identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
03A087D88374FFC9B2EA4F0E20E4F81E.text	03A087D88374FFC9B2EA4F0E20E4F81E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus hellmayri Berlepsch 1902	<div><p>Catharus hellmayri Berlepsch 1902</p><p>Talamanca nightingale-thrush</p><p>(Fig. 13)</p><p>Catharus fuscater hellmayri Berlepsch 1902: 69; Ridgway 1907: 24; Carriker 1910: 748; Nelson 1912: 24; Hellmayr 1934: 464; Slud 1964: 301; Wetmore et al. 1984: 155; Ridgley and Gwynne 1989: 352; Phillips 1991: 112; Clement 2000: 299; Collar 2005: 700; Halley 2020; Halley 2021.</p><p>Catharus fuscater Salvin 1867: 132; Lawrence 1868: 9; Frantzius 1869: 289; Salvin 1870: 180; Salvin and Godman 1879: 5 (in part); Ridgway 1881: 333; Zeledon 1885: 104; Bangs 1902: 50; Meyer de Schauensee 1964: 317 (in part); Powell 1979; Stiles and Skutch 1989: 368; Loiselle and Blake 1991; Young et al. 1993; Wolfe et al. 2009; Bueter et al. 2009; Jankowski et al. 2010; Kounek et al. 2013.</p><p>Catharus hellmayri Sharpe 1902: 182; Davis 1972: 176.</p><p>Type material</p><p>Monotypic species. SMF 53156 (holotype), study skin, adult male, collected by E. Gounelle in ‘ Panama, Veragua, Chiriquí’. This specimen was not examined in this study. Label data were provided by G. Mayr (in litt.) .</p><p>Geographic range</p><p>Northern mountains of Costa Rica (Rincón de la Vieja, Miravalles, Tenorio) to west-central Panama (Parque Nacional Santa Fé, Veraguas).</p><p>Adult specimens examined</p><p>Catharus hellmayri (N = 64): Costa Rica (N = 34): Cartago (five males, two females): ‘ Aqinares’ (= Aquiares): AMNH 391543 (male); Cervantes: USNM 49152 (female) ; Juan Vinas: CM P28231 (male) ; Navarro: AMNH 391541 (male) , AMNH 391542 (female); Santa Cruz de Turrialba: AMNH 391544 (male) ; Tierra Blanca: CM P11045 (male) ; Heredia (seven males, one female): Cariblanco de Sarapiquí: ANSP 67249, CM P25349, P25362, P25368, USNM 199697 (males), AMNH 503920 (female); unknown site about 24 km. N. of San José: AMNH 391539, AMNH 391540 (males); San José (13 males, six females): Carrillo: AMNH 503916–503918 (males); Guayabo: AMNH 391536 (male) ; La Guaria de Dota: USNM 209929 (male) , USNM 209928 (female); La Hondura: AMNH 391537 , CM P26384, P26393, P26414, P26423, P26434, P26445, P26535 (males), AMNH 391538, CM P26108, P26550, P26581 (females); unknown locality (one female): USNM 68172 . Panama (N = 30): Chiriquí (11 males, seven females): Boquete: AMNH 503911–503913, FMNH 24305 207592 207595, USNM 188572 188573 (males), FMNH 53580, USNM 188570, USNM 188571, AMNH 503814 (females); 12.6 km north of Los Planes, Gualara-Chiriquí Grande Road: USNM 607605 (male) , USNM 607614 (female); unknown site 23.3 km north of Los Planes, Gualara-Chiriquí Grande Road: USNM 607617 (male) ; Reserva Forestal Fortuna: FMNH 470768 (male) , FMNH 470767 (female); Veraguas (seven males, six females): AMNH 246360–246366 (males), AMNH 187643, AMNH 246367–246371 (females).</p><p>Immature specimens examined</p><p>Catharus hellmayri (N = 8): Costa Rica (N = 6): Heredia (three females): Cariblanco de Sarapiquí: AMNH 503919, CM P25344, P25363 (females) ; San José (two males): Guayabo: AMNH 391534, 391535 (males); Unknown Province: Cusoua, La Palma: USNM 85546 (male) . Panama (N = 2): Chiriquí (two females): Boquete: FMNH 207593 207594 (females) .</p><p>Audio recordings examined</p><p>Catharus hellmayri (N = 63): Costa Rica (N = 51): Alajuela: Bosque de Paz Biological Reserve: XC 107128; Monteverde: XC 220593; Parque Nacional Juan Castro Blanco, sector Volcán Viejo: ML 64132091; Volcán Tenorio: XC 166201, 166202 ; Cartago: Orosi: XC 169247; Parque Nacional Tapantí: ML 48406801, XC 274444, 274445, 274447, 274448; Heredia: Braulio Carrillo, Carretera: ML 28234 ; Puntarenas: Monteverde: ML 28202, 28263, 28264, 28269, 32268, 32300, 32309, 32327, 32328, 32330, 32701, 32715, 53950, 53951, 56198, 72831, 72839, 72844, 72850, 72854, 74195, 76697, 76699, 76703 209231 209234 209242 209244, 219846, 220560, 220582, 220611, 220633, XC 45226, 137776, 593699; San José: Monserrat de Coronado: ML 59585321; San Gerardo de Dota: XC 107125, 107127 . Panama (N = 12): Chiriquí: Boquete: XC 133595; Continental Divide, along Gualaca-Chiriquí Rd.: ML 54410; Kakintú: XC 112998; Los Planes, Fortuna Field Station: ML 54207, 54225, 54391, 54427; Reserva Forestal de Fortuna: XC 271146, 271147, 367649 ; Ngäbe-Buglé: Cerro Colorado: XC 78388, 145618.</p><p>Diagnosis</p><p>Genetics: In both phylogenetic reconstructions, samples from the range of C. hellmayri formed a clade that was sister to the rest of the ingroup. ABGD and ASAP analyses both identified C. hellmayri as an independent genetic cluster. The estimated divergence time between C. hellmayri and the rest of the complex was 3.2 Mya (range = 2.6–3.8) and it was highly divergent from its closest geographic neighbours, C. [f.] mirabilis (mean uncorrected p -distance = 0.075) and Undescribed 1 (mean uncorrected p -distance = 0.076).</p><p>Morphology: Study skins of C. hellmayri have darker ventral plumage than any other taxon, except C. [f.] sanctaemartae (Fig. 13); less black on the chin than C. [f.] sanctaemartae; and shorter tails on average than any other taxon (see: Hellmayr 1934).</p><p>Voice: Catharus hellmayri is distinguished from the South American taxa by its Type 1 punctuation call structure (Fig. 9), ‘tightly spaced’ contact call structure (Fig. 10), and greater diversity of song contours (Fig. 12). It was the only taxon with a narrow-bandwidth blurred call that rose in pitch (Fig. 11). The song contour DACB accounted for 15% of songs in C. hellmayri (N = 27/179), but was not detected in any other clade.</p><p>Etymology</p><p>The proposed English name references the Cordillera de Talamanca, which encompasses nearly the entire range of C. hellmayri .</p></div>	https://treatment.plazi.org/id/03A087D88374FFC9B2EA4F0E20E4F81E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
