ORNITOLOGIA NEOTROPICAL 21: 397–408, 2010 © The Neotropical Ornithological Society

THE NEST OF THE BAY-RINGED TYRANNULET ( SYLVIOLUS), A LITTLE-KNOWN ATLANTIC FOREST ENDEMIC, SUPPORTS A CLOSE RELATIONSHIP BETWEEN PHYLLOSCARTES AND

Guy M. Kirwan1, Alejandro Bodrati2,3,4, & Kristina Cockle2,3,5

174 Waddington Street, Norwich NR2 4JS, UK. E-mail: [email protected] 2Proyecto Selva de Pino Paraná, Vélez Sarsfield y San Jurjo S/N, San Pedro, Misiones, CP 3352, Argentina. 3Fundación de Historia Natural Félix de Azara, Departamento de Ciencias Naturales y Antropología, Universidad Maimónides, Valentín Virasoro 732, Buenos Aires (C1405BDB), Argentina. 4Grupo FALCO, Los Ceibos 1695 (1607), Villa Adelina, Prov. de Buenos Aires, Argentina. 5Center for Applied Conservation Research, Faculty of Forestry, University of British Columbia, 2424 Main Mall, Vancouver, BC, V6T 1Z4, Canada.

Resumen. – El nido de la Mosqueta Cara Canela (Phylloscartes sylviolus), una especie poco conocida y endémica de la Mata Atlântica, apoya el estrecho parentesco entre Phylloscartes y Pogonotriccus. – Existen pocos datos sobre la biología reproductiva de la Mosqueta Cara Canela Phyl- loscartes sylviolus. Aquí describimos cuatro nidos encontrados en varios momentos durante la etapa de construcción, en el este de Paraguay, noreste de Argentina y sudeste de Brasil. Todos los nidos se pueden describir como cerrado/esférico/lateral o quizás cerrado/retorta/colgado, según el esquema reciente para la clasificación de los nidos de las aves Neotropicales. Los nidos eran mayormente verdes, construidos de musgo vivo, pelusa de semillas, otras fibras vegetales, tela de araña, y líquenes. Los dos adultos contribuyeron con la construcción del nido. El único huevo que observamos era blanco limpio. Recientes estudios moleculares han encontrado fuerte apoyo para un parentesco cercano entre Phyllo- scartes y el género Pogonotriccus, e incluso han sugerido que un mayor muestreo podría apoyar la reunificación de estos géneros; un estudio reciente de arquitectura de nidos, en contraste, propone que Phylloscartes se asemeja más a Leptopogon y Mionectes. Nuestra revisión de los datos de arquitectura de nidos para estos géneros sugiere un grado mucho mayor de plasticidad que lo que ha sido reconoc- ido, por lo menos dentro de Phylloscartes, y similitudes llamativas entre los nidos de Phylloscartes y Pogonotriccus. Otros géneros cercanos, especialmente Mionectes y Leptopogon, construyen nidos obvi- amente diferentes. El sistema para clasificar nidos, presentado por Simon & Pacheco (2005), aunque no es perfecto, merece mayor uso por los ornitólogos describiendo nidos, para facilitar las comparaciones.

Abstract. – There are few data concerning the breeding biology of the Near Threatened Atlantic Forest endemic, the Bay-ringed Tyrannulet Phylloscartes sylviolus. Here, we describe four nests from eastern Paraguay, northeast Argentina and southeast Brazil, all found at various stages of construction. All four nests can be described as closed/globular/lateral, or perhaps closed/retort/pensile, according to the recent classification scheme for nests of Neotropical . Nests were mostly green, constructed of live moss, seed down, other plant fibers, spider webs, and lichen. Both adults contributed to nest-building. The single egg we observed was clean white. Recent molecular studies have found strong support for a close relationship between Phylloscartes and the Pogonotriccus, and have even suggested that

397 KIRWAN ET AL. additional sampling might support their reunification; a recent study of nest architecture, in contrast, pro- poses that Phylloscartes is more similar to Leptopogon and Mionectes. Our review of nest architecture data for these genera suggests a much greater degree of plasticity than has heretofore been recognized, at least within Phylloscartes, and striking similarities between the nests of Phylloscartes and Pogonotric- cus. Other genera, especially Mionectes and Leptopogon, build obviously different types of nests. The system for categorizing nests of Neotropical birds devised by Simon & Pacheco (2005), while not always perfect, merits greater use by ornithologists describing nests, in order to facilitate future comparisons. Accepted 8 August 2010.

Key words: Atlantic Forest, Bay-ringed Tyrannulet, nesting, Phylloscartes sylviolus, Pogonotriccus.

