Proposal
(59) to South American Classification
Committee
Split Celeus
obrieni from C. spectabilis
Effect on South American
CL: This proposal would elevate a taxon to species rank that
we currently treat as a subspecies on our baseline list.
Background: Prior
to 1973, Celeus spectabilis (Rufous-headed Woodpecker) was
considered to be a single polytypic species consisting of two described
subspecies:
C. s.
spectabilis Sclater & Salvin, 1880 -- E Ecuador & NE
Peru.
C. s.
exsul Bond & Meyer de Schauensee, 1941 -- SE Peru,
extreme W Brazil & N. Bolivia.
In 1973, Short described a
new subspecies, C. s. obrieni, from a single unidentified specimen
(pointed out to him by Charles O'Brien) in the AMNH: an adult female, collected
16 August 1926 by E. Kaempfer at Iruçui, state of Piauí, Brazil, elevation 124
m, on the Rio Parnaiba. Short refers only generally to the habitat in
which obrieni was collected, saying that it was from "dry
forested country", and speculating that it was probably widespread in the
Piauí-Maranhão region.
Short's diagnosis of the
specimen follows: "Differs from C. s. spectabilis and C.
s. exsul in its smaller size (wings, tail, bill, tarsus), and in
several color features, especially the greatly reduced barring dorsally, and
reduced markings ventrally. Also whiter above and below; small outer rectrices
mainly cinnamon (nearly all black in exsul and spectabilis); and
secondaries paler, more buffy (less chestnut), especially on tertial feathers.
The bill appears yellower, less white, compared with both older and more
recently collected specimens of other races."
Measurements given by
Short (1973) are as follows:
Wing Chord:
C. s. obrieni 136 mm
C. s. spectabilis (n = 2) 150 mm, 147 mm
C. s. exsul (n = 9) range of 138-153 mm
Tail:
C. s. obrieni 95
mm
C. s. spectabilis (n = 2) 92 mm, 101 mm
C. s. exsul (n = 9) range 99-108
Exposed Culmen:
C. s. obrieni 24.3
mm
C. s. spectabilis (n = 2) 28.3
C. s. exsul (n = 9) range 28.7-31.2
Tarsus:
C. s. obrieni 21.1
mm
C. s. spectabilis (n = 2) 23.4 mm, 23.1 mm
C. s. exsul (n = 9) range 22.1-24.2
Short concluded that the
"new form clearly represents a race of C. spectabilis, rather
than some other species of Celeus, by virtue of its fully rufous
head, its black, shield-like breast patch, it mainly clear rufous secondaries,
and its black, unbanded tail. Its bill, although small, matches spectabilis in
the slight curvature of the culmen, and in the breadth across the nostrils. In
its reduced markings obrieni bears the same relation to other races
of C. spectabilis that C. torquatus torquatus does
to other races of C. torquatus, and that C. flavescens
ochraceus does to other races of C. flavescens."
Subsequent authors have
followed Short's lead in treating obrieni as a subspecies
of spectabilis: Sibley & Monroe (1990), Sick (1993), Winkler et
al (1995), Parker et al (1996), Clements (2000) and Winkler & Christie
(2002).
F. C. Novaes conducted
surveys in Piauí in the region of Uruçui-una close to the type locality
of obrieni in 1980, but was unsuccessful in relocating the
taxon (Novaes 1992). Novaes described the dominant habitat of the area as
cerrado intermixed with caatinga, but with other kinds of vegetation, such as
dry scrub forest, low riverine scrub forest, and swamp vegetation dominated by
burití palms (Mauritia flexuosa).
The holotype (and lone
specimen) of obrieni remained the sole basis for the inclusion
of C. spectabilis on the Brazilian list, until 1995, when A.
Whittaker, as part of a Goeldi Museum expedition, located and tape-recorded
several individuals fitting the plumage characters of C. s. exsul at
various localities in the upper Rio Juruá drainage in Acre (Whittaker &
Oren 1999). All of the Acre birds were found either in stands of bamboo, or in
humid second-growth (Cecropia dominated) bordering rivers; the
typical habitats in which the species is found in Peru and Ecuador (e.g.