03A087D8836BFFCBB2AE4FA024A5FD1B.text	03A087D8836BFFCBB2AE4FA024A5FD1B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus arcanus Halley & Catanach & Klicka & Weckstein 2023	<div><p>Catharus arcanus sp.nov.</p><p>Darién nightingale-thrush</p><p>(Figs 13, 14; formerly Undescribed 1)</p><p>urn:lsid:zoobank.org:act: 25B3DB71-FE5E-4FBF-A2CE-8D270CBA7250.</p><p>Catharus fuscater Meyer de Schauensee 1964: 317 (in part).</p><p>Catharus fuscater fuscater Wetmore et al. 1984: 157; Ridgley and Gwynne 1989: 352; Clement 2000: 299 (in part); Collar 2005: 700 (in part); Renjifo et al. 2017.</p><p>Catharus fuscater hellmayri Clement 2000: 299 (in part).</p><p>Catharus fuscater mirabilis Clement 2000: 299 (in part).</p><p>Catharus fuscater subsp.? Phillips 1991: 112.</p><p>‘Unnamed-1’ Halley 2021: 132.</p><p>Type material</p><p>Monotypic species. FMNH 470769 (holotype), study skin, adult male, collected by J. Klicka (preparator), G.M. Spellman, and J.M. DaCosta on Cerro Chucantí, 17 km south-west of Chimán, Panamá province, Panama (8.7958°, –78.4630°, elev. 1200 m), on 16 February 2006 .</p><p>Geographic range</p><p>Serranía de Majé, Panama, and along the Serranía del Darién from Cerro Azul in the west, to Cerro Tacarcuna in the east (Wetmore et al. 1984, Cuervo 2013, Renjifo et al. 2017).</p><p>Adult specimens examined</p><p>Catharus arcanus sp. nov. (N = 25): Panama: Panamá (one male, one female): Cerro Chucantí: FMNH 470769 (male); NW slope of Cerro Jefe: LSUMZ 164300 (female) ; Darién (14 males, nine females): unspecified locality on Cerro Tacarcuna, eastern slope: AMNH 136159–136165 (males), AMNH 136166 (female); unspecified locality on Cerro Tacarcuna, slope unknown: AMNH 136171, 811713, LSUMZ 80201 (males) , AMNH 136167– 136170, AMNH 811714 (females); La Laguna Camp on Cerro Tacarcuna: USNM 469735 (male) , USNM 486474 (female); unspecified locality on Cerro Malí: USNM 484791, USNM 484792 (males), USNM 484793 (female); unspecified locality on ridge west of Cerro Malí: USNM 486473 (male).</p><p>246 • Halley et al.</p><p>Immature specimens examined</p><p>Catharus arcanus sp. nov. (N = 1): Panama: Darién (one male): La Laguna Camp, Cerro Tacarcuna: USNM 486475 (male) .</p><p>Audio recordings examined</p><p>Catharus arcanus sp. nov. (N = 6): Panama: Darién: Cerro Chucantí: ML 137736401, XC 2976–2979, 164780 .</p><p>Diagnosis</p><p>Genetics: In both phylogenetic reconstructions, samples of C. arcanus sp. nov. formed a clade that was sister to C. [f.] mirabilis of Cerro Pirre (mean uncorrected p -distance = 0.04 ± &lt;0.01). In the UCE tree, these taxa were reciprocally monophyletic and formed a clade that was sister to all South American taxa. ABGD and ASAP analyses both identified C. arcanus sp. nov. and C. [f.] mirabilis as independent genetic clusters. The estimated divergence time between C. arcanus sp. nov. and C. [f.] mirabilis was 1.3 Mya (95% HPD = 0.9–1.7).</p><p>Morphology: Adult study skins of C. arcanus sp. nov. are paler on the dorsal surface than similarly-aged skins of C. hellmayri and C. [f.] mirabilis, in both sexes (Fig. 13). Unlike adult females of C. [f.] mirabilis, which had darker crowns than mantles, there was no contrast between the pale crown and mantle of the C. arcanus sp. nov. adult female (LSUMZ 164300), which had an enlarged ovary (11 × 5 mm), 100% pneumatized skull, and no bursa. Ventrally, study skins of C. arcanus sp. nov. and C. [f.] mirabilis were paler than C. hellmayri (Fig. 13). Like C. [f.] mirabilis (and to a lesser extent C. hellmayri, see: Ridgley and Gwynne 1989), C. arcanus sp. nov. has a yellowish wash on the ventral surface in life, that renders the breast ‘olive’ and abdomen ‘pale yellow’ (fide E. Eisenmann, on the label of AMNH 811714). Similar notes were written on the labels of LSUMZ 80201 (‘Yellowish olive instead of grey underneath in fresh specimens’) and AMNH 811713 (‘olive yellowish instead of grey underneath’). This character distinguishes these three taxa from all South American populations, but the yellow wash fades quickly after death. After a few years, museum skins of C. arcanus sp. nov. have no trace of yellow in the ventral plumage and the ‘olive’ fades to cooler than Light Drab (119C). To our knowledge, no fresh (&lt;5 years old) material of C. arcanus sp. nov. or C. [f.] mirabilis exists in any collection.</p><p>Voice: Catharus arcanus sp. nov. is distinguished from the South American taxa by its Type 1 punctuation call structure (Fig. 9), and from C. [f.] mirabilis by its ‘tightly spaced’ contact call structure (Fig. 10). Our dataset lacked recordings of blurred calls from C. arcanus sp. nov., and too few recordings were available to adequately determine whether C. arcanus sp. nov. and C. [f.] mirabilis are divergent in song. Notwithstanding, the only triadic song contour (BCA) detected in C. arcanus sp. nov., which accounted for 43% of songs (N = 3/7), was not detected in C. [f.] mirabilis . Of the three tetradic contours detected in C. arcanus sp. nov. (CABD, CDAB, DBAC), only one (CDAB) was detected in C. [f.] mirabilis (Fig. 12).</p><p>Description of the holotype</p><p>FMNH 470769 weighed 37.9 g before it was prepared. The testes were slightly enlarged (2 × 1 mm) and the skull was completely pneumatized. In 2022, the entire dorsal surface of FMNH 470769 was darker than Sepia (119), with the crown being slightly darker than the back and rump. The undertail coverts were Light Drab (119C) and the flanks were darker and cooler than Vandyke Brown (121). The breast and throat were darker than Light Drab (119C). A fine black line connected the malar regions across the throat.