Proposal
(29) to South American Classification
Committee
Remove Euphonia
and Chlorophonia from Incertae Sedis and place as subfamily Euphoniinae in
Fringillidae
Effect on South American
CL: We currently treat the genera Euphonia and Chlorophonia as
Incertae Sedis within the 9-primaried oscines. This proposal would move them to
Fringillidae, as a subfamily (and therefore also force recognition of subfamily
Carduelinae for the fringillids in South America).
Background:
Although Euphonia and Chlorophonia have always been
placed in Thraupidae, several aspects of their biology have marked them as
"weird." The complex, melodious songs of some species have been noted
as superficially "goldfinch-like" and unlike that of most tanagers.
The presence of pronounced vocal copying in E. violacea, E. laniirostris,
and E. pectoralis (Snow 1974, Morton 1976, Remsen
1976, Sick 1997) is unlike any tanager but reminiscent of several fringillids
(e.g., Mundinger 1970, Remsen et al. 1982, Taylor 1979). The domed
nest with side entrance is unlike that of any typical tanager (Isler and Isler
1987). The vagility of some (e.g., E. chlorotica, C. cyanea, C. flavirostris)
seems unusual for tanagers but typical of fringillids, as do reports of large
single-species flocks in at least one species, C. flavirostris.
The near-vegetarian diet recalls that of many fringillids, and their stomach
"structure" is bizarre. They feed their young by regurgitation,
unlike (?) tanagers but like many fringillids. None of these features, however,
has been accorded any phylogenetic significance.
New data: Four
labs have independently shown that Euphonia does not belong in
Thraupidae but clusters with Fringillidae (Burns 1997, Burns et al. 2002,
Klicka et al. 2000, Sato et al. 2001, and Yuri and Mindell 2002). My only
hesitation is that these results are based mostly on one gene, cytochrome b.
Although Sibley & Ahlquist (1990) retained Euphonia within
their Thraupini, that branch was the deepest in the group, which also
included Piranga, Habia, Coereba, and Sicalis.
Recommendation: I
recommend a YES on this one. The AOU Checklist Committee has unanimously
approved a similar proposal by Jim Rising, and created a subfamily taxon,
Euphoniinae, for these two genera within Fringillidae. Although I would feel
more comfortable if some sequence data came from a nuclear gene, I see no
reason other than historical momentum to keep these two genera in Thraupidae.
The additional phenotypic, albeit anecdotal, data is also consistent with a
fringillid connection.
Lit Cit:
BURNS,
K. J. 1997. Molecular systematics of tanagers (Thraupinae): Evolution and
biogeography of a diverse radiation of Neotropical birds. Molecular
Phylogenetics and Evolution 8:334-348.
BURNS,
K. J., S. J. HACKETT, & N. K. KLEIN. 2002. Phylogenetic relationships and
morphological diversity in Darwin's finches and their relatives. Evolution 56:
1240-1252.
ISLER,
M., AND P. ISLER. 1987. The tanagers, natural history, distribution, and
identification. Smithsonian Institution, Washington, D.C.
KLICKA,
J, K. P. JOHNSON, & S. M. LANYON. 2000. New World nine-primaried oscine
relationships: constructing a mitochondrial DNA framework. Auk 117: 321-326.
MORTON,
E. S. 1976. Vocal mimicry in the Thick-billed Euphonia. Wilson Bull. 88:
485-487.
MUNDINDER,
P. C. 1970. Vocal imitation and individual recognition of finch calls. Science
168: 480-482.
REMSEN,
J. V., JR., K. GARRETT, & R. ERICKSON. 1982. Vocal copying in Lawrence's
and Lesser goldfinches. West. Birds 13: 29-33.
SATO,
A., H. TICHY, C. O'HUIGIN, P. R. GRANT, B. R. GRANT, & J. KLEIN. 2001. On
the origin of Darwin's finches. Mol. Biol. Evol. 18: 299-311.
SIBLEY,
C. G., AND J. E. AHLQUIST. 1990. Phylogeny and classification of birds. Yale
Univ. Press, New Haven, Connecticut.
SICK, H.
1997. Ornitologia Brasileira. Ed. Nova Fronteira, Rio de Janeiro.
TAYLOR,
P. 1979. Interspecific vocal mimicry by Pine Grosbeaks. Can. Field Nat. 93:
436-437.
YURI,
T., & D. P. MINDELL. 2002. Molecular phylogenetic analysis of Fringillidae,
"New World nine-primaried oscines" (Aves: Passeriformes). Molecular
Phylogenetics & Evolution 23: 229-243.
Van
Remsen, 5 June 2003
________________________________________________________________________________________
Comments from Zimmer:
"YES. The preponderance of vocal, ecological, morphological and molecular
data all suggest this otherwise."
Comments from Schulenberg:
"YES. That said, I don't see that Burns 1997 or Burns et al. 2002 include
any fringillids among their outgroups. So, these two papers support the idea
that Euphonia and Chlorophonia are not tanagers (whatever a
tanager is), but do not directly address the current proposal. And, therefore,
we are down to three (not four) laboratories that report this result, not that
this matters much.
"Otherwise, I also
note that these studies generally compare a euphonia to a cardueline (although
a different species of Euphonia n each case), so our sampling
isn't very deep here. (Yuri and Mindell do a little better in this regard.)
Klicka et al. were cautious in suggesting that their results, for example,
indicate that "Euphonia (and by association Chlorophonia)
represents either a derived cardueline form or a basal, previously unrecognized
radiation within the nine-primaried oscine clade." So, I can approve the
proposal based on current knowledge. But I also wouldn't be surprised if down
the line, after the relationships of Euphonia and Chlorophonia
become the subject of an investigation (and not just a side issue to some other
question), we are asked to vote on recognizing a family Euphonidae, or
something along those lines. Time will tell."
Comments from Stotz:
"Tom's suggestion that we might eventually have a separate Euphoniidae someday
is interesting, and could be true. However, Yuri and Mindell did have stronger
taxon sampling relating to this question (six Fringillid genera) and Euphonia
did not end up basal in this group. Since that paper is based on just
mitochondrial DNA, it certainly is not the last word, but it does suggest that Euphonia
and Chlorophonia belong within Fringillidae proper. Another problem is
that based on Yuri and Mindell, Euphonia and Chlorophonia could
cluster within Fringillidae, in which case we'd have to do away with
Euphoniinae. Since the SACC list does not include subfamilies, I guess we don't
need to worry about this. Given that Euphonia clusters with Fringilla
in Yuri and Mindell, I suggest that Euphonia and Chlorophonia be
placed at the start of the of Carduelinae in their lists."
Comments from Stiles:
"YES (the evidence now seems solid enough to justify this change)"