CYPSELOMORPHAE Huxley, 1867
The biggest chunk of Metaves is the Cypselomorphae. Although it is tempting to group all of the remaining nightbird families in a single order, it is hard to find convincing evidence that they are even constitute a clade (e.g., Barrowclough et al., 2006; Braun and Huddleston (2009), Mariaux and Braun, 1996; Mayr, 2002). The divisions between them are very deep, and DNA studies have not only not come to a clear conclusion about the 4 orders are related, but many have not even shown they are each other's closest relatives! The oilbirds are particularly problematic. Because of this, I have listed them in separate orders. Hackett et al. (2008) and Ericson et al. (2006a) are two of the few genetic analyses that place them all in the same clade, which is augmented by the Adopiformes. The Apodiformes may be found on the next page, with the remainder of the Cypselomorphae on this one.
In fact, the monophyly of the Cypselomorphae is only supported by analyses that include β-fibrinogen, which also support the Metaves hypothesis. When β-fibrinogen was excluded in Ericson et al. (2006a), the Cypselomorphae ended up in the Anomalogonatae. Moreover, the nighthawks did not group with the other Cypselomorphae, being closer to a clade containing the owls and others. One of Braun and Huddleston (2009) analyses also placed the nighthawks closer to the owls.
The orders are arranged as in Hackett et al. (2008). However, the evidence for this is rather weak. Recent morphological studies (e.g. Mayr, 2010; Nesbitt et al., 2011) suggest that the Steatornithiformes and Podargiformes are either basal branches or a sister clade to the remaining 3 orders.
STEATORNITHIFORMES Sharpe 1891
The results of both Hackett et al. (2008) and Braun and Huddleston (2009) suggest the oilbirds and potoos are sisters, which makes sense geographically. Note however, that oilbirds were once more widely distributed. An oilbird fossil is known from Wyoming (Olson, 1987).
Steatornithidae: Oilbird Bonaparte, 1842
1 genus, 1 species HBW-5
- Oilbird, Steatornis caripensis
No one seems to have recongized the Nyctibiiformes at an ordinal level before. I've been using it on these webpages since at least March 2008. If had I published it in print instead of a web page I could have my name on it!
Nyctibiidae: Potoos Chenu & des Murs, 1851
1 genus, 7 species HBW-5
- Rufous Potoo, Nyctibius bracteatus
- Great Potoo, Nyctibius grandis
- Long-tailed Potoo, Nyctibius aethereus
- Common Potoo, Nyctibius griseus
- Northern Potoo, Nyctibius jamaicensis
- Andean Potoo, Nyctibius maculosus
- White-winged Potoo, Nyctibius leucopterus
PODARGIFORMES Sharpe 1891
The Frogmouths are sometimes divided into two families, Podargidae and Batrachostomidae. Molecular evidence indicates they are each other's closest relatives, and I think that is best indicated by placing them in a single family. One new species, the Solomon Islands Frogmouth, was recently discovered (Cleere et al., 2007). It seems to be closer to Podargus than to Batrachostomus.
Following Cleere (2010), Blyth's Frogmouth, Batrachostomus affinis and Palawan Frogmouth, Batrachostomus chaseni) are separated from Javan Frogmouth, Batrachostomus javensis, based on differences in vocalizations. Although Cleere separates it as a separate species, there seems to be some question concerning the vocalizations of the race continentalis (see Marshall, 1978). Until that is clarified, preferably in a journal article, I continue to include it as a subspecies of Blyth's Frogmouth, Batrachostomus affinis.
Although the extant frogmouths are restricted to the Oriental and Australasian regions, they were not always so. Frogmouth fossils have been found in both Europe and the United States (Nesbitt et al., 2011).