</p><p>Etymology</p><p>The scientific name is derived from masculine Latin adjective arcanus (mysterious, secret), referring to the retiring nature of the species. The proposed English name references the Serranía del Darién, which encompasses most of its geographic range.</p><p>Comments</p><p>The sister-species C. arcanus sp. nov. and C. [f.] mirabilis are separated by only 125 km on the Isthmus of Panama and there is no major water barrier between them. The palynological record in lowland Panama, from the Last Glacial Maximum (LGM), includes elements of the montane flora that now occur&gt; 1000 m higher in elevation (Bush and Colinvaux 1990). Given these facts, and the age of the divergence, it seems likely that these mountain-bound species had opportunities for secondary contact in the Darién lowlands during the LGM, and have already passed historical ‘tests’ of sympatry.</p></div>	https://treatment.plazi.org/id/03A087D8836BFFCBB2AE4FA024A5FD1B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
03A087D88369FFCCB2884FE62114F930.text	03A087D88369FFCCB2884FE62114F930.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus mirabilis Nelson 1912	<div><p>Catharus mirabilis Nelson 1912</p><p>Pirre nightingale-thrush</p><p>(Fig. 13)</p><p>Catharus fuscater mirabilis Nelson 1912: 24; Deignan 1961: 430; Ridgley and Gwynne 1989: 352; Phillips 1991: 112; Clement 2000: 299 (in part); Collar 2005: 700; Halley 2020; Halley 2021.</p><p>Catharus fuscater fuscater Hellmayr 1934: 465 .</p><p>Catharus fuscater Meyer de Schauensee 1964: 317 (in part); Robbins et al. 1985.</p><p>Type material</p><p>Monotypic species. USNM 232933 (holotype), study skin, adult male, collected by E.A. Goldman (original no. 15534) at 1585 m elevation on Cerro Pirre, Darién, Panama, near the head of the Río Limón, on April 18 1912 (see: Deignan 1961: 430) . Nelson (1912) also listed the following specimens, which are paratypes (N = 11): USNM 232600, 232927–232932, 232934–232936. The type series was examined by M. R.H. on 23 October 2013 .</p><p>Geographic range</p><p>Endemic to Cerro Pirre, Darién province, Panama, where it is fairly common on the eastern slope from 1000 to 1500 m (Robbins et al. 1985) .</p><p>Adult specimens examined</p><p>Catharus mirabilis (N = 14): Panama: Darién (10 males, four females): Cerro Pirre: USNM 232927–232933, 232936, ANSP 131889, 131890 (males), USNM 232600, 232934, 232935, ANSP 131891 (females) .</p><p>Audio recordings examined</p><p>Catharus mirabilis (N = 15): Panama: Darién: Cerro Pirre: ML 25790, 25807, 25814, 25835, 25872, 60762, 105199, 286837, 286854, 286865, XC 10493, 220586–220588, 361694 .</p><p>Diagnosis</p><p>Genetics: In both phylogenetic reconstructions, samples of C. mirabilis formed a clade that was sister to C. arcanus sp. nov. (mean uncorrected p -distance = 0.04 ± &lt;0.01). In the UCE tree, these taxa were reciprocally monophyletic and formed a clade that was sister to all South American taxa. ABGD and ASAP analyses of ND2 data both identified C. mirabilis and C. arcanus sp. nov. as independent genetic clusters. The estimated divergence time between these sister-taxa was 1.3 Mya (95% HPD = 0. 9–1.7).</p><p>Morphology: Study skins of C. mirabilis were darker, and in adult females showed more contrast between the mantle and crown, than C. arcanus sp. nov (Fig. 13). Study skins of C. mirabilis and C. arcanus sp. nov. were ventrally paler than C. hellmayri specimens collected the same year (Fig. 13). As mentioned previously, the ventral surface of C. mirabilis is ‘distinctly yellowish … on the under parts of the head, neck, and body’ (Nelson 1912), similar to C. arcanus sp. nov. and (to a lesser extent) C. hellmayri . This rapidly fading character distinguishes these three taxa from the South American taxa.</p><p>Voice: Catharus mirabilis was distinguished from all other taxa by the combination of a Type 1 punctuation call structure (Fig. 9) and ‘widely spaced’ contact call structure (Fig. 10). Our dataset lacked recordings of blurred calls from C. mirabilis . Too few recordings were available to determine whether C. mirabilis and C. arcanus sp. nov. are divergent in song. Of the three triadic contours (ACB, CAB, CBA) detected in C. mirabilis, none were shared with C. arcanus sp. nov .. Of the three tetradic contours (BADC, CDAB, DCAB) detected in C. mirabilis, only one (CDAB) was detected in C. arcanus sp. nov. (Fig. 12).</p><p>Comments</p><p>According to published descriptions, the eggs of C. mirabilis (‘very pale greenish white, spotted lightly with dots of cinnamon-brown, which are grouped to form a faintlyindicated cap on the summit of the blunt end’, Wetmore et al. 1984: 157) are patterned differently than the eggs of C. hellmayri (‘pale blue, thickly speckled and dotted and blotched over the entire surface with light chestnut-rufous’, Carriker 1910: 748). Egg colour and pattern have been used previously to distinguish morphologically similar Catharus species (e.g. C. occidentalis and C. ffantzii; Phillips 1969). More research is needed.</p><p>Etymology</p><p>The proposed English name refers to Cerro Pirre, where C. mirabilis is endemic.</p></div>	https://treatment.plazi.org/id/03A087D88369FFCCB2884FE62114F930	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
