Proposal (996) to South
American Classification Committee
Treat Xenops minutus
as three species
That Xenops
minutus may include multiple biological species has long been suspected
(Remsen 2003). It is the objective of the present proposal to suggest that
currently available evidence is sufficient to split X. minutus into
three biological species.
Relevant
information:
1. Xenops
minutus includes three populations that differ markedly in vocalizations. An analysis of geographic acoustic
trait variation documented three distinct vocal groups in Xenops minutus:
one comprising the nominate taxon in the Atlantic rainforest south of the São
Francisco River, one comprising all taxa in lowland Amazonia plus X. m.
alagoanus of the Atlantic rainforest north of the São Francisco River, and
one comprising trans-Andean birds from Central America to northwestern South
America (Boesman 2016). Differences between
the first two groups are particularly remarkable. Vocal differences between the
nominate taxon and the Amazonian taxa sound as great or greater than those
among the three species currently recognized in the genus Xenops.
2. Vocal
differences between Xenops minutus populations generate behavioral discrimination. Field playback experiments between X.
m. obsoletus (part of the Amazonian vocal group) and X. m. littoralis
(part of the trans-Andean vocal group) revealed strong behavioral
discrimination between these two vocal groups (Freeman & Montgomery 2017).
Given that the vocal differences between these two vocal groups translate into
behavioral discrimination (and presumably into premating isolation), the
relatively greater vocal differences between the nominate subspecies and these
two vocal groups should also expected to act as effective premating barriers if
they were to come into contact.
3. The three
vocally distinct Xenops minutus populations show high levels of
genome-wide differentiation.
A phylogenomic analysis of Xenops minutus found three deeply divergent
clades experiencing little to no gene flow, entirely congruent with the three
groups delineated by vocal variation (Harvey & Brumfield 2015).
Currently
available evidence suggests that Xenops minutus is best treated as three
biological species:
Xenops
minutus — monotypic, in
the Atlantic rainforest south of the São Francisco River.
Xenops
genibarbis — polytypic,
including obsoletus, ruficaudus, remoratus, and alagoanus.
Xenops
mexicanus — polytypic,
including ridgwayi, littoralis, olivaceus, and neglectus.
Note: Two of
the three resulting species taxa are paraphyletic in mtDNA gene trees (Burney 2009; Smith et al. 2014, Harvey &
Brumfield 2015). However, they are all
reciprocally monophyletic in species trees inferred from genome-wide nuclear
markers (Harvey & Brumfield 2015). Importantly, nuDNA variation is much more
consistent with phenotypic variation than is mtDNA. For example, the nominate
taxon, which is the most distinct in plumage and song, was not recovered as the
most genetically distinct lineage in mtDNA gene trees, but was so in nuDNA
trees. The latter are more likely to reflect the true species tree.
References:
Boesman, P. (2016). Notes on
the vocalizations of Plain Xenops (Xenops minutus). HBW Alive
Ornithological Note 85. In Handbook of the Birds of the World Alive.
Lynx Edicions. https://doi.org/10.2173/bow-on.100085
Burney, C. W. (2009). Comparative
phylogeography of Neotropical birds. PhD thesis. Louisiana State
University. https://repository.lsu.edu/gradschool_dissertations/2682/
Freeman, B. G., &
Montgomery, G. A. (2017). Using song playback experiments to measure species
recognition between geographically isolated populations: A comparison with
acoustic trait analyses. The Auk, 134(4), 857–870. https://doi.org/10.1642/AUK-17-63.1
Harvey, M. G., &
Brumfield, R. T. (2015). Genomic variation in a widespread Neotropical bird (Xenops
minutus) reveals divergence, population expansion, and gene flow. Molecular
Phylogenetics and Evolution, 83, 305–316. https://doi.org/10.1016/j.ympev.2014.10.023
Remsen, J. V., 2003. Family
Furnariidae (Ovenbirds). In: del Hoyo, J., Elliott, A., Sargatal, J., Christie,
D.A. (Eds.), Handbook of the Birds of the World. Lynx Edicions, Barcelona,
Spain, pp. 162–357.
Smith, B. T., et al. (2014).
The drivers of tropical speciation. Nature, 515(7527), 406–409. https://doi.org/10.1038/nature13687
Rafael D. Lima, April 2024
Note
from Remsen: Here is a photograph of specimens that includes the three major
groups, from top to bottom: X. m. minutus, two from the mostly
Amazonian genibarbis group (X. m. remoratus and X.
m. ruficaudus), and X. m. mexicanus.
Note
from Remsen: Here is the BirdLife International/HBW rationale for their 2-way
split of X. genibarbis from X. minutus provided by Marshall
Iliff:
“Hitherto treated as conspecific with X.
genibarbis, but differs in its bold plain white chin and throat connecting
to white submoustachial streak with relatively little
brown on malar below it (prominent and completely separating chin from-submoustachial streak in X. genibarbis, all races of
which have brown-streaked whitish throat) (2); less pale streaking-extending
onto breast (1); loudsong a series of 4–5 upslurred notes, the first slightly
lower-pitched and subdued, vs much-faster-delivered (2), more numerous and overslurred (3) notes (1). Claimed smaller size (in HBW)
not apparent vs (at least)-nominate X. genibarbis, but further study
needed; molecular evidence supports the split (2). Monotypic.-Distribution-E
& SE Brazil (Bahia S to Santa Catarina), E Paraguay (E from Canindeyú,
Caaguazú) and NE Argentina (Misiones).”
HBW/BLI
did not mention splitting mexicanus group from genibarbis
group. The vocal information also comes
from the Boesman analysis cited in the proposal.
Note
on English names:
If the proposal passes, we will do a separate one on English names. HBW/BLI used “White-throated Xenops” for minutus
and left everything else as Plain Xenops.
___________________________________________________________________
Comments
from Robbins:
“YES. Based on vocal and genetic data this seems to be a straightforward split.
Additionally, the clear white throat of nominate appears to be distinct
from the other two taxa (based on the photo included in this proposal and Marshall's
assessment in BirdLife/HBW. Thus, I vote to treat the current Xenops minutus as
three species.”
Comments
solicited from Mike Harvey: “I
believe this proposal is well justified and the recommendation is eminently
reasonable. Although our 2015 genetic study was one of the first to use RAD-Seq
for phylogeography/phylogeny, recent years have established the utility of the
approach for this purpose. I think this fact, combined with the subsequent
vocal and playback analyses, provides more-than-sufficient support for the
existence of these three species.”
Comments from Del-Rio: “YES. Based on
nuclear genome structure and phenotypic differentiation. Vocalizations are also
pretty distinctive.”