Proposal (955) to South American Classification Committee

Treat Pachyramphus salvini as a separate species from P. albogriseus

This is a revamp of proposal 906 after publication of new information.

Effect on SACC: This would raise to species rank a taxon mainly occurring in western Colombia, west Ecuador, northwestern Peru and the Marañón drainage, but also found, perhaps only seasonally, on the Amazonian slope in N and C Peru, Ecuador, and possibly S Colombia. It is currently treated as a subspecies of Pachyramphus albogriseus, which occurs from Venezuela and E Colombia south through E Ecuador to S Peru, and disjunctly in Costa Rica and W Panama.

BACKGROUND: Proposal 906 was met by a committee that agreed in that there was evidence that Pachyramphus albogriseus is polyphyletic, but most committee members believed that a closer study was needed in order to clarify nomenclature and distribution of the two species before accepting the split, just as a study of vocalizations was asked for. It was mainly the occurrence of the western species on the east slope of the Andes in S Colombia (genetics) and C Peru (vocalizations) that was cause of concern, but also the use of the name salvini was questioned by Areta, who put forward the hypothesis that the name salvini should be applied to the clade including the samples of guayaquilensis, given its priority.

NEW INFORMATION: Musher et al. (2023) published a study that used both genetics, morphology, vocalizations and photographs from the Macaulay Library to clarify the distributions and diagnoses of the two species to circumscribe them more precisely. They examined type specimens to ensure a correct nomenclature. They found a fine correlation between vocal, morpho- and genotypes. Surprisingly, they found no less than 13 records of the western form from the east slope, and one record (2 specimens) of the nominate form from the Marañón drainage. The east slope records of the western form were from all months of the year except February and March (the peak breeding season in the west), so it remains possible that the western form is partly migratory and does not breed on the east slope. The two specimens of nominate P. albogriseus from the Marañón drainage, however, strongly suggest that the two species breed sympatrically or parapatrically, perhaps occupying different habitats, the western form tolerating drier and more disturbed habitats, the nominate form perhaps in undisturbed humid forest higher on the slope, this needing further investigation. All 5 specimens from E Ecuador that Zimmer (1936) had examined pertained to the western species, whereas the single specimen that Musher had sequenced from E Ecuador pertained to nominate albogriseus. This led Musher & Cracraft (2018) to apply the name salvini to the eastern form, which left only the name guayaquilensis available for the western form. A close examination of a large series of specimens, including the ten specimens in the type series of salvini, showed that the diagnostic characters of guayaquilensis are not statistically significant, and that guayaquilensis is but a junior synonym of salvini. Thus, the correct name for the western form is P. salvini.

A more detailed account of the approaches:

Genetics: Seven more specimens were sequenced (marked with red stars below). The resulting phylogenetic tree based on concatenated nuclear (UCE) data and was similar to previous trees (Musher & Cracraft 2018, Musher et al. 2019), confirming beyond doubt that Pachyramphus albogriseus sensu lato is polyphyletic.

Vocalizations: Two song types were found, consistent with songs described for E and W Ecuador (Ridgely & Greenfield 2003) and for E and W Peru (Schulenberg et al. 2007); note however that due to nomenclatural confusion, the names to which these vocalizations were ascribed have been mixed (see Proposal 906 and spectrograms below). No specimen with a voice attached was available, but one of the two vocal types was found in Venezuela and C America, the other in W Ecuador and NW Peru, each being areas where only one genotype, one vocal type, and one morphotype had been recorded. Besides song, the two also appear to have different calls. Both vocal types had been recorded in E Ecuador and E Peru, the eastern type mainly in undisturbed forest, the western type mainly in more open habitat (see

spectrograms below).

Morphology: Two morphotypes were found. These were consistent with the two genotypes and vocal types. One, P. albogriseus, is a large species (average 22 g) with a broad upper wingbar, heavy bill, blackish loral spot, uniform alula, a faint pale collar, female with a bright chestnut crown surrounded by a broad black band, male with a mostly black upper tail. The other, P. salvini, is a smaller species (average 17 g) with a slender bill, a relatively narrow upper wingbar, a pale-edged alula, no or faint loral spot, no pale collar, female with a light brown or dull chestnut crown surrounded by a narrow or no black band, and male with much gray on the upper tail. In C America and Venezuela only the large species has been recorded, in W Ecuador and NW Peru only the small species. Both types have been found in E Ecuador and E Peru, in the Marañón drainage, and possibly in SE Colombia. Photographic records are consistent with these morphotypes, adding that the small species also occurs in W Colombia. A comparison of specimens from the Perijá mountains (described as a subspecies coronatus) with specimens from other parts of Venezuela showed that the diagnostic feature of coronatus (the darker crown of the female) does not hold and that coronatus should be treated as a junior synonym of albogriseus. The C American form, however, differs from nominate albogriseus in a number of respects and should be treated as a valid subspecies: P. albogriseus ornatus. To illustrate the differences between these taxa we show some specimen comparisons copied from the supplementary data to the paper.

