Proposal (178) to South American Classification Committee
Abandon the Hinkelmann-Schuchmann classification of the hermit hummingbirds (Phaethorninae), and specifically their classification of the Phaethornis superciliosus-malaris-longirostris species group
This proposal would endorse the classification of these hummingbirds used in our current baseline list (and for the most part in HBW), which represents a departure from most previous comprehensive classifications.
Recognition of the hermit hummingbirds as a separate subfamily Phaethorninae goes back nearly 150 years. At the generic level, the classification of the hermits has remained fairly stable over the last century, save that biochemical and genetic evidence (Gill & Gerwin 1989, Bleiweiss et al.1994, 1997) dictated the transfer of Doryfera and Androdon to the Trochilinae. The classification of the "true" hermits in five genera (Eutoxeres, Ramphodon, Glaucis, Threnetes, Phaethornis) has changed relatively little since Cory (1918) save that Hinkelmann & Schuchmann (1997) split Anopetia from Phaethornis. In the most detailed and comprehensive study to date, Hinkelmann & Schuchmann (1997) presented a classification of the Phaethorninae, which was followed for the most part by Schuchmann (1999). This study, based on analysis of 96 morphological and plumage characters, involved nearly 6000 study skins and all taxa currently recognized in the subfamily. A somewhat unusual variant of cladistic methodology was used to polarize characters: the reconstruction of a hypothetical "common ancestor" of the Phaethorninae, deemed necessary because so few character states were clearly shared with the Trochilinae. The supposed features of this ancestor, plus the characters of Ramphodon naevius (the extant form judged to show the greatest number of plesiomorphic character states) were used to polarize characters in the rest of the subfamily. Although some circularity might be suspected (it appears that the "ancestor" was constructed at least in part with Ramphodon in mind), the arrangement and composition of genera produced are generally consistent with those found in recent genetic studies (e.g., Bleiweiss 200?, McGuire et al. 200?). Among the results of these studies are the transfer of "Ramphodon" dohrnii to Glaucis and the recognition of the close affinities of Ramphodon and Glaucis; species limits and nomenclature in Threnetes have shifted somewhat as well (see SACC proposal no. 77).
Species limits in the large genus Phaethornis have been especially fluid, and nowhere more than in the "long-tailed hermit" complex: the superciliosus-malaris-longirostris species group. Different authors have recognized two to six species over the last century, with two or three species being most frequent. Cory (1918) recognized the three aforementioned forms as separate species, but at the subspecies level his taxonomy differed from that of others in considering some cis-Andean taxa as races of the trans-Andean longirostris, and in different assignment of some races to malaris vs. superciliosus. Peters (1945) included nearly all members of the complex in P. superciliosus (except for nominate malaris, sympatric with nominate superciliosus in NE Brazil and the Guianas), an arrangement essentially followed by most authors since (e. g., Meyer de Schauensee 1966, Sibley & Monroe 1990). Zimmer (1950a) disagreed, and on morphological grounds placed all members of the complex in malaris except for superciliosus (and muelleri of NE South America), an arrangement not generally accepted, perhaps in part due to the isolated distribution of malaris itself. Hinkelmann (1996) presented a much more detailed study, incorporated into the arrangement of Hinkelmann & Schuchmann (1997) and Schuchmann (1999). Based mainly upon plumage morphology and coloration, they divided the complex into three species, the main difference from the classification of Zimmer being their recognition of the trans-Andean taxa as a separate species, longirostris, which in effect represents a partial return to the classification of Cory, but based on a much more thorough analysis.
Various other points of the Hinkelmann-Schuchmann classification of Phaethornis represent departures from previous classifications, but the reasons are generally set forth explicitly in Hinkelmann & Schuchmann (1997). In general, I feel that this is the most thorough attempt at classifying the hermits based entirely upon morphology and plumage (which, after all, is what the older classifications were based on as well) in this difficult group, in which divergence in these characters is much more limited than in the Trochilinae. As intimated by others, it is entirely possible that genetic evidence will eventually dictate changes in this classification, but I know of no genetic study currently in progress with a comparably thorough taxon sampling. Therefore, until such a study is published, I recommend that we follow the Hinkelmann-Schuchmann classification of the hermits in general (except for species limits in Threnetes as mentioned above), and of the genus Phaethornis in particular: that is, a NO vote on this proposal.
BLEIWEISS, R., S. L. HENDRICKSON, M. E. BERRES, Y. O. ONIKI, AND E. O. WILLIS. 2003. Affinities of the Saw-billed Hermit (Ramphodon naevius) determined by cytochrome-b sequence data. Wilson Bull. 115: 1-10.
Bleiweiss, R., J. A. W. Kirsch & J. C. Matheus. 1994. Auk 111:8-19.
Bleiweiss, R., J. A. W. Kirsch & J. C. Matheus. 1997. Mol. Biol. Evol. 14:325-343.
Cory, C. B. (1918), Catalogue of birds of the Americas, part 2 vol. 1.
Gill, F. B. & J. A. Gerwin. 1989. Proc. ANSP 141:409-421.
Hinkelmann, C. 1996. Ornitologia Neotropical 7:119-148.
Hinkelmann, C. & K.-L. Schuchmann. 1997. Stud. Neotrop. Fauna Envir. 32:142-163.