Podargidae: Frogmouths G.R. Gray, 1847
3 genera, 16 species HBW-5
- Solomons Frogmouth, Rigidipenna inexpectata
- Marbled Frogmouth, Podargus ocellatus
- Papuan Frogmouth, Podargus papuensis
- Tawny Frogmouth, Podargus strigoides
- Large Frogmouth, Batrachostomus auritus
- Dulit Frogmouth, Batrachostomus harterti
- Philippine Frogmouth, Batrachostomus septimus
- Gould's Frogmouth, Batrachostomus stellatus
- Sri Lanka Frogmouth, Batrachostomus moniliger
- Hodgson's Frogmouth, Batrachostomus hodgsoni
- Short-tailed Frogmouth, Batrachostomus poliolophus
- Bornean Frogmouth, Batrachostomus mixtus
- Javan Frogmouth, Batrachostomus javensis
- Blyth's Frogmouth, Batrachostomus affinis
- Palawan Frogmouth, Batrachostomus chaseni
- Sunda Frogmouth, Batrachostomus cornutus
CAPRIMULGIFORMES Ridgway, 1881
The thesis by Han (2006) and the follow-up paper (Han et al., 2010) have done much to clarify the situation within the Caprimulgidae. It was clear from previous work (Mariaux and Braun, 1996; Barrowclough et al. 2006; Larsen et al. 2007) that Caprimulgus itself was paraphyletic and that the Caprimulgidae were not simply divided into Nighthawks and Nightjars. It also seemed that the Eared-Nightjars are completely separate from Caprimulgidae. Han sampled many more taxa, and has shown the general shape of the family, and provided evidence against separating some species into a separate family Eurostopodiae. Indeed, looking at Han's results, it does not appear that the wider Caprimulgidae can even be usefully divided in subfamilies. Such a division would not be particularly informative.
Han found that the Great Eared-Nightjar, formerly placed in Eurostopdus is actually closer to the rest of Caprimulgidae. The Malaysian Eared-Nightjar is morphologically similar, and we are guessing that it is closely related to the Great Eared-Nightjar. Both are moved to the genus Lyncornis. The Papuan Nightjar has sometimes been placed in Lyncornis, but it is more closely related to the other Eurostopodus. Since it's thought to be close to the Cloud-forest and Satanic Nightjars, I leave them in Eurostopodus too.
The next surprise is the Collared Nightjar, formerly Caprimulgus enarratus, which seems to be sister to the remaining nightjars and nighthawks. Han et al. (2010) established the new genus Gactornis for it.
The remaining Caprimulgidae seem to fall into 4 groups, a group of American nightjars, a group of nighthawks and American nightjars, the remaining nighthawks, and a complex group of Old World nightjars. Han (2006) examined 3 genes, while Barrowclough et al. (2006) examined another. There doesn't seem to be consensus on how these four groups fit together, and as you can see on the diagram, I am currently treating it as a unresolved polytomy.
The ordering is based on the overall trees in Han (2006) and Han et al. (2010), where the upper groups successively branch off.
The first group consists of Siphonorhis, Nyctiphrynus, Phalaenoptilus, and the possibly unfamiliar genus Antrostomus. This name, due to Bonaparte (1838, type species A. carolinensis), is being applied to one of the New World Caprimulgus nightjars clades. As you can see, it is rather separated from the main group of Caprimulgus.
I have split the Whip-poor-will into Mexican Whip-poor-will, A. arizonae, and Eastern Whip-poor-will, A. vociferus. The Mexican Whip-poor-will does not appear to even be sister to A. vociferus, being closer to the Dusky Nightjar, Antrostomus saturatus. Further, it is distinctive vocally and differs somewhat in plumage from the Eastern Whip-poor-will, having often been viewed as a separate species (e.g. Howell and Webb, 1995; Navarro-Sigüenza and Peterson, 2004).
The second group contains Lurocalis through Hydropsalis. I had been using different generic limits while the AOU's South American Checklist Committee deliberated on how to best incorporate Han et al. (2010). See SACC proposals #465, #501, and #522. Now that they have decided, I am following the generic limits they've established.
The group starts out with two basal nighthawk genera, Lurocalis and Nyctiprogne. This is followed by an expanded Nyctidromus (gaining the Scrub Nightjar from Caprimulgus) together with Nyctipolus (Ridgway 1912, type nigrescens), which includes the Blackish and Pygmy Nightjars.