INTRODUCTION Picinguaba, east of Ubatuba, São Paulo, in southeast Brazil. It is not possible to gain a Limited data are available concerning the clear impression of the materials used, but the breeding biology of the Bay-ringed Tyrannu- closed nest was high above the ground, well let Phylloscartes sylviolus, a poorly known and attached to a ‘pocket’ in a branch, at a point Near Threatened endemic of the Atlantic where the latter was broadly vertical, and the Forest, which ranges from Espírito Santo nest was obviously longer top to bottom than south to Santa Catarina, Brazil, as well as it was wide, with a lateral entrance, and was through the province of Misiones in north- pensile but not pyriform. Finally, Aleixo & east Argentina and eastern Paraguay (Fitz- Galetti (1997) report adults with fledged patrick 2004, BirdLife International 2009). young in February in southeastern Brazil. Narosky & Salvador (1998) briefly reported Here, we report on four nests of P. sylviolus observations of two nests under construction and present more detailed information than in Parque Nacional Iguazú, Misiones, Argen- has appeared in the literature previously. We tina, the first in October 1970 (M. Rumboll) review knowledge concerning the nest archi- and the second in August 1991 (C. Saibene). tecture of the genus Phylloscartes in general, Nests were 10 m and 16 m above ground, and that of its apparent closest relative, attached to horizontal branches, and con- Pogonotriccus. We offer remarks on the relation- structed mainly of moss with a lateral ship between these two genera based on our entrance hole; the first nest was apparently findings and contrast these with that of ëhangingí but the second was attached to the another recent publication that addressed this branch and hidden within a bunch of Rhipsalis problem (Greeney 2009). (Cactaceae). At the second nest , both mem- bers of the pair were observed carrying nest METHODS materials. Lowen et al. (1996) report a nest, also under construction, in the canopy of a Nests were found between 2001 and 2009 at 30-m-tall Aspidosperma polyneuron (Apocy- three sites in the Atlantic Forest. The first naceae) tree at Reserva Natural Privada Itabó, nest was at Reserva Privada Itabó (Rivas) eastern Paraguay, in mid-October 1995. No (24o30’S, 54o38’W), a 5000-ha private forest details on materials or structure were pre- reserve in the department of Canindeyú, Par- sented, but one of the observers, M. Pearman aguay (the same site as the nest mentioned by (in litt. 2010), recalled that it was placed on Lowen et al. 1996). The second and third top of a large horizontal branch c. 25 m above nests were at Parque Provincial Cruce Cabal- the ground. Remold (2002) presents a photo- lero (26o31’S, 53o59’W), a 600-ha protected graph of a nest found in mid-October 1999 at area situated between a large privately owned

398 NEST OF BAY-RINGED TYRANNULET secondary forest (in the valley of the Arroyo adult feeding a juvenile in the native canopy Alegría) and an area of small farms with rem- of the same plantation. nant forest, in the province of Misiones, The second nest was discovered by KC on Argentina. The fourth nest was at Parque 22 October 2006, at c. 600 m a.s.l., 300 m Estadual Intervales (24o15’S, 48o10’W), a from the camping area in Parque Provincial 41,700-ha conservation unit in southern São Cruce Caballero. The nest was in primary for- Paulo state, Brazil. We found all nests during est, 18 m above the ground on the trunk of a the construction phase and watched them 30-m tall Paraná Pine (Araucaria angustifolia; using binoculars, making documentary photo- Araucariaceae). It could only be observed graphs of the nests and adults, and sound- from the bottom, and was in the early stages recordings of the adults in several instances. of construction. It appeared as a thin, partial These photographs and sound-recordings are sphere of soft, pale green materials, attached available on request from the authors. We among small ferns and moss on the vertical climbed to one of the Argentine nests and tree trunk, well below the lowest branches. measured it using a 15-cm wing-rule. Both members of the pair added material inside the nest, entering through the bottom. RESULTS On 23 October, KC watched the nest from 08:14 to 08:26 h and saw the pair enter the The first nest was discovered at c. 300 m a.s.l. nest six times, but she was unable to ascertain and c. 400 m from the Arroyo Pozuelo at whether the birds were bringing new material Reserva Privada Itabó in late November 2001. or only adjusting what was already there. On This nest was attached to the trunk of a 27 October, the nest had changed little and large cedro (Cedrela fissilis; Meliaceae) that was still far from completion. The adults were formed part of the native tree canopy over an not seen in this area again, despite weekly vis- 80-ha plantation of shade-grown yerba its throughout November. mate within the reserve (see Cockle et al. The third nest (Fig. 1A) was discovered by 2005). The nest was c. 17 m above the AB at 600 m a.s.l. in the camping area of ground, at the base of a small epiphytic Parque Provincial Cruce Caballero, c. 300 m guembé (Philodendron sp.; Araceae) surrounded from the second nest and c. 20 m from a by epiphytic bromeliads. The nest was hidden small creek, on 16 November 2008 and but we could see that it was globular, with a observed daily until 22 November. It was 15 closed top and a lateral hole in the top third of m above the ground, behind a dead epiphytic the side, pointing 90º from the tree trunk. The cactus (Rhipsalis sp.; Cactaceae), 2 m above an adults made repeated trips to a nearby cedro active ant nest, attached firmly to a large, to collect nest material. One of the adults mossy, vertical branch of a Grapia (Apuleia would search for and collect nest material leiocarpa; Fabaceae) c. 25 m tall. Some from behind some epiphytic bromeliads on branches of the cactus passed through the this tree, then after some time among these underside of the nest, helping to support it. plants it would fly to the nest. AB & KC On 16 November, the nest was discovered observed this behavior on several different when both adults arrived together at 17:35 h. days, in the last week of November 2001. One The nest would later be closed and globular, of the adults made the trips to collect nest with a very short lateral entrance ‘tunnel’ in material and the other sometimes accompa- the top third of the side pointing 90º from the nied it, but did not seem to directly assist. In tree trunk (Fig. 1B); however, on 16 Novem- late February 2002, the same observers saw an ber the adults were still visible inside the nest

