Revise Phrygilus classification
Proposal: If passed, this proposal would divide the existing genus Phrygilus into four genera: Phrygilus + two genera resurrected from synonymy + two species merged into Idiopsar.
Background: The heterogeneity of members of the genus Phrygilus is widely recognized among field researchers. Thus, it is no surprise that genetic data reveal that it is highly polyphyletic. Here’s what our Note says:
6. The genus Phrygilus is highly polyphyletic (Klicka et al. 2007, Campagna et al. 2011). Campagna et al. (2011) found that the genus consists of at least four lineages: (1) gayi, patagonicus, punensis, and atriceps, which comprise the sister group to Melanodera (including extralimital Rowettia goughensis); (2) fruticeti, alaudinus, and carbonaria; (3) plebejus and unicolor, which are sister to Haplospiza; and (4) dorsalis and erythronotus, which are sister to Idiopsar. The type species for Phrygilus is gayi; Hellmayr’s (1938) synonymy indicates that Rhopospina Cabanis is available for Group 2, with fruticeti the designated type species, and that Geospizopsis Bonaparte is available for Group 3, with unicolor the designated type species. Clearly, major taxonomic revisions are needed but additional taxon sampling is needed within the Thraupidae. SACC proposal needed minimally to change linear sequence.
New information: The Campagna et al. (2011) paper (let me know if you need pdf) sampled all species in the genus plus a decent set of related taxa. The gene sampling was good, also: -- from their paper:
“We amplified a total of 3925 base pairs (bp) from five gene fragments using polymerase chain reaction (PCR). We included three mitochondrial regions, COI (694 bp), Cyt b (922 bp) and the control region (CR, 1050 bp); one Z-linked marker, intron 3 of the muscle skeletal receptor tyrosine kinase gene (MUSK, 678 bp); and one autosomal intron, intron 5 of the b-fibrinogen gene (Fib5, 581 bp). This combination comprises genes with a range of substitution rates with the objective of achieving resolution at both deep and recent nodes.”
The tree from their combined data is pasted below, and the results make sense from the standpoint of plumage, morphology, and distribution:
Analysis and Recommendation: Clearly, Phrygilus contains four separate lineages, and rather than expanding the existing genus to include an absurdly heterogeneous group, three new group names are needed. Two can be resurrected from synonymy, as indicated in the Note above. I think I might actually favor merging group 3 into Haplospiza, but for now, I think we should just resurrect Geospizopsis. Dick Banks has confirmed that both this and Rhopospina are available for groups 2 and 3. That leaves Group 4, for which recommend a merger into Idiopsar. Although I know I. brachyurus, I am not familiar with “P.” erythronotus or “P.” dorsalis, but I do know they share that same high-elevation, southern Andean, rocky puna and have roughly similar plumages. Idiopsar’s only striking feature is its big, long bill, but bill shape is an extremely labile character that has been over-emphasized in taxonomy.
In summary, I recommend we change the classification as follows:
Rhopospina fruticeti Mourning
Rhopospina alaudina Band-tailed Sierra Finch
Rhopospina carbonaria Carbonated Sierra Finch
Phrygilus patagonicus Patagonian Sierra Finch
Phrygilus gayi Gray-hooded Sierra Finch
Phrygilus punensis Peruvian Sierra Finch
Phrygilus atriceps Black-hooded Sierra Finch
Melanodera melanodera White-bridled Finch
Melanodera xanthogramma Yellow-bridled Finch
Haplospiza rustica Slaty Finch
Haplospiza unicolor Uniform Finch
Geospizopsis unicolor Plumbeous Sierra Finch
Geospizopsis plebejus Ash-breasted Sierra Finch
Idiopsar brachyurus Short-tailed Finch
Idiopsar dorsalis Red-backed Sierra Finch
Idiopsar erythronotus White-throated Sierra Finch
Diuca speculifera White-winged Diuca-Finch
Diuca diuca Common Diuca-Finch
This sequence rearranges the genera near Phrygilus in our classification according to the Campagna et al. tree in accordance with conventions of linear sequencing. Further changes are not warranted because of limited taxon sampling in Campagna et al. I also resequenced species with the genera in accord with conventions (less diverse lineage first; sister species NW-SE)
Idiopsar is masculine, so no endings change. Rhopospina is feminine, and so two variable endings change. Geospizopsis is feminine, but plebejus is invariable (see BBOC 131: 107). This has all been confirmed by Normand David (in litt.).
