Proposals (222) to South American Classification Committee

Split Atlapetes nigrifrons from A. latinuchus

Summary of Proposal: This proposal, if passed, would result in the splitting of one of the many taxa currently found in "Yellow-breasted Brush-Finch" / "Northern Rufous-naped Brush-Finch" Atlapetes latinuchus. Atlapetes l. nigrifrons is endemic to Serranía de Perijá of northern Colombia and Venezuela and is the northernmost taxon in A. latinuchus. It was formerly known as A. l. phelpsi. Cladistic analysis using a limited number of morphological (plumage) characters carried out in connection with the recent description of A. l. yariguiorum suggested that A. l. nigrifrons was more closely related to Santa Marta Brush-Finch A. melanocephalus than to other forms. It was further suggested that A. l. nigrifrons is probably more closely related to Moustached Brush-Finch A. albofrenatus than to any A. latinuchus taxa (Donegan & Huertas 2006).

Taxonomic note: A. l. nigrifrons was described by Phelps & Gilliard (1940). It was later assigned the epithet A. l. phelpsi by Paynter (1970) because A. torquatus nigrifrons was then senior. However, with A. torquatus now assigned to Buarremon (see further Hackett 1993; Remsen & Graves 1995a; Remsen & Graves 1995b and followed by almost all recent authors), phelpsi is a junior synonym of nigrifrons (Dickinson 2004).

Discussion: The following discussion is cribbed from the discussion of species limits in northern Atlapetes in Donegan & Huertas (2006). We carried out an analysis of species limits in northern Atlapetes based on morphological data (plumage characters and biometrics) in connection with the description of new taxon Atlapetes (latinuchus) yariguierum with the objective of describing that taxon at a suitable taxonomic level. This analysis was also undertaken to analyse the relationships of a new species, undescribed colour morph or hybrid from the Perijá mountains found in a museum during the study ("Perijá bird") which will be discussed and described in more detail in a future publication.

SACC committee members should note that the plate of A. l. nigrifrons in the only major reference work in which it is illustrated (Hilty 2003) is poor. For those not familiar with this form, A. l. nigrifrons is akin to Santa Marta Brush-Finch A. melanocephalus but with a light grey back and rufous on the rear and central section of the crown. Its grey cheeks, black forehead and black face (malar merged with crown) present a very different facial pattern to that of all other A. latinuchus but is broadly similar to A. melanocephalus (which lacks red on the crown). Its back colour (light grey) also contrasts with that of all northern forms, especially the closest populations, A. l. yariguierum and A. l. elaeoprorus, which have almost jet black backs in adult plumage.

García-Moreno & Fjeldså (1999) recently re-evaluated species limits within Atlapetes using molecular data. The taxonomy of the group was previously based on morphological analyses (Paynter 1972, 1978), with the taxa Slaty Brush-Finch A. schistaceus and Rufous-naped Brush-Finch A. rufinucha delimited largely on the presence of grey (schistaceus group) or yellow underparts (rufinucha; pileatus group). Previously, Remsen & Graves (1995a) had suggested that several geographically close taxa with alternating grey and yellow underparts may be more closely related to one another than to more geographically distant taxa with similar underpart coloration. García-Moreno & Fjeldså's phylogeny supported Remsen & Graves' propositions, and suggested further that many sister taxa should be ranked specifically.

The northern taxa of Yellow-breasted Brush-Finch A. latinuchus, Slaty Brush-Finch A. schistaceus, Moustached Brush-Finch A. albofrenatus and Santa Marta Brush-Finch A. melanocephalus were not studied in detail by García-Moreno & Fjeldså (1999), but various taxa formerly considered part of A. rufinucha, namely baroni, caucae, chugurensis, comptus, elaeoprorus, latinuchus, nigrifrons (=phelpsi: see above), simplex(=spodionotus: Donegan & Huertas 2006) and spodionotus, were tentatively reassigned to the grouping A. latinuchus. García-Moreno & Fjeldså noted 'Our study did not include various more richly coloured forms in the northern Andes However, it is evident that the current sequence does not reflect natural groupings'.

We conducted a cladistic analysis on the northern taxa based on an admittedly small number of plumage characters (17). Analyses of southern Atlapetes taxa using this data set showed phylogenies generally consistent with García-Moreno & Fjeldså (1999) contra Paynter (1978). The authors recognised the limitations of a study involving so few characters, especially in a group in which pigmentation can be phylogenetically less informative than in other groups. However, an analysis encompassing a broad range of plumage characters (cf. Paynter, 1978) can be useful, if interpreted appropriately and conservatively.

