Ukaa 003
Ukaa 003
Ukaa 003
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RESEARCH ARTICLE
ABSTRACT
Tropical mountains feature marked species turnover along elevational gradients and across complex topography,
resulting in great concentrations of avian biodiversity. In these landscapes, particularly among morphologically
conserved and difficult to observe avian groups, species limits still require clarification. One such lineage is Scytalopus
tapaculos, which are among the morphologically most conserved birds. Attention to their distinctive vocal repertoires
and phylogenetic relationships has resulted in a proliferation of newly identified species, many of which are restricted
range endemics. Here, we present a revised taxonomy and identify species limits among high-elevation populations of
Scytalopus tapaculos inhabiting the Peruvian Andes. We employ an integrated framework using a combination of vocal
information, mitochondrial DNA sequences, and appearance, gathered from our own fieldwork over the past 40 yr and
supplemented with community-shared birdsong archives and museum specimens. We describe 3 new species endemic
to Peru. Within all 3 of these species there is genetic differentiation, which in 2 species is mirrored by subtle geographic
plumage and vocal variation. In a fourth species, Scytalopus schulenbergi, we document deep genetic divergence and
plumage differences despite overall vocal similarity. We further propose that an extralimital taxon, Scytalopus opacus
androstictus, be elevated to species rank, based on a diagnostic vocal character. Our results demonstrate that basic
exploration and descriptive work using diverse data sources continues to identify new species of birds, particularly in
tropical environs.
Keywords: integrated taxonomy, Neotropics, systematics, tapaculo, vocalizations
Copyright © American Ornithological Society 2020. All rights reserved. For permissions, e-mail: [email protected].
2 New Scytalopus tapaculos from Peru N. K. Krabbe, T. S. Schulenberg, P. A. Hosner, et al.
Los datos fueron obtenidos mediante nuestro trabajo de campo en los últimos 40 años, complementados con información
de archivos de sonidos y especímenes de museo. Describimos tres nuevas especies endémicas de Perú. Dentro de las
tres existe diferenciación genética, que en dos casos se asocia con variación geográfica sutil en plumaje y vocalizaciones.
Documentamos que en una cuarta especie, Scytalopus schulenbergi, existe divergencia genética profunda y variación en
plumaje a pesar de que las poblaciones son similares vocalmente en general. Además, proponemos que un taxón de otra
región, Scytalopus opacus androstictus, se eleve al rango de especie, con base en un carácter vocal diagnóstico. Nuestros
resultados demuestran que las exploraciones básicas y un trabajo descriptivo basado en fuentes de datos diversos sigue
permitiendo identificar nuevas especies de aves, particularmente en ambientes tropicales.
1997, Gill and Donsker 2019, Remsen et al. 2019, Cadena METHODS
et al. 2020). When omitting S. griseicollis, which belongs
in a separate clade (see Avendaño et al. 2015, Cadena Fieldwork and Study Sites
et al. 2020), S. [magellanicus] is a monophyletic group The fieldwork was carried out through large parts of the
comprising nearly a quarter of Scytalopus taxonomic di- central Peruvian Andes between 1978 and 2018. Details on
versity: 10 species with 2 additional described subspecies. study sites, dates, and individual authors’ participations are
Most forms replace each other geographically, although 2 given in Appendix C.
Morphological Comparisons space using the prcomp function in R 3.5 (R Core Team
We performed visual assessments of plumage and 2013). We described plumage coloration using Munsell
took morphological measurements from Scytalopus Soil Color Charts (1994, 2000).
