The Condor 109:399–407 # The Cooper Ornithological Society 2007

BEHAVIOR AND PHYLOGENETIC POSITION OF BARBTAILS (FURNARIIDAE)

JUAN IGNACIO ARETA1,2,3 1CICyTTP-CONICET, Materi y Espan˜a, 3105, Diamante, Entre Rı´os, Argentina 2Grupo FALCO, El Coronillo, Reserva Natural Punta Lara, 1915, Ensenada, Buenos Aires, Argentina

Abstract. The assemblage includes four genera: Margarornis, Premno- plex, Premnornis, and Roraimia, all thought to be closely related. Differences in vocalizations and habitat use between Premnoplex brunnescens () and P. tatei (White-throated Barbtail) are consistent with their full status. In light of the weak anatomical (hindlimb musculature) evidence supporting the inclusion of Premnornis in the Margarornis assemblage, the convergence-prone nature of characters associated with climbing habits, and the differences in their nests and foraging behavior, I propose that Premnornis is not a member of the Margarornis assemblage and that Premnoplex is closely related only to Margarornis. These results are supported by recent molecular phylogenetic analyses. Although Roraimia differs in plumage, tail shape, and song from other members of the Margarornis assemblage, it must be provisionally included in the assemblage until evidence clarifies its phylogenetic position. Key words: barbtails, behavior, convergence, morphology, nest, Premnoplex, Premnornis. Comportamiento y Posicio´n Filogene´tica de los Premnoplex (Furnariidae) Resumen. El ensamble Margarornis incluye cuatro ge´neros: Margarornis, Premnoplex, Premnornis y Roraimia, aparentemente cercanamente emparentados. Las diferencias de ha´bitat y voces de Premnoplex brunnescens ydeP. tatei son consistentes con su estatus especı´fico. A la luz de la de´bil evidencia anato´mica (musculatura apendicular) apoyando la inclusio´n de Premnornis en el ensamble Margarornis, la naturaleza convergente de los caracteres asociados a los ha´bitos trepadores y las diferencias en comportamiento y nidificacio´n, propongo que Premnornis no es un miembro del ensamble Margarornis y que Premnoplex estarı´a solamente cercamente emparentado a Margarornis. Estos resultados son congruentes con recientes estudios moleculares. Aunque Roraimia difiere en plumaje, forma de la cola y canto de otros miembros del ensamble Margarornis, debe ser provisionalmente incluido en e´l hasta que ma´s evidencia aclare su posicio´n filogene´tica.

INTRODUCTION 1938, Vaurie 1980) have considered them con- There are two species of Premnoplex barbtails, specific. Some classifications (e.g., Meyer de little-known ovenbirds (Furnariidae) that in- Schauensee 1966, Vaurie 1980) merge Premno- habit humid montane forests in South and plex into Margarornis. Both Premnoplex species Central America. P. brunnescens (Spotted Barb- are included in the so-called ‘‘Margarornis tail) comprises five mountain-dwelling subspe- assemblage,’’ which consists of eight species in cies present in Costa Rica, Panama, and from the genera Margarornis, Roraimia, Premnornis, the Coastal Cordillera of Venezuela through the and Premnoplex (Vaurie 1980, Rudge and Andes to Bolivia, whereas P. tatei (White- Raikow 1992a, 1992b). This grouping is largely throated Barbtail) is endemic to northeastern based on structural (size, wing shape, tarsus Venezuela and consists of two distinctive sub- proportions), anatomical (hindlimb muscula- species: P. t. tatei in the Turimiquire Massif and ture), and plumage pattern and coloration P. t. pariae in Penı´nsula de Paria (Phelps and features (Vaurie 1980, Rudge and Raikow Phelps 1949, 1963, Hilty 2003, Remsen 2003). 1992a, 1992b). Relationships among members Evidence for species status of both taxa is of the assemblage are not clear, and hypotheses considered to be weak (Remsen 2003, Remsen derived from structural and anatomical features et al. 2006), and some authors (e.g., Hellmayr conflict with those suggested by natural history and molecular data (Rudge and Raikow 1992b, Dobbs et al. 2003, Irestedt et al. 2006). Manuscript received 11 July 2006; accepted 9 January 2007. Natural history data have proven useful for 3 E-mail: [email protected] predicting the phylogenetic relationships of