Distribution: The integration of genetic, morphological and vocal data result in the following distributions (type localities are shown by yellow stars, including for the synonymized taxa guayaquilensis and coronatus):

• Pachyramphus albogriseus ornatus: Humid montane forest of Costa Rica and W Panama (600-1200 m).

• P. a. albogriseus: Humid montane forest of Venezuela and N Colombia south through E Ecuador to S Peru (500-2200 m). Apparently also locally in the Río Marañón drainage ("Lomo Santo" = Loma Santa, Jaen district, 1500 m).

• P. salvini: Humid montane and dry deciduous forest in W Colombia, W Ecuador, NW Peru and the Río Marañón drainage. Also, perhaps seasonally only, on the Amazonian slope of S Colombia(?), E Ecuador and E Peru (0-2450 m).

English names: Both species have large ranges, so none of them should bear the name Black-and-white Becard. The authors’ preferred name for P. salvini was Cryptic Becard, but there is such a widespread reluctance against the use of the name Cryptic, that reviewers would not let us publish it. We don’t really think it was their place to interfere with what vernacular name we preferred, and in fact the name is in line with Cryptic Honeyeater (Microptilotis imitatrix) and Cryptic Flycatcher (Ficedula crypta), but there it is. We also had to change the word ”cryptic” with ”look-alike” in the title of the paper. So in the end, we applied the name ”Slender-billed Becard” to P. salvini, the slenderer bill being fairly obvious from some angles. We suggested the name ”Broad-banded Becard” for P. albogriseus, a name we are fine with, as it highlights the most easily seen character in the field. ”Black-tailed Becard” was suggested too, but was given up because some individuals, especially of the C American subspecies do have gray on the upper tail.

We can divide the proposal:

A) Treat Pachyramphus salvini as a separate species. We strongly recommend a yes vote.

B) Use the English name Cryptic Becard for P. salvini. This is our preferred choice.

C) Use the English name Slender-billed Becard for P. salvini. Probably the best alternative to Cryptic Becard.

D) Use the English name Broad-banded Becard for P. albogriseus. We recommend a yes vote.

References

Musher, L. J., and Cracraft, J. (2018). Phylogenomics and species delimitation of a complex radiation of Neotropical suboscine birds (Pachyramphus). Molecular Phylogenetics and Evolution 118:204–221.

Musher, L.J., Ferreira, M., Auerbach, A.L. and Cracraft, J. (2019). Why is Amazonia a ’source’ of biodiversity? Climate-mediated dispersal and synchronous speciation across the Andes in an avian group (Tityrinae). Proceedings of the Royal Society B: Biological Sciences 286:20182343.

Musher, L.J., Krabbe, N.K. and Areta, J.I. (2023) Underestimated Neotropical diversity: Integrative taxonomy reveals two unrelated look-alike species in a suboscine bird (Pachyramphus albogriseus). Ornithology 140: 1–17. Online version published 2022: https://doi.org/10.1093/ornithology/ukac047.

Ridgely, R.S., and Greenfield, P.J. (2001). Birds of Ecuador: Status, distribution and taxonomy. Cornell University Press, Ithaca, NY, USA.

Schulenberg, T. S., D. F. Stotz, D. F. Lane, J. P. O’Neill, and T. A. Parker III (2007). Birds of Peru. Princeton University Press, Princeton, NJ, USA.

Zimmer, J.T. (1936) Studies of Peruvian birds XXIVL. Notes on Pachyramphus, Platypsaris, Tityra and Pyroderus. American Museum Novitates 894.

Niels K. Krabbe, J. Ignacio Areta, and Lukas J. Musher, January 2023

Comments from Lane: “A) YES. The authors make a compelling case for the species status of Pachyramphus salvini, and have done an elegant job of untangling the taxonomic conundrum that has plagued us for so long!

