70 (3): 319 – 331

© Senckenberg Gesellschaft für Naturforschung, 2020. 2020

Alpha of stictothorax group (Aves: Passeriformes: Furnariidae): Synallaxis chinchipensis Chapman, 1925 as a valid , with a lectotype designation

Renata Stopiglia 1, 2, 3,*, Flávio A. Bockmann 3, 4, Claydson P. de Assis 2 & Marcos A. Raposo 2

1 Museu de História Natural do Ceará Prof. Dias da Rocha, Centro de Ciências da Saúde, Universidade Estadual do Ceará, Av. Dr. Silas Mun- guba, 1700, Campus Itaperi, 60741 – 000, Fortaleza, CE, Brazil — 2 Setor de Ornitologia, Departamento de Vertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s/n, 20940 – 040 Rio de Janeiro, RJ, Brazil — 3 Laboratório de Ictiologia de Ribeirão Preto, Departamento de Biologia, FFCLRP, Universidade de São Paulo, Av. dos Bandeirantes 3900, 14040 – 901 Ribeirão Preto, SP, Brazil — 4 Programa de Pós-Graduação em Biologia Comparada, FFCLRP, Universidade de São Paulo, Av. dos Bandeirantes 3900, 14040 – 901 Ribeirão Preto, SP, Brazil — * Corresponding author; email: [email protected]

Submitted July 10, 2019. Accepted July 6, 2020. Published online at www.senckenberg.de/vertebrate-zoology on August 7, 2020. Published in print Q3/2020. Editor in charge: Martin Päckert

Abstract The Synallaxis stictothorax group comprises poorly understood South American Furnariidae. This paper aims to present the morphological and nomenclatural aspects of this group, re-describing its valid species and to propose a fresh nomenclatural treatment for group members. Our analysis corroborated the specifc status of the disputed taxon Synallaxis chinchipensis and refuted the diagnostic characteristics of the subspecies Synallaxis stictothorax maculata. Following the recommendations of the Code, a lectotype was designated to the nominal species Synallaxis hypochondriaca. We also draw attention to the need for continued review of the taxonomy of polytypic species, as this is the most effcient way of distinguishing natural groups from those that are merely historical artefacts of taxonomy.

Key words Synallaxis chinchipensis, Synallaxis hypochondriaca, subspecies, Taxonomy.

Introduction

Within the genus Synallaxis Vieillot, 1818, nearly 90 taxa discussion, and the status of subspecies of some taxa re- are considered valid (e.g. Dickinson & Christidis, 2014), mains disputed. including those belonging to the clade composed of The frst discussions involving the relationships of Synallaxis stictothorax Sclater, 1859 and Synallaxis hy­ S. stictothorax to the other representatives of the family pochondriaca (Salvin, 1895) (Derryberry et al., 2011). Furnariidae began with Peters (1951), who retained it According to Derryberry et al. (2011), the Synallaxis in this genus as a valid species despite some misgivings. stictothorax species group encompasses Synallaxis sticto­ The author noted that the tail shape and plumage colour thorax Sclater, 1859, Synallaxis hypochondriaca (Salvin, patterns were discordant with those of other species then 1895) and Synallaxis zimmeri Koepcke, 1957. The hy- assigned to Synallaxis, calling attention to the similarity potheses of relationships between these species and other between S. stictothorax and S. hypochondriaca, at that genera of the family Furnariidae have a long history of time non-congeneric species. Although currently treated

ISSN 1864-5755 | eISSN 2625-8498 | DOI: 10.26049/VZ70-3-2020-05 319 Stopiglia, R. et al.: Taxonomy of Synallaxis stictothorax species group

in the genus Synallaxis, S. hypochondriaca is the type Since there is fagrant disagreement regarding the species by original designation of the monotypic genus validity of species and subspecies belonging to S. stic­ Siptornopsis Cory, 1919, a formally valid genus until tothorax species group, the main objective of this study Ohlson et al. (2013). Vaurie (1971, 1980) agreed that the is to evaluate the taxonomic status of each of them based relationships between S. stictothorax and the other spe- on morphology, re-describing those considered valid. In cies of genus Synallaxis required further consideration line with the results achieved, we undertook a review of although he was quite certain that S. stictothorax belongs their nomenclatural aspects, respecting the guidelines ex- to Synallaxis in stating “…that this species may not be pressed in the the Code (Anonymous, 1999). a Synallaxis seems totally unwarranted to me” (Vaurie, 1971: 21). Later, Remsen (2003) emphasized that S. stic­ tothorax may be more closely related to Siptornopsis or Reichenbach, 1853 than to Synallaxis, but Materials and Methods did not suggest any amendments to the nomenclature of the group. Derryberry et al.’s account (2011) was the frst to Both morphometric characteristics and plumage colour examine the relationships between Furnariidae from a were analysed from 130 specimens, including the type phylogenetic perspective including specimens of the specimens of Synallaxis stictothorax stictothorax Sclater, S. stictothorax group. In that study, the clade formed 1859, Synallaxis stictothorax maculata Lawrence, 1874, by S. stictothorax and S. zimmeri was inferred to be the Synallaxis stictothorax piurae Chapman, 1919, Synallax­ sister group of S. hypochondriaca. Later, as mentioned is stictothorax chinchipensis Chapman, 1925, Synallaxis below, Ohlson et al. (2013) merged Siptornopsis with hypochondriaca (Salvin, 1895) and Synallaxis zimmeri Synallaxis, thereby solving the issue of paraphyly of Koepcke, 1957 housed at nine scientifc institutions as Synallaxis. detailed in the Appendix. A total of 29 institutions were When investigating evolutionary trends in the pheno- consulted for analysis of specimens, but only nine had types and of furnariid taxa, Tobias et al. (2014) skins available for the study. used the same taxa as Derryberry et al. (2011), but add- The morphometric analysis included the following ed the subspecies Synallaxis stictothorax chinchipensis measurements: bill length (taken from exposed culmen Chapman, 1925 and Synallaxis stictothorax maculata to tip of bill), bill height (taken at level of the nostril), Lawrence, 1874, yielding results that were relevant to the wing length (chord), and tail length (length of central rec- taxonomy of the S. stictothorax species group. Among trices). The frst three measurements were obtained using these results, S. chinchipensis appeared as the sister a calliper (precision 0.05 mm), whereas the last measure- group of S. hypochondriaca, which was unexpected con- ment was obtained using both a calliper and millimetre sidering that S. chinchipensis is usually referred to as a ruler (precision 0.1 mm). The number of rectrices was subspecies of S. stictothorax. also recorded. For the plumage analysis, we documented At the intraspecifc level, although the majority of the colour for 14 areas of the body (ventral, dorsal, and catalogues considered S. chinchipensis as a subspecies of lateral sequence: throat, lateral throat, breast, abdomen, S. stictothorax (e.g. Peters, 1951; Vaurie, 1980; Remsen, side, fank, rectrices, upper tail coverts, back, crown, 2003; Dickinson & Christidis, 2014), popular handbooks forehead, superciliary line, wing coverts and remiges) us- (e.g. Sibley & Monroe, 1990; Ridgely & Tudor, 1994; ing Smithe (1975, 1981) coding to describe the colours, Ridgely & Greenfield, 2001; del Hoyo et al., 2016) and the anatomical topography of Proctor & Lynch overwhelmingly treated S. chinchipensis as a distinct (1993). species. Synallaxis s. maculata, in turn, was considered a Descriptive statistics (mean, standard deviation, min- junior synonym of S. stictothorax by Sclater (1874) un- imum and maximum values), normality tests and homo- til Cory & Hellmayr (1925), who considered this taxon scedasticity tests, scatterplots, principal component anal- as a subspecies of S. stictothorax, an opinion followed ysis (PCA type: covariance; matrix data log-transformed; by Peters (1951). Vaurie (1980), in his revision of the and pairwise deletion for missing data) and discriminant family Furnariidae, stated, “In Synallaxis stictothorax I analysis, were generated using the xlstat 2020.1.3 soft- recognise nominate stictothorax ranging from southwest- ware (Addinsoft, 2020) considering a signifcance level ern Ecuador to north-western Peru, and chinchipensis of 5%. In addition, qgis 2.4.0 was used to generate dis- in the valleys of Cajamarca”. Ridgely & Tudor (1994) tribution maps, which were based on georeferenced data, followed the decision by Vaurie (1980) in not consid- according to Stephens & Traylor (1983), and Paynter ering S. s. maculata a valid taxon, until Remsen (2003) (1993). Specimens that were unsexed, juvenile, moulting reversed such assessment. Synallaxis stictothorax piurae or damaged were not included in either the morphometric Chapman, 1919 was considered a junior synonym of S. s. or plumage analyses, as well as the specimen of S. zim­ maculata by Chapman (1925) and Cory & Hellmayr meri due to the fact that only one specimen was analysed (1925). This decision was followed by all subsequent au- (paratype AMNH 461650). thors (e.g. Peters, 1951; Ridgley & Tudor, 1994; Rem- sen, 2003; Dickinson & Christidis, 2014; del Hoyo et al., 2016).