399 KIRWAN ET AL.

FIG. 1. Nest of Bay-ringed Tyrannulet Phylloscartes sylviolus in different phases: A) Adult P. sylviolus with Nest 3 in construction on 19 November 2008 at Parque Provincial Cruce Caballero, Argentina (M. Lam- mertink), B) Nest 3 completed and containing one egg on 22 November 2008 at Parque Provincial Cruce Caballero, Argentina (K. Cockle). because the tunnel was not constructed and nessed two further visits, with both adults the top of the nest chamber not fully closed. adding or adjusting material inside the nest. AB watched the nest until 18:55 h and wit- On one of the visits the adults copulated

400 NEST OF BAY-RINGED TYRANNULET briefly c. 2 m above the nest. On 17 Novem- 1A). On 20 November, during 2.5 h of obser- ber, AB watched the nest from 06:25 to 08:30 vation, only one individual was seen and it did h, observing 19 visits to bring or adjust mate- not enter the nest. On 21 November there rial in the nest. All construction occurred was no activity in 3 h of observation by J. Seg- inside the nest, which was still relatively open ovia and E. Jordan, and we suspected the nest around the entrance and roof. Sometimes might have been abandoned. On 22 Novem- both individuals entered the nest. Other ber, the adults were not seen during the morn- times, only one individual entered while the ing and KC inspected the nest. It measured 86 other waited in a smaller nearby tree. mm tall × 65 mm side to side × 85 mm front During this period, they copulated twice. One to back. Its internal horizontal depth was 73 of the individuals seemed to have a paler cin- mm from the inside of the entrance tunnel to namon color to the face. This individual the back of the nest chamber. Although the appeared to be more involved in the nest con- entrance was covered completely by the tun- struction and spent more time in the nest. In nel roof, the lower lip of the entrance tunnel the evening, the nest was observed for 1 h 40 was shorter (c. 3 cm). The nest did not swing min, during which time the adults visited it freely but was set firmly among living and three times. One individual brought material dead epiphytes attached to the tree trunk (Fig. each time, and the other only once. On 18 1B). The outside of the nest was constructed November, AB watched the nest from 06:40 of moss, spider web, some relatively hard to 07:40 h. The nest had advanced consider- plant fibres, and white and orange lichen. The ably and the entrance tunnel appeared to be egg chamber was covered in soft white plant under construction. At 06:45 h one of the fibres, probably seed down. The nest con- adults entered the nest, remaining there for tained one white egg (presumably an incom- six minutes. It was no longer visible inside the plete clutch, given that clutch size is nest because the entrance had been built up. apparently three in P. ventralis: Dabbene 1919, The other bird remained nearby. Both vocal- Smyth 1928, Narosky & Salvador 1998). ized, including the bird inside the nest. They The fourth nest was discovered, by GMK, seemed to maintain vocal contact even at c. 650 m a.s.l. beside the Carmo road, in though the individual outside the nest flew to Parque Estadual Intervales on 30 October about 20 m away. A pair of Piratic Flycatchers 2009, and was observed again on 31 October, Legatus leucophaius had a nest with two chicks, as well as more extensively on 1 November, c. 13 m away. At one point the pair of Legatus for a total of c. 2 hours. The nest, which was foraged for insects among the epiphytes along still under construction, was sited c. 15 m the branch less than 30 cm from the nest of P. above the ground on one of the principal sylviolus. A Phylloscartes in the nest did not react. trunks of a “bico do pato” (Machaerium sp., That evening the nest was watched from Fabaceae) tree at a point where it was near- 16:25 to 17:40 h. One individual was heard vertical, smooth, and covered in live lichens vocalizing away from the nest. On 19 Novem- and Tillandsia (Bromeliaceae). The tree was ber, the nest appeared complete. AB observed sited on the steep slopes of a valley. The nest it from 06:25 to 09:20 h. The pair arrived was closed and globular, and its outer layer three times; each occasion one individual appeared to be mainly constructed of moss entered the nest for 3–11 min, maintaining and or Tillandsia. Its dimensions were difficult vocal contact with the other. AB sound- to determine at the distance (c. 25 m) from recorded these vocalizations and M. Lam- which observations were made, but it was c. mertink photographed one of the adults (Fig. 10 cm in overall length and approximately

