Ornitología Colombiana No.6 (2008):24-65

24

NOTES ON TAPACULOS (PASSERIFORMES: RHINOCRYPTIDAE) OF THE EASTERNANDES OF COLOMBIA AND THE VENEZUELAN ANDES, WITH A NEW SUBSPECIES OFSCYTALOPUS GRISEICOLLIS FROM COLOMBIA
Notas sobre tapaculos (Passeriformes: Rhinocryptidae) de la Cordillera Oriental de Colombia
y los Andes venezolanos, con una nueva subespecie de Scytalopus griseicollis de Colombia
Thomas M. Donegan
ProAves Foundation, Caversham, Reading, UK. tdonegan@proaves.org, thomasdonegan@yahoo.co.uk
Jorge Enrique Avendaño-C.1
Escuela de Biología, Universidad Industrial de Santander, Bucaramanga, Colombia
jorgeavec@gmail.com

ABSTRACTWe analysed biometrics, plumage and voice and inspected specimens to study the taxonomy ofvarious high elevation tapaculos Scytalopus of the Eastern Andes of Colombia and the Mérida
Andes of Venezuela. In light of a lack of any diagnostic vocal, plumage or biometric character,we propose treating S. infasciatus as a subjective junior synonym of S. griseicollis. S.
fuscicauda and S. meridanus are indistinguishable by morphology, but we propose treating S.
fuscicauda as a subspecies of S. meridanus in light of small observed differences inintroductions to songs, which require further investigation. As the names were publishedcontemporaneously, we propose priority for S. meridanus over S. fuscicauda. S. meridanus and
S. griseicollis as redefined are each diagnosable vocally, supporting species rank for both ofthem. S. griseicollis gilesi subsp. nov. is described from the Yariguíes mountains. The newsubspecies differs from S. griseicollis in its darker plumage, lower acoustic frequency scolds
and longer tail. The recently discovered Eastern Andes population of S. spillmanni differs from
Ecuadorian populations in its shorter tarsus length and slower song, meeting the requirements
for some, but not all, subspecies concepts. With the status of S. griseicollis, S. meridanus and S.
spillmanni populations clarified, it becomes apparent that two undescribed Scytalopus
populations of the lower montane zone of the west slope of the Eastern Andes and Venezuelan
Andes each await formal description. Other undescribed taxa apparently related to S.
griseicollis or S. meridanus are present in Serranía del Perijá and the northern East Andes.
Notes on geographical variation in S. latrans and Myornis senilis in the Eastern Andes arepresented. Various proposed species and subspecies concepts for allopatric populations are
discussed.
Key words: Scytalopus, tapaculo, new subspecies, species and subspecies limits, Colombia,
taxonomy.

RESUMEN
Analizamos morfometría, plumaje, voces y especímenes para estudiar la taxonomía de varios
tapaculos de altas elevaciones del género Scytalopus de la Cordillera Oriental de Colombia y laCordillera de Mérida de Venezuela. Dada la ausencia de algún carácter diagnóstico en voz,
plumaje o morfometría, proponemos tratar a S. infasciatus como un sinónimo subjetivo más
joven de S. griseicollis . S. fuscicauda y S. meridanus son indistinguibles en su morfología, pero
1

= Present address: Departamento de Ciencias Biológicas, Universidad de los Andes, A.A. 4976 Bogotá, Colombia.

Ornitología Colombiana No.6 (2008)

25

proponemos tratar a S. fuscicauda como una subespecie de S. meridanus debido a posibles
pequeñas diferencias en la introducción de sus cantos, las cuales requieren mayor investigación.
Como sus nombres fueron publicados contemporáneamente, proponemos la prioridad de S.
meridanus sobre S. fusciacuda. S. meridanus y S. griseicollis (cada uno, como redefinidos) son
vocalmente distinguibles, soportando su tratamiento como especies distintas. Describimos S.
griseicollis gilesi subsp. nov. de la Serranía de los Yariguíes. La nueva subespecie difiere de S.
g. griseicollis en su plumaje más oscuro, la frecuencia acústica más baja de los llamados y cola
más larga. La recientemente descubierta población de S. spillmanni en la Cordillera Oriental
difiere de las poblaciones ecuatorianas en su tarso más corto y canto más lento, cumpliendo losrequerimientos de algunos, pero no todos, los conceptos de subespecie. Con la situación de las
poblaciones de S. griseicollis, S. meridanus y S. spillmanni clarificada, es evidente que dos
poblaciones de Scytalopus de la zona montana baja de la ladera occidental de la Cordillera
Oriental y los Andes de Venezuela esperan ser descritas formalmente. Otro taxón sin describir,
aparentemente relacionado con S. griseicollis o S. meridanus, se encuentra en la Serranía de
Perijá. Se presentan notas sobre la variación geográfica en S. latrans y Myornis senilis en la
Cordillera Oriental. Se discuten conceptos de especie y subespecie propuestos para poblaciones
alopátricas.
Palabras clave: Scytalopus, tapaculo, nueva subespecie, límites de especie y subespecie,
Colombia, taxonomía.
INTRODUCTION
Despite some recent research and new taxon descriptions, the tapaculos in the genus Scytalopus
Gould, 1836 of the northern Andes remain among
the most poorly understood bird taxa. Reasons forthe taxonomy of Scytalopus being particularly difficult, such as morphological homogeneity, collecting difficulties and the foxing of skins, have been
widely discussed (Chapman 1915; Whitney 1994;
Arctander & Fjeldså 1994; Krabbe & Schulenberg
1997 and 2003; Cuervo et al. 2005). In a landmark
study, Krabbe & Schulenberg (1997) demonstrated
that various morphologically similar but vocallydifferent populations replace one another by elevation or in different habitats and are best consideredseparate species Three new Scytalopus species
have been described from Colombia in the last decade: Chocó Tapaculo S. chocoensis, Upper Magdalena Tapaculo S. rodriguezi and Stiles’ Tapaculo S.
stilesi (Krabbe & Schulenberg 1997; Krabbe et al.
2005; Cuervo et al. 2005). Such descriptions, together with those of taxa from elsewhere in the
Neotropics (Whitney 1994; Krabbe & Schulenberg
1997; Bornschein et al. 1998, 2007; Mauricio 2005;
Raposo et al. 2006) and the elevation of many races
to species status (principally in Krabbe & Schulenberg 1997; also Fjeldså & Krabbe 1990, Arctander& Fjeldså 1994, Ridgely & Tudor 1994, Born-schein et al. 1998, Coopmans et al. 2001) have increased the number of recognized species in thegenus from around 10 to over 40 since the 1990s.
Birds reach among their greatest levels of diversity,
intraspecific variation and terrestrial ‘endemism’ inthe northern Andes (Stattersfield et al. 1998; Ormeet al. 2005; Phillimore et al. 2007), but recent advances in Scytalopus taxonomy have largely resulted from research in Ecuador, which is only asmall part of this region (Krabbe & Schulenberg
1997). Due to lack of recent field studies, our understanding of the genus Scytalopus in the Eastern
Andes of Colombia remains based largely on early
20th century studies of museum specimens without
associated vocal data (Chapman 1915, Hellmayr
1922, Zimmer 1939). In this paper, we assess thestatus of the taxa S. griseicollis, S. infasciatus, S.
meridanus and S. fuscicauda, and discuss geographical variation in S. spillmanni, with particularreference to the Eastern Andes. We also describe a
new subspecies of S. griseicollis from Serranía delos Yariguíes in Colombia and lay the foundations
for several further new taxon descriptions.
STUDY AREAThe Eastern Andes (Cordillera Oriental) are one ofColombia’s three principal mountain ranges, ex-26Taxonomy of Scytalopus tapaculos

tending from just north of the Equator to the Caribbean coast. They comprise one of the world’s
greatest centers of terrestrial avian endemism, withat least 35 bird species restricted to the range(Stattersfield et al. 1998). Our field surveys wereconcentrated in Serranía de los Yariguíes, with additional fieldwork in Santander, Boyacá and Cundinamarca departments of Colombia. Serranía de los
Yariguíes forms a spur of the Cordillera Oriental,
extending 100 km northwestward, rising to around3400 m. The range is isolated from the rest of thecordillera by the río Sogamoso valley to the northand east, and to a lesser extent by depressions associated with the ríos Horta, Quirola and Opón andtheir tributaries to the south from around 2500m
elevation. The high elevations of the Yariguíes had
not been subject to ornithological study until our
surveys (discussed in Donegan & Huertas 2005,
Donegan et al. 2005 and 2007 and Huertas &Donegan 2006). The region was subsequently declared a National Park (see “Conservation” section
below).
The following Scytalopus taxa were encounteredduring the Yariguíes study: White-crowned
Tapaculo S. atratus (western slope 1300-1600 m); a
presumably undescribed taxon related to S. rodriguezi (western slope 1700-2100 m; eastern slope
2000 m); Blackish Tapaculo S. latrans subsp.
(western slope 1900-2900 m and eastern slope 2600
-2750 m); Pale-bellied Tapaculo S. griseicollis(western slope 2500-3000 m and eastern slope 3100
-3200 m); and a population related to Spillmann’s
Tapaculo S. spillmanni (western slope 2450-2900
m and eastern slope 2700 m). Data relating to
specimens and sound recordings are set out in Appendix 1.

METHODSMUSEUM STUDIES: In addition to studying the material we obtained in the field at Serranía deYariguíes, we inspected all northern Andean Scytalopus specimens at the following museums: Museode Historia Natural, Universidad de la Salle, Bogotá, Colombia (MLS); Instituto de Ciencias Naturales, Universidad Nacional, Bogotá, Colombia(ICN); Colección Ornitológica Phelps, Caracas,
Venezuela (COP); Instituto Alexander von Hum-

Donegan &Avendaño-C.

boldt, Villa de Leyva, Colombia (IAVH); Natural
History Museum, Tring, UK (BMNH); and Museum National d'Histoire Nacional, Paris, France(MNHN). We were provided with photographs of
specimens held at American Museum of Natural
History (AMNH); Museum of Comparative Zoology, Harvard University (MCZ); Field Museum of
Natural History, Chicago (FMNH); Academy of
Natural Sciences, Philadelphia (ANSP) and Carnegie Museum (CM), USA; and Museum fürNaturkunde, Berlin, Germany (MFNU). The specimens (see Appendix 1) included 194 skins labelled
as either “S. griseicollis”, “S. infasciatus”, “S. meridanus” or “S. fuscicauda” from the Colombian and
Venezuelan Andes (excluding specimens bearing
such labels but clearly of other species), as well as
photographs of the type specimens of all of them.
Plumage differences were noted by comparing
specimens directly or by comparing photographs of
specimens, in some cases taken with the same lightbox and camera alongside a standard white to blackcolor array and correcting for darkness in Adobe
Photoshop. In order to evaluate the degrees of isolation vs. overlap between certain taxa, specimen
and sound recording localities were plotted, andmodels of potential distribution built using MAXENT 3.0 (Phillips et al. 2006) based on climate dataobtained from Worldclim (Hijmans et al. 2005).
BIOMETRICS: We took the following measurementsof each specimen: chord of closed wing, tail length(to nearest 0.5 mm), tarsus length, culmen fromskull to tip of upper mandible (to nearest 0.1 mm)
and, from specimen labels, mass (g) (Appendix 2).
Data from birds in juvenile or immature plumage,undergoing moult from juvenile to adult plumage,or undergoing primary moult were excluded. Juveniles of these species were easily identified by theirstrongly and densely barred dark brown / whitish
plumage throughout or in part, compared to a lackof barring (other than rufous / dark brown in theundertail region) in adults. Data for males only
were subjected to principal components analysis(PCA) and analysis of diagnosability “levels” discussed further below, to exclude sex biases.
VOCAL ANALYSIS: Sound spectrograms, generatedusing the default settings of Raven Lite 1.0(sometimes adjusted for brightness, expanded toshow up to c. 5kHz and 2-5 seconds), were studied

Ornitología Colombiana No.6 (2008)
from unpublished and published sound recordings
from plausible localities of S. spillmanni, S. griseicollis, S. infasciatus, S. meridanus and S. fuscicauda in order to assess the taxonomic status of
new populations of S. spillmanni and S. griseicollis
discovered during our Yariguíes fieldwork, giventhat any description could potentially otherwise be
criticized as a result of the uncertain status of S.
meridanus, S. fuscicauda and S. infasciatus. Songsand calls subject to study were defined as sequences of notes broken by gaps significantly
longer than the intervals between individual notesof a sequence. Multiple calls were measured fromsingle recordings and localities when available in27

order to determine the full range of variables. Recordings of scolds of individuals recorded in natural conditions and following playback were included for the same reason. Only scolds, rattles
and alarm calls (not songs) were given directly inresponse to playback by the taxa subject to vocal
analysis in the Yariguíes; thus, song data are likely
not to include a significant number of unnaturally
stimulated individuals. Data on numbers of recordings and spectrograms, together with a gazetteer of recording localities, are presented for eachtaxon and call studied in Appendix 3.
The taxa studied make various calls, all of which

a

d

b

e

f
c
Figure 1: Spectrograms of scolds of S. griseicollis and S. g. gilesi. a) S. griseicollis undescribed subsp. PNN Tamá, Apure, Venezuela (C. Parrish recording no. 6079 on www.xeno-canto.org); b) S. g. griseicollis (=“S. infasciatus”) PNN Chingaza, Cundinamarca, Colombia (J. Parra-B. recording); c) S. g. griseicollis Rogitama, Boyacá, Colombia (J. Zuluaga recording). d) S. griseicollis gilesi, Filo Pampona, Serranía de los Yariguíes (TMD recording); e) S. griseicollis gilesi, Alto Cantagallos (TMD recording). f) curtailed scold of S. griseicollis undescribed subsp., La Pajita, Bucaré Suratá, Santander 3,300 m (JEAC recording,
15 March 2006. Variation in song shape is individual.

28

Taxonomy of Scytalopus tapaculos

b

a

c

Donegan &Avendaño-C.

de

Figure 2: Spectrograms of reeling songs of S. griseicollis and related taxa (or extracts thereof): a) rising reeling song of S. griseicollis undescribed subsp. (JEAC recording Suratá, Santander, 15 March 2006, at 3300 m elevation at La Pajita Farm, Vereda Bucaré, Santander) with accentuated first overtone and flat "zz" at start of call; b) rising reeling song of S. griseicollis gilesi (TMD
recording at Filo Pamplona type locality, Yariguíes, July 2005) which lacks the accentuation of the second overtone but includes
the initial "zz", barely visible on spectrogram at start of call; c) flat reeling song of S. griseicollis (“S. infasciatus”) from PNN
Chingaza, Cundinamarca, Colombia (J. Parra recording), also with introductory "zz"; d) flat reeling song of S. griseicollis gilesi
with initial "zz" (TMD recording, Lepipuerto, Yariguíes, January 2005); e) reeling song of S. meridanus for comparison(Boesman 1999: Páramo de Batallón, Táchira state, Venezuela, 2700-2900 m).

Figure 3: Scolds of S. spillmanni taxa and “S. cf.
canus opacus sp.”. a) S. spillmanni Eastern Andes (a slower example: J.E.A.C. recording, La
Aurora type locality, Yariguíes, Santander, Colombia; b) S. spillmanni Eastern Andes (a fasterexample: TMD recording, Lepipuerto, Yariguíes,
Santander, Colombia); c) S. spillmanni Central
Andes (O. Laverde recording: Reserva Natural
Ibanasca, Ibagué, Tolima, Colombia, IAVH
24245); d) S. cf. canus opacus sp. (N. Krabbe
recording, east slope of Cerro Toledo, ZamoraChinchipe, Ecuador: track 10.38 in Krabbe &Nilsson 2004); e) S. spillmanni Ecuador (N.
Krabbe recording; Quebrada Las Ollas, Sucumbíos, Ecuador: track 44.3 in Krabbe et al. 2001);f) S. spillmanni Ecuador (N. Krabbe recording,
San José, Tandayapa ridge, Pichincha, 2300m:
track 9.16 in Krabbe & Nilsson 2004).

a

b

c

d

e

f

Ornitología Colombiana No.6 (2008)
consist of rapidly repeated up-down, up, or down
strokes. The most common calls comprise: (i) shortscolds which rise and then fall in acoustic frequency (“scold”) (e.g. Figs. 1 & 3); (ii) rising or
flat reeling songs which either rise or are relatively
constant in acoustic frequency and which may bevery short or long (up to over a minute) in duration(“reeling song”) (e.g. Figs. 2, 4 & 5). Scytalopus
spillmanni taxa also make explosive, short “brzk”
calls and females give what Krabbe & Schulenberg(1997) term an “advertising call”. Scytalopus meridanus has been reported making short call notes(Krabbe & Schulenberg 1997, Boesman 1999).
Scytalopus griseicollis rarely makes a slower,
higher frequency version of its scold (Fig. 1). S.
griseicollis and S. spillmanni also give unmusical29

rattles, similar to the scold but with less clearlydefined and higher frequency individual notes.
Only songs and scolds were subject to detailed
analysis, on account of small sample sizes for othervocalizations. Scolds are given in response to playback and apparently have a territorial function.
Songs appear likely to be more relevant to mateselection but are given year-round and may also
have a territorial function.
Various vocal variables described by Isler et al.
(1998, 1999, 2006, 2007, 2008) were considered
for further analysis in a preliminary study of spectrograms. Variables which showed variation
between individuals, spectrograms, or populations
were subject to further analysis, whilst apparently

a

d

b

e

c

f

Figure 4: Reeling songs of S. spillmanni. Left: elongated rising reeling songs of a) S. spillmanni Eastern Andes (J.E.A.C. recording, La Aurora, Yariguíes, Santander, Colombia); b) S. spillmanni Central Andes (L. E. Urueña recording, Ibagué, El Rancho,
road to Nevado del Tolima, Tolima, Colombia); and c) S. spillmanni in Ecuador (N. Krabbe recording, Las Palmas, Cotopaxi;
recording 44.1 in Krabbe et al. 2001). Right: elongated “flat” reeling songs of d) S. spillmanni Eastern Andes (O. Laverde recording: Fusagasugá, Cundinamarca, Colombia, IAVH 17856); e) S. spillmanni Central Andes (O. Laverde: Reserva Natural Ibanasca, Ibagué, Tolima, Colombia, IAVH 24179); and f) S. spillmanni Ecuador (N. Krabbe recording: San Jose, Tandayapa ridge,
Pichincha, 2300m). Variation in acoustic frequency over time between different spectrograms and other differences are subject toindividual, not geographic, variation. However, note the lower frequency Central Andes recordings compared to the Eastern Andes (compared top two and middle two recordings); and the elaborate introduction to the rising reeling song in Ecuador (bottom
left) not found to such an extent in an any of our Eastern or Central Andes recordings (although this introduction is often absent in
Ecuadorian recordings also).

