,Accipiter ventralis
Sharp-shinned Hawk,Accipiter velox
White-breasted Hawk,Accipiter chionogaster
Cuban Hawk,Accipiter fringilloides
Hispaniolan Hawk,Accipiter striatus
Puerto Rican Hawk,Accipiter venator

The oldest named subspecies of the Sharp-shinned Hawk wasstriatus.As that is now the Hispaniolan Hawk, we have to turn to the next oldest namefor the Sharp-shinned Hawk,Accipiter velox. This paper has also led me to rearrange the South American species.

They also note an intriguing connection between the Rufous-thighed Hawk,Accipiter erythronemius and the Eurasian Sparrowhawk,Accipiter nisus.

The arrangement of the harriers follows Oatley et al. (2015). I have splitthe Hen and Northern Harriers. Oatley et al. found that the Northern Harrierwas closer to the Cinereous Harrier than to the Hen Harrier. Oatley et al. also found that the last 5 harriers are very closely related, possibily conspecific.However, I distrust their estimated divergence dates. They are markedly shorterthan comparable dates in Nagy and Tökölyi (2014), which already seema bit short compared to Jarvis et al. (2015).

Accipitrini: Accipiters, Harriers

Chestnut-shouldered Goshawk,Erythrotriorchis buergersi
Red Goshawk,Erythrotriorchis radiatus
Chestnut-flanked Sparrowhawk,Aerospiza castanilius
Red-chested Goshawk,Aerospiza toussenelii
African Goshawk,Aerospiza tachiro
Red-thighed Sparrowhawk,Tachyspiza erythropus
Little Sparrowhawk,Tachyspiza minulla
Spot-tailed Sparrowhawk,Tachyspiza trinotata
Japanese Sparrowhawk,Tachyspiza gularis
Besra,Tachyspiza virgata
Dwarf Sparrowhawk,Tachyspiza nana
Vinous-breasted Sparrowhawk,Tachyspiza rhodogaster
Rufous-necked Sparrowhawk,Tachyspiza erythrauchen
Collared Sparrowhawk,Tachyspiza cirrocephala
New Britain Sparrowhawk,Tachyspiza brachyura
Moluccan Goshawk,Tachyspiza henicogramma
Brown Goshawk,Tachyspiza fasciata
Variable Goshawk,Tachyspiza hiogaster
Gray Goshawk,Tachyspiza novaehollandiae
Shikra,Tachyspiza badia
Levant Sparrowhawk,Tachyspiza brevipes
Nicobar Sparrowhawk,Tachyspiza butleri
Chinese Sparrowhawk,Tachyspiza soloensis
Black-mantled Goshawk,Tachyspiza melanochlamys
Pied Goshawk,Tachyspiza albogularis
White-bellied Goshawk,Tachyspiza haplochroa
Fiji Goshawk,Tachyspiza rufitorques
Frances's Sparrowhawk,Tachyspiza francesiae
Slaty-mantled Goshawk,Tachyspiza luteoschistacea
Imitator Goshawk,Tachyspiza imitator
Gray-headed Goshawk,Tachyspiza poliocephala
New Britain Goshawk,Tachyspiza princeps
Gray-bellied Hawk,Accipiter poliogaster
Ovambo Sparrowhawk,Accipiter ovampensis
Madagascan Sparrowhawk,Accipiter madagascariensis
Eurasian Sparrowhawk,Accipiter nisus
Rufous-breasted Sparrowhawk,Accipiter rufiventris
Rufous-thighed Hawk,Accipiter erythronemius
Plain-breasted Hawk,Accipiter ventralis
Sharp-shinned Hawk,Accipiter velox
White-breasted Hawk,Accipiter chionogaster
Cuban Hawk,Accipiter fringilloides
Hispaniolan Hawk,Accipiter striatus
Puerto Rican Hawk,Accipiter venator
Doria's Goshawk,Megatriorchis doriae
Bicolored Hawk,Astur bicolor
Cooper's Hawk,Astur cooperii
Gundlach's Hawk,Astur gundlachi
Northern Goshawk,Astur gentilis
Meyer's Goshawk,Astur meyerianus
Black Sparrowhawk,Astur melanoleucus
Henst's Goshawk,Astur henstii
Spotted Harrier,Circus assimilis
Pallid Harrier,Circus macrourus
Black Harrier,Circus maurus
Hen Harrier,Circus cyaneus
Northern Harrier,Circus hudsonius
Cinereous Harrier,Circus cinereus
Long-winged Harrier,Circus buffoni
Montagu's Harrier,Circus pygargus
Pied Harrier,Circus melanoleucos
Western Marsh-Harrier,Circus aeruginosus
African Marsh-Harrier,Circus ranivorus
Swamp Harrier,Circus approximans
Papuan Harrier,Circus spilothorax
Reunion Harrier,Circus maillardi
Eastern Marsh-Harrier,Circus spilonotus
Malagasy Harrier,Circus macrosceles

Buteoninae: ButeosVigors 1824

The Buteoninae are the largest subfamily of the Accipitridae.According to Nagy and Tökölyi (2014), the clade is roughly 20million years old. There are three deep divisions (15-18 million yearsago) in the Buteoninae which I have recognized as tribes: Harpagini,Milvini, and Buteonini.