[magellanicus] specimens from central and southern Peru
(Appendix A). DFL took additional measurements (bill Molecular Identification and Phylogeny
depth) from many of the same specimens as well as from A comprehensive Scytalopus phylogeny based on mito-
some material in AMNH and USNM. Most specimens chondrial NADH dehydrogenase 2 (ND2), ultraconserved
were gathered together for assessment at LSUMZ, but a elements (UCEs), and exon sequences was recently
few specimens were reviewed and measured separately by completed by Cadena et al. (2020). We sequenced 16 older
PAH and MBR. We measured wing (flat), tail, tarsus, and Scytalopus specimens lacking vocal data, including the holo-
bill (from distal edge of the operculum to tip and depth type of S. altirostiris (ANSP 115273 from Atuén, Amazonas),
at base). In addition to comparing measurements individ- and we obtained ND2 sequences using methods designed for
ually, we also performed principal components analysis UCE capture and sequencing, to confirm their identities by
to determine if taxa occupy unique areas of multivariate comparison with the Cadena et al. (2020) reference database.
Briefly, we sampled dried tissue from specimen toepads and be S. [magellanicus] sequences (GenBank accession num-
prepared them for sequence capture and Illumina sequencing bers MN729326-34), and were added to the 56 individuals
following Salter et al. (2019). From demultiplexed samples, previously sequenced (Cadena et al. 2020) along with 2
we quality-controlled reads using Trimmomatic (Bolger outgroup species (S. acutirostris, S. diamantinensis), totaling
et al. 2014) default settings, and then readmapped to pu- 66 sequences. This alignment represented all named taxa in
tative conspecific Scytalopus ND2 sequences in Geneious S. [magellanicus] as well as unassigned populations from
6 (Kearse et al. 2012). Following readmapping, we checked central Peru.
sequence quality manually and extracted consensus ND2 From the new combined dataset, we inferred a ND2 gene-
sequences. Nine of these 16 specimens were confirmed to alogy using maximum likelihood (ML, RAxML, Stamatakis
Amazonas, Peru, grouped with a specimen referred to genetically and vocally but not in morphology or plumage.
altirostris from Unchog, Huánuco, and was only distantly The southern population is also identifiable by plumage
related to a vocally distinct population also traditionally characteristics. We formally describe these 3 species here.
thought to represent altirostris and occurring in Amazonas,
San Martín, and Huánuco north of the Río Huallaga Scytalopus krabbei sp. nov.
(Cadena et al. 2020). The affinities of the S. altirostris [T. S. Schulenberg, D. F. Lane, A. J. Spencer, F. Angulo,
type confirm that the latter population represents a third and C. D. Cadena]
undescribed and unnamed taxon. White-winged Tapaculo
Each of the 3 unnamed populations fulfills criteria ex-
pected of Scytalopus species. They are each vocally diag- Holotype
nosable and have monophyletic mitochondrial sequences. LSUMZ 174041; adult male, collected by Daniel F. Lane
The northern population (Amazonas, San Martín, and (DFL 1753) in humid temperate shrub forest on June 29,
Huánuco north of the Río Huallaga) is identifiable by 2002, in Peru: San Martín/Amazonas border: east slope of
plumage characteristics and morphological measurements, Cerro Patricia, ~22 km ENE of Florida, Camp Buena Vista
and occurs locally in parapatry with S. altirostris. The Social Club, Bosque de Proteccion Alto Mayo, coordinates:
central population is distinguishable from S. altirostris 5.723°S, 77.754°W, elevation: 2,975 m. Vocalizations audio
TABLE 1. Some measurements (range, mean ± SD, number of individuals sampled) of songs of some members of Scytalopus
[magellanicus] from Peru and immediately adjacent Bolivia. Note the rapid pace of krabbei, the few and slow-paced strokes in churrs
of S. frankeae, birds from Junín with fewest strokes and highest pitch. Note also the single-noted song of S. whitneyi, highest pitched in
birds from Ayacucho. The slow-paced strokes and high pitch in churred song from Ayacucho (n = 1) might also prove to be a constant
difference from Apurímac birds.