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The Condor cond-109-02-14.3d 14/3/07 12:31:17 399 Cust # 8210 400 JUAN IGNACIO ARETA furnariids (Kratter 1994, Rodrigues et al. 1994, Nacional Yacambu´, Premnoplex brunnescens Whitney and Pacheco 1994, Kratter and Parker was always found close to or at ravines or 1997), but the ecology and behavior of the rivers, both during the breeding and nonbreed- Premnoplex barbtails have been poorly described ing seasons. The species is known to favor and are almost unknown for the localized P. tatei humid areas in montane evergreen rainforests, (Remsen 2003). Herein I document habitat use particularly dense undergrowth with vines and analyze the systematic significance of vocal- alongside watercourses (Slud 1964, Skutch izations, behavior, and morphology of Premno- 1967, Vaurie 1980; JIA, pers. obs.). In 2.7 km plex barbtails. I also reanalyze plumage, mor- of suitable habitat along the Quebrada Negra, I phological, and nesting characters of members in found 22 territories, giving a density of 8.2 pairs the Margarornis assemblage in an attempt to per linear km, or one territory every 120 m. evaluate the validity of the grouping and the Premnoplex tatei was found in mossy, epi- position of Premnoplex within it. phyte-laden montane forest in Parque Nacional Peninsula de Paria, usually in flat mountaintop METHODS areas with large stands of palms and unidenti- I studied Premnoplex brunnescens (probably fied Heliconia-like plants (see Hilty [1999, 2003] rostratus) for ca. 100 days from for habitat in Turimiquire). Only once did I find March 28 to June 25 2005 in Parque Nacional P. tatei in typical P. brunnescens habitat: a pair Yacambu´, Estado Lara, Venezuela (9u249N, was located by a creek with extreme humidity, 69u309W, 1200–1800 m asl), during both pre- ferns, and decaying wood as outstanding breeding and breeding seasons. Premnoplex features. In 1 linear km of palm-dominated tatei pariae was studied for two days during 8 habitat I found five P. tatei territories, or one and 9 July 2005 in Parque Nacional Peninsula territory every 200 m. de Paria, Estado Sucre, Venezuela (10u419N, VOCALIZATIONS 62u369W, 700–1150 m asl), apparently during the breeding season. I recorded data on habitat, The noisy environment of Premnoplex brunnes- behavior, and vocalizations for both species. I cens made it extremely difficult to obtain clear studied nesting only in P. brunnescens. Record- recordings of its vocalizations. I was only able ings of P. brunnescens vocalizations were made to obtain recordings of its most commonly used with a Sennheiser ME-66 directional micro- call: a short, descending, trilled ‘‘prriiip’’ phone and a Sony TCM-5000 tape recorder. P. (Fig. 1A). Three other vocalizations were also tatei vocalizations were recorded with an heard: a sharp and high-pitched ‘‘pssiit’’ and Audio-Technica at-835b directional micro- a cricket-like, high-frequency ‘‘ti-ti-ti-ti-ti-ti-ti- phone and a Marantz PMD-222 tape recorder. ti-ti-ti-ti’’ slowly descending in pitch and tempo. I examined and measured 189 specimens Only once did I hear its complex and long song, housed at the Coleccio´n Ornitolo´gica Phelps which reminded me of the songs of some in the four genera comprising the Margarornis Cranioleuca spinetails. Slud (1964), Fjeldsa˚ assemblage (29 P. brunnescens brunnescens, nine and Krabbe (1990), and Hilty (2003) described P. b. rostratus,26P. tatei tatei,49P. t. pariae, the same vocalizations, although using a differ- 42 Margarornis squamiger perlatus [Pearled ent phonetic notation. The most common call Treerunner], 12 Premnornis guttuligera venezue- was heard roughly 95% of the time, and the lana [Rusty-winged Barbtail], and 22 Roraimia other two vocalizations were heard 5% of the adusta adusta [Roraiman Barbtail]). Bill length time. I also obtained recordings of the most (exposed culmen) was measured to the nearest common call of Premnoplex tatei. This was the 0.05 mm with digital calipers, and wing chord only vocalization that I heard, although its and tail length were measured with a metal ruler repertoire is surely greater. Its call is reminis- to the nearest 0.5 mm. cent in pattern to that of P. brunnescens, but differs by being lower in pitch and deeper and RESULTS somewhat bubbled with an underwater quality (Fig. 1B). This vocalization was highly variable HABITAT in length, intensity, and tempo, varying from Both Premnoplex barbtails frequented cool, two to five notes, from mildly low to strong in epiphyte-laden, dark rainforest. In Parque volume, and from a leisurely series of notes to

