taxonomia upucerthia

8/13/2019 taxonomia upucerthia

1/16

BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and res

funders in the common goal of maximizing access to critical research.

Natural History, Morphology, Evolution, and Taxonomic Status of the Earthcreeper

Upucerthia saturatior(Furnariidae) from the Patagonian Forests of South America

Author(s): Juan I. Areta and Mark Pearman

Source: The Condor, 111(1):135-149. 2009.

Published By: Cooper Ornithological Society

DOI: http://dx.doi.org/10.1525/cond.2009.080009

URL: http://www.bioone.org/doi/full/10.1525/cond.2009.080009

BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental

sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies,

associations, museums, institutions, and presses.

Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOnes Terms of

Use, available at www.bioone.org/page/terms_of_use.

Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and

permissions requests should be directed to the individual publisher as copyright holder.
http://www.bioone.org/page/terms_of_usehttp://www.bioone.org/http://www.bioone.org/doi/full/10.1525/cond.2009.080009http://dx.doi.org/10.1525/cond.2009.080009


2/16

135

The Condor111(1):135149 The Cooper Ornithological Society 2009

The Condor,Vol. 111, Number 1, pages 135149. ISSN 0010-5422, electronic ISSN 1938-5422. 2009 by T he Cooper Orn ithological Society. All rights reserved. Please directall requests for permission to photocopy or reproduce art icle content through the University of California Presss Rights and Permissions website, http://www.ucpressjournals.com/reprintInfo.asp. DOI: 10.1525/cond.2009.080009

Resumen. Upucerthia saturatiores un furnrido bien diferenciado que habita los bosques patagnicos delcentro-oeste de Argentina y Chile, dentro del rea de endemismo deNothofagus. Inicialmente descripta como unaespecie distinta en 1900, U. saturatiorfue rpidamente incluida dentro de U. dumetariasin una explicacin o estu-dio que lo avalaran; este tratamiento ha sido seguido por la mayora de los autores posteriores. Basados en la varia-cin geogrfica aparentemente clinal de U. dumetariay en un re-anlisis de la morfologa y plumaje de individuosque se supona representaban gradacin entre U. dumetariay U. saturatior, concluimos que no hay evidencia degradacin entre estos taxones. Upucerthia saturatiordifiere de U. dumetariaen canto (p-p-tirik-tirik-tirik-tirik-tirik-tiruk vs. pli-pli-pli-pli-pli . . .) y llamados (pep vs. kiip), morfologa (ms pequea y oscura, con piconegro y corto vs. ms grande y plida, con pico largo y marrn) y patrn de la cola, hbitat reproductivo (bordesde bosque vs. estepa arbustiva y habitats abiertos de altura) y patrn migratorio (trans-andino vs. nortesur). Es-tas diferencias son mayores a las que se encuentran entre U. jelskiiy U. albigula, y son mucho mayores que las queencontramos entre U. jelskiiy U. validirostris. Nuestros datos apoyan decididamente el estatus de especie plena

para U. saturatior. La existencia de una Upucerthiade bosque en parapatr a con una especie de ambientes abiertosprovee de una oportunidad para poner a prueba el papel de los cambios de hbitat entre ambientes ridos abiertosy ambientes boscosos durante la especiacin.

NATURAL HISTORY, MORPHOLOGY, EVOLUTION, AND TAXONOMIC STATUSOF THE EARTHCREEPERUPUCERTHIA SATURATIOR(FURNARIIDAE)

FROM THE PATAGONIAN FORESTS OF SOUTH AMERICA

Historia Natural, Morfologa, Evolucin, y Estatus Taxonmico de la Bandurrita Upucerthiasaturatiorde los Bosques Patagnicos de Amrica del Sur

Abstract The Patagonian Forest Earthcreeper (Upucerthia saturatior) is a distinctive furnariid that inhabitsthe Patagonian forests of central-western Argentina and adjacent Chile within the NothofagusCenter of Ende-mism. After its description as a species in 1900, U. saturatiorwas quickly subsumed, without comment or study,as a subspecies of the Scale-throated Earthcreeper (U. dumetaria), a treatment followed by most subsequent au-thors. On the basis of an apparent geographical cline within U. dumetariaand a reanalysis of the morphology and

plumage of reported intergrades between U. dumetariaand U. saturatior, there is no evidence of intergradationbetween these taxa. Upucerthia saturatiordiffers from U. dumetariaby its song (p-p-tirik-tirik-tirik-tirik-tirik-tiruk vs. pli-pli-pli-pli-pli . . .), which is also three times faster in dumetaria, call (pep vs. keep), morphol-ogy (smaller and darker with a short black bill vs. larger and paler with a long brown bill), distinctive tail pattern,

breeding habitat (forest borders vs. shrubby steppe and open highland habitats), and migration patterns (trans-

Andean vs. northsouth). These differences exceed those between U. jelskiiand U. albigulaand are far greaterthan those between U. jelskiiand U. validirostris; they overwhelmingly support ranking U. saturatioras a fullspecies. The existence of a forest-dwelling species of Upucerthiaparapatric to an open-country Upucerthiapro-vides an opportunity for testing the role of habitat shift between d ry exposed habitats and forest habitats (and viceversa) during speciation.

Manuscript received 29 August 2008; accepted 12 November 2008.4E-mail: [email protected]: [email protected]

JUANI. ARETA1,2,4 ANDMARKPEARMAN3,5

1CICyTTP-CONICET, Materi y Espaa, 3105 Diamante, Entre Ros, Argentina2Grupo FALCO, Calle 117 Nro 1725 e/67 y 68, 1900 La Plata, Buenos Aires, Argentina

3Aves Argentinas/Asociacin Ornitolgica del Plata, Matheu 1246/8 (C1249 AAB), Capital Federal, Argentina

INTRODUCTION

The ground-dwelling and curve-billed earthcreepers of thegenus Upucerthiahave recently been the focus of two phylo-genetic studies (Chesser et al. 2007, Fjelds et al. 2007). Both

Key words: Andes, evolution, natural history, Nothofagus forest, Patagonian Forest Earthcreeper,systematics, taxonomy, Upucerthia saturatior.

studies consistently showed that Upucerthiais the sister groupof Cinclodesand that the four species that should belong to Up-ucerthiaare the Scale-throated Earthcreeper (U. dumetaria,type species of Upucerthia), Buff-breasted Earthcreeper (U.validirostris), Plain-breasted Earthcreeper (U. jelskii), and


3/16

136 JUAN I. ARETA AND MARK PEARMAN

White-throated Earthcreeper (U. albigula). Additionally, theStriated Earthcreeper (U.serrana) was found to be basal tothe monophyletic group of Upucerthiaplus Cinclodesin oneof the studies (Fjelds et al. 2007), whereas it was found to

be distantly related to all of these furnariids in a phylogenysampling a broader range of taxa (Chesser et al. 2007). This

pattern is consistent with the lack of close relatives sharedby other bird species with the same relictual distribution ofU. serranain Peru (Fjelds et al. 2007) and suggests that the

position of U. serranaclose to Upucerthiaand Cinclodesre-

ported by Fjelds et al. (2007) should be interpreted as an arti-fact of insufficient taxon sampling. For the present, U. serranais provisionally retained in Upucerthia.

The species-level taxonomy of Upucerthia sensu strictoremains confusing, leading to differing treatments (Zotta1938, Esteban 1951, Meyer de Schauensee 1966, Vaurie 1980,Remsen et al. 2008). Geographical variation in U. dume-tariais not clearly understood. Historically, eight taxa (someof them originally described as species) have been includedwithin U. dumetaria(Geoffroy Saint-Hilaire 1832, Reichen-

bach 1853, Scott 1900, Chubb 1911, Chapman 1919, Zimmer1954), and between two and five subspecies have been recog-nized by various authors (Hellmayr 1932, Navas 1971, Vaurie1980, Fjelds and Krabbe 1990, Remsen 2003). Modern treat-ments acknowledge four subspecies, dumetaria, hypoleuca,

saturatior, andperuana(Remsen 2003). The distinction be-tween dumetariaand hypoleuca, however, frequently provesdifficult both with specimens and in the field (Navas 1971),and peruana is known from only two specimens (Zimmer1954). By far the most distinctive subspecies within U. dume-tariais saturatior(Fig. 1), originally described as a species(Scott 1900) and so treated occasionally (Chubb 1911, Esteban

1951). Nevertheless, this treatment has not been followed bymost authors, who have classified saturatioras a subspeciesof U. dumetaria(e.g., Chapman 1919, Hellmayr 1935, Navas1971, Vaurie 1980, Remsen 2003, Remsen et al. 2008). Evi-dence for either taxonomic treatment is weak, and more datafor a convincing rationale are necessary.