Three principal multi-taxa clades, each rooted from the same node, were identified in the strict consensus tree for northern taxa as follows: (i) A. schistaceus, subspecies castaneifrons, fumidus, schistaceus, taczanowskii and tamae; (ii) A. latinuchus, subspecies baroni, caucae, chugurensis, comptus, elaeoprorus, latinuchus, simplex (=spodionotus), spodionotus and (new taxon) yariguierum; and (iii) A. melanocephalus, A. latinuchus nigrifrons and the "Perijá bird". The grouping of A. l. nigrifrons with A. melanocephalus and the Perijá bird was supported by bootstrap (83%); the others were not. All A. fuscoolivaceus, A. tricolor and A. albofrenatus taxa were unresolved in the strict consensus tree at the same level. A tree was not reproduced as we did not propose a phylogeny given the small number of characters used in the study.

We refrained from suggesting a wholesale revision and splitting of A. latinuchus, whilst noting that probably several biological and phylogenetic species are involved. We described A. l. yariguiorum within the A. latinuchus complex as a result rather than as a separate species. We, however, suggested splitting of A. l. nigrifrons as a clear anomaly in the group.

From the analysis above, A. melanocephalus and A. nigrifrons would provisionally appear to be more closely related to one another than either is to A. latinuchus taxa, a proposition supported by biogeographical and morphological considerations. Atlapetes nigrifrons and A. melanocephalus are both restricted to the northernmost Colombian and Venezuelan mountains: the Perijá and Santa Marta ranges. Studies by Remsen & Graves (1995a) and García-Moreno & Fjeldså (1999) suggest that some geographically close but morphologically distinctive Atlapetes taxa are more closely related to one another than they are to superficially similar but more geographically distant taxa. Paynter (1978) drew attention to the morphological similarity of the forms A. l. nigrifrons (then A. l. phelpsi) and A. melanocephalus, which share a black forehead (extending to a black crown in A. melanocephalus), black chin, lack of light moustachial markings (the malar merged with the mask), distinctly greyish cheeks and a lighter grey back, features not found in northern A. latinuchus taxa.

A lineage as follows:

A. a. albofrenatus (green back; moustachial stripe; black forehead; red crown)

A. sp. (Perijá bird) (green back; moustachial merged into malar; black forehead; red crown)

A. l. nigrifrons (grey back; moustachial merged into malar; black forehead; red crown)

A. melanocephalus (grey back; moustachial merged into malar; black forehead; black crown), although not borne out in phylogenetic analysis where A. albofrenatus taxa behaved rather counter-intuitively, appears a more plausible hypothesis than any closer relation between A. l. nigrifrons and northern A. latinuchus taxa.

The assignment of A. l. nigrifrons to the A. latinuchus species grouping appears to be a clear example of the current sequence not reflecting natural groupings, per García-Moreno & Fjeldså (1999). If it is not to be assigned species rank, a better placement for nigrifrons would be in either Santa Marta Brush-Finch A. melanocephalus or Moustached Brush-Finch A. albofrenatus. Other possible approaches, lumping A. melanocephalus and/or A. albofrenatus into A. latinuchus, would not be sensible. Significant morphological differences exist between A. latinuchus (less A. l. nigrifrons), A. l. nigrifrons, A. melanocephalus and A. albofrenatus, these being considerably greater than those between various southern forms now considered separate species. Under the comparative approach for assessing allopatric populations of birds under the biological species concept (Helbig et al. 2001), the split would appear to be warranted. The status of A. l. nigrifrons under the phylogenetic species concept should also be beyond any doubt.

English name: We proposed Perijá Brush-Finch as an appropriate vernacular name for A. nigrifrons, given that it is restricted to that mountain range and morphologically driven names are notoriously difficult to coin in this group.

Arguments against splitting A. l. nigrifrons:

Arguments against splitting of this taxon could include: (i) the small number of characters used in the morphological study; (ii) the lack of molecular data; (iii) the lack of vocal data; and (iv) general lack of agreement with or suspicion over evidence for the Remsen & Graves / García-Moreno & Fjeldså model for speciation in this group (evidenced in certain comments on SACC proposals 86 and 87).

