Proposal (943) to South American Classification Committee

Recognize Lepidothrix velutina as a separate species from Lepidothrix coronata

Background information:

Many authors have suggested that west-of-Andes populations of Lepidothrix coronata (L. c. velutina and L. c. minuscula) may deserve species-level recognition apart from east-of-Andes populations found in Amazonia and the adjacent Andean foothills (L. c. coronata, L. c. caquetae, L. c. carbonata, L. c. exquisita, L. c. caelestipileata, and L. c. regalis; Hilty, 2021; Kirwan and Green, 2011; Ridgely and Tudor, 1994; Snow, 2004). However, factors including the lack of extensive genomic data from across the large geographic range of L. coronata and the complex plumage variation within Amazonian populations of L. coronata have posed a challenge to clarifying the classification of this species group.

Moncrieff et al. (2022) published a phylogenetic hypothesis of the genus Lepidothrix with widespread sampling within the L. coronata species group, including all currently recognized subspecies (Dickinson and Christidis, 2014). Below is a sampling map, which includes the proposed L. velutina marked with an asterisk.

Moncrieff et al. (2022) used both coalescent and concatenation methods to estimate phylogenies, which consistently identified a deep divergence within L. coronata corresponding to west-of-Andes and east-of-Andes clades. Below is a concatenated tree (estimated with IQ-TREE2 and based on 5,025 SNPs) that is representative of the results found using other methods and data filtering schemes. Numbers at the end of the tip labels refer to locality numbers shown on the sampling map above.

Moncrieff et al. (2022) also pointed out that populations west of the Andes differ markedly in plumage (males have a much deeper black and more extensive black on forehead) and in voice from populations east of the Andes. Below we highlight the main vocal differences with sonograms.

A single trilled primary call from Lepidothrix coronata velutina. By contrast, east-of-Andes populations of L. coronata have a sweet, rising whistle as their primary call. Recording by Jay McGowan from Sendero Ibe Igar, Kuna Yala, Panama (ML202732201).

An abbreviated “ti’ti’t’t’t’t’t’t’t’t” call followed by two “ti’t’t’t’t’t, chu’WAK” advertisement songs from Lepidothrix coronata velutina. The trilled calls and trilled introductions to the advertisement song are unique to west-of-Andes populations of L. coronata. Recording by David L. Ross, Jr. from Parque Nacional Corcovado, Puntarenas, Costa Rica (ML55245).

Populations of L. coronata east of the Andes have males with either a paler black plumage or variations of green with a yellow belly; they also have a whistled primary call and whistled introductory note to the advertisement song that is highly distinctive and consistent.

A series of three “swee” primary calls from Lepidothrix coronata. This “swee” call given by east-of-Andes populations contrasts with the trilled call of west-of-Andes populations. Recording by Curtis Marantz from Parque Nacional do Jaú, Amazonas, Brazil (ML117006).

A single “swee” primary call followed by a series of three “swee chí-wrr” advertisement songs from Lepidothrix coronata. The sweet, whistled calls and whistled beginning to the advertisement song is unique to east-of-Andes populations of L. coronata. Recording by Gregory Budney from Bushmaster Trail, Yanamono Camp, Iquitos, Loreto, Peru (ML34194).

Using locally sympatric species of Lepidothrix (L. coronata and L. coeruleocapilla) as a benchmark, Moncrieff et al. (2022) also pointed out that vocalizations of west-of-Andes and east-of-Andes L. coronata differ more than apparently necessary for maintenance of reproductive barriers within the genus.

Finally, to provide some more context for the level of genetic divergences involved, the mean sequence divergence between west-of-Andes and east-of-Andes L. coronata populations at mitochondrial gene ND2 is 4.25%, which is greater than that observed between species in the L. nattereri + L. vilasboasi + L. iris clade (1.4–3.1%) and between L. suavissima and L. serena (3.7%).