Duration
Pace of Number Pace of strokes Pace of of note in Loudest pitch
churrs Duration of of strokes in churr Loudest pitch 1-noted song 1-noted of 1-noted
recorded by Daniel F. Lane (ML531469/XC237624). Tissue of S. altirostris and several other S. [magellanicus] spe-
samples preserved (LSUMZ B-43813). ZooBank registra- cies. The head, mantle, throat, and breast of fully adult
tion A77E1A34-6D32-420B-9BE0-2C54E5769E0B; ND2 S. krabbei are generally grayer and less brownish than those
sequence GenBank MN692513. of S. altirostris, and its rump and back are less prominently
barred. The tail of Scytalopus krabbei is dusky finely mottled
Diagnosis with tawny markings, rather than clearly pale brown with
Scytalopus krabbei has long been confused with dark barring or vermiculations as in S. altirostris. Outside
S. altirostris, but these 2 species differ from each other in of the S. [magellanicus] complex, S. krabbei differs from
many respects and are only distantly related to each other. syntopic S. acutirostris from Huánuco and Amazonas in
Most noticeable, all known S. krabbei specimens have being lighter gray and in having brown flanks with sparse
a small white wing patch formed by white outer webs of and relatively broad (1.4 mm) blackish barring, whereas
the outer 2 or 3 greater primary coverts, which differen- specimens of S. acutirostris in this region have the flanks
tiate it from all other Scytalopus except for some males of dark with little or no brown and no barring. The tail of
the allopatric S. opacus androstictus (Krabbe and Cadena S. acutirostris also differs from S. krabbei in being uni-
2010). Additionally, S. krabbei averages larger (Table 2), form blackish. The back of S. krabbei is usually paler and
with a thinner bill throughout its length. By contrast, the browner, but in fully adult specimens (including the holo-
base of the mandible is deep in S. altirostris, as its name type) it is as dark as in S. acutirostris. Within the Cordillera
indicates. Scytalopus krabbei lacks the whitish supercilium Colán in Amazonas department, S. krabbei differs from
flattened wing, bill measure from fore edge of operculum to tip. There was no suggestion of mensural differences between simonsi from Cusco (n = 3 mm, 2ff ) and Puno (n = 10 mm, 6ff ).
Taxon Sex Weight (g) Wing (mm) Tail (mm) Tarsus (mm) Bill (mm)
affinis mm 13–18.6 (15.3 ± 2.1), n = 5 50–57 (53.1 ± 2.7), n = 7 35–38.2 (37.0 ± 1.3), n = 6 18.2–22.8 (20.3 ± 1.7), n = 7 4.9–7.4 (6.3 ± 1.0), n = 7
krabbei mm 18.0–20.8 (19.1 ± 1.5), n = 3 55–59 (57.0 ± 2.0), n = 3 36.8–40.8 (39.1 ± 2.1), n = 3 23.0–24.6 (23.9 ± 0.8), n = 3 5.8–6.5 (6.1 ± 0.4), n = 3
altirostris mm 17–21 (18 ± 2), n = 4 47–59 (56 ± 4), n = 9 34–39 (36 ± 2), n = 9 18.9–21.8 (20.8 ± 0.9), n = 9 4.9–6.5 (5.8 ± 0.5), n = 9
Frankeae Huánuco mm 16.5–19 (17.6 ± 0.7), n = 19 48–56 (53.9 ± 2.0), n = 20 26.8–36.7 (34.2 ± 2.3), n = 20 18.8–21.4 (20.3 ± 0.6), n = 20 5.4–6.6 (5.8 ± 0.3), n = 20