“As for B-D, I am not enamored of their selected names of the daughter species, and vote NO for all. Honestly, those names seem to highlight features that are not immediately obvious (which band is broad?), and do not allow the unaware user to know that the two species had been considered one for so long, nor that they are extremely easily confused! We can argue that the name “Black-and-white” isn’t exactly accurate, as the base color for most of the (male’s) plumage is actually gray, not strictly black and white, but if we can agree that we have lived with it without much chafing, allow me to suggest alternative names. Another way to say “black-and-white” is “pied.” So why not draw attention to the fact that these two species are large and small versions of nearly identical plumage patterns? I would suggest “Greater Pied Becard” for P. albogriseus (sensu stricto) and “Lesser Pied Becard” for P. salvini. That way, it is clear that they are very similar in appearance (like, e.g., the Yellowlegs), but that size is one of the important characters distinguishing them. Furthermore, these names do not necessarily require sister relationship, as the Yellowlegs case illustrates, but do make clear that the two species have been closely tied for most of their existence.”

Comments from Stiles: “YES. Me gusta la nueva propuesta sobre Pachyramphus salvini vs. albogriseus so YES for my vote! My only doubt is the E-name "Broad-banded" for the latter: to which band does the name refer to?

Comments from Remsen: “A. YES. I’ve been following this case from the sidelines since my initial proposal (906), and I echo Dan’s comments – outstanding job of working this one out, and indisputable evidence in my opinion for species rank for salvini.”

Comments from Gary Rosenberg (voting for Areta on B, C, D): “I agree that the authors have shown using multiple methods that salvini should be considered distinct from albogriseus - not only is it genetically distinct, but both the vocal, size, and plumage differences support this treatment. I also don’t really like “Broad-banded” for the name for “Black-and-white” - a bit like “Orange-banded “ Flycatcher - which I assume refers to the “wing-bars” but have always wondered, as they usually are not orange, and no one really calls wing bars “bands"? I guess there are other English names that use “banded” but it is not always referring to the wing-bars - so not so obvious to the observer. I don’t really like the name Cryptic - as I agree with others that it is not hiding in plain sight from a similar species that it co-occurs with - I think there are enough clear differences now that we are aware of them that the name Cryptic doesn’t really apply. Salvini does have a more slender bill - so I am not against that as a common name - although is it really more “slender billed” than other becards? It is when compared to albogriseus - but no one is going to see the two together. I like Dan’s idea of “Pied” Becard - so maybe a compromise - Pied and Slender-billed - as opposed to Greater and Lesser Pied Becards? Therefore:”

“B: NO

“C: YES

“D: NO

Comments from Don Roberson (voting for Claramunt): “Like Dan Lane, I vote note “no” on the English name choices.

“As a tiny bit of background, Bret Whitney showed those of us on a Madagascar tour in 1992 a newly discovered species of “warbler” in eastern Madagascar, which I was able to see, listen to, and photograph. It had unique habitat along ridge lines in lowland rainforests, see: Goodman, S.M., Langrand, O. and Whitney, B.M. (1996). A new genus and species of passerine from the eastern rainforest of Madagascar. Ibis 138(2): 153-159.

“I thought at the time that “Ridgeline Warbler” would be a good English name; at that time it was thought to be in the Sylviidae before that family got split into a dozen or more. It proved to be in Family Bernieridae. In the meantime, its proposed English name was Cryptic Warbler, and eventually became Cryptic Warbler Cryptosylvicola randrianasoloi, and it remains that today. However, it has a distinctive song and was only “cryptic” in the sense that it looked somewhat like a bunch of Phylloscopus warblers at first glance.

“Ever since, that English name proved so disappointing to me that — given that it proved to be in an entirely different family than Phylloscopus, from which it was supposed to be so “cryptic,” -- I’ve thought the English name “Cryptic” should be used with caution.

“The proposed “Slender-billed” and “Broad-banded” are both a bit of a mouthful, and the differences not all that apparent without a lot of background. “Broad-banded” appears to be restrict to the broad v. narrow black band adjacent to the chestnut crown in female, if I understand this correctly. But I don’t understand why it is not proposed as “Slender-billed” versus “Thick-billed,” or “Broad-banded” vs “Slender-banded” for the two taxa, or similar comparative names, which focuses the observer on the same character in each species, rather than two different traits, one of which is only in females? This seems needlessly confusing. With prions there is a Slender-billed and Broad-billed Prion —even if those characters might be hard to determine in the field — but at least it focuses the observer on the bill, not the bill of one and the crown of the other (if a female). So preliminarily a “no” on all these.

“I do like Dan Lane’s proposed Greater Pied and Lesser Pied Becard — which not only gets us to something like “Black-and-white” (the long-standing English name), but a Yellowlegs like comparison.”