320 VERTEBRATE ZOOLOGY — 70 (3) 2020

Table 1. Plumage colour of Synallaxis stictothorax, S. chinchipensis and S. hypochondriaca according Smithe (1975, 1981) and topogra- phy according Proctor & Lynch (1993). Asterisk (*) for the diagnostic character.

S. stictothorax S. chinchipensis S. hypochondriaca Character Character state Smithe’s colours (%) (%) (%) Throat White — 100 100 100 Lateral White — 89 63 100 throat White + spot cinnamon 39 11 37 0 White (± macula or stretch Breast (± 121 or 221) 100 100 100 marks brown) Abdomen White — 100 100 100 Cinnamon 39 100 0 0 White (± macula or stretch (± 121 or 221) 0 100 0 Side* marks brown) White + umber (± stretch 123 (± 121 or 221) 0 0 100 marks brown) Cinnamon 39 100 0 0 White + cinnamon (± maculas Flank* 39 (± 121 or 221) 0 100 0 or stretch marks brown) White + umber 123 0 0 100 Rufous (± Brown) 340 or 136 (± 221) 100 100 0 Rectrices Dark drab (± Brown) 119B (± 221) 0 0 100 Upper tail Rufous 340 or 136 100 100 0 coverts Dark drab 119B 0 0 100 Back Dark drab (± Rufous) 119B (± 340) 100 100 100 Crown Dark drab (± Brown) 119B (± 119A) 100 100 100 White (± stretch marks brown) (± 121 or 221) 100 0 0 Dark drab + cinnamon 119B + 39 Forehead* 0 100 0 (± stretch marks brown) (± 121 or 221) Dark drab 119B 0 0 100 Superciliary White — 100 0 100 line Cinnamon 39 0 100 0 Wing Rufous 340 or 136 100 100 100 coverts Remiges Brown 121 or 221 100 100 100

Results Therefore, when considered together, the two principal components accounted for 97% of the total morphomet- ric variation. The plumage analysis clearly distinguished Synallaxis Concerning the discriminant analysis, 100% of the stictothorax chinchipensis Chapman, 1925 from the re- S. stictothorax and S. hypochondriaca specimens (previ- maining taxa, based on the colour patterns of the side, ously identifed by plumage characters) could be correct- fank, and forehead (Table 1 and Figs 1 to 3), which are ly identifed on the basis of morphometric measurements, unique character states and, therefore, fully diagnostic. whereas 94% of S. chinchipensis specimens were cor- Table 2 presents the descriptive statistical analysis of rectly identifed, with just one individual (MVZ 163833) the morphometric data of the studied taxa. The results being incorrectly identifed as S. stictothorax. Therefore, of normality and homoscedasticity tests indicated the 99% of the specimens could be correctly identifed as use of data log-transformed for analyses. Despite the S. stictothorax, S. chinchipensis and S. hypochondriaca morphometric similarity between S. stictothorax and based on the discriminant analysis of the morphometric S. chinchipensis when compared to S. hypochondriaca, data. Those data strongly corroborate the hypothesis of the principal component analysis (Fig. 3) and discrimi- the existence of three different morphological groups, as nant analysis corroborated the division of the taxa into suggested by plumage colour. three distinct groups: S. stictothorax, S. chinchipensis There was no overlap between the ranges of bill and S. hypochondriaca. In the principal component anal- height and wing length when comparing S. hypochondri­ ysis, the frst component (PC1) accounted for 92% of the aca to either S. stictothorax or S. chinchipensis (Table 2), morphometric variation and was mainly infuenced by the values of which were consistently smaller than those tail and bill length, whereas PC2 was responsible for 5% of S. hypochondriaca. The results also indicated that the of the variation, being mainly infuenced by bill length. bill length of S. stictothorax was diagnostically smaller

321 Stopiglia, R. et al.: Taxonomy of Synallaxis stictothorax species group

A B

C D

Fig. 1. Specimens that illustrate the morphological variation between Synallaxis chinchipensis (left) and S. stictothorax (right): A) ventral view, MVZ 163831 and MVZ 163683; B) dorsal view, MVZ 163831 and MVZ 163683; C) lateral view MVZ 163831 and MVZ 163683; and D) forehead detail, AMNH 182060 and AMNH 129789. than that of S. hypochondriaca; however, such a distinc- In addition, the means of the morphometric measure- tion was not observed for the bill lengths of S. chinchi­ ments of S. stictothorax were always smaller than those pensis and S. hypochondriaca as the ranges of values of S. chinchipensis, but these could not be used to dis- observed for the two species overlapped. criminate the species as they displayed overlapping rang-

322 VERTEBRATE ZOOLOGY — 70 (3) 2020

A B C

Fig. 2. Specimens that illustrate the morphological variation between Synallaxis hypochondriaca (left) (AMNH 523868, paralectotype of Siptornis hypochondriacus Salvin, 1895) and Synallaxis stictothorax (right) (AMNH 129789): A) ventral view; B) dorsal view; and C) side view.