Frankeae Junín mm 16.1–17.7 (16.9 ± 0.6), n = 5 51–54 (53.0 ± 1.2), n = 5 32.8–35.1 (33.8 ± 0.9), n = 5 17.7–20.8 (19.4 ± 1.4), n = 5 5.0–7.0 (5.9 ± 0.7), n = 5
whitneyi mm 14.7–17.2 (15.6 ± 1.0), n = 5 47.2–53 (51.0 ± 2.0), n = 8 34.5–38.8 (37.3 ± 1.6), n = 8 18.6–20.8 (19.6 ± 0.7), n = 8 5.5–6.4 (5.9 ± 0.4), n = 5
urubambae mm 14.6–16.9 (15.5 ± 0.9), n = 5 49–53 (50.9 ± 1.5), n = 7 32–38 (35.1 ± 2.5), n = 6 19–21 (20.2 ± 0.7), n = 7 4.6–5.7 (5.2 ± 0.6), n = 3
schulenbergi (Peru) mm 12–17 (15.2 ± 1.5), n = 8 52–56 (53.9 ± 1.2), n = 9 31.5–40 (36.5 ± 2.9), n = 9 21.3–23.2 (22.1 ± 0.6), n = 9 5.4–6.2 (5.8 ± 0.2), n = 9
simonsi (Peru) mm 15–17.2 (16.0 ± 0.7), n = 10 46–54 (51.7 ± 2.3), n = 13 34.6–42.2 (37.2 ± 2.4), n = 12 18.6–21 (20.2 ± 0.6), n = 13 5.1–6.5 (5.8 ± 0.4), n = 9
affinis ff 16, n = 1 51–53 (52 ± 1), n = 3 32.6–37.4 (34.8 ± 2.4), n = 3 18.9–20.3 (19.5 ± 0.7), n = 3 6.1–6.4 (6.3 ± 0.2), n = 2
krabbei ff 16.5–18 (17.2 ± 0.6), n = 4 54–61 (58.0 ± 3.2), n = 4 35.8–41.1 (37.7 ± 2.5), n = 4 22.5–25.8 (23.7 ± 1.5), n = 4 5.2–6.9 (6.2 ± 0.7), n = 4
New Scytalopus tapaculos from Peru
ff 21, n = 1 52–56 (54 ± 2), n = 5 31–38 (35 ± 3), n = 5 20.0–21.1 (20.4 ± 0.4), n = 5 5.0–6.2 (5.8 ± 0.5), n = 5
©
Frankeae Huánuco ff 17–18 (17.4 ± 0.4), n = 5 52–57 (54.0 ± 1.7), n = 9 31.2–38.7 (35.1 ± 2.6), n = 9 19.4–22 (20.2 ± 0.8), n = 9 5.5–6.2 (5.9 ± 0.3), n = 9
Frankeae Junín ff 13.7–17.4 (15.5 ± 1.4), n = 5 50–55 (52.5 ± 2.1), n = 4 31.3–33.3 (32.6 ± 1.1), n = 3 15.7–18.3 (16.9 ± 1.1), n = 4 5.9–7.5 (6.7 ± 0.8), n = 4
whitneyi ff 14.2, n = 1 48, n = 1 31.2, n = 1 18.7, n = 1 5.5, n = 1
urubambae ff 17.4, n = 1 53–55 (54 ± 1.4), n = 2 33–34 (33.5 ± 0.7), n = 2 18.8–20 (19.4 ± 0.8), n = 2 5.3, n = 1
schulenbergi (Peru) ff 11.5–16 (14.1 ± 1.9), n = 6 50–54 (51.8 ± 2.1), n = 4 31.8–38.2 (35.8 ± 3.0), n = 4 21–23 (22.1 ± 1.0), n = 4 5.3–5.8 (5.5 ± 0.2), n = 4
simonsi (Peru) ff 13–17 (15.1 ± 1.4), n = 7 49–57 (52.6 ± 2.7), n = 7 31.7–38.1 (34.9 ± 2.4), n = 7 18–21.5 (19.9 ± 1.1), n = 7 5.2–6.1 (5.6 ± 0.3), n = 7
(see below).
Vocalizations
onto the lower belly, and the bill and feet are lighter.
tail 40.8, tarsus 24.6, bill from fore edge of operculum 5.8.
N. K. Krabbe, T. S. Schulenberg, P. A. Hosner, et al.
straight, dusky bars. Iris dark brown, bill slate black, tarsus
and preocular area blackish. Lower back, rump, and upper
narrower than in parvirostris (1 vs. 2.5 mm). Apart from
feather tips on the belly are present in S. krabbei, they are
lacking broad pale feather tips on central underparts, and
TABLE 3. Range, mean, standard deviation, and sample size for depth at base of bill (mm) of some Peruvian Scytalopus species (both
sexes). Species abbreviations are: acuti = acutirostris, affin = affinis, krabb = krabbei, altir = altirostris, frank = frankeae, whitn = whitneyi,
uruba = urubambae, simon = simonsi, and schul = schulenbergi. Scytalopus simonsi and S. schulenbergi specimens are from Cusco, Puno,
and w La Paz. Note that all taxa overlap. The deepest bill measured was of S. altirostris, the thinnest of S. acutirostris.