Winkler et al 1995, Parker et al 1996, Ridgely & Greenfield 2001, Winkler
& Christie 2002). Whittaker & Oren (1999), commenting on the
distinctiveness of obrieni, combined with the huge range
disjunction from other populations of C. spectabilis, and its very
different habitat, suggested that obrieni should be accorded
full species status. They also suggested the English name of "Caatinga
Woodpecker" for obrieni, to highlight its fairly unique habitat
(among Celeus). Winkler and Christie (2002), noted that "obrieni
differs significantly in plumage, and data on habitat indicate major
distinction from other races; possibly a separate species, but no further
information available." These authors later go on to say "it has to
be assumed either that the taxon is extinct or that it represents a highly
aberrant form of another species."
Analysis: This
is a difficult case, primarily because of lack of information. The taxon obrieni is
described from a single specimen, there are no tape recordings of voice, and
habitat preference is essentially inferred from the general habitat surrounding
the type locality. In general, obrieni appears to be smaller than
other Rufous-headed Woodpeckers, although I am largely unimpressed by the
degree of difference (except possibly for culmen length) given a sample size of
N = 1. On the other hand, plumage distinctions between obrieni and
the other two subspecies of C. spectabilis are striking (as
illustrated by dorsal and ventral photos in Short 1973). Short's comments that
the differences, although major, are in order of magnitude similar to those
found between various subspecies in the C. torquatus and C.
flavescens complexes are well-taken. However, in both torquatus and flavescens,
the different subspecies, although strongly divergent in plumage, have
intergrading populations that are known to be vocally similar or identical to
one another. With obrieni, we are talking about a highly
distinctively plumaged bird that is separated from the nearest known population
of spectabilis by ca. 3,150 km, and which occurs in a dry
forest-caatinga-cerrado biome, whereas both spectabilis and exsul are
birds of humid forest habitats. Whittaker and I searched for obrieni near
the type locality in February of this year (without luck): we saw no habitat
approaching the habitats in which we have seen C. spectabilis in
Peru or w. Brazil. Aside from a few remnant patches of humid (semi-deciduous)
terra firme forest (which were occupied by C. flavescens ochraceus),
the habitats encountered were notably xeric. It seems safe to assume that
regardless of the precise habitat in which obrieni was
collected, it was not even remotely similar to that occupied by C.
spectabilis in the remainder of its range. Given the extreme range
disjunction, very different habitat types, and quantum plumage differences of
the taxa involved, I am somewhat surprised that obrieni was
not described as a separate species in the first place, but Short (1973) makes
no mention that this was even under consideration.
Recommendation: In
spite of my general reluctance to describe anything on the basis of a single
specimen (especially lacking vocal data), I recommend splitting obrieni as
a separate species from C. spectabilis on the basis of a
hugely disjunct range, occupation of a different biome, and quantum differences
in morphology. The caatinga region of northeast Brazil is a noted region of
endemism, and recognition of obrieni as a separate
species-level taxon would fit an established biogeographic pattern. Although I
don't believe that conservation considerations should drive taxonomic
decisions, elevation of this obscure form as a species-level taxon would have
the added benefit of focusing more attention on locating an extant population,
and on conservation initiatives for the caatinga region of Piauí in general.
Literature Cited:
CLEMENTS,
J. F. 2000. Birds of the world: A checklist. Ibis Publishing Company, Vista,
California.
NOVAES,
F. C. 1992. Bird observations in the state of Piauí, Brazil, Goeldiana Zool.
17:15.
PARKER,
T. A., III, D. F. STOTZ, AND J. W. FITZPATRICK. 1996. Ecological and
distributional databases. Pp. 132436 in: STOTZ, D. F., J. W. FITZPATRICK, T.
A. PARKER III, AND D. K. MOSKOVITS. Neotropical birds: Ecology and
conservation. University of Chicago Press, Chicago, Illinois.
RIDGELY,
R.S., AND P. J. GREENFIELD. 2001. The birds of Ecuador. Vol. 1. Status,
distribution, and taxonomy. Cornell University Press, Ithaca, New York.
SHORT,
L. S. 1973. A new race of Celeus spectabilis from eastern
Brazil. Wilson Bulletin 85:465467.
SIBLEY,
C. G., AND B. L. MONROE, JR. 1990. Distribution and taxonomy of birds of the
World. Yale University Press, New Haven, Connecticut.