As regards (i), it is worth re-iterating that the A. nigrifrons/A. melanocephalus / Perijá bird clade nonetheless had strong bootstrap support in our study. Each of the relationships in our study of southern taxa that were supported by bootstrap involved taxa that molecular studies suggest are more closely related to one another than to other taxa (per García-Moreno & Fjeldså, 1999).

As regards (ii), a molecular study of this group is in progress led by J. Klicka, C. D. Cadena & J. L. Pérez-Emán, some preliminary results of which relating to A. l. yariguierum were kindly shared with us for the description. Following review of our manuscript, these researchers have recently sequenced the Perijá bird and A. l. nigrifrons to clarify the relations of these taxa. Whilst unpublished data is apparently not to be taken into account by SACC members, C. D. Cadena has recently informed us (and consented to us noting in this proposal) that their molecular data also support the proposition that A. l. nigrifrons is more closely related to A. albofrenatus and the Perijá bird than it is to A. latinuchus taxa. This is mentioned not to buttress the proposal, which should stand or fall on the morphological analysis presented, but to reassure committee members that the re-lumping of A. l. nigrifrons into A. latinuchus is not likely to be recommended in the future.

Atlapetes l. nigrifrons is much more different morphologically from A. latinuchus (and A. melanocephalus and A. albofrenatus) than many sister Atlapetes now treated as separate species by SACC (whether or not all members concur fully with the approach taken to southern taxa). Further, A. latinuchus would certainly be paraphyletic if A. l. nigrifrons is maintained within it. And no-one has ever seriously advocated lumping either A. albofrenatus or A. melanocephalus with A. latinuchus in the past due to the marked differences between these taxa (including A. l. nigrifrons in A. latinuchus).

The approach we took in Donegan & Huertas (2006) is a conservative one, proposing the splitting of just one clearly anomalous member from the A. latinuchus complex when one could conceive arguably strong arguments for much more aggressive splitting of A. latinuchus (and A. albofrenatus), even on the basis of comparative morphological evidence. However, we felt that it would be better to await the molecular study in advance of further splitting.

Proposal: A "YES" vote would split Perijá Brush-Finch A. nigrifrons from A. latinuchus. A "NO" vote would retain it within A. latinuchus (or move it into A. albofrenatus or A. melanocephalus given that subspecies are not listed on SACC). I would recommend a "YES" vote for the reasons stated in Donegan & Huertas (2006) and set out above.

References:

Donegan T.M. & Huertas B.C. 2006. A new brush-finch in the Atlapetes latinuchus complex (Passeriformes: Emberizinae) from the Yariguíes mountain range and adjacent Eastern Cordillera of Colombia. Bulletin of the British Ornithologists' Club 126(2): 94-116.

For a .pdf, go to: http://www.proaves.org/IMG/pdf/Donegan_Huertas_Atlapetes_latinuchus_yariguierum-2.pdf

Other references are all cited in Donegan & Huertas (2006) and most are on SACC baseline.

Thomas Donegan, June 2006

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Comments from Steve Hilty: "Comments to voting committee members on the Donegan proposal to split Atlapetes latinuchus nigrifrons as a separate species.

"There are some confusing and unresolved taxonomic issues with the Donegan and Huertas paper (2006, BBOC) and specifically with the SACC proposal #222 which relies solely on a weakly constructed set of plumage characters for taxonomic conclusions. The authors may (or may not) have a point in elevating one or more of these northern populations of the Atlapetes latinuchus complex to species level (SACC proposal #222), but the evidence, presented in part in the BBOC paper, and more fully articulated in the SACC proposal, does not seem convincing. Here's why:

"The facts:

1) A new subspecies, Atlapetes latinuchus yariguierum is described from a western spur of north end of Eastern Andes which has black back and richer coloration overall than allies to the south.

"2) Several forms of Atlapetes latinuchus more or less from Bogotá southward are grouped together because they are mostly paler in coloration than the new yariguierum. Some of these might eventually be regarded as species using somebody's concept. Donegan and Huertas include yariguiorum in this "swarm" of taxa.

"3) Another population of Atlapetes latinuchus now designated as A. l. nigrifrons (formerly A. l. phelpsi) occurs in the Serranía de Perijá. Nobody knows a thing about it in life and probably nobody will for some time to come. This is the taxon (SACC proposal #222) being proposed as a full species.

"The description of the new subspecies (item 1) and other conclusions in the BBOC paper appear to be well researched. However, the SACC proposal, which takes these conclusions a step further, does not appear well supported by the evidence presented.