Fig. 3. Scatterplots showing the results of Principal Component Analysis for the morphometric data of Synallaxis stictothorax, S. chinchipen­ sis and S. hypochondriaca. *Illustrations from del Hoyo, J., Elliott, A., Sargatal, J., Christie, D.A. & de Juana, E. (2017). Handbook of the of the World Alive, Barcelona, Lynx Edicions (retrieved from http://www.hbw.com on 10.10.2017).

323 Stopiglia, R. et al.: Taxonomy of Synallaxis stictothorax species group

es. The means of female specimens were also consistent- ly smaller than those of the male specimens, regardless of

12 species. n , number

Rectrices Regarding their geographical distribution, S. sticto­ thorax, S. chinchipensis and S. hypochondriaca are allo­ 4 12 6 12 7 10 n 11 12 10 19 10 742348 10 10 10 patric, with no known overlap (Fig. 4). Synallaxis sticto­ thorax was found in southwest Ecuador and northwestern Peru, below 400 meters, whereas S. hypochondriaca and S. chinchipensis were both found in the Río Marañón valley, Cajamarca, Peru. However, S. hypo­chon­driaca min – max and S. chinchipensis were separated by more than 2,000 meters of altitude, as S. chinchipensis was only collected

Tail length (mm) Tail in the margins of the Chinchipe River, between 400 and del oyo ± SD 600 m (according to H , 2020) while S. hypochon­

85.00± 7.91 69.00 – 94.00 driaca specimens occur between 2,000 and 2,800 meters elevation (according to Remsen, 2003 and Lloyd, 2020). The altitudinal difference between distributions of the 4 85.50± 10.41 71.00 – 94.00 6 87.33± 1.51 85.00 – 89.00 n 11 15 67.38± 3.14 60.00 – 72.00 10 64.64± 2.53 61.00 – 68.00 25 66.37± 3.16 60.00 – 72.00 8125 55.55± 3.5953 48.00 – 69.00 55.15± 3.35 50.00 – 60.00 55.85± 3.76 48.00 – 69.00 species of this group was, was, apparently, not consid- ered by BirdLife International (2020) when trying to stablish the geographical occurrence of S. chinchipensis. Apparently BirdLife International (2020) estimated the area of occurrence of S. chinchipensis as the entire min – max Marañón River Valley, disregarding the fact that the spe- cies is restricted to a small portion of the valley between

Wing length (mm) Wing 400 and 600 meters. Fig. 4 illustrates the distribution of . , mean; SD, standard deviation; min, minimum values; max, maximum values; maximum max, values; minimum min, deviation; standard SD, mean; , .

± SD the species, based on altitude information and specimens from scientifc collections. This spatial distribution of the species in the Andean Mountains may correspond to the differences found in 3 66.09± 2.57 63.18 – 68.57 6 61.22± 1.01 60.47 – 63.11 9 63.33± 2.91 60.47 – 68.57 n 15 51.64± 1.32 49.41 – 53.65 10 50.40± 1.88 47.89 – 53.63 25 51.14± 1.65 47.89 – 53.65 7224 50.58± 1.6145 46.01 – 55.11 49.68± 1.26 47.67 – 52.11 51.05± 1.61 46.01 – 55.11 the two species’ body size, a pattern already well known in that region (e.g. Winger & Bates, 2015). This corre-

and S . hypochondriaca lates with the predictions of Bergmann’s rule, which pro- poses that greater body size is benefcial to individuals that face lower temperatures (Bergmann, 1847; Graves, min – max 1985). Bill height (mm) , S . chinchipensis

± SD Synallaxis stictothorax Sclater, 1859 4.43± 0.25 4.19 – 4.97 3.43± 0.14 3.16 – 3.63 Synallaxis stictothorax Sclater, 1859: 191. Syntypes, by original des- stictothorax

ignation: NHMUK 1841.4.2.471 and NHMUK 1886.6.24.455, 4 4.64± 0.36 4.25 – 4.97 6 4.32± 0.10 4.19 – 4.45 n aynter 11 11 15 3.64± 0.10 3.43 – 3.81 26 3.56± 0.16 3.16 – 3.81 7725 3.19± 0.1549 3.16± 0.17 2.82 – 3.52 3.20± 0.14 2.82 – 3.50 2.85 – 3.52 both from Guayaquil, Ecuador; according to P (1993) the coordinates are 02°10′S, 79°50′W, at sea level. Lectotype, by present designation: NHMUK 1841.4.2.471, adult male (see Remark 1). Synallaxis stictothorax maculata Lawrence, 1874: 186. Holotype, by original monotypy: Museum of Vassar College nº 2437,

min – max from Tumbes, Peru; according to Stephens & Traylor (1983) the latitude and longitude are 03°34′S, 80°28′W, respectively (see Remarks 2 and 3). Bill length (mm) Synallaxis stictothorax piurae Chapman, 1919: 257. Holotype, by

± SD original designation: AMNH 163085, adult female from Chila- co, near Samate (= Somate), on the Río Chira, Piura, Peru, ca. 0.4611.51± 10.35 – 12.77 0.4411.57± 10.39 – 12.39 0.4511.51± 10.35 – 12.77 18.00± 1.91 15.46 – 19.66 16.64± 0.91 15.48 – 17.93 16.99± 1.55 15.01 – 19.66 14.64± 0.74 12.93 – 15.84 14.25± 0.99 12.15 – 15.40 14.48± 0.86 12.15 – 15.84 100 m elevation; the coordinates are 04°43′S, 80°31′W, ac- cording to Stephens & Traylor (1983) and LeCroy & Sloss (2000) (see Remark 3).

Diagnosis. Synallaxis stictothorax is distinguishable from S. hypochondriaca and S. chinchipensis by the colour of the lower part of the side of the body and fanks, which is cinnamon (39), contrasting with the umber (123) lower Males Females S. hypochondriaca General Males Females S. stictothorax General Females Males S. chinchipensis General Descriptive statistics of morphometric data of Synallaxis of morphometric statistics 2. Descriptive Table of specimens; mm, millimeters. part of the sides of the body with brown spot-streaks (121

