Proposal (434) to South American Classification Committee

Place all members of Schizoeaca and Oreophylax in Asthenes

Effect on SACC: Species in the genera Schizoeaca and Oreophylax would be placed within Asthenes. The linear classification of these species would change.

Background & New information (much of this was copied from Derryberry et al. 2010 --RTB):

A recently published phylogeny of the Furnariidae provided the first genetic evidence of lack of monophyly in Asthenes (Irestedt et al. 2006). This phylogeny included two species of Asthenes, one of which—A. cactorum (Cactus Canastero) —was sister to Pseudoseisura, whereas the other—A. urubambensis (Line-fronted Canastero) —formed a clade with Oreophylax and Schizoeaca. Gonzalez and Wink’s (2008) phylogeny of the Synallaxinae included three species of Asthenes. They found that A. cactorum and A. humicola (Dusky-tailed Canastero) formed a clade that was sister to Pseudoseisura, whereas A. urubambensis formed a clade with Schizoeaca and Oreophylax. In a broader genus-level study of the infraorder Furnariides, Moyle et al. (2009) found Asthenes to be paraphyletic with respect to Schizoeaca in that S. helleri (Puna Thistletail) was nested within a group that contained A. humilis (Cabanis) (Streak-throated Canastero), A. urubambensis, and A. baeri (Berlepsch) (Short-billed Canastero). These findings suggested the need for a new phylogenetic classification for taxa currently included in Asthenes and related genera.

As part of a project to reconstruct the phylogenetic relationships of all species in the Furnariidae from DNA sequences (mitochondrial and nuclear), extensive taxon-sampling allowed Derryberry et al. (2010) to determine conclusively that the genus Asthenes consists of two groups that are not sister taxa. One group consists of four species of Asthenes (A. cactorum, A. steinbachi), A. patagonica (Patagonian Canastero), and A. humicola), whereas the second group consists of all remaining species of Asthenes as well as all species sampled from the genus Schizoeaca and Oreophylax moreirae (Itatiaia Spinetail), the sole member of its genus. The type species of Asthenes (A. sordida, currently considered a subspecies of A. pyrrholeuca [Sharp-billed Canastero]) belongs to the large second group. Because no generic name is available for the clade consisting of A. cactorum, A. steinbachi, A. patagonica, and A. humicola (Cory 1919, Cory & Hellmayr 1925, see classification below) Derryberry et al. (2010) described the new genus Pseudasthenes for these four species – see Proposal 433.

After transferring the four species to Pseudasthenes, Asthenes remains paraphyletic because Oreophylax moreirae and all species of Schizoeaca are nested within it. Thus, it is proposed that species currently in Oreophylax and Schizoeaca be moved to Asthenes. More details on rationale can be found in Derryberry et al. (2010).

Derryberry (2010) recommended a provisional classification of Asthenes based on their phylogeny (see below). The relationships of the three species not included in their study were inferred tentatively from phenotypic characters. Asthenes berlepschi is almost certainly closely related to A. dorbignyi and may be a subspecies of A. dorbignyi (Cory & Hellmayr 1925: Bond & Meyer de Schauensee 1942; Fjeldså & Krabbe 1990). The second missing species, A. heterura, has been considered closely related to (Cory & Hellmayr 1925), sister species to (Vaurie 1980), or conspecific with A. pudibunda. Pearman (2001), however, noted that A. heterura is sufficiently similar to A. pyrrholeuca in plumage that they can easily be confused in the field and even in the hand. Because A. pudibunda and A. pyrrholeuca are not sister species the specific placement of A. heterura is better regarded as uncertain, although it probably belongs to the long-tailed Asthenes/Schizoeaca/Oreophylax clade. Schizoeaca coryi is similar to other Schizoeaca thistletails in plumage, tail structure, and habitat (Remsen 1981, 2003), and it presumably forms part of the same clade.

