Proposal (947) to South
American Classification Committee
Split
Bronze-brown Cowbird (Molothrus armenti) from Bronzed Cowbird (M.
aeneus)
Effect on SACC: The South American subspecies of our current Molothrus aeneus
would be treated as a separate species
Description of the
problem:
The
Bronze-brown Cowbird Molothrus [aeneus] armenti has remained little-known until recently, and its status as
a subspecies of Bronzed Cowbird Molothrus
aeneus rests upon the authority of one scientist, Herbert Friedmann, who,
beginning with his review of cowbirds in 1927, considered it a full species,
and only in 1963 decided otherwise, seemingly being swayed by his observations
of a single captive individual, but without providing a rationale.
Molothrus [a.] armenti Cabanis, 1851
(based on Lichtenstein’s 1826 label name) was long known from only four
specimens from Cartagena and “Savanilla”, Colombia
(=Sabanilla, near Baranquilla), though Friedmann
(1957) considered two further specimens (from Demerara, Guyana and Venezuela)
to be doubtfully identified, correctly it would appear as no further reports
have come from these countries. Two of the Cartagena specimens were lost and no
further specimens or living individuals were found for over a hundred years,
despite special effort by e.g., Dugand (1947) and Carriker (in Friedmann 1957),
and the exact collection localities were uncertain. When a live adult male
turned up in a shipment of cagebirds from Leticia, Amazonian Colombia, it was
initially heralded as a rediscovery (Friedmann 1957), and Leticia was accepted
as a genuine locality by Meyer de Schauensee (1964, 1966, 1970), who then
considered the specimens from Cartagena of doubtful provenance. Subsequently,
however, it became evident that the Leticia origin of the shipment was instead
highly doubtful because the many collectors operating there never obtained any
cowbirds, and the likelihood is that the birds were brought on board during
refueling in Barranquilla, where a Leticia-based bird dealer had another
operation (Camacho & Rodríguez-Mahecha 1986).
Although usually
treated as a full species (e.g., Friedmann 1927, 1933, Hellmayr 1937, Meyer de
Schauensee 1966, 1970, Camacho & Rodríguez-Mahecha
1986, Hilty & Brown 1986, Sibley & Monroe 1993, Rada Quintero 2002,
Gill & Wright 2006, Fraga 2011, del Hoyo & Collar 2016, BirdLife
International 2020, Hilty 2021), some of these authors expressed doubt as to
its species status, and it was considered a subspecies by others (e.g., Meyer
de Schauensee 1964, Parkes & Blake 1965, Blake 1968). Then, after its 1969
rediscovery in the wild at P.N. Isla de Salamanca by Gilberto Toro-García,
Dugand & Eisenmann (1983; as interpreted by Friedmann, who at least
finished, or perhaps wrote the paper because both authors of record were by then
deceased) examined a series from the rediscovery site and deemed that it
“cannot be looked upon as other than a southernmost race of … M. aeneus.” Dugand & Eisenmann
(1983) then went on to reiterate some of the known characters by which armenti differs from aeneus, but their posthumously published
opinion as to its subspecific status has been widely followed (Ridgely &
Tudor 1989, AOU 1983, 1998, Dickinson 2003, Dickinson & Christidis 2014,
McMullan & Donegan 2014, McMullan et al. 2018, Avendaño et al. 2017, Ayerbe-Quiñones 2019, Clements et al. 2019, Remsen et al.
2020). By 1963, Friedmann (1963) was treating armenti as a subspecies of aeneus,
although without any comment as to why and without benefit of the larger series
later available. The less extravagant plumage and structural features of armenti compared to aeneus were in fact used as evidence by Parkes & Blake (1965)
to sink the genus Tangavius into Molothrus, with the argument that armenti bridged the gap between Bronzed
and typical cowbirds. Nevertheless, later in the same paper they stated
‘Although its differences in color and proportions suggest that it might be
specifically distinct, armenti is now
considered by Dr. Friedmann to be conspecific with aeneus. We propose to follow his recommendation, based in part on
observations of a living adult male armenti
in the National Zoo…’ (Parkes & Blake 1965). Presumably these observations
of the living bird were Friedmann’s, and it seems unlikely that Bronzed Cowbirds
were also present simultaneously in the zoo for direct comparison.