SICK, H.
1993. Birds in Brazil. Princeton University Press, Princeton, New Jersey.
WHITTAKER,
A., AND D. C. OREN. 1999. Important ornithological records from the Rio Juruá,
western Amazonia, including twelve additions to the Brazilian avifauna. Bull.
B. O. C. 119:235260.
WINKLER,
H., D. A. CHRISTIE, AND D. NURNEY. 1995. Woodpeckers: An identification guide
to woodpeckers of the world. Houghton-Mifflin, Boston, Massachusetts.
WINKLER,
H., AND D. A. CHRISTIE. 2002. Family Picidae (Woodpeckers). Pp. 296558 in: DEL
HOYO, J., A. ELLIOTT AND J. SARGATAL eds. Handbook of birds of the world. Vol.
7. Jacamars to Woodpeckers.
Kevin J.
Zimmer, August 2003
P.S.: If the proposal
passes, then I'll follow up with another one on the English names.
_____________________________________________________________________________________________________
Comments from Robbins:
"Although it seems likely that obrieni is a valid species
given the distinct plumage and the large distributional disjunction from Celeus
spectabilis, I'm against elevating it to species status based solely on a
single specimen. I'd even support this if Kevin and Andy had seen birds that
matched the description of obrieni, but at this point we can't rule
out the possibility, however unlikely, that the holotype is an aberrant or
hybrid individual."
Comments from Jaramillo: "YES.
The great difference in plumage, habitat, and massive range disjunction all
suggest that this is a good species. It is unfortunate that we have only one
specimen to work from, and no recent sightings of this taxon. However, the
hybrid theory for this specimen is exceedingly unlikely in my opinion. The
question would be hybrid of what? Particularly with respect to the place it is
known from. I don't know that part of the world but is Celeus
flavescens the only possibility there? Is there any other Celeus there
which could be the potential other parent? I think that in this case the
possibility that this single specimen is a hybrid is so remote that at least
for me it becomes a non-issue."
Comments from Silva:
"YES. obrieni remains as a big problem. I have a student
(Marcos Pérsio Santos) living in Piauí. He spent lots of time looking for this
taxon and did not find anything. Certainly, it is a distinctive taxon and I do
not think that it represents a case of hybridization. The type-locality is a
mosaic composed by cerrado, caatinga and dry forests. Unfortunately, most of
the tall (25 m) dry forests that once was widespread in this region has been
already completely modified. If obrieni is associated with this kind of
habitat, it is certainly under very strong threat. I agree with Kevin
that obrieni does not belong to Celeus spectabilis, a
completely different species. Although the weak evidence (one specimen
and no recent record) I will vote yes in this proposal."
Comments from Stotz:
"YES. Celeus obrieni is clearly related to spectabilis.
Given that spectabilis doesn't occur anywhere close to the
type locality of obrieni and the very distinctive plumage of obrieni,
it seems essentially impossible to me that obrieni could be
explained as a hybrid or aberrant spectabilis. Although the
published justification is not very strong, I think it is sufficient to hang
our hat on, and recognize obrieni."
Comments from Stiles:
"[YES] Somewhat reluctantly, I will recommend species status. Reluctantly,
because I dislike recognizing taxa from single specimens, especially when no
reliable field observations are available. However, to my knowledge Kaempfer
was a reliable collector and the data on the label are specific as to date and
locality. The morphological differences in combination with the wide range
disjunction and presumably very different habitat (here is where I am least
content with the lack of field data, but if the locality is accurate the
habitat must be different from spectabilis) all point to obrieni being
something different."
Comments from Schulenberg:
"My vote is NO. But that is a vote on principle, pure principle.
“Kevin Zimmer does a good
job of convincing me that obrieni deserves species rank. My
only concern is that Kevin's well-reasoned arguments are not published. I don't
see any substantive difference between Peters, or a field guide author,
publishing a taxonomic decision with little or no available rationale; and our
committee publishing or reaching a taxonomic decision on the basis of
"inside" information, with little or no available (to anyone outside
of SACC) rationale. In both cases, from the perspective of anyone on the
outside, the decision is made inside of a "black box."
