Proposal (472) to South American Classification Committee
Recognize
Thalurania nigricapilla as a valid
species
The taxonomy of the trans-Andean
members of the genus Thalurania has
long been somewhat controversial. In NW
Colombia and adjacent areas, two putative groups of taxa are now recognized as
distinct species: the purple-crowned (in adult males) colombica group and the green-crowned fannyi group, based upon the study of Escalante & Peterson
(1992). Their study involved only adult
males, and the character that separated the groups completely was precisely
crown color. However, other characters,
particularly those of the females, may not support this split, as noted by
Schuchmann (1999), who cautioned against “excessive reliance on crown color” in
the classification of this group (with which I agree). Therefore, it is somewhat ironic that Valdés
and Schuchmann (2009) have now described a new species in this group, based
mainly upon crown color!
This
putative new species is described as Thalurania
nigricapilla on the basis of two male specimens. The type locality is the
vicinity of Lake Calima (actually a reservoir) on the Pacific slope of the
Western Andes of Colombia, some 50 km N of Cali at an elevation of ca. 1300
m. The paratype is from Alto del Toro
near Yotoco at 1500 m, near the continental divide above the reservoir. Two taxa of the fannyi group are known from this general region: nominate fannyi from the lowlands and foothills
of the Pacific slope, and subtropicalis from
the eastern slope of the Western Andes and the western slope of the Central
Andes (both sides of the upper Cauca valley) and, according to their paper,
extending locally onto the Pacific slope NW of Cali, occupying higher
elevations than fannyi. The two forms of fannyi are stated to differ in the more bronzy, less blackish nape
and occiput and shorter, less deeply forked tail of subtropicalis compared to fannyi.
Their new species is stated to
differ from subtropicalis (and fannyi, although all detailed
comparisons in their paper involve only subtropicalis)
in the lack of a glittering green crown; both are described as having the crown
and forehead as black, when viewed frontally.
No female specimens are mentioned, although they describe a male of the
putative species displaying to a female.
They also compared the tail shape of the new form to that of subtropicalis, stating that it is longer
and more deeply forked.
There are two reasons to question whether T. nigricapilla is a valid taxon. One is a series of specimens in the ICN
labeled “Restrepo, Riobravo, Embalse de Calima, 800-850 m,” and the second is
the possibility that the specimens of nigricapilla
actually might represent old first-year males of subtropicalis – or fannyi. I have been informed that the expedition
of the Instituto de Ciencias Naturales expedition to the Embalse de Calima in
February 1984 had its base camp close to the reservoir itself, where the
collecting was conducted; if so, because the reservoir itself is between 1200
and 1300 m elevation, the elevations on the specimen labels must be erroneous
(the reservoir is in the jurisdiction of the town of Riobravo, which is at a
lower elevation, so perhaps the elevations refer to the town and not the
collecting site). In any case, it is the
plumages of the male specimens in this series that are interesting: these
include two adult males with fully green crowns, one possible adult with a
mostly green crown but with some dull, blackish feathers on one side, and one
immature male with a nearly fully green throat and chest but only a very
restricted green area on the forecrown, most of the crown being dull
dusky-bronze. Valdés and Schuchmann
(2009) actually described the crowns of the holotype and paratype as
“green-bronze” and “bronze-greenish forehead and bronze-colored crown and nape”
respectively. Because virtually all of
the iridescent feathering of hummingbirds looks black if viewed from an angle different
from that of maximal reflectance, the description given for the new species as
“black from frontal view” might be somewhat misleading depending upon the
precise angle of viewing. Moreover, the
green of the throat and chest of older first-year males (as well as the crown)
is often less a pure green than a paler golden-green, evident in the immature
male, and one of the putative adults has black feathering immediately adjacent
to the bill on the anterior forehead.
Thus, different birds in this series show varying degrees of approach
towards nigricapilla. One of the adult males has a decidedly bronzy
nape and occiput; the other has these areas more blackish, suggesting that this
population may be to some degree intergrades between fannyi and subtropicalis (which
in turn might explain the variability apparent in this small series). I note that very young immature males of the
entire colombica complex lack
iridescent feathering on the crown and have at best much duller green on the
throat and chest confined to the feather tips giving a mottled appearance; the
molt to a more adult-like plumage apparently begins soon after fledging (the
young male described above has remnants of this plumage on the chest). The crown appears to molt somewhat later than
at least the throat and anterior chest and the specimens of nigricapilla could well represent
extreme cases of this - I note that in
at least T. c. venusta, this is also
the pattern of the adult males in their annual molt (see Stiles 1995). The
males of this series also show variation in tail lengths and forks; one has a
tail fork exceeding that of the one specimen of nigricapilla measured (the other lacked the outer rectrices), again
suggesting the influence of fannyi. The other details of the color comparison
between nigricapilla and subtropicalis do not rule out immaturity
or influence of fannyi as well. I note that further south, according to the
map of localities given by Valdés and Schuchmann, fannyi apparently reaches elevations similar to those from whence
come the specimens of nigricapilla and
could well extend up the deep valley below the Embalse de Calima to elevations
at least approaching the reservoir. The
description of a male nigricapilla displaying
at and mounting a female is strikingly similar to what I have seen in young
males of Calypte anna and a few other
species; thus, play behavior of a young bird cannot be entirely ruled out.