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FIGURE 1. Spectrograms of the most common vocalizations of Premnoplex barbtails. A) P. brunnescens call, recorded at Parque Nacional Yacambu´, 11 April 2005. B) P. tatei pariae calls, recorded at Parque Nacional Penı´nsula de Paria, 9 July 2005. Note the higher pitch and faster delivery rate of each note of P. brunnescens. The interval between successive calls is shorter in P. tatei than in P. brunnescens, thus two consecutive calls are shown for P. tatei. These differences in their calls support their treatment as two separate species.

a frantic bubbling that was never as fast as that flicking was performed while the body was held of P. brunnescens. Vocalizations of P. tatei have in any position, from upright perching to previously been described as an ‘‘almost bubbly clinging to vegetation while hanging upside ser. of rather low, soft, reedy whistles, we-whu´r, down. Horizontal hopping was a behavior that we-whu´r, we-hee´t…, varied to be-be-bu´r, be-be- both species displayed when alarmed. bu´r…orpi, pr-pr-pr-prip!… in long ser. when would perch in an upright position on a hori- excited’’ (Hilty 2003:490). Both species used zontal branch and hop from side to side while their calls routinely while foraging, moving, facing the intruder. Each hop was accompanied perching, or when alarmed, which, when added by wing-flicking and the most common call. to song structure, strongly suggests homology. Individuals usually hopped many times to one side before hopping in the other direction. In BEHAVIORAL OBSERVATIONS the case of Premnoplex brunnescens, this behav- Wing-flicking and horizontal hopping. Two ior was performed each time I approached a nest distinctive behaviors of the Premnoplex barb- with nestlings, and both parents would hop tails that I observed have not previously been with their beaks full of worms for their noted in the literature: constant high wing- nestlings. I did not witness this behavior outside flicking and horizontal hopping. Although the breeding season, thus I consider this some furnariids occasionally flick their wings behavior to be an indication of breeding. (Whitney and Pacheco 1994), this behavior is Therefore, although I did not find a nest of taken to an extreme in Premnoplex. Almost Premnoplex tatei, the horizontal hopping be- constantly, while foraging, perching, or simply havior they directed at me suggested that they moving, both species flicked their wings high were nesting (see Breeding). over their backs, in the manner of many Foraging. Premnoplex brunnescens foraged Neotropical flycatchers (Tyrannidae). Both mostly by probing and gleaning while climbing wings were flicked at the same time, and this small mossy trees, moving along horizontal

The Condor cond-109-02-14.3d 14/3/07 12:31:18 401 Cust # 8210 402 JUAN IGNACIO ARETA branches and vines, hanging upside down, or cades or in areas reached by water spray. The creeping over and under logs. The species only nests were mossy balls with the entrance usually occasionally foraged on the ground (Slud 1964, facing downward, and had an inner platform Wetmore 1972, Remsen 2003; JIA, pers. obs.). on which the eggs were laid (not visible from While foraging, individuals could spend several the outside). The platforms bent upwards close minutes carefully searching for arthropods to the nest entrance, preventing the eggs from among mosses and epiphytes or in tree bark. falling through the vertical tunnel entrance (see All birds were found feeding alone or in pairs, drawing in Skutch [1967]). Nestlings of P. never as part of mixed-species flocks. In brunnescens showed a complex behavior asso- contrast