401 KIRWAN ET AL. two-thirds as wide, i.e. probably broadly simi- determined, but included a c. 2 cm-long plant lar to the nests at Parque Provincial Cruce fibre, some seed down, parts of dead leaves, Caballero and a nest of Serra do Mar Tyran- and twice parts of live leaves, suggesting that nulet Phylloscartes difficilis (Kirwan 2009), and the birds were finalizing the egg chamber. had a lateral side entrance. Both members of The altitude at which this nest was discovered the pair brought materials, usually alone but is marginally higher than the published eleva- occasionally together (twice during a one- tional range of the (Fitzpatrick 2004, hour observation period on 1 November, but Ridgely & Tudor 2009). also on other occasions on the other days). However, when this happened, only one indi- DISCUSSION vidual would visit the nest, whilst the other remained ‘on guard’ in the same tree approxi- The nests of P. sylviolus we found were broadly mately 5 m away (‘mate guarding’). As at nest similar to those previously reported for the 3, both members of the pair vocalized quite species. They were sited high on large vertical regularly. Visits were usually at a rate of one branches or trunks of large live trees. The two per ten minutes, but during each period when that could be observed closely were globular, a bird or birds were at the nest they would constructed mostly of moss, and at least three visit 2–3 times either with different items or of the four had lateral entrances. return to adjust material already present. Dur- In recent decades, especially, there have ing periods when both birds were present been several large-scale and numerous they would stay longer in the vicinity and visit smaller-scale contributions to the debate over the nest more frequently. On each occasion a generic relationships within the Tyrannidae. bird visited the nest, it was usually for < 10 In addition to the obvious studies of DNA, seconds, but occasionally longer. On c. 50% anatomy and morphology, voice, behavior of visits the birdís tail would be visible when and nest architecture, amongst other charac- it was entering the nest. Usually the birds ters, are being increasingly mobilized in sought materials some distance from the nest efforts to create a robust phylogeny for this (flying out of sight when leaving the vicinity) diverse family. Treatment of the genus Phyllos- but sometimes they collected materials in the cartes has varied during this period; it some- immediate vicinity, especially in the nest tree times has been expanded to include itself and two adjacent Cecropia (Urticaceae) Pogonotriccus (bristle tyrants). Relatively few trees (including once some vegetable matter species of Phylloscartes have been sampled from within a clump of dead leaves). Regular genetically to date; the recent nuclear DNA perches were utilized to approach or stay in study by Tello et al. (2009) included just three the general vicinity of the nest, c. 5 of these species (Black-fronted Tyrannulet P. nigrifrons, were in the most adjacent Cecropia, and two Mottle-cheeked Tyrannulet P. ventralis, and were in the nest tree, all of them within a 5-m Oustaletís Tyrannulet P. oustaleti), as well as radius of the nest. When approaching from one species of Pogonotriccus, Marble-faced Bris- long distance, the birds always staged in two tle Tyrant P. ophthalmicus. Tello et al. (2009), or three trees en route to the nest, and some- like Ohlson et al. (2008), recovered evidence times hovered briefly in front of it before of a sufficiently close relationship between entering, but generally the final approach was these two genera to suggest that Traylor direct. However, a small snag immediately (c. (1977: 154) might have been correct to con- 5 cm) below the nest was utilized only occa- sider them congeneric. Traylor also consid- sionally. Items brought generally could not be ered Leptotriccus (see below) part of his