Many of the changes between versions 2.00 and 2.01 of this pagewere due to the publication of Lerner et al. (2008). Their results havesubsequently been refined by Amaral et al. (2009) and Nagy andTökölyi (2014). I have additionally consulted Amaral et al. (2006),Barrowclough et al.\ (2014), Griffiths et al.\ (2007), Kocum (2006),Kruckenhauser et al.\ (2004), Ong et al.\ (2011), and Riesing et al.(2003). Although these sources are not in 100% agreement, and there arestill a few lacuna, I think the Buteoninae are now in pretty good shapeeven at the species level.

There might still be a little modification of species boundaries togo, particularly inPseudastur and nearButeo buteo. I'vemodified the generic limits quite a bit in the Buteoninae, but lessdrastically than suggested by Riesing et al. (2003) and by Lerner et al.(2008). Most of these changes were also adopted by Amaral et al.(2009), which prompted some further changes in version 2.15. This hasbeen further refined by considering Nagy and Tökölyi (2014) inversion 2.61.

HarpaginiBonaparte, 1854

Harpagini consists of the twoHarpagus kites. Nagy andTökölyi (2014) put Harpagini in the basal position. Griffithset al. (2007) also included them in their analysis. They haveHarpagus basal to Accipitrinae + Buteoninae. However, resolutionwas poor, and other basal raptors from Griffiths et al. have moved toAccipitridae.

Double-toothed Kite,Harpagus bidentatus
Rufous-thighed Kite,Harpagus diodon

Milvini: Sea Eagles and Milvine KitesVigors 1824

Buteoninae tree Milvini consists of the milvine kites and sea and fish eagles. Thekites are the second of the four kite clades in Buteoninae.

The two-species version ofIcthyophagaturns out to be nested withinHaliaeetus. It could either besubmerged intoHaliaeetus, or expanded to include fourHaliaeetus species. I've taken the second option. This putsall of the northern fish-eagles and sea-eagles inHaliaeetus, whilethe tropical fish/sea-eagles are inIcthyophaga. The two groups are notonly geographically separated, but are visually distinct—compare therelatively longer, narrower wings ofHaliaeetus to the shorter,broader wings ofIcthyophaga. There's a spelling issue here:Icthyophaga orIchthyophaga? Both Dickinson (2003) andPeterson (zoonomen.net) give the former, which is the original spelling. Peterson also argues thatIcthyophaga is correct. Nonetheless,Ichthyophaga is also in wide usage (e.g., HBW-2).

Brahminy Kite,Haliastur indus
Whistling Kite,Haliastur sphenurus
Red Kite,Milvus milvus
Black Kite,Milvus migrans
Yellow-billed Kite,Milvus aegyptius
Pallas's Fish-Eagle,Haliaeetus leucoryphus
Steller's Sea-Eagle,Haliaeetus pelagicus
White-tailed Eagle,Haliaeetus albicilla
Bald Eagle,Haliaeetus leucocephalus
White-bellied Sea-Eagle,Icthyophaga leucogaster
Sanford's Sea-Eagle,Icthyophaga sanfordi
African Fish-Eagle,Icthyophaga vocifer
Madagascan Fish-Eagle,Icthyophaga vociferoides
Lesser Fish-Eagle,Icthyophaga humilis
Gray-headed Fish-Eagle,Icthyophaga ichthyaetus

Buteonini: ButeosVigors 1824

This brings us to Buteonini. The first branch consists ofButastur buzzards and the second branch is theIctinia kites(third of the four Buteoninae kite clades).The next clade containsBusarellus,Geranospiza, and the final kite clade,Rostrhamus and presumablyHelicolestes. Except possibly for the last two, they are all long-separated anddeserving of genus status.

There have been big changes forButeogallus. The traditionalButeogallus hawks turned out to be closely related toHarpyhaliaetus and three of theLeucopternis hawks.Prior to version 2.15, I was rolling them all intoButeogallus. Then,I followed the recommendations of Amaral et al. (2009), who create two newgenera and also slightly adjust the taxonomy. After several rounds of debate,the SACC has settled on another solution, which I follow as of version 2.55.They recognize the new genusCryptoleucopteryx (Amaral et al, 2009)for the Plumbeous Hawk (formerly inLeucopternis).Buteogallus now applies to the oldButeogallus plusSlate-colored Hawk and White-necked Hawk (fromLeucopternis) andthe twoHarpyhaliaetus solitary-eagles. Amaral et al. (2009) and Nagy and Tökölyi (2014) estimate the commonButeogallus ancestorat about 4-6 million years ago, so putting them all in one genus is reasonable.In contrast, they estimate that the originalButeogallus separated fromCryptoleucopteryx somewhere between 7 and 10 million yearsago.