Discussion:

Based on my ongoing genetic work within the east-of-Andes populations of L. coronata, further splits within the species group do not seem warranted. Although L. c. exquisita of the Andean foothills of central Peru is highly distinctive in its plumage around the type locality, it appears to intergrade with L. c. coronata near the Marañón River in the San Martín and Amazonas Regions. More sampling in that area is highly desirable. Also, the voice of Amazonian and east-slope Andean foothill populations is remarkably consistent. If any further splits are considered in the future, these would only involve east-of-Andes populations and should therefore not hold up a split of west- vs. east-of-Andes populations, which are monophyletic and differ in the various ways pointed out above.

Recommendation:

This proposal has three parts:

A. Recognize Lepidothrix velutina (Berlepsch, 1883) as a separate species from Lepidothrix coronata. Lepidothrix velutina would include L. velutina minuscula (Todd, 1919).

B. Use the English name Velvety Manakin for west-of-Andes populations, following use by various authorities (e.g., Hilty, 2021; Snow, 2004).

C. Use the English name Blue-capped Manakin for east-of-Andes populations, following the use of this common name by Hellmayr (1929). This would avoid ambiguity in usage of “Blue-crowned Manakin”.

References:

von Berlepsch, H., 1883. Descriptions of six new species of birds from Southern and Central America. Ibis 5, 487–494.

Dickinson, E.C., Christidis, L. (Eds.), 2014. The Howard and Moore complete checklist of birds of the world, fourth. ed. Aves Press, Eastbourne, U.K.

Hellmayr, C.E., 1929. Catalogue of birds of the Americas. Part 6. Oxyruncidae-Pipridae- Cotingidae-Rupicolidae-Phytotomidae. Publications of the Field Museum of Natural History No. 266.

Hilty, S.L., 2021. Birds of Colombia. Lynx and Birdlife International field guides, Lynx Edicions, Barcelona.

Kirwan, G.M., Green, G., 2011. Cotingas and manakins. Princeton University Press, Princeton.

Moncrieff, A.E., B. C. Faircloth, and R. T. Brumfield. 2022. Systematics of Lepidothrix manakins (Aves: Passeriformes: Pipridae) using RADcap markers. Molecular Phylogenetics and Evolution 173. https://doi.org/10.1016/j.ympev.2022.107525

Ridgely, R.S., Tudor, G., 1994. The suboscine passerines. Volume II. The birds of South America. University of Texas Press, Austin.

Snow, D.W., 2004. Family Pipridae (Manakins), in: Del Hoyo, J., Elliott, A., Christie, D. (Eds.), Handbook of the Birds of the World. Vol. 9. Cotingas to Pipits and Wagtails. Lynx Editions, Barcelona, pp. 110–169.

Todd, W.E.C., 1919. Descriptions of apparently new Colombian birds. Proc. Biol. Soc. Washington 32, 113–118.

Andre E. Moncrieff, June 2022

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Comments from Remsen:

A.YES. The differences in vocalizations alone are sufficient evidence for me, regardless of degree of genetic differentiation.

B. YES. Not only does Velvety have a long track record, but it also is an apt description that is also memorable. The word “velutinus” means “velvety” in Latin, so that’s nice.

C. YES. Good idea. This maintains the connection with “Blue-crowned” and has the bonus advantage of already being used historically. Retaining Blue-crowned for either daughter species would be contrary to our guidelines because this is a classic parent-daughter split with both daughters having large ranges; therefore, retaining “Blue-crowned” for one of the daughters would lead to perpetual confusion. This is a case in which “stability” (retaining Blue-crowned) is disadvantageous because the species classification itself has been destabilized --- time to learn new names to go along with a new taxonomic concept.

Comments from Areta: “A. YES. The deep split, different calls/introduction to song, and distinct plumages argue in favor of the recognition of L. velutina. Regarding the common names, I am fine with Velvety. Changing the name Blue-crowned by Blue-capped seems to create unnecessary instability, for a bird that encompasses more than 90% of the range and for which no species-level name change has been made. Whether these are sister or not is immaterial to me.”

Comments from Lane: “A. YES. Phylogenetic and vocal datasets warrant this split.

”B and C: YES.”

Comments from Donsker:

“B. YES. Velvety Manakin is an excellent English name for the west-of-Andes populations with roots extending at least back to Hellmayr. 

“C. YES. Use Blue-capped Manakin for the east-of-Andes populations by restoring a very appropriate Hellmayr name. “Blue-crowned” Manakin would best be retired, and reserved for the broader species concept, as it was by Meyer de Schauensee, if the split is accepted.”