The remaining clade starts with 4 former Caprimulgus nightjars: Todd's, Little, Roraiman, and Cayenne. These are placed in the genus Setopagis (Ridgway 1912, type parvulus). It's followed by a subgroup containing the Band-winged Nightjar, now in Systellura (Ridgway 1912, type longirostris) and the Sickle-winged and White-winged Nightjars. The latter two are placed in Eleothreptus (GR Gray 1840, type anomalus), with the White-winged moved from Caprimulgus. Then come the two Uropsalis nightjars, the Swallow-tailed and Lyre-tailed. The genus Uropsalis was named by Miller in 1915 with type lyra. The last four species in the group go in Hydropsalis (Wagler 1832, type torquata).
The third group consists of the Chordeiles nighthawks and the Nacunda Nighthawk, Podager nacunda. It's not clear whether the Lesser Nighthawk belongs in Podager or Chordeiles. It doesn't affect the linear order. One option is to put all of them in the same genus. In that case, I think Chordeiles has priority. It was established by Swainson in part II of the “Fauna Boreali-Americana” (Richardson and Swainson), dated 1831, but apparently published Feb. 4, 1832. The genus Podager is due to Wagler from heft 3 of Isis von Oken in 1832, which is apparently the March issue. That's pretty close to Feb. 4, and some issues of Isis von Oken seem to have been published prior to their cover date. So far, I have been unable to confirm when it was actually published.
I've put the Brown Nightjar, Veles binotatus, in group four on the grounds that is a Old World species sometimes considered part of Caprimulgus. However, others have put it elsewhere (e.g., closer to Chordeiles or Lyncornis), which is also reflected in this arrangement. Han (2006) found considerable structure in the remaining Caprimulgus. As a general rule, the Asian species appear to be relatively basal in Caprimulgus. This is not reflected in the current arrangement of the species, which is fairly conventional. Note that Macrodipteryx needed to be submerged into Caprimulgus.
Finally, ABA and AOU use the name Gray Nightjar for Caprimulgus indicus. Here C. jotaka has been split from C. indicus and keeps the name Gray Nightjar while C. indicus becomes Jungle Nightjar.
Caprimulgidae: Nightjars, Nighthawks Vigors, 1825
20 genera, 94 species HBW-5
- Spotted Nightjar, Eurostopodus argus
- White-throated Nightjar, Eurostopodus mystacalis
- Satanic Nightjar, Eurostopodus diabolicus
- Papuan Nightjar, Eurostopodus papuensis
- Archbold's Nightjar, Eurostopodus archboldi
- Malaysian Eared-Nightjar, Lyncornis temminckii
- Great Eared-Nightjar, Lyncornis macrotis
- Collared Nightjar, Gactornis enarratus
- Jamaican Pauraque / Jamaican Poorwill, Siphonorhis americana
- Least Pauraque / Least Poorwill, Siphonorhis brewsteri
- Choco Poorwill, Nyctiphrynus rosenbergi
- Eared Poorwill, Nyctiphrynus mcleodii
- Yucatan Poorwill, Nyctiphrynus yucatanicus
- Ocellated Poorwill, Nyctiphrynus ocellatus
- Common Poorwill, Phalaenoptilus nuttallii
- Eastern Whip-poor-will, Antrostomus vociferus
- Puerto Rican Nightjar, Antrostomus noctitherus
- Mexican Whip-poor-will, Antrostomus arizonae
- Dusky Nightjar, Antrostomus saturatus
- Buff-collared Nightjar, Antrostomus ridgwayi
- Tawny-collared Nightjar, Antrostomus salvini