Despite its uniqueness and Cinclodes-like aspect,satura-tiorwas not explicitly included in the recent molecular phylo-genetic studies of Upucerthia(Chesser et al. 2007, Fjelds etal. 2007), hampering our understanding of the phylogenetic

relationships, biogeography, evolution, and alpha taxonomy ofUpucerthia.In this ar ticle we provide newly obtained data on the dis-

tribution, seasonal movements, morphology, habitat use, vo-calizations, and behavior of the little-knownsaturatiorfromthe province of Neuqun (Argentina), and we analyze previ-ously published data on morphology and distribution. Thesedata overwhelmingly suggest that saturatiorshould be ele-vated to the rank of species. We also discuss the evolutionaryand biogeographic importance of the existence of a Patago-nian forest-dwelling Upucerthia.

METHODS

We studied Upucerthia saturatioras follows: 28 December1999, 614 March 2000, 26 January 2007, 1115 January2008, and 616 March 2008 at La Fragua, Arroyo Ranqui-leo, Reserva Turstico Forestal Lagunas de Epulauqun, La-guna Vaca Lauqun, Paso Pichachn, Pino Hachado, 20 kmwest of Rahue, and 10 km west of San Martn de los Andes,all in western Neuqun province, west-central Argentina (seeFig. 2). For each field observation we took notes on behavior,

FIGURE 1. A, Adult Patagonian Forest Earthcreeper (Upucerthia saturatior) photographed at Lagunas de Epulauquen, Neuqun, Argen-

tina. Note the distinctive short black bill, dark dorsal coloration, and indistinct breast markings. Photo: Juan I. Areta. B, Adult Scale-throatedEarthcreeper (Upucerthia dumetaria) photographed at Sierra Baguales, Santa Cruz, Argentina. Note the distinctive long bill, lighter dorsalcoloration, large undertail markings, and distinct breast markings. Photo: Santiago Imberti.


4/16

SYSTEMATICS AND NATURAL HISTORY OFUPUCERTHIA SATURATIOR 137

habitat, plumage, and the color, shape, and size of the bill. Werecorded vocalizations with a Sennheiser ME-62 microphonein a Telinga Universal parabola and a Sony TCD5-ProII taperecorder and with a Sennheiser ME-67 microphone and aSony TCM 5000 tape recorder. Recordings made by Areta aredeposited at the Macaulay Library of Natural Sounds (CornellLab of Ornithology), those by Pearman will be deposited inthe British Library (National Sound Archive). To determinequalitative differences in the vocalizations ofsaturatiorfromthose of other species of Upucerthia, we examined our record-ings aurally and then visually with the aid of spectrograms

built with Syrinx 2.6h (www.syrinxpc.com, John Burt). Weexamined 49 recordings of all the currently recognized spe-cies of Upucerthia(includingserrana) and of three of the fourcurrently recognized subspecies of U. dumetaria (nominatedumetaria, hypoleuca, andsaturatior; there are no known re-cordings of subspeciesperuana ; Appendix 2).

We carried out eight playback experiments betweenlate September and mid October 2008 (early breeding sea-son): three in El Chocn (Ro Negro, Argentina), three inEl Porvenir (Tierra del Fuego, Chile), and two in Farellones(Regin Metropolitana, Chile). All the experiments tested the

FIGURE 2. Distribution and migration patterns of the Patagonian Forest Earthcreeper (Upucerthia saturatior) and Scale-throated Earth-creeper (U. dumetaria) in Argentina and Chile. A, Distribution of Upucerthia dumetariaand U. saturatiorin Argentina and Chile. Whitecircles, U. saturatior; black circles, U. dumetaria; gray circles, possible areas of breeding parapatr y of dumetaria /saturatiorbased on speci-mens [16/112, 2022/114115, 26/118, 31/121, 67/117]. Numbers: dumetaria, 144; hypoleuca, 4564; saturatior, no numbers, see B fordetails. B, Details of the distribution of U. saturatiorin Argentina and Chile. White circles, U. saturatior; gray circles, dark specimens ofU. dumetariapresumed to be intergrades dumetaria/saturatiorby Navas (1971). Numbers:saturatior, 74121; dark dumetaria, 6573. SeeAppendix 3 for names and coordinates of localities. In nominate dumetaria, southern populations migrate to the pampas of Argentina duringwinter. Insaturatior, while most northern and some coastal Chilean records fall from April to September, some southern Chilean and mostArgentine records fall from September to March. These highly divergent migration routes of U. saturatiorand U. d. dumetariaenhance theirspatial segregation and argue for species status of both forms.


5/16

138 JUAN I. ARETA ANDMARK PEARMAN

response of U. d. dumetaria/hypoleucato voices of U. d. du-metaria/hypoleucaandsaturatiorduring the breeding season(the response ofsaturatiorto voices of dumetaria/hypoleuca

was not tested by this method). Each experimental individ-ual of U. dumetariawas exposed to three stimuli: one songof U. d. dumetaria, one song of hypoleuca, and one song of

saturatior. The order of stimuli was randomized to avoid se-quence effects, and the songs were sampled from our database(see Appendix 2) to avoid pseudoreplication (Kroodsma 1986,1989a). We categorized the results of each experiment as si-lent approach, vocalization without approach, and approachwith vocalization.

We examined and measured 35 specimens of U. d. du-metaria, 21 specimens of hypoleuca, and 45 specimens of

saturatiordeposited at the Fundacin Miguel Lillo, Tucumn(FML), Museo Argentino de Ciencias Naturales, Buenos Ai-res (MACN), and Museo de La Plata, La Plata (MLP) (Appen-dix 1). Bill length (exposed culmen) and tarsus length weremeasured to the nearest 0.05 mm with digital calipers, andflattened wing and tail length were measured with a metalruler to the nearest 0.5 mm. We also used distributional in-formation from specimens at the Field Museum of NaturalHistory, Chicago (FMNH), Louisiana State University, Ba-ton Rouge (LSU), and Yale Peabody Museum, New Haven(YPM) (Appendix 3).

RESULTS

DISTRIBUTION AND MOVEMENTS

Distribution.In Argentina, U. d. saturatior is known fromnorthwestern Neuqun, western Ro Negro, and northwest-ern Chubut provinces (Zotta 1938, Esteban 1951, Navas 1971,Fig. 2). Specimen and sight records range from 250 to 1800 mabove sea level and are restricted to Patagonian forest bordersin the Andes of central Chile and central-western Argentina(this Andean form is unknown from the Patagonian steppe,seeHabitatbelow). We recorded U. d. saturatiorat 10 locali-ties in western Neuqun province (Fig. 2). The closest distancefrom U. d. saturatiorat which we could find U. d. dumetaria/hypoleucawas ca. 25 km (saturatiorat Epulauquen and du-metaria/hypoleucaat Las Ovejas). However, U. d. dumetariahas been also reported at Epulauquen (Veiga et al. 2005), anidentification which, if correct, would indicate that both U. d.

saturatiorand U. d. dumetariacan be found at the same lo-cality. Additionally, Navas (1971) reported specimens of U.d. saturatiorand U. d. dumetariacollected as close as 4 km

apart, and specimens of both subspecies have been collectedat Nahuel Huapi (saturatiorMACN-8389 67ae and dume-tariaMACN-8389). Their geographic ranges may be parap-atric or with minor overlap on a wide geographic scale (seeEsteban 1951 and Navas 1971, Fig. 2, and Appendix 3 for otherlocalities of possible overlap). Sympatry, however, does notimply syntopy (seeHabitatbelow).

The absence of U. d. saturatiorfrom seemingly suitablehabitat further south of its known restricted distribution inArgentina and Chile is remarkable and remains a mystery.