acuti affin krabb altir frank whitn uruba simon schul
3.0–4.6 3.7–4.6 3.7–4.0 3.8–5.9 3.7–5.5 4.0–4.9 3.5–4.5 3.3–4.5 3.2–4.4
(4.0 ± 0.3) (4.2 ± 0.3) (3.9 ± 0.1) (5.0 ± 1.0) (4.5 ± 0.3) (4.4 ± 0.3) (4.0 ± 0.4) (3.9 ± 0.3) (4.0 ± 0.3)
would represent S. altirostris, the type locality of which Scytalopus frankeae sp. nov.
(Atuén) is a mere 140 km to the south-southeast. In 2002, [K. V. Rosenberg, T. J. Davis, G. H. Rosenberg,
DFL and TV encountered a S. [magellanicus] tapaculo at P. A. Hosner, M. B. Robbins, T. Valqui, and D. F. Lane]
Cerro Patricia, a location southeast of Colán and north of Jalca Tapaculo
Atuén, and obtained poor audio recordings. Again, it was
easy to infer that the Cerro Patricia population was the Holotype
same as that at both Colán and Atuén, although DFL was LSUMZ 128615; adult male, collected by G. H. Rosenberg
supercilium beginning, narrowly, above the anterior edge with a blackish subterminal bar, and the primaries are
of the eye, broadening behind the eye, and extending cinnamon-brown with the inner primaries having a small
posteriorly just above the auriculars, although becoming cinnamon-brown spot on the tip of each outer web. Four
quite narrow again at the posterior end. Nape, mantle, and of 8 females from Junín, however, are similar to males in
scapulars dark gray with a wash of dark reddish brown coloration and pattern except that 2 are lighter gray, richer
(5YR3/2). Lower back, rump, and upper tail coverts cin- brown, and have more pronounced paler gray tips to the
namon brown (7.5YR3/6), each feather with 2 or 3 subter- belly feathers.
his specimens in 1922. Nearly every accessible part of Huánuco and Junín differ by 2.9–3.0% (uncorrected pair-
this region, south to the Chipa, Pasco location, has been wise divergence). Because of this sampling bias, it is diffi-
heavily grazed and burned to manage pastures, and suit- cult to ascertain if variation is gradual or clinal, or whether
able bunchgrass habitat likely occurs today only on the S. frankeae is composed of 2 differentiated populations. If
steepest rocky slopes and cliffs. In addition, no protected the latter, the Junín population could warrant naming, and
areas exist within its range in this region. The same applies northern S. frankeae would occupy a much smaller distri-
for parts of Junín, but the southern population of the spe- bution. We recommend that future survey efforts focus
further south in Apurímac, as it was found to be absent threatened by grazing and burning, thus leaving the spe-
from Polylepis forests at 4,360 m near Laguna Antanay cies vulnerable. The species is widespread within Santuario
despite targeted searching (14.061°S, 73.001°W; Benham Nacional de Ampay (Bosque Ampay), a 36.4 km2 large area
et al. 2011). It might also occur in some of the many unex- declared a wildlife sanctuary in 1987, about half of which
plored patches of seemingly suitable habitat between Río is forested. Unless heavy deforestation or natural disaster
Chalhuanca and Río Pampas. Recorded at elevations of occurs, the population in Bosque Ampay should be vi-
3,500–4,200 m in Ayacucho, 3,150–4,500 m in Apurímac. able (based on observed density we estimate 450–1,800
by 1.2–1.4% in ND2 sequences (uncorrected pairwise di- given by S. frankeae. In all churrs, the following strokes are
vergence). This species could also occur in a limited area of connected and fairly similar to each other, have loudest
treeline forest that occurs south of the Río Mantaro on the second harmonic (as most other Scytalopus songs), and
east slope in Huancavelica, adjacent to Ayacucho. are variably rising, level, or falling in average pitch. The
pace of strokes in the churr is relatively fast, on average
Vocal Diagnosis and Geographic Variation of Other faster than in S. urubambae and slower than in S. affinis,
Peruvian Scytalopus [Magellanicus] but overlapping with both. No calls have been recorded.