402 NEST OF BAY-RINGED TYRANNULET expanded Phylloscartes. Traylor also included ton 1985, Narosky & Salvador 1998, Smith & Capsiempis (Yellow Tyrannulet C. flaveolus), but Betuel 2006), Alagoas (Long-tailed) Tyrannu- this suggestion has been refuted (e.g., Lanyon let P. ceciliae (B. M. Whitney in Collar et al. 1988). 1992), P. sylviolus (Lowen et al. 1996, Narosky The genus Phylloscartes is currently usually & Salvador 1998; this paper), Restinga Tyran- treated as constituting 16 species of primarily nulet P. kronei (Remold & Ramos Neto 1995), circum-Amazon distribution, one of which, P. roquettei (Kirwan et al. 2004), and Serra do Yellow-green Tyrannulet P. flavovirens, is Mar Tyrannulet P. difficilis (Kirwan 2009). restricted to eastern Panama (Fitzpatrick Only the eggs of P. ventralis (Dabbene 1919, 2004, Ridgely & Tudor 2009). Intra-generic Smyth 1928, Belton 1985, Narosky & Salva- relationships within this circumscribed view dor 1998) and P. sylviolus (this paper) have of Phylloscartes also have intrigued taxonomists been described (both are white). working with Tyrannidae. Several superspecies groupings have been suggested, most notably Nests of Phylloscartes. The nests of Phylloscartes that P. nigrifrons, Rufous-browed Tyrannulet P. have provided us with a problem of categori- superciliaris, Rufous-lored Tyrannulet P. flaviven- zation according to the classification system tris, Cinnamon-faced Tyrannulet P. parkeri, devised by Simon & Pacheco (2005). P. roquettei, and P. syl- Although all appear to be closed with a lateral violus might form an expanded superspecies, entrance, their attachment points may be con- although it is possible that this close relation- sidered ‘pensile’ (attached from above), ‘lat- ship should be confined to P. flaviventris and P. eral’ (attached laterally to substrate) and parkeri (Raposo et al. 2002, Fitzpatrick 2004, sometimes even ‘fork’ (sitting in a fork). To Maldonado-Coelho 2009). On the other hand, some extent, the P. sylviolus nests described P. sylviolus has sometimes been removed to its here present a midway point between closed/ own genus, Leptotriccus Cabanis & Heine, globular/lateral, or closed/retort/pensile, 1859, on the basis of minor structural features because the nests were somewhat pensile but (e.g., Hellmayr 1927), and a separate genus is also supported laterally (by the tree trunk). also available for P. difficilis, Guracava, H. von Nests 3 and 4 also were supported by vegeta- Ihering & R. von Ihering, 1907. However, tion on the trunk and did not move freely. A Rheindt et al. (2008) recovered molecular evi- recently discovered nest of P. difficilis can also dence to suggest that P. sylviolus is closely be considered ëmidwayí between pensile and related to P. superciliaris and, to a lesser extent, lateral, and was also constructed on a vertical P. ventralis, but his sampling of the genus was section of trunk (Kirwan 2009). Complicating limited to these three taxa. the classification of nest architecture, at least Given that nest architecture is recognized some species of Phylloscartes may show consid- to be a taxonomically informative character erable intraspecific variation in nest construc- among some suboscines (Sheldon & Winkler tion, as had already been noted by Simon & 1999, Zyskowski & Prum 1999, Miller & Pacheco (2005: 147) for some other Tyran- Greeney 2008), it is worthwhile to reconsider nidae. some of these suggested relationships in the Among the Phylloscartes, nest architecture is light of recently published breeding data. To best known for P. ventralis. We know of ten date, the nests of just six species of the genus nests of this species, all pertaining to the nom- Phylloscartes have been described, in varying inate subspecies and P. v. tucumanus, and levels of detail: Mottle-cheeked Tyrannulet P. described as closed with lateral entrances, usu- ventralis (Dabbene 1919, Klimaitis 1984, Bel- ally protected by a slight overhang, and