McGuire, Altshuler & Dudley 2004? Hummingbird phylogeny paper -- in press.
Meyer de Schauensee, R. 1966. The species of birds of South America.
Peters, J. L. 1945. Checklist of birds of the world, vol. 5.
Schuchmann, K.-L. 1999. HBW, vol. 5.
Sibley, C.G. & B. L. Monroe 1990. Distribution and taxonomy of birds of the world.
Zimmer, J. T. 1950a. Studies on Peruvian birds, no. 55. AMNH Novitates no. 1449.
Zimmer, J. T. 1950b. Studies on Peruvian birds, no. 56. Ibid, no. 1450.
Gary Stiles, May 2005
Comments from Remsen: "NO. The alternative is to revert to Meyer de Schauensee (1966, 1970). Clearly, the Hinkelmann-Schuchmann classification is supported by explicit published rationale and should be the starting point for our classification."
Comments from Zimmer: "NO, for reasons summed up by Gary."
Comments from Robbins: "NO, based on the rationale provided by Gary."
Comments from Stotz: "YES. I can't argue that Hinklemann's analysis is the most detailed for this group. Yet I have to vote for a return to Mayer de Schauensee arrangement for the superciliosus complex. I am just not convinced by Hinklemann’s arguments. I am least concerned about whether longirostris should be recognized as a distinct species. I could live with either treatment. I would note however, that Hinklemann’s sole character supporting longirostris that they share the characteristic of having 3 light ochraceous bands and 2 blackish bands on the outermost feathers of the upper tail coverts versus 2 and 2 in the cis-Andean forms. Whether that is a valid specific-level character is up to you to decide. Seems pretty weak to me, but I voted for the split from the other AOU committee and I am not opposed here.
"What I can't handle is Hinklemann’s return to Zimmer's arrangement, where all of the Amazonian taxa go with the strictly Guianan malaris, while splitting superciliaris and muelleri from north and south of the eastern Amazon off. We have at least two species east of the Andes. the question is do most of the taxa go with malaris or superciliosus? The characteristics that Hinkelmann uses to join malaris and the rest of Amazonian forms are two: 1) the color of the rectrical margins (pale ochraceous in superciliosus, and light brownish in malaris); 2) superciliosus has distinct mid-throated streaking in both sexes, while it is pronounced in females of malaris and "distinctly less pronounced or lacking" in males. I personally think he is wrong about the first character. We have a pretty good series (20-ish) of superciliosus, and to my eye they look much like moorei from Colombia, dull and not quite white. Pale brown seems reasonable. The one that stands out is ochraceiventris, which has strongly ochraceous edges. Those of you with access to northern Peruvian specimens, take a look. On the throat character, I will agree that at least some specimens of males of the Amazonian taxa Hinkelmann places in malaris lack the midline pale streak on the throat, and that in our superciliosus series, males and females look the same. However, the throat character is variable in males, and in all of the taxa that we have a reasonable number of males, we have at least one that looks just like the females with a strong midline streak.
"We have only one nominate malaris. It is a male and has a dark throat. the thing I would say is that the nature of the throat marking looks very different to me from that shown by the Amazonian forms. It is very dark and extensive and does not have pale edges to the dark throat feathers. In the Amazonian forms the dark patch is less extensive and the feathers are narrowly edged with pale, even in those that lack the pale midline. I should note that we lack insolitus, the form that sounds like it most resembles malaris in plumage.
"So, what sets malaris apart from the Amazonian forms. The most obvious is its size. It is substantially larger than any other form in the complex. In terms of plumage (based on 1 specimen) it is generally greener and darker than any of the other forms which show varying degrees of brownish. I will admit that my argument for superciliosus rather than malaris is weak, not really much stronger than Hinklemann’s. But biogeographically it seems like superciliosus fits better than malaris. It abuts other Amazonian forms along the Negro, Amazon, and Tapajos, while malaris is isolated from everybody else in the Guianas. With superciliosus as part of that species, the distribution looks like any number of widespread Amazonian species. With malaris you have this strange disjunction.
"I recognize that this rant will not result in changing our treatment of malaris and superciliosus, but I think what Eisenmann was quoted as saying in Meyer de Schauensee (1966) is still true -- "the arrangement of Peters and earlier authors [malaris monotypic, others joined with superciliosus] is adopted here "because it seems more consistent with the zoogeographic situation (and at least equally consistent with the morphological evidence)." While I don't really expect us to change this, I am convinced that genetic work will someday show that malaris is the outlier."
Additional comments from Stiles: "NO. While I agree that the evidence for Hinlkelmann's rearrangement of the superciliosus-malaris group is thin, it is explicit and I consider it better than the "gut feelings" of other authors - how often have we been led astray by superficial similarities in pattern or the vagaries of distribution, etc.? (Geositta seems to be a case in point?) The last word has yet to be said on this group, but for now published evidence seems to favor the Hinkelmann arrangement."
Comments from Silva: "NO, based on Gary's comments."
Comments from Jaramillo: "NO - Doug Stotz has a good point, but it is not enough to keep me from accepting the arrangement as suggested by Hinkelmann and summarized by Gary."
Comments from Nores: "NO. Encuentro razonable el criterio de Gary para seguir a Hinkelmann y Schuchmann hasta tanto haya algún estudio genético o de otro tipo que aclare las cosas."