FIGURE 6. Scytalopus [magellanicus] ND2 mitochondrial genealogy inferred with BEAST. Node support values are BEAST posterior
probabilities followed by RAxML maximum likelihood bootstrap values. Color highlighted selected populations correspond to those
used in Appendix Figure 7. Although the 3 newly described species, S. krabbei, S. frankeae, and S. whitneyi, together form a nearly
continuous distribution and replace each other geographically from Amazonas to Apurímac, they are not sister taxa, each being
related to another nearby S. [magellanicus] taxon.
birdsong archives and museum specimens. The exist- (Fjeldså and Krabbe 1990, Schulenberg et al. 2010), but
ence of at least 2 unnamed Peruvian tapaculos in the S. until now the affinities of these taxa had remained elusive,
[magellanicus] complex has been known for many years preventing formal description. The presence of a third new
species “hiding in plain sight” in the northern Peruvian Colán in northern Amazonas, where S. altirostris is ab-
Andes was more surprising, however. In addition to these sent, S. krabbei is common at treeline, and S. acutirostris
newly described species, application of our species rec- inhabits closed-canopy forest. At Cerro Patricia in San
ognition criteria also results in the elevation of an extra- Martín, where both S. altirostris and S. acutirostris are ab-
limital taxon from subspecies to species, S. androstictus of sent, S. krabbei occupies treeline scrub and closed-canopy
southern Ecuador and northernmost Peru. Our approach forested habitats. However, at Bosque Unchog, Huánuco,
has significantly increased the known diversity within the where all 3 species co-occur, S. acutirostris occupies
rendering unique populations unidentifiable without the after their initial field discoveries of new taxa. As addi-
aid of genotyping. Alternatively, genetic structure could tional data were collected, the idea expanded to com-
reflect past history, such as signatures of once isolated bine these descriptions into a single collaborate paper.
populations that have since merged, resulting in strong mi- All authors, except CDC and JFS, took part in the field-
tochondrial structure in populations without nuclear dif- work. Specimens were obtained by DFL, GRR, JF, KVR,
ferentiation (Block et al. 2015, Kearns et al. 2018). To date, MBR, NKK, PAH, TJD, and TSS, voice recordings by AJS,
geographic sampling within Scytalopus species has been DFL, FA, GRR, JF, MBR, MJA, NKK, PAH, and TSS. Most
Satellite imagery reveals new critical habitat for endangered Greeney, H. F. (2008). Additions to our understanding of Scytalopus
bird species in the high Andes of Peru. Endangered Species tapaculo reproductive biology. Ornitología Neotropical
Research 13:145–157. 19:463–466.
Bickford, D., D. J. Lohman, N. S. Sodhi, P. K. Ng, R. Meier, K. Winker, Guindon, S., J. F. Dufayard, V. Lefort, M. Anisimova, W. Hordijk, and
K. K. Ingram, and I. Das (2007). Cryptic species as a window O. Gascuel (2010). New algorithms and methods to estimate
on diversity and conservation. Trends in Ecology & Evolution maximum-likelihood phylogenies: assessing the performance
22:148–155. of PhyML 3.0. Systematic Biology 59:307–321.
Block, N. L., S. M. Goodman, S. J. Hackett, J. M. Bates, and Ho, S. Y., and S. Duchêne (2014). Molecular‐clock methods for
selecting partitioned models of evolution for molecular and Terborgh, J. (1977). Bird species diversity on an Andean elevational
morphological phylogenetic analyses. Molecular Biology and gradient. Ecology 58:1007–1019.
