Proposal
(73) to South American
Classification Committee
Split Cacicus
microrhynchus from C. uropygialis
Effect on South American
CL: This proposal would split Cacicus uropygialis into
two species, with recognition of Central American microrhynchus,
including pacificus (Panama to Pacific slope Ecuador) as a
separate species.
Background: The
bird we treat as one species, Cacicus uropygialis has a disjunct
distribution, with nominate uropygialis restricted to the
subtropical zone of the Andes in a patchy distribution from W. Venezuela to C.
Peru; microrhynchus occurs from E. Honduras to E. Panama,
whereas pacificus is found from W. Panama south to El Oro, Ecuador.
Both latter subspecies are restricted to the tropical zone.
Ridgway (1902)
listed microrhynchus as separate from uropygialis.
In his description of pacificus, Chapman (1915) describes it as
similar in size to microrhynchus, but with larger bill, swollen at
base. Measurements show a substantial difference in size from the larger uropygialis.
He notes the following regarding the swelling at the base of the bill of pacificus and microrhynchus:
"In others it is less prominent, and it seems probable that the large and
small-billed forms merge somewhere between the Canal Zone and the Colombian
boundary." This suggests a possible cline between pacificus and microrhynchus.
Measurements of uropygialis and pacificus show no
tendency towards a cline between those two allopatric taxa, and appear
distinctly different in size, even though pacificus is
relatively large-billed. Hellmayr considered them all conspecific, with the
following footnote: ""This form combines the general dimensions
of C. u. microrhynchus with the powerful bill of C. u.
uropygialis, thus occupying in its characters an intermediate position as
it does geographically." Peter's Checklist (Blake 1968) puts all three
taxa under uropygialis, without comments. Wetmore et al. (1984),
noted difference in size between nominate and microrhynchus/pacificus,
but mentioned that pacificus seems intermediate in bill shape
and geographic position. Pacificus has a larger bill than microrhynchus,
with the mandibular rami distinctly swollen. Wetmore et al. (1984) do note that
an occasional male of microrhynchus shows a faint swelling on
the outer face of the base of the mandibular rami, an indication of approach to
the condition found in C. u. pacificus, but this is unusual. They
also note that these two races are similar in size. Ridgely and Tudor consider
all conspecific, but note that the lowland western taxa probably deserve
species status, separate from the highland uropygialis. More recently,
Ridgely and Greenfield (2003) considered pacificus a subspecies of microrhynchus,
but both separate from uropygialis as did Hilty (2003).
We did the same, separating the lowland taxa from the highland taxon (Jaramillo
and Burke 1999).
In terms of plumage the
three taxa are quite similar, black with a restricted red rump patch. However
it is in their size and structure that they differ noticeably. Other than in
bill size, pacificus is similar to microrhynchus. On the
other hand, uropygialis is substantially larger than the
lowland forms, and is proportionately longer-tailed, whereas the microrhynchus group
is proportionately longer-winged (absolute wing lengths still substantially
smaller than in uropygialis though). Note that in turn all
red-rumped caciques are quite similar, with the eastern lowland haemorrhous differing
from uropygialis largely in size, proportions, size of rump patch, voice
and strength of plumage iridescence. I don't think that the species status of haemorrhous has
been questioned seriously in the literature.
New information: There
isn't much new information per se, but there has been recent publication of the
vocalizations of these taxa. Hardy et al. (1998) includes songs of microrhynchus, pacificus and uropygialis.
More recordings of uropygialis are available on Moore and Lysinger (1997).
Analysis: The
problem we have here is of taxa that are visually similar, which would not
necessarily pop out at you as being but mere varieties of each other by looking
at the museum skins. In real life the two groups, highland uropygialis and
western lowland microrhynchus group are quite different.