30

Taxonomy of Scytalopus tapaculos

constant variables and those inappropriate for study
were discarded. For example, due to calls andsongs being composed of a number of individual
notes of the same volume but different acoustic
frequencies, frequency variables defined with reference to a “principal” note were not amenable tostudy. Measurements of frequency bandwidthsabcFigure 5: Examples of curtailed reeling songs of S. spillmanni. a) S. spillmanni Eastern Andes (J.E.A.C. recording,
La Aurora, Serranía de los Yariguíes, Santander, Colombia);b) S. spillmanni Central Andes (O Laverde recording, Río
Blanco, Caldas, Colombia); c) S. spillmanni Ecuador (N.
Krabbe recording, Ecuador, Krabbe et al. 2001). Note thedownstrokes in the Ecuadorian recording, not found in East
Andes recordings, but Ecuador recordings include songs ofsimilar note shape to East and Central Andes birds.

Donegan &Avendaño-C.

were avoided because individual notes in the callsof several taxa have short, quiet “tails” to the upstroke, downstroke, or both, meaning that the lowest recorded point on a spectrogram is biased by thequality of recording equipment and distance of abird from the observer. The maximum point provides a more consistent reference point for thesebirds. The following variables were selected forfurther study as apparently varying between differrent populations or individual calls: total number ofnotes in call (n); total song duration (s); song speed
or pace (average number of notes per second: by
dividing number of notes in song by song duration); maximum acoustic frequency of lowest note;maximum frequency of highest note; variation in
frequency (the difference between the latter two
measurements, all in kHz); and subjective description of note shape. Of the various selected variables, number of notes, song duration and frequency variation showed far greater intra- than interspecific variation and a maximum frequency oflowest note was correlated within populations withthe maximum frequency of highest note. None ofthese variables were subject to further analysis.
In order to assess the validity of the putative taxon“S. infasciatus” and differences between otherpopulations, S. griseicollis / infasciatus recordings
were split into four subsets: (i) eastern Cundinamarca and Boyacá (Iguaque region); (ii) Bogotá
region, western Cundinamarca and Boyacá; (iii)
Serranía de los Yariguíes; and (iv) northern sites(Santander, Norte de Santander, and Venezuela).
Data for S. fuscicauda (Lara state) and S. meridanus (other states in Venezuela) were also considered separately. For S. spillmanni populations,data were split into the following geographical subsets, after excluding data from inter-Andean sites in
Ecuador: (i) Colombian Central Andes; (ii) Colombian Eastern Andes; (iii) western slope of Ecuador;(iv) eastern slope of Ecuador. Western Andes recordings and biometrics for S. spillmanni were notconsidered in detail as such populations are thesubject of separate studies (Cuervo et al. 2004;
Krabbe et al. 2006). Some of the above subsets ofdata were subject to further splitting and analysis toassess possible clinal and elevational variation.
STATISTICAL TESTS: For means and standard deviations (Appendix 3) and diagnosability analyses

Ornitología Colombiana No.6 (2008)(Appendix 4) we used a reduced data set with only
the first three spectrograms for which full data wereavailable from each recording. Various diagnosability tests were assessed for pairs of vocal
variables of different populations for song speed
and maximum acoustic frequency of songs andscolds and biometric data.
LEVEL 1: Statistically significant differences at
p<0.05. A Bonferroni correction was applied fo
both vocal data (6 variables) and biometrics (5 variables), to produce testwise significance levels of
p<0.0083 and p<0.01 respectively. For acoustic
frequency data, an unequal variance (Welch’s) ttest was used; for speed data, both KolmogorovSmirnov and Mann-Whitney U tests were used.
These calculations consider statistical significanceof differences, but tolerate a considerable degree ofoverlap.

31

LEVEL 4: Diagnosability based on recorded values(first part of Isler et al. 1998’s diagnosability test).
LEVEL 5: 95% / 95% diagnosability or sample
means four standard deviations apart controlling forsample size (second part Isler et al. 1998’s diagnosability test):|(x1 - x2)| > s1(t1 @ 97.5%) + s2(t2 @ 97.5%)|(x1 - x2)| > s1(t1 @ 99%) + s2(t2 @ 75%)

CRITERIA APPLIED FOR ASSESSING SPECIES ANDSUBSPECIES RANK: Helbig et al. (2002) advocated
the general principle, previously discussed by someother authors cited therein, that in order to assess
the rank of allopatric populations, a comparison
with closely-related sympatric or parapatric populations known to be good species should be undertaken. Isler et al. (1998, 1999, 2006, 2007, 2008),
in studies of species limits using voice for suboscine passerines, suggested assigning species rank toallopatric antbird populations where three or moremeasured vocal variables for loudsongs differeddiagnosably, based on observed differences between closely related sympatric species in that family. A useful comparison for assessing Scytalopus
species limits is provided between the S. spillmanni
population recently discovered in the Eastern Andes (Donegan et al. 2007) and S. griseicollis, whichhave similar calls and replace one another in elevational or habitat parapatry and for which we have alarge sample of recordings. Isler et al.’s diagnosability test (Levels 4 and 5) was met for S.
griseicollis and the Eastern Andes population of S.
spillmanni, but for one vocal variable; and only for
scolds, not songs (cf. Bornschein et al. 2007). For
songs, some shape features are likely to assist identification: for example, introductory notes to songs(if present), but these are not found on all recordings. Scytalopus griseicollis and Eastern Andes S. spillmanni also differ in plumage. We propose supplementing Isler et al.’s diagnosability testwith consideration of the statistical significance ofdifferences (Level 1). We therefore ranked allopatric populations as species where they show Levels 1, 4 and 5 diagnosability for songs or scolds
based on at least one vocal variable and treat populations as conspecific if they do not meet this test.

and|(x2 - x1)| > s2(t2 @ 99%) + s1(t1 @ 75%)

Possible criteria for defining subspecies limits in
Scytalopus have been little explored. Vocalizations

Various further calculations described below werealso carried out to measure differences between
populations in the context of various proposed or
possible species and subspecies concepts. In theformulae used below, x1 and s1 are the sample meanand sample standard deviation of Population 1; x2
and s2 are the sample mean and sample standard
deviation of Population 2; and t is the value of t
using one-sided confidence intervals at the percentage specified for the lower of the degrees of freedom of the two populations for the relevant variable, with t1 referring to Population 1 and t2 referring to Population 2.
LEVEL 2: Hubbs & Perlmutter (1942)’s now littleused subspecies concept, as modified. Sample
mean of Population 1 falls outside range of 95% ofPopulation 2 (or sample means two standard deviations apart, controlling for sample size):|(x1 - x2)| > (s1(t1 @ 97.5%) + s2(t2 @ 97.5%))/2
LEVEL 3: The traditional 75%/99% test for subspecies diagnosability (Amadon 1949; Patten & Unitt
2002), modified to control for sample size:

32

Taxonomy of Scytalopus tapaculos

in suboscines such as Scytalopus are considered to
be innate and stereotypical (Kroodsma 1984; Whitney 1994; Cuervo et al. 2005). Differences in voice
between populations may therefore have a genetic
basis. Isler et al. (op. cit.) applied a “diagnosabilityof one character” test (whether vocal, biometrics orplumage) for subspecies in Thamnophilidae (Level
4/5: essentially a phylogenetic species with small
differences not meriting species rank). The traditional test in ornithology for diagnosing subspecies
is our Level 3 “99+%/75%” test. Luckow (1995)
has alternatively proposed ranking allopatric populations with statistically significant means (ourLevel 1 diagnosability) as subspecies. This approach has been applied to some recent bird subspecies descriptions where statistically significant
differences are shown for a range of biometric andplumage variables (e.g. Stiles & Caycedo 2002).
We do not propose new criteria to delimit species
or subspecies for allopatric taxa. We therefore conservatively propose the recognition of new subspecies only if allopatric taxa meet Levels 1, 2, 3 and4/5 diagnosability for at least one character (i.e.
satisfy all mentioned subspecies definitions). We
propose synonymy at subspecies level only if two
allopatric populations fail to achieve any level ofdiagnosability. Other putative subspecies are discussed but are not described, pending further discussion of the use of subspecies concepts in ornithology. We also consider in this paper the statusof an isolated population that is phenotypically differentiated but not genetically diagnostic from allopatric populations. Proponents of a biological species concept (Helbig et al. 2002; Remsen 2005)
recommend ranking such taxa as subspecies. Proponents of phylogenetic species concepts have proposed ranking such taxa as species (Zink 2005).

SCYTALOPUS TAPACULOS IN THE
EASTERN ANDES OF COLOMBIAThe status of Scytalopus infasciatus and
S. griseicollis
The Pale-bellied (or Matorral or Rufous-rumped)
Tapaculo S. griseicollis (Lafresnaye, 1840) and
Colombian (or Cundinamarca) Tapaculo S. infasciatus Chapman, 1915 are both recorded from theDonegan &Avendaño-C.

Eastern Andes of Colombia. Both have rather lightgray plumage compared to congeners and occur
generally at high elevations. Zimmer (1939) treated
S. infasciatus as a synonym of S. griseicollis andwas followed by Peters (1951). However, S. infasciatus has been ranked as a species by some authors(e.g. Chapman 1915; Krabbe & Schulenberg 1997;
Asociación Bogotana de Ornitología 2000; Gill &Wright 2006), but Krabbe & Schulenberg (1997)
noted that comparison of material from the Eastern
Andes with the type of infasciatus would be necessary before its status could be fully resolved. Thesesame authors later (2003) treated S. infasciatus as asubspecies of S. griseicollis, but restricted the former to a single site and noted that the two taxa
might be synonymous. Remsen et al. (2008) andSalaman et al. (2007) followed this treatment.
The type specimens of S. griseicollis (MCZ 76330
and 76331) are two "Bogotá" skins, an adult and ajuvenile, and are typical of birds currently ascribedto this taxon in leading texts. The juvenile hasstrong barring throughout. The other specimen has
acquired almost complete adult plumage but retains
strong barring on the flight feathers. Although it isnever possible to be certain with "Bogotá" skins,these specimens may have originated from near
Bogotá itself, as S. griseicollis remains common in
hills above Colombia's capital today. The type locality of S. infasciatus (AMNH 132328) is Páramo
de Beltrán, on the eastern slope of the Eastern Andes in Cundinamarca department, some 15 km eastof Bogotá and now in Parque Nacional Natural(PNN) Chingaza. The only unique distinguishingfeature of S. infasciatus per the original description
is its darker underparts compared to S. griseicollis.
Other features of the holotype, such as barring on
the vent, were considered probably due to agerelated variation as such features were also observed in S. griseicollis (Chapman 1915). It has
been hypothesized that some of the various calls
given by S. griseicollis may have been calls of S.
infasciatus, that the two taxa may separate by extent of barring on the underparts, and that S. griseicollis is more common in drier or secondary habitats, with S. infasciatus present in more humid
habitats or better-conserved forest (Asociación Bogotana de Ornitología 2000). The darkness hypothesis is considered further below. The “barring”
hypothesis has some support from the type speci-

Ornitología Colombiana No.6 (2008)
men but was rejected by Chapman (1915) in thetype description and appears to involve merely agerelated variation. Having observed and sound recorded individuals of identical song on different
mountain slopes and in drier, more humid, primaryand disturbed habitats of the Eastern Cordillera, wecan also reject the “habitat” hypothesis.
Although we were unable to make direct comparisons, the S. infasciatus holotype appears unexceptional in its plumage. Its dorsal and ventral plumage are within the range of variation of specimens
at ICN and other museums assigned to either S.
griseicollis or S. infasciatus. A recent plate of "S.
infasciatus" (Krabbe & Schulenberg 2003) exaggerates considerably the darkness of the plumage ofthe type. Twelve recently-collected and older Eastern Andes Scytalopus specimens, including fromPNN Chingaza / Páramo de Beltrán region do notdiffer significantly in their plumage from Bogotá
region specimens in the ICN. Darker skins previously labelled S. infasciatus and lighter ones previously labelled S. griseicollis at ICN come from avariety of localities and regions. Dark / light plumage is not strongly linked to sex differences, withintra-sex variation exceeding inter-sex variation.
There is no clear division of the series inspected
into “darker” and “lighter” specimens, nor into
barred or unbarred specimens; rather, there is acontinuum of plumage variation. A similar extent
of variation in underpart coloration and barring isevident in the series of S. meridanus inspected andwithin our three specimens of the Yariguíes population of S. griseicollis. No statistically significant(Level 1) differences in biometrics were noted between specimens from Iguaque and the Bogotá region.
Flat and rising reeling songs and scolds are given
by S. griseicollis / infasciatus populations throughout the Eastern Andes with no geographical separation of such song types. A sample including 12
recordings of reeling songs and 11 of scolds (withmany more spectrograms) from PNN Chingaza region are not diagnosable (even to Level 1) fromwestern Cundinamarca or Boyacá recordings.
Also, no known geographic barrier exists between
páramos in PNN Chingaza and those in hills above
Bogotá that might act to isolate S. griseicollis from
“S. infasciatus”. Given the lack of statistically33

significant differences in plumage, biometrics orvoice between populations from the region of the S.
griseicollis and S. infasciatus type localities, thesimilarity of type specimens and lack of any obvious isolating barrier, we concur with Krabbe &Schulenberg (2003) that S. griseicollis and S. infasciatus should be treated as conspecific. In addition,
since it fails to meet the requirements for any of thesubspecies concepts mentioned above, we consider
infasciatus to be a subjective junior synonym of S.
griseicollis.

A new subspecies of Scytalopus griseicollis
The isolated Serranía de los Yariguíes populationof S. griseicollis shows diagnostic plumage andbiometric differences and statistically significant
mean vocal differences from populations in themain Eastern Andes. We therefore describe it as:
Scytalopus griseicollis gilesi, subsp. nov.
HOLOTYPE.– Adult male (Fig. 6), collected by theauthors and prepared by J. Avendaño-C. (field
number JEAC 297) on 13 July 2005 in páramo at
Filo Pamplona above La Aurora, vereda San Isidro,
Municipality of Galán, Santander Department, Colombia (06º38’N; 73º24’W; 3200 m) on the eastern
slope just below the main ridgeline of the Yariguíes
massif. It is deposited at the Instituto de Ciencias
Naturales, Universidad Nacional, Bogotá, Colombia (no. ICN 35609). Tissue samples (heart) and a

Figure 6: The holotype of S. g. gilesi. Photographs by B.
Huertas/Proyecto YARE.

34

Taxonomy of Scytalopus tapaculos

skeleton were deposited at the Museo de Historia
Natural at Universidad de los Andes in Bogotá.
Sound recordings of the holotype prior to capture
are deposited at IAVH and the British Library andare available at www.xeno-canto.org (XC 18452,
18453, 18454, 18455 and 18456).
DIAGNOSIS.– Scytalopus griseicollis gilesi exhibits
all the characteristics of the genus Scytalopus(Ridgway 1911; Krabbe & Schulenberg 1997;
Cuervo et al. 2005). It appears to be most closely
related to S. griseicollis and S. meridanus on account of its rather gray plumage and orange-rufous
vent, similar calls and a páramo and subpáramo
habitat of the Eastern Andes of Colombia. A preliminary molecular study found a specimen of eachof S. g. griseicollis and S. g. gilesi to share the same
mtDNA haplotype for the second subunit of the
NADH deshydrogenase gene (ND2; C.D. Cadena
in litt. 2007).
Scytalopus griseicollis gilesi is diagnosable from S.
griseicollis populations by its darker and lessbrownish back and tail and its darker, more slate
gray, underparts (Fig. 7). It also differs from thenominate in its longer tail (Levels 1, 2 and 4; or 4
only compared to the undescribed northern subspecies discussed below), slower and lower song
(Level 1) and lower frequency scold (Level 1; Levels 1, 2 and 4 compared to the northern subspecies:
Fig. 1). Yariguíes specimens cluster separatelyfrom those of other Eastern Andes populations in
PCA (Fig. 9), but a larger sample might show someoverlap. We did not hear long reeling songs (cf. up
to 15 seconds given by nominate S. griseicollis) at
any of our study sites (Level 1; also F-test, p<0.01).
The new subspecies does not appear to represent
the extreme of a cline. We have inspected specimens and sound recordings of nominate S. griseicollis from the Eastern Andes at sites adjacent toSerranía de los Yariguíes at Suratá in Santander(ICN 36121; J. Avendaño-C. sound recordings) andthe Boyacá/Santander border at Arcabuco (ICN
10852; R. Chavarro & J. Zuluaga sound recordings), each of which are c.60 km from Serraníade los Yariguíes in the main section of the Eastern
Andes. These sound recordings and specimens are
consistent with those of S. g. griseicollis and theunnamed northern subespecies respectively.

Donegan &Avendaño-C.

Scytalopus griseicollis gilesi can be distinguishedfrom S. meridanus of the Venezuelan Andes by its
lower song and faster and lower scold (up to Level
5, in each case), different note shape (Level 4),
longer wing (Levels 1, 2, and 4), and longer tail(Levels 1 and 4) (Fig. 8). It differs from S. spillmanni populations to varying degrees in its lighter
plumage, slower reeling song, faster trill, shorter
wing, shorter tarsus, and lower mass (see Appendix
4 for details). Scytalopus g. gilesi shares a numberof morphological and vocal features with the Central Andes population of S. spillmanni, the songs ofwhich overlap for all measured variables. The only
recent Central Andes specimen of S. spillmanni
studied is lighter plumaged than southern andeastern populations of the same species, whilst S. g.
gilesi is a darker version of S. griseicollis. However, the diagnostic difference in scold speed requires species rank between these populations.
DESCRIPTION OF THE HOLOTYPE.– Color nomenclature follows Munsell Color (1977; 2000). Bill black(not coded); ventral proximal lower mandible
slightly lighter at base (Gley 1 7/N); iris darkbrown (10R 3/3); head, mantle and tail dark graybrown (7.5YR 2.5/2); underparts to breast gray
(Gley 1 3/N), becoming slightly lighter (Gley 1 4/N) on lower belly; wing coverts and flight feathers
dark gray (7.5YR 2.5/2) with tertials tipped
ochraceous brown (5YR 3/4), with dark gray
(7.5YR 2.5/2, but darker than other wing feathers)
subterminal bar; wing rounded with wing point
hard to discern but close to fifth (sixth from outermost) primary; no emarginations or primary
notches noted; flanks and undertail coverts
ochraceous brown (7.5YR 3/4) barred black (notcoded) on flanks; rump brown (5YR 3/4) thinlybarred dark gray (7.5YR 2.5/2); tarsus reddish darkgray (5YR 4/3) frontally, behind lighter (10R 3/1);
Foot soles grayish yellow (2.6Y 8/2). All feathers
fresh with no moult noted; testes rather enlarged(left testis: 5.4 x 2.2 mm; right testis 4.2 x 2.7 mm);stomach contents not determined. Measurementsof holotype: maximum flattened wing (field) 62,
wing chord (skin) 58, tail 45, tarsus 22.5, total culmen 13.5, exposed culmen 11.0; mass 18.0g.
PARATYPES AND OTHER MATERIAL.– We designate
the following paratypes: (1) Immature male no.
ICN 35610, collected by the authors and preparedOrnitología Colombiana No.6 (2008)

35

Figure 7: Dorsal and ventral views of (left to right): (i) S. g. gilesi holotype; (ii) S. g. gilesi Pamplona paratype; (iii) Specimens of
S. g. griseicollis ICN 34780 (male, Cundinamarca, Bogotá, Cerros Orientales, La Aurora, collected on 15/11/2003 by F.G.
Stiles); (iv) ICN 34492 (female, Cundinamarca, Bosque de Toma, Cerros Orientales de Bogotá, 3050m, collected on 1 April 1993
by A. Guttiérez & S. Rojas, labelled "griseicollis" and an example of one of the lighter specimens); (v) ICN-31235 (male, Cundinamarca, Chingaza, Piedras Gordas; collected by F.G. Stiles on 1 November 1991, a relatively dark bird formerly labelled"infasciatus"); (vi) ICN 19623 (male, Cundinamarca, Bogotá, Monserrate, collected by P. Bernal on 19 June 1968). Photographs
at ICN by TMD.