324 VERTEBRATE ZOOLOGY — 70 (3) 2020

Fig. 4. Map of northwestern South America with the plotted distributions of Synallaxis stictothorax (yellow diamond), S. hypochondriaca (green diamond) and S. chinchipensis (blue diamond) based on examined skins, being that one symbol may represent more than one speci- men. or 221) in S. hypochondriaca, and the white with brown from 46.01 to 55.11 mm in S. stictothorax, versus from spot-streaks (121 or 221) of S. chinchipensis. Addition- 60.47 to 68.57 mm in S. hypochondriaca. Synallaxis ally, the white forehead with spot-streaks (121 or 221) of stictothorax is still diagnosable from S. chinchipensis S. stictothorax contrasts with the dark drab (119B) fore- by having: a cinnamon (39) fank, which is white and head of S. hypochondriaca and the dark drab (119B) or cinnamon (39) with brown spot-streaks (121 or 221) in cinnamon (39) with brown stretch marks (121 or 221) S. chinchipensis; and a white superciliary line, versus the of S. chinchipensis. Synallaxis stictothorax is further di- cinnamon (39) one of S. chinchipensis. Finally, S. sticto­ agnosable from S. hypochondriaca by possessing: fve thorax is also distinguishable from all other species of pairs of rectrices and rufous (340 or 136) upper tail cov- the genus Synallaxis by its white breast with brown spot- erts, compared to S. hypochondriaca, which presents six streaks (121 or 221; see Figs. 1 and 2 and Tables 1 and 2). pairs of rectrices and dark drab (119B) upper tail coverts; bill length varying from 10.35 to 12.77 mm, compared to Description. White throat; lateral part of the throat white S. hypochondriaca with bill length ranging from 15.01 with cinnamon spot-streaks (39) (on each side) on some to 19.66 mm; bill height varying from 2.82 to 3.52 mm, individuals; white breast with a greater or lesser presence compared to S. hypochondriaca with bill height ranges of brown spot-streaks (varying between 121 or 221); from 4.19 to 4.97 mm; and by wing length, which ranges white abdomen; sides and upper part white with more or

325 Stopiglia, R. et al.: Taxonomy of Synallaxis stictothorax species group

less presence of brown spot-streaks (varying between 121 tion, comes from the J. Orton (1830 – 1877)’s collection, or 221) and cinnamon underside (39); cinnamon fanks which is originally preserved at the Museum of Vassar (39) without spot-streaks; fve pairs of rufous rectrices College, now Warthin Museum of Geology & Natural (varying between 340 or 136), in some individuals the History, Vassar College, Poughkeepsie, USA. The cur- vexilla are partially brown (221), giving a two-colour ap- rent curator, after consulting the museum fles, informed pearance to the rectrices; rufous upper tail coverts (vary- us that Orton’s specimen of S. s. maculata was likely tak- ing between rufous 340 or 136); dark drab back (119B) en to have been destroyed on January 3, 1943, probably with greater or lesser presence of the rufous (340); dark due to its advanced stage of degradation (Lois Horst, in drab crown (varying between 119B or 119A); forehead litt.). We have also unfruitfully searched for this speci- white with brown spot-streaks (varying between 121 or men at the American Museum of Natural History, New 221); white superciliary line; rufous wing coverts (vary- York, USA, and at the New York State Museum, Albany, ing between 340 and 136); remiges brown (varying be- USA, whose ornithological collections historically re- tween 121 and 221). Bill length varying from 10.35 to ceived material from the Vassar College. 12.77 mm; bill height varying from 2.82 to 3.52 mm; wing length varying from 46.01 to 55.11 mm; and tail Remark 3. Synallaxis stictothorax piurae Chapman, length from 48.00 to 69.00 mm. No sexual dimorphism 1919 was considered a junior synonym of S. stictothorax was detected regarding plumage colour, but in the mor- maculata Lawrence, 1874 by Chapman (1925: 8) who phometric data, the wing length was smaller in females stated that “Dr. Hellmayr calls my attention to the fact that than in males (see Figs. 1 to 3, Tables 1 and 2). maculata Lawr. (1874, Ann. Lyc. Nat. Hist., N. Y., p. 186) antedates my piurae.”. Besides the fact that the author of Distribution. Synallaxis stictothorax occurs in southwest the species (Chapman, 1925) has considered S. s. piurae Ecuador (central Manabí, south to west Guayas, Isla Puna Chapman, 1919 a junior synonym of S. s. maculata Law- and south Loja) to northwestern Peru (Tumbes, Piura, rence, 1874 our analysis of type series and topotypes (in Lambayeque, La Libertad) as shows Fig. 4 and according the case of S. s. maculata, see Remark 2), corroborates to Remsen (2003: 289) and Dickinson & Christidis (2014: the absence of characters useful to distinguish between 139). According to (Lloyd, 2020) it lives in “dry scrub those taxa, as indicated at Table 3. The same can be said and open woodland below 200 meters in elevation.” and about S. s. maculata in relation to S. stictothorax. Law- according to del Hoyo (2020) in “arid lowland scrub, also rence (1874) in describing S. s. maculata compared his deciduous woodland edge; below 400 m.” new species with S. gujanensis (Gmelin, 1789), in spite of comparing it with S. stictothorax, probably, because he Remark 1. Sclater (1859), in the original description of was not aware of the description previously published by S. stictothorax, mentioned a specimen from the William Sclater (1859). This situation was resolved by Sclater Jardine (1800 – 1874)’s collection and another specimen (1890) who considered S. maculata a junior synonym de that was caught in Guayaquil, Ecuador, presented by S. stictothorax. Cory & Hellmayr (1925: 99) mentioned G. W. Barclay, and sent to the British Museum (= Natu- the importance of the variation on rufous extension in ral History Museum, Tring, England, NHMUK). Accord- remiges and rectrices in distinguishing S. s. maculata ing to Warren & Harrison (1971), the second one is the from S. stictothorax “Synallaxis stictothorax maculata specimen NHMUK 1841.4.2.471, obtained during the Lawrence: Similar to S. s. stictothorax, but back more voyage of the H.M.S. Sulphur, an English vessel utilized rufescent brown; cinnamon rufous area at base of remiges to explored the Pacifc coast of the Americas from 1836 more extensive; tail almost wholly rufous, only the medi- to 1837 (for details see Gray et al., 1844). Neverthe- an pair of rectrices being washed with dusky on terminal less, Warren & Harrison (1971) cited another syntype portion of inner web.” This variation is also reported by preserved at the Natural History Museum but they did Vaurie (1980: 123), but he considered that “Although the not provide any further detail on it. During our analy- population of stictothorax which inhabits northwestern sis of the NHMUK bird collection, it became clear that Peru is clearly differentiated from that of southwestern this syntype is NHMUK 1886.6.24.455, also a specimen Ecuador, their ranges are continuous and the differences from Guayaquil, Ecuador, from Jardine’s collection, with between the two populations are relative only,… In Synal­ the reference to William Jameson (1796 – 1873) and date laxis stictothorax I recognize nominate stictothorax rang- of July 1850. Both specimens are syntypes under the pro- ing from southwestern Ecuador to northwestern Peru, and visions of article 73.2 of the Code (Anonymous, 1999), chinchipensis in the valleys of Cajamarca.” Ridgely & and according to Article 74.6 (Fixation of lectotype by Tudor (2009) pointed out that the variation of the amount inference of holotype or “the type” before 2000) we be- of rufous in rectrices as the primary cause of the bicolored lieve that specimen NHMUK 1841.4.2.471 should be rectrices effect (see Figs. 1B e 2B) presented by the ma- considered the lectotype of Synallaxis stictothorax Sclat- jority of individuals of S. stictothorax from Ecuador and er, 1859, as Sclater (1890: 49) strictly indicated G. W. absent in individuals from southern Ecuador and Peru. Barclay’s specimen as the type of this species. Notwithstanding that, these authors made no taxonomic distinction based on such characters, considering it as Remark 2. The holotype of Synallaxis stictothorax ma­ a simple plumage variation. Our analysis indicated that culata Lawrence, 1874, according to its original descrip- such a variation in rectrices and remiges exists along the

326 VERTEBRATE ZOOLOGY — 70 (3) 2020

Table 3. Plumage colour of lectotype of S. stictothorax, topotype of S. s. maculata and holotype of S. s. piurae, according Smithe (1975, 1981) and topography according Proctor & Lynch (1993).