This brings us to point C, one place where Lerner et al. suggest therest be consideredButeo. I think this idea hides more taxonomythan it reveals and prefer a somewhat different arrangement. As ithappens, Amaral et al. (2009) came to similar conclusions. Amaral etal. and Nagy and Tökölyi (2014) estimate thatMorphnarchus separated fromButeo about 8-11 million yearsago.Morphnarchus is anotherLeucopternis refugee, whilethe Roadside Hawk (Rupornis) is sometimes considered part ofButeo. The same is true of the White-rumped Hawk, which hasjoined Harris's Hawk inParabueto. Nagy and Tökölyi (2014)and Barrowclough et al. (2014) putRupornis in a clade withParabueto, but I believe that Amaral et al. (2009) are used moredata here and found them on sequential branches.

The clade starting at B is also flagged by Lerner et al. as a goodstarting point forButeo. The formerLeucopternis, White,Gray-backed, and Mantled Hawks are placed inPseudastur while thesome of the remainingLeucopternis are put inGeranoaetus. This includes the White-tailed, and Variable (sometimes split as Red-backed andPuna) Hawks, formerly inButeo. It should be noted that the speciesboundaries inPseudastur seriously need adjustment, but it seems thatfurther study will be needed to clarify the situation.

The clade at C is also a plausible way to delimitButeo. The SACC prefers to retainLeucopternis. The balance of evidence suggestsLeucopternis is more basal than the Gray and Gray-lined Hawks(subgenusAsturina). Amaral et al. (2009) findAsturinasister toButeo and Nagy and Tökölyi (2014) put theAsturina hawks a little deeper in the buteos. I now follow bothAOU committees by includingAsturina inButeo.

Based on Millsap et al. (2011), the Gray Hawk, formerlyAsturinanitida, has been split into Gray Hawk,Asturina plagiata andGray-lined Hawk,Asturina nitida.

As defined below,Buteo consists primarily of species breedingin the Old World and Nearctic, with a few Neotropical species. Theother genera that might be included inButeo —Leucopternis (point A),Pseudastur,Geranoaetus(point B),Parabuteo,Rupornis, andMorphnarchus(point C) — are primarily South and Middle American breeders.

Amaral et al. (2009) and Nagy and Tökölyi (2014) help clearup the situation with the remainingButeo species, although someissues remain. The New World species split out nicely, up to thelagopus/regalis pair. Then we get into the Old Worldbuteos, whose taxonomy remains somewhat murky. Two species were notconsidered in their analysis, and the true species boundaries among theOld World buteos remain somewhat uncertain. I expect that the treatmenthere will subject to a bit of revision as more is known.

Riesing et al. (2003, Fig.5) found evidence that Archer's Buzzard,Buteo archeri is not part of the Augur Buzzard,Buteoaugur, but is sister to a clade consisting of the Red-necked, Augurand Jackal Buzzards, and possibly the Madagascan Buzzard,Buteobrachypterus. Other evidence in Riesing et al. (Fig.4), and in Nagyand Tökölyi (2014) putsbrachypterus in a slightly morebasal position, out of this clade.

Another apparently misplaced buteo was the Forest Buzzard,Buteotrizonatus. It was considered a subspecies of the Mountain Buzzard,Buteo oreophilus, but genetic data (esp. Kruckenhauser et al., 2004)suggests it is closer to the Common Buzzard,Buteo buteo.Although the genetic distance is small, its distant separation from theCommon Buzzard's breeding range and distinct plumage suggest it shouldbe considered a distinct species.

Although the breeding buzzards of the Cape Verde and Socotra Islandshave been considered races of the Common Buzzard, genetic data (Clouetand Wink, 2000) suggests they are more closely related to theLong-legged Buzzard,Buteo rufinus. Once again, the geneticdistances are small, but this suggests treating them as distinctspecies: Cape Verde Buzzard,Buteo bannermani, and SocotraBuzzard,Buteo socotraensis. Oddly, although it had been studiedfor over a century, the Socotra Buzzard had not been formally nameduntil 2010 when Porter and Kirwan dubbed itButeo socotraensis.This replaces the informal name ‘B. socotrae’ that some haveused.