This area is poorly studied, however, and possiblysaturatiorhas been overlooked there. From minimal data Navas (1971)estimated the breeding season of saturatior in Argentina asSeptember to February; the breeding season could extend intoMarch (pers. obs.). Hellmayr (1932) reported recently fledged

birds in central Chile in March; however, Zimmer (1954)clearly demonstrated that Psslers observations ofsaturatior

breeding on the coast of Chile, quoted by Hellmayr (1932),were based on misidentifications (very likely of the coastalCinclodes nigrofumosus). The lack of definite breeding re-cords of U. d. saturatiorin Chile and the abundance of recordsduring the nonbreeding period could suggest that saturatior

breeds exclusively in Argentina and not in Chile. Some re-cords from the Andes of Chile, however, might refer to breed-ing birds (see Fig. 2, Appendix 3).

Movements.It has been suggested that U. d. saturatiorcould migrate north in Argentina and Chile during winter(Hellmayr 1932, Goodall et al. 1946, Johnson 1967, Vau-rie 1980). All specimens and sight records from Argentina,however, are from September to March, and most specimensfrom Chile are from late March to August (Hellmayr 1932,

Navas 1971, Fig. 2, Appendix 1, Appendix 3). We suspect,as did Navas (1971), that U. d. saturatior is a trans-Andeanmigrant because of (1) the geographic and temporal distribu-tion of specimens, (2) the mapping by Jaramillo (2003) of anexclusive winter range (unspecified to subspecies), north ofthe regular Chilean range ofsaturatior, in dry deciduous for-est and matorral along the coast of central Chile between thelake district and Valparaiso. Furthermore, Hellmayr (1932)

mentionedsaturatioras only a winter visitor in the Angol andNilahu valleys of Chile, at the same latitude at which birdscan be found breeding in Argentina but across the Andes. De-spite this, Jaramillo (2003) mapped the Angol Valley as partof the resident range of U. d. saturatior. This trans-Andeanmigration pattern is shared by other species (e.g.,Phytotomarara) and differs strongly from the well-known migration un-dertaken by nominate dumetaria, from the arid Patagoniansteppe north and northeast principally to the pampas region(Fig. 2, Navas 1971, Fjelds and Krabbe 1990). Outside of the

breeding season,saturatiorand dumetariaare segregated forat least five months, since there is no overlap in Chile.

HABITAT USE

In the province of Neuqun we found U. d. saturatiorat forestborders, primarily close to bodies of water but also away fromthem, both during the breeding season and at the onset of fallmigration. All our observations were at the edges of both largeand small patches ofNothofagusforest, at a clearing at least1.5 km insideNothofagusforest, or at the edge of a denselywooded mosaic of rivers and pools with introduced Salixand


6/16


other trees (some possibly native) but withoutNothofagus. Al-though surrounding vegetation was almost always Nothofa-

gusforest, the birds were never found inside the forest. At the

same time, during the breeding season, we found several ter-ritories of U. d. dumetaria/hypoleuca(e.g., one pair singingand holding territory at Laguna Varvarco Campos on 14 Janu-ary 2008), and during mid-March 2008 (probably just priorto migration), we noted ten individuals (five pairs) of U. d.dumetaria/hypoleuca, all in dry areas, without surroundingforest, in habitats of Monte desert and Patagonian steppe inwestern Neuqun.

Navas (1971) mentioned the collection, in February, of anominate dumetariaat Confluencia Traful Limay and of a typ-icalsaturatiorin Valle Encantado del Ro Limay. These twolocalities are only 4 km apart and have the same mixed habitatdominated by subantartic forest intermixed with Patagoniansteppe. Although syntopy of the two forms while breeding is

possible, it has yet to be confirmed. In the Patagonian Andeanforest region open habitats are always close to forests. There-fore, it is almost impossible to determine the precise habitat inwhich specimens of U. dumetariawere collected from localitydata on older specimen labels only. Thus, for example, birdscollected around San Martn de los Andes cannot be safelyattributed to either steppe or forested habitats.

MORPHOLOGY

General morphology.The most detailed description of satu-ratiorin the literature is by Hellmayr (1932:165), who wrote,U. d. saturatiordiffers at a glance from the other races byvery much darker, olive or sepia brown, upper par ts and mid-dle tail feathers without paler tips to the wing coverts; tawnyrather than cinnamomeous basal portion of remiges; gener-

ally wider, deeper cinnamon rufous tips to lateral rectrices;much duller, avellaneous rather than buffy, under parts withthe blackish margins on foreneck and chest much more pro-nounced, and the flanks dark buffy brown or olive-brown.Besides, the bill is shorter, stouter, and more blackish. Thisdescription is an accurate synthesis, although we contest thesuggestion thatsaturatiorhas breast scaling more pronouncedthan in the other races (Fig. 1, Table 2). To the contrary,satu-ratiorshows broader, often brown (not black) scaling, usuallyrestricted to the upper breast, creating a more diffuse and lessconspicuous scaled pattern than in U. d. dumetaria/hypoleuca(Fig. 1). Upucerthia d. saturatioris therefore distinctly darker.It is also smaller with a shorter (distinctly black, not brown)

bill than in other subspecies of U. dumetaria, and its tail is also

proportionately shorter, with a distinct pattern (Fig. 1, Tables1 and 2, Hellmayr 1925, 1932, Zotta 1938, Esteban 1951, Zim-mer 1954, Navas 1971, Vaurie 1980). These differences areeasily noticed in the field. At first glance, U. d. saturatiorlooks intermediate between U. d. dumetariaand one of thedark species of Cinclodesof Patagonia (e.g., C. patagonicus)in overall shape, size, and plumage. For example, a foraging

U. d. saturatiorseen facing away could easily be mistakenfor a Cinclodes, a scenario most unlikely with any other spe-cies of Upucerthia. This similarity is interesting (although not

unexpected) given the close relationship between Cinclodesand Upucerthia(Chesser et al. 2007, Fjelds et al. 2007, seeBehavioral notesbelow).

We assume, from the examination of numerous speci-mens, that juveniles of both U. dumetaria/hypoleucaand U.d. saturatiorhave whitish shaft streaks on the nape and up-

per back, are more extensively scaled on the breast, and showbroken brown barring on the lower abdomen, especially atthe sides. Likewise, immatures may retain a few pale nuchalstreaks but always show some barring on the lower abdomen,which is lacking in adults. In all of these plumages,saturatiormaintains its distinct tail pattern (as mentioned by Hellmayr1932; see above), which allowed us to immediately distinguishyoung specimens ofsaturatiorfrom U. dumetaria/hypoleuca(seeEvidence of Intergradationbelow). It is noteworthy that

juveniles of U. d. dumetaria/hypoleuca, like adults ofsatura-tior, have black bills. Measurements of juvenile dumetariaav-erage larger than those of juvenilesaturatiorand are similarto those of adultsaturatior(Table 1).

Ecomorphological variation and clines. Most species ofUpucerthiaare believed to fit Glogers rule (i.e., birds inhab-iting more humid regions have darker pigmentation; Remsen2003). The northern subspecies U. dumetaria hallinaniwassynonymized with the more southern U. dumetaria hypoleucaon the basis of weak morphological differentiation, local vari-ation in color intensity, and the presence of paler individu-als toward drier areas (Hellmayr 1932, Esteban 1951, Johnson1967). All authors agree that U. d. dumetariaand U. d. hypo-leucaare very similar (Navas 1971, Remsen 2003, Jaramillo

2003). Pigmentation varies greatly, regardless of sex, in a se-ries of nine adult specimens of hypoleuca taken in August1898 (one probably in 1897) at an unspecified location in SanLuis province (MLP-425 through 433). Although some indi-viduals appear indistinguishable from nominate dumetaria,others show a slightly paler and more rufescent dorsal color-ation and paler ventral coloration suggesting U. d. hypoleuca.A specimen from Ro Seco, Concarn (also from San Luis

province), taken in mid-May is referable to nominate dume-taria(MLP-10734). Zotta (1938) argued that hypoleucawasthe form intermediate between hallinaniand dumetaria, andHellmayr (1932) suggested that hallinanishould probably bemerged with hypoleuca. Both authors indirectly argued thatthe variation is clinal. We suspect that this clinal variation,

combined with strong individual variation, might have beenthe basis for Vauries (1980) decision to merge all subspeciesof dumetaria(exceptsaturatior) under nominate dumetaria(see Table 2). The darker individuals of dumetariaclose tothe Andes add another dimension of complexity to the vari-ation within dumetaria (seeEvidence of intergradationbe-low). Also, the large, dark subspecies peruana, known only


7/16


(continued)

TABLE 1. Measurements of bill length, tail length, and wing chord of the Scale-throated Earthcreeper (Upucerthiadumetaria dumetaria, U. d. hypoleuca) and Patagonian Forest Earthcreeper (Upucerthia saturatior) according to variousauthors and specimens listed in Appendix 1. Values are given in mm as mean SD with sample size in parentheses andrange in brackets, if available. Measurements of specimens suggested by Navas (1971) to be intergrades appear under dark

adults. The measurements support full species status for U. saturatiorand lack of evidence of intergradation between U.saturatiorand U. dumetaria. Note the consistent size differences between juvenile and adults of both forms.