03A087D8836EFFCEB1A84BD320CAFC21.text	03A087D8836EFFCEB1A84BD320CAFC21.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus fuscater (Lafresnaye 1845)	<div><p>Catharus fuscater (Lafresnaye 1845)</p><p>Cordilleran nightingale-thrush</p><p>(Fig. 13)</p><p>Myioturdus fuscater Lafresnaye 1845: 341 .</p><p>Catharus fuscater Sclater 1859b: 324 (in part); Salvin and Godman 1879: 5 (in part); Seebohm 1881: 285 (in part); Bangs 1899: 108; Allen 1900: 183; Berlepsch 1902; Sharpe 1902: 181; Meyer de Schauensee 1966: 412 (in part); Meyer de Schauensee 1970: 342 (in part); Hilty and Brown 1986; Ridgley and Tudor 1989: 109 (in part); Boesman 1998; Hilty 2003, Naveda-Rodríguez and Bisbal 2008; Biancucci and Martin 2010; López-O. et al. 2014; Remsen et al. 2023 (in part).</p><p>Catharus fuscater sanctae-martae Ridgway 1904: 112; Todd and Carriker 1922: 405; Hellmayr 1934: 465; Meyer de Schauensee 1950: 922; Meyer de Schauensee 1964: 317 (in part).</p><p>Catharus fuscater fuscater Bangs 1930: 332; Hellmayr 1934: 465; Meyer de Schauensee 1950: 922; Meyer de Schauensee 1964: 317; Ginés et al. 1953: 188; Wetmore 1955; Hilty and Brown 1986: 543; Fjeldså and Krabbe 1990: 554 (in part); Clement 2000: 299 (in part); Collar 2005: 700 (in part); Halley 2020; Halley 2021.</p><p>Catharus fuscater sanctaemartae Cracraft 1985: 57; Hilty and Brown 1986: 543; Ridgley and Tudor 1989: 109; Fjeldså and Krabbe 1990: 554; Clement 2000: 299; Collar 2005: 700; Strewe and Navarro 2003; Halley 2020; Halley 2021.</p><p>Type material</p><p>Catharus f. fuscater: MCZ 76525 (holotype), study skin, collected ‘ad Bogotám’ (= Bogotá, Colombia) and exported to France, where it was accessioned (original no. 3544) into the private collection of Lafresnaye (see: Bangs 1930: 332; Halley 2021: 64). The precise type locality remains uncertain, because many ‘Bogotá’ trade skins were not collected in the immediate vicinity of the city. This specimen was not examined in this study.</p><p>Catharus f. sanctaemartae: CM 8797 (holotype), study skin, adult male, collected by ‘ Mrs H. H. Smith’ (=Amelia ‘Daisy’ Woolworth Smith) in ‘Elheibano’ (=El Líbano, see: Paynter 1997: 132), Magdalena, Colombia, on 22 April 1899; chosen from a series of 10 skins reported by Allen (1900), of which the remaining nine skins are paratypes: AMNH 72219–72224, 72226, 72227, 97770. For this study, MRH examined the holotype on 8 February 2023.</p><p>Geographic range</p><p>Catharus f. fuscater: Sierra de Perijá of Venezuela (Zulia), northern Andes of Venezuela (southern Táchira to southern Lara) and Colombia (Cesar and La Guajira), eastern Andes of Colombia (Norte de Santander south to Bogotá); inhabits montane forests between 1500 and 2900 m (Hilty 2003).</p><p>Catharus f. sanctaemartae: Sierra Nevada de Santa Marta of north-eastern Colombia; inhabits forests of the Subtropical Zone from 600 to 2100 m (Ridgway 1904, Todd and Carriker 1922).</p><p>Adult specimens examined</p><p>Catharus f. fuscater (N = 24): Colombia (N = 20): Boyacá (three males): Peña Blanca: CM P59638, P59738, P59739 (males); Casanare (one male): Río Negro: CM P59940 (male) ; La Guajira (four males, one female): Laguna de Junco, Cerro Pintado, Sierra Perijá: USNM 374606, 374607 (males); La Africa, south of Villanueva, Sierra Perijá: USNM 374611, 374612 (males), USNM 374609 (female) ; Magdalena (two males, one female): Monte Elias, Sierra Negra, south-east of Fonseca: USNM 369688, 369689 (males); Sierra Perijá, above Airoca: USNM 374608 (female); Norte de Santander (five males, two female): Buenos Aires, on the Abrego-Sardinata highway: USNM 397776, 398778, 398779 (males), USNM 398780 (female); Las Ventanas: CM P57684 (female); Ocaña: CM P55137 (male); Pueblo Nuevo: CM P55260 (male); Santander (one male): Cachirí: CM P58600 (male); Unknown Department: ANSP 16106 (unsexed) . Venezuela (N = 2): Mérida (one male): La Cutata: AMNH 503904 (male); Trujillo (one male): Guamito: CM P88902 (male) .</p><p>Catharus f. sanctaemartae (N = 33): Colombia (N = 33): Magdalena (eight males, eight females): Cuchilla de San Lorenzo: CM P41731, USNM 387888, 387889 (females); El Líbano: AMNH 72224, 72226, CM P8797 (males), AMNH 72220, 72227, CM P8834 (females); La Cumbre: FMNH 72426 (male), FMNH 72427 (female); Las Taguas: CM P37834 (male); Pueblo Viejo (N slope on Río San Miguel): CM P44941 (female); San Miguel: ANSP 63298, CM P45092 (males); unspecified locality: CM P38599 (male); Cesar (nine males, eight females): Chenducua, on the Río Guatapurí: USNM 387891, 387893–387896, 387900 (males), USNM 384177, 384179, 387892, 387897–387899 (females); Pueblo Bello, Chinchicua, San Sebastián Track: USNM 387902, 387903 (males), USNM 387901 (female); San José, on the Río Guatapurí: USNM 384178 (male); Valparaíso: AMNH 72219 (female) .</p><p>Immature specimens examined</p><p>Catharus f. fuscater (N = 5): Colombia: Magdalena (two males, one female): Monte Elias, Sierra Negra, south-east of Fonseca: USNM 369692, 369691 (males), USNM 369690 (female); La Guajira (one female): La Africa, south of Villanueva, Sierra Perijá: USNM 374610 (female) ; Norte de Santander (one female): Buenos Aires, on the Abrego-Sardinata highway: USNM 398777 (female) .</p><p>C. f. sanctaemartae (N = 0).</p><p>Audio recordings examined</p><p>Catharus f. fuscater (N = 37): Colombia (N = 17): Boyacá: Garagoa: XC 562817; Miraflores: Reserva Sucuncuca: ML 178609131; Cesar: Serrania del Perijá: Chamicero del Perijá Proaves Reserve: ML 104944281, 104944491, XC 693383, 693385; Manaure: ML 204647 204669 204676 192628511, XC 511581; Sabana Rubia Rd.: XC 201876; Meta: San Juanito: XC 523041; Santander: ElTope: ML89839571. Venezuela (N = 20): Lara: Parque Nacional Yacambú: XC 220582, 220583, 220591; Mérida: Cardenal Quintero: XC 387113; El Morro ( Aricaqua Rd.): XC 220580; La Azulita Rd (near Mérida): ML 55724, 66350; La Carbonera: ML 66351–66356; Pico Humboldt Trail: ML 55730; UAL Forest Trail: ML 64833; Santo Domingo: XC 105423; Táchira: Matamula: XC 220579; Páramo Zumbador: ML 66348, 66349; Trujillo: Parque Nacional Guaramacal: ML 151295441, 157081451, XC 220581, 471156.