“There may be other good potential names, but at this point, I like Greater Pied and Lesser Pied for what is currently up for offer.”

Comments from Donsker (voting for Bonaccorso): “I am reluctant to accept the English names proposed by the authors of this proposal. Rather than trying to select English names that attempt to distinguish these two very similar-appearing species as the authors have valiantly attempted to do, I would suggest that, instead, we consider English names that help to distinguish P. albogriseus (s.s.) and P. salvini rom the three other similar “black-and-white” or “pied” species in the genus Pachyramphus, i.e., White-winged Becard P. polychopterus, Black-capped Becard P   marginatus, and, in Central America, Gray-collared Becard P. major.

“In comparison to those other three ‘black-and-white’ becards, I believe that only P. albogriseus and P. salvini have uniquely gray or slaty-gray mantles. The mantles of the other three are black. I would propose that the name ‘Gray-backed Becard’ be incorporated into the English names of both species. This unique plumage feature is already reflected in the vernacular German name for P. albogriseus (s.l.). which is ‘Graurückenbekarde’.

“My suggestion would be to apply Greater Gray-backed Becard to P. albogriseus (s.s.) and Lesser Gray-backed Becard to P. salvini. As Dan has already pointed out, using the same basic English term for these two species doesn’t necessarily imply a sister relationship any more than it does for the two yellowlegs, the two black-backed gulls or, for that matter, Greater and Lesser Flamingos which aren’t even in the same genus. Retaining a similar English name for both species emphasizes their historically confounding similarity.

“Although I think that “pied” becard is actually more suitable for the three becards with black mantles, if gray-backed” becard is unacceptable, I would support Dan’s suggestion of Greater and Lesser Pied Becard for the two species. ‘Pied Becard’ would not be a unique application since the French name for P. albogriseus (s.l.) is already ‘Bécard pie’.”

Comments from Hilty (voting for Pacheco): “Geez, they have unraveled quite a convoluted puzzle here. And, interesting regarding the songs. This bird is regular in the coastal cordillera of northern Venezuela, and in the Mérida Andes of Venezuela and always seemed quite vocal. However, it seems decidedly scarce (or perhaps not recognized?) in Colombia. At least in my experience, I rarely hear it.

“I don't much care for either name suggested. But, if the name Black-and-white Becard has to go, then Dan Lane's suggested alternatives, Greater and Lesser Pied Becard (eastern and western forms respectively) are better and clearly show originality. As both Dan and Gary pointed out, the name suggested (the name Broad-banded is confusing and unhelpful); and Cryptic really doesn't provide any helpful information.

“I would, however, suggest a slight variation on Dan's names, to make it easier for people to keep the geographical distributions in mind (and this is important with so many new name changes occurring): thus Eastern Pied Becard, and Western Pied Becard. I am aware that there has been some push-back in the past over adding "geographical adjectives" to names, but these are really very helpful for people who English names. The use of "Greater" and "Lesser" in the two names suggests a distinction that I don't think is very obvious at all (at least not in the field, and that is where these English names will be used).”

Comments from Josh Beck (voting for Remsen): “"This is a really cool result, and it shakes up things in a group of similar looking Becards. For better or worse, the name Black-and-white, even if not particularly accurate, and now needing to be retired, was pretty entrenched and well understood; this is not an extremely obscure bird. Looking at possible new English names, while Broad-banded and Slender-billed might be technically correct, I don't feel that they best help a birder / user of English names. They don't provide any way to relate back to the prior name. Based on field experience and having also looked through a good number of photos, I don't think either field mark is particularly easy to interpret in the field. So I follow others in voting NO on B/C/D. I do like Dan's suggestion, and like the idea of "Eastern vs Western" Pied Becard about as well or perhaps a bit better than Greater vs Lesser. In a group of quite similar birds that are all difficult to assign uniquely identifying names to, something that ties back to what they used to be named and that helps distinguish between the two new species does seem helpful and desirable."