"The facts subject to interpretation:

In SACC proposal #222 Donegan and Huertas propose elevating the Perijá population of Atlapetes latinuchus nigrifrons to a full species, stating that it is closer in appearance to Atlapetes melanocephalus, a Santa Marta endemic, and perhaps also to Atlapetes albofrenatus (which itself is composed of a non-overlapping population in the Mérida Andes of Venezuela, and another in the north end of the Eastern Andes of Colombia), than to other subspp. of latinuchus southward. One of the reasons for the split is their claim that nigrifrons looks more like Santa Marta birds than other Atlapetes latinuchus populations to the south. SACC voting members are specifically told to note that ."the only major reference work in which it [A. l. nigrifrons] is illustrated (Hilty 2003) is poor. They state (twice) that A. l. nigrifrons, has a "light gray back" which they say distinguishes it from yariguiorum and other taxa southward (and apparently by implication aligns it more closely to A. melanocephalus). This is incorrect. The Phelps collection in Caracas has a considerable number of specimens of nigrifrons (formerly phelpsi) and, unless they are all misidentified, the back color of the adults is very dark gray or slate gray; in Ridgway nomenclature, it matches chaetura black and dark mouse gray fide R. Restall, and this shows well in Restall's excellent paintings of the various subspp. of A. latinuchus in his forthcoming book on the birds of northern South America. Furthermore, in my judgment (and Restall's description of a large series in COP fide Restall), the back color of nigrifrons has a slight (but distinct, fide Restall) olive tone, which is denied in Appendix 3 of Donegan and Huerta's BBOC paper.

"In any case, calling the back color light gray is a stretch and I believe Gwynne's illustration is accurate although the narrow black forehead does not show as well as it could. In any case, my text description is correct and should eliminate any reader/viewer misconceptions regarding the plate. The similarity mentioned by Donegan and Huertas of A. l. nigrifrons to A. melanocephalus is tenuous. Although both taxa have grayish cheek patches, Perijá birds have rufous crowns with black foreheads while Santa Marta birds have no rufous on the crown at all, a dramatic difference. Picky details aside, where does this get us?

"I suggest that some of the interpretations above are premature and ought to include supporting molecular data and/or vocal comparisons before embarking on piecemeal schemes to redraw taxonomic boundaries in a group of birds this complex. Dawn songs of Atlapetes are quite distinctive and should be helpful in an analysis of this type although in their BBOC paper a sound recording (apparently not a dawn song) was more or less dismissed as . . ."The call is typical of the genus." I have personally recorded dawn songs of most Venezuelan Atlapetes and some other species southward, and while day vocalizations are often variable, as these authors suggest, dawn songs have highly species specific patterns." An analysis of these songs, along with molecular data, could help support or refute some of the questions Donegan and Huertas are attempting to answer and this information even might strengthen their case.

"At this point one might just as easily align the Perijá birds with the new Atlapetes latinuchus yariguiorum because both have blackish or dark-backs, both share all or mostly rufous crowns, neither has a white wing speculum, and the two populations are not separated by a geographical barrier that is as significant as either is to the Santa Marta birds. Alternatively, one could suggest other revisions-but until appropriate supporting data are mustered, such suggestions are unlikely to have much traction and do not provide a solid basis for advancing taxonomic understanding. Now, any volunteers for a one-way expedition up the Perijás to get some tissue and that critical dawn song?"

Response to Hilty comments from Donegan: pdf (which contains color photos of specimens).

Comments from Cadena: "YES. This proposal is somewhat "unorthodox" in that it uses only phylogenetic analyses of plumage characters to propose that a taxon currently ranked as a subspecies merits species status, something that I don't think has been done in any other case before. Also in contrast with many recent proposals, this one does not present any data on vocalizations, and the genetic information it refers to has not been published.