U. saturatior U. d. dumetaria U. d. hypoleuca U. d. peruana Source

Bill length29.1 0.9 (17)

[2831] 35.7 1.6 (15)

[3437.5]Hellmayr 1932a

33.05[30.835.4]

35.55[3338.6]

38.2[34.441.5]

Esteban 1951

27.9 (22)[2529]

31.8 (34)[2935]

32.95 (16)[2935]

Navas 1971

34 (10)[3137]

38.5 (10)[3543]

lumped withdumetaria

Vaurie 1980

30.5 1,3 (12)[28.531]

35.82.8 (5)[3138.5]

Zotta 1938b

35 Zimmer 1954

Adults28.37 1.08

(33)[26.8230.56]

Juveniles23.773.13 (4)[19.4526.38]

Adults31.57 1.49

(17)[30.4134.7]Juveniles

30.8 1.1 (5)[29.2430.95]Dark adults33.9 2.1 (3)[32.2936.19]

Dark juveniles28.6 2.5 (6)[24.9130.37]

Adults32.98 2.15

(17)[27.5234.48]

Appendix 1

Tail length79.9 2.3 (17)

[7784] 83.03.0 (15)

[7788]Hellmayr 1932a

74.85

[69.578.7]

79.65

[75.586.2]

83.6

[76.987.9]

Esteban 1951

78.2 (23)[7381]

83.4 (35)[8093]

83.8 (20)[7990]

Navas 1971

83.6 4.6 (12)[7786]

88.2 3.5 (5)[8493]

Zotta 1938b

85 Zimmer 1954Adults

72.25 3.02(38)

[6677.5]Juveniles

69.38 3.44 (7)[68.578.5]

Adults79.37 3.74

(19)[7384.5]Juveniles

75.8 2.6 (6)[7380]

Adults78.03 4.11

(20)[67.583]

Appendix 1

Dark adults79.7 6.1 (3)

[7385]Dark juveniles

75.8 3.0 (6)[70.578.5]


8/16


from two specimens, seems to fit the pattern of ecomorpho-logical variation with humidity (Table 1). Clearly, more studyis needed to ascertain whether hypoleucaandperuanaare di-agnosable subspecies and to elucidate patterns of genetic andmorphological geographical variation in the U. dumetariacomplex.

The comparatively dark color and small size of U. d. satu-ratior, of forest borders, compared to the larger U. d. dumetaria,

of brushland, parallels the color differences between U. albig-ulaand U. jelskii(Schulenberg 1987). In northern Chile U. al-bigulainhabitsPolylepiswoodlands and shrubbery along rivercourses, whereas the rather similar U. jelskii inhabits driershrub-steppe and river beds with sparser and lower shrubs(pers. obs., Schulenberg 1987). The two can be found within500 m of one another along the same river in Putre, Chile (pers.obs.), where albigulais slightly darker above, and their songs

TABLE 1. (Continued).

TABLE 2. Summary of features distinguishing the Patagonian Forest Earthcreeper (Upucerthia saturatior) and Scale-throatedEarthcreeper (Upucerthia dumetaria).

U. saturatior U. dumetaria/hypoleuca

Geographic variation None or minor Apparent northto-south clineBreast scaling Mild Strong

Dorsal color Very dark brown Variable, from ferruginous (N) to light brown (S),and dark brown (indiscriminate distribution)

Tail pattern Reduced chestnut terminal markings Large light terminal markingsMeasurements (Table 1) Smaller Larger Habitat Forest borders Open steppeFall /winte r migration Trans-Andean (Argentina to Ch ile) South to nor theastSong Syncopated series of couplets or triplets Continuous series of single notesPlayback experiments Not tested Ignores voice ofsaturatior, strong response to own voice

U. saturatior U. d. dumetaria U. d. hypoleuca U. d. peruana Source

Wing chord99.6 3.4 (17)

[95106] 101.03.4 (15)

[95106]Hellmayr 1932a

97.45[92.2104.9]

99.85[94.4109.2]

100.85[96.2108.5]

Esteban 1951

99.7 (23)[95106]

101.4 (35)[95109]

102.5 (20)[96110]

Navas 1971

95.6 2.7 (12)[91101]

101.3 3.5 (5)[98106.5]

Zotta 1938b

108 Zimmer 1954Adults

96.82 3.47(32)

[92.5103.5]Juveniles

91.903.30 (7)[91.5100]

Adults101.23 4.37

(20)[93111]

Juveniles99.2 1.8 (6)

[96.5101]

Dark adults100.0 0.5 (3)[99.5100.5]

Dark juveniles97.0 3.0 (6)[94101.5]

Adults101.48 4.51

(21)[92110.5]

Appendix 1

Tarsus length 26.5 Zimmer 1954

Adults24.42 0.68 (6)[23.4125.31]

Adults26.02 0.96 (8)[24.4427.4]

Adults25.96 0.93 (10)

[24.5727.71]

Appendix 1

aHellmayr (1932) referred these measurements to hallinani, but we include them with hypoleuca.bZotta (1938) measured hallinaniand hypoleucaseparately, here measurements are pooled according to modern taxonomicuse. See text for discussion of clinal var iation.


9/16


differ in pitch and speed. A cline involving both plumage andmeasurements has been suggested to link U. validirostrisandU. jelskii(Vaurie 1980, Mazar Barnett et al. 1998), although

they are currently treated as separate species (Remsen 2003).Hellmayr (1925) suggested a similar explanation for increasedpigmentation in relation to habitat humidity in U. dumetariaand in a number of other Chilean furnariids that inhabit dryand open areas, such as the Common Miner (Geositta cunic-ularia), Crag Chilia (Chilia [Ochetorhynchus] melanura),Plain-mantled Tit-Spinetail (Leptasthenura aegithaloides),and Cordilleran Canastero (Asthenes modesta).

Evidence of intergradation. Although U. d. hypoleucaofnorthwestern Argentina ranges south to the steppe region ofcentral Neuqun (Navas 1971; 3 MACN specimens), no inter-mediates between U. d. hypoleucaand U. d. saturatiorhave

been reported. The reference by Navas (1971) to supposedlyintermediate individuals between saturatior and dumetariaclose to the Andean forests is confusing and inconclusive. Onone hand, he mentioned widely scattered specimens from LaPampa, Buenos Aires (both far from the Andes), Ro Negro,and Chubut that could be regarded as dark (toward sepia) in-dividuals of U. d. dumetaria. On the other hand, he later con-sidered that Andean specimens of three dark juveniles (twofrom orquinco and one from El Bolsn in western Ro Ne-gro) and two dark adults from El Maitn (western Ro Negro),whose bill measurements (34 and 35 mm) are well outside hisrange forsaturatior, provided evidence of intergradation be-tweensaturatiorand dumetaria(Fig. 2). Similarly, four darkAndean birds from Cushamen, northwestern Chubut (oneadult and three juveniles), and one adult from Alto Ro Sen-guerr, southwestern Chubut (both adults with bill lengths of32 and 34 mm, typical of dumetaria) were also considered in-