to Wetmore (1972), I never saw P. ciated with defecation: both nestlings would brunnescens searching dead leaves, despite their turn inside the nest and, while perching on the prevalence in their habitat. As pointed out by inner platform, would defecate through the nest Slud (1964) and Hilty (2003), P. brunnescens entrance; most feces were removed by the water only rarely used its tail for support while flowing underneath the nest. This behavior stationary or as a prop while climbing tree frees parents from fecal-sac removal duties trunks. Foraging height ranged from ground and is also shown by Margarornis (Mennill level to 5 m, but birds were usually found and Doucet 2005). The nest structure and nest- between 2 and 4 m. In contrast, Premnoplex site locations of Premnoplex tatei remain tatei foraged mostly on the ground and less undescribed. Although riverine nesting cannot commonly on tree trunks and branches. While be excluded, there were no permanent rivers in hopping on the ground, its wings usually the habitat where P. tatei was found at Parque remained closed, without the characteristic Nacional Peninsula de Paria. Available habitat wing-flicking. When foraging in vegetation, it use data suggest that its nest might be built in moved much like its congener, but when preexisting clumps of moss, under tree foraging on the ground, it stuck its head into branches, or in crevices in habitat somewhat holes in fallen trunks and rotten logs and different from that of P. brunnescens.The frequently tossed leaves aside with its bill to finding of apparent fledglings and display of look for arthropods. P. tatei, like P. brunnescens, lateral hopping behavior by adults suggests that typically foraged alone or in pairs, although I breeding was going on while fieldwork was once observed four P. tatei (a pair and two carried out. probable fledglings) foraging in a mixed-species flock composed of Basileuterus tristriatus COMPARATIVE MORPHOLOGY OF THE (Three-striped Warbler) and Myrmotherula MARGARORNIS ASSEMBLAGE schisticolor (Slaty Antwren). Foraging height General morphology. Three main results emerge ranged from ground level to 2 m, but most birds when measurements of tail length, wing chord, were found on the ground or below 1 m. Hilty and exposed culmen of some members of the (1999) reported a single foraging on the Margarornis assemblage are compared (Ta- ground or on logs (never climbing) at heights ble 1). First, geographic variation in the from 0.2 to 3 m, loosely associated with a pair of Premnoplex barbtails is evident when subspecies Basileuterus griseiceps (Gray-headed Warbler). are analyzed separately (a fact obscured due to A video of a foraging P. tatei on the ground by extreme species lumping in Vaurie [1980]). D. Ascanio (Ascanio Birding Tours) is avail- Secondly, Premnoplex barbtails have relatively able at ,http://www.hbw.com/ibc/phtml/especie. longer bills and shorter tails and wings than the phtml?idEspecie54178.. remaining genera, accounting for their wren-like Breeding. I found and examined over 35 nests aspect and behavior (Wetmore 1972, Vaurie of Premnoplex brunnescens. All were built of 1980). Lastly, Premnornis is the only in moss and were attached to rocks, trees, or roots which tail length exceeds wing chord. usually in dark, hidden, and extremely wet Tail shape. Tail shape is extremely variable areas, but never in what could strictly be and plastic in the Furnariidae and can be considered a cavity. This coincides with pre- considered a rough indicator of phylogenetic vious descriptions (Skutch 1967, Zyskowski and relationships, with closely related species often Prum 1999). Nests were usually placed in sites having similar tails (Remsen 2003, Fjeldsa˚ et al. above fast-flowing water, often close to cas- 2005). Tails of all members of the Margarornis

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TABLE 1. Wing chord, tail length, and exposed culmen measurements of some members of the Margarornis assemblage. Values are given in mm as mean 6 SD (with number of individuals measured in parentheses). Specimens were housed at the Coleccio´n Ornitolo´gica Phelps.