</p><p>Catharus f. sanctaemartae (N = 28): Colombia: Magdalena: Cuchilla de San Lorenzo: ML 211710, 211778, 515400, 515410, XC 45455, 45456, 56808, 56810, 74777, 220584, 235616; Reserva Natural El Dorado: ML 215345031, XC 16014, 43477, 43478, 43550, 75357, 101689, 143821, 154657, 165105, 312974, 312975, 316751, 465174, 481613 191687 191688.</p><p>Diagnosis</p><p>Genetics: In the ND2 reconstruction, samples from the range of C. f. fuscater formed a clade that was sister to a single sequence of C. f. sanctaemartae (mean uncorrected p -distance = 0.02 ± &lt;0.01), with an estimated divergence of 0.7 Mya (95% HPD = 0.31– 1.21). This corroborates Cuervo (2013), who found that C. f. fuscater and C. f. sanctaemartae were sisters, with broader sampling of ND2 sequences from within the range of C. f. fuscater . ABGD analysis of ND2 data identified C. f. sanctaemartae as an independent genetic cluster, but ASAP analysis merged it with C. f. fuscater . We lacked UCE data for C. f. fuscater and C. f. sanctaemartae, so further research will be needed to confirm their sister relationship, to determine their placement in the broader phylogeny of the complex, and to evaluate whether C. f. sanctaemartae is deserving of species rank.</p><p>Morphology: Unlike C. f. sanctaemartae, the phenotype of C. f. fuscater shows puzzling patterns of dichromatism. An unsexed specimen (ANSP 16106) from ‘Santa Fe [de Bogotá]’ in ‘New Grenada’ closely resembles ‘brown type’ males and female specimens of C. [f.] mentalis (e.g. FMNH 433738, 433740) and an individual from Cesar, Colombia, has a brownish dorsal surface (ML 4670402810). In contrast, C. f. fuscater adults in the CM collection, collected in Santander and Norte de Santander, Colombia, show no brown on the dorsal surface, although in only one case were multiple individuals collected at the same site. Specimens of C. f. sanctaemartae have darker ventral plumage than any other taxon, including C. f. fuscater (Fig. 13), rivalled only by C. hellmayri, from which it is distinguished by having more black on the chin (Todd and Carriker 1922) and a longer tail (Hellmayr 1934). We lacked fresh specimens of C. f. sanctaemartae, so were unable to thoroughly evaluate the claim that males are ‘darker and more richly coloured’ than females (i.e. a reported difference in brightness, not hue; Allen 1900, Todd and Carriker 1922). However, no clear differences in plumage colour or brightness are evident in male and female specimens collected by Carriker on 16 May 1925, at La Cumbre, Magdalena, Colombia (FMNH 72426, 72427).</p><p>Voice: The combination of Type 2 punctuation calls (Fig. 9) and ‘widely spaced’ contact calls (Fig. 10) in recordings of C. f. fuscater and C. f. sanctamartae distinguish them from C. hellmayri, C. arcanus sp. nov., and C. mirabilis . Punctuation calls were more elaborate and variable in C. f. fuscater than any other South American taxon (Fig. 9). An exceptional set of C. f. fuscater recordings (ML 66348 –66356) by P. Schwartz features individual birds responding to playback by cycling through repertoires of multiple, distinct Type 2 (‘long/ sinuous’) punctuation calls, and one recording features a Type 3 (‘short/simple’) punctuation call, reminiscent of the calls of C. [f.] opertaneus, Undescribed 2, Undescribed 3, and C. [f.] mentalis, but with a greater frequency range (see Fig. 9). The descending blurred call detected in C. f. fuscater recordings was similar in frequency to, but wider in bandwidth than, the blurred calls of C. [f.] berlepschi (Fig. 11); no recordings of blurred calls were available for C. f. sanctaemartae. In song, the ascending dyadic contour (AB) was more prevalent in C. f. fuscater and C. f. sanctaemartae than in any other taxa (Fig. 12).</p><p>Of the four triadic contours detected in C. f. fuscater (ABC, ACB, BAC, CAB), two were shared with C. f. sanctaemartae (ABC, BAC). Of the three triadic contours detected in C. f. sanctaemartae (ABC, BAC, BCA), two were detected in C. f. fuscater (ABC, BAC). The only tetradic contour detected in C. f. fuscater (DABC) was not detected in C. f. sanctaemartae, which had three tetradic contours of its own (ABCD, BCAD, CDAB). The rare contour BCAD, detected in only one recording of C. f. sanctaemartae (ML 215345031), was not detected in any other taxon.</p><p>Comments</p><p>Lafresnaye’s (1845) holotype (MCZ 76525) was evidently a grey type based on his description, ‘une coloration noirâtre et grise sans la moindre nuance de brun ou de roux’ (‘a blackish and grey colouring without the slightest hint of brown or of red’). Study skins of C. f. fuscater from Boyacá, Colombia at the Instituto de Ciencias Naturales, Universidad Nacional de Colombia (ICN), and at the Instituto Alexander von Humboldt (IAvH), have ‘dark brown to greyish brown’ backs (Beltrán and Kattan 2001), and thus probably represent both colour types. The within-population frequencies of each type are difficult to determine without more data. More study is needed with fresh material.</p></div>	https://treatment.plazi.org/id/03A087D8836EFFCEB1A84BD320CAFC21	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