36

Taxonomy of Scytalopus tapaculos

Donegan &Avendaño-C.

Figure 8: Principal Component Analysis using bill, tail, tarsus and wing data for adult males only of S. griseicollis ("A": all Eastern Andes north to Santander), S. meridanus ("M": all Venezuelan Andes), S. sp. of Perijá ("P": all Perijá range of Eastern Andes), S. griseicollis undescribed subsp. from the Santander-Tamá region ("T": Eastern Andes in Norte de Santander, Colombia
and Apure, Venezuela) and S. g. gilesi ("Y").

by T. Donegan on 14 July 2005 at the Filo Pamplona type locality. Tissue samples (heart) and askeleton have also been deposited at Universidad
de los Andes. (2) Adult male no. ICN 36175, collected and prepared by J. Avendaño-C. (JEAC 377)
on 12 November 2006 on a stunted ridge above
Alto Cantagallos, above Santo Domingo farm, Municipality of San Vicente de Chucurí, Santander
Dept, Colombia (06º48’N; 73º21’W; 2450 m) on
the western slope of the Yariguíes massif. Tissue
samples (heart and liver) have also been depositedat Universidad de los Andes (ver Fig. 7).
The Pamplona paratype had smaller testes (left: 3 x
1.5mm; right: 2.5 x 1.5mm) than the holotype. As
the latter was collected just a few days earlier andwas in breeding condition, it is possible that theparatype was a younger bird. It has darker grayunderparts than the holotype (Gley 1 3/N), notlightening so significantly on the lower belly; andmore brownish (7.5YR 3/4) plumage on the upper
mantle, neck and crown. Measurements are as follows: maximum flattened wing (field) 61, wing
chord (skin) 56, tail 44, tarsus 22.0, total culmen
13.5, exposed culmen 11.0, mass 17.5 g. Stomach
contents included Coleoptera exoskeleton remains.
The Alto Cantagallos male had rather large testes(left: 4.6 x 2.7mm; right: 4.2 x 3.2mm). It is similar to the holotype in its coloration but lacks barringon flanks and undertail coverts. Barring on the ventis a character that is also variable in nominate S.
griseicollis (Chapman 1915) and S. meridanus (seeabove). Measurements in mm are as follows: maximum flattened wing (field) 57, wing chord (skin)57, tail 44, tarsus 23.0, full culmen 13.2, exposed
culmen 9.3.
We, E. Briceño and B. Huertas have observed individuals consistent with the type series at very closequarters on approximately 20 occasions during
three weeks' total fieldwork at sites at which thespecies was relatively common in the Yariguíes.
ETYMOLOGY.– This bird is named for O. A. Robert
Giles of Wimbledon, London, UK. His name isformed as a fictional first declension masculine
Latin noun and declined in the genitive singular.
Robert is a keen birder and conservationist and afriend of the first author. He is among the founders
and a member of the Consejo of Fundación
ProAves, a bird conservation NGO based in Colombia, and is chairman of Ecoturs S.A., an ecotourism organization operating in Colombia that
donates all its profits to ProAves. Robert has sup-Ornitología Colombiana No.6 (2008)37

ported many conservation initiatives in Colombia.
In particular, Robert's support was crucial in providing seed funding for various ProAves nature
reserve purchases, including for the reserve in
Serranía de los Yariguíes near where this new
tapaculo is found.

was heard once from a peculiar primary bamboo
forest just below the ecotone, characterized by tall
emergent trees, an open canopy with little midstorey and thick bamboo (Poaceae: Chusquea) understory. Below this elevation, it was again replaced
by S. latrans and S. spillmanni.

ECOLOGY.– S. g. gilesi is a skulking bird observed
most frequently foraging in dense shrubs of páramo
and subpáramo ridgetop habitat up to 2 m above
ground level. Stomach contents suggest it is insectivorous, typical of the genus. It seems to be mostnumerous in slightly taller scrub found in shelteredalcoves, along exposed ridges and just above thetreeline. The area is subject to quite extraordinarylevels of precipitation and near-constant groundlevel cloud (fog). Our own rainfall readings takenin the field and data in Worldclim (Hijmans et al.
2005) reveal higher levels of precipitation for sites
where S. g. gilesi is predicted to be present in
Serranía de los Yariguíes than the average for sites
where S. g. griseicollis is predicted to be present in
the Eastern Andes (average 1400 vs. 1900 mm/
year; Welch’s t, p<0.001, d.f.=523). The humid
habitats of S. g. gilesi may have given rise to adarker population, per "Gloger’s rule" (Zink &Remsen 1986). Plants found in S. g. gilesi's habitat
include Blechnaceae (Blechnum schomburgkii),
Eriocaulaceae (Paepalanthus), Ericaceae, Clusiaceae, Bromeliaceae, Melastomataceae, Orchidaceae, unusual Asteraceae (Espeletia) and some isolated tall palms (Arecaceae). At Alto Cantagallos(2450m) on the western slope, S. g. gilesi was present along an exposed ridgetop only, which presented stunted vegetation (2m high “canopy”) including short, gnarled, winding trees and Espeletia
of apparently the same species as in the páramo.

DISTRIBUTION.– We recorded S. g. gilesi at highelevations along almost the entire length ofSerranía de los Yariguíes in three municipalities(Fig. 9). However, no skins of this taxon exist incollections from the main Eastern Andes: apparently it is endemic to the Yariguíes range. Although S. griseicollis was recently sound recorded
as low as 1300 m in Santander (A. Hernández-J. inlitt. 2007), this was in a human-altered habitat. In
primary forest, we have not recorded S. griseicollis
below 2,450 m in Serranía de los Yariguíes. Above
this elevation, the Yariguíes massif is separated
from the main Andean cordillera to the south bysome 50km of depressions associated with the rivers Opón and Horta. To the north, west and east,
the massif is isolated below the 800m contour byarid low valleys of the Magdalena, Sogamoso andSuárez rivers. Scytalopus griseicollis has been collected and sound recorded at sites close to this ecological divide in the main Eastern Andes: Arcabucoon the Boyacá /Santander border (c.60 km from our
Lepipuerto site) and Suratá and Piedecuesta in
Santander (c.60 km from our Alto Cantagallos site).
Cerro de las Armas is the only locality in the intervening region of suitable elevation but has not beensurveyed for birds (beyond brief observations inwhich no Scytalopus were recorded; Fundación
Natura 2003). Figure 10 suggests elevational connectivity of that site with localities where S. g.
griseicollis populations are found.

At Alto Cantagallos and Lepipuerto, S. g. gilesi washeard calling only in the ridgetop páramo andtreeline habitat in small bushes and shrubs; inmossy montane cloud forest below the treeline, it
was replaced sharply by S. latrans and S. spillmanni. Calling males of S. g. gilesi on the one
hand and S. latrans and S. spillmanni on the otherwere sometimes audible within 10 m of one anotherat the treeline, but never were found together in thesame habitat. Scytalopus g. gilesi was common in
the páramo of Filo Pamplona, but was not present
in the montane forest at La Aurora below, though it

CONSERVATION.- Although bird conservation assessments often consider only species level taxa
(e.g. BirdLife International 2004), distinctive subspecies such as S. g. gilesi are of conservation relevance (e.g. Peterson & Navarro-Sigüenza 1999;
Zink 2003). An assessment of the new taxon’s
conservation status therefore follows. The area ofoccurrence of S. g. gilesi is rather difficult to determine because it occurs principally in the ecotone
between páramo and forest, which does not correspond to any elevational band but rather varies inwidth and elevation with topography and on differ-

38

Taxonomy of Scytalopus tapaculos

Donegan &Avendaño-C.

Figure 9: Potential distribution map by J. Velázquez using MAXENT 3.0 (Phillips et al. 2006) showing locations of records of S.
griseicollis in the Eastern Andes Colombia, with potential distribution based on topography and climate layers available fromWorldclim (Hijmans et al. 2005). Localities in Serranía de los Yariguíes are north to south: Alto Cantagallos; Filo Pamplona andLepipuerto. Note the apparent isolation of the Yariguíes and Perijá populations; and lack of geographic barrier between “S. infasciatus” type locality and “S. griseicollis” sites.

Ornitología Colombiana No.6 (2008)
rent slopes. It is absent from the more windswept
páramo of highest elevations (which consists principally of stunted bromeliads and reeds) and fromforest below the treeline. Based on our estimate of
the area of páramo-forest ecotone at appropriate
elevations for the species (IGAC 1995) and forest
coverage information (IGAC 1999), there should beapproximately 12.6km2 of suitable habitat for S. g.
gilesi (7.5 and 5.1km2 on east and west slopes, respectively). It is abundant in such habitats, with anestimated density of 2.65 individuals/ha or 265 individuals/km2 based on data from our fieldwork.
Assuming 70-100% occupancy in suitable habitat
produces a total estimated population of ca. 23003300 individuals. Despite its small range, S. g.
gilesi would not qualify for Vulnerable under category D1 on a basis of its population alone, but
could be considered Near-Threatened. It technically qualifies as Vulnerable under D2 on accountof being recorded in less than three localities but
habitat is fairly continuous in the Yariguíes range,
meaning that there must be many other sites. The
relative intactness of the remaining forest at highelevations in the Yariguíes suggests that the population is unlikely to be declining by >30% over ten
years (criterion A), nor is the habitat is "severely
fragmented" (required to qualify for criterion B).
Deforestation continues in lowlands and foothill topremontane regions of the Yariguíes mountains, but
the highest elevations remain remote.
In light of the inaccessibility of S. g. gilesi’s habitat, its high abundance where found, lack of hunting or trade threats for Scytalopus and the adaptability of S. griseicollis to secondary habitats, we are
not particularly concerned for the long-term conservation of this taxon at present. We also note that
the Serranía de los Yariguíes National Park was
declared on 16 May 2005 (Ministerio del Medio
Ambiente, Desarrollo y Vivienda 2005), following
biological justification provided by our and otherstudies. NGOs have declared the region an Alliance for Zero Extinction site (Ricketts et al. 2005)
and an Important Bird Area (Franco & Bravo
2005), and Fundación ProAves has established anature reserve in the region (discussed further inDonegan & Huertas 2005; Huertas & Donegan2006). If these areas receive effective protection, it
will assure the survival of S. g. gilesi as well as anumber of other species endemic to the Eastern

39

Andes, since they include the largest remaining
block of forest in this EBA (Stattersfield et al.
1998).
Other northern populations of or related to
S. griseicollis
North-south variation in S. griseicollis is evident
and requires further research. Recent sound recordings (e.g. Fig. 1) and various specimens(Appendix 1; Fig. 9) confirm the presence of S.
griseicollis in Tamá National Park, on both Colombian and Venezuelan sides of the national borderand other localities in Santander and Norte deSantander departments. This population has a notably browner back in adult plumage than any of theother populations. S. g. griseicollis juveniles also
have a brown back, but of a rather different shade.
This undescribed population also has on average
higher frequency scolds (Level 1). Although no
obvious isolating barrier exists between these populations and those to the south (Fig. 9), several other
understory highland species include a diffe-rent
subspecies in the Tamá-Santander region (e.g. Atlapetes schistaceus, Henicorhina leucophrys). This
population will be described elsewhere with othercoauthors. In any event, this population is not S.
infasciatus because the infasciatus type does nothave a brown back.
Various specimens, most labelled “Scytalopus meridanus”, have been collected in the Serranía de
Perijá (Appendix 1; Figs. 9 & 10). Although
Krabbe & Schulenberg (2003) did not assign thispopulation to any described taxon, Hilty (2003) andSalaman et al. (2007) treated it provisionally as
related to S. meridanus. Plumage and biometrics
suggest strongly that this population is related toeither S. meridanus or S. griseicollis (Fig. 10). Although the Perijá population has been sound recorded (C. Sharpe in litt. 2007) and collected recently (Appendix 1), recordings were not available
for this study such that we cannot make strong conclusions about its status. Perijá birds cluster closer
with those from the Venezuelan Andes for biometrics (Fig. 8) and are differ from Eastern Andes
specimens up to Level 1 (and, for southern populations, Level 2) in bill and wing length; and from the
Yariguíes population in their shorter tail (Levels 1,
2, and 4). The Perijá population is isolated from

40

Taxonomy of Scytalopus tapaculos

nominate S. griseicollis populations and S. meridanus by the narrow, low-elevation section of the Andes in the Ocaña region which is a formidable barrier to high elevation fauna (Stattersfield et al.
1998). An undescribed Scytalopus taxon is clearly
involved.
VENEZUELAN ANDES: THE STATUS OF S.
MERIDANUS AND S. FUSCICAUDA
Recent authors recognise two light-plumaged Scytalopus in the Venezuelan Andes: Mérida Tapaculo
S. meridanus Hellmayr, 1922 is found across muchof the range; and Lara Tapaculo S. (griseicollis)
fuscicauda Hellmayr, 1922 is found in Lara stateand on the Trujillo state border (e.g. Krabbe &Schulenberg 1997, 2003; Hilty 2003). Scytalopus
fuscicauda is generally described as a high elevation bird with at most only traces of barring on its
lower underparts (Hellmayr 1922, Zimmer 1939,
Krabbe & Schulenberg 2003, Hilty 2003). S. meridanus has been considered a species with morestrongly barred lower underparts, with higher elevation records having been doubted and some texts
illustrating a rather dark-plumaged bird (Fjeldså &Krabbe 1990, Krabbe & Schulenberg 2003, Hilty
2003). Krabbe & Schulenberg (2003) ranked fuscicauda as a species, concluding that it was unlikely
to be related to S. griseicollis. However, they notedthat S. fuscicauda might be synonymous with S.
meridanus and recommended further studies. Remsen et al. (2008) followed this approach and also
treated S. fuscicauda as a species. BirdLife International (2007) recently declined to recognise S.
fuscicauda, based on some of the data presented
herein. Our analysis of specimens suggests that
two species or distinctive color morphs exist within
what is currently referred to as “S. meridanus” inthe Venezuelan Andes (see below). However, as
suspected by Krabbe & Schulenberg (2003), we
will argue that S. meridanus and S. fuscicauda
should be treated as conspecific.
The type of S. meridanus (AMNH-492377) is agray Scytalopus with a strongly barred orangerufous vent. The specimen label states that it wascollected at 4000m elevation at La Culata in
Mérida state (08°45'N; 71°05’W, per Paynter
1982). Another S. meridanus specimen from thislocality is labelled as having been collected at

Donegan &Avendaño-C.

3000m (Appendix 1). The S. fuscicauda type wascollected by Carriker at Páramo de Rosas, close to
the Trujillo state border in Lara state (09°35'N, 70°
07'W, 3245 m, per Paynter 1982). It differs fromthe S. meridanus type in lacking strong barring on
the vent (Hellymar 1922; Fjeldså & Krabbe 1990;
Hilty 2003), although Zimmer (1939) discerned
weak barring not visible from the photographs we
inspected. Hellmayr (1922) provisionally included
in S. meridanus some specimens from Santa Marta,
southern Colombia and Peru whilst noting that suchspecimens might refer to other taxa (as is now
known to be the case), as well as at least one specimen from SW Lara state (Anzoátegui, Lara: 09°
36'N 69°54'W, per Paynter 1982). One BMNH
skin (1914.11.26.507) labelled "S. fuscicauda", indistinguishable from others labelled as this form at
BMNH, lacks strong barring and was collected at
La Culata, Mérida, the type locality of S. meridanus. Zimmer (1939) also included a specimen from
Guamito, Trujillo, within S. fuscicauda.
Individual morphological variation within the S.
meridanus series at COP and elsewhere is remarkable, and the S. fuscicauda type falls within therange of plumage in this series. Specimens withstrong barring on the vent have been collected in
Táchira, Mérida, and Trujillo states and over a wide
range of elevations (2200-3300 m). Specimens
without strong barring are less frequent in collections but are found in series from Mérida, Trujillo,
and Lara (S. fuscicauda type) states, and over asimilar range of elevations (2400-3300 m). Birds
with intermediate barring are also found in allstates, with no discontinuity in this plumage featureor clear sex differences. Strongly barred and lessstrongly barred or unbarred birds thus do not separate out according to sex, elevation, or distribution.
Individuals from different states and elevations in
Venezuela similarly show no diagnostic differencesin biometrics. We were only able to inspect one
specimen from Lara state, which falls within therange of measurements of S. meridanus in biometrics. We tested Zimmer (1939)’s tentative hypothesis of bill size differences between his five
“fuscicauda” and other S. meridanus, but we foundno significant (Level 1) differences in any measurement (d.f.=7), with sample means of measurementsof Lara and Trujillo specimens differing from thoseof Mérida specimens by <0.5 mm.