Subsequently, however, this treatment was followed by others, including the AOS
classification committees. Dugand & Eisenmann (1983), who did examine the
larger series unavailable at the time Friedmann decided on subspecific status,
but it is not clear whether the statement about its being a race of aeneus was theirs or the subsequent
opinion of Friedmann, who finished the manuscript for the predeceased Dugand
and Eisenmann after having been sent their notes and correspondence by F.
Vuilleumier (Friedmann’s footnote in Dugand & Eisenmann 1983). In any case,
it does not appear that convincing argumentation for the conspecificity of armenti with aeneus has been published.
No DNA-based phylogenetic
studies appear to have thus far included armenti.
Nevertheless, though it seems clear that armenti
is closely related to aeneus, it is
quite morphologically divergent from Bronzed Cowbird, as noted in various
sources (especially Jaramillo & Burke 1999):
· Much
smaller size (the smallest cowbird). For males (n=8), wing vs. the three races
of aeneus averaged 96.5 vs. 116.5, 119.0, 108.7; culmen 16.7 vs. 23.0,
23.0, 21.3; tail 61.4 vs. 79.6, 85.0, 73.0; for females (n=8), wing 87.9 vs.
108.7, 104.5, 102.6; culmen 16.2 vs. 20.6, 19.8, 19.1; tail 59.4 vs. 73.0,
73.5, 73.5; measurements from Jaramillo & Burke 1999);
· relatively
smaller bill (see Friedmann’s 1933 Fig. 6) and feet; see above measurements;
· greatly
reduced sexual size dimorphism;
· redder
iris in adult female;
· head
and body of adults of both sexes glossy brown, much like head color of male
Brown-headed Cowbird;
· neck
ruff present in males but much reduced;
· outer
primary not emarginate, unlike Bronzed Cowbird;
· much
reduced hairlike texture of breast feathers in males.
New
information:
We have not had the
opportunity of independently studying specimens of armenti. However, several photos of armenti are now available online on eBird, and these show that iris
color of males is not consistently different from Bronzed Cowbird, although
perhaps not as brilliant red, and irides of female armenti can be nearly as red as in males (brownish to pale orange
in female aeneus).
Thirteen recordings of armenti are available as of 26 August
2022 on xeno-canto (https://www.xeno-canto.org/species/Molothrus-armenti?view=3), and although somewhat variable and similar in quality to
the plentiful material there and on Macaulay Library of North to Middle
American M. aeneus, none share the
same pattern. Its vocal distinctiveness has already been noted by Fraga (2011)
and del Hoyo & Collar (2016), and it seems to be relatively quiet.
For years after its
rediscovery, armenti was mainly
reported from P.N. Isla de Salamanca (e.g., Gochfeld et al. 1980), but remained
relatively little-known, and from few localities (Ridgely & Tudor 1989).
However, with the growth of birding in Colombia that has changed, as can be
seen from the eBird map (https://ebird.org/species/brocow; Panama records are aeneus),
and further data and records have been enumerated in Arzuza
Buelvas et al. (2017), and thus although now much
better known, major areas of its life history are still mysterious such as its
host species (Arzuza Buelvas
et al. 2017). Donegan et al. (2016) stated that “Bronze-brown Cowbird Molothrus armenti is not seen east of
Salamanca”, but Freeman et al. (2011) and others have recorded it east to SFF
Los Flamencos. It has also been reported to the west in Córdoba Department (Estela & López-Victoria 2005). Given
its range restriction and evidently small population size, the species was
considered Threatened (Rada Quintero 2002) or even endangered (Fraga 2011), but
has now been downgraded to Vulnerable (Renjifo et al. 2016). However, it does
occur in degraded and pastoral areas and over a broader area than earlier believed.
Because the range of the Bronzed Cowbird is expanding in Panama with
deforestation, it seems only a matter of time before secondary contact is
established between aeneus and armenti.