“Admittedly, outsiders
might stumble across our SACC proposal website, or we might even direct people
to it. But what is the long term prospect of the website? To the greatest
extent possible, I'd prefer for our decisions to be based on published information
and published rationales. Kevin's proposal looks to me to be a great first
draft of a short note for publication on this taxonomic change; if such a
manuscript had been submitted already, then I would vote in favor of this
proposal in a heartbeat.
“I too am uncomfortable,
of course, with species known from only a single specimen. Hybridization seems
very unlikely to me in the instance, however: what would the parent combination
be? And outside of some well known instances of secondary contact (e.g., Colaptes),
so far as I know woodpeckers are not particularly prone to hybridization
anyway.
“It would be interesting
to know what other specimens were collected the same day and place as the type
of obrieni, to see whether this might give some clues as to what habitat the
bird might be found in (or once occupied?)."
Comments from Whitney: "Celeus
spectabilis obrieni is known only from the type at AMNH. That proposal in
the literature to split it as a species is ludicrous (nothing more than a
statement that it should be elevated to species rank because it's apparently
unique). This question deserves careful analysis, which Kevin should do and
publish before the SACC accepts it as a species.
“I looked at the specimen
a few years ago, and I wish I could lay may hands on the video and notes I
made, but it's late and this may be a done deal now. Anyway, it's a very
strange specimen, I think. I was not convinced, on looking at it, that it could
not be a mutant individual showing a combination of characters that are all
individually present across the genus, or perhaps a cross or backcross between C.
flavus and another species (flavescens, even torquatus
potentially involved). C. flavus has a lot of rufous in the
wing up in that region. Finally, I went to Uruçuí back in about March 1999. If
the bird is at all like spectabilis, it is almost certainly going to be
in a river-created habitat. I saw no sign anywhere near (above or below) Uruçuí
of patches of Gynerium, bamboo, other primitive grasses, other
river-edge veg that looked like it might be OK for something similar to C.
spectabilis. Who knows what the river looked like when Kaempfer was there
-- but I can report that the entire region is basically cerrado, often right
down to the bank of the river, and it's in amazingly good shape, I imagine very
much like it was back then. All of this means next to nothing, of course -- but
I do believe that obrieni is best left alone until someone finds it
alive or gets a molecular analysis of it done. At least find out what its
mother was. If she was not any of the species we have on the books, then that's
enough to convince me that it can be elevated to species rank. (Not that this
info is recoverable from this specimen, but it certainly could be tried).
“What's the point of
elevating it without clear reason? Feel free to forward this message to others
if you think appropriate."
Comments from Remsen:
"NO. In view of tom and Bret's comments above, I change my vote from YES
to NO. When someone with Bret's experience and instincts has actually handled
the type specimen and has his doubts, that's enough for me, and Tom's
philosophical points are on target."
New comments from Zimmer:
"Both Tom and Bret's points are well-taken, and indeed, I think I conveyed
my own general reluctance to recognize anything based on only a single
specimen. I also want to state that I respect Bret's informed views on this
particular situation. However, I think a few of his points call for
clarification. I don't think that proposals in the literature to elevate obrieni to
species status are "ludicrous". Rather, they seem natural given the
uniqueness of the type specimen and the huge range disjunction from any known
population of Celeus spectabilis, the species with which obrieni was
lumped by Short. Short also examined the type of obrieni, and
compared it with all other Celeus at AMNH. He obviously
thought the specimen was different enough (i.e. not clearly a mutant or hybrid)
to warrant the creation of a new subspecies. Given the range disjunction and
the biome from which the specimen originated, I'm surprised that he decided to
describe obrieni as a subspecies rather than as a distinct
species, but my point is the same: an experienced taxonomist looking at the
specimen against a backdrop of all other Celeus still decided
it represented a distinctive taxon rather than a hybrid or mutant of some type.
“I think that all too
often we tend to invoke the hybrid theory when confronted with single specimens
of distinctive taxa. For some groups, in which hybridization is a proven
frequent phenomenon, such speculation may be justified. But for most other birds,
I think it is less likely that someone collected the ultra-rare hybrid than
that the specimen is of a valid taxon whose range and/or microhabitat we simply
haven't managed to pin down. The validity of Pithys castanea was
constantly questioned until it was rediscovered by LSU personnel. Ditto for Hemitriccus
inornatus, Hemitriccus (Todirostrum) senex and Pipra vilasboasi.