tergrades between dumetaria and saturatior (Fig. 2). How-ever, (1) dark individuals seem to occur in various parts of therange of U. d. dumetariawithout U. saturatior(e.g., widelyscattered localities in Buenos Aires, La Pampa, and Ro Ne-gro provinces;fideNavas 1971, MACN specimens), and paleindividuals of U. d. dumetariaalso seem to occur without aconsistent geographic pattern (e.g., several localities in BuenosAires, La Pampa, San Luis, and Ro Negro fideNavas 1971,MACN specimens). (2) No intermediate measurements have

been reported for U. d. dumetariacloser to the Patagonian for-ests because bill measurements of juveniles cannot be directlycompared to those of adults and measurements of purportedlyintermediate individuals fall within the range of U. d. dume-taria, being larger than those of U. d. saturatior. (3) There may

be confusion regarding the significance of dark birds close tothe Andean forests; these birds might exemplify variation ac-cording Glogers rule within a species (U. dumetaria) indepen-dent of the existence of a darker species (U. saturatior), notnecessarily intergradation with U. saturatior. Additionally,Alto Rio Senguerr (one of the localities with birds presumedintermediate between U. d. saturatiorand U. d. dumetaria) is

outside of, and a long distance from, the known distribution ofU. d. saturatior(Fig. 2). We examined the supposedly inter-mediate specimens mentioned by Navas (1971) and found that

they are dark variants of dumetaria. Most are juveniles (withmeasurements larger than those of juvenilesaturatiorbut closeto those of adultsaturatior, Table 1). Others are adults, refer-able to dumetariabecause their measurements are well outsidethe range ofsaturatior (Table 1). Although these individualsare indeed darker than some of dumetaria, others of dumetariafrom distant locations are similarly dark, and all have the tail

pattern typical of dumetariaand distinct from that ofsatura-tior(see General morphologyabove, see Table 2).

VOCALIZATIONS

We tape-recorded three different vocalizations of U. d. sat-uratior(n8 indviduals): the song (and subsongs partialsongs), a continuous song, and one contact call. The song ofU. d. saturatiordiffers structurally from the apparently ho-mologous song of U. d. dumetaria. The complete song of U.d. saturatiorconsists of one or two sharp, short, and lower-

pitched initial notes followed by either a syncopated or acontinuously delivered series of triplets, at three per second,each of which starts with one fast metallic, abrupt, and almostvertical note followed by two shorter, flatter, longer variablyv-shaped or shallowly u-shaped notes (only the latter two dis-tinguished by the human ear). Occasionally, it ends with a dis-tinct triplet including a lower-pitched final note (Fig. 3A). Acomplete version of the song of U. d. saturatiorcan be de-scribed as p-p-tirik-tirik-tirik-tirik-tirik-tiruk (Figs. 3A,B). Both the soft introductory ps and the disyllabic tiriks(triplets) vary in number. After playback, the song is faster,and an extra u-shaped note may be added to the triplets (Fig.

3C). Birds heard along forest borders around San Martn delos Andes, towards Paso Hua-Hum, were not tape-recorded

but sounded identical to birds in other localities of westernNeuqun province. In contrast (n13 individuals), the songof U. d. dumetariais a series of fairly homogeneous similarlyshaped notes delivered in rapid succession (Figs. 4A, B), withnine notes per second (i.e., three times faster than in satura-tior), and again the precise number of notes is highly variable.The song of hypoleuca(n7 individuals) appears indistin-guishable from that of nominate dumetaria(Fig. 4C). A com-

plete version of the song of U. d. dumetaria/hypoleucacan bedescribed as pli-pli-pli-pli-pli . . . which increases slightlyin pitch, sometimes descending toward the end (Figs. 4AC).Bothsaturatiorand dumetaria/hypoleucaalso sing a form of

continuous song in which successive songs are linked bymany rapidly delivered notes that gradually change in pitch(Figs. 3D, 4D). The continuous song of U. d. dumetaria/hypo-leucaand U. d. saturatiorwas given by agitated birds, eithernaturally excited during territorial encounters or after play-

back (this type of continuous song is present at least also inU. jelskiiandU. validirostris). Both, the song and the continuous


10/16


song, can be performed in conjunction with the wing-raisingdisplay. The contact call of U. d. saturatioris a hollow pep,

peaking at just over 4 kHz, uttered alone or as a series of notesspaced with intervals of ca. 1.5 to 3 sec between successivenotes (Figs. 3EG). The contact call of U. d. dumetariais asofter and squeaky sounding keep, peaking at just over 6kHz (Fig. 4E).

Previously published descriptions of the voice of U. du-metariahave not specified the subspecies or locality on which

they were based and do little to aid our understanding of vo-calizations in the complex. For example, Fjelds and Krabbe(1990) wrote, Song a rapid chippy chippy chippy chip. Calla wheezy, abrupt keet (p. 331), to which Remsen (2003:247)added a sharp, dry dzit. Jaramillo (2003:158) wrote, A se-ries (1.53 sec) of scolding, ringing, syncopated notes givenat a speed just fast enough to be uncountable. Usually risesin pitch in middle and trails off at end: chwip chwip chwip

chwip chwip chwiip chwiip chwip chp chp, none of which weare able to identify to a taxon. Ridgely and Tudor (1994:31)stated, The song in cen. Chile is a spritely, rather musical tr,tr, trreetrreetr reetrrritrritrri with a slightly ascending effect,and Canevari et al. (1991:301) described a harsh voice whichis emitted while perching in bushes Cli cli cli cli . . . , whichundoubtedly refers to U. d. dumetaria/hypoleuca.

FIGURE 3. Spectrograms of songs and calls of the Patagonian For-est Earthcreeper (Upucerthia saturatior). A, Song of U. saturatior

(13 March 2008, Epulauquen, Neuqun, JIA 1); B, song of U. satu-ratior(13 March 2008, Epulauquen, Neuqun, JIA 2); C, song of U.saturatiorafter playback (January 2007, west of Rahue, Neuqun,MS/MP A6); D, continuous song of U. saturatiorafter playback(8 December 1999, Pino Hachado, Neuqun, MP B38/39); E, calls ofU. saturatior(8 December 1999, west of Pino Hachado, Neuqun, MPB34/35); F, call of U. saturatior(7 December 1999, Pino Hachado,

Neuqun, MP B29); G, call of U. saturatior(13 March 2008, Ep-ulauquen, Neuqun, JIA 3). Time scale reduced by 6 in Fig. 3Dand augmented by 3 in Figs. 3F and 3E to show details. Voices of U.

saturatiordiffer markedly from those of U. dumetariaand supportfull species status for U. saturatior.

FIGURE 4. Spectrograms of songs and calls of the Scale-throatedEarthcreeper (Upucerthia dumetaria/hypoleuca). A, Song of U. d.dumetaria (November, Pennsula Valds, Chubut, from Straneck1990); B, song of U. d. dumetaria(22 October 2004, Ea. La Queren-cia. Santa Cruz, S. Imberti); C, song of U. d. hypoleucaafter play-

back (1 October 2001, El Infierni llo, Tucumn, L. Macaulay, MLNS115989); D, continuous song of U. dumetariaafter playback (28 Sep-tember 2008, El Chocn, Ro Negro, JIA); E, call of U. d. dumetaria(5 February 1984, Laguna Los Escarchados, Santa Cruz, N. Krabbe,XC 15997). Time scale reduced by 2 in Fig. 4C and by 6 in Fig. 4D toshow overall pattern , and augmented by 3 in Fig. 4E to show details.Voices of U. dumetariadiffer markedly from those of U. saturatiorand support full species status for U. saturatior.


11/16


Songs and calls of nominate dumetaria and hypoleucadiffer from those ofsaturatiorin note shape, pattern, and paceand are consistently distinctive to the human ear (Figs. 3, 4).

Our data on the vocalizations of U. d. saturatiorspan most ofits known distribution in Argentina and are not limited to asmall geographic area (see Remsen 2005). Geographic varia-tion in voice within a species is relatively small in the Furnari-idae (e.g., Zimmer 1997, 2008), and we do not expect voicesof U. d. saturatiorat other localities outside Neuqun to differsubstantially from our recordings.