One putative ecological
difference mentioned by several authors, the seeming lack of affinity of armenti for cattle (e.g., Hilty &
Brown 1986, Jaramillo 1999), does not appear to hold. A known site for this
species is a cattle pen along the highway near Salamanca, where D. Donsker’s guide took him to see six individuals of armenti along with other blackbirds on
15 Mar 2012. It has even been reported in a park in downtown Magdalena (“red
eye seen clearly”; https://ebird.org/checklist/S30013686), and may feed among domestic fowl (https://ebird.org/checklist/S53399078; Fraga 2011).
Most recent treatments
other than HBW/BirdLife and Gill & Wright (2006) and those following these
lists continue to treat armenti as a
subspecies of aeneus, including
in-country sources (e.g., McMullan & Donegan 2014, McMullan et al. 2018
[not seen], Donegan et al. 2015, Ayerbe-Quiñones
2019). This is likely to be largely due to historical inertia. We consider that
there is a strong case for the treatment of armenti
as specifically distinct, given its morphological and vocal differentiation,
that a strong case for conspecificity was never made, and that continued
subspecific status would need to be based upon evidence of which we are
unaware.
English name.—The name “Bronzed Cowbird” is vastly more familiar and
entrenched for the widespread North and Middle American M. aeneus, which was also long known as Red-eyed Cowbird (Friedmann
1957). The name Bronze-brown Cowbird is the most-used name for armenti (and continues to be used in the most recent field guide to Colombia,
Hilty 2021), but it has also been known as Arment’s Cowbird (in e.g.
Hellmayr 1937), Cabanis’s Cowbird (Brabourne and Chubb 1914), and Colombian
Red-eyed Cowbird (Friedmann 1957). Although Friedmann (1957) quoted E.
Stresemann as having informed him that armenti
stems from the Latin word for armentum, meaning a drove of cattle, Jobling (2010)
indicates it was named after T. Arment, a “collector in Colombia”.
Recommendation:
We recommend that M. armenti be
considered specifically distinct, for the reasons stated above, and we
recommend that it continue to be known as Bronze-brown Cowbird, which is highly
entrenched. If voting yes for the split (part A), please also vote for an
English name (part B).
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Pamela C. Rasmussen and David B. Donsker, June 2022
Note from Remsen: This is from
a manuscript that is in review; normally, we would just wait for it to appear,
but the IOC’s Working Group in Avian Classification (on with Nacho is our
representative) could use our input ASAP.
Also,
for voting, for those of you who vote yes who also vote on English name issues,
I am going to assume that use of the historical name Bronze-brown Cowbird is
acceptable unless you state otherwise. I
don’t see any point in doing a separate subproposal on this unless someone
objects.
_______________________________________________________________________________
Comments
from Stiles:
[YES]. I have a small set of data for this species, but some interesting
points can be made. For one thing, by weight armenti is less than half
as heavy as aeneus, sex for sex. Mean weights: for males, armenti:
34.65 g, n=2; aeneus: 67.76 g, n=6 (from Costa Rica; for females, armenti:
24.60 g; aeneus: 55.12 g (n=5, again from Costa Rica). Sexual dimorphism
in weights is thus actually greater for armenti than for aeneus,
the opposite from plumage differences, which are less for armenti. My
field experience with armenti is limited, but in both my contacts with
this bird near Cartagena, my impression of its voice was that it differed
considerably from what I had often heard from aeneus in Costa Rica,
although I was unable to obtain recordings.
Another observation: in both of my observations of small flocks of armenti.
I noted an unusual plumage, presumably of young males, in which the heads and
breasts were blackish, concolor with the upperparts, but sharply separated from
the slightly paler brown of the abdomen (similar to one of the females in the
ICN collection), giving a “hooded” aspect; I had never seen such a plumage in
years with aeneus in Costa Rica. Unfortunately, this plumage is not
represented in the small series in ICN. In sum, I think that these observations
further tip the balance for species status of armenti, so YES to this
proposal.”
Comments
from Claramunt:
“NO. I really don’t see the evidence. I only see anecdotal
information that is difficult to interpret.”