03A087D8836CFFC3B1AD4EDD2140FA24.text	03A087D8836CFFC3B1AD4EDD2140FA24.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus berlepschi Lawrence 1888	<div><p>Catharus berlepschi Lawrence 1888</p><p>Trans-Andean nightingale-thrush</p><p>(Figs 15–18)</p><p>Catharus fuscater Sclater 1859a: 136, 1859b: 324 (in part); Berlepsch and Taczanowski 1884: 283; Salvin 1895, Goodfellow 1901: 309; Meyer de Schauensee 1966: 412 (in part); Meyer de Schauensee 1970: 342 (in part); Ridgley and Tudor 1989: 110 (in part); Clements and Shany 2001: 193; Solano-Ugalde et al. 2007; Schulenberg et al. 2007 (in part); Kikuchi 2009; Schmitt et al. 2013; Astudillo et al. 2015; Remsen et al. 2023 (in part).</p><p>Catharus berlepschi Lawrence ‘1887’ (=1888; see: LeCroy 2005: 39): 503; Ridgway ‘1887’: 504; Sharpe 1902: 182.</p><p>Catharus fuscater caniceps Chapman 1924: 14; Hellmayr 1934: 466; Bond 1956: 241; Fjeldså and Krabbe 1990: 554 (in part); Collar 2005: 700 (in part); Halley 2020.</p><p>Catharus fuscater fuscater Fjeldså and Krabbe 1990: 554 (in part); Clement 2000: 299 (in part); Ridgley and Greenfield 2001: 660 (in part); Collar 2005: 700 (in part).</p><p>Catharus fuscater berlepschi Halley 2021: 132 .</p><p>Catharus berlepschi nebulus ssp. nov. (formerly Undescribed 3).</p><p>urn:lsid:zoobank.org:act: EAE392CC-A763-4E5E-8F6F-68921515C9A2.</p><p>Catharus fuscater Taczanowski 1874: 504; 1879: 222; 1882: 4; 1884: 483.</p><p>Catharus fuscater caniceps Fjeldså and Krabbe 1990: 554 (in part); Clement 2000: 299 (in part); Collar 2005: 700 (in part).</p><p>‘Unnamed-3’ Halley 2021: 133.</p><p>Type material</p><p>Catharus b. berlepschi: AMNH 39096 (holotype), study skin, adult male, collected by Joseph Siemiradzki in Cayandeled (approximately –2.117°, –78.98°; see: Paynter 1993: 34), Chimborazo, Ecuador, on 23 January 1883 (see: LeCroy 2005: 38). Siemiradzki collected four males and a female, which Berlepsch and Taczanowski (1884: 283) classified as C. fuscater . Berlepsch then sent one of the males (AMNH 39096) to Lawrence, who described it as a new species. For this study, M. R.H. examined the holotype on 20 June 2013 .</p><p>Catharus b. caniceps: AMNH 175530 (holotype), study skin, adult male, collected by H. Watkins (no. 6011) at Palambla (approximately –5.23°, –79.37°; see: Stephens and Traylor 1983: 153), Piura, Peru, on 16 September 1922 (see: LeCroy 2005: 39). For this study, M . R.H. examined the holotype on 20 June 2013 .</p><p>Catharus b. nebulus ssp. nov.: FMNH 474413 (holotype), study skin, adult male, collected and prepared by David E. Willard on Cerro Monte Alegre (–6.5850°, –77.5617°, elev. 2400 m), Amazonas, Peru, on 23 June 2010. When it was prepared, the holotype weighed 37.5 g. Its testes were not enlarged (1 × 0.5 mm) and the skull was 50% pneumatized. In 2022, the entire dorsal surface was darker than Sepia (119), with the crown slightly darker than the back and rump. The undertail coverts were Dark Drab (119B) and the flanks were darker and cooler than Vandyke Brown (121). The breast was darker than Glaucous (79) with faint mottling. The throat was cooler than Drab-Grey (119D), contrasting with the breast.</p><p>Geographic range</p><p>Catharus b. berlepschi: Western Andes in Ecuador (Carchi, south to El Oro).</p><p>Catharus b. caniceps: Western Andes in Ecuador (El Oro), south to north-western Peru (Cajamarca, La Libertad), and the Huancabamba and Chunchuca valleys, west of the Río Marañón .</p><p>Catharus b. nebulus ssp. nov.: Eastern slope of the Peruvian Andes from Amazonas (east of Río Marañón) south to Cusco (north-west of Río Apurímac), including Chachapoyas (Amazonas), Tambillo ( Huamanga), and Chilpes (Junín) between 1770 and 2290 m (Taczanowski 1884).</p><p>Adult specimens examined</p><p>Catharus b. berlepschi (N = 11): Ecuador (N = 11): Carchi (two males, one female): west slope near Maldonado-Tulcán Rd, along Río La Plata: ANSP 181213 181214 (males), ANSP 181212 (female) ; Chimborazo (three males): Chunchi: ANSP 59755–59757 (males); Pinchincha (three males): Chiriboga: DMNH 59252, 62883 (males); Mindo: AMNH 503903 (male); El Oro (two males, two females): Salvias: AMNH 167890 (male); Zaruma: ANSP 83643 (male), AMNH 130154, ANSP 83644 (females); El Chiral: AMNH 167888 (male) .</p><p>Catharus b. caniceps (N = 33): Ecuador (N = 6): Loja (three males, three females): Alamor: AMNH 167894 (female); Celica: AMNH 167891, 167893 (males), AMNH 167892 (female); San Bartolo: AMNH 172116 (male), AMNH 172117 (female) . Peru (N = 27): Piura (four males, four females): Palambla: AMNH 175530–17533 (males), ANSP 116437, 116438, AMNH 175534, 175535 (females) ; Lambayeque (one female): Porculla: ANSP 116441 (female); Cajamarca (six males, one female): Chira: ANSP 116442, 116443 (males); Quebrada Lanchal: LSUMZ 170101, 170102 (males), LSUMZ 170103 (female); Quebrada La Palma: LSUMZ 170104 (male); Cajabamba: AMNH 503909 (male) . La Libertad (eight males, three females): Seques, north-east of Pacasmayo: AMNH 236066, 236068–236070 (males), AMNH 236067 (female); Taulis, north-east of Pacasmayo: AMNH 236071, 236072, 236075, 236076 (males), AMNH 236073, 236074 (females) .</p><p>Catharus b. nebulus ssp. nov. (N = 19): Peru (N = 19): Amazonas (two males, four females): 33 km NE of Ingenio: LSUMZ 82159 (female); Cerro Montealegre: FMNH 474413 (male); La Lejia,north of Chachapoyas: AMNH 235051 (male); Leymebamba: ANSP 109278 (female); 20 km E of La Peca: LSUMZ 88630, 88631 (females); Huánuco (three males): Cayumba Alto: FMNH 299304 (male); CordilleraCarpish: LSUMZ 78972, 80744 (males); Junín (onemale): Rumicruz: AMNH 174319 (male); Pasco (four males, four females): Playa Pampa: LSUMZ 129006, 129007, 129011 (males), LSUMZ 129002, 129005 (females); Santa Cruz: LSUMZ 106496 (male), LSUMZ 106495, 106497 (females); San Martín (one female): 22 km ENE of Florida Pomacochas: LSUMZ 174203 (female) .</p><p>Immature specimens examined</p><p>Catharus b. berlepschi (N = 5): Ecuador (N = 5): Carchi (two females): west slope near Maldonado-Tulcán Rd., along Río La Plata: ANSP 181215 181216 (females); Pinchincha: Chiriboga: DMNH 62882 (male); Mindo Arriba: AMNH 180632 (male); El Oro (two males): Salvias: AMNH 167889 (male); Zaruma: AMNH 130153 (male) .</p><p>Catharus b. caniceps (N = 3): Peru (N = 3): Piura (one male, one female): Palambla: ANSP 116439 (male), CM 119947 (female); Cajamarca (one male): Chugur, 65 km north-west of Cajamarca: AMNH 236065 (male); Quebrada Lanchal: LSUMZ 170100 (female) .