What about Conothraupis mesoleuca? It remains known from a single
specimen (less distinctive than Celeus obrieni). Is it ludicrous that we
continue to recognize it as a distinct species? Had Short described obrieni as
a distinct species rather than as a distinct subspecies, it would be in the
same boat as the Conothraupis -- an enigma that no one has been
able to re-find despite much searching in the vicinity of the type locality.
“We also have to ask what
are the likely parents that would produce this hybrid. The only Celeus that
we found in the region was Celeus flavescens ochraceus. The
other Celeus species that Bret mentions (torquatus, flavus)
are humid forest or gallery forest bird, and this habitat is lacking from the
region. To accept the hybrid theory, we must not only accept that this rare
occurrence (i.e. two Celeus species interbreeding and
producing a spectabilis-looking hybrid) took place, but that the
resulting hybrid was some sort of vagrant that made its way out into
inhospitable habitat that was inconsistent with the habitat occupied by its
parent forms.
“While Bret found the type
specimen of obrieni odd (i.e. possibly a mutant, hybrid, or
backcross), his examination of it obviously wasn't enough to deter him from
making the effort to search extensively for the bird near the type locality
(which isn't exactly on your way to anywhere). Based on my own time in that
region, I agree with Bret's assessment that there is no habitat typical of Celeus
spectabilis. This proves nothing, because we don't have any real reason to
think that obrieni would be ecologically similar to spectabilis
anyway. Short placed obrieni with spectabilis on
the basis of plumage characters alone. I don't agree with Bret regarding the
lack of any real alteration to the native habitat since the time of Kaempfer.
As Jose Maria has noted, much of the tall forest from Piauí has disappeared. A
similar assumption that habitat loss could not explain the disappearance of
Spix's Macaw predominated for some time, based on the abundance of caatinga
extant within its former range. It is now known that the macaw was intimately
tied to a critical abundance of caraiba trees; a microhabitat within a habitat.
For a big woodpecker, for which potential nest trees could be a limiting
factor, it seems entirely conceivable that destruction of tall, semi-deciduous
forest or systematic removal of larger trees from remaining stands of such
forest could explain a massive decline in a specialized species. I found
caatinga and dry forest to be present over large areas of Piauí. At the same
time, I found only remnant stands of taller forest, which supports Jose Maria's
observations.
“None of this proves that obrieni is
a distinct species, but I do think that the only reason we are having this
conversation is because of the "accident" that Short described it as
only a subspecies in the first place. If the hybrid/mutant argument prevails,
then I see no more justification for recognizing obrieni as a
subspecies of spectabilis than I do for recognizing it as a
distinct species. To subscribe to the hybrid/mutant theory or the "can't
recognize anything based on a single specimen argument" would invalidate
recognition of obrieni at any level. If the bird is a legitimate taxon, I don't
see how it can possibly belong with spectabilis. To maintain the
status quo until the bird is discovered in life, is to essentially validate
Short's description of the bird as a distinct taxon. Once you've accepted that
part, the elevation of obrieni to species-level is not such a
leap.”
New comments from Silva:
"After reading carefully all new comments, I think I will keep my vote on obrieni."
More comments from Bret: "I
guess I'll add a few more thoughts on the Celeus obrieni proposal,
especially since Kevin called for some clarification of my comments.
First, I apologize for the word "ludicrous". When I saw
it written back at me, it did look harsh.
“Hey, I don't know what
this specimen represents. Without a molecular analysis of it, I doubt we
are going to "find out". Now, if someone does manage to prove
that it exists in the wild, that's great. But that hasn't happened despite
some good observers looking at the single known locality of occurrence (and
Kaempfer's locality *can be trusted*; as I recall, his notes showed that he was
at the end of an extended stay at Uruçuí when he collected it, and it looked to
me like he must have been close to town). For sure, Kevin, I thought it was
worth going to the type locality to look for the bird. It is a distinctive
looking bird. And Celeus woodpeckers, especially those in relatively
open habitats, are almost certainly going to be heard or seen within three
days. (However, I wouldn't call that
specimen, or most other Celeus, a "big woodpecker").