Species Wing chord Tail length Exposed culmen Premnoplex tatei tatei 58.2 6 1.7 (26) 56.0 6 5.0 (26) 15.4 6 0.9 (25) P. t. pariae 60.6 6 1.3 (49) 51.9 6 5.7 (49) 16.0 6 0.6 (41) P. brunnescens brunnescens 59.9 6 1.4 (29) 58.7 6 2.9 (28) 14.3 6 0.5 (28) P. b. rostratus 60.0 6 2.5 (9) 55.6 6 3.0 (7) 15.4 6 0.2 (7) Margarornis squamiger perlatus 75.6 6 3.0 (42) 70.2 6 3.7 (42) 11.2 6 0.6 (38) Roraimia adusta adusta 63.6 6 3.2 (22) 60.9 6 4.2 (22) 12.7 6 0.5 (20) Premnornis guttuligera venezuelana 65.8 6 1.2 (12) 66.8 6 2.5 (12) 13.6 6 0.4 (12)

assemblage are composed of 12 rectrices that Margarornis rubiginosus (). are more or less graduated (Vaurie 1980). I Notably, Premnoplex tatei pariae shows a dif- distinguished three different tail types: the ferent pattern: the dark margin of each breast Premnoplex-Margarornis type (Fig. 2A), the is limited to the sides, hence there is no Roraimia type (Fig. 2B), and the Premnornis drop-shaped center, but a light, broad, central type (Fig. 2C). Tails of Margarornis and stripe, bordered by a narrow dark line to each Premnoplex are very similar, showing side. The ventral pattern of Roraimia is similar with well-integrated webs ending abruptly by to that of P. tatei pariae but the central, light the asymmetrical reduction of vanes, and part of each feather is wider, thus giving it prolonged in a protruding spine with reduced a strongly flammulated appearance. The facial (and sometimes absent) vanes. The tail of pattern of Roraimia adusta is strikingly similar Roraimia has notably less-integrated webs, to that of Synallaxis [Poecilurus] candei (White- and feather ends are pointed (‘‘v-shaped tip’’) whiskered Spinetail), and Synallaxis [Poeci- due to the more gradual reduction of the vanes lurus] kollari (Hoary-throated Spinetail): all and the lack (or near lack) of spines in tails in have a boldly patterned throat, dark facial good condition. The tail of Premnornis differs stripe and auriculars, and a broad reddish- strikingly from the others in rectrix shape: the orangish eyebrow that extends downward tips are much more rounded, show no asym- passing behind the facial stripe, which differs metrical vane reduction toward the tip, and lack notably from other members of the Margar- barbs or spines. Integration of webs is in- ornis assemblage. Moreover, when the throat termediate between that of Roraimia and feathers of Roraimia are closely examined, Premnoplex-Margarornis. Tail shapes change black is obvious at their bases exactly where due to abrasion, sometimes so much that they the black gular spot of both previously men- lose diagnostic features (Fig. 2), which must be tioned Synallaxis (Poecilurus) spp. appears, taken into account when studying tail shape in suggesting a close relationship among these the group. forms. Premnornis differs from the other three Plumage. An exhaustive plumage analysis of genera in that the color of the tail strongly the Margarornis assemblage is beyond the scope contrasts with the color of the back, but of this paper, but some features and ideas are otherwise it is similar in plumage to the worth noting (see Zimmer [1934], Wetmore Premnoplex-Margarornis barbtails. Without [1972], Vaurie [1980], and Remsen [2003] for a formal phylogenetic analysis, it is not clear detailed plumage descriptions). Margarornis whether the plumage similarities reflect a close and Premnoplex share a common ventral relationship or homoplasy. pattern, with the feather edges enclosing a drop-shaped paler center. This pattern is well DISCUSSION developed in Margarornis bellulus (Beautiful Premnoplex tatei was initially described as a new Treerunner), Margarornis squamiger, Premno- species by Chapman (1925), but was sub- plex tatei tatei, and all subspecies of Premno- sequently considered a subspecies of Premno- plex brunnescens, but less developed in Margar- plex brunnescens by Hellmayr (1938) and ornis stellatus (Fulvous-dotted Treerunner) and Vaurie (1980). Meyer de Schauensee (1966)