"I must say I felt a bit uneasy when Donegan decided to employ plumage characters in a phylogenetic analysis of Atlapetes, considering that based on work by García-Moreno and Fjeldså (1999) it appeared that plumage variation is a poor indicator of phylogenetic relationships in this group. However, it is reassuring that some of the results are consistent with relationships inferred using molecular data, suggesting that when multiple plumage characters are considered and attention is given to patterning vs. general coloration (i.e. not simply grey vs. yellow overall appearance), these traits may be phylogenetically informative. This, of course, was foreshadowed by Remsen and Graves (1995), when they predicted that pairs of yellow-bellied ("rufinucha") and grey-bellied ("schistaceus") birds were likely each other's closest relatives based on plumage patterns. In other words, the problem has not really been that plumage variation is useless, but that it needs to be assessed properly. Thus, I think I can accept the conclusion based on morphology that nigrifrons is more closely allied to other species of Atlapetes than to latinuchus, which would imply that nigrifrons should be ranked at the species level unless one lumps multiple species into one. Steve Hilty's comment raises several important concerns, but I believe those related to plumage are satisfactorily addressed by Donegan in his rebuttal (I will comment on other concerns such as lack of additional data below). I would also add that Hilty's emphasis on the distinctive rufous crown of nigrifrons in comparison to melanocephalus is not a very strong argument because according to the framework applied by Donegan (i.e. a phylogenetic, parsimony analysis) what really matters in establishing the purported close relationship between these taxa are characters representing shared and derived traits (e.g. moustachial markings). I should note, however, that the taxon sampling by Donegan and Huertas is not ideal: the set of taxa they worked with does not form a monophyletic group, and the use of Buarremon as an outgroup, which followed Garcia-Moreno and Fjeldså, is clearly inadequate (Buarremon is certainly not the sister group of Atlapetes, these two genera are actually quite far apart from each other in the working phylogeny of the Emberizinae by John Klicka et al. I don't know whether the sparse sampling or the inadequate outgroup choice may influence the outcome of the analyses my hunch is that it does not.

"In his proposal, Donegan mentioned our (i.e. John Klicka, Jorge Perez, Garth Spellman, and myself) unpublished mtDNA work on Atlapetes, which indeed suggests that nigrifrons is more closely allied to albofrenatus (and other taxa, including melanocephalus and albofrenatus as suggested by the morphological analysis but also semirufus, which is very distinct in plumage) than it is to members of the latinuchus complex occurring in Colombia (e.g. elaeoprorus, yariguierum, spodionotus). We now have sequence of nigrifrons from both the Colombian and Venezuelan side of the Perija (total = 3 individuals) and it appears that this taxon does not form a reciprocally monophyletic mtDNA group with respect to nominate A. albofrenatus (we do not have data for A. a. meridae yet but should have some very soon). Considering how different albofrenatus and nigrifrons are in plumage (with the caveat that Donegan's "Perijá bird" may be a hybrid or intermediate), this probably indicates that these taxa split recently from each other (i.e. incomplete lineage sorting), not that they are conspecific. However, it would be interesting to know whether albofrenatus and nigrifrons differ vocally, but this is not possible owing to the lack of recordings of the latter. In any event, nigrifrons definitively does not belong with latinuchus.

"Our work, which so far includes complete sequences of cytb and ND2 for all but three species of Atlapetes and many distinctive subspecies, suggests that many (most?) of the currently recognized polytypic species of Atlapetes (i.e. not only the "rufinucha" and "schistaceus" types) are poly- or paraphyletic at the mtDNA level. Either traditional taxonomy in this genus is substantially flawed (being misled by highly labile plumage traits), or mtDNA variation does not track phylogeny properly owing to rampant hybridization or rapid radiation and incomplete sorting. I tend to think that the problem is more related to incorrect taxonomy for reasons we shall expose once the relevant publications are out. This is all to say that Atlapetes taxonomy is a mess that will require multiple changes to be consistent with evolutionary history. The proposal by Donegan, although it is not yet backed up by published genetic data (eventually it will), seems like a logic first step in the right direction. The proposal might have been a bit premature in that the genetic data are not yet published, but I am sure that if this proposal is rejected because of this, we will need to revisit it and overturn such a decision in the near future when the genetic data are out. Thus, I would recommend a YES vote on this one."

Comments from Jaramillo: "YES - Donegan makes a good argument, and supported by soon to be published molecular data by Cadena. That is satisfactory for me, but I appreciate Steve Hilty's concerns on the subject. Given what a mess Atlapetes is, any change or analysis raises troubling issues, this one is no different. However, the evidence is good for making this split."

Comments from Stiles: "NO for now; in this case, best to wait until more data are in and then do it right. I suspect that Donegan is correct, but share Hilty's reluctance to tinker piecemeal with what clearly is a more complex situation. As I understand it, Daniel (and others?) are working on genetic data for this group, and I prefer to await this before making changes that might have to be modified again within a few months or years. The case is much like that of Momotus, in which we voted down recognizing aequatorialis because it represented only one piece of a more complicated puzzle."

Comments from Robbins: "NO. Steve Hilty provides information that questions Donegan and Huerta's interpretations, and I agree with his rationale for not splitting this species until there is unequivocal data to support this change."