Comments from Remsen: “YES. Santiago is correct in that the evidence is a
collection of anecdotes. Obviously, a
formal, quantitative study would be needed if this were a novel split,
especially on male displays, which would obviously be critical to assessing
potential gene flow in this promiscuous species. But in cases like these, I think we should
also evaluate the rationale for their treatment as conspecific as well, which
in this case rests on the following (from Dugand and Eisenmann):
“With 12 specimens now available for study in the museums of New York
and Bogota, it becomes clear that armenti cannot be looked upon as other
than a southernmost race of the species M. aeneus. M. armenti
agrees with aeneus in the emargination of the primaries in the adults.
Inasmuch as no adult females of armenti have been described until now,
it may be put on record that in coloration they resemble, but are not quite as
glossy as, female M. aeneus assimilis. The adult males of armenti
resemble those of aeneus but are less bronzy, more brownish. The brown
color of the head and body of armenti, while more iridescent, is not
unlike that of the head of male ater (Parkes and Blake 1965). The bill
of adult armenti is somewhat shorter and slenderer than that of aeneus.”
“Personally,
I don’t see this as compelling logic for taxon rank either way. Note that when Dugand and Eisenmann (or was
it Friedmann? See proposal) wrote their paper, armenti was barely known,
and a main theme of the paper was where armenti actually occurred,
because an imported zoo bird was thought to have come from Leticia on the
Amazon. The other main point of the
paper is that the species was previously known from just two extant specimens,
and Dugand and Eisenmann now had ca. 10 new specimens collected by Gilberto
Toro-Garcia, a Colombian who was in-charge of the newly created Parque Nacional
de la Isla de Salamanca. (That
Toro-Garcia was not included as a co-author on this paper is unfortunate but
reflects different attitudes on authorship of that era.)”
“So,
what we have here is competing interpretations of anecdotes. In my opinion, those assembled by Rasmussen
and Donsker place burden-of-proof on treating armenti as just a
subspecies, especially since some of them might be relevant to mate selection.
“The
only thing I might add it that using a comparative approach, my superficial
assessment of subspecies differences in the other two widespread polytypic Molothrus,
ater and bonariensis, is that they are minor compared to those
between armenti and aeneus.
In M. ater, the three subspecies are defined based on size and
female plumage; male plumages are evidently indistinguishable. In M. bonariensis, subspecies are also
defined primarily on female plumage and size.
Variation among the other three subspecies of aeneus is largely
in female plumage and size (Jaramillo & Burke). With all appropriate caveats concerning these
comparisons, armenti would seem to be an outlier in terms of Molothrus
subspecies-level differentiation.”
Comments
from Areta:
“YES. A borderline case. In the face of uncertainty and the lack of consensus on
how to treat armenti, the
accumulation of structural, size, plumage, and vocal differences occurring over
a short geographic distance and across an avifaunal break tips the scale
towards species recognition for armenti.
This is a judgment call, and the case begs for an integrative study.”
Additional
comments from Stiles:
“Ridgway also gaves the same set of measurements for M.
aeneus, so I made the corresponding bill measurements for armenti. (note
that the appropriate comparison is with the nominate and largest race of aeneus
(called robustus in Ridgway); the two smaller races are resident in
extreme SW US to Jalisco, and Guerrero-Oaxaca in Mexico and are thus not likely
to approach Colombia, whereas M. a. aeneus is known southeast to
central-western Panama and is expanding its range eastward. An interesting
feature of both M. a. aeneus and armenti is that the bills of the
females average larger than those of the males and, comparing the sexes, show
differences in shape. The sexual dimorphism in the CtB/
BHb ratios is: for aeneus
= 0.992; for armenti = 0.968 – bills of armenti average more
different from those of males (being relatively longer and slenderer). Another
difference: Ridgway´s diagram of the wing of male aeneus clearly shows
that the three outermost primaries have the inner webs distinctly deeply
notched; in male armenti, none of these primaries are notched.
Thus, the sound of the courtship flight of aeneus will be distinctly
different from that of armenti (another difference in favor of species
status for armenti).