</p><p>Catharus b. nebulus ssp. nov. (N = 1): Peru (N = 1): San Martín (one male): 22 km ENE of Florida Pomacochas: LSUMZ 174204 (male) .</p><p>Audio recordings examined</p><p>Catharus b. berlepschi (N = 20): Ecuador (N = 20): Azuay: Cuenca: Molleturo, Corona de Oro Rd.: XC 602205; Yunguilla Reserve: ML 135902, 135926, 76216901, XC 19999; Loja: near Celica: XC 67875; Reserva Utuana: ML 141950781, XC 67874; Pichincha: Bellavista, Tandayapa: XC 128440, 13227, 4214; Mindo: XC 545139, 546525; Mindo Cloud Forest: ML 139115; Old Mindo-Nono Rd., before Tandayapa: ML 63456; Recinto 23 de Junio: ML 216745971, XC 186913; Reserva Paz de Aves: XC 30933; Tandayapa Bird Lodge: XC 32386, 54902.</p><p>Catharus b. caniceps (N = 4): Peru (N = 4): Cajamarca: Chotas, Bosque Protector Pagaibamba: XC 108771; Sallique: Quebrada Lanchal: ML 515357, 515369 ; Piura: Ayabaca, Bosque de Cuyas: XC 231 .</p><p>Catharus b. nebulus ssp. nov. (N = 18): Peru (N = 18): Amazonas: Camino Atuen: ML 398834941; Camp Lowbrow: ML 304312761; Fundo Alto Nieva: ML 72665721; La Llantera cerca de Afluente: ML 272483101; above Leimebamba: XC 175873; Utcubamba: Bagua Grande, ACP Bosque Berlín: ML 129649691, XC 123919; Huánuco: Paty Trail: XC 243435, 243436, 243437; Pasco: Bosque Shollet: ML 228217101, 289516811; Oxapampa: ML 269963401, 376022481; San Martín: Abra Patricia: ECOAN Lodge: XC 30494, 144628; Ucayali: Oventeni: ML 138773; Ulcumano Ecolodge: ML 287691191.</p><p>Diagnosis</p><p>Genetics: In the UCE tree, samples from (i) western Ecuador and north-western Peru, encompassing the type localities of C. berlepschi Lawrence 1888 and C. f. caniceps Chapman 1924, and (ii) eastern Peru from Amazonas south to the Río Apurímac (C. b. nebulus ssp. nov.), formed reciprocally monophyletic clades. Together, these clades formed the sister-group of a clade composed of samples of C. [f.] opertaneus and Undescribed 2. We treat these major sister-groups as polytypic species: C. berlepschi (including C. b. berlepschi, C. b. caniceps, and C. b. nebulus ssp. nov.) and C. opertaneus (including C. [f.] opertaneus and Undescribed 2, see below). The subspecies C. b. berlepschi and C. b. caniceps were not genetically diagnosable, despite their plumage colour differences. A sample of C. b. berlepschi from the Yunguilla Valley, Azuay, Ecuador (ZMUC 128360) was more similar to a paratype of C. f. caniceps (AMNH 175531) than a sample of C. b. berlepschi from Imbabura, Ecuador (ZMUC 119570). In the ND2 tree, these three samples, plus one sample each from the Huancabamba and Chunchuca valleys (i.e. east of the Abra de Porculla, an east–west pass through the Western Cordillera at 2145 m) and a sample from Loja, Ecuador, formed a clade with low within-clade divergence (mean uncorrected p -distance = 0.005 ± 0.004). Samples from across the range of the eastern Andean group (C. b. nebulus ssp. nov.) also formed a clade in the ND2 tree with low within-clade divergence (mean uncorrected p -distance = 0.002 ± 0.001). The divergence between these sister-clades (C. b. berlepschi + C. b. caniceps, vs. C. b. nebulus ssp. nov.) was an order of magnitude greater (mean uncorrected p -distance = 0.022 ± 0.002) than the mean divergence within either clade. ABGD analysis of ND2 data merged C. berlepschi with Undescribed 2 (uncorrected p -distance = 0.01 ± &lt;0.01), although these were not sister-groups in the UCE tree, while treating C. b. berlepschi and C. b. nebulus ssp. nov. as independent genetic clusters. ASAP analysis merged C. b. berlepschi, C. b. nebulus ssp. nov., and Undescribed 2.</p><p>Morphology: Study skins from the range of C. berlepschi in the western Andes were sexually monochromatic in the adult plumage and exhibited a north–south colour cline. At the northern end of the cline, adults had dark dorsal plumage (darker than Sepia, 119), medium-light ventral plumage (Figs 15, 16; Table 3), and completely black maxillae (Table 4), whereas southern adults (C. b. caniceps), including in the Huancabamba and Chunchuca valleys, were paler on the flanks, breast, and throat (Figs 15, 16; Table 3) and had dichromatic irides (Table 5). Specimens from Zaruma, El Oro (ANSP 83643, 83644) had somewhat intermediate characters between the endpoints of the plumage colour cline, and are here listed under the nominate form, C. b. berlepschi . Adult study skins of C. b. nebulus ssp. nov. were dichromatic in iris colour and more similar in plumage colour to C. b. caniceps than C. b. berlepschi . A juvenile specimen identified as C. b. nebulus was morphologically divergent from C. b. berlepschi and C. b. caniceps (see below), but this requires further investigation to confirm.</p><p>Voice: The blurred calls of C. b. berlepschi and C. b. nebulus ssp. nov. had a ‘check-like’ sonogram shape, which distinguished them from all other taxa in the complex, including Undescribed 2, which had a downward sloping blurred call that was narrower in bandwidth (Fig. 11). The subspecies C. b. berlepschi and C. b. nebulus ssp. nov. differed in punctuation call structure (Type 2 vs. Type 3, respectively). In song, of the five triadic contours (ABC, ACB, BCA, CAB, CBA) detected in C. b. berlepschi, four (ABC, ACB, BAC, BCA) were shared with C. b. nebulus ssp. nov.; of the four tetradic contours (ACDB, BCDA, CABD, CDBA) detected in C. b. berlepschi, none was shared with C. b. nebulus ssp. nov., and only one (BDAC) was shared with Undescribed 2.</p><p>Etymology</p><p>We propose the English name ‘Trans-Andean nightingale-thrush’ for the polytypic C. berlepschi because it is the only species in the complex that occurs on both slopes of the Andes. The scientific name C. b. nebulus ssp. nov. is derived from Latin feminine noun nebula (mist, fog), referring to its montane forest habitat.