Ornitología Colombiana No.6 (2008)
S. meridanus has an unusually wide elevational
range, with skins from 2200m to 4000m and observations in PN Yacambú in Lara state down to 1600m (P. Boesman in litt. 2006). Such an elevational
range is similar to that of S. griseicollis in the Eastern Andes – which is present from 2450 m in forest(Yariguíes) or 1300 m in secondary growth (recent
sound recording by A. Hernández-J. in IBA La
Judía, Santander) up to 3900m at Laguna de Chisacal, Cundinamarca. We segregated Mérida range
biometric data at arbitrary mid-elevation points totest the hypothesis that fuscicauda and meridanus
might be elevational replacements, but found no
Level 1 or greater differences. It is unlikely that
two morphometrically and vocally indistinguishable Scytalopus species would co-occur across sucha broad elevational and geographical range: indeed,
Krabbe & Schulenberg (1997) found sharp elevational or habitat replacements to be frequent in thisgenus. The only case of which we are aware oftrue sympatry in the genus in Colombia is between
S. spillmanni and S. latrans, which are two species
with rather different plumage, biometrics, and vocalizations.
For voice, as for S. griseicollis and S. infasciatus,
we inspected various recordings and assigned themto either of “S. fuscicauda” or “S. meridanus”
based on proximity to the relevant type locality.
Recordings in Boesman (1999) suggest that most of
the vocalizations given in Táchira, Mérida, Trujillo,
and Lara states are indistinguishable, including thescold (represented by spectrograms 68/71 in
Krabbe & Schulenberg 1997) and some call notes(e.g. spectrograms 69/72). Reeling songs, similar
to spectrogram 70, have also been recorded
throughout the Venezuelan Andes. There is considerable intrapopulation variation in song speed,with calls often including short gaps between
groups of two, three, or more notes (cf. some callsof S. parkeri), rather than in long, uninterrupted
sequences (cf. S. griseicollis and S. spillmanni). In
Táchira, Mérida, and Trujillo states, reeling songs
often have introductory notes which are longer induration than notes comprising the reeling song,
though in one recording there is no such introduction. The two recordings we have inspected ofreeling songs from Lara state (presumably “S. fuscicauda”) have shorter introductory notes (noted by
P. Boesman in litt. 2006). However, considerable

41

individual variation in introductory phrases to reeling songs is evident within S. griseicollis and S.
spillmanni. A larger sample of S. meridanus and S.
fuscicauda recordings would be needed to assess
whether these differences are due to individual or
geographical variation. Even if such differences inintroductory notes were to be confirmed with agreater sample, variables of song speed and acoustic frequency overlap considerably between westernand eastern recordings for reeling songs, not reaching Level 1 diagnosis, let alone Level 4/5 thatwould be required for species rank.
In conclusion, no morphometric, biogeographic,plumage, or vocal data support the treatment of S.
fuscicauda as a species. Further, such a treatment
should not be regarded as a "status quo" (contra
e.g. Remsen et al. 2008) given that S. fuscicauda
was lumped with either S. magellanicus or S.
griseicollis until 2003, including by Hilty (2003) inthe leading field guide for the region. Whilst we
agree with Krabbe & Schulenberg (2003) that S.
fuscicauda is not conspecific with S. griseicollis,
the most conservative approach at present would beto treat it as a subspecies of S. meridanus. We suspect that the two taxa are synonyms but we decline
to go so far, pending analysis of a greater sample ofvocalizations from Lara state and other regions.

Scytalopus fuscicauda and S. meridanus were described contemporaneously by Hellmayr (1922).
Neither therefore has priority in the event that they
are treated as conspecific (International Code for
Zoological Nomenclature, Article 23). Pursuant to
ICZN Article 24A, it is recommended that thename, spelling or nomenclatural act that will best
serve stability and universality of nomenclature
should be adopted. Universality considerations
could be thought to include “position precedence”
or “line priority” (under ICZN Article 69A.10,
which applies to the fixation of type species forgenera, a broadly analogous situation, all otherthings being equal). Such an approach would favour use of the species name that appears first in
Hellmayr (1922) which is S. fuscicauda. However,
all other things are not equal here. Because meridanus is the name which has been assigned to Scytalopus occurring in most of the Venezuelan Andesin three recent landmark publications (Krabbe &42Taxonomy of Scytalopus tapaculos

Donegan &Avendaño-C.

Taxonomic rank of Scytalopus meridanus and
S. griseicollis
Due to morphological similarities, all populationsof gray tapaculos occurring in the northern Colombian and Venezuelan Andes have formerly beentreated as conspecific (Zimmer 1939). The scolds
and reeling songs of S. griseicollis taxa and S. meridanus taxa (as redefined above) are diagnosable toLevels 1 through 5 (Appendix 4), whilst note shapeis different (Fig. 2). Scytalopus meridanus reelingsongs become slower over time, being delivered ingroups of 2-7 notes with spaces between them (vs.
uniform delivery, with gradual slowing over time
but no gaps, in S. griseicollis). Introductory notes
are of a different note shape to the “zz” sometimes
found in S. griseicollis recordings. Vocal differences between S. griseicollis and S. meridanus exceed those between sympatric S. griseicollis and S.
spillmanni, requiring species rank for each of themunder the Helbig et al. (2002) and Remsen (2005)
guidelines.
THE POPULATION OF SCYTALOPUS
SPILLMANNI IN THE EASTERN ANDES OF
COLOMBIA

Figure 10: Photographs showing various populations labelled"S. meridanus" from Venezuela. From left to right: (i) Scytalopus sp. of Perijá: COP 72580, Perijá, Zulia, Venezuela,
3000m, 6 July 1974; male; (ii) S. griseicollis undescribedsubsp. COP 11101, collected at Páramo de Tamá, Venezuela(Colombian border), 2800m on 27 February 1941; sex not
determined; (iii) S. meridanus COP 49296 La Honda, Santo
Domingo, Mérida, Venezuela, 6 December 1949; male; (iv)apparently undescribed Scytalopus species COP 65395, La
Azulita, Mérida, 2300m, 25 November 1959; male. Note themorphological similarities of the first three skins. Birds represented by the second and third skins have strikingly different
voices from one another. Photographs at COP by B. Huertas.

Schulenberg 1997 and 2003 and Hilty 2003), we
formally propose that meridanus be given precedence over fuscicauda in the event that the two are
considered conspecific or synonymous (for the purposes of ICZN Article 24.2).

As discussed in Donegan et al. (2007), Spillmann’s
Tapaculo S. spillmanni was recently found in the
Eastern Andes for the first time. The populationshows differences from other populations in darker
plumage (Fig. 11), tarsus length, song speed andacoustic frequency. However, it does not meet therequirements of all modern subspecies concepts. A
preliminary molecular study of various S. spillmanni specimens including the Eastern Andes
population is being undertaken, but results were notavailable for this paper. The Eastern Andes population is described and discussed below but is notformally named.
Localities for the Eastern Andes population of S.
spillmanni and its potential range are shown in Figure 12; a model of potential distribution constructedusing MAXENT suggest this population may beisolated from more southern and western populations due to the low elevations of the Eastern Andesridge line in the Serranía de los Picachos region or
further south. Specimens we ascribe to this population on the basis of morphology include two recent

Ornitología Colombiana No.6 (2008)

43

Figure 11. Dorsal and ventral photographs of (left to right) two S. spillmanni Eastern Andes population (ICN 35605 (male) and
35608 (female) (both collected by the authors, Finca Pamplona/La Aurora, vereda San Isidro, municipality of Galán, Serranía delos Yariguíes, Santander, Colombia, 06º38’N; 73º24’W, 2700 m); one S. spillmanni Central Andes: ICN-33166 (male, Risaralda,
Reserva Ucumarí, la Pastora, 2450 m, collected by J.A. Mobley on 5 January 1996); five S. spillmanni Western Andes: ICN35034 (male, Antioquia, Jardín, Vda. Meseria, 05°29'N, 75°54W, 2200 m, collected by N. Krabbe on 23 August 2004), ICN35033 (female, as previous but 2300 m, 22 August 2004), ICN-35032 (male, as previous); ICN-35029 (Antioquia, Urrao, Páramo
Frontino, 06°25'N, 70°04'W, 2600 m, collected by N. Krabbe on 16 August 2004), ICN-35026 (as previous, 3150 m, 14 August2004). Photograph at ICN by TMD.

44

Taxonomy of Scytalopus tapaculos

Yariguíes specimens we collected, two MLS specimens labelled “S. latebricola meridanus” (Fig. 11)
from near Pamplona, Norte de Santander, and a
BMNH “Bogotá” skin labelled “Scytalopus sylvestris” (see Appendix 1).
Various Scytalopus species including S. spillmanni
were previously treated as conspecific with S. latebricola (Zimmer 1939), but S. latebricola is now
considered a monotypic endemic of the Santa
Marta mountains of Colombia (Krabbe & Schulenberg 1997), with S. meridanus endemic to the
Venezuelan Andes (see above). The type locality
of “S. sylvestris” (now not considered a valid
taxon) is Pallaypampa, Peru (Chapman 1915), so
the BMNH specimen appears to have been misidentified. By biometrics, the MLS and BMNH
specimens all fall within the range of the Eastern
Andes S. spillmanni population. In Donegan et al.
(2007), the MLS and BMNH skins mentioned
above were thought possibly to relate to S. rodriguezi, but JEAC recently collected several specimensof the Yariguíes population that is related to S. rodriguezi, which has enabled comparison and rejection of this hypothesis. Two other MLS skins from
Pamplona labelled “S. latebricola meridanus” areof the undescribed Tamá subspecies of S.
griseicollis.
Individuals of the Eastern Andes S. spillmanni
population were observed, sound recorded, or captured at three montane forest sites in Serranía de los
Yariguíes: La Aurora, Lepipuerto, and Alto Cantagallos (details above). Other sound recordings
were made at Finca La Carbonera, Los Robles, La
Aguadita, Fusagasugá, Cundinamarca (4°25’N, 74°
19’W, 2450-2550m) by O. Laverde and (probably,
although, as it is a song, we cannot exclude the possibility that the recording is not an unusual recording of S. griseicollis) at Vereda Carrizal,
Sisavita, Cucutilla, Norte de Santander by S. Córdoba-C. (IAVH BSA 8887, 07°26’20N, 72°
50’27W, 2380m). Birds tentatively identified as S.
spillmanni have been reported based on field observations at Rogitama, Boyacá (05°47’N, 73°31’W)
and Santuario de Fauna y Flora Iguaque, Boyacá(05º40’N, 73º27’W) (J. Zuluaga in litt. 2006). No
other skins of the Eastern Andes S. spillmanni
population were found in any of the collections inspected nor are any observations reported by the

Donegan &Avendaño-C.

members of the network of Colombian bird observers (RNOACOL).
DIFFERENCES FROM OTHER POPULATIONS.– The
Eastern Andes population is clearly referable to S.
spillmanni due to its broadly similar morphology
and voice. Although both Yariguíes specimens arein molt and lack various tail feathers, the positionof the feathers is consistent with the birds each having 12 or more rectrices, which is typical for S.
spillmanni, whereas other Scytalopus of the Eastern
Andes have only 10 rectrices (Krabbe & Schulenberg 1997). The type of S. spillmanni is from Volcán Illiniza, on the Pichincha/Cotopaxi border on
the western slope of Ecuador (Stresemann 1937)
and shares broadly similar dark plumage to theEastern Andes population.
The Eastern Andes population differs from populations in Ecuador in its shorter tarsus length (Levels
1 & 2) and from populations in Eastern Ecuador by
its slower song speed (Level 1; marginally missing
Level 2). Due to slowing of song speed over timein longer calls, some S. spillmanni recordings attainslower speeds not reflected in average song speed
data: the above analysis is based on average song
speeds. Reeling songs from Western Ecuador are
higher on average in acoustic frequency than in
Eastern Ecuador (Level 1), with the Eastern Andes
population intermediate (Level 1 from both). No
recordings of the Eastern Andes population inspected include elaborate introductory sequences tosongs nor are any composed of downstrokes. Suchfeatures are also rare or reduced in Central Andes
recordings, but both are found in western and eastern slope S. spillmanni populations in Ecuador, including downstrokes in 80% of flat reeling songs.
The Eastern Andes population appears smallbodied compared to Ecuadorian populations (butYariguíes specimens had appreciable subcutaneousfat). A comparison of photographs suggests that
the Ecuadorian and Eastern Andes populations
might not be diagnosable by plumage, but directcomparisons of series have not been possible.
The Eastern Andes population can be distinguishedfrom populations of S. spillmanni in the Central
Andes by the lower acoustic frequency of its reeling songs (Levels 1 & 2: Figs. 4 & 5) and scolds(Levels 1 & 2: Fig. 3). Although insufficient data

Ornitología Colombiana No.6 (2008)

45

Figure 12. Potential distribution map by J. Velázquez using MAXENT 3.0 (Phillips et al. 2006) showing locations of records ofS. spillmanni in the Eastern Andes Colombia, with potential distribution based on topography and climate layers available fromWorldclim (Hijmans et al. 2005).

46

Taxonomy of Scytalopus tapaculos

are available to assess biometrics, the two specimens we have measured of the Central Andes
population are indistinguishable from Ecuadorian
populations for tarsus length, tail length and mass,
but fall outside the range of the Eastern Andes
population. We have not considered in detail thestatus of Western Andes populations assigned to S.
spillmanni, for which vocal differences from otherS. spillmanni populations have been asserted(Krabbe et al. 2006). Our Yariguíes specimens aredarker on the underparts and upperparts than recent
Central and Western Andean specimens at ICN,
although an old BMNH Central Andes skin is indistinguishable from the BMNH Eastern Andesskin.
An undescribed but well-known Scytalopus population in S Ecuador and Peru (“S. cf. canus opacus”)
has similar scolds to those of S. spillmanni. Suchcalls are diagnosably different from Ecuadorian
populations of S. spillmanni but are not fully diagnosable from the Eastern Andes population (Fig. 3),
leading to Donegan & Avendaño-C. (2006) erroneously referring to the population as “Scytalopus sp
nov (canus)”. Although scolds are very similar, theflat reeling songs of “S. cf. canus opacus” have a
different note shape (essentially, a thick horizontalline), are of higher acoustic frequency than otherpopulations and apparently have a different ratio
between note duration and gap duration, whereas
the female “brzk” call of “S. cf. canus opacus”
bears little resemblance to that of S. spillmanni.
Further research into this population and its affinities with S. spillmanni and true S. canus opacus iswarranted. However, as this population differs diagnosably in its song from S. spillmanni, species
rank with respect to S. spillmanni is not an issue.
The similarity of scolds of S. cf. canus opacus andthe Eastern Andes population of S. spillmanni begs
further investigation.
It seems unlikely that the Eastern Andes population
represents a point on a broad north-south cline ofslowing song speed from populations in Ecuador.
Comparison of vocal data between the 250 km distant Cundinamarca and Norte de Santander recordings of the Eastern Andes population (d.f.=4);and separately between Napo and Sucumbíos department populations on the east slope in Ecuador(d.f.=5) shows no statistically significant differ-

Donegan &Avendaño-C.

ences in song speed, with relatively high t (>0.5)
values returned in each case and averages <0.7
notes/s different. The Tamá recording, if confirmed to be of S. spillmanni, would fall within therange of other Eastern Andes recordings for measured variables but specimens have lighter plumage
than Yariguíes birds. East slope Ecuador recordings were slightly faster on average than west
slope Ecuador recordings (but not to Level 1),
meaning that the Eastern Andes population was
marginally more distinct vocally from the geographically closer population (App. 4). Slower
song speed appears to be a feature of populations in
all three Colombian cordilleras, with the few Western Andes recordings we have inspected also having slower songs (as noted by Krabbe et al. 2006) .
Though we lack vocal data from Nariño to Caquetá,
it would be inconsistent with variation elsewhere
for a broad cline to occur in this region. The single
Central Andes Huila recording in our sample is acurtailed rising reeling song at 29.5 notes/s, suggesting that it is related to the Central Andes population (slower than 97% of the Ecuador population’s songs; and curtailed songs typically are fasterthan elongated songs due to a lack of slowing overtime more pronounced in longer songs). The differentiation of a slower (vs. Ecuador) and higher(vs. Central Andes) song in the Eastern Andes isnotable given sympatry in this region with S. griseicollis, a species with a similar but faster-paced andlower frequency song. No such similarly-calling,
apparently closely related, species are sympatricwith S. spillmanni elsewhere in the northern Andes.
Acoustic frequency appears to be more plastic thansong speed in these Scytalopus. Variation in acoustic frequency between S. griseicollis populations is
also evident. It is intriguing that the Eastern Andes
population has higher frequency calls and the Central Andes population lower frequency calls, representing the reverse west-east pattern to that observed in Ecuador and again possibly influenced by
S. griseicollis.
NOTES ON OTHER NORTHERNRHINOCRYPTIDAE
Our studies in the field and of skins reveal the presence of possibly two undescribed Scytalopus atlower montane elevations (around 2000 m) of Colombia's Eastern Andes and the Venezuelan Andes,

Ornitología Colombiana No.6 (2008)
all specimens of which have been labelled "S. meridanus" or "S. femoralis" in museums (Donegan etal. 2007). One such population is found at around2000 m elevation in Serranía de los Yariguíes (seeDonegan et al. 2007 and specimens listed in Appendix 1).
This population is apparently not
closely related to either S. meridanus or S. griseicollis, having darker plumage and a different voice.
It has a longer tail (Levels 1,2 and 4, d.f.=2) than S.
rodriguezi. Separately, four COP specimens collected at 2200-2400m elevation in the Venezuelan
Andes (listed in Appendix 1) have a darker shadeof gray plumage overall, a darker and brownershade of rufous on the underparts (which are allstrongly barred: Fig. 11, plate of “S. meridanus” in
Hilty 2003), and a longer tail than S. meridanus(Levels 1 & 2, d.f.=2). These specimens appear
likely to be of an undescribed color morph or species (possibly related to S. spillmanni, S. micropterus or S. rodriguezi). These populations will beconsidered in further detail in future publications.
Blackish Tapaculo S. latrans is another widespread
species of the northern Andes. Our specimen from
La Aurora and others observed at the same site (butnot others in the Yariguíes) had a pinkish white billbase, a feature not previously observed for the species and perhaps a local leucism. Central Andes
specimens of S. latrans, including live and recently-collected specimens, are not as dark as West andEastern Andes specimens. Although songs andsome calls are similar to those of other populations,
our recording of the “brzk” call of S. latrans in
Serranía de los Yariguíes shows small differencesfrom populations in Ecuador. Krabbe & Schulenberg (1997) reported differences in plumage between west and east slope populations of this species in Ecuador and recordings in Krabbe & Nilsson (2004) evidence geographic variation in voicein Ecuador. Intraspecific variation in S. latrans is amatter in great need of further research.
Other Scytalopus taxa present in the Eastern Andes
include Long-tailed Tapaculo S. micropterus,
which is present apparently on the east slope(Salaman et al. 2002) and replaced by S. rodriguezi
at similar elevations on the west slope. The foothill
species White-crowned Tapaculo S. atratus is alsopresent on both slopes of the Eastern Andes(Donegan et al. 2007; Cuervo et al. 2007) and in47

the Venezuelan Andes. Finally. S. caracae is present in the Coastal Cordillera of Venezuela. A recent Scytalopus record in the Caripe mountains further to the east (Hilty 2003) requires urgent attention in light of the status of the region as a center ofavian endemism and high threat levels (Stattersfield
et al. 1998).
The monotypic Ash-colored Tapaculo Myornis senilis occurs at high elevation sites (mostly páramos)
in Ecuador and all three Andean ranges of Colombia. Some variation in darkness of plumage andbiometrics is evident among Colombian populations. As with S. spillmanni, Central Andes birds
are similar morphometrically to those in Ecuador;whilst East Andes and West Andes birds show
small differences. Whether such variation is of asimilar nature to that in S. spillmanni or requires
the description of new taxa is a matter requiring
further investigation.