Genus Asthenes Reichenbach 1853

Asthenes coryi (Berlepsch)

Asthenes perijana (Phelps)

Asthenes fuliginosa (Lafresnaye)

Asthenes griseomurina (Sclater)

Asthenes pudibunda (Sclater)

Asthenes heterura (Berlepsch)

Asthenes vilcabambae (Vaurie)

Asthenes palpebralis (Cabanis), type of Schizoeaca Cabanis

Asthenes ottonis (Berlepsch) type of Pseudosiptornis Cory

Asthenes helleri (Chapman)

Asthenes harterti (Berlepsch)

Asthenes moreirae (Ribeiro), type of Oreophylax Hellmayr

Asthenes pyrrholeuca (Vieillot), type of Asthenes

Asthenes modesta (Eyton)

Asthenes humilis (Cabanis)

Asthenes wyatti (Sclater & Salvin)

Asthenes sclateri (Cabanis)

Asthenes anthoides (King), type of Eusiptornoides Cory

Asthenes hudsoni (Sclater)

Asthenes urubambensis (Chapman)

Asthenes flammulata (Jardine), type of Siptornoides Cory

Asthenes virgata (Sclater)

Asthenes maculicauda (Berlepsch)

Asthenes luizae Vielliard

Asthenes dorbignyi (Reichenbach)

Asthenes berlepschi (Hellmayr)

Asthenes baeri (Berlepsch)

Figure 1 from Derryberry et al. (2010) --- A simplified majority-rule Bayesian consensus tree of the Furnariidae that highlights the lack of a sister relationship between Pseudasthenes and Asthenes as well as the paraphyly of Asthenes, Schizoeaca, and Oreophylax. Asterisks represent nodes with a posterior probability of 1.0.

References

Bond, J., & R. Meyer de Schauensee. (1943) The birds of Bolivia, Part II. Proceedings of the Academy of Natural Sciences, Philadelphia, 95, 167–221.

Cory, C. B. (1919) A review of Reichenbach's genera Siptornis and Cranioleuca, with descriptions of new allied genera and subgenus. Proceedings of the Biological Society of Washington, 32, 149–160.

Cory, C. B. & Hellmayr, C. E. (1925) Catalogue of birds of the Americas and the adjacent islands, Part IV, Furnariidae-Dendrocolaptidae. Field Museum of Natural History, Zoological Series, 13(3), 1–390.

Derryberry, E., S. Claramunt, K. E. O’Quin, A. Aleixo, R. T. Chesser, J. V. Remsen, Jr., and R. T. Brumfield. 2010. Pseudasthenes, a new genus of ovenbird (Aves: Passeriformes: Furnariidae). Zootaxa 2416:61-68.

Fjeldså, J. & Krabbe, N. (1990) Birds of the High Andes. Zoological Museum, University of Copenhagen, and Apollo Books, Svendborg.

Gonzalez, J. & Wink, M. (2008) Phylogenetic position of the monotypic Des Murs' Wiretail (Sylviorthorhynchus desmursii, Aves : Furnariidae) based on mitochondrial and nuclear DNA. Journal of Ornithology, 149, 393–398.

Irestedt, M., Fjeldså, J. & Ericson, P. G. P. (2006) Evolution of the ovenbird-woodcreeper assemblage (Aves : Furnariidae) - major shifts in nest architecture and adaptive radiation. Journal of Avian Biology, 37, 260–272.

Moyle, R. G., Chesser, R. T., Brumfield, R. T., Tello, J. G., Marchese, D. J. & Cracraft, J. (2009) Phylogeny and phylogenetic classification of the antbirds, ovenbirds, woodcreepers, and allies (Aves: Passeriformes: Furnariides). Cladistics, 25, 386–405.

Pearman, M. (1990) Behaviour and vocalizations of an undescribed canastero Asthenes sp. from Brazil. Bulletin of the British Ornithologists’ Club, 110, 145–153.

Reichenbach, H. G. L. (1853) Handbuch der speciellen Ornithologie (Icon. Syn. Av. No. 10) 146, 168.

Remsen, J. V., Jr. (1981) A new subspecies of Schizoeaca harterti with notes on taxonomy and natural history of Schizoeaca (Aves: Furnariidae). Proceedings of the Biological Society of Washington, 94, 1068–1075.