Grasshopper Buzzard,Butastur rufipennis
White-eyed Buzzard,Butastur teesa
Rufous-winged Buzzard,Butastur liventer
Gray-faced Buzzard,Butastur indicus
Mississippi Kite,Ictinia mississippiensis
Plumbeous Kite,Ictinia plumbea
Black-collared Hawk,Busarellus nigricollis
Crane Hawk,Geranospiza caerulescens
Snail Kite,Rostrhamus sociabilis
Slender-billed Kite,Helicolestes hamatus
Plumbeous Hawk,Cryptoleucopteryx plumbea
Slate-colored Hawk,Buteogallus schistaceus
Rufous Crab Hawk,Buteogallus aequinoctialis
Common Black Hawk,Buteogallus anthracinus
Cuban Black Hawk,Buteogallus gundlachii
Savanna Hawk,Buteogallus meridionalis
White-necked Hawk,Buteogallus lacernulatus
Great Black Hawk,Buteogallus urubitinga
Solitary Eagle,Buteogallus solitarius
Chaco Eagle,Buteogallus coronatus
Barred Hawk,Morphnarchus princeps
Roadside Hawk,Rupornis magnirostris
Harris's Hawk,Parabuteo unicinctus
White-rumped Hawk,Parabuteo leucorrhous
White-tailed Hawk,Geranoaetus albicaudatus
Variable Hawk,Geranoaetus polyosoma
Black-chested Buzzard-Eagle,Geranoaetus melanoleucus
Gray-backed Hawk,Pseudastur occidentalis
White Hawk,Pseudastur albicollis
Mantled Hawk,Pseudastur polionotus
Semiplumbeous Hawk,Leucopternis semiplumbeus
Black-faced Hawk,Leucopternis melanops
White-browed Hawk,Leucopternis kuhli
Gray Hawk,Buteo plagiatus
Gray-lined Hawk,Buteo nitidus
Red-shouldered Hawk,Buteo lineatus
Ridgway's Hawk,Buteo ridgwayi
Broad-winged Hawk,Buteo platypterus
Hawaiian Hawk,Buteo solitarius
White-throated Hawk,Buteo albigula
Short-tailed Hawk,Buteo brachyurus
Galapagos Hawk,Buteo galapagoensis
Swainson's Hawk,Buteo swainsoni
Zone-tailed Hawk,Buteo albonotatus
Red-tailed Hawk,Buteo jamaicensis
Rufous-tailed Hawk,Buteo ventralis
Rough-legged Hawk / Rough-legged Buzzard,Buteo lagopus
Ferruginous Hawk,Buteo regalis
Madagascan Buzzard,Buteo brachypterus
Archer's Buzzard,Buteo archeri
Red-necked Buzzard,Buteo auguralis
Augur Buzzard,Buteo augur
Jackal Buzzard,Buteo rufofuscus
Eastern Buzzard,Buteo japonicus
Upland Buzzard,Buteo hemilasius
Himalayan Buzzard,Buteo burmanicus
Mountain Buzzard,Buteo oreophilus
Common Buzzard,Buteo buteo
Forest Buzzard,Buteo trizonatus
Long-legged Buzzard,Buteo rufinus
Cape Verde Buzzard,Buteo bannermani
Socotra Buzzard,Buteo socotraensis

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AFROAVES II

ACCIPITRIMORPHAE: Cathartiformes, Accipitriformes

Afroaves

Coliiformes

Accipitrimorphae

Strigiformes

Picimorphae

Leptosomiformes

Trogoniformes

Bucerotiformes

Coraciiformes

Piciformes

ACCIPITRIMORPHAEVieillot, 1816

CATHARTIFORMESCoues, 1884

Cathartidae: New World Vulturesde Lafresnaye, 1839

5 genera, 7 speciesHBW-2

ACCIPITRIFORMESVieillot, 1816

Sagittariidae: SecretarybirdFinsch & Hartlaub, 1870 (1825)

1 genus, 1 speciesHBW-2

Pandionidae: OspreyBonaparte, 1854

1 genus, 1 speciesHBW-2

Accipitridae: Hawks, Kites, EaglesVigors, 1824

73 genera, 255 speciesHBW-2

Elaninae: Elanine KitesBlyth, 1851

Gypaetinae: African Vultures, Harrier-HawksBonaparte, 1831

Perninae: Bazas, Honey-BuzzardsBlyth, 1851

Gypinae: Old World VulturesBlyth, 1851

Circaetinae: Serpent EaglesBlyth, 1851

Lophospizinae: Crested GoshawksInformal

Harpiinae: Harpy EaglesLesson, 1828

Aquilinae: Booted EaglesVigors, 1825

Accipitrinae: Accipiters, HarriersVigors, 1824

Melieraxini: Chanting Goshawks and alliesInformal

Notes on Accipitrini: Accipiters & Harriers

Accipitrini: Accipiters, Harriers

Buteoninae: ButeosVigors 1824

HarpaginiBonaparte, 1854

Milvini: Sea Eagles and Milvine KitesVigors 1824

Buteonini: ButeosVigors 1824