Comments from Steve Hilty:

“A. Aside from firm genetic evidence (see Moncrieff 2022, gene tree) the vocal differences between west-of-the Andes, and east of the Andes populations is striking. Displaying western birds give a soft rattling trill, this often (but not always) followed by a couple harsh notes. Displaying eastern birds commonly give a slightly rising two-noted "cha-vick" repeatedly (which I have always assumed to be advertising, but could be given in other context); and a simple, rather soft rising "pweeet!" repeated a few times (also advertising, or given in other context?). In any case, the differences between the vocalizations of these western and eastern populations are hard to miss. In fact, to my ears, the two-note call of the eastern birds (carbonata) sounds most like that of Dwarf Tyrant-Manakin, Tyranneutes stolzmanni, although the latter's call is harsher, and will always come from mid-levels or higher in forest (not understory as in case of Blue-crowned (Blue-capped) Manakin.

Moncrieff (2022) also points out plumage differences between western and eastern birds. Although these differences can be discerned in the hand in direct comparison, they are (in my opinion) of minimal value at best for a field observer—especially in the typically low light conditions of tropical forest understory, where individuals of both populations occur. It is likely that these relatively subtle plumage differences (if they are important) are better appreciated in life by the birds themselves, who undoubtedly have sharper color resolution, than we (humans) do. However, because these two populations do not overlap, the importance of these differences, in life, may be minimal.

(voting for Areta)

“B & C: YES”

Comments from Josh Beck (voting for Claramunt):

“B & C: YES. I am in favor of the proposed names (Velvety and Blue-capped). Velvety has precedence, is unique, is apt, and is memorable. For coronata (sensu stricto), although the range is, as pointed out, at least an order of magnitude larger than for velutina, a 10 minute dig into eBird shows that about 65% of observations are trans-Andean, and about 35% are cis-Andean. This is obviously tilted by bias in where birders go, where there are greater numbers of domestic birders, and where eBird is used more, but still clearly shows that there is observation bias towards velutina, which to me is a strong argument against retaining Blue-crowned, despite the instability that will result.”

Comments from Bonaccorso: “YES. The combination of genetic differentiation, voice differentiation, and very modest plumage differences (which, I agree with Steve, seem very difficult to discern in the field), make the case for giving species status to Lepidothrix velutina. Fortunately, the species do not overlap geographically! Nice proposal, by the way. Proposals are much easier to evaluate when all the available graphic information (maps, trees, songs) are included directly in the proposal.”

Comments from Claramunt: “YES. Genetic, plumage, and vocal evidence point to the species status of velutina. MtDNA shows instead velutina sister to the N Amazonian forms of coronata (Smith et al., 2014), but maybe just a case of incomplete lineage sorting.”

Comments from Robbins: “YES for recognizing Lepidothrix velutina as a species given the dramatic differences in vocalizations and genetics between it and east of the Andes populations as thoroughly documented in the Moncrieff et al. (2022) paper.”

Comments from Stiles: “YES on A,B, and C. The genetic and vocal evidence, as well as geography strongly support splitting velutina from the cis-Andean coronata group. I find the claim that retaining  Blue-crowned for velutina would go against SACC policy: the only way this could be possible would be to use Eastern and Western Blue-crowned for the cis- and trans-Andean species, which would go over like a lead balloon for SACC, and I very much doubt that the name Blue-capped for coronata S.S. would cause irremediable confusion for northern birders in Brazil!”

Comments from Schulenberg: YES on B and C. I am totally onboard with Velvety (velutina) and Blue-capped (coronata) manakins.”

Comments from Pacheco: “YES. Genetic and vocal repertoire data sets provide satisfactory support for this division.”

Comments from Jaramillo: “A. YES – The vocalization differences are key in this situation. B. YES – Fits the darker black plumage. Good name. C. YES – I don’t mind keeping Blue-crowned to tell the truth. If used more commonly, and affecting a large portion of the range, then keeping Blue-crowned would be fine. But in this case, as Josh points out the usage is biased to the western population (vetulina), so that makes it more complex.”