- Yucatan Nightjar, Antrostomus badius
- Silky-tailed Nightjar, Antrostomus sericocaudatus
- Chuck-will's-widow, Antrostomus carolinensis
- Rufous Nightjar, Antrostomus rufus
- Greater Antillean Nightjar, Antrostomus cubanensis
- Short-tailed Nighthawk, Lurocalis semitorquatus
- Rufous-bellied Nighthawk, Lurocalis rufiventris
- Band-tailed Nighthawk, Nyctiprogne leucopyga
- Bahian Nighthawk / Plain-tailed Nighthawk, Nyctiprogne vielliardi
- Common Pauraque / Pauraque, Nyctidromus albicollis
- Scrub Nightjar / Anthony's Nightjar, Nyctidromus anthonyi
- Blackish Nightjar, Nyctipolus nigrescens
- Pygmy Nightjar, Nyctipolus hirundinaceus
- Todd's Nightjar, Setopagis heterurus
- Little Nightjar, Setopagis parvulus
- Roraiman Nightjar, Setopagis whitelyi
- Cayenne Nightjar, Setopagis maculosus
- Band-winged Nightjar, Systellura longirostris
- White-winged Nightjar, Eleothreptus candicans
- Sickle-winged Nightjar, Eleothreptus anomalus
- Long-trained Nightjar, Macropsalis forcipata
- Swallow-tailed Nightjar, Uropsalis segmentata
- Lyre-tailed Nightjar, Uropsalis lyra
- Spot-tailed Nightjar, Hydropsalis maculicaudus
- White-tailed Nightjar, Hydropsalis cayennensis
- Ladder-tailed Nightjar, Hydropsalis climacocerca
- Scissor-tailed Nightjar, Hydropsalis torquata
- Nacunda Nighthawk, Chordeiles nacunda
- Least Nighthawk, Chordeiles pusillus
- Sand-colored Nighthawk, Chordeiles rupestris
- Lesser Nighthawk, Chordeiles acutipennis
- Common Nighthawk, Chordeiles minor
- Antillean Nighthawk, Chordeiles gundlachii
- Brown Nightjar, Veles binotatus
- Red-necked Nightjar, Caprimulgus ruficollis
- Jungle Nightjar, Caprimulgus indicus
- Gray Nightjar, Caprimulgus jotaka
- European Nightjar, Caprimulgus europaeus
- Sombre Nightjar, Caprimulgus fraenatus
- Rufous-cheeked Nightjar, Caprimulgus rufigena
- Egyptian Nightjar, Caprimulgus aegyptius
- Sykes's Nightjar, Caprimulgus mahrattensis
- Vaurie's Nightjar, Caprimulgus centralasicus
- Nubian Nightjar, Caprimulgus nubicus
- Golden Nightjar, Caprimulgus eximius
- Jerdon's Nightjar, Caprimulgus atripennis
- Large-tailed Nightjar, Caprimulgus macrurus
- Mees's Nightjar, Caprimulgus meesi
- Andaman Nightjar, Caprimulgus andamanicus
- Philippine Nightjar, Caprimulgus manillensis
- Sulawesi Nightjar, Caprimulgus celebensis
- Donaldson Smith's Nightjar, Caprimulgus donaldsoni
- Black-shouldered Nightjar, Caprimulgus nigriscapularis
- Fiery-necked Nightjar, Caprimulgus pectoralis
- Montane Nightjar, Caprimulgus poliocephalus
- Ruwenzori Nightjar, Caprimulgus ruwenzorii
- Indian Nightjar, Caprimulgus asiaticus
- Madagascan Nightjar, Caprimulgus madagascariensis
- Swamp Nightjar, Caprimulgus natalensis
- Nechisar Nightjar, Caprimulgus solala
- Plain Nightjar, Caprimulgus inornatus
- Star-spotted Nightjar, Caprimulgus stellatus
- Savanna Nightjar, Caprimulgus affinis
- Freckled Nightjar, Caprimulgus tristigma
- Bonaparte's Nightjar, Caprimulgus concretus
- Salvadori's Nightjar, Caprimulgus pulchellus
- Prigogine's Nightjar, Caprimulgus prigoginei
- Bates's Nightjar, Caprimulgus batesi
- Long-tailed Nightjar, Caprimulgus climacurus
- Slender-tailed Nightjar, Caprimulgus clarus
- Square-tailed Nightjar, Caprimulgus fossii
- Standard-winged Nightjar, Caprimulgus longipennis
- Pennant-winged Nightjar, Caprimulgus vexillarius