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In furnariids, morphological features associ- ated with foraging are far more plastic than nest-building behaviors (including nest shape, nest placement, and construction materials), as shown by Limnoctites-Cranioleuca and Limnor- nis-Phleocryptes (Olson et al. 2005), Geositta (Cheviron et al. 2005), and Anumbius-Coryphis- tera (Zyskowski and Prum 1999, Fjeldsa˚ et al. 2005). Thus, a weighting of the characters to be used for phylogenetic reconstruction becomes justified and necessary: more weight should be given to the characters that convey more accurate phylogenetic information (e.g., nesting characters) and less weight given to plastic or convergent-prone characters (e.g., feeding adaptations). Morphological convergence seems a pervasive phenomenon among trunk-climbers, and the role of morphology in elucidating phylogenetic relationships is usually limited, because conver- gent adaptations can mask actual phylogenies (Feduccia 1973, Irestedt et al. 2004). Rudge and Raikow (1992b) nonetheless based their phylo- genetic analysis of the Margarornis assemblage on myiological hindlimb characters that could represent convergent adaptations for foraging (McCracken et al. 1999, Irestedt et al. 2004). Rudge and Raikow (1992b) proposed a closer relationship between Premnoplex and Premnor- nis within the Margarornis assemblage. How- ever, Dobbs et al. (2003) described the nest of Premnornis guttuligera as a cup made of tree- fern petiole scales inside a cavity, contrasting with the mossy ball-shaped nest attached to trees or banks built by Premnoplex and Margarornis (Feduccia 1970, Fjeldsa˚and Krabbe 1990, Skutch 1996, Remsen 2003, FIGURE 2. Tail shapes of members of the Margar- Mennill and Doucet 2005). This marked con- ornis assemblage. A) Premnoplex–Margarornis,B) trast in nest structure prompted Dobbs et al. Roraimia, and C) Premnornis. Tails in good condition (2003) to suggest that Premnornis was mis- are shown on the left side, worn tails on the right side. Note the loss of diagnostic features in the worn tails of placed within the Margarornis assemblage. Premnoplex–Margarornis and Roraimia, which could Some furnariid species that usually have domed suggest a false similitude among their tails. Sketches nests have been reported to build cups when were based on the examination of individuals housed nesting in cavities (Johnson 1967, Zyskowski at the Coleccio´n Ornitolo´gica Phelps. and Prum 1999), but to date no nest of Premnoplex or Margarornis has been found and Remsen (2003) treated them as separate totally within a cavity (contra Irestedt et al. species, but published evidence for either 2006; JIA, pers. obs.), a fact that strengthens treatment is weak (Remsen et al. 2006). the use of a cup-shaped cavity nest in Differences presented here regarding vocaliza- Premnornis as evidence for its exclusion from tions, foraging mode, and habitat use support the Margarornis group (Dobbs et al. 2003). the validity of P. brunnescens and P. tatei as Moreover, the peculiar behavior of defecating separate species. through the nest entrance shown by nestlings of

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Premnoplex and Margarornis would be pre- strikingly from Premnoplex. In the light of the cluded if nests were placed wholly in a cavity. convergence-prone nature of characters associ- A reanalysis of the two anatomical synapo- ated with climbing habits that were suggested as morphies in Rudge and Raikow (1992b) that supporting the inclusion of Premnornis in the support the Premnoplex-Premnornis clade re- Margarornis assemblage, and the differences in veal their weaknesses as phylogenetic indica- tail shape, external morphology, nests, and tors. First, the origin of Tendon M could be foraging behavior, I propose that Premnornis a symplesiomorphy or