For all eight playback experiments the response was iden-tical: individuals of U. d. dumetaria/hypoleuca answeredstrongly to voices of U. d. dumetariaand U. d. hypoleuca, ap-

proaching and vocalizing to the sound source, regardless ofthe order of stimuli. Likewise, all individuals of U. d. dume-taria/hypoleucatested ignored the voices of U. d. saturatior.On the other hand, our uncontrolled experiments with U. d.

saturatiordemonstrate that they answer to the playback oftheir own voices, unlike U. d. dumetaria/hypoleuca.

BEHAVIORAL OBSERVATIONS

Upucerthia dumetaria saturatiorforaged exclusively on theground near tall trees, around small bushes in sandy soils,on boggy ground close to forest edge, and in forest clear-ings by vigorously and rapidly probing its bill into the groundobliquely (video available at ibc.lynxeds.com). Occasionally,it removed large clumps of earth and stones with the aid of thefeet or pushed them aside with lateral bill movements. Aftersome minutes of foraging, the ground was fur rowed, the birdsleaving several small holes with mounds around their borders.While foraging, members of a pair frequently gave their con-tact call (Figs. 3EG).

Between foraging bouts, U. d. saturatior commonlyperches in trees, unlike other forms of U. dumetaria, to preenand to sing (thus making important use of an element not pres-ent in the habitat of U. dumetaria; pers. obs., Johnson 1967).A territorial pair of U. d. saturatiorstudied in a clearing 1.5km inside Patagonian forest dominated by Lenga (Nothofagus

pumilio), Pehun (Araucaria araucana), and ire (Nothofa-gus antarctica) spent much of their time perching in two deadtrees ofN. antarctica, where they preened and rested, withone bird (presumably the male) singing 24 m up, sometimesaccompanied by a wing-rotating display, usually from the ex-tremity of dead branches. Most prey was consumed on theground, although on one occasion a bird returned to one of thefavored trees to consume a large grub. When delivering con-

tact calls, the birds shifted its tail from side to side or flickedit up and down.

Once, a single bird also used a large unidentified deadtree over marshy ground with some standing water for rest-ing and preening before and after terrestrial foraging. Thisindividual sang and displayed 7 m up in a live N. antarcticatree, but also from nearby fence posts. In contrast, the song

of U. d. dumetaria/hypoleucais delivered from bushtops. Inthe height of the breeding season birds begin singing from a

bush and then launch themselves downslope while singing in

flight.Another U. d. saturatiorwas observed foraging on theground when an immature Chilean Hawk (Accipiter chilen-

sis) landed noisily in a Roble Pelln (Nothofagus obliqua) treeca. 30 m from the bird, upon which two White-throated Tree-runners (Pygarrhichas albogularis) started calling excitedly.The U. d. saturatior immediately adopted a frozen posture,

positioning its legs to the side of the body and flattening itsbody toward the ground. At this moment we could approachthe bird to ca. 1.5 m while it remained frozen before eventu-ally flying off (video available at ibc.lynxeds.com).

In flight, U. d. saturatior exhibits fast wingbeats, fastturns, lateral movements of the body, and abrupt takeoffs thatare characteristic of the true Upucerthiaand Cinclodes, sup-

porting its placement in this group. Wing-raising and wing-rotating displays in Upucerthiaand Cinclodesare also similar

but are not present in any other furnariid genus of which weare aware.

DISCUSSION

TAXONOMIC REMARKS

Upucerthia dumetaria saturatior was originally describedas Upucerthia saturatiorby Scott (1900). Soon after, Chap-man (1919) listedsaturatioras a subspecies of U. dumetaria,under the misspelled name Upucerthia dumetoria saturatior,without providing any rationale. Most subsequent authors(Hellmayr 1925, 1932, Wetmore 1926, Peters 1951, Meyerde Schauensee 1966) have followed Chapmans classifica-tion without providing an explicit explanation for adoptingthe modified taxonomy. It is of interest to note that Upucer-thia tamucoensisChubb, 1911 (presumably a typographicalerror in the original description for temucoensis), was also de-scribed as a full species, although thereafter Hellmayr (1925)and Wetmore (1926) considered it a synonym of U. dumetaria

saturatior (i.e., both Scott and Chubb ascribed species sta-tus tosaturatior). Since then, even the most conservative ap-

proaches have retainedsaturatioras a subspecies within U.dumetaria(e.g., Hellmayr 1932, Vaurie 1980). Notwithstand-ing, various works (e.g., Fjelds and Krabbe 1990, Remsen2003, Jaramillo 2003, Kovacs et al. 2006) have illustrated

saturatior, desirable in aiding its field identification, given itsstriking differences from U. d. dumetaria.

Zotta (1938:5556), who treated U.(jelskii)pallidaasa subspecies of U. validirostris, wrote, The dif ferences of-fered by saturatiorcompared to the rest of the dumetariasubspecies are so noteworthy that they make one suspect itis a distinct species and went on to say the established tri-nominal division does not appear to be very justified. Re-flecting his opinion, Zotta (1938, 1944) twice referred to the


12/16


13/16


BIOGEOGRAPHY AND EVOLUTION

The recognition of U. saturatioras a distinct species providesinsight into speciation patterns and endemic bird areas in the

Patagonian forests. Cracraft (1985) did not include U. d. sat-uratioramong the endemic birds of the Nothofagus Center.The distributional pattern of U. saturatior, however, is con-sistent with this center of endemism and should be consideredas belonging to it (see Vuilleumier 1985, 1991 for mappingof Patagonian habitats). In his study of the ecological geog-raphy, speciation, and endemism of Patagonian forest birds,Vuilleumier (1985) did not include U. d. saturatioras a for-est form. This can be fully appreciated in his Appendix I (p.289), where U. dumetaria is listed only as a nonforest spe-cies. He also did not mention Upucerthiain his survey of spe-ciation in Patagonian birds (Vuilleumier 1991). Hence, nodiscussion is provided as to the evolutionary meaning of the

presence of forms of forest (saturatior) and dry open habitats

(dumetaria /hypoleuca) within the species complex. Pygar-rhichas albogularis, a Nothofagus Center forest endemic, hasbeen found to be closely related and phylogenetically basal toa group of species living in dry open habitats, the Crag Chilia(Chilia melanura), Band-tailed Earthcreeper (Eremobius

phoenicurus), and two species formerly classified in Upuc-erthia, the Straight-billed Earthcreeper (Ochetorhynchus ru-

ficauda) and Rock Earthcreeper (O. andaecola) (Chesser etal. 2007, Fjelds et al. 2007). This divergence has been asso-ciated with the creation of dry open habitats during the An-dean uplift in the Miocene (Fjelds et al. 2007). In the broadercontext of repeated shifts from forested to open habitats in theFurnar iidae (Fjelds et al. 2005, 2007), the existence of a for-est-dwelling Upucerthiaprovides an opportunity for furthertesting of this hypothesis in closely related birds. We can onlyspeculate as to the phylogenetic position of U. saturatior, but ifit proves to be basal to U. dumetaria, jelskii, validirostris, andalbigula, the pattern would provide further support for a for-est origin of another lineage that now occupies open habitats.Regardless of the direction of evolutionary change, the exis-tence of the forest-based U. saturatiorimplies a habitat shiftduring speciation at some point in the history of Upucerthia,

but whether it originated following a secondary incursion intoforest habitats by an ancestral open-habitat Upucerthiaor thereverse can be tested only through complete taxon-samplingof the group in a formal phylogenetic analysis. Knowledge ofthe exact phylogenetic position of U. saturatiormight also

prove instrumental in fully solving the relationships betweenCinclodesand Upucerthia.

The voucher specimen of U. dumetariaused by Fjeldset al. (2007) is from Meseta Strobel (900 m above sea level),Santa Cruz, Argentina (J. Fjelds in litt., ZMUC 1984-02-14),and undoubtedly therefore refers to U. d. dumetaria. Thevoucher specimen used by Chesser et al. (2007) was collectedat Cerro Perito Moreno (ca. 800 m above sea level), ca. 2 km

below Refugio Perito Moreno, ca. 20 km NE of El Bolsn,

Dpto. Bariloche, Ro Negro, in a steppe like area within theforest, but since it was later prepared as skeleton it is impossi-

ble to attach a subspecific designation to this specimen on the

basis of current criteria (P. Sweet in litt., AMNH RTC 368).From the habitat description the specimen could be eitherdumetariaorsaturatior.