403 KIRWAN ET AL. attached to trees or lianas at a height of 1.5– province (erroneously omitted from the 7.0 m above the ground. Four of the nests speciesí range by Fitzpatrick) and therefore appeared somewhat longer than they were pertained to P. v. ventralis. Equally, the nest wide (Dabbene 1919, Klimaitis 1984, specifically mentioned for P. v. tucumanus Narosky & Salvador 1998). Three were con- seems to have been taken from Klimaitis structed using a variety of plant materials (1984), which too was based on observations including small twigs, liana fibres, spider in Buenos Aires. Fitzpatrick (2004) consid- webs, dry leaves, lichens, and moss (Klimaitis ered the season to be October–December in 1984, Narosky & Salvador 1998). Two Argentina, but Narosky & Salvador (1998) appeared to be constructed mostly of Tilland- also mention a January nest, and, for Brazil, sia usneoides (Bromeliaceae; Dabbene 1919, Belton (1985) reported various breeding data Belton 1985); in one of these the bird for most months between late August and appeared to have simply adapted a hanging December. The date of the only Uruguayan clump of Tillandsia for its own use (Belton nest reported to date is unclear, but probably 1985). This latter nest was suspended from “a refers to the same general period (Rocha small tree” and another was suspended from 2000). “a thin branch” (Belton 1985). In contrast, Few or no nests have been described for the nest found by Klimaitis (1984) was placed the remaining Phylloscartes species. Only one within an accumulation of dead leaves and nest of P. roquettei has been described, and it other litter between interlaced lianas and sur- was closed/retort/pensile (Kirwan et al. rounded by Rhipsalis (Cactaceae), and the one 2004). However, photographs of two recently found by Narosky & Saibene (per Narosky discovered nests of P. roquettei posted on the and Salvador 1998) was placed among internet, in both of which the identification of branches to which it was attached using abun- the adults appears correct, show moss-like dant spider webs above and in various places. nests supported against trunks and within Likewise, two nests found by AB (pers. tree forks (www..org and www.wikia- observ.), in Buenos Aires and Entre Ríos, ves.com), suggesting a structure that is Argentina, were both attached laterally to closed/globular/lateral or even closed/globu- trees at points where branches or twigs joined lar/fork. One nest of P. kronei has also been the main trunk, and a nest found by Smith & described (Remold & Ramos Neto 1995); the Betuel (2006: 21) was “placed on a thin, hori- side walls and back of the nest were sup- zontal moss-covered branch with two further ported by a total of five thin twigs (H. vertical branches providing a crotch-like sup- Remold in litt. 2010), so it can be considered port for the entrance”. Some of these nests closed/globular/lateral. Finally, there is a very are difficult to characterise on the basis of the brief description of a nest of P. ceciliae in Col- details presented, but most appear to be lar et al. (1992). The nest was a 45-cm pensile closed/globular/lateral. All were obviously structure sited 6 m above the ground and closed and none sounds truly pyriform, consisting of three parts: a 10-cm attachment although some do appear to have been pen- to the branch of the tree, the ball-like nest sile. Contra Fitzpatrick (2004: 299), we are chamber with a side entrance, and a 20-cm tail unaware of any description of the nest of P. v. of dangling material, all of them constructed angustirostris (of eastern Peru to Bolivia). The of moss-like material (B. M. Whitney pers. nest ascribed to that race in the latter work is comm.). This description suggests the nest transcribed from Dabbene (1919), whose was pensile, and quite different to all of the observations were made in Buenos Aires other nests of Phylloscartes species described to

404 NEST OF BAY-RINGED TYRANNULET date. Unlike the round nest of P. roquettei (Kir- P. poecilotis nest filmed in eastern Ecuador, wan 2009) this nest appears to have been pyri- archived on the Internet Bird Collection form, albeit with an additional tail of material (https://ibc.lynxeds.com) clearly shows a nest hanging below the nest chamber. To date, this similar in form and structure to those of Phyl- description is by far the most disparate of all loscartes sylviolus reported here and that of those available for the genus Phylloscartes. Pogonotriccus ophthalmicus reported by Greeney (2009, of which video evidence is also avail- Nests of Pogonotriccus. The nests of Pogonotric- able on the Internet Bird Collection web site). cus also are largely unknown. No nest descrip- tions are available for four species (Fitzpatrick Nest architecture and the relationship between Phyl- 2004) and the description for a fifth species loscartes and Pogonotriccus. Greeney (2009) appears erroneous, leaving only two species - considered relationships between Phylloscartes, Southern Bristle Tyrant P. eximius (the type Pogonotriccus, and other related small tyrannid species of Pogonotriccus) (Bertoni 1901) and genera. Contra the genetic data (cited above), Marble-faced Bristle Tyrant P. ophthalmicus nest architecture and behavioral innovations (Greeney 2009) - with valid nest descriptions. led him to speculate that Phylloscartes could be The nest of P. eximius found in Paraguay (Ber- united with Leptopogon and Mionectes, whereas toni 1901) was a mossy oven-shaped ball con- he considered Pogonotriccus to be closer to structed against a tree trunk, and clearly could Corythopis and Pseudotriccus in nest characters. be considered closed/retort/lateral in archi- Greeney (2009) regarded Phylloscartes as build- tecture. The nest of P. ophthalmicus discovered ing solely pendant nests. However, as by Greeney (2009) was placed 7 m above the reviewed above, Phylloscartes ventralis (Dabbene ground and in most general respects of form 1919, Klimaitis 1984, Belton 1985, Narosky & and structure closely matched the nests of Salvador 1998, Smith & Betuel 2006; AB pers. Phylloscartes sylviolus (described here) and other obs.), P. difficilis (Kirwan 2009), P. kronei (H. Phylloscartes, as did that of another nest of Remold in litt. 2010), and P. sylviolus (this Pogonotriccus ophthalmicus collected in Peru by T. paper) often (but not exclusively) build nests S. Schulenberg (also described in Greeney that are supported laterally, in the same man- 2009). We concur with Greeney (2009) that ner as the nest of Pogonotriccus ophthalmicus the nest of P. ophthalmicus mentioned in Hilty Greeney (2009) described. Like the nests of & Brown (1986) must either have been misi- Pogonotriccus, they are also globular structures dentified to species, or was described so built more or less well above the ground (this imprecisely (“a mossy cup … on a small seems to vary with the foraging strata of the forked branch”) as to unwittingly invite subse- species concerned, but nests are never sited quent confusion. There is also confusion below overhanging banks within root masses), about the nest of the Variegated Bristle Tyrant and frequently using green materials such as P. poecilotis. The nest reported to belong to this live moss. In contrast, the nests of Leptopogon species by Londoño & Muñoz (2006) was and Mionectes are more pyriform structures more likely that of a Slaty-capped Flycatcher that hang from roots or low plants close to Leptopogon superciliaris (see Greeney 2009, the ground, and are often constructed of whose conclusions we support, for a thor- brown materials such as dry plant fibres, ough discussion of this nest). Hilty & Brown though live moss can also be utilized to cover (1986) mentioned a ‘July nest’ of this species the outside, whilst those of Mionectes are sited that had been reported to them, but without toward the tip of long branches (e.g., Skutch additional information. However, a video of a 1960, Aguilar et al. 2000, Fitzpatrick 2004,