DISCUSSION
Allopatric populations in the process of speciation,
which have reached a certain level of differentiation, are cases where subspecies rank is likely to bean appropriate taxonomic treatment. The recognition of isolated populations such as the East Andes
population of S. spillmanni as subspecies would
facilitate communication and draw attention to thedifferentiation of the population. On the other
hand, populations showing only small average differences should not be recognized as subspecies.
The various proposed subspecies tests consideredin this paper each seek to balance these concerns.
Several antbird taxa have been ranked as valid subspecies on a basis of qualitatively diagnostic plumage characters but show only non-diagnostic, vocal
differences from proposed conspecifics (Isler et al.
1998, 1999, 2006, 2007). For Scytalopus, plumage
and biometrics are often not diagnostic even among
species (Krabbe & Schulenberg 1997), and a diagnostic difference in a single vocal variable is regarded herein as sufficient for species rank. Various tapaculo species have allopatric populations
that show some geographic variation in voice but
no obvious plumage differences (e.g. S. latrans:
Krabbe & Nilsson 2004; S. atratus: Cuervo et al.

48

Taxonomy of Scytalopus tapaculos

2007; Donegan et al. 2007; S. spillmanni: thisstudy). “Relatively minor or few diagnostic characters” (per the Isler et al. model for antbirds) may
therefore be an illusive criterion for the ranking ofsubspecies within some Scytalopus species. S.
griseicollis gilesi and the undescribed Tamá subspecies of S. griseicollis are rare cases of Scytalopus populations that have diagnostic plumage
characters but which are not diagnosable by voiceor biometrics. The Eastern Andes population of S.
spillmanni shows similar differentiation of vocal
and biometric characters as S. griseicollis subspecies, suggesting similar isolation, but is not capableof formal description under our conservative approach owing to the lack of a single diagnostic(plumage) character.
Our Level 1 test of statistical significance tolerates
a considerable degree of overlap, meaning that, if itwere used as a subspecies test, the placement of aparticular individual with one taxon or another may
be difficult; and “subspecies” may involve narrower distinctions, risking a destabilizing taxonomic inflation. Our Levels 1 and 2 involve agreater level of differentiation and reflect variationin vocal and biometric variables among ‘good’ S.
griseicollis subspecies. However, the Hubbs &Perlmutter (1942) test is rarely used today in favorof the Amadon (1948) test, raising universality issues and risking taxonomic inflation.
The traditional Level 3 “99+%/75%” test for subspecies (Amadon 1948; Patten & Unitt 2002;
Cicero & Johnson 2006) is an arbitrary one and has
been criticized (e.g. Zink 2003). As noted by
Amadon (1948), the 99+%/75% test is “roughly
equivalent” to 97%/97% diagnosability – which (ifrepeated for multiple characters) is essentially the
benchmark of current tests of species rank under
biological species concepts (e.g. Isler et al. 1998).
Under phylogenetic species concepts, two allopatric phylogenetic species would be recognized
when, for a continuous variable with infinite sample size, sample means are 4.00 standard deviations
apart (Level 4/5). The assessment of a subspecies
where sample means are between 3.92 (=t99.9%@ ∞ +t75%@ ∞) and 4.00 standard deviations (=2t97.5% @ ∞)apart would result in very few populations (fallingwithin a range of 0.08 standard deviations’ difference for a continuous variable with infinite sample

Donegan &Avendaño-C.

size) being ranked as subspecies under some approaches.
A separate difficulty arises where
smaller samples are involved, which is a hazard forcomparative studies of some Neotropical bird
populations. Where t values are used rather thanrecorded standard deviations, t99.9% becomes increasingly large compared to t97.5% with smaller
samples. Where d.f.<8, Isler et al. (1998)’s diagnosability test for species (Level 5) may be met
where the traditional subspecies test (Level 3) isnot. Where 99%/75% diagnosability is used (ashere), rather than 99.9%/75%, the availability ofsubspecies rank includes cases where sample
means fall between 3.00 (rather than 3.92) and 4
standard deviations apart at infinity and the scopefor ‘species not being subspecies’ is restricted toeven smaller samples (d.f.<4), giving the conceptof “subspecies” more meaning.
The East Andes population of S. spillmanni meetsthe Luckow (1995) “Level 1” test; and the Hubbs &Perlmutter (1942) Level 2 test for subspecies but
fails all others based on our current data set: insummary, observed differences in voice and biometrics are statistically significant and sample
means of at least one variable fall between 2 and 3
standard deviations apart (controlling for samplesize). This is not sufficient for ranking of a subspecies under modern concepts. Biometrics of S. spillmanni populations should be re-evaluated once agreater sample size is available and we await theresults of molecular studies with interest.

ACKNOWLEDGMENTS
Special thanks are due to Blanca Huertas who obtained funding for and directed the YARE Project,organized much of the logistics for the EBA Project
research, obtained various bibliographic materials
and took some of the photographs used in our research and to Elkin Briceño, who accompanied
TMD on bird fieldwork at Alto Cantagallos. John
Jairo Arias, Martin Donegan, Laura Rosado, Diana
Villanueva, Diana Montealegre, Cristobal Ríos andguides José Pinto, Hernando Figueroa, Alonso
Masías and Fabio Aleán accompanied us in fieldwork at sites where we recorded the new subspecies. Nick Athanas, Peter Boesman, Jurgen Beckers, Diego Calderón, Roberto Chavarro, Sergio

Ornitología Colombiana No.6 (2008)
Córdoba-Córdoba, Oswaldo Cortés, Walter Halfwerk, Alejandro Hernández, Don Jones, Doug
Knapp, Oscar Laverde, Jorge Parra, Chris Parrish,
Andrew Spencer, Luis Eduardo Urueña and Johana
Zuluaga each kindly provided sound recordings(some through IAVH and xeno-canto); and P. Pulgarín, photographs. Niels Krabbe and Tom
Schulenberg made available their biometric data on
S. spillmanni from Ecuador. We further acknowledge the substantial work of Niels Krabbe in soundrecording Scytalopus species in Ecuador; and Oscar
Laverde in Colombia. Mauricio Álvarez and David
Mejía shared IAVH’s database of Scytalopus recordings. Jorge Velásquez kindly collaborated in
producing the distribution maps. V. Heinrich, Nils
Hoff and Sylke Franhert (MFNU), Nathan Rice(ANSP), Guillermo Ramirez (MLS), Mary Hennen(FMNH), Peter Capainolo, Margaret Hart & Paul
Sweet (AMNH), Steve Rogers & Mindy
McNaugher (CM) and Jeremiah Trimble (MCZ)
provided us with photographs of specimens. C.
Daniel Cadena sequenced various specimens andwill publish results elsewhere. We are grateful toPeter Boesman, Mort Isler, José Gregorio Moreno Patiño, J.Van Remsen, Paul Salaman, F. Gary
Stiles and various anonymous reviewers for theircomments on earlier versions of this manuscript orparts thereof. Jean François Voisin provided information about the types of S. griseicollis. Paul Salaman and Juan Carlos Verhelst provided specimen
data from Biomap Project. Miguel Lentino provided specimen data from the COP database. We
are grateful to Robert Prŷs-Jones, Mark Adams andDouglas Russell (BMNH), Enrique Castillo, Fernando Forero, Diego Perico and Socorro Sierra(IAVH), José Gregorio Moreno Patiño (UIS), Hno.
Roque Casallas and Arturo Rodríguez (MLS), Jean
François Voisin and Claire Voisin (MNHN), F.
Gary Stiles (ICN), Miguel Lentino (COP) and eachof the institutions involved for access to specimens.
CAS (resolution 832 of 2004) and the mayoralties
of San Vicente de Chucurí, Galán and El Carmen
each provided permissions for fieldwork. The EBA
expeditions to Serranía de los Yariguíes were madepossible through generous financial support of theRoyal Geographical Society (with Rio Tinto plc),
Duke of Edinburgh, Fondo para Acción Ambiental,
Fundación Omacha, Conservation International
Colombia (Becas Iniciativa de Especies
Amenazadas – Jorge Ignacio “El Mono” Hernández

49-Camacho), the Percy Sladen Memorial Fund
(Linnean Society) and ProAves Foundation.
YARÉ Project took place with the support of the
BP Conservation Programme (BirdLife International, Conservation International, Flora and Fauna
International, Wildlife Conservation Society),
Game Conservancy Trust, Carter Ecological, Tropical Andean Butterfly Diversity Project, ProAves
Foundation, World Pheasant Association, Carter
Ecological, Universidad Industrial de Santander,
Universidad de Caldas, Universidad de Tolima and
Gobernación de Santander. Idea Wild and The Explorers Club’s Youth Activity Fund assisted withequipment and financial support for JEAC for fieldwork in Yariguíes and CDMB at Suratá and AsoDiviso at Piedecuesta.

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50

Taxonomy of Scytalopus tapaculos

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Recibido: 25 mayo 2007
Aceptado: 18 febrero 2008

Ornitología Colombiana No.6 (2008)

53

APPENDIX 1: SPECIMENS EXAMINED All specimen localities are in Colombia unless otherwise stated.

Myornis senilis Ecuador: AMNH 186368, 492365,
173001, 492364, 124376, 180947, 180948, 184748,
39575 (all, Ecuador, no details taken); BMNH
1916.8.24.58 (Baeza, E. Ecuador), 1916.8.24.60,
1916.8.24.69 (both, “E. Ecuador”), 1940.12.5.1005,
1940.12.5.1006 (both, Volcan Pichincha, Ecuador).
Western Andes: ICN 35024, 35027 (both, Frontino,
Urrao, Antioquia). Central Andes: AMNH 11851(Laguneta, Quindio); ICN 26257 (Páez, PNN Nevadodel Huila, Cauca). Eastern Andes: ANMH 121829(El Peñon, Cundinamarca); AMNH 492366, 492367(both, “Colombia”); BMNH 86.6.24.515,
86.6.24.516, 89.9.20.687, 89.9.20.688, 89.9-10.972,
89.9-10.974 (all “Bogotá”); ICN 12002-12006 (allSan Miguel, Cundinamarca); MLS 3984 (PNN Chingaza, Boyacá), 3986 (San Miguel, Sibaté, Cundinamarca).
Scytalopus latrans: BMNH 89.9.20.684 to 89.9.20.686(all, Santa Elena, Antioquia), 89.9.20.702, 89.710.969, 89.7-10.970 (all Santa Elena or Medellín,
Antioquia), 89.7-10.970 (“Colombia”) 89.7-10.972(“Bogotá”), 1916.8.24.61, 1921.12.29.179 to
1921.12.29.181 (all, Munchique or Popayán, Cauca),1940.12.5.1014 to 1940.12.5.1016 (all, Ecuador);
COP (all Venezuela) 9088 and 9382 (both, Queniquea, Táchira, 1900m and 1600m), 10911 (Páramo
de Tamá, Villa Páez, Táchira, 2200m), 24554 (Bocade Monte; Pregonero, Táchira, 1950-2000m), 61231
to 61234, 62232 to 62235, 62238 to 62243 (Río Chiquito, Hacienda La Providencia, Táchira, 18002250m); IAVH 2476 (Cauca), 6723 (Quebrada Bomobona, Finca Indostán, Anzoategui, Tolima), 8220
(Santa Leticia, km 129, Cauca), 8221 (San Rafael,
PNN Puracé, Cauca), 10257 (PNN Los Picachos, San
Vicente del Cagüán, Caquetá), 10654 (Sector Orocué, PNN Tamá, Herrán, Norte de Santander), 11682(Bojacá, Cundinamarca); ICN 3500 (Alto de Ventanas, Jardín, Antioquia, 2900m), 4763 to 4767, 4846,
11992, 11998, 11999, 12000, 12968 (all Aguabonita,
Silvania, Cundinamarca), 11989, 11990, 11993 to
11997, 11999 (all, San Miguel, Cundinamarca,
2800m), 22186 (vereda Farralorado, Choachí, Cundinamarca, 2300m), 25906, 26032, 26048, 26074 (all:Corea, PNN Farallones de Cali, Valle del Cauca),
30933, 30934 (both, 3km NE of Monterredondo,
Camino a San Juanito, Güayabetal, Cundinamarca,
2050m), 31788 (Cuchillas de Santa Bárbara, Bojacá,Cundinamarca, 2600m), 33460, 33477 (both, Torre
Telecom, Serranía de los Churumbelos, Cauca,
2400m), 34704 (El Escobero, San Sebastian, Retiro,

Antioquia), 35023 (Frontino, Urrao, Antioquia,
3159m), 35607 (La Aurora, Galán, Serranía de los
Yariguíes, 2750m; sound recording), 36177 (El Talismán, Serranía de los Yariguíes, Santander,
2100m), unnumbered (2) (La Lana, San Pedro de losMilagros, Antioquia, Colombia, 2700 m; sound recording); MLS 2977 (Alturas de Medellín, Antioquia), 3978 to 3981, 3983 (all, Pamplona, Norte deSantander), 3982 (Tequendama, Cundinamarca),
4798 (Yarumal, Antioquia), 8070 (Boquerón, Medellín, Antioquia), 8583 (Medellín, Antioquia);
MNHN 522, 523, 1334, 1996 (all, “Ecuador”), 1336
(Pichincha, Ecuador), 1387 (Cerro Majanda, Ecuador), 2003.885 (“Colombia”); AMNH series inspected, no details taken.
S. micropterus: BMNH 89.9-10.983 (“Ecuador”),
1940.12.5.343 (Cututucu, Macas, Ecuador),
1953.68.679 (Soldados Range, Azuay, Ecuador),
1953.68.879 (Macuina, Ecuador); ICN 33263, 33267(both, Villa Igüana, Serranía de los Churumbelos,
Cauca); MNHN 1391 (E. Ecuador), 1933
(Alucincho, Ecuador); AMNH series inspected, nodata taken.
S. atratus (includes specimens labelled “confusus” and“nigricans”): BMNH 89.9.20.699 (“Bogotá”); COP(all Venezuelan) 54928 and 54929 (Sierra de Perijá,
Cerro Pejochaina, Falda Oeste, Zulia, 1900-1950m),
61224 to 62237 (Río Chiquito, Hacienda La Providencia, Táchira, 1800-1900m), 60640A to 60640L(Burguam, Cumbre, Cerro El Teteo, Táchira, 12001300m); ICN 32621 (vereda Aguas Claras, Serraníade Aguas Claras, Cubarral, Meta), 34387 (Anorí,
Antioquia), 35613 (Alto Honduras, Serranía de los
Yariguíes, Santander, 1600m; sound recording);
MNHN 386 (no data); AMNH series inspected, nodata taken.
S. panamensis: BMNH 1921.7.3.69 (Tacarcuna, Panama); AMNH series inspected, no data taken.
S. rodriguezi: BMNH 1921.12.29.190 (La Palma, Huila); ICN 34845, 35234 (both, Finca Merenberg, Huila).
S. rodriguezi undescribed subspecies or related but undescribed species: ICN 35821 (La Luchata, Galán,Serranía de los Yariguíes, 2000m) 36178, 36179(both, El Talismán, San Vicente de Chucurí, Serraníade los Yariguíes, 2100m; sound recording).
S. stilesi: ICN 34420, 34505, 34512, 34615, 34584,
34609, 34610 (all, Amalfi, Antioquia).
S. vicinior: ICN 31207, 31208 (both, 8km NE of Jeguadas, Alto de Pisones, Mistrató, Risaralda), 34840

54

Taxonomy of Scytalopus tapaculos(finca La Minga, vereda Chicoral, Cato Bitaco, La
Cumbre, Valle); AMNH series inspected, no datataken.
S. latebricola: ICN 23338 (San Lorenzo, Santa Marta,
Magdalena).
S. meridanus (all Venezuela): AMNH 492377 (La Culata, Mérida, 4000m: type), rest of series inspected but
no data taken; COP 9441 (Páramo Zumbador, Táchira, 2600m), 14205 to 14208 (all, Llano Rucio, Mérida, 2500m), 14523 (Quintero, Mérida, 2800m),
14577 (El Escorial, Mérida, 2800m), 20177 to 20181(all, Cerro Niquitaz, El Rincón, Trujillo, 22002300m), 24547 to 24553 (all Boca de Monte, Pregonero, Táchira, 1950-2400m), 26240 (Güamito, Trujillo), 45378 to 45384 (all, Páramo Aricagüa, El Muerto, Falda Norte, Mérida, 3000-3140m), 49294 to
49298 (Santo Domingo, Mérida, 2300-2700m),
64259 to 64272 (Páramo La Negra, Mérida, 32003250m), 65206 (Güaraque, Mérida, 2250m or
1600m), 65312 (La Montaña Teleférico, Mérida,
2600-2650m), 65394 (La Azulita, Mérida, 2300m),
65397 (La Azulita, Mérida, 2100m), 71525 (35 km S
of Mucuchíes, Barinas, 2500m), MNHN 523, 599(both, Mérida). Specimens labelled "S. fuscicauda":
BMNH 89.9.20.695 (Mérida), 89.9-10.999 (Sierra
Nevada, Mérida), 1914.11.26.506 (Mérida),
1914.11.26.507 (La Culata, Mérida, 3000 m),
1914.11.26.505 (Mérida); COP 19963 to 19967 (allPáramo de Cendú, Cendú, Trujillo, 2700-2960m),
26240 (Güamito, Trujillo); CM 37221 (type Páramo
de Rosas, Lara), .
S. caracae: (all Venezuela) BMNH 47.7.16.12(“Caracas”); COP 226, 13047 to 13051, 56781 to
56787, 58459 to 58472, 61644 to 61648 (all, El Junquito, Distrito Federal, 1900-1950m), 1469 to 1470,
13384 to 13384 (all, Colonia Tovar, Aragua, 19001950m), 3345 to 3346 (all, Cerro El Ávila; Plan deLos Lirios, San Isidro, Distrito Federal, 16001700m), 13152 to 13155, 58455 to 58458, 61642 to
61643 (Caracas, León, Distrito Federal, 20002100m), 18850 to 18852 (all, Güarenas, Hacienda
Izcaragüa, Miranda, 1400-1880m), 62603 to 62614(all, Cerro el Ávila, San Isidro, Distrito Federal,
2000m), 62615 to 62617 (both, Cerro el Ávila, San
Antonio de Galipán, Distrito Federal, 1600m), 62618(Cerro el Ávila, Estación Teleférico, Distrito Federal,
2100m), 62619 to 62621 (Cerro el Ávila, Cerro Papelón, Distrito Federal, 1850-1900m), 75775 and78732 (both, Colonia Tovar; Fundo Jeremba, Aragüa, 2300m).
Scytalopus spillmanni Ecuador and southern ColombianAndes: AMNH 176050 (Baeza, Ecuador), 180944(Tambillo, Río Upano, Ecuador, 8000 ft.); (BMNH
1902.3.13.1261 (Porvenir, Bolívar, Ecuador); MFNU
1937.11.76 (type: Volcán Iliniza, Pichincha / Coto-

