Background: Threnetes leucurus (Pale-tailed Barbthroat) with a range through most of Amazonia, has been long considered a separate species from Threnetes niger (Sooty Barbthroat), found in French Guiana and adjacent Brazil (Amapa). Grantsau (1969) described a new species in this complex, Threnetes loehkeni from Amapá. Hinkelmann (1988) suggests that Threnetes loehkeni is best treated as a subspecies of leucurus, but at that time does not include Threnetes niger in the mix. In this paper, he also nixes a couple of Ruschi taxa (Threnetes cristinae and T. niger freiri described from specimens from Amapá). In HBW, Hinkelmann (1999) lumps all of the Amazonian Threnetes into one species under the oldest name niger. He states "niger represents a melanistic form of geographically limited distribution, based on apomorphic characters within the range of variation found in leucurus; otherwise the two are apparently inseparable." Because of Hinkelmann's uncertainty about Threnetes niger and the fact that Hinkelmann and Schuchmann (1997) did not correctly apply the names in this case, the taxonomic issues did not really become generally appreciated until the publication of HBW, Vol. 5.
Analysis:
The taxon niger is as Hinkelmann states a melanistic bird.
I have seen two specimens, one at the Field Museum, and one in
Sao Paulo. On these specimens, the basic Threnetes plumage
pattern is visible underlying the dark coloration of the underparts.
The basic plumage pattern is that of loehkeni. This led
Parker et al (1996) to suggest that loehkeni was probably
a pale morph of niger. The logic of Hinkelmann's treatment
of loehkeni and leucurus as conspecific with niger,
is not made explicit, but seems to be based on loehkeni as
a subspecies of leucurus and niger being a melanistic
population allied to one or the other of these (see Hinkelmann
and Schuchmann 1997 for the most explicit discussion, although
they mess up the name there).
It looks to me like Hinkelmann is correct that niger and
loehkeni are basically melanistic and normal morphs of the
same thing. It isn't completely clear that whether there is a
geographical element to this, so that they should be considered
two subspecies, or whether they are morphs within the same subspecies.
In HBW, Hinkelmann takes the former view. In Hinkelmann and Schuchmann
(1997) they suggest that the latter view is more likely. The question
is whether the treatment of niger plus loehkeni
as conspecific with the widespread leucurus is appropriate.
Hinkelmann (1988) suggests that the plumage characters that Grantsau
(1969) mentions for loehkeni are invalid, except for the
tail pattern. In loehkeni (and in niger), the tail
is dark green, with very narrow pale tips to the outer four rectrices
on each side. In leucurus, the outer four tail features
are mostly pale (ranging from white to ochraceous depending on
the subspecies), with a black subterminal band followed by pale
tips. The pale on the tail is broadest on the innermost of the
four outer feathers, so nearly completely pale, and gets progressively
more black as you move outward, so that the outermost feather
is about 50% black. The result is a very different and distinctive
tail pattern from that of niger and loehkeni (the
plate in HBW does a good job on these birds).
Hinkelmann (1988) treats loehkeni as a "distinctive
subspecies" of leucurus, stating "the colouration
of loehkeni does not go beyond the variation found
in all subspecies of T. leucurus, variation which is probably
due to age and sex." I would agree with Hinkelmann on this,
except for the tail pattern. Although females and young birds
of leucurus have less white in their tails than do adult
males, in the 100 +/- specimens of leucurus at the Field
Museum, none approach the condition of loehkeni.
Whether to treat niger as a distinct species from leucurus
or as conspecific is difficult, however, I think that recognizing
two species is the best course. The two taxa approach one another
moderately closely, and may occur together. Typical leucurus
occurs in Surinam and southern Amapá, at least (Hinkelmann
1988). Tostain et al. (1992) list both species from French Guiana,
and gives both from two localities, St. Georges and Saul. However,
since they do not give subspecies, I don't know whether the leucurus
may refer to loehkeni rater than nominate. As an aside
I would note that the range of niger within French Guiana
includes most of the border with Surinam, so it should occur in
Surinam, but I a unaware of any records.
Given the close approach of niger and leucurus and
the possibility of sympatry without any indication of intermediate
tail-patterned birds being found, and the lack of any reports
of melanistic birds in leucurus from near the range of
niger, I would argue that there is no evidence of intergradation
occurring between niger and leucurus. Further, the
other species in the genus, Threnetes ruckeri, allopatric
to these birds in Trans-Andean forests, is very similar to leucurus
in plumage, except for its distinctive tail pattern. The tail
pattern of niger is even more distinctive than that of
ruckeri in my view.
Proposed change: Recognize Threnetes leucurus as a distinct species from Threnetes niger, treating loehkeni as a subspecies of niger. The English name Pale-tailed Barbthroat would apply to leucurus. Threnetes niger has been called Sooty Barbthroat, except when treated as conspecific with leucurus. This name is not particularly appropriate when loehkeni is lumped into it. Loehken's Barbthroat seems inappropriate as well. Maybe Dark-tailed Barbthroat?
Recommendation:
I recommend treating Threnetes niger as a distinct species
from Threnetes leucurus, treating loehkeni as a
subspecies of niger.
References:
Grantsau, R. 1969. Uma nova espécie de Threnetes
(Aves, Trochilidae). Pap. Avuls. Zool. 22:245-247.
Hinkelmann, C. 1988. Comments on recently described new species of hermit hummingbirds. BBOC 108:159-169.
Hinkelmann, C. 1999. Pale-tailed Barbthroat
(Threnetes niger) [species account], in del Hoyo,
J., Elliot, A. & J. Sargatel, eds. Handbook of the Birds of
the World. Vol. 5. Page 539.
[How should you cite species accounts in this?]
Hinkelmann, C. and K. Schuchmann. 1997. Phylogeny of the hermit hummingbirds (Trochilidae: Phaethornithinae). Studies on Neotropical Fauna and Environments 32:142-163.
Parker, T. A., III, D.F. Stotz, and J. W. Fitzpatrick. 1996. Ecological and Distributional Databases, in Stotz, D. F., J. W. Fitzpatrick, T. A. Parker III, and D. K. Moskovits, Neotropical Birds: Ecology and Conservation. University of Chicago, Chicago.
Tostain, O., J.-L. Dujardin, Ch. Érard,
and J.-M Thiollay. Oiseaux de Guyane. Société d'Etudes
Ornithologiques, Brunoy, France.
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Comments from Robbins: "[YES.] Doug has done a good job of providing details to the convoluted treatment of the T. niger complex."
Comments from Stiles: "NO. Sounds reasonable but lacks solid published evidence - if Doug were to publish something on this, I would probably go for it."
Comments from Zimmer: "A reluctant "NO". I think Doug's arguments are persuasive, but once again, I think there needs to be some published analysis, particularly since the historical treatments have been so muddled."
Comments from Jaramillo: "YES although reluctantly. A quick note stating what has been stated here with illustrations of tail pattern and a map would be highly encouraged."
Comments from Nores: "YES. Acepto reconocer a Threnetes leucurus como una especie distinta de Threnetes niger y tratar a loehkeni como una subspecie de niger. Pienso que el patrón de la cola de leucurus es concluyente."
Comments from Remsen: "YES. In retrospect, and after reading the information assembled by Doug, it seems premature in my opinion to consider leucurus and niger conspecific -- the burden-of-proof in this case falls on the treatment that would consider them conspecific."