The 'Caprimulgiformes' is an
order of
birds that includes a number of
birds with global distribution (except
Antarctica). They are generally
insectivorous and
nocturnal. The order gets its name from the Latin for "goat-sucker", an old name based on an erroneous view of the
European Nightjar's feeding habits.
Systematics
The classification of the various birds that make up the order has long been controversial and difficult, particularly in the case of the nightjars. All things considered, the nightjar order would probably best be limited to potoos, nightjars, and eared-nightjars, all other lineages being elevated to
order level, and the
owlet-nightjars being altogether distant:
★ Family Steatornithidae (
Oilbird) - probably distinct order N.N. ("Steatornithiformes"
[1])
★ Family Podargidae (
frogmouths, 12 species in 2 genera) - probably distinct order Podargiformes
★ Family
Nyctibiidae (Potoos, about 5 species in 1 genus)
★ Family
Caprimulgidae
★
★ Subfamily
Chordeilinae (New World nighthawks)
★
★ Subfamily
Caprimulginae (typical nightjars)
★ Family
Eurostopodidae (eared-nightjars)
Traditionally, they were regarded, on morphological grounds, as being midway between the
owls (Strigiformes) and the
swifts. Like the owls, they are nocturnal hunters with a highly developed sense of sight, and like the swifts they are excellent flyers with small, weak legs. At one time or another, they have allied with owls, swifts,
kingfishers,
hoopoes,
mousebirds,
hornbills,
rollers,
bee-eaters,
woodpeckers,
trogons and
hummingbirds.
Based on analysis of
sequence data - notably β-
fibrinogen intron 7 -, Fain & Houde (2004) considered the families of the Caprimulgiformes to be members of the proposed clade
Metaves, which also includes the
hoatzin,
tropicbirds,
sandgrouse,
pigeons,
kagu,
sunbittern,
mesites,
flamingos,
grebes and
swifts and
hummingbirds. This clade was also found by the expanded study of Ericson ''et al.'' (2006), but support was extremely weak.
While only the latter study recovered monophyly of the
Cypselomorphae (see below) within Metaves, the former was based on only a single locus and could not resolve their relationships according to standard criteria of statistical confidence. No robust
apomorphies have been found that unite Metaves, except a similarity in the β-fibrinogen DNA sequence, for which there is some indication that it may be due to chance
convergent evolution. Ericson ''et al.'' (2006) concluded that if valid, the "Metaves" must originate quite some time before the
Paleogene, but this is rather hard to reconcile with the fossil record.
While the relationships of cypselomorphs are a subject of ongoing debate, the
phylogeny of the individual lineages is better resolved. Much of the remaining uncertainty regards minor details.
Initial
mtDNA cytochrome ''b'' sequence analysis (Mariaux & Braun 1996) agreed with earlier morphological (Cracraft 1981) and
DNA-DNA hybridization (Sibley & Ahlquist 1990) studies insofar as that the oilbird and the frogmouths seemed rather distinct. The other lineages appeared to form a
clade, but this is now known to have been caused by methodological limitiations.
The Aegothelidae (
owlet-nightjars) with about a dozen living species in one genus are apparently closer to the
Apodiformes (Mayr 2002); these and the Caprimulgiformes are closely related, being grouped together as
Cypselomorphae. The oilbird and the frogmouths seem quite distinct among the remaining Caprimulgiformes, but their exact placement cannot be resolved based on osteological data alone (Mary 2002).
Even the study of Ericson ''et al.'' could not properly resolve the oilbird's and frogmouths' relationships beyond the fact that they are quite certainly well distinct. It robustly supported, however, the idea that the owlet-nightjars should be considered closer to Caprimulgiformes, unlike the methodologically weaker studies of Mariaux & Braun (1996) and Fain & Houde (2004).
Alternatively, Mayr's
phylogenetic taxon Cypselomorphae might be placed at order rank and substitute the two present orders Caprimulgiformes and Apodiformes. Such a group would be fairly uninformative as regards its evolutionary history, as it has to include some very
plesiomorphic and some extremely derived lineages (such as hummingbirds) to achieve monophyly.
Evolution
The
fossil record of caprimulgiform birds (in the loose sense) is rather scant. Nonetheless, it supports the emerging consensus phylogeny well. The genus ''
Paraprefica'', probably from the
Early Eocene (though this is somewhat uncertain), seems to be a
basal form that at times has been allied with the oilbird and the potoos, but cannot be assigned to either with certainty. In the consensus scenario, it would represent a record of the initial divergence of the three lineages.
This nicely agrees with fossils suggesting that the basal divergence of the owlet-nightjar and apodiform branch also occurred during that time. In addition, ''
Eocypselus'', a
Late Paleocene or Early
Eocene genus of north-central Europe, cannot be assigned to any one cypselomorph lineage with certainty but appears to be some ancestral form.
These
Paleogene birds strongly suggest that the 2 main extant lineages of cypselomorphs separated about 60-55
mya (
Selandian-
Thanetian), and that some time around the
Lutetian-
Bartonian boundary, some 40 mya, the common ancestors of Nyctibiidae, Caprimulgidae and Eurostopodidae diverged from those of oilbird and frogmouths.
References
★ (1981): Toward a phylogenetic classification of the recent birds of the world (Class Aves). ''
Auk'' '98'(4): 681–714.
PDF fulltext
★ (2006): Diversification of Neoaves: integration of molecular sequence data and fossils. ''
Biol. Lett.'' '2'(4): 543-547.
PDF fulltext
★ (2004): Parallel radiations in the primary clades of birds. ''
Evolution'' '58'(11): 2558-2573.
PDF fulltext
★ (2002): Osteological evidence for paraphyly of the avian order Caprimulgiformes (nightjars and allies). ''Journal für Ornithologie'' '143'(1): 82–97.
HTML abstract
★ (1996): A Molecular Phylogenetic Survey of the Nightjars and Allies (Caprimulgiformes) with Special Emphasis on the Potoos (Nyctibiidae). ''Molecular Phylogenetics and Evolution'' '6'(2): 228–244.
(HTML abstract)
★ (1990): ''Phylogeny and classification of birds''. Yale University Press, New Haven, Conn.
Footnotes
1.
Apparently not a valid taxon (yet), and thus it must not be written without quotation marks.