“Where does this leave us?
Well, someone could write a paper actually evaluating the specimen and
all associated data and reasoning. The conclusion could be that it is a
species-level taxon, or that it is a mutant/hybrid. I think that's it for
possibilities (unless the conclusion is that it's not possible to pin it down
on present knowledge, surprise). Short
decided it was a lot like spectabilis and called it a subspecies;
nobody seems very happy with that. But Tom's got it quite right when he
says that elevating it to species level without anything more than "it's
disjunct and looks distinctive" is too arbitrary (not that we can't be a
little arbitrary in some cases). So, I continue to think that if anyone wants
to elevate obrieni to species level, then they should publish a
well-balanced, objective argument for it. That argument will have nothing
to do with whether or not it was originally described as a species or
subspecies.
“Some points to be
considered, among many others: Yes, if it is a simple F1 hybrid, it cannot be
explained. I don't think it could be this kind of hybrid, and I'm not
arguing that it is a more complicated (and infinitely rarer) type of hybrid. I will comment, however, that a wandering
individual torquatus or flavus in this region is not too
far-fetched. There is a mosaic of
habitats, and I suspect that populations of these species could be found not
terribly far away. Should such an
individual end up mating with a "resident" flavescens, and if
this were to happen again with another species... dream on. At least this can be checked! So, let's see *who it's mother was* -- if it
is clearly a named species, then the case for obrieni as a valid
taxon at any level is out... with one exception. If the mother was especially close to spectabilis,
there is a good argument for either splitting obrieni as a
species [disjunct, distinctive sounds good now] or leaving it as a well-marked
subspecies, depending on where one draws that line (and it looks like most/all
of us, including me, would call it a species). I am not arguing that it
is a mutant displaying an odd combination of Celeus-like characters,
though I think that is possible. I *am* arguing that we cannot be sure
what it is, *especially in the absence of a good, detailed analysis of the
situation*.
“It should be noted that
there is no caatinga anywhere near the type locality. There is no
"transition to caatinga" anywhere near the type locality. This entire region of Piauí and Maranhão is a
mosaic of cerrado, cerradão, palms, and somewhat more mesic woodland. This is reflected clearly on vegetation maps
and in the avifauna present there -- right, Kevin (or am I missing something)?
In 1999 (I think that's when I was there), it was all in pretty darned
good shape, with extensive cleared areas only around Uruçuí itself (along with
some nice habitat): basically, I was impressed. The area was on the verge of massive
alteration, however. The tractors were already at work. Farmers from Paraná, in particular, had bought
up vast tracts of land and were planning to level it all for soybeans within
the next few years; highly mechanized operations. I talked with several
of these guys, independently, always by accident, always without really wanting
to. (Never stop for a beer where there are 0 women present. Or only 1 woman.
And certainly don't do it more than once.) Thus, aligning this locality with caatinga
areas of endemism and suggesting that obrieni could be a species
that fits some caatinga-related pattern, calling it "Caatinga
Woodpecker" is inappropriate at best.
“The idea that we are
worried about this at all is due to it being described as a subspecies doesn't
look at the other side of the coin: One might well have submitted a proposal to
suggest that there are some reasonably good points in favor of C.
obrieni not being a good species. Lumping? Yes, that's always right there besides
splitting.
“Finally, somewhat
sheepishly: I saw a female Conothraupis
mesoleuca at P. N. Noel Kempff Mercado back in about March 1993. It
sat in good view for more than a minute. his was not far from the type
locality, and I feel confident that it couldn't have been a speculigera though
it sure looked a lot like it, mostly being grayer, less greenish as I recall. Since the female is undescribed, and I've been
meaning to get back for a more careful look at the area, I've just let it
slide. I am a strong proponent of
getting documentation for stuff like this; I still don't have it. No
further excuse.
“Thanks to all for
considered opinions, but this is the last from me on this one."
Comments from Nores:
"[YES] Si. Las razones para reconocer a un
nuevo taxa son similares, ya sea para especies o subespecies. Si uno acepta que
el único ejemplar conocido tiene características diferentes como para
aceptarlo, y estas características son a nivel de especies, me parece que lo
lógico es considerarlo como especie."