convergence of Premno- is not a member of the Margarornis assemblage. plex and Premnornis. Rudge and Raikow Similarities in vocalizations (high-frequency (1992b:765) stated that in Premnoplex (brun- trilled songs; Moore and Lysinger 1997, Mayer nescens) and Premnornis, ‘‘Tendon M originates 2000), nests (mossy balls with a lateral or entirely from the tibial lobe of M. flexor cruris underside entrance; Skutch 1967, Feduccia lateralis pars pelvica,’’ but in the next sentence 1970, Fjeldsa˚ and Krabbe 1990, Mennill and write, ‘‘A similar modification has also been Doucet 2005), foraging behavior (scansorial found among the dendrocolaptines.’’ Assum- habits and epiphyte-searching; Ridgely and ing, rather than testing, monophyly of the Tudor 1994, Sillett 1994, Remsen 2003), clade, they go on to conclude that ‘‘given the nestling behavior (autonomous defecation; monophyly of the treerunners, [the character] is Mennill and Doucet 2005), and tail shape derived for the clade consisting of Premnoplex (feathers with barbed tips and asymmetrically brunnescens and Premnornis guttuligera.’’ In reduced vanes; Wetmore 1972, Vaurie 1980) short, Rudge and Raikow (1992b) seem to point to a close relationship between Margar- disregard information suggesting that this ornis and Premnoplex (Remsen 2003). Data on character is not necessarily a synapomorphy, osteology and pterylosis are compatible with but that it can also be interpreted as a symple- this view (Wetmore 1972, Clench 1995). No siomorphy or as a convergent character. Sec- other furnariid genera show such a combination ond, the absence of the third belly of the M. of features; thus, Premnoplex and Margarornis lumbricalis could be an artifact. Rudge and are probably sister taxa, and the core of the Raikow (1992b) stated that the presence of Margarornis assemblage. a third belly in the M. lumbricalis is a synapo- The phylogenetic relationships of Premno- morphy of Premnoplex (brunnescens)and plex, Margarornis, and Premnornis suggested by Premnornis. However, they noted that the natural history, external measurements, and character is not present in every examined tail shape agree with phylogenetic hypotheses individual. The lack of the third belly in some based on DNA sequencing data, which found individuals is accounted for by suggesting that Premnornis to form a clade with Pseudoco- it could be due to a preparation artifact because laptes, and Premnoplex to group with Margar- the structure is so small. An alternative ornis in a distant clade (Irestedt et al. 2006). explanation seems equally likely: the lack of These independently derived and coincident the third belly in the other species could also be hypotheses support each other and disagree a preparation artifact, not being preserved in with the sister relationship of Premnoplex and any of the few studied individuals in other Premnornis proposed by Rudge and Raikow genera. Additionally, reduced taxon sampling (1992b) on anatomical grounds, suggesting that in Rudge and Raikow (1992b) renders their the guttate ventral pattern has evolved many work unsuitable to test for monophyly of the times in the furnariids (Irestedt et al. 2006). Margarornis assemblage. The relationships of Roraimia adusta are Premnornis is not scansorial and almost half enigmatic. Originally described as a Synallaxis of its foraging attacks were directed toward spinetail, a monotypic genus was later created prey in or on suspended dead leaves (Dobbs et for it, despite the lack of any diagnostic al. 2003), unlike Premnoplex and Margarornis, character, based on number of rectrices (12 in which are scansorial and seldom forage in dead Roraimia,10inSynallaxis), tail shape (pointed leaves (Fjeldsa˚ and Krabbe 1990, Sillett 1994; in Roraimia, usually rounded in Synallaxis), JIA, pers. obs.). Finally, the reversed tail to and hind-toe claw size (long in Roraimia, wing chord ratio of Premnornis, and its mildly Margarornis, and Premnoplex, short in Synal- acuminate tail shape with no spines, differs laxis; Chapman 1929). Roraimia has a compos-