405 KIRWAN ET AL.

Greeney et al. 2006, Greeney 2009). To date, between Pogonotriccus and Phylloscartes, distinct of the available descriptions, the only individ- from Mionectes and Leptopogon. ual nest of Phylloscartes that could be described More information concerning the nests of as hanging freely and pyriform is the sole nest all Phylloscartes and Pogonotriccus species is found of P. ceciliae (Collar et al. 1992). It is also needed, not only for the role it can play in worth remarking in this respect that even the expanding our knowledge of their life histo- pensile nest of P. roquettei described in the lit- ries, and aiding their conservation (several erature (Kirwan et al. 2004) was still quite dif- species are considered to be globally threat- ferent from those of any known Leptopogon or ened), but also to help inform our knowledge Mionectes, in being placed high above the of inter- and intra-generic relationships. We ground suspended (but not swinging wholly suspect that Phylloscartes might prove to be free) from a tree branch (in perhaps the same polyphyletic, but that in terms of some manner as some of the nests of P. sylviolus aspects of nest architecture the genus, as cur- mentioned in the introduction), and in being rently constituted, appears to sit somewhere strongly globular rather than pyriform. While in the middle of a continuum, with Pseudo- few green materials, and no mosses, were triccus (which builds nests closely affixed to a used in its construction, this probably largely trunk) and Mionectes (which constructs highly reflected their relative lack in the vicinity of pensile nests) at opposite ends of this spec- the nest at this season (e.g., early October is trum. To this end, further data for those spe- still very dry in the region concerned). cies whose nests are already known, to a Greeney (2009) also drew attention to greater or lesser extent, also will be valuable, what he considered to be probably a second if only to evaluate the degree of plasticity in distinction, in how the nest is attached to the their breeding behavior (which is already evi- substrate: ‘stuffing’ (in Pogonotriccus) versus dent for at least two species of Phylloscartes, P. ‘draping’ (Phylloscartes) the nest. However, it ventralis, and P. roquettei). We believe that seems certain from our observations of robust (correctly identified to species) and Phylloscartes sylviolus and P. difficilis that these large samples are probably needed to infer two species are also ‘stuffers’, i.e. rather than patterns from nest architecture in at least attaching their nest by draping material over some groups, especially genera of wide geo- an attachment point and building the nest graphical range and preferences. We chamber within the resulting curtain (as in also encourage nest finders and describers to Leptopogon), they stuff material into small use the system proposed by Simon & Pacheco spaces between epiphytic plants, effectively (2005) for describing nests and to document creating a nest that is part of an existing sub- the species whenever possible by sound- strate. This is certainly true of P. difficilis, and recording or photographs, thereby providing seems very likely to have been the case in ready means for researchers to compare nest most or all of the P. sylviolus nests we architecture between different species and observed. The obviously pensile nest of P. syl- genera. violus found by Remold (2002) and at least one of the nests of P. ventralis found by Belton ACKNOWLEDGMENTS (1985) appear also to have been constructed using the advantages afforded by an already We thank Kaspar Delhey, Ignacio Roesler, suitable substrate. As such, contra Greeney Emilse Merida, and Juan Ignacio Areta for (2009) we consider that nest architecture, like help with references. Hadoram Shirihai, Faus- the genetic data, points to a close relationship tino Avelino Ribeiro, Emilio Jordan, José Seg-