Lectotype of Topotype of Holotype of Character S. stictothorax S. s. maculata S. s. piurae (NHMUK 1841.4.2.471) (MVZ 163779) (AMNH 163085) Throat White White White Lateral throat White White White White ± macula or stretch marks White ± macula or stretch marks White ± macula or stretch marks Breast brown (121) brown (121) brown (121) Abdomen White White White Side Cinnamon (39) Cinnamon (39) Cinnamon (39) Flank Cinnamon (39) Cinnamon (39) Cinnamon (39) Rectrices Rufous (± Brown) Rufous (± Brown) Rufous (± Brown) Upper tail coverts Rufous (340) Rufous (340) Rufous (340) Back Dark drab (119B) Dark drab (119B) Dark drab (119B) Crown Brown (119A) Brown (119A) Brown (119A) White ± stretch marks brown White ± stretch marks brown White ± stretch marks brown Forehead (221) (221) (221) Superciliary line White White White Wing coverts Rufous (340) Rufous (340) Rufous (340) Remiges Brown (221) Brown (221) Brown (221) distribution of S. stictothorax, but it is also present in in- (121 or 221) in S. chinchipensis. Synallaxis hypochon­ dividuals of S. chinchipensis, with no indication of use- driaca is still diagnosable relative to S. stictothorax by: fulness as diagnostic character between taxa (see Fig. 1B bill length, which varies from 15.01 to 19.66 mm versus and Table 1). 10.35 to 12.77 mm in S. stictothorax; bill height, varying between 4.19 and 4.97 mm versus 2.82 to 3.52 mm in S. stictothorax; and wing length, varying between 60.47 Synallaxis hypochondriaca (Salvin, 1895) and 68.57 mm, while S. stictothorax presents variation Siptornis hypochondriacus Salvin, 1895: 14. Syntypes, by origi- from 46.01 to 55.11 mm. Synallaxis hypochondriaca is nal designation: NHMUK 1899.6.1.573, adult male; NHMUK still diagnosable relative to S. chinchipensis by: a white 1899.6.1.574, adult female; NHMUK 1899.6.1.579 adult fe- superciliary line versus the cinnamon (39) one of S. chin­ male; AMNH 523869, adult female; from “Malea”, Cajabam- chipensis; bill height that varies from 4.19 to 4.97, while ba, Peru (8,000 ft.), collected by O. T. Baron from 16 to 18 in S. chinchipensis it ranges from 3.16 and 3.81 mm; April 1894; and AMNH 523868 from Cajabamba, Peru (9,000 ft.) collected by O. T. Baron on April 2 1894. Lectotype, by and by wing length varying from 60.47 to 68.57 mm in present designation: AMNH 523869 (see Remark 4). Stephens S. hypochondriaca and between 47.89 and 53.65 mm in & Traylor (1983) suggest that “Malea” is a variation of the S. chinchipensis. Synallaxis hypochondriaca is diagnos- locality Malca, Department of Cajamarca, Peru. Therefore, as able relative to all other Synallaxis by the white breast previously mentioned by Cory & Hellmayr (1925), the correct with brown spot-streaks (121 or 221; see Figs 1 and 2, reference is Malca and Cajabamba, Department of Cajamar- Tables 1 and 2). ca, Peru. The coordinates are 07°35′S, 78°09′W (Stephens & Traylor, 1983), with an altitude of nearly 2,500 – 3,000 m (8,000 – 9,000 ft.), according to Salvin (1895). Description. Throat and lateral throat white; white breast with greater or lesser presence of brown spot-streaks Diagnosis. Synallaxis hypochondriaca is diagnosable (varying between colours 121 and 221); white abdomen; relative to S. stictothorax and S. chinchipensis by the sides of the body white and umber with greater or lesser colours white and umber (123) with brown spot-streaks presence of brown spot-streaks (varying between 121 (121 or 221) at the lower part of the sides of the body and and 221); fanks white and umber (123) without spot- fanks, while S. stictothorax presents the lower part of the streaks; six pairs of dark rufous rectrices (119B); in some sides of the body and fanks in cinnamon (39), and in individuals the vexilla are partially brown (221), giving a S. chinchipensis the sides and fanks are white with brown two-colour appearance for the rectrices; dark drab (119B) spot-streaks (121 or 221). Synallaxis hypochondriaca is upper tail coverts; dark drab back (119B) with greater also distinguishable by its dark drab (119B) rectrices and or lesser presence of the rufous (340); dark drab crown upper tail coverts, versus the rufous rectrices (340 or (varying between 119B and 119A); dark drab forehead 136) of S. stictothorax and S. chinchipensis, and by its (119B); white superciliary line; rufous (varying between dark drab forehead (119B), which is white with stretch 340 and 136) wing coverts; brown remiges (varying be- marks brown (121 or 221) in S. stictothorax and cinna- tween 121 and 221). Bill length varying from 15.01 to mon (39) and dark drab (119B) with stretch marks brown 19.66 mm; bill height from 4.19 to 4.97 mm; wing length