A
final reason why the status of nigricapilla
as a valid species might be considered doubtful is the simple fact that in
the limited area described for it occurs a second species of Thalurania virtually identical in
morphology that surely would be a competitor for resources; I consider the
evolution of such a species in sympatry with a potential congeneric competitor
to be extremely unlikely and no other comparable case in hummingbirds occurs to
me. Hence, I must conclude that without
more evidence, ideally including a longer series of males of all ages as well
as females, the evidence for considering nigricapilla
as a valid taxon is inconclusive and I therefore recommend a NO vote on
this proposal.
References
Griscom, L. The ornithology of the Caribbean coast in
extreme eastern Panama. Bull. Mus. Comp. Zool. 72:303-372 (Pp. 334-337 include
a review of the T. colombica complex
as well as more data on subtropicalis,
which is described on P. 337).
Stiles, F. G. 1995. Intraspecific and interspecific
variation in the molt patterns of some tropical hummingbirds. Auk 112:113.132.
Valdes-Velásquez, A. & K.-L. Schuchmann. 2009. A new
species of hummingbird (Thalurania,
Trochilinae, Trochilidae) from the western Colombian Andes. Ornithol. Anz.
48:143-149.
F. Gary Stiles, October 2010
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Comments from Remsen: “NO.
If this is indeed a valid taxon, then more specimens are needed. Thalurania
are typically common and frequently netted, so obtaining more specimens should
be easy. In addition to Gary’s excellent
points, I also note: (1) N = 2 specimens; (2) even if a valid taxon, whether it
should be assigned species rank is a separate question, because its diagnostic
feature, crown color (in this case perhaps just an absence of iridescent
feathers?), is one that varies among other taxa ranked at the subspecies level,
at least within subspecies currently assigned to T. furcata – see the plate in Schuchmann’s chapter in HBW and look
at T. f. nigrofasciata vs. T. f. baeri, for example. Resolving this one should be easy with a
little more fieldwork.”
Comments from Nores: “NO. El hecho de que sean sólo dos ejemplares y que las características que definirían a esta nueva especie son variables, no soporta la validación de la misma, lo cual ha sido señalado con gran precisión por Gary. Además, resulta poco convincente describir a una especie sobre la base de una indicación tal como “negro desde una vista frontal”. Si los autores piensan que realmente se trata de una nueva especie, deberían capturar más machos adultos que muestren claramente las características que separan a la nueva especie.”
Comments from Robbins: “NO. Gary provides very solid comments for questioning the validity of this taxon.”
Comments from Zimmer: “NO. The case
for “nigricapilla” being a distinct
species appears weak at best, for reasons nicely summarized by Gary. I echo Van’s comments, both with respect to
N, and as regards reliance on crown color.
And, I think Gary’s final paragraph regarding sympatry of the proposed
new species with a nearly identical presumed ecological competitor really hits
the nail on the head. This flies in the
face of the essentially allopatric species-distribution pattern of Thalurania.”
Comments
from Graves: “Gary's summary
nicely outlines the problems with the supposed new taxon. The two specimens of
"Thalurania nigricapilla"
were collected by Professor Schuchmann in 1978 and deposited in Bonn. During the subsequent decades, Schuchmann
published many dozens of papers on hummingbirds. The fact that they were undescribed for 30 years
suggests that he didn't ascribe any unusual significance to their appearance
until recently. I have not examined the
specimens, but plumage descriptions suggest they represent young males in
prebasic molt or in first basic plumage.
Wing length (about 4 mm shorter than Thalurania
furcata subtropicalis) also suggests a subadult plumage. The authors did not mention the presence or
absence of fine striations on the maxillary ramphotheca. These would almost certainly be present in
specimens in juvenal plumage and in many specimens undergoing prebasic
molt. Most species in that clade lack
striations in first definitive plumage, however, so the absence of striations
does not necessarily indicate "adult" plumage. Why aren't they subadult (or aberrantly
plumaged) Thalurania furcata
subtropicalis? In any event, the age
class of the specimens is unclear and this ambiguity lobbies against the
recognition of the proposed taxon. This
issue should be relatively easy to resolve with a few days of mist-netting at
the type locality.”
Comments from Pacheco: “NO. A validade do táxon nigricapilla não pode ser assumida a partir dos questionamentos bem construídos por Gary. A julgar pelo conhecimento em Thalurania, as possibilidades de validade do táxon são de fato muito reduzidas.”
Comments from Cadena: “NO. Gary Stiles's arguments are persuasive, and as indicated by Van and Gary Graves, a few days worth of fieldwork in the area would go a long way towards resolving the issues. I visited the Rio Bravo area in 1998 and have Thalurania in my notes as being common then.”
Comments from Jaramillo: “NO. It is
prerequisite to have more specimens from this locality to determine if this is
indeed a new taxon, or an age class, individual variation or subspecies.
Definitely more fieldwork is necessary.”