Donegan &Avendaño-C.

paxi, Ecuador) (photographs); ANSP 8374, 8442,
8448, 8540 (all, Buena Vista, Huila, Colombia, 2300m), 176867, 176868, 176869, 176870, 176871,
176872 (all, west slope Andes, road from Quito to
Mindo, 2.5 km south of Tandayapa, Pichincha, Ecuador, 2300 m), 178099, 178101, 178102 (all, SW slope of Volcán Pichincha, new road to Mindo, Pichincha, Ecuador, 2100m), 178100 (as previous, 2300m), 181226, 181228 (both, west slope near road between Maldonado and Tulcán, along Río La Plata,
Carchí, Ecuador, 2525 m, 0°48’N, 78°02’W)(photographs); FMNH 292133 (both, La Victoria,
Nariño, Colombia, 2700m) 292135, 292138 (both, asprevious, 2800m), 292139 (Llorente, Nariño, 1800m)(photographs). Western Andes: ICN 35026, 35029(both, Páramo Frontino, Antioquia, 2600m), 35032,
35033, 35034 (all, vereda Meseria, Jardín, Antioquia,
2300m).
Central Andes population: BMNH
1921.12.29.189 (La Guneta, Quindio, 10,300ft.); ICN
33166 (Ucumarí, Risaralda). Eastern Andes: BMNH
89.9-10.995 (“Bogotá”); ICN 35605 (male) and
35608 (female) (Finca Pamplona/La Aurora, vereda
San Isidro, municipality of Galán, Serranía de los
Yariguíes, Santander, Colombia, 06º38’N; 73º24’W,
2700 m; sound recording: tissue samples (heart) andskeleton at Universidad de los Andes); MLS 3990
(Fontibón, Pamplona, Norte de Santander, Colombia), 3991, (Alturas de Pamplona, Norte de Santander, Colombia).
S. sp. (all Venezuela): COP 14363 (El Valle, Mérida,
2200m), 62230 (Río Chiquito, Hacienda La Providencia, Táchira, 2180-2250m), 65395 and 65396 (LaAzulita, Mérida, 2300m).
S. g. griseicollis (including specimens labelled "S. infasciatus"): AMNH 132328 (type of S. infasciatus:
Páramo de Beltrán) and rest of series inspected, nodata taken; BMNH 44.12.31.16, 44.12.31.24,
69.8.16.31, 80.4.30.33, 80.4.30.34, 89.9.20.691 to
89.9.20.694, 89.9-10.978, 89.9-10.996, 89.9-10.999(all “Bogota”), 2002.3.1014 (“New Grenada”);
IAVH 10305 (Mamaramos, SFF Iguaque, Boyacá),
12282 (vereda Ermitaño, Sutamarchán, Boyacá),
12586 (vereda La Capilla, near SFF Igüaque, Villa deLeyva, Boyacá), 12701, 12712, 12716, 12717, 12720(all, vereda San Francisco, Güasca, PNN Chingaza,
Cundinamarca); ICN 10852 (vereda Ruparita, 2km E
of Arcabuco, Boyacá), 10853 (vereda Puente Boyacá, Ventaquemala, Boyacá), 11982 to 11984 (all,Páramo de Güasca, Cundinamarca, 3500m), 11985 to
11988 (all, Alto Onzaga, Soatá, Boyacá), 12001(Güasca, Cundinamarca), 12007 (La Mercedes, km
25, Carretera La Herrera - La Mesa, Cundinamarca),
12008 to 12009 (Boquerón de Chipaque, Cundinamarca), 12010 (Laguna de Chisacal, Cundinamarca,
3900m), 12011 to 12012 (Choachí, Cundinamarca),

Ornitología Colombiana No.6 (2008)
12013 (Subachoque, Cundinamarca), 12014 to 12016(both, Páramo de Guasca, Cundinamarca) 13015 to
13017 (Páramo de Palacio, Cundinamarca), 14074(hacienda El Otoño, Sabana de Bogotá, Cundinamarca), 19963 (Monserrate, Bogotá, Cundinamarca),
24920 (Boquerón de Juan Viejo, Pasca, Cundinamarca), 31235, 31236 (both, Páramo de Chingaza, Piedras Gordas, La Calera, Cundinamarca), 31798(Suba, Cerro La Conejera, Bogotá, Cundinamarca),
34492 (Bosque de Torca, cerros orientales de Bogotá, Cundinamarca, 3050m), 34780 (vereda Aurora,
altos cerros orientales, Bogotá, Cundinamarca),
35441 (vereda Molinos, Soatá, Boyacá), MLS 3985
(Nomocón, Cundinamarca), 3988 (Tabio, Cundinamarca), 3989 (La Calera, Cundinamarca); MCZ
76330, 76331 (both, types of S. griseicollis:“Bogotá”); MNHN 518, 2001.519, 2003-894 (all,“Colombia”).
S. griseicollis undescribed Santander-Tamá subspecies:
COP 11101 to 11103 (all, PNN Tamá, Apure, Venezuela 2800m), 62227 to 62229, 62231, 62244, 62245(Río Chiquito, Hacienda La Providencia, Táchira,
2180-2300m), 73946 to 73952 (La Revancha, Cumbre, Cerro El Retiro, Táchira, 2700-2800m)); IAVH
10625, 10664, 10728 (all, Sector Orocué); Herrán,
PNN Tamá, Norte de Santander), 12068, 12123,

55

12125 (all, vereda Carrizal, Sector Sisavita, Cucutilla, Norte de Santander); ICN 36121 (Suratá, Santander), ICN 36416 (Piedecuesta, Santander); MLS
3992 (Alturas de Pamplona), 3993 (Fontibón, Pamplona).
S. griseicollis gilesi: ICN 35609 (sound recording),
35610, 36175 (details of each in text above).
S. sp. Perijá: COP (all Venezuela) 54930 to 54933,
54945 to 54947 (all, Sierra de Perijá, Cerro Pejochaina, Cumbre, Zulia, 1900-2350m), 54934 to 54944,
54948 (all, Sierra de Perijá, SE Cerro Tetari, Zulia,
2900m), 57708 (all, Sierra de Perijá, Pie Nudo 4 deFebrero, Zulia, 2600m), 57709 to 57710 (all, Sierrade Perijá, Fila Macoita, Campamento Avispa, Zulia,
2175m), 72846 to 72855 (all, Sierra de Perija, Frontera, Zulia, 2750-3050m), 74170 to 74172 (all, Sierrade Perijá, Cerro Viruela, Zulia, 3100m); ICN 36125
and 36126 (both vereda El Cinco, Manaure, Serraníade Perijá, Cesar, Colombia, 2600m).
S. canus canus: AMNH series inspected, no data taken;
BMNH 1921.7.3.70 (Paramillo, Antioquia, 12,500ft); ICN 35020 (Páramo Frontino, Urrao, Antioquia,
3500m).
S. cf. canus opacus: BMNH 1953.68.680 (Pichincha,
Ecuador); MNHN 1390 (Cerro Mojanda, Ecuador).

APPENDIX 2: BIOMETRICS OF NORTHERN RHINOCRYPTIDAE
In the table below, all data are from Colombia and Venezuela, with species marked * also including datafrom NHM and MNMH specimens from Ecuador. For each species’ measurement, data are presented asfollows: mean ± standard deviation (minimum-maximum) (n= number of specimens). Species order andnomenclature follows Remsen et al. (2008), as modified herein.
Tail (mm)Tarsus (mm)Myornis senilis*

Wing chordfrom skins(mm)
57.5 ± 2.5 (55.0-65.0) (n=28)

58.8 ± 4.2 (53.0-63.0) (n=28)

M. senilis
Ecuador59.4 ± 3.0 (55.0-65.0) (n=9)

59.1 ± 2.3 (56.0-63.0) (n=10)

M. senilis W
Andes

56.0 ± 0.0 (56.0-56.0) (n=2)

68.5 ± 0.7 (68.0-69.0) (n=2)

M. senilis C
Andes

59.0 ± 1.4 (58.0-60.0) (n=2)

59.5 ± 4.9 (56.0-63.0) (n=2)

M. senilis E
Andes

56.4 ± 1.6 (54.0-60.0) (n=15)
56.0 ± 2.7(52.0-60.0)(n=48)

57.1 ± 3.6 (52.0-63.0) (n=14)

21.9 ± 0.7 (21.0-24.0) (n=28)
22.1 ± 0.8(21.0-24.0)(n=10)
22.0 ± 0.0 (22.0-22.0) (n=2)
21.5 ± 0.7(21.0-22.0)(n=2)
21.8 ± 0.6 (21.0-23.0) (n=14)
22.5 ± 0.9(21.0-24.0)(n=45)
22.3 ± 0.5(22.0-23.0)
(n=4)

Taxon

Scytalopus latrans*
S. latrans W
Andes

54.0 ± 2.2 (52.0-57.0) (n=4)

39.6 ± 2.2 (35.0-43.0) (n=43)39.3 ± 1.7 (37.0-41.0) (n=4)

Full culmen(mm)Body mass (g)15.7 ± 0.9 (14.0-17.5) (n=24)

19.0 ± 0.7 (18.5-19.5) (n=2)

15.9 ± 1.1 (14.0-17.5) (n=9)/

16.8 ± 0.4 (16.5-17.0) (n=2)

19.0 ± 0.7 (18.5-19.5) (n=2)//

15.4 ± 0.6 (14.5-16.5) (n=13)
13.6 ± 0.9(12.0-14.5)(n=45)
12.8 ± 1.1(12.0-13.5)(n=2)/
18.1 ± 1.7(16.0-21.0)(n=12)/

56

Taxonomy of Scytalopus tapaculos

Donegan &Avendaño-C.

Taxon

Wing chordfrom skins(mm)

Tail (mm)Tarsus (mm)S. latrans CAndes

57.5 ± 2.5 (53.0-60.0) (n=9)

38.6 ± 2.0 (36.0-41.5) (n=9)

22.7 ± 0.9(21.0-24.0)(n=10)

S. latrans E
Andes

55.0 ± 2.1 (52.0-60.0) (n=28)

39.7 ± 2.2(36.043.0) (n=25)

22.4 ± 0.9(21.0-24.0) (n=27)

S. micropterus*

59.2 ± 2.9 (55.0-61.0) (n=6)

49.6 ± 4.8 (44.0-56.0) (n=5)

24.5 ± 1.1 (23.0-26.0) (n=5)

13.7 ± 0.6(12.0-14.5)(n=10)
13.4 ± 0.6(12.0-14.5)(n=27)
16.0 ± 0.7 (15.0-16.5) (n=4)

S. atratus(nominate andS. a. confusus)

57.0 ± 1.4 (55.0-59.0) (n=6)

40.0 ± 2.6 (36.0-43.0) (d.f.=6)

22.5 ± 0.9 (21.0-23.5) (n=6)

14.6 ± 0.6(14.015.5) (n=6)/

S. panamensis

51 (n=1)

39 (n=1)/

14.5 (n=1)/

S. rodriguezi

54.3 ± 2.1 (52.0-56.0) (n=3)

47.0 ± 1.7 (45.0-48.0) (n=3)

22.5 ± 0.9 (22.0-23.5) (n=3)

14.8 ± 0.3 (14.5-15.0) (n=3)

21.5 ± 0.1 (21.4-21.6) (n=3)

55.7 ± 1.7 (52.0-57.0) (n=7)

39.2 ± 1.8 (36.0-40.2) (n=5)

20.8 ± 0.5 (20.0-21.5) (n=5)

14.1 ± 0.4 (13.5-15.0) (n=3)

17.1 ± 1.7 (15.1-19.0) (n=4)

53.0 ± 1.8 (51.0-54.0) (n=4)
56.5 ± 0.8 (55.0-57.0) (n=6)
57.0 ± 1.7(56.0-59.0)(n=3)

42.7 ± 0.6 (42.0-43.0) (n=3)
43.3 ± 1.4 (41.0-45.0) (n=6)

22.4 ± 0.9 (21.5-23.5) (n=4)
22.8 ± 0.6 (22.5-24.0) (n=7)

13.3 ± 0.3 (13.0-13.5) (n=3)
15.1 ± 0.5 (14.5-16.0) (n=6)

46.0 ± 2.6 (43.0-48.0) (n=3)

22.7 ± 0.8(22.023.5) (n=3)

14.7 ± 0.3 (14.5-15.0) (n=3)

24.2 ± 2.6 (21.5-26.7) (n=3)

S. latebricola

58 (n=1)

36 (n=1)

23.0 (n=1)

14.0 (n=1)/

S. caracae

52.8 ± 1.6 (50.0-54.0) (n=5)

39.8 ± 1.9 (37.0-42.0) (n=5)

21.9 ± 0.7 (21.0-23.0) (n=5)

14.5 ± 0.5 (14.0-15.0) (n=5)/

60.0 ± 2.9 (58.0-65.0) (n=5)

39.4 ± 3.4 (36.0-45.0) (n=5)

22.5 ± 0.5 (22.0-23.0) (n=5)

13.2 ± 1.1 (11.5-14.5) (n=5)

21.7 ± 1.8 (20.4-23.0) (n=2)

60.7 ± 3.8 (58.0-65.0) (n=3)

40.0 ± 4.6 (36.0-45.0) (n=3)

22.8 ± 0.3 (22.5-23.0) (n=3)

13.0 ± 1.5 (11.5-14.5) (n=3)

23.0 (n=1)

58.0 (n=1)

39.0 (n=1)

22.0 (n=1)

13.5 (n=1)

21.7 (n=1)

61.8 ± 2.4 (57.0-67.0) (n=43)45.2 ± 3.0 (39.0-54.0) (n=39)

24.4 ± 1.0 (22.2-26.0) (n=21)

14.5 (n=1)

25.1 ± 2.2 (20.5-30.0) (n=43)62.2 ± 2.2 (58.0-67.0) (n=37)

45.5 ± 3.1 (39.0-54.0) (n=33)

24.5 ± 1.0 (22.2-26.0) (n=18)/

25.1 (20.530.0) (n=37)

58.7 ± 1.5 (57.0-60.0) (n=4)

43.0 (41.0-45.0)(n=4)

23.8 (23.5-24.1)
(n=4)

/

24.9 (22.029.5) (n=5)

59.0 ± 1.4 (58.0-60.0) (n=2)

41.5 ± 2.1 (40.0-43.0) (n=2)

24.5 ± 0.7 (24.0-25.0) (n=2)

15.0 ± 0.7 (14.5-15.5) (n=2)

27.0 (n=1)

S.rodriguezi
sp.
(E. Andes:Col.)
S. sp.
(Ven. Andes)
S. stilesi
S. vicinior

Scytalopus
spillmanni (EAndes) All
Scytalopus
spillmanni (EAndes) Males
Scytalopus
spillmanni (EAndes) Females
Scytalopus
spillmanni Ecuador All

Scytalopus
spillmanni Ecuador Males

Scytalopus
spillmanni Ecuador Females

Scytalopus
spillmanni (CAndes) Males

Full culmen(mm)Body mass (g)19.4 ± 1.5 (18.0-21.0) (n=3)
17.4 ± 1.7 (16.0-19.3) (n=7)//
21.3 ± 0.8 (20.0-22.0) (n=6)

Ornitología Colombiana No.6 (2008)
Taxon
Scytalopus
spillmanni (WAndes) Males
Scytalopus
spillmanni (WAndes) Females
S. g. griseicollis
S. g. griseicollis
Males
S. g. griseicollis
FemalesS. griseicollis(SantanderTamá)
S. griseicollis(SantanderTamá) Males
S. griseicollis(SantanderTamá) Females
S. griseicollis
gilesi Males
S. sp.
(Perijá)S. sp.
(Perijá) Males
S. sp.
(Perijá) Females
S. meridanus(inc. "S. fuscicauda")
S. meridanus
Males
S. meridanus
FemalesS. canus canus
S. cf. canus
opacus57

Wing chordfrom skins(mm)

Tail (mm)Tarsus (mm)Full culmen(mm)Body mass (g)59.3 ± 0.6 (59.0-60.0) (n=3)

44.0 ± 4.2 (41.0-47.0) (n=2)

24.0 ± 1.0 (23.0-25.0) (n=3)

15.0 ± 0.9 (14.0-15.5) (n=3)

24.4 ± 2.1 (23.0-26.8) (n=5)†

56.5 ± 0.7 (56.0-57.0) (n=2)

42.5 ± 0.7 (42.0-43.0) (n=2)

23.0 ± 0.0 (23.0-23.0) (n=2)

14.0 ± 0.0 (14.0-14.0 (n=2)

23.0 ± .14 (22.0-24.0) (n=2)†22.0 ± 0.9(20.0-23.5)(n=43)
22.2 ± 0.9 (20.0-23.5) (n=22)
21.5 ± 0.7 (20.5-22.5) (n=8)

12.7 ± 0.7 (11.0-14.0) (n=38)

17.1 ± 2.6 (11.0-21.0) (n=12)55.5 ± 2.5 (52.0-61.0) (n=22)
54.8 ± 2.5 (50.0-59.0) (n=9)

39.8 ± 1.9(36.5-43.5)(n=28)
39.9 ± 1.6 (38.0-43.5) (n=15)
38.7 ± 1.8 (36.5-41.0) (n=7)

12.9 ± 0.7 (11.0-14.0) (n=22)
12.3 ± 0.7 (11.5-13.0) (n=8)

17.5 ± 3.0 (11.0-21.0) (n=8)
16.5 ± 1.9 (16.0-18.0) (n=4)

55.5 ± 2.6 (52.0-60.0) (n=11)

40.3 ± 3.1 (34.0-44.0) (n=11)

21.4 ± 0.8 (20.5-22.5) (n=10)12.6 ± 0.6 (11.5-13.5) (n=10)

16.3 ± 0.7 (15.0-17.0) (n=6)

57.0 ± 2.0 (55.0-60.0) (n=5)

41.2 ± 2.5 (38.5-44.0) (n=5)

21.9 ± 0.8(21.5-22.5)(n=5)

12.8 ± 0.5 (12.0-13.5) (n=5)

16.8 ± 0.4 (16.3-17.0) (n=3)

54.3 ± 2.6 (52.0-58.0) (n=6)

39.6 ± 3.5 (34.0-43.5) (n=6)

21.0 ± 0.5 (20.5-22.0) (n=5)

12.4 ± 0.6 (11.5-13.0) (n=4)

15.8 ± 0.8 (15.0-16.0) (n=3)

57.0 ± 1.0 (56.0-58.0) (n=3)
51.9 ± 2.5 (48.0-55.0) (n=28)
52.7 ± 2.6 (49.0-55.0) (n=11)

44.3 ± 0.6 (44.0-45.0) (n=3)
38.5 ± 1.5 (36.0-42.5) (n=26)
39.4 ± 1.7 (37.0-42.5) (n=10)

22.5 ± 0.5 (22.0-23.0) (n=3)
21.1 ± 1.0 (18.5-22.5) (n=27)
21.6 ± 0.9 (19.5-22.5) (n=11)

13.6 ± 0.4 (13.0-14.0) (n=3)
14.0 ± 0.6 (13.0-15.5) (n=22)
14.3 ± 0.5 (13.5-15.5) (n=10)

17.8 ± 0.4 (17.5-18.0) (n=2)

51.9 ± 2.3 (48.0-54.0) (n=12)38.3 ± 1.2 (37.0-40.0) (n=11)

20.9 ± 0.9 (19.5-22.5) (n=12)13.8 ± 0.5 (13.0-14.5) (n=9)/

50.8 ± 2.2 (46.0-55.0) (n=45)

38.4 ± 2.2 (34.0-43.0) (n=37)

21.3 ± 0.9 (20.0-23.0) (n=45)13.0 ± 0.6 (12.0-14.0) (n=42)/50.9 ± 2.5 (48.0-54.0) (n=8)
50.1 ± 2.3 (44.0-53.0) (n=22)49.0 ± 2.9 (47.0-51.0) (n=2)
50.5 ± 2.1 (49.0-52.0) (n=2)

40.2 ± 1.9 (38.0-43.0) (n=6)
37.3 ± 2.0 (34.0-43.0) (n=16)34.0 ± 1.4 (33.0-35.0) (n=2)
36.5 ± 0.7 (36.0-37.0) (n=2)

21.5 ± 0.8 (20.5-23.0) (n=8)
20.9 ± 0.7 (19.5-22.5) (n=22)
21.5 ± 0.7 (21.0-22.0) (n=2)
20.8 ± 1.1 (20.0-21.5) (n=2)

13.4 ± 0.6 (12.5-4.0) (n=8)
12.9 ± 0.7 (12.0-14.0) (n=20)
12.5 ± 0.7 (12.0-13.0) (n=2)
11.8 ± 0.4 (11.5-12.0) (n=2)

54.9 ± 2.5 (50.0-61.0) (n=41)† A recent capture apparently of this population (data not included above) massed 20 g (Pulgarín 2007).