327 Stopiglia, R. et al.: Taxonomy of Synallaxis stictothorax species group

from 60.47 to 68.57 mm; and tail length varying between Remark 5. There is one specimen mentioned above, that 69.00 and 94.00 mm. No sexual dimorphism was de- deserves a special mention: NHMUK 1899.6.1.578, tected regarding plumage colour, but in the morphomet- from Chusgon, Peru. According to Stephens & Traylor ric data, the bill height, bill length, and wing length are (1983), this specimen was collected at 2,593 m, 12 km smaller in females than in males (see Figs. 1 to 3, Tables from Sarin on Río Chusgon, left bank affuent of the Río 1 and 2). See Remark 5 for additional information. Marañón, by O. T. Baron on 13 February 1895. It pre- sents a plumage more similar to S. chinchipensis than to Distribution. Northcentral Peru, upper Río Marañón val- S. hypochondriaca, including the dark drab (119B) fore- ley in north Ancash, south Cajamarca, south-west Ama- head with cinnamon (39) and brown spot-streaks, as well zonas and La Libertad, between 2,000 and 2,800 meters as a cinnamon (39) superciliary line. Nevertheless, it is elevation, restricted to the Marañón Valley as shows Fig. not totally morphologically congruent with that species 4, according to Remsen (2003: 290) and Dickinson & considering the presence of cinnamon (39) in the breast, Christidis (2014: 138). while in S. chinchipensis it is white. The morphometric data are similar to those of S. hypochondriaca (bill length Remark 4. The original description of Synallaxis hy­ 17.17 mm, bill height 4.23 mm, wing length 63.58 mm, pochondriaca (Salvin, 1895) mentions male and female tail length 87.00 mm, and 12 rectrices; see Table 2). This without details. During our analysis of the NHMUK and specimen is under investigation. For the purposes of this AMNH collections, we encountered fve supposed syn- study, it will be identifed as a morphological variant types, according to the information on their labels. The of S. hypochondriaca, but in future analysis it may be catalogue of types from NHMUK (Warren & Harrison, shown to be more than a simple variation of the plumage. 1971) indicated two syntypes: NHMUK 1899.6.1.573, adult male; and NHMUK 1899.6.1.574, adult female. The catalogue of types of the AMNH (LeCroy & Sloss Synallaxis chinchipensis Chapman, 1925 2000) do not point out any type for S. hypochondriaca Synallaxis stictothorax chinchipensis Chapman, 1925: 8. Holotype, (Salvin, 1895), however LeCroy (2017, in litt.) informed by original designation: AMNH 182062, adult male from Per- us that the determination of those syntypes at AMNH ico, Río Chinchipe, near the Marañón, Cajamarca, Peru, coor- preceded the publication of the referred catalogue. Ac- dinates are 05°15′S, 78°45′W with an altitude of nearly 200 m cording to LeCroy (2017, in litt) the AMNH specimens (Stephens & Traylor, 1983; LeCroy & Sloss, 2000). should be regarded as syntypes, as well as those speci- mens housed at NHMUK, given the fact that Salvin Diagnosis. Synallaxis chinchipensis is distinguishable (1895) wrote in his introductory pages that part of the from S. stictothorax and S. hypochondriaca by the lower material which has been collected by O. T. Baron in part of the sides of the body and fanks white with brown Peru had been sent to him (NHMUK specimens) and the (121 or 221) spot-streaks, while the lower part of the other part (those indicated by asterisk in Salvin, 1895) sides of the body and fanks are white and umber (123) to the Rothschild Museum (now the AMNH specimens). with brown spot-streaks in S. hypochondriaca, and the In addition to these four specimens recognized by their lower part of the sides of the body and fanks are cinna- respective safeguard institutions as syntypes, during our mon (39) in S. stictothorax. Synallaxis chinchipensis also studies at NHMUK we found at least one more specimen, differs from S. stictothorax and S. hypochondriaca by its NHMUK 1899.6.1.579, that should also be considered white and cinnamon (39) fanks with brown spot-streaks. one of the syntypes following article 73.2.1. of the Code. The two other species present fanks that are cinnamon This specimen is an adult female and, as the other two (39), and white and umber, respectively. Synallaxis chin­ that are recognized as syntypes by Warren & Harrison chipensis is also distinguishable by its dark drab (119B) (1971), was collected by O. T. Baron in Malea, Cajabam- and cinnamon (39) forehead with brown spot-streaks ba, Peru on April 1894. There are also other four speci- (121 or 221), versus the white with brown spot-streaks mens of S. hypochondriaca that were collected by O. T. (121 or 221) of S. stictothorax, and the dark drab (119B) Baron, however, these specimens are from different lo- forehead of S. hypochondriaca. Synallaxis chinchipen­ calities of Peru (NHMUK 1899.6.1.575, Malea, Chusgon; sis presents a cinnamon (39) superciliary line, while in NHMUK 1899.6.1.576, Sueca, Huamachuro; NHMUK S. stictothorax and S. hypochondriaca it is white. Synal­ 1899.6.1.577, Cajabamba; NHMUK 1899.6.1.578, Chus­ laxis chinchipensis is still diagnosable relative to S. hy­ gon), collected on February 1895, instead of April 1894, pochondriaca by its rufous rectrices and upper tail cov- and since the original description was published on Febru- erts, which are dark drab (119B) in S. hypochondriaca, ary 1895, we did not consider them types. In this respect, and by its measurements: bill height varying from 3.16 considering the Remark 4 and the recommendations of to 3.81 mm, while in S. hypochondriaca it ranges from the Code, here we designate the specimen AMNH 523869 4.19 to 4.97 mm; and wing length varying from 47.89 as the lectotype of the nominal species S. hypochondri­ to 53.65 mm, while in S. hypochondriaca it varies from aca (Salvin, 1895) and state that the remaining speci- 60.47 to 68.57 mm. Synallaxis chinchipensis is diagnos- mens (NHMUK 1899.6.1.573, NHMUK 1899.6.1.574, able relative to all other Synallaxis by its white breast NHMUK 1899.6.1.579, AMNH 523868) be considered with brown spot-streaks (121 or 221; see Figs. 1 and 2, from now on as its paralectotypes. Tables 1 and 2).