Remsen, J. V., Jr. (2003) Family Furnariidae (ovenbirds). In: del Hoyo, J., Elliot, A. & Christie, D. A. (Eds), Handbook of the Birds of the World. Vol. 8. Lynx Edicions, Barcelona, pp. 162–357.

Vaurie, C. (1980) Taxonomy and geographical distribution of the Furnariidae (Aves, Passeriformes). Bulletin of the American Museum of Natural History, 166, 1–357.

Robb T. Brumfield, April 2010

Comments from Nores: “YES, pero con prácticamente las mismas consideraciones que para la propuesta anterior, en la cual puse ‘Todos los fundamentos morfológicos y biogeográficos que tenía con este grupo quedan desactualizados por los análisis moleculares. Ningún otro cambio me ha producido una desorientación tan grande como este caso, no sólo por el nuevo género sino también por las inesperadas relaciones entre las especies. Así, por ejemplo, Asthenes baeri y a A. dorbignyi, que son tan parecidos a A. steinbachi, resultan muy diferentes, y en cambio se agrupan con Schizoeaca y Oreophylax, un grupo que parecía tan distinto morfológica y biogeográficamente, restringido a los páramos. Como si esto fuera poco, Pseudasthenes está relacionado con Pseudoseisura, algo que parecía imposible de ser. Es todo para mi tan inesperado, que no descarto totalmente la posibilidad que haya habido algún error en los análisis moleculares’.”

Comments from Stotz: “NO. I can’t quite bring myself to vote for this, given the bizarre topology within the Schizoeaca + piece of the clade. Schizoeaca is usually treated as a superspecies, and it has occasionally been treated as a single species. It seems like a clear specialized group of paramo Asthenes. Oreophylax seems clearly the Atlantic forest version of this group. I have no problem with some of the current Asthenes being related to Schizoeaca. So far so good. But to have several ex-Asthenes scattered through the genus doesn’t make a lot of sense. The geography of the Schizoeaca also doesn’t seem to fit. I’m going to need something more than this tree to convince me that Schizoeaca is not a monophyletic unit (don’t know about coryi, not included here).

“Before I got concerned about the Asthenes buried within Schizoeaca, I was leaning toward recognizing two clades here. The big Asthenes clade (which looks good), plus the Schizoeaca + clade. The problem is that the type of Asthenes is pyrrholeuca, so the Schizoeaca clade would become Asthenes, and the big clade of just “Asthenes” would be Siptornoides or Eusiptornoides. We’d still need to know what to do with luizae, baeri and dorbignyi (like Manuel, I can’t quite get my brain around dorbignyi and baeri as sisters.)”

Comments from Zimmer: “NO. I have to agree with Doug’s take on this. I have no problem with Schizoeaca, Oreophylax and some Asthenes being considered as comprising a separate “thistletail” group distinct from other Asthenes, but it’s hard to buy into the proposed sequence, which has members of these two groups scattered all over the place. It doesn’t make much biogeographic sense to me, and it seems counter-intuitive biologically, when you consider that, as Doug points out, the various Schizoeaca have variously been considered as either a single species or as forming a superspecies.”

Additional comments from Remsen: “I understand completely where Doug and Kevin are coming from. When I first saw that tree, I thought “no way” (as did the others in our group, whose first reaction was “lab screw-up”). After all, it took a publication from me back in ’81 to break up Vaurie’s broadly defined Schizoeaca fuliginosa back into its Meyer de Schauenseenean components. But I’m now “at peace” with this startling result:

“First, the genetic data are solid. Multiple individuals and multiple genes have been analyzed, and the nodes are strongly supported. We are not going to get any better or different tree anytime soon. Like it or not, “Schizoeaca” is hopelessly polyphyletic with respect to Asthenes and Oreophylax – maintaining it as is untenable under modern concepts of classification.