Comments from Pacheco: "NO. Fortemente influenciado pelos argumentos de Hilty e até que estudos (abrangendo dados adicionais) sejam tornados disponíveis."

Additional comments from Thomas Donegan: "For fear of flogging a dead horse, I find certain of Stiles, Robbins and Pacheco's comments as baffling as Hilty's.

"First in relation to "difficult parts of a more complicated puzzle" per Stiles, such a comment would apply were we to have proposed A. l. yariguierum or any of the other races for species rank. We did not. The situation regarding many Atlapetes taxa is indeed a complicated puzzle and the latinuchus group is particularly difficult. This is why we described A. l. yariguierum as a subspecies, against the advice of some peer reviewers, despite it clearly being a species under some species concepts (e.g. some versions of PSC) and, as future research may or may not demonstrate, possibly under others. Although Atlapetes latinuchus is a difficult puzzle generally, in the case of A. l. nigrifrons, one piece is quite clearly in the wrong box. This is therefore rather different from Momotus.

"As for endorsement of Hilty's morphological-based assertions by Robbins and Pacheco, could I please encourage those committee members to re-consider with an open mind the attached .pdf file that includes photographs of the relevant taxa and a discussion. With one more "no" vote, SACC would seek to perpetuate Phelps & Gilliard's and Paynter's proposed relation between A. l. nigrifrons and an apparently little-related Peruvian taxon A. melanolaemus, both formerly in A. rufinucha. Especially in the light of Cadena's comments, there seems no logic in doing so other than on grounds of pedantry / technicality (i.e. do it again in a few months when the published molecular study is out). In any event, what is the point in perpetuating a "status quo"? When peer reviewed research is published, surely that becomes the status quo until someone shows otherwise. Like the molecular study, Hilty's comments are also unpublished. Rejection for technicality is fine with me. However, could I please request that Robbins and/or Pacheco respond to the rebuttal stating what they disagree with in it and why they consider the interpretations to be erroneous and those of Hilty to be better? Even off-line would be much appreciated.

"I should also clarify that Cadena et al.'s molecular and our morphological results in relation to this taxon were entirely independently reached."

Additional comments from Cadena: "I see that several committee members are voting against Donegan's proposal to split A. nigrifrons on the basis of (1) comments by Steve Hilty and (2) lack of additional data. Like I said in my comments, I think that Donegan responded to Hilty's comments satisfactorily, and I would encourage committee members to read Donegan's response once again. Regarding the lack of additional data, it seems as though Gary had not yet seen my comments when he submitted his vote. We do have sequence data to address this question, and these data certainly show that A. nigrifrons does not belong in A. latinuchus: it is more closely allied to A. albofrenatus. I realize that our molecular data set is not out yet, so perhaps this is a reasonable argument to turn down the proposal. However, I would like to reiterate that if turned down now, we will need to revisit the proposal and accept it once our molecular data set is out. I can understand that committee members may feel this is best, but I'd rather do this change now, knowing it is well-justified, than have to go through all the process once again a few months down the road (Note that this proposal has been posted for several months already). I know that conservation considerations should not drive taxonomic decisions, but here we are talking about a restricted-range taxon occurring in area posing multiple threats. Now that the SACC is the official authority for organizations such as BirdLife, ranking nigrifrons at the proper taxonomic level (i.e. as a species) sooner than later can only help conservation action (or at least the development of proper categorizations of threatened species)."

Comments from Zimmer: "YES. The unpublished molecular work of Cadena et al. is the tipping point for me. While I appreciate the desire to have a clearer picture of the entire complex rather than making piecemeal changes, I think that sometimes you have to take a piecemeal approach just to move things along. Large, complex groups such as this one tend never to get worked out, simply because the task of taking on the entire complex is so daunting (see for example Sittasomus or Tolmomyias sulphurescens). In the present case, we have indication that molecular evidence supports the morphological analysis, so I think we should go with it."

Comments from Stotz: "YES. While I am sympathetic to the idea that we shouldn't work on a group piecemeal, we've already started doing that with Atlapetes (see proposals 86-87, which are a change from the status quo in my view), and Kevin's. Given that Daniel's molecular data appears to agree with Donegan's treatment, and he recommends a yes vote, despite it not being published, I think we should follow that recommendation."

Comments from Remsen: "YES. I find Donegan's rationale consistent, and I am comforted by that the genetic analyses confirm it."