406 NEST OF BAY-RINGED TYRANNULET ovia, Andrea Norris, and Martjan Lammertink Wege. 1992. Threatened birds of the Americas: shared in field observations. Heinz Remold the ICBP/IUCN Red Data book. International kindly and swiftly clarified the non-pensile Council for Bird Preservation, Cambridge, UK. nature of the P. kronei nest he found in Brazil, Dabbene, R. 1919. Nido y huevos del tiránido and informed us of his observations of a P. Phylloscartes ventralis angustirostris [sic]. Hornero 1: 292. sylviolus nest. Mark Pearman sent information Fitzpatrick, J. W. 2004. Genera Phylloscartes, Pogonot- concerning his personal observations. We riccus, Leptopogon and Mionectes species accounts. thank John Bates and an anonymous referee Pp. 299–305 in del Hoyo, J., A. Elliott, & D. A. for their useful comments on the submitted Christie (eds.). Handbook of the birds of the version of the manuscript. The work in Para- world. Volume 9: Cotingas to pipits and wag- guay was funded by a Canadian International tails. Lynx Edicions, Barcelona, Spain. Development Agency Innovative Research Greeney, H. F. 2009. A nest of the Marble-faced Award, NSERC Post Graduate Scholarship, Bristle Tyrant (Pogonotriccus ophthalmicus) with Bergstrom Memorial Research Award, and in- comparative comments on nests of related gen- kind contributions from Guayakí Yerba Mate. era. Wilson. J. Ornithol. 121: 631–634. The work in Argentina was funded by the Greeney, H. F., C. Dingle, R. C. Dobbs, & P. R. Martin. 2006. Natural history of Streak-necked Neotropical Bird Club, Rufford Foundation, Flycatcher Mionectes striaticollis in north-east Killam Predoctoral Award, and NSERC Can- Ecuador. Cotinga 25: 59–64. ada Graduate Scholarship, and was authorized Hellmayr, C. E. 1927. Catalogue of birds of the by the Ministry of Ecology, RNR y Turismo Americas and the adjacent islands. Part 5, Tyr- of the province of Misiones. annidae. Field Mus. Nat. Hist., Publ. 242, Zool. Ser. 13, Chicago, Illinois. REFERENCES Hilty, S. H., & W. H. Brown. 1986. A guide to the birds of Colombia. Princeton Univ. Press, Prin- Aguilar, T. M., M. Maldonado-Coelho, & M. Â. ceton, New Jersey. Marini. 2000. Nesting biology of the Grey- Kirwan, G. M. 2009. Notes on the breeding biol- hooded Flycatcher (Mionectes rufiventris). Ornitol. ogy and seasonality of some Brazilian birds. Neotrop. 11: 223–230. Rev. Bras. Ornitol. 17: 121–136. Aleixo, A., & M. Galetti. 1997. The conservation of Kirwan, G. M., J. Mazar Barnett, M. F. Vasconcelos, the avifauna in a lowland Atlantic forest in M. A. Raposo, S. D. Neto, & I. Roesler. 2004. south-east Brazil. Bird Conserv. Internat. 7: Further comments on the avifauna of the mid- 235–261. dle São Francisco Valley, Minas Gerais, Brazil. Belton, W. 1985. Birds of Rio Grande do Sul, Bra- Bull. Br. Ornithol. Club 124: 207–220. zil. Part 2. Formicariidae through Corvidae. Klimaitis, J. F. 1984. Nota sobre un nido de la Bull. Am. Mus. Nat. Hist. 180: 1–242. Mosqueta Vientre Amarillo (Phylloscartes ventra- Bertoni, A. de W. 1901. Aves nuevas del Paraguay. lis) en Punta Lara, Ensenada, Buenos Aires, H. Kraus, Asunción, Paraguay. Argentina. Hornero 12: 203–204. BirdLife International. 2009. Species factsheet: Lanyon, W. E. 1988. A phylogeny of thirty-two Phylloscartes sylviolus. Downloaded on 9 Decem- genera in the Elaenia assemblage of tyrant fly- ber from http://www.birdlife.org. catchers. Am. Mus. Novit. 2914. Cockle, K., M. Leonard, & A. Bodrati. 2005. Pres- Londoño, G., & M. C. Muñoz. 2006. Primera ence and abundance of birds in an Atlantic for- descripción del nido del Atrapamoscas Varie- est reserve and adjacent plantation of shade- gado (Tyrannidae: Pogonotriccus poecilotis). Orni- grown yerba mate. Biodivers. Conserv. 14: tol. Colombiana 4: 66–69. 3265–3288. Lowen, J. C., L. Bartrina, R. P. Clay, & J. A. Tobias. Collar, N. J., L. P. Gonzaga, N. Krabbe, A Madroño 1996. Biological surveys and conservation pri- Nieto, L. G. Naranjo, T. A. Parker, & D. C. orities in eastern Paraguay. CSB Conservation

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