Evolution 34:772–773. Tschudi, J. J. (1844). Avium conspectus quae in Republica
Lægaard, S. (1992). Influence of fire on the grass páramo Peruana reperiuntur et pleraeque observatae vel collectae
vegetation of Ecuador. In Páramo: An Andean Ecosystem sunt in itinere a Dr. I. I. de Tschudi. Archiv für Naturgeschichte
under Human Influence (H. Balslev and J. L. Luteyn, Editors). 10:262–317.
Botanical Institute, University of Aarhus, Denmark. pp. Whitney, B. M. (1994). A new Scytalopus tapaculo (Rhinocryptidae)
151–170. from Bolivia, with notes on other Bolivian members of the
(m), 80036 (m); Pasco: LSUMZ 128586 (m), 128587 (m), from the recordist B. M. Whitney [[email protected]]),
128588 (f ); Junín: KU 113886 (m), 113885 (m), 113883 primary song s, secondary song ss, call c, male m, female f.
(m), 113884 (m), 113882 (u), 125653 (u), LSUMZ 127654 acutirostris: (100): Amazonas: XC480722s, 144s, 145s;
(m), 179574 (m), CORBIDI 2558 (m; MBR), 2570 (m; San Martín: XC468108s, 468109c; La Libertad: ML17254s,
MBR), 2603 (m; MBR). 17224s; Huánuco: ML28767excited call,s, 28774c, 28777c,
affinis: (9) Ancash: ANSP 115281 (f ), 115282 (f ), 115283 36057s, 40123c,s, 40145c, 40149c, 40150c, 40154s, 40155s,
(u), 115284 (m), LSUMZ 80597 (m), 80598 (u), 82008 (f ). 40157c, 168647c/s, 168659s, 195137s, 195140s, 195151s,
whitneyi (33): Apurímac: ML128964c, 128965s,fc, Huánuco: In November 1983, NKK and JF (Natural
128966c, 129544ss,s, 129545ss,s, 27249131s (also 27249641 History Museum of Denmark; NHMD) recorded
and 27250361), 27250391s, 27257931s, 75262521ss,s, Scytalopus vocalizations at Bosque Unchog. This site was
75262571s, XC20898s, 33861ss, 36107ss, 36108s, 74412– visited by several LSUMZ expeditions in the 1970s–1980s;
13s, 90570s, 102365s, 102366s, 333300ss, 404224s, in July 1984, GHR collected S. [magellanicus] here. In July–
XC436055/ML230279(1st indiv.)s (also XC436058), August 1985, KVR, GHR, TJD, and TSS (LSUMZ) collected
XC436055/ML230279(2nd indiv. Holotype)s (also specimens and recorded vocalizations of an unknown S.
A B 2
0 strokes
per churr
churr pace
−2
kH
−2 0 2
z
win
g
tartsauils
0.0
bill
−4
−2 −1 0 1 2 3
PC1 (61% variation) −2.5
APPENDIX FIGURE 7. Principal component biplots of selected central Peruvian Scytalopus [magellanicus] vocal and morphological
characteristics. Ellipses around point clouds define 95% confidence. (A) Principal component biplot of S. [magellanicus] churred songs,
including the variables frequency at peak volume (kHz), pace (number of churrs per second), duration of churr, duration of each stroke
in a churr. Species generally occupy unique areas of PC1/PC2 space, and those which overlap substantially (e.g., S. simonsi, S. urubambae)
differ in additional qualitative characteristics. S. schulenbergi is not included because its song is structurally distinctive and difficult to
compare meaningfully with other taxa represented. (B) Principal component biplot of primary (single-noted) songs of S. whitneyi from
Apurímac and from Ayacucho. There is considerable overlap, but Ayacucho songs were on average higher pitched and slower paced.
Variables include frequency at peak volume (kHz), pace (number of notes per second), and duration of single notes. (C) Principal
component biplot of central Peruvian Scytalopus [magellanicus] morphological characteristics. Wing, tail, and tarsus measurements are
highly correlated. All taxa overlap almost entirely, with the exception of S. krabbei, which is slightly larger than other taxa.