First, looking at what you would see in the museum skins the following is the
case:
- uropygialis much
larger than microrhynchus group. Uropygialis males:
Wing 156.5 (145-165); tail 129.6 (107-142), culmen 30.8 (29-33); tarsus 33.0
(32 36). Microrhynchus males: Wing 129.0 (122.0-136.5); tail
89.7 (83.3 96.5); culmen 29.6 (28.8 30.2); tarsus 28.8 (27.4 ?). pacificus males:
Wing 132.2 (122.2 137.2); tail 91.1 (87.8 94.9); culmen 29.6 (28.2-31.0);
tarsus 29.5 (28.2 31.5).
-
Structurally, uropygialis is very long-tailed. The microrhynchus group
is relatively long-winged, something that is lost in the measurements due to
their smaller size.
Second, are the behavioural and ecological differences.
- Uropygialis is
quite consistently a species of the subtropical zone of the Andes, restricted
to the east slopes south of Colombia. The microrhynchus group
is found in the tropical zone, west of the Andes. The other component, if one
enlarges this complex is haemorrhous, which is restricted to the
tropical zone east of the Andes.
-
Vocally the uropygialis and the microrhynchus group
differ noticeably. Songs and vocalizations can be complex in this group, and
certainly have a learnt component, other Cacicus species show
noticeable changes in songs each year, and some are even mimics. Even so, it
needs to be pointed out that the vocalizations of these two groups differ
substantially, with uropygialis having more harsh and screechy
notes, the microrhynchus group being more musical and
pleasant. The exact structure and notes given in songs and other vocalizations
differ clearly to the ear, but are extremely difficult to describe on paper.
Calls are harsher in uropygialis, resembling haemorrhous more
than the more pleasant and whistled notes of microrhynchus.
-
Nesting behaviour and social structure is not well
known for uropygialis. The smaller microrhynchus has
a strong tendency to nest solitarily although small colonies are known, and has
even been seen to have male help at the nest. Most unlikely cacique behaviour. Flocks of uropygialis tend to
be larger than the pairs or presumed family groups of microrhynchus,
suggesting that it may have a more standard cacique social structure. Hilty
(2003) reports that uropygialis nests in small colonies.
What we have here is a
situation that does not make much sense from a biogeographical stand point, a
western lowland population is here lumped with a patchily distributed
subtropical Andean taxon. No intermediates between the two are known, they are
entirely allopatric. I imagine that the two forms may come relatively close
geographically in parts of Colombia. Comments that the form pacificus may
in fact be intermediate between microrhynchus and uropygialis are
in error. The more southern pacificus is larger than microrhynchus,
and supposedly proportionally larger in bill size (though my measurements do
not bear this out), but in reality it is still a much smaller and
shorter-tailed bird than uropygialis. Vocal differences between the two
groups are clearly diagnosable, although the complexity of cacique voices makes
comparison difficult. What I take to be clear differences in the call notes are
suggestive of groups that have not been in contact for some time, assuming that
the call notes are not learnt. The form pacificus has also
been suggested to comprise a separate species (Ridgely and Tudor 1989) but this
seems much more of a hard sell, particularly since little data is available on
where the two come into contact, and all differences that have been noted could
potentially be clinal. In fact, Chapman (1915) in his description of pacificus suggests
that there may be an area where microrhynchus and pacificus merge
into each other between the Canal Zone and Colombia. Ridgely and Greenfield
(2001) do not propose species status for pacificus.
So we have two groups that
differ in size and structure, voice, life zone that they inhabit and perhaps
nesting behaviour and social structure. Early on they
were considered different species, but were lumped more recently without any
reasoning as to why this made sense. From my perspective on these caciques if
you are going to put these two together, then you need to seriously consider
why haemorrhous should not be lumped in as well. I don't think
that lumping all red-rumped caciques is logical. I am just making the case that
if taxa as different as uropygialis and the microrhynchus are
lumped then why not lump all? The differences in voice, size, structure, and
distribution of these taxa suggest that the lowland microrhynchus/pacificus pair
should be split from the highland uropygialis.
Recommendation: YES –
divide microrhynchus from uropygialis.
Maintain pacificus as a subspecies in microrhynchus.