Notice that because “Sierra-Finch” no longer refers to a monophyletic group, we have to remove hyphens by our rules. For stability, I would recommend NOT inserting “Sierra” into Short-tailed Finch.
Note: I asked Leonardo Campagna if the proposal fairly represents his findings and if he agreed with the proposed taxonomy, and he replied ”I think you have summarized the relevant information from our paper and at this point I have no further suggestions.”
CAMPAGNA, L., K. GEALE, P. HANDFORD, D. A. LIJTMAER, P. L. TUBARO, AND S. C. LOUGHEED. 2011. A molecular phylogeny of the Sierra-Finches (Phrygilus, Passeriformes): extreme polyphyly in a group of Andean specialists ). Molecular Phylogenetics and Evolution 61: 521–533.
KLICKA, J., K. BURNS, AND G. M. SPELLMAN. 2007. Defining a monophyletic Cardinalini: A molecular perspective. Molecular Phylogenetics and Evolution 45: 1014-1032.
Van Remsen, October 2011
Comments from Stiles: “YES. I did pretty much the same exercise upon receiving the Phrygilus paper and came up with the identical changes to the taxonomy.”
Comments from Pacheco: “YES. A proposta representa um real avanço (todos as espécies do gênero foram amostrados) e se coaduna com a visão prévia e amplamente reconhecida pelos pesquisadores em campo.”
Comments from Robbins: “NO, only because I believe that we should include plebejus and unicolor in Haplospiza. This is consistent with placing dorsalis and erythronotus in Idiopsar.”
Additional comment from Remsen: “Mark has a good point (above). If the proposal passes, I’ll write a separate one to merge those two into Haplospiza, and if it doesn’t pass, I’ll rewrite this one to add that in.”
Comments from Stiles: “An additional point, with respect to Mark’s comment: I didn’t recommend merging these two with Haplospiza if only because that would require renaming the original H. unicolor (the species epithet unicolor for the “Phrygilus” has priority, and there seems to be no other name available for the “Haplospiza” unicolor). In any case, the genus name would still be Geospizopsis, which has priority over Haplospiza, and the type species would now be the “Phrygilus”, not the “Haplospiza” unicolor! If this seems not to be too much bother for someone, I agree with Mark’s comment. Anyone have a good name in mind?”
Comments from Stotz: “YES. Although sympathetic to Mark’s suggestion that we should place plebejus and unicolor in Haplospiza, I think the strong ecological and biogeographic differences argue for creating a separate genus for the ex-Phrygilus. Haplospiza are humid montane forest bamboo specialists, whereas plebejus and unicolor are dry open-habitat species in the Altiplano and west side of the Andes. In contrast, Idiopsar and the two ex-Phrygilus share habitat and biogeography.”
Comments from Pérez: “YES. Proposed classification is consistent with genetic data, morphology, and distribution. I also thought about merging plebejus and unicolor into Haplospiza, but Doug’s comments argue against it.”