Additional comments from Lane: “I appreciate that my comments on this proposal seem to have resonated with others here, and I am inspired to make a few additional ones on the topic of English names. As several have pointed out “pied” isn’t really all that accurate (at least, not more so than “Black-and-white”… but most English names simply fall short of perfect accuracy, as we all know). I considered “pied” to be a shorter word that still retains the basic meaning. David’s suggestion of “Gray-backed” could work, but males of Pachyramphus rufus and some populations of Pachyramphus major also share the gray back, so it isn’t as unique as David’s proposal suggested to these two species within the “black-and-white” group of Pachyramphus. In addition, the more syllables a name has, the less I like it (harder to spit out when trying to get a group on a bird, for example!). So I’d prefer “pied” over “gray-backed” for succinctness if nothing else. As for Eastern and Western… well, I'd urge a review of the map above. P. albogriseus has a representative in Costa Rica and Panama, which is entirely WEST of P. salvini’s range. One could see “Eastern” in Costa Rica and “Western” on the eastern slope of the Andes in Peru! In essence, P. albogriseus is east, west, north, and south of the distribution of P. salvini, so I don’t personally feel using geographic terms really fits well here unless it was something like “Central Pied Becard” and “Peripheral Pied Becard.” Returning to the usefulness of “Greater” and “Lesser” here: yes, it is nearly impossible to judge size of an individual becard in the canopy, but the bill size (in proportion to the head) may prove to be useful in field identification, and P. albogriseus has a proportionally larger bill than P. salvini (again in a surprising parallel to the yellowlegs!), so to me “Greater” and “Lesser” actually do fit fairly well and could help birders sort out which they are watching while in the field. These remain my first choices, but I am certainly open to seeing what other options are put forth.”

Comments from Areta: “YES. As an author of the work, I would of course vote yes to the recognition of P. salvini as separate from P. albogriseus, as I did in Proposal #906. Indeed, it was that proposal by Van the one that launched the whole process, which lead to the Musher et al. work that forms the basis of this proposal.”

Comments from Areta on English names: “Regarding the common names, I would like to stress that the whole point of our naming suggestions is to have names aiming to diagnostic features of each taxon. Greater and Lesser perpetuate the myth that one can judge size differences in the field even when lacking comparative views: it is easy to see the size differences of Lesser and Greater yellowlegs when they are side-by-side, but side-by-side comparisons on the same branch of P. salvini and P. albogriseus would be a miracle. However, they may coexist at least seasonally indicating that being aware of how to distinguish them is key to a better understanding of their distributions, and the only field characters that we were able to consistently assess based on several hundred pictures where the amount of white on the upper wing coverts (i.e., THE band) and the relative bill width. So, to me, even if not perfect, Slender-billed and Broad-banded indicate the two key features that most observers under field conditions will be able to evaluate. Slender-billed is also slender-banded, and Broad-banded is also broad-billed (indeed, Slender-billed is among the most slender-billed Pachyramphus). Piece of cake, easy to remember, "easy" to use in the field, and informative. As Dan argued, using Eastern and Western does not work, given the complicated distributions. As for X and Y Pied Becard, I think there is just too much wording in there, which adds little to their ID, and these are not more pied than other becards. Songs are also diagnostic...”

Comments from Robbins: “YES. A very thorough proposal that made this a straightforward decision. Yes to treating salvini as a separate species.

“Although I don’t vote on English names, if I did, I would support Dan’s “pied” names!”

Additional comments by Lane: “I recognize that Nacho and all had reasons for the English names they proposed in their paper and this proposal, but I find the reasons Nacho has put forward above to be unsatisfactory. For the average user, it will be unclear that "Slender-billed" and "Broad-banded" are meant to distinguish the two former Black-and-white groups from one another specifically, and *not* from all other Pachyramphus! There are other Pachyramphus (P. versicolor, for example) that are slenderer-billed, and several that have similarly broad upper wingbars. I further have looked at several specimens, and a ream of photos on Macaulay, and concluded that the upper wingbar is not a particularly helpful character to distinguish these two species in the field. Bill size (proportional length with respect to head size, especially) seems to be better and voice better still! As a tour guide who uses English names a lot, and regularly gets questioned by clients about "why did they change the names?" I believe it is best to select names that make clear that these two species are nearly identical -- they are basically large and small copies of one another with very few clear phenotypical characters to separate them. In that vein, Greater and Lesser Pied Becard does manages to convey these several ideas about as well as I can fashion. And as Nacho says that "Slender-billed" also has a "slender band" and "Broad-banded" has a "broad bill"... well "Lesser" has a smaller bill AND band, and "Greater" has a larger bill and band, so these two names seem to better satisfy the issue than the more confusing ones originally proposed. But in the end, this will have to be decided in a separate proposal anyway, methinks.”

Comments from Claramunt: “YES. The new study further clarifies the situation.”

Comments from Stiles: “YES to split salvini from albogriseus; and I consider Greater and Lesser Pied Becard to be the most digestible E-names.”

Comments from Del-Rio: “YES.”