If we vote to split these
two, using the Ridgely name Subtropical Cacique for uropygialis does
work for me. It is one of the features that clearly separate it from all other
red-rumped caciques. Note that microrhynchus has been known as
"Small-billed Cacique" (Ridgway 1902), and this is an option, but
retention of Scarlet-rumped Cacique for microrhynchus may be
the way to go as this is the name that has been used recently by folks that are
separating these taxa out. I don't know that microrhynchus is
all that small-billed, it just is a small cacique.
Literature Cited:
BLAKE,
E. R. 1968. Family Icteridae. Pp. 138-202 in "Check-list of birds of the
World, Vol. 14" (Paynter R. A., Jr., ed.). Museum of Comparative Zoology,
Cambridge, Massachusetts.
CHAPMAN,
F.M. 1915. Diagnoses of apparently new Colombian birds IV. Bull. American Mus.
Nat. Hist. XXXIV: 657-659.
HARDY,
J.W., G.B. REYNARD, AND T. TAYLOR. 1998. Voices of the Troupials, Blackbirds
and their Allies. ARA Records, Gainesville, FL.
HILTY,
S. L. 2003. Birds of Venezuela. Princeton University Press, Princeton, New
Jersey.
JARAMILLO,
A. AND P. BURKE. 1999. New World Blackbirds, the Icterids. Princeton University
Press, Princeton, NJ.
MOORE,
J.V. AND M. LYSINGER. 1997. The bird of Cabañas San Isidro, Ecuador. John V.
Moore Recordings, San Jose, CA.
RIDGWAY
1902. The birds of North and Middle America. Bull. U.S. Natl. Mus., 50, pt. 2.
RIDGELY
, R. S., AND P. J. GREENFIELD. 2001. The birds of Ecuador. Vol. II. Field
guide. Cornell University Press, Ithaca, New York.
RIDGELY,
R. S., AND G. TUDOR. 1989. The birds of South America, vol. 1. Univ. Texas
Press, Austin.
WETMORE,
A., R.F. PASQUIER, S.L. OLSON. 1984. The birds of the Republic of Panama, Part
4.
Alvaro
Jaramillo, October 2003
________________________________________________________________________________________
Comments from Silva:
"NO. I think that this set of arguments could be published before the
change may be accepted."
Comments from Robbins:
"YES. Long overdue".
Comments from Stiles:
"NO, no properly published analysis. Having said this, I might add that I
agree with the proposal, having had experience with all three in the field - if
Alvaro wishes to publish a note incorporating measurement stats and sonograms,
I will certainly go for it (might I suggest Ornitología Colombiana??)"
Comments from Zimmer:
"NO. I agree completely with Alvaro that two species are involved.
However, lack of any published analysis is the monkey wrench. I'd switch in a
heartbeat if there was even a brief published analysis."
Comments from Nores: "NO estoy de acuerdo con considerar a Cacicus
microrhynchus como especie y a pacificus como una
subespecie de microrhynchus. Aunque las diferencias entre los
caciques de rabadilla roja marcadas por Alvaro parecen válidas, antes de tomar
una decisión habría que ver una nota publicada al respecto. Particularmente
importante en su argumento pararse ser lo de las vocalizaciones y el hecho de
que haemorrhous no difiere más de uropygialis que microrhynchus de uropygialis."
Comments from Remsen:
"NO. Reluctantly. This is one of many in which our unpublished or
qualitative knowledge strongly indicates that at least two species are
involved. However, I regard our mission as evaluating published evidence for
taxonomic changes, and with that mission in mind, the evidence is insufficient,
in my opinion. Most of the evidence summarized by Alvaro is of interest and
consistent with 2-species treatment, but in my opinion, also consistent with
1-species treatment, i.e., similar patterns of geographic variation exist
within polytypic species, including Icteridae. The exception would be the vocal
differences, but these are not formally described or presented. All that is
needed for a YES vote from me on this is publication of sonograms of homologous
vocalizations from critical areas."