Comments from Jaramillo: “NO – But a no on the details, not the major decision to re-classify members of Phrygilus. I do think we need to revise and separate several of these lineages in Phrygilus. Rhopospina is clear -- that is a fine resolution to that clade, although I would not be surprised if Porphyrospiza caerulescens is in that group, and I have no idea if that is an older name than Rhopospina and we need not worry about that now. Similarly, I think Geospizopsis is the way to solve the issue with unicolor and plebejus, and that was a suggestion I wrote in the HBW account dealing with these species. I do not think they should be lumped in Haplospiza because the two groups differ in their ecology, as well as in song type. Haplospiza have these high-pitched buzzes, unicolor whereas plebejus trill or have sweet melodious songs, not high-pitched buzzes. Also these two subgroups are from what I could gather not all that closely related (although clearly a clade); I am more comfortable having them in two different genera. Acanthidops may fit in here as well, and I don’t know how that would change the pattern of relationships between Haplospiza and Geospizopsis. Quite a similar issue is that of Idiopsar. To me Idiopsar is a very, very strange creature. It is perhaps just an unusual member of a larger and more uniform group (like having the Fan-tailed Warbler in Basileuterus), but it may be different enough to warrant its own genus. Idiopsar is a very large and bulky finch with rather strange proportions in the field, including big legs and feet; it also has a unique habitat of liking boulder fields. Surely its morphology is tied to its ecology, but it is unusual and unique enough for any highland finch that I would rather keep it in a monotypic genus. Furthermore, isn’t it actually somewhat distant in the genetic data from the Phrygilus pair? There is a second part to my concern over lumping within Idiopsar, and that is that other species may be in this group and what their inclusion might do to shift any of the results here, in terms of relationship or nomenclature. My main concern is Diuca speculifera. As far as I can recall this species has not been sampled in any of these recent molecular papers on these finch-like tanagers. Having had a substantial amount of time with both species of Diuca in the field, I am thoroughly convinced that they share a plumage pattern but are entirely unrelated; vocally they are very different, with speculifera being rather silent and mainly uttering simple short call like whistles, whereas D. diuca is a loud songster. Diuca diuca has an odd quail-like walking style unlike speculifera. I am making the educated guess that speculifera will be in the clade now being considered under Idiopsar. I have not researched if there is any generic name associated with speculifera that could be used if it is confirmed that it is in the clade with erythronotus and dorsalis. I do think it will be part of this group though; they are all species not only of the highlands but also the extreme highlands, and they are vocally simple, giving weak little short or slurred whistles. Songs are unknown yet for these birds, they may sing very rarely – unlike all others previously in Phrygilus.
“Perhaps unrelated is where Piezorhina and Xenospingus fit in – this is anybody’s guess; they could be entirely unrelated to this group (perhaps closer to Incaspiza?) or embedded in it; I gather that Acanthidops is in with Haplospiza based on the available data?
“To sum up, yes separate Phrygilus. But keep Idiopsar separate, figure out what to do with erythronotus and dorsalis (is there a previously published genus available for them?), similarly resurrect Geospizopsis for unicolor and plebejus, and do not lump in Haplospiza.”
Comments from Zimmer: “YES on changing the classification of Phrygilus, which is clearly polyphyletic. As with Alvaro’s comments, the devil is in the details. The proposed Rhoposina makes perfect sense, as does the proposed 4-species treatment for Phrygilus and the proposed Melanodera. I believe strongly that Geospizopsis is the way to go for unicolor and plebejus, and DO NOT fold those into Haplospiza. As Doug and Alvaro have already pointed out, Haplospiza has very different vocalizations and both species are nomadic bamboo specialists within wet forest – completely different ecologically from unicolor/plebejus. I don’t know Idiopsar in life, but I’ve heard enough about it to make me somewhat queasy about folding dorsalis and erythronotus into that genus. Maintaining Idiopsar as a monotypic genus may be the way to go. Finally, Alvaro’s comments on Diuca are compelling, although my experience with these birds is pretty limited. So, perhaps we need to vote piecemeal on this proposal? I would say definitely “YES” on changing the classification as proposed for Rhoposina, Phrygilus, Melanodera and Geospizopsis, and for restricting Haplospiza to rustica and unicolor. Maintain Diuca pending more published analysis, which, I’m guessing will bear out Alvaro’s thinking. Find a new name for dorsalis and erythronotus, and maintain Idiopsar as monotypic.”
Additional comments from Remsen: “Comments from Alvaro and Kevin above, plus some counsel from Kevin Burns, have convinced me to change my vote and recommendation to NO.”
Comments from Nores: “NO. Although I fully agree with Van that Phrygilus clearly contains four separate lineages I am not in accord in folding dorsalis and erythronotus into Idiopsar. As neither of two species is the type of a previously described genus would have to describe a new genus for them. Moreover, I do not think that unicolor and plebejus should be lumped in Haplospiza as suggested by some members. The strong ecological and biogeographic differences argue for resurrecting Geospizopsis for them.”