/
/

/
/
/
/

58

Taxonomy of Scytalopus tapaculos

Donegan &Avendaño-C.

APPENDIX 3: VOCAL VARIABLES OF TAXA AND POPULATIONS OF SCYTALOPUS DISCUSSED IN THIS PAPER. For each species and call, data is presented as follows: mean ± standard
deviation (minimum - maximum) (n= number of specimens). Numbers in the second column refer tonumber of recordings and spectrograms inspected for the purposes of minimum and maximum data recorded values. XC refers to www.xeno-canto.org catalogue numbers. IAVH refers to IAVH’s sound archive catalogue numbers.
A. Gazzetteer of recording localities:
S. g. griseicollis: Rogitama, mun. Arcabuco, Boyacá,
Colombia (05°47’N, 73°31’W, 2500m) (J. Zuluaga,
R. Chavarro & J. Beckers recordings); Soatá, Boyacá, Colombia (05°07’N, 73°07’W) (O. Cortés, A.
Hernández & O. Laverde recordings); SFF Igüaque,Boyacá, Colombia (Sector Carrizal) (05º44'N,
73º31'W, 2800 m) (J. Zuluaga recordings); Reserva
Privada el Secreto, Vereda Ciénaga-Valvanerra,
mun. Garagoa, Boyacá, Colombia (2000-2200 m)(O. Laverde recordings); finca San Cayetano, vereda Fute, mun. Bojacá, Cundinamarca, Colombia(4º38’N, 74º18’ W, 2650 m) (O. Laverde recording); PNN Chingaza, Cundinamarca, Colombia (J.
Parra & O. Laverde recordings); Monserrate, Bogotá, Cundinamarca, Colombia (J. Beckers recordings); Quebrada La Vieja, Bogotá, Colombia(2900m) (D. Knapp recording: XC 10866); finca La
Carbonera, vereda el Roble, inspección La Aguadita, mun. Fusagasugá, Cundinamarca, Colombia (4°25’N, 74°19’W, 2450-2550 m) (O. Laverde recordings).
S. g. gilesi: Alto Cantagallos, mun. San Vicente de Chucurí, Serranía de los Yariguíes, Santander, Colombia (06°49'N, 73°22'W, 2450m) (TMD recordings:
XC 18457, 18458, 18459 and others); Lepipuerto,
mun. El Carmen / Simacota, Serranía de los Yariguíes, Santander, Colombia (6°28’N, 73°28’W,
2900m) (TMD recordings: XC 18471, 18472,
18473, 18475, 18476, 18477, 18478 and others);
Filo Pamplona, Mun. Galán, Serranía de los Yariguíes, Santander, Colombia (06º38’N, 73º24’W,
3200m) (TMD & JEAC recordings XC 18452,
18453, 18454, 18455, 18456 and others).
S. griseicollis Santander-Tamá subspecies: Tamá, Río
Oría, Apure state, Venezuela (2350-2410 m) (C.
Parrish recordings: XC 6079, 16655, 16656, 16657,
16658, 16659, 16660, 16661); Páramo el Judio,
Apure (2730 m) (C. Parrish recording: XC 16654);
Suratá, Santander (07°23'N, 73°00W, 3000m)(JEAC recordings); Vereda Carrizal, Norte de Santander (S. Córdoba-C. recording: IAVH 8812).
S. meridanus: 10 km SE of La Azulita, Mérida, Venezuela, 2300m (C. Parrish & A. Spencer recordings:
XC 6234, 6235, 6236, 14790); Pico Humboldt
Trail, Parque Nacional Sierra Nevada, Mérida, Venezuela, 2500m (N. Athanas & A. Spencer recor-

dings: XC 8249 and 11243); Páramo de Zumbador,Táchira, Venezuela (2500-2700m) (Boesman1999); Yacumbu NP, Lara, Venezuela (1700m)(Boesman 1999); Laguna Mucubaji area, Mérida,
Venezuela (3200m) (Boesman 1999); Páramo de
Batallón, Táchira, Venezuela (2700-2900m)(Boesman 1999); Humucaro Alto, Lara, Venezuela(2650m) (Boesman 1999); Güaramacal NP, Trujillo, Venezuela (2400-2600m) (Boesman 1999).
S. spillmanni Ecuador recordings: Bellavista, Tandayapa(2200m) (W. Halfwerk recordings: XC 1480, 4217,
4218, 4219; N. Athanas recording: XC 11544); Old
Chiriboga Road, Pichincha (1800m) (D. Jones recording: XC 3997); Las Palmas, Cotopaxi (00°35’S,
79°00W) (recording 44.1 in Krabbe et al. 2001);
Las Palmas, Cotopaxi (2340m) (track 9.1 in Krabbe& Nilsson 2004), Hotel Bellavista, SW above Tandayapa, Pichincha (2250-2300m) (tracks 9.2, 9.3,
9.9, 9.10, 9.12, 9.14, 9.21 and 9.25 in Krabbe &Nilsson 2004; track 44.2 in Krabbe et al. 2001), San
José, Tandayapa ridge, Pichincha (2300m) (tracks
9.4, 9.5, 9.16, 9.19 and 9.32 in Krabbe & Nilsson
2004), Cordillera Güacamayos, Napo (17002300m) (tracks 9.6, 9.13, 9.15, 9.18, 9.26, 9.28 and9.29 in Krabbe & Nilsson 2004; track 44.4 in
Krabbe et al. 2001), Orregán, Chimborazo (3145m)(tracks 9.7 and 9.8 in Krabbe & Nilsson 2004),
Quebrada las Ollas, Santa Bárbara - La Bonita
Road, Sucumbíos (2150m) (tracks 9.11, 9.23 and9.31 in Krabbe & Nilsson 2004; track 44.3 in
Krabbe et al. 2001), Playa, Napo (3300m) (track
9.17 in Krabbe & Nilsson 2004), Loma Bahamonte,
old Santo Domingo road, Pichincha (2450-2900m)(track 9.20 in Krabbe & Nilsson 2004), NW slope
Volcán Tungurahua, Tungurahua (3100m) (track
9.22 in Krabbe & Nilsson 2004), Tambo deAshilán, upper Río Upano, Morona-Santiago (30303150m) (tracks 9.24 and 9.27 in Krabbe & Nilsson
2004), SW slope, Volcán Corazón, Pichincha (3150-3850m) (track 9.30 in Krabbe & Nilsson 2004);
Quebrada el Recreo, Imbrarura (00°24’N, 78°27’W) (track 44.5 in Krabbe et al. 2001).
S. spillmanni Western Andes recordings: (not analysed
in detail due to small sample size): PNN Munchique, Cauca (2600m) (O. Cortés recordings);
Cuchilla La Linea-La Cumbre, Pueblo Rico, Risar-

Ornitología Colombiana No.6 (2008)
alda (05°14’N 76°02’W) (IAVH 15768: Álvarez-R.
et al. 2007).
S. spillmanni Central Andes recordings: Ibagué, El Rancho, road to Nevado del Tolima, Tolima (L.E.
Urueña recordings); Reserva Natural Ibanasca, Ibagué, Tolima (04°36’N; 75°15W, 2300-2600 m) (O.
Laverde recordings: IAVH 24179, 24188, 24237,
24242, 24245, 24251, 24284, 24287); Reserva Natural Río Blanco, Manizales, Caldas (05°05’N 75°
21’W, 2550-3600m) (O. Laverde recordings: IAVH
24504, 24516, 24624, 24632, 24672; D. Bradley
recordings: XC 17623, 17624; IAVH 30903: Álvarez et al. 2007; Álvarez-R. & Córdoba-C. 2002);
Parque Regional Ucumarí, Pereira, Risaralda (04°43’N 75°35’W) (IAVH 6807: Álvarez-R. et al.
2007); Finca los Molinos, corregimiento de Dantas,
Ibagué, Tolima (04°34’N 75°16’W; 2000-2200 m)(O. Laverde recording: IAVH 17271); Vereda El
Laurel, Hacienda Termópilas, Aranzazu, Caldas (5°14 N, 75°29’ W, 2250m) (S. Córdoba-C recording:
IAVH 7900); Reserva Los Yalcones, San Agustín,
Huila (01° 49 N, 76°21’W, 2322-2430m) (D. Calderón recording: IAVH 16625); RNA El Mirador,

59

Génova, Quindío (2900 m) (F. Lambert recording:
XC 17623).
S. spillmanni Eastern Andes recordings: La Aurora, finca Pamplona, Serranía de los Yariguíes, vereda San
Isidro, municipio Galán, Santander Dept, Colombia(06º 38’ N; 73º 24’ W; 2700 m) (TMD & JEAC
recordings: XC 18486, 18687, 18488 and others),
Lepipuerto, mun. El Carmen / Simacota, Serraníade los Yariguíes, Santander, Colombia (6°28’N, 73°
28’W, 2900m) (TMD recordings: XC 18489, 18490
and others); Finca La Carbonera, Los Robles, La
Agüadita, Fusagasugá, Cundinamarca (4°25’N, 74°
19’W, 2450-2550m)(O. Laverde recordings:
IAVH 17854, 17855, 17856, 17857, 17884, 17917,
17939, 17940); Vereda Carrizal, Norte de Santander(S. Córdoba-C. recording: IAVH 8645).
“S. cf. canus opacus undescribed southern race”: see
Krabbe et al. (2001) and Krabbe & Nilsson (2004)
for localities.
S. latrans, S. atratus and S. rodriguezi sp.: Various
Serranía de los Yariguíes recordings by the authors
and published recordings in Álvarez et al. 2007 andKrabbe & Nilsson (2004).

No. of calls / no. of recordings

Duration(s)

Flat: 45/15: Chingaza(23/9), Iguaque (4/1), Soatá(3/1), Fusagasugá (6/1)
Monserrate (2/1), La Vieja(4/1) and Rogitama (3/1).

Flat: 6/3: Yariguíes (1/1: La
Aurora), Fusagasugá (1/1)
and Carrizal (4/1).
111/24:
Rising: 66/7: Bogotá (1/1),
Chingaza (42/3), Iguaque(5/1) & Soatá (18/2).

43/17:
Rising: 37/14: Yariguíes(17/8: all La Aurora) andFusagasugá (20/6).

Flat: 48/12: Los Molinos(1/1), Ibanasca (5/3), Rio
Blanco (17/5), Ucumarí(1/1), Ibagué road to Nevado (2/1) and Aranzazu(22/1)

Flat: 48/11: Chiraboga (3/1),
Güacamayos (3/2), Tandayapa (34/6), Playa (5/1), Las
Ollas (3/1).
68/19:
Rising: 20/7: Ibagué road to
Nevado (1/1), Ibanasca(3/1), Ucumarí (1/1), Río
Blanco (12/2), San Agustín(2/1) and Génova (1/1).

2.22 ±
3.19 (0.1513.35)(n=57)

5.54 ±
4.83 (0.6213.50)(n=18)

5.86 ±
4.88 (0.3514.10)(n=31)

4.12 ±
9.03 (0.0868.75)(n=57)

29.07 ± 2.12
(25.33-32.98)(n=17)

24.98 ± 3.20
(20.00-32.00)(n=57)

53.39 ±
73.77
(4.00294.00)(n=56)

28.72 ± 3.01
(22.00-36.21)(n=33)

34.02 ± 3.36
(27.00-42.86)(n=57)

Pace (Notes/s)

169.00 ±141.23(20.00369.00)(n=26)

155.01 ±
130.15(10.00504.00)(n=34)

130.35 ±263.66(3.001856.00)(n=66)

No. ofnotes

2.37 ± 0.24
(1.70-2.90)(n=55)

3.13 ± 0.26
(2.80-3.60)(n=26)

2.76 ± 0.25
(2.35-3.25)(n=34)

3.11 ± 0.39
(2.40-4.20)(n=66)

Max. frequency of lowest note (kHz)2.80 ± 0.20
(2.50-3.40)(n=57)

3.68 ± 0.25
(3.20-4.10)(n=25)

3.01 ± 0.31
(2.55-3.90)(n=34)

3.68 ± 0.47
(2.70-4.90)(n=66)

Max. frequency of highest note(kHz)

0.42 ±
0.21 (0.101.20)(n=54)

0.53 ±
0.19 (0.200.90)(n=25)

0.26 ±
0.16 (0.100.70)(n=34)

0.57 ±
0.32 (0.101.60)(n=66)

Frequency
variation(kHz)

Up-down strokes (100%)

Up-down strokes with somedouble-notes (100%).

Flat: Upstrokes, some with faintleading upstroke (20%), up-down
strokes with some double notes(80%)

Rising: Up-down strokes withsome double-notes (100%).

Rising: Up-down strokes withsome double-notes (56%); downstrokes, some with faint leading
upstroke (44%).
Flat: Downstrokes, some with
faint leading upstroke (78%),
updownstrokes (including broad
blobs with up and down tails andsoe double notes) (22%).

Note shape

Taxonomy of Scytalopus tapaculos

Scytalopus
griseicollis
griseicollis(Eastern Andes)

Scytalopus
spillmanni
subsp.
(Eastern Andes)

Scytalopus
spillmanni(C Andes)Scytalopus
spillmanni(Ecuador)

171/32:
Rising: 123/21: Chiriboga(3/1), Mindo (1/1), Tandayapa (36/9) Güacamayos(57/5), Las Palmas (1/1),
Orregán (1/1), Las Ollas(3/1), Volcán Corazón(20/1), Las Palmas (1/1).

Reeling songs (data for flat and rising songs combined)

Call type /Taxon andregion60
Donegan &Avendaño-C.

B. Measurements of vocal variables of taxa and populations of Scytalopus discussed in this paper.

0.94 ± 0.27
(0.24-1.49)(n=29)

1.04 ± 0.17
(0.77-1.32)(n=15)

94/10: Güacamayos (9/1),
Tandayapa (10/1), Bahamonte (2/1), Volcán Tunurahua(10/1), Playa (3/1), Las Ollas(21/2), San José (13/2) andTambo de Ashilán (26/1).

41/5: Ibagué road to Nevado(3/1), Ibanasca (32/3) and
Río Blanco (6/1).

Scytalopus spillmanni(Ecuador)

Scytalopus spillmanni(C Andes)Scolds“Scytalopus
canus opacus
undescribedsouthernrace” (S. Ecuador & Peru)

16.53 ± 3.71
(10.87-23.26)(n=29)

15.81 ± 2.06
(12.57-18.18)(n=15)

15.41 ±
4.86 (5.00-22.00)(n=29)
16.13 ±
1.92
(13.0020.00)(n=15)

31.99 ± 5.64
(21.33-41.00)(n=11)

Flat: 21/5: see Krabbe et al.
(2001) and Krabbe & Nilsson (2004).

S. meridanus
Slower Song229.27 ±160.26(6.00534.00)(n=12)

7.98 ± 1.74
(6.19-11.50)(n=7)

180.86 ±147.54(42.00441.00)(n=7)

23.30 ±
20.25 (5.20-61.20)(n=7)

7/5: La Azulita (2/2) Páramo
de Zumbador (1/1) and Güaramacal NP (1/1).

7.14 ± 4.72
(0.2022.25)(n=11)

14.10 ± 3.60
(8.50-19.80)(n=9)

S. meridanus

23.31 ± 3.67
(19.00-30.00)(n=23)

26.11 ±
10.31
(18.0060.00)(n=19)

1.12 ± 0.34
(0.65-2.00)(n=21)

214.86 ±117.72(68.00479.00)(n=9)

27.19 ± 4.13
(22.00-35.42)(n=27)

Pace (Notes/s)

32.49 ±
11.80
(16.0071.00)(n=25)

No. ofnotes

1.25 ± 0.59
(0.46-3.10)(n=25)

Duration(s)

15.46 ±
8.00 (7.8031.90)(n=9)

Flat: 6/3: Lepipuerto (3/1)
and Filo Pamplona (3/2).

Flat: 11/3: Tamá Venezuela(8/2) and Suratá (3/1).
39/11:
Rising: 33/8: Lepipuerto(32/7) and Filo Pamplona(1/1).

60/9:
Rising: 49/6 Tamá Venezuela (23/4) Suratá (26/2).

No. of calls / no. of recordings

Flat: 10/7: La Aazulita (1/1),
Pico Humboldt (2/2), Páramo de Zumbador (1/1) Yacumbu NP (4/1), Guaramacal
NP (1/1), Mucubaji (1/1) and
Humucaro Alto (1/1).