328 VERTEBRATE ZOOLOGY — 70 (3) 2020

Description. White throat; white lateral part of the S. hypochondriaca and S. chinchipensis. Such a varia­ throat, with cinnamon spot-streak (39) on some indi- tion has, historically, caused much confusion, and result­ vidual; white breast with greater or lesser presence of ed in the descriptions of the subspecies S. s. maculata brown spot-streaks (colour varying between 121 and Lawrence, 1874 and S. s. piurae Chapman, 1919, which 221); white abdomen; side of the body white, with were based on simple variations that correspond to usual greater or lesser presence of brown spot-streaks (varying fuctuations in plumage colour patterns, which occur in from 121 to 221); fanks white with cinnamon (39) inva- virtually all taxa, and do not correspond to diagnostic sion and brown spot-streaks (varying from 121 to 221); characters. These are particularly noticeable with respect fve pairs of rufous rectrices (varying from 340 to 136), to the lateral throat, with some individuals of S. sticto­ and in some individuals the vexilla are partially brown thorax and S. chinchipensis presenting the cinnamon (221), giving a two-colour appearance for those feath- spot-streak, and with respect to the breast, sides of the ers; rufous (varying from 340 to 136) upper tail coverts; body, and fanks, in which the shape of the spot-streaks is dark drab back (119B) with greater or lesser invasion of sometimes more elongated and sometimes more rounded rufous (340); dark drab crown (varying from 119B to (like macules), as well as variation in the number of spot- 119A); cinnamon (39) and dark drab (119B) forehead streaks. The spot-streaks of S. stictothorax and S. chin­ with few brown spot-streaks; cinnamon superciliary line chipensis vary in amount, location, and tone of brown (39); rufous wing coverts (colours varying between 340 (121 and 221), as well as in relation to the presence of and 136); brown remiges (varying from 121 to 221). rufous tones (340 or 136). The back of S. stictothorax, Bill length varying from 12.15 to 15.84 mm; bill height S. chinchipensis, and S. hypochondriaca also vary in from 3.16 to 3.81 mm; wing length varying from 47.89 regards to the presence of rufous, and the crown varies to 53.65 mm; and tail length from 60.00 to 72.00 mm. No depending on the presence of brown (119A). The fore- sexual dimorphism was detected regarding plumage col- head is also variable in S. stictothorax and S. chinchipen­ our, but morphometric data shows consistent differences sis, showing a greater or lesser presence of brown spot- among sexes, with smaller females overall (see Figs. 1 to streaks and varying in tone (121 or 221). Wing coverts 3, Tables 1 and 2). vary in the kind of rufous (340 or 136), and remiges in the tone of brown (121 or 221). Distributiom. Northwest Peru, River Chinchipe and mid- Given their patterns of distribution and degree of dle Río Marañón valley in Cajamarca as shows Fig. 4, ac- overlap, most of these variations were interpreted by us cording to Remsen (2003: 289) and Dickinson & Chris- as polymorphisms, without relation to geography, sexual tidis (2014: 139). It lives in “Arid lowland scrub, also dimorphism, or ontogeny, as variation was observed in deciduous woodland edge; mainly at 400 – 600 m.” (del specimens from the same locality (e.g. AMNH 129790 Hoyo et al., 2020). and AMNH 129791 from Guayaquil, Ecuador), in both male and female adults. Finally, we call attention to the importance of con- tinuing the revision of polytypic taxa in order to identify Discussion valid species, which are sometimes hidden at the subspe- cies level. Indeed, some subspecies that are commonly recognised as valid are clearly worthy of specifc status; As mentioned previously, Derryberry et al. (2011) were however, others may not even correspond to geographi- the frst to contribute to the elucidation of the intra- and cal variation. This was the case for S. s. maculata, which interspecifc relationships in the S. stictothorax species was not supported as a valid taxon by this study. Howev- group from a phylogenetic perspective. In that study, er, S. chinchipensis, which was previously recognized as S. hypochondriaca was included in the genus Synallaxis, a subspecies of S. stictothorax, was diagnosed morpho- as the sister group of the clade comprising S. stictothorax logically and classifed as a close relative of S. hypochon­ and S. zimmeri. After Derryberry et al. (2011), it became driaca, according to a molecular analysis (Tobias et al., evident that the monotypic genus Siptornopsis should be 2014). The precise distinction between subspecifc taxa considered synonymous with Synallaxis, as adopted by representing natural units and those that are the result of Ohlson et al. (2013). Tobias et al. (2014), repeated the historical sample artefacts is of ultimate importance even analysis of Derryberry et al. (2011) adding S. chinchi­ today, as those entities will serve as the basis for all our pensis and S. s. maculata to the dataset. The resulting to- efforts in conservation and for studies on biogeography, pology included S. stictothorax and S. s. maculata as the comparative anatomy, and others (Mayr, 1982; Crac- sister clade of S. zimmeri and also indicated that S. chin­ raft, 1983; Wheeler et al., 2004; Peterson, 2006; Ag- chipensis was more closely related to S. hypochondriaca narsson & Kuntner, 2007; de Queiroz, 2007; Padial et than to S. stictothorax. This relationship is better under- al., 2010). The more careful and scientifc the taxonomic stood and represented now with the elevation of S. chin­ reviews are, the more reliable will be any inference made chipensis to species status, as indicated by our analysis of from their conclusions. plumage colour and morphometry. It should also be noted that a high degree of morpho- logical variation was observed within the S. stictothorax,