Comments
from Lane:
“NO. I remain rather unimpressed by the plumage and
voice of M. armenti as being sufficiently distinct from M. aeneus
as to suggest different species. The differences in size may be enough to move
me, but I'd like to see a far more explicit comparison (molecularly, vocally,
behaviorally, morphologically) of the two groups of taxa to feel that they
merit species status. I will point out that the "13" cuts of M.
armenti in X-c are fraught with duplicates and
even one or two triplicates, so there are more like 6 cuts, from only two days
and two localities, so perhaps only representing two individuals... and my
experience from M. aeneus is that a single individual can give several
different song types and fragments of songs, so any perceived differences based
on the cuts on X-c are, to me, fairly meaningless. A
far larger sample would be necessary for me to sit up and take notice. Indeed,
I hear vocalizations I would not be able to separate from common vocalizations
I regularly hear from M. aeneus here in Louisiana and Texas.
Furthermore, the "hooded" plumage Gary mentioned in his comments is
one I have seen among M. aeneus in the USA (e.g.: https://macaulaylibrary.org/asset/251320751, https://macaulaylibrary.org/asset/56040011, https://macaulaylibrary.org/asset/102409151, https://macaulaylibrary.org/asset/442149081), and is
presumably a stage in the aging process and so should not be seen as a
distinctive plumage class that would merit difference between the two groups.”
Comments from Robbins: “NO. I had looked at the Molothrus proposal
a month or so ago and came away with the same impression as Dan. As a result, I decided not to vote on the
proposal at that time. As Dan noted, the
sample size for the vocalizations and photos is quite small, i.e., people on a
tour had photos taken by one person put on all the eBird checklists. I didn't appreciate what Dan stated concerning
the "hooded" plumage appearing among other aeneus populations. Finally, there is considerable variation in
morphometrics among the aeneus taxa, so if armenti is indeed
smaller, that may be meaningless for ascertaining species status. Given these issues, I vote NO for recognizing
armenti until we have more information.
Comments from Bonaccorso: “NO. I would love to say yes,
because of the conservation concerns. However, the problem is that size
differences could result from local adaptation, and plumage differences seem to
be subtle (“lacking the bronzy cast of other subspecies” (Ellison and Lowther
2020)). Also, since cowbird songs are so variable, a rich comparison would be
needed to identify homologous vocalizations. On top of that, there are no DNA
data. So, we are deciding based on very little data.”
Comments from Zimmer: “A
very tepid YES. I don’t have strong
feelings about this one way or the other, and agree with Dan and Mark that much
of the evidence for splitting is shaky.
However, what I come back to, is that the evidence for the original lump
was even shakier. The status quo becomes
harder to defend, when we consider that it became the entrenched position only
through the unjustified overturning of the prior status quo. The anecdotal accounts of vocal differences
and the limited biometric and plumage data, imperfect as they may be, are
enough to tip the scales toward the original status quo in my mind, and by
doing so, may generate enough interest for someone to get out there and really
nail this thing down, one way or the other.”
Comments from Pacheco: “NO. I have read the arguments for and against the split and agree
that the best decision requires waiting for DNA data.”
Additional comments from Stiles: “First, the evidence for
conspecificity with M. aeneus, based principally on Friedmann, who in
turn used the unpublished information from Dugand & Eisenmann posthumously.
Friedmann stated that there was no difference in the degree of emargination
between aeneus and armenti; however, this is incorrect (see above).
Second, the size difference is dismissed by Dan as a useful criterion. However,
if one uses as a yardstick the differences among the three subspecies of aeneus,
these are far less than that between these and armenti, especially given
the different degrees of sexual dimorphism between armenti and nominate aeneus.
Also, I have looked at the 4 photos of supposedly hooded aeneus given by
Dan; only one approaches the hooded plumage of armenti that I described
and in even that one, the match is by no means perfect (head to upper breast
black, breast a brighter brown that shades into black on the abdomen, vs. a black head to breast, more sharply set off
from a dark brown abdomen). So in sum, the evidence for conspecifity is
definitely not wholly satisfactory.”