The lack of attention to the distinctive U. saturatiorinthe literature is somewhat surprising given its unique aspect,ecology, and distribution, and its potential to contribute to-ward the understanding of evolution and biogeography ofUpucerthiaand Cinclodes. Further studies on the systematicsof the U. dumetaria/saturatiorcomplex should focus on theintegration of genetic, morphological, and vocal data in theecotone between Andean forests and open steppe habitats.Until this issue is finally elucidated, the recognition of U. sat-uratioras a distinct species is fully supported by our data.

ACKNOWLEDGMENTS

We thank A. Bodrati, G. Bieber, K. Cockle, A. Grigoli, E. Lang-lois, G. N. Maurcio, L. Pagano, I. Pearman, M. Repenning, K.Roesler, J. Segovia, and M. Sokol for sharing fieldwork, C. Darrieu,A. Echevarra, J. R. Navas, J. V. Remsen, P. Sweet, P. Tubaro andK. Zyskowski for providing access to, and information on speci-mens held in their care, and to J. Fjelds for data on voucher speci-mens of Upucerthiaused in his study. J. A. Areta, A. Camperi, P.Capllonch, D. Ortiz, and T. Rosenberry provided bibliographicalassistance. I. Holzmann, L. Pagano and R. Jensen helped with bio-metric data in various ways. We also thank N. Athanas, T. Bishop, S.Imberti, B. Knapp, N. Krabbe, I. Roesler, and M. Sokol for provid-ing access to their recordings and to recordings held at the MacaulayLibrary of Natural Sounds and at the British Museum (NationalSound Archive). J. V. Remsen and K. J. Zimmer made useful criticalreviews of our manuscript.

LITERATURE CITED

ARETA, J. I.2007. Behavior and phylogenetic position ofPremnoplexbarbtails (Furnariidae). Condor 109:399407.

CHAPMAN,F.1919. Descriptions of proposed new birds from Peru,Bolivia, Argentina, and Chile. Bulletin of the American Museumof Natural History 41:323333.

CHESSER,R. T.2004. Systematics, evolution, and biogeography ofthe South American ovenbird genus Cinclodes. Auk 121:752766.

CHESSER, R. T., F. K. BARKER,AND R.T. BRUMFIELD. 2007. Fourfoldpolyphyly of the genus formerly known as Upucerthia, with noteson the systematics and evolution of the avian subfamily Furnarii-nae. Molecular Phylogenetics and Evolution 44:13201332.

CHUBB, C.1911. Exhibition and description of a new species of bi rd(Upucerthia tamucoensis) from southern Chili. Bulletin of theBritish Ornithologists Club 27:99106.

CRACRAFT, J.1985. Historical biogeography and patterns of differen-tiation within the South American avifauna: areas of endemism.Ornithological Monographs 36:4984.

ESTEBAN, J. G.1951. Furnariinae de la Republica Argentina. ActaZoologica Lilloana 12:377441.

FJELDS, J., ANDN. K. KRABBE. 1990. Birds of the High Andes. Zoo-logical Museum, University of Copenhagen, and Apollo Books,Svendborg, Denmark.


14/16


15/16


APPENDIX 1. List of specimens examined and measured for thisstudy.

U. saturatior: FML 4106, 4157, 4231, 4248, 4254, 7751, 7758. MACN

nn, 65g, 1478a, 1478a, 2280-295a, 2282, 2285, 2331a, 3611, 4340a,4483, 4626a, 4797, 4822, 4824, 4842, 4927, 4821, 5262, 5263, 5264,5371, 8389-67a, 8389-67b, 8389-67c, 8389-67d, 8389-67e, 33414,38056, 38058, 38059, 38075, 42461. MLP10162, 10163, 10164, 10165,10166, 10167.

U. d. dumetaria: FML 1586, 8285, 12114, 12115, 12116, 12117, 12118,12120, 12121. MACN 8339-65c, 8339-65e, 8339-65d, 8389, 31518,49226, 49241, 51491, 52219. MLP 5730, 5731, 10734, 12412, 12455,12873, 13015, 13530, 13656.

U. d. hypoleuca: FML 1570, 1571, 1572, 5451, 5614, 8202, 8203,8204, 11314, 11315, 12119. MLP 425, 426, 427, 428, 429, 430, 431,432, 433, 7917.

Specimens considered intermediates betweensaturatiorand dume-tariaby Navas (1971). Adults: MACN 42765, 48572, 48575. Juve-niles/immatures: MACN 40763, 40764, 42459, 48570, 48571, 48573.

List of specimens not examined but used to plot geographical distri-

bution and migration patterns.U. saturatior: FMNH 61140, 61141, 61142, 61143, 61144, 61145,61146, 61147, 61148, 61149, 61150, 61151, 61152, 61153, 62457, 316676.LSU 31036, 69968. YPM 21731, 21732, 21797, 21828, 22680, 82818,82819, 82820, 82821, 82822, 82823, 82824, 82825, 82826, 82827,82828, 82829, 82830, 82831, 82832, 82833, 82834, 82835, 82836,82837, 82838, 112658, 112659, 112660.

U. d. dumetaria: FMNH 56810, 56811, 58944, 120658, 120659,120660, 262641, 303506, 303507, 303508, 303509, 303510. LSU63628, 69967, 178756, 178757. YPM 82790, 82791, 82792, 82793,82794, 82795, 82796, 82797, 82798, 82799, 82800, 82801, 82802,82803, 82804, 82805, 82806, 82807, 82808, 82809, 82810, 82811,82812, 82813, 82814, 82815, 82816, 82817, 115046, 115047.

U. d. hypoleuca: FMNH 54124, 54125, 58945, 61154, 61155, 61156,61157, 61158, 61159, 61160, 61161, 61162, 61163, 61164, 61165, 61166,61167, 61168, 180034. LSU 37595. YPM 21730, 21794, 21795, 38925,

38926, 38927, 38928.

APPENDIX 2. List of recordings used for this study. Capitalizednames refer to Argentine provinces, Chilean regions, and Bolivianor Peruvian departments. Each locality is followed by name of therecordist or catalogue number, and the number of individual birds re-

corded at each locality is in parentheses. Acronyms: MLNS, MacaulayLibrary of Natural Sounds, Cornell Lab of Ornithology; NSA, Na-tional Sound Archive, British Museum; XC, www.xeno-canto.org.

U. saturatior: NEUQUN. Lagunas de Epulauquen (Areta 3, Roesler 1),Paso Pichachn (Pearman 1), Pino Hachado (Pearman 2), 20 kmwest of Rahue (Sokol/Pearman 1).

U. d. dumetaria: CHUBUT. Pennsula Valdez (Straneck 1). RIONEGRO. Embalse El Chocn (Areta 5) . SANTA CRUZ. EstanciaLas Buitreras [60 km from Ro Gallegos] (Imberti 1), Estancia LaQuerencia [northern limit of Lago Argentino] (Imberti 1), Lago Car-diel (Pearman 1), Laguna Los Escarchados (Krabbe 1 [XC-15997,16226]), Monumento Natural Bosque Petrificado (Imberti 1). XIIREGION. El Porvenir (Areta 1), Sierra Baguales (Imberti 1).

U. d. hypoleuca: MENDOZA. Cuesta de los Terneros (Areta 1), SanRafael (Areta 1), Refugio San Antonio (Areta 2). SAN JUAN. Las

Flores to La Serena road (Pearman 1). TUCUMN. El Infiernillo(Athanas 1, Macaulay 1 [MLNS-115989]). REGION METROPOLI-TANA. Farellones (Knapp 2 [XC-15774, 17124]).

U. albigula: AREQUIPA.15 km e from Arequipa on road to Puno(Parker 2 [MLNS-33870, 33879]), ca 10 km w of Arequipa by roadwest of Chiguata (Schulenberg 1 [MLNS-36056]). I REGION. Putre(Whitney 2 [MLNS- 46451, 46452], N. Krabbe 1 [NSA 23462]).

U. jelskii: JUJUY. Azul Pampa (Pearman 1). REGION I. Putre(Pearman 1, Areta 2). LA PAZ. Apolobamba National Park (Hen-nessey 1 [MLNS-110776]).