Scytalopus
griseicollis gilesi

Scytalopus griseicollis subsp.
(SantanderTamá)

Call type /Taxon andregion3.17 ± 0.28
(2.55-3.50)(n=15)

3.73 ± 0.38
(3.10-4.90)(n=29)

4.11 ± 0.31
(3.55-4.60)(n=12)

3.41 ± 0.46
(2.70-4.20)(n=7)

3.38 ± 0.24
(3.00-3.90)(n=9)

2.33 ± 0.36
(2.30-3.00)(n=16)

2.33 ± 0.39
(1.80-2.90)(n=26)

Max. frequencyof lowest note(kHz)

3.60 ± 0.18
(3.25-3.90)(n=15)

4.27 ± 0.36
(3.75-5.15)(n=29)

4.43 ± 0.28
(4.00-4.85)(n=12)

4.64 ± 0.26
(4.20-4.90)(n=7)

3.96 ± 0.23
(3.60-4.40)(n=9)

2.61 ± 0.23
(2.00-3.30)(n=26)

2.88 ± 0.20
(2.45-3.15)(n=27)

Max. frequencyof highest note(kHz)

0.43 ± 0.16
(0.20-0.75)(n=15)

0.54 ± 0.28
(0.05-1.30)(n=29)

0.31 ± 0.18
(0.10-0.65)(n=12)

1.23 ± 0.49
(0.50-1.90)(n=7)

0.48 ± 0.16
(0.30-0.80)(n=9)

0.37 ± 0.16
(0.10-0.60)(n=14)

0.57 ± 0.30
(0.10-1.10)(n=26)

Frequency
variation(kHz)

Up-down stroke (100%).

Up-down stroke (90%), blob with
faint upstroke (10%).

Downstrokes, sometimes with thin
upstroke (60%); lines (40%).

Fast downstrokes or lines (100%).

Fast downstrokes or lines (100%).

Up-down strokes (100%)

Up-down strokes (100%)

Note shape

Ornitología Colombiana No.6 (2008)61

1.07 ± 0.11
(0.85-1.23)(n=15)

1.19 ± 0.30
(0.55-2.00)(n=53)

0.99 ± 0.11
(0.80-1.30)(n=15)

1.11 ± 0.17
(0.70-1.50)(n=33)

1.16 ± 0.30
(0.75-1.60)(n=7)

60/8: Yariguíes (26/6, of
which Lepipuerto 5/2 and La
Aurora 21/4) and Fusagasugá(34/2).

152/20: Rogitama (38/5),
Chingaza (92/11); Soatá(4/2), Garagoa (7/1), Bojacá(8/1), Fusagasugá (2/1) andMonserrate (1/1).

49/6: Tamá Colombia (3/1)
and Tamá Venezuela (46/5).

75/13: Alto Cantagallos(19/3), Lepipuerto (30/7) andFilo Pamplona (26/4).

14/3: Páramo de Batallón(7/1), Güaramacal NP (6/1)
and La Azulita (1/1).

64/8: see Krabbe et al. (2001)
and Krabbe & Nilsson(2004).

Scytalopus spillmanni subsp.
(Eastern Andes)

Scytalopus
griseicollis
griseicollis(Eastern Andes)

Scytalopus griseicollis subsp.
(SantanderTamá)

Scytalopus
griseicollis gilesi

S. meridanus“S. cf. canus
opacus undescribed southernrace” (S. Ecuador & Peru)
0.94 ± 0.14
(0.65-1.26)(n=21)

Duration(s)No. of calls / no. of recordings

Call type /Taxon and region
19.00 ±
1.81
(14.0022.00)(n=19)
32.36 ±
8.35
(16.0052.00)(n=42)
28.80 ±
5.56
(23.0046.00)(n=15)
31.36 ±
4.94
(21.0043.00)(n=32)
21.14 ±
6.47
(12.0031.00)(n=7)
23.89 ±
4.27
(17.0033.00)(n=21)

No. ofnotes

25.47 ± 2.54
(20.59-30.00)(n=21)

18.06 ± 1.04
(16.00-19.23)(n=7)

28.10 ± 2.64
(21.82-33.64)(n=32)

29.05 ± 2.42
(26.36-35.38)(n=15)

28.21 ± 1.89
(24.83-32.31)(n=42)

17.74 ± 1.90
(13.93-21.78)(n=15)

Pace (Notes/s)

3.97 ± 0.27 (3.20-4.40) (n=21)

4.96 ± 0.23 (4.70-5.30) (n=7)

2.39 ± 0.31 (1.70-2.90) (n=33)

2.90 ± 0.23 (2.40-3.30) (n=14)

2.91 ± 0.28 (2.20-3.70) (n=53)

3.83 ± 0.20 (3.30-4.25) (n=16)

Max. frequencyof lowest note(kHz)

4.44 ± 0.19 (3.95-4.85) (n=21)

5.44 ± 0.20 (5.20-5.70) (n=7)

3.20 ± 0.26 (2.60-3.60) (n=33)

3.97 ± 0.14 (3.70-4.50) (n=18)

3.72 ± 0.26 (3.00-4.20) (n=53)

4.17 ± 0.25 (3.70-4.60) (n=17)

Max. frequencyof highest note(kHz)

0.48 ± 0.21
(0.05-1.05)(n=21)(0.20-0.80)(n=7)

0.81 ± 0.31
(0.30-1.70)(n=33)

1.03 ± 0.27
(0.60-1.75)(n=14)

0.81 ± 0.26
(0.20-1.50)(n=53)

0.37 ± 0.11
(0.10-1.00)(n=16)

Frequency
variation(kHz)

Up-down stroke with stronger
downstroke (100%)

Downstrokes (100%).

Up-down stroke with stronger
upstroke (100%).

Up-down stroke with stronger
upstroke (100%).

Up-down stroke with stronger
upstroke (100%).

Up-down stroke (60%); blobwith faint downstroke (40%).

Note shape

62
Taxonomy of Scytalopus tapaculos
Donegan &Avendaño-C.

X

X

X

Scold
(d.f.=20/20)

Plumage(d.f.=15)

Biometrics(d.f.=22/19/8/0/23)

Slower (1)Lower (1,2)

Lower (1,2)

N/A?

Reeling song(d.f.=31/33)

Scold(d.f.14/14)

Plumage(d.f.=1)

S. spillmanni Central Andes

Shorter Tail?

N/APlumage(d.f.=2)

Biometrics(d.f.=12/12/8/0/12)

N/A

Lower (1)Scold(d.f.=3/7)

Reeling song(d.f.=21/28)

S. spillmanni Eastern Ecuador

X

Reeling song(d.f.=23/35)

S. spillmanni Western Ecuador

S. spillmanni
Western Ecuador

N/A?

Lower?

Slower (1)Lower (1,2)

X

X

X

X

Longer Tail?

N/A

N/A

Higher (1)S. spillmanni
Eastern Ecuador

X

X

X

N/A?

N/A?

Higher?

Faster (1)Higher (1,2)

N/A?

N/A?

Higher (1,2)

Faster (1)Higher (1,2)

S. spillmanniCentralAndes

Lighter?

Lower (1,2)

Lower (1,2)

Longer Tarsus? Greater
Mass?

N/A

N/A

Faster (1)Lower (1)Longer Tarsus(1,2)
Greater Mass?

N/A

N/A

Faster (1)Higher (1)S. spillmanni
Eastern Andes

Darker (4)Slower(1,2,3,4,5)

Darker (4)Slower(1,2,3,4,5)Higher (1)Faster (1,2)
Higher (1)Longer Wing
(1,2) Longer
Tarsus (1)Greater Mass
(1,2,4)Longer Wing
(1,2) Longer
Tail (1)Longer Tarsus? Greater
Mass (1,2)

Faster (1)Higher (1)Darker (4)Slower (1,2,4)Higher (1,2,4)

Darker (4)Slower(1,2,4)

Faster (1,2,3,5)Higher (1,2)

Longer Wing
(1,2) Longer
Tarsus (1,2,4)
Greater Mass
(1,2,3,4)Longer Wing
(1,2) Longer
Tail (1,2)
Longer Tarsus (1,2)
Greater Mass
(1,2,3,4)

Faster (1,2)
Higher (1,2)

Darker (4)Slower (1,2,3)
Higher(1,2,3,4,5)

Slower(1,2,3,4,5)Higher (1)Darker (4)Faster (1,2,3,5)Higher (1,2,3)

S. g. gilesi

Faster (1,2)
Higher (1,2)

S. g. griseicollis

Darker (4)Slower(1,2,3,4,5), Lower(1,2)

Longer Wing (1)Longer Tarsus (1)Greater Mass
(1,2,3,4,5)

Darker (4)Slower (1,2,4)Faster (1) Higher(1)Longer Wing
(1,2)
Longer Tail (1)Longer Tarsus(1,2,4) Greater
Mass (1,2,3,4,5)

Darker (4)Slower (1,2,3,4,5)Higher (1)Faster (1) Higher(1,2)

S. griseicollis(SantanderTamá subsp.)

Darker (4)Faster (1,2,3,4,5)
Lower (1,2,3) Note
Shape (4)Slower (1) Lower(1,2,3,4,5) Note
Shape (4)Longer Wing
(1,2,3,4,5) Longer
Tail (1) Longer
Tarsus (1,2)
Greater Mass?

Darker (4)Faster (1,2,3,4,5)
Lower (1) Note
Shape (4)Slower (1) Lower(1,2,4) Note Shape(4)

Longer Wing
(1,2,3,4,5) Longer
Tail (1,2) Longer
Tarsus (1,2,3,4)
Greater Mass?

Darker (4)Lower (1,2,3,4,5)Note Shape (4)Faster (1,2,3,4,5)Note Shape (4)S. meridanus /S. fuscicauda

Ornitología Colombiana No.6 (2008)
63

APPENDIX 4: DIFFERENCES BETWEEN S. SPILLMANNI, S. GRISEICOLLIS AND S. MERIDANUS Shading shows proposed species limits resulting from this article, defined by populations diagnosable to at least Level 5 by voice (for S. spillmanni, using data from the Eastern Andes population only).
Differences marked with a question mark or without Level 1 did not pass test of statistical significance.
Degrees of freedom for vocal data are presented for (song speed/acoustic frequency. Degrees of freedom
for biometric data are presented for (wing/tail/tarsus/culmen/mass).

N/A?

N/A

Plumage(d.f.=4)

Shorter Wing (1,2)
Biometrics
Shorter Tarsus(d.f.=2/2/2/2/1)(1,2,4) Lower
Mass (1,2,3,4)
S. griseicollis Santander-Tamásubsp.
Slower (1) Lower
Reeling song
(1,2)(d.f.=26/26),

Lighter (4)Faster (1,2,3)
Lower (1,2,3,4,5)

Scold(d.f.=31/32)

Slower(1) Lower(1)

Shorter Wing (1,2)
Shorter Tarsus (1)Lower Mass
(1,2,4)

Lighter (4)Faster (1,2,4)
Lower (1,2,4)

Slower (1,2,3,5)
Lower (1,2)

Longer Tail (4)Shorter Tarsus(4) Lower Mass
(4)

Lower (1,2,4)

Lower Mass (4)Lighter (4)Faster (1,2,3,4,5)
Lower (1,2,3,4)Faster(1,2,3,4,5)
Lower (1)Lighter (4)Slower (1,2)
Lower (1,2,3,5)

Shorter Tarsus?
Lower Mass (4)Lighter (4)Faster (1,2,3,4,5)
Lower (1)

Slower (1,2)
Lower (1)

Shorter Tarsus(4), Lower
Mass (4)Lighter (4)Faster(1,2,3,4,5)

Slower (1)Lower (1,2,3)

X

Slower (1)Lower (1)X

Darker?

X

Longer Tail
(1,2,4)

Darker (4)Lower (1)

Slower (1)Lower (1)

X

X

X

X

Longer Tarsus?
Greater Mass
(4)

Faster (1) Higher
(1)

X

X

X

X

Shorter Tail
(1,2,4)

Lighter (4)Higher (1)Faster (1) Higher(1)

Greater Mass (4)Darker (4)Slower (1,2,3,4,5)Higher (1,2,3,4)

Slower(1,2,3,4,5)Higher (1)Darker (4)Faster (1,2) Higher(1,2,3,5)

Shorter Tail (4),Longer Tarsus (4)Greater Mass (4)S. g. gilesi

Faster (1)Higher (1,2,3)Longer Tarsus(4) Greater
Mass (4)Longer Tarsus(4) Greater Mass
(4)

X

S. g. griseicollis

S. spillmanni
Eastern Andes

Shorter Tarsus(4) Lower Mass
(4)

Higher (1,2)

Higher (1,2)

X

S. spillmanniCentral AndesX

Longer Tail (4)Grayer Back (4)Lower (1,2,4)

Slower (1) Lower (1)Grayer Back (4)Lower (1)Longer Tarsus (4)Greater Mass (4)Slower (1,2,3,4,5)Higher?,Higher (1,2,4)Longer Tarsus (4)Greater Mass (4)S. griseicollis(Santander-Tamásubsp.)

Faster (1,2,4) Lower(1,2,3,4,5)Note Shape (4)Longer Wing (1,2,4)Longer Tail (1,2,4)Longer Tarsus?

Faster (1,2) Lower(1,2,3,4,5) Note Shape(4)
Faster (1,2,3,4,5)
Lower (1,2,3,4,5) Note
Shape (4)Direct comparison notpossible

Longer Wing (1)Longer Tarsus?
Longer Bill?

Faster (1,2,4) Lower(1,2,3,4,5) Note Shape(4)
Faster (1,2,3,4,5)
Lower (1,2,3,4,5) Note
Shape (4)Direct comparison notpossible

Longer Wing (4)Longer Tarsus?
Greater Mass?

Darker (4)Lower (1,2,3,4,5) Note
Shape (4)Faster (1,2,3,4,5)
Lower (1) Note Shape(4)

Longer Wing (1,4)
Longer Tarsus (4)Greater Mass?

S. meridanus /S. fuscicauda

Taxonomy of Scytalopus tapaculos

Plumage(d.f.=2)

Slower (1,2,3,5)
Lower (1,2,3)

Shorter Wing (1,2)
Shorter Tail (1)Shorter Tarsus?
Lower Mass (1,2)

Shorter Wing (1,2)
Shorter Tail (1,2)
Shorter Tarsus(1,2) Lower Mass
(1,2,3,4)

Reeling song(d.f.=41/44)

S. g. gilesi

Biometrics(d.f.=21/14/21/2
1/7)

Lighter (4)Faster (1,2,4)

Slower (1,2)
Lower (1,2)

Shorter Tarsus?
Lower Mass?

N/A

N/A

Slower (1) Higher
(1)

N/A?

S. spillmanni
Eastern Ecuador

Lighter (4)Faster (1,2,3,4,5)
Lower (1)Scold(d.f.=41/52)

Plumage(d.f.=56)

Slower (1,2)
Lower (1,2)

Reeling song(d.f.=56/56)

S. g. griseicollis

Shorter Tarsus(1,2) Lower Mass?

N/A

Scold
(d.f.=17/19)

Biometrics(d.f.=2/2/2/2/0)

Slower (1) Lower(1)

Reeling song(d.f.=16/25)

S. spillmanni Eastern Andes

Biometrics(d.f.=1/1/1/1/0)

S. spillmanni
Western Ecuador

64
Donegan &Avendaño-C.

Biometrics(d.f.=6/4/5/6/0)

Shorter Wing
(1,2,3,4,5) Shorter
Tail (1) Shorter
Tarsus (1,2)
Lower Mass?

Shorter Wing
(1,2,3,4,5) Shorter
Tail (1,2) Shorter
Tarsus (1,2,3,4)
Lower Mass?
Shorter Wing
(1,4) Shorter
Tarsus (4)Lower Mass?

Lighter (4)Shorter Wing (4)Shorter Tarsus?
Lower Mass?

Lighter (4)Scold (d.f.=6/6)

Lighter (4)Higher(1,2,3,4,5) Note
Shape (4)Faster (1)Higher(1,2,3,4,5) Note
Shape (4)Faster (1) Higher(1,2,4) Note Shape(4)

Higher (1,2,3,4,5)Note Shape (4)Lighter (4)Slower(1,2,3,4,5)Higher (1) Note
Shape (4)Slower(1,2,3,4,5)Higher (1,2,3)Note Shape (4)Slower (1,2,3,4,5)Higher (1) Note
Shape (4)Slower (1,2,3,4,5)Note Shape (4)Reeling song(d.f.=8/8)

Plumage(d.f.=58)

Shorter Tarsus(4) Lower Mass
(4)

Lighter (4)Faster (1,2,3,4,5)
Lower?

Shorter Tarsus(4) Lower Mass
(4)

Lighter (4)Faster(1,2,3,4,5)Higher (1,2)

S. spillmanni
Eastern Andes

Shorter Wing (1)Shorter Tarsus (1)Lower Mass
(1,2,3,4,5)

Lighter (4)Faster (1,2,4)Faster (1,2,3,4,5)
Lower (1)S. spillmanniCentral Andes

Shorter Wing (1,2)
Shorter Tail (1)Biometrics
Shorter Tarsus(d.f.=4/4/4/4/2)(1,2,4) Lower
Mass (1,2,3,4,5)
S. meridanus / S. fuscicauda

Plumage(d.f.=10)

Scold(d.f.=14/17)

S. spillmanni
Eastern Ecuador

S. spillmanni
Western Ecuador

Shorter Wing
(1) Shorter
Tarsus? Shorter
Bill?

Slower (1,2,4)Higher(1,2,3,4,5) Note
Shape (4)Slower(1,2,3,4,5)Higher(1,2,3,4,5) Note
Shape (4)Direct comparison not possible

Browner Back
(4)

Higher (1)S. g. griseicollis

Shorter Wing
(1,2,4) Shorter
Tail (1,2,4)
Shorter Tarsus?

Direct comparison
not possible

Slower (1,2,3,4,5)Higher (1,2,3,4,5)Note Shape (4)Slower (1,2)
Higher (1,2,3,4,5)Note Shape (4)Shorter Tail (4)Browner Back (4)Higher (1,2,4)

S. g. gilesi

Shorter Wing (1,2)
Shorter Tarsus?
Shorter Bill?

Grayer Back (4)Slower (1,2,3,4,5)Higher (1,2,3,4,5)Note Shape (4)Slower (1,2,4)Higher (1,2,3,4,5)Note Shape (4)

X

X

X

S. griseicollis(Santander-Tamásubsp.)

X

X

X

X

Longer Wing (1,2)
Longer Tarsus?
Longer Bill?

Browner Back (4)Faster (1,2,3,4,5)
Lower (1,2,3,4,5)Note Shape (4)S. meridanus /S. fuscicauda

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