The Condor cond-109-02-14.3d 14/3/07 12:31:20 405 Cust # 8210 406 JUAN IGNACIO ARETA ite plumage pattern, with facial features of some CLENCH, M. H. 1995. Body pterylosis of wood- Synallaxis (Poecilurus) spp., a ventral pattern creepers and ovenbirds (Dendrocolaptidae and Furnariidae). Auk 112:800–804. reminiscent of Premnoplex tatei pariae but even DOBBS, R. C., H. F. GREENEY, AND P. R. MARTIN. more flammulated, and tail shape intermediate 2003. The nest, nesting behavior, and fora- between that of Premnoplex-Margarornis and ging ecology of the Rusty-winged Barbtail Premnornis. The song of Roraimia differs in (Premnornis guttuligera). Wilson Bulletin 115: pattern, tempo, and harmonic structure from 367–373. FEDUCCIA, A. 1970. Natural history of the avian songs of Premnoplex and Margarornis (D. families Dendrocolaptidae (woodhewers) and Ascanio and JIA, unpubl. data). The situation Furnariidae (ovenbirds). Journal of the Gradu- of Roraimia is analogous to that of Gyalophy- ate Research Center 38:1–26. lax, which is deemed to be intermediate FEDUCCIA, A. 1973. Evolutionary trends in the Neotropical ovenbirds and woodhewers. Orni- between Asthenes and Synallaxis (Whitney thological Monographs 13:1–69. and Pacheco 1994). The placement of Roraimia FJELDSA˚ , J., M. IRESTEDT, AND P. G. P. ERICSON. in the Margarornis assemblage should be 2005. Molecular data reveal some major adap- considered provisional until data on genetics, tational shifts in the early evolution of the most natural history, vocalizations, and nesting are diverse avian family, the Furnariidae. Journal of Ornithology 146:1–13. published. FJELDSA˚ , J., AND N. KRABBE. 1990. Birds of the High Andes. Zoology Museum, University of ACKNOWLEDGMENTS Copenhagen, Denmark. I wish to thank the many Venezuelan partners who HELLMAYR, C. E. 1938. Catalogue of birds of the stimulated my work with the Premnoplex barbtails: Americas and adjacent islands. Part IV. Field Violeta Go´mez Serrano was a wonderful field partner Museum of Natural History Publications 430, both in Yacambu´ and at Las Melenas, and David Zoology Series, Volume XIII, Chicago. Ascanio provided information on the places were P. HILTY, S. 1999. Three bird species new to Venezuela tatei could be found and shared his recordings of and notes on the behaviour and distribution of Roraimia. Laura Herna´ndez contributed some papers other poorly known species. Bulletin of the when the study was beginning. Miguel Lentino and British Ornithological Club 119:220–235. Margarita Herna´ndez at the Coleccio´n Ornitolo´gica HILTY, S. 2003. A guide to the birds of Venezuela. Phelps kindly guided me through their library and Princeton University Press, Princeton, NJ. grantedpermissiontostudythecollectionand IRESTEDT, M., J. FJELDSA˚ , AND P. G. P. ERICSON. consult the useful notes of the Phelps. The drawings 2004. Phylogenetic relationships of were skillfully prepared by Robin Restall, who also (Aves: Dendrocolaptinae) – incongruence be- provided crucial help, becoming an important tween molecular and morphological data. Jour- contributor to this publication despite the geograph- nal of Avian Biology 35:280–288. ical distance. The comments of Jorge Pe´rez and IRESTEDT, M., J. FJELDSA˚ , AND P. G. P. ERICSON. James Van Remsen greatly improved the structure 2006. Evolution of the – and content of this paper, which also benefited from assemblage (Aves: Furnariidae) – major shifts in suggestions made by Kristof Zyskowski and an nest architecture and adaptive radiation. Journal anonymous reviewer. This study was made possible of Avian Biology 37:260–272. in part by support from National Science Foundation JOHNSON, A. W. 1967. The birds of Chile and grants DEB-9981527 and DEB-0543178 to T. 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