U. validirostris: TUCUMAN. El Infiernillo (Pearman 5, Macaulay1 [MLNS-115979]).

U. serrana: ANCASH. Llanganuco (Parker 1 [MLNS-33953], Vanden Berg 2 [MLNS-28713, 28717]). LA LIBERTAD. Tayabamabaoutskirt s (Parker 1 [MLNS-17462]).


16/16


APPENDIX 3. List of localities and dates used to build distribu-tion maps in this study. Numbers of each locality refer to Figure 2. Lat-itude (S) and longitude (W) in parentheses; months in brackets.

U. d. dumetaria: 1, Chilecito (2910, 6730) [May]; 2, Ro Seco, Con-carn (3234, 6515) [May]; 3, Embalse El Yeso (3338, 7000)[Nov]; 4, Laguna Llancanelo (3535, 6909) [Sep]; 5, CuramalnChico (3728, 6206) [Aug]; 6, DOrbigny (3741, 6143) [Aug]; 7,Sierra Auca Mahuida (3753, 6831) [Nov]; 8, Lihuel Calel (3802, 6533) [Jun]; 9, Caleu Caleu, Laguna Colorada (3815, 6345)[May]; 10, Chasico (3820, 6239) [Jul]; 11, Ro Sauce Grande,Monte Hermoso (3855, 6133) [May]; 12, Plaza Huincul (3855,6909) [?]; 13, 40 km N Choele Choel (3856, 6541) [Feb]; 14,Cipolletti (3856, 6759) [Sep]; 15, Mayor Buratovich (3915, 6237) [Aug] 16; 18 km N San Martin de los Andes (39 59, 7120)[Nov]; 17, Baha San Bls (4033, 6215) [May]; 18, San AntonioOeste (4044, 6456) [?]; 19, Carmen de Patagones (4048, 6259) [Apr]; 20, Nahuel Huapi (4058, 7130) [Apr]; 21, Departa-mento Pilcaniyeu, near Neneo Ruca (4109, 7024) [NovJan]; 22,Departamento Bariloche (4109, 7118) [Feb]; 23, Ingeniero Jaco-

bacci (4118, 6935) [Mar]; 24, Estancia Huanu-luan (4122, 69

52) [Nov]; 25, Departamento orquinco (imprecise locality) (4151, 7054) [Feb]; 26, El Hoyo (imprecise locality) (4204, 7130)[Sep]; 27, Peninsula Valdez (4230, 6400) [Feb]; 28, 100 km ENEEsquel (4238, 7008 ) [?]; 29, 7 km S Puerto Madryn (4250,6503) [Dec]; 30, Punta Tombo (4403, 6511) [?]; 31, Ro Pico(4414, 7159) [Jan]; 32, 8 km S Fitzroy (4706, 6715) [Nov];33, 43 km N Tres Lagos (4913, 7132) [Jan]; 34, 21 km N TresLagos (4924, 7132) [Jan]; 35, 1 km N Tres Lagos (49 34, 7132) [Jan]; 36, Ro Chico (4956, 6832) [Apr]; 37, Estancia La Ju-lia (5016, 7144) [Jan]; 38, Coy Inlet (5057, 6910) [Apr]; 39,Lago Sarmiento (5102, 7246) [AugOct]; 40, Ro Gallegos (5138, 6913) [Jul]; 41, Estancia Federica (5141, 7158) [Nov]; 42,

between Estancia Punta Alta and Estancia Federica (5141, 7158)[Sep]; 43, Ro Ciaike, near Argentine boundary (5213, 7029 )[Dec]; 44, Estancia Monte Dinero (5217, 6832) [Dec].

U. d. hypoleuca: 45, Ro Loa (2214, 6836) [Jul, Nov]; 46, Mai-mara, Lagunas del Cerro Colorado (2337, 6524) [Apr]; 47, Ra-

madilla Copiap Valley (2721, 7018) [Nov]; 48, Monte Amargo(2721, 7019) [Aug]; 49, Domeyko (2857, 7053) [Jul]; 50,Romero (2958, 7118) [FebJun]; 51, Baos del Toro (2958,71 18) [FebMar]; 52, Aiquina (2217, 68 20) [Jul]; 53, SanPedro de Atacama (2330, 6815) [Jul]; 54, Tilcara (2334, 6522) [Aug]; 55, Cafayate (2605, 6558) [Aug]; 56, Tucumn (2649, 6513) [May, Oct]; 57, Taf del Valle (2652, 6541) [May];58, Los Nacimientos de Abajo, Hualfin (2711, 6644) [Sep]; 59,Gualfin (2714, 6650) [Aug]; 60, Ischigualasto, Valle de la Pea(3008, 6751) [AprMay]; 61, Pampa de Achala (3135, 6450)[May]; 62, General Lavalle (3243, 6835) [Sep]; 63, Cordillera deLa Punta (3400, 7038) [SepOct]; 64, Manantiales (3410, 7043) [Jul].

U. d. dumetaria(dark specimens, intergradessensuNavas [1971]):65, 66, orquinco (4151, 7054) [Feb]; 67, El Bolsn (4158, 7131) [Dec]; 68, El Maitn (4203, 7110) [Sep]; 6972, Cushamen(4212, 7050 ) [DecFeb]; 73, Alto Ro Senguerr (4502 , 7050) [Feb].

U. saturatior: 74, La Ligua (3227, 7114) [Apr]; 75, La Calera(3245, 7112) [Apr]; 76, 77, Concon (3255, 7131) [Jul]; 78, RoAconcagua (3256, 7131) [?]; 79, Olmue (3300, 7112) [Aug];80, Costa Valparaso (3301, 7133) [?]; 81, Quilpu (3302, 71

24) [Jun]; 82, Aconcagua (3302, 7138) [Apr]; 83, Casablanca(3303, 7136) [?]; 84, Lampa (3317, 7052) [Jun]; 85, Cerrosde Curacavi (3324 , 7109 ) [Jun]; 86, Alrededores de Santiago(3328, 7038) [May]; 87, Santiago (3328, 7038) [Aug]; 88,Melipilla (3342, 7113) [Aug]; 89, Paine (3348, 7044) [Apr];90, San Francisco del Mostazal (3359, 7043) [May]; 91, Gran-eros (3404, 7044) [Aug]; 92, Rancagua (3409, 7044) [Jun];93, Peleuquen (3409, 7045) [Jan]; 94, Machali (3411, 7038)[Sep]; 95, Teno (3451, 7109 ) [Sep]; 96, Cordillera de Curic(3459, 7059) [May]; 97, Concepcin (3608, 7301) [Apr]; 98,Quirihue (3617, 7232) [Jan]; 99, Penco (3644, 7249) [Aug];100, Hacienda Gualpencillo (3647, 7304) [AugFeb]; 101, La-gunas de Epulauquen (3648, 7104) [DecMar]; 102, Paso Pich-achn (3726, 7106) [Dec]; 103, Angol (3747, 7242) [Oct];104, Lonquimay Pedregoso (3826, 7113) [Dec]; 105, Ro Lolen,Lonquimai Valley (3826, 7114) [AugOct]; 106, Villa Portales(3827, 7122) [Feb]; 107, Reaco (3835, 7250) [Jul]; 108, Pino

Hachado (3840 , 7053) [Dec]; 109, Laguna Gualletue (3842,7116) [Mar]; 110, Temuco (3843, 7234) [Jul]; 111, 20 km westof Rahue (3922, 7109) [Dec]; 112, Sierra de Pilpil, San Martinde los Andes (4010, 7121) [Dec]; 113, Valle Encantado del RoLimay (4043, 7105) [Feb]; 114, Nahuel Huapi (4058, 7130)[SepJan]; 115, Lago Moreno (4104, 7132) [Mar]; 116, Lago Es-condido on the Ro Foyel (4136, 7143) [?]; 117, El Bolson (4158,7131) [Feb]; 118, Laguna Espejo, El Hoyo (4204, 7130) [Oct];119, Cholila (4231, 7127) [May]; 120, Esquel (4254, 7119)[?]; 121, Ro Pico (4414, 7159) [Jan].

Documents

taxonomia upucerthia