329 Stopiglia, R. et al.: Taxonomy of Synallaxis stictothorax species group

Acknowledgements Brumfield, R. T. (2011). Lineage diversifcation and morpho- logical evolution in a large-scale continental radiation: the Neotropical ovenbirds and woodcreepers (Aves: Furnariidae). Evolution, 65, 2973 – 2986. We would like to acknowledge the following supports provid- Dickinson, E. C. & Christidis, L. (2014) The Howard and Moore ed to RS: FUNCAP Proc. # 02853404/2019, FAPESP Proc. # Complete Checklist of the Birds of the World. Eastbourne, Aves 2013/26609-1 and 2016/18963-8; CAPES; the Ernst Mayr Travel Press Limited , 752 pp. Grants in Systematics, Museum of Comparative Zoology, Graves, G.R. (1985). Elevational correlates of speciation and in- Harvard University, Cambridge, USA; the Collection Study Grant, traspecifc geographic variation in plumage in Andean forest birds. The Auk: Ornithological Advances, 102, 556 – 579. American Museum of Natural History, New York, USA; and the Gray, J. E., Gould, J. & Richardson, J. (1844). Mammalia, Birds Visiting Scholarship Grant, Field Museum of Natural History, and Fish. in: Hinds, R. B. (ed.) The Zoology of the Voyage of Chicago, USA. We also would like to acknowledge the CNPq and H.M.S. Sulphur, under the Command of Captain Sir Edward FAPERJ by the support given to the project “Catálogo de tipos de Belcher, R.N., C.B., F.R.G.S., etc. Vol. I. London, Smith, Elder espécies brasileiras” (Proc. # 479049/2006-8) and the fellowship & Co. Lawrence, G. N. (1874). Descriptions of New Species of Birds of (Proc. # 310384) provided to MAR. FAB is supported by CNPq the Genera Icterus and Synallaxis. Annals Lyceum of Natural scholarship (Proc. # 312687/2018-4). History of New York, 10, 184 – 187. LeCroy, M., & Sloss. R. (2000). Type specimens of birds in the American Museum of Natural History. Part 3: Passeriformes: Eurylaimidae, Dendrocolaptidae, Furnariidae, Formicariidae, References Conopophagidae and Rhinocryptidae. Bulletin of the American Museum of Natural History, 257, 1 – 88 pp. Lloyd, H. (2020). (Synallaxis hypochondriaca), Addinsoft (2020). xlstat statistical and data analysis solution. version 1.0. in: Billerman, S. M., Keeney, B. K., Rodewald, P. New York, USA. https://www.xlstat.com. G. & Schulenberg, T. S. (eds) Birds of the World. Ithaca, NY, Agnarsson, I. & Kuntner, M. (2007). Taxonomy in a changing USA, Cornell Lab of Ornithology. world: seeking solutions for a science in crisis. Systematic Bio­ Mayr, E. (1982). The Growth of Biological Thought. Cambridge, logy, 56, 531 – 539. MA, Harvard University Press, 974 pp. Anonymous [International Commission on Zoological Nomencla- Ohlson, J. I., Irestedt, M., Ericson, P. G. P. & Fjeldså, J. (2013). ture] (1999). International Code of Zoological Nomenclature. Phylogeny and classifcation of the New World suboscines Fourth edition. London, International Trust for zoological No- (Aves, Passeriformes). Zootaxa, 3613, 1 – 35. menclature, 306 pp. Padial, J. M., Miralles, A., De La Riva, I. & Vences, M. (2010). Bergmann C. (1847). Ueber die Verhältnisse der Wärmeökonomie The integrative future of taxonomy. Frontiers in Zoology, 7, der Thiere zu ihrer Grösse. Göttinger Studien, 3, 595 – 708. 1 – 16. BirdLife International (2020) Species factsheet: Synallaxis hy­ Paynter, R. A. (1993). Ornithological Gazetteer of Ecuador, Sec­ pochondriaca. Downloaded from http://www.birdlife.org on ond Edition. Cambridge, Harvard University, 247 pp. 11/05/2020. Recommended citation for factsheets for more Peters, J.L. (1951). Check-list of birds of the world. Vol. VII. Cam- than one species: BirdLife International (2020) IUCN Red bridge, MA, Museum of Comparative Zoology. List for birds. Downloaded from http://www.birdlife.org on Peterson, A. T. (2006). Uses and requirements of ecological niche 11/05/2020. models and related distributional models. Biodiversity Infor­ Chapman, F. M. (1925). Descriptions of one new genus and of spe- matics, 3, 59 – 72. cies of birds from Peru and Ecuador. American Museum Novi­ Proctor, N. S. & Lynch, P. J. (1993). Manual of Ornithology: tates, 205, 1 – 11. Avian Structure and Function. New Haven and London, Yale Cory, C. B. & Hellmayr, C. E. (1925). Catalogue of birds of the University Press, 340 pp. Americas and the adjacent islands in Field Museum of Natural Remsen, J. V. (2003). Family Furnariidae (ovenbirds), pp. 162 – 357 History including all species and subspecies known to occur in in: del Hoyo, J., Elliott, A., Sargatal, J., Christie, D. A. & North America, Mexico, Central America, South America, the de Juana, E. (eds) Handbook of the Birds of the World, Vol. 8. West Indies, and islands of the Carribean Sea, the Galapagos Broadbills to Tapaculos. Barcelona, Lynx Edicions. Archipelago, and other islands which may be included on ac- Ridgely, R. S. & Greenfield, P. J. (2001). The birds of Ecuador. count of their faunal affnities. Part IV, Furnariidae – Dendro- Status, Distribution, and Taxonomy, vol. I. Ithaca, New York, colaptidae. Publications of the Field Museum of Natural His­ Cornell University Press. tory (Zoological Series), 13, 1 – 390. Ridgely, R. S. & Tudor, G. (1994). The Birds of South America, Cracraft, J. (1983). Species concept and speciation analysis, pp. vol. 2. Austin, University of Texas Press. 159 – 187 in: Johnston, R. F. (ed.) Current Ornithology. v. 1. Salvin, O. (1895). On birds collected in Peru by Mr. O. T. Baron. New York, Plenum Press. Novitates Zoologicae 2(1), 1 – 22. de Queiroz, K. (2007). Species concepts e species delimitation. Sclater, P. L. (1859). On some new species of Synallaxis, and on Systematic Biology, 56, 879 – 886. the geographical distribution of the genus. Proceedings of the del Hoyo, J., Collar, N. J., Christie, D. A., Elliott, A., Fishpool, Zoological Society of London, 27, 191 – 197. L. D. C., Boesman, P. & Kirwan, G. M. (2016). HBW and Sclater, P. L. (1874). On the species of the genus Synallaxis of the BirdLife International Illustrated Checklist of the Birds of the family Dendrocolaptidae. Proceedings of the Scientifc Meet­ World. Volume 2: Passerines. Barcelona, Spain and Cambridge, ing of the Zoological Society of London, 2 – 28 pp. United Kingdon, Lynx Edicions and BirdLife International. Sclater, P. L. (1890). Catalogue of the Passeriformes in the Col­ del Hoyo, J., Remsen, J. V., Kirwan, G. M. & Collar, N. (2020). lection of the British Museum. Volume 15. London, Taylor and (Synallaxis stictothorax), version 1.0. in: Francis, 371 pp. Billerman, S. M., Keeney, B. K., Rodewald, P. G. & Schulen- Sibley, C. G. & Monroe, B. L., (1990). Distribution and Taxonomy berg, T. S. (eds) Birds of the World. Ithaca, NY, USA, Cornell of Birds of the World. New Haven, Connecticut, Yale Univer- Lab of Ornithology. https://doi.org/10.2173/bow.necspi1.01 sity Press. Derryberry, E. P., Claramunt, S., Chesser, R. T., Remsen, J. V., Smithe, F. B. (1975). Naturalist’s Color Guide. New York, Ameri- Cracraft, J., Aleixo, A., Pérez-Emán, J., Remsen, J. V., & can Museum of Natural History, 119 pp.

330 VERTEBRATE ZOOLOGY — 70 (3) 2020

Smithe, F. B. (1981). Naturalist’s Color Guide. New York, Ameri- Warren, R. M., & Harrison, C. J. O. (1971). Type-specimens of can Museum of Natural History, 37 pp. Birds in the British Museum (Natural History), vol. 2. Passer­ Stephens L. & Traylor, M. A. (1983). Ornithological Gazetteer of ines. London, Trustees of the British Museum (Natural His- Peru. Cambridge, Harvard University, 273 pp. tory), 628 pp. Tobias, J. A., Cornwallis, C. K., Derryberry, E. P., Claramunt, S., Wheeler, Q. D.; Raven, P.; & Wilson, E. O. (2004). Taxonomy: Brumfield, R. T. & Seddon, N. (2014). Species coexistence impediment or expedient? Science, 303, 285. and the dynamics of phenotypic evolution in adaptive radia- Winger, B. M. & Bates, J. M. (2015). The tempo of trait diver- tion. Nature, 506, 359 – 363. gence in geographic isolation: Avian speciation across the Vaurie, C. (1971). Classifcation of the Ovenbirds (Furnariidae). Marañón Valley of Peru. Evolution, 69, 772 – 787. London, H. F. Witherby. Vaurie, C. (1980). Taxonomy and geographical distribution of the Furnariidae (Aves, Passeriformes). Bulletin of the American Museum of Natural History, 166, 1 – 357.

Zoobank Registration at http://zoobank.org

This published work and the nomenclatural acts it contains have been registered in ZooBank, the online registration system for the Interna- tional Commission on Zoological Nomenclature (ICZN). The Zoo- Bank LSIDs (Life Science Identifers) can be resolved and the associ- ated information can be viewed through any standard web browser by appending the LSID to the prefx http://zoobank.org. The LSID for this publication is as follows: urn:lsid:zoobank.org:pub: act:CDBF4996-1B5C-4EE8-BC55- 447F65BF9FC3

331