“Second, the oddity of the phenotype is really not as odd as it might first seem. The only thing that really unites Schizoeaca is the disintegrated tail vane. Otherwise, they vary greatly in terms of face pattern, body size, and color. The three Asthenes nested within that part of the tree are also fairly close to certain Schizoeaca in terms of having eyebrows, throat patches, and varying degrees of obsolete striping on chest. They are generally paler, but remember that the classic thistletails occur in more humid habitat (a Gloger’s Rule sort of thing). If you put skins of them all together with Schizoeaca, as I have, the only thing that really stands out is the tail; dorsally they are essentially identical. As for biogeography, A. ottonis and A. pudibunda cluster with the southern Schizoeaca types, and so there’s some biogeographic sense there; further, A. pyrrholeuca, which is likely partly migratory, is sister to Oreophylax, and these two cluster with the southernmost thistletail, S. harterti.

“Back to the tail. What the phylogeny says to me is that thistletail tail morphology is a plastic character, clearly associated with paramo habitat. Perhaps the loss of barbs is not only very easy, developmentally, but an “adaptation” to dealing with constantly wet vegetation.

“Whether to include all of the remaining post-Pseudasthenes species in a broad Asthenes is a separate question, one that could/should be addressed in separate proposals. One could make a case that the hudsoni-to-modestus branch and dorbignyi-baeri branch of the tree each should be a separate genus. With the type species of Asthenes being pyrrholeuca, those other two branches are the ones that would need new names. Within the part of the tree under discussion, to maintain Schizoeaca, one would have to move harterti to Asthenes, and you would still have to include ottonis and pudibunda in Schizoeaca.”

Comments from Jaramillo: “YES. I did not know exactly how to vote on this one, but sticking to the main point of the proposal, whether to include Schizoeaca and Oreophylax in Asthenes, I am fine with that. The issue for me is that there are at least three, maybe four, clades within this larger Asthenes. I see the classic streaked canasteros, the Schizoeaca-Oreophylax-Asthenes “long-tailed” group, and then Short-billed (baeri), Creamy-breasted (dorbignyi) and Cipo (luizae). Short-billed and Creamy-breasted makes a great deal of sense to me, in many ways Short-billed is a washed out looking version of Creamy-breasted. Both of them have aspects in behavior and some aspects of vocalization, and even bill coloration that evokes a thornbird (Phacellodomus). Vocally they are similar in having a song type that is a long and drawn out trill, not speeding up or moving much in pitch … at least this applies to southern dorbignyi, as well as arequipae within the Creamy-breasted group. In some ways having this long trill song, as well as some (all?) in the dorbignyi group, which has a stuttering song type, parallels some of the vocal aspect of Pseudasthenes! I don’t know about luizae, but listening to the few vocalizations available there are suggestions of the dorbignyi – baeri group and I wonder if the relationship is really those three in a group? Looking at the tree, this is not inconceivable. I would prefer to separate Asthenes into at least three genera and can see that vocal, and morphological aspects that support such a move. But keeping them all in Asthenes does not seem like a good course of action, because it makes the genus too heterogeneous.”

Comments from Pacheco: “YES. Na minha opinião, eu vejo dois arranjos possíveis. Em ambos, contudo, Schizoeaca e Oreophylax são submergidos em Asthenes, conquanto o typus de Asthenes é A. pyrrholeuca. A segunda composição possível – diante da árvore em Derryberry et al. (2010) – seria reconhecer pelo menos um segundo gênero Siptornoides Cory, 1919 para os Asthenes entre A. hudsoni e A. dorbignyi da sequência obtida na filogenia.”

Comments from Stiles: “YES – something of a mess … the degree of paraphyly is rather disconcerting, especially given the high posterior probabilities.. I se little hope for maintaining Schizoeaca, and it is a bit frustrating that recognizing two (or four, if luizae and dorbignyi-baeri are accorded generic status) genera in this clade would leave the name Asthenes for a group composed mainly of ex-Schizoeaca while the bulk of Asthenes species would require a new name! Perhaps the best and least disruptive course is indeed to put the whole bunch into Asthenes – there really is a common theme running through the group in terms of plumage pattern, and since the most obvious difference (streaky vs. non-streaky plumage) doesn’t fall out neatly in separate clades, putting all the eggs in one basket has its logic.”