</p><p>Comments</p><p>We placed the name ‘ C. f. caniceps ’ into the synonymy of C. berlepschi because, in both reconstructions, genetic samples from western Ecuador and north-western Peru formed a clade that encompassed the type localities of both taxa, and the older name has priority. Based on the shallow ND2 divergence between samples from the western Andes, on one hand, and the eastern side of the Abra de Porculla in the Huancabamba (LSUMZ 170103) and upper Chunchuca valleys (LSUMZ 170104) on the other, we hypothesize that C. berlepschi crossed the Abra de Porculla, from the west, sometime after the LGM.</p><p>Despite its monophyly in both genetic datasets, we treat the taxon from the eastern Andes as a subspecies of C. berlepschi, instead of its sister-species, because additional sampling is needed to assess the degree of vocal and morphological divergence. We note that C. b. nebulus ssp. nov. is apparently divergent from C. b. berlepschi in juvenile plumage and punctuation call structure, but this may be an artefact of low sample size. Study skins of C. b. nebulus ssp. nov. also exhibit subtle sexual dichromatism, unlike the sexually monochromatic (but clinally variable) C. b. berlepschi + C. b. caniceps clade. Males of C. b. nebulus ssp. nov. (e.g. FMNH 474413, LSUMZ 129006) were darker and less brown on the dorsal surface than adult females, including on the crown, and some adult males of C. b. nebulus ssp. nov. approached the brownish plumage of the adult female (e.g. LSUMZ 106496, with enlarged testes: 9 × 15 mm). This suggests that C. b. nebulus ssp. nov. may exhibit polychromatic plumage ‘types’ like C. [f.] mentalis (see below). Notably, the strongest colour differences between C. b. nebulus ssp. nov. and the C. b. berlepschi + C. b. caniceps clade occur where their ranges approach each other in northern Peru, which may be indicative of character displacement.</p><p>The plumage of a juvenile from central Peru (LSUMZ 174204), identified as C. b. nebulus ssp. nov., was boldly marked relative to a juvenile C. b. berlepschi (ANSP 181216). LSUMZ 174204 has tawny streaks on the mantle, tawny spots on the breast, a whitish throat and abdomen, and a pale mandible (Fig. 19). At first glance, this specimen resembles the juvenile of C. maculatus, but there are multiple reasons to doubt that identification. CM 85107, a genuine C. maculatus juvenile from Cochabamba, Bolivia, has a dark mandible (vs. pale with dusky tip in LSUMZ 174204), a darker and more heavily marked belly (vs. white), and a brown ‘collar’ across the upper breast (vs. no collar). LSU 174204 was also collected at 2300 m, at the same site as an adult female C. b. nebulus ssp. nov. (LSUMZ 174203), and about 500 m upslope of the known upper range limit of C. maculatus in central Peru (Schulenberg et al. 2007). As noted by Ridgely and Tudor (1989: 110), C. maculatus and C. [ fuscater] populations in South America ‘seem rarely if ever actually to occur together, rather, replacing each other in an as yet not understood way’.</p><p>Freeman et al. (2022) played recordings of C. b. nebulus ssp. nov. in ‘territories’ of C. b. berlepschi and recorded the response of the ‘territorial bird’ (i.e. how closely the playback target approached the speaker, and whether and how much it vocalized). In each of eight different experiments, one recording was played in the vicinity of a singing individual, and three different stimuli were used overall (XC 175873 for two experiments, XC 30494 for three experiments, XC 144268 for three experiments). We reiterate that interpreting the results of these experiments (and ‘sympatric’ experiments using audio playback of C. b. berlepschi recordings in the range of C. b. berlepschi) is not straightforward, because the function of song (and, therefore, any response to it) is not clearly defined in species that exhibit cooperative parental care and/or facultative territoriality (e.g. Goetz et al. 2003, Halley 2014, Greeney et al. 2015, Halley et al. 2016).</p></div>	https://treatment.plazi.org/id/03A087D8836CFFC3B1AD4EDD2140FA24	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
03A087D88361FFC5B05948A520C4FC2E.text	03A087D88361FFC5B05948A520C4FC2E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus opertaneus Wetmore 1955	<div><p>Catharus opertaneus Wetmore 1955</p><p>Antioquia nightingale-thrush</p><p>(Figs 15, 16 19)</p><p>Catharus fuscater opertaneus Wetmore 1955: 46; Deignan 1961: 430; Meyer de Schauensee 1964: 317; Hilty and Brown 1986: 543; Fjeldså and Krabbe 1990: 554; Clement 2000: 299; Collar 2005: 700; Acevedo-Charry 2014; Halley 2020; Halley 2021.</p><p>Catharus fuscater Meyer de Schauensee 1966: 412 (in part); Meyer de Schauensee 1970: 342 (in part); Hilty and Brown 1986 (in part); Ridgley and Tudor 1989: 110 (in part); Beltrán and Kattan 2001; Krabbe et al. 2006; Cuervo et al. 2008; Molina Martinez 2014; Greeney et al. 2015; Dyrcz et al. 2015; Remsen et al. 2023 (in part).</p><p>Catharus fuscater fuscater Ridgley and Greenfield 2001: 660 (in part); LeCroy 2005: 38.</p><p>Catharus opertaneus tenebris ssp. nov. (formerly Undescribed 2).</p><p>urn:lsid:zoobank.org:act: B1F7F134-2EB8-4FB9-A1EB-31E2CC7AB4D7.</p><p>Catharus fuscater Parker et al. 1985: 192 196; Rasmussen et al. 1996: 40.</p><p>Catharus fuscater fuscater Clement 2000: 299 .</p><p>Catharus fuscater caniceps M.B. Robbins in Ridgley and Greenfield 2001: 660.</p><p>‘Unnamed-2’ Halley 2021: 133.</p><p>Type material</p><p>Catharus o. opertaneus: USNM 427029 (holotype), study skin, adult male, collected by M.A. Carriker Jr. at ‘ Haciendo Potreros’ (1981 m elev.), ‘on the Río Herradura, 15 miles south-west of Frontino’, Antioquia, Colombia (approximately 6.39°, –76.09°; see: Paynter 1997: 190) on 10 June 1950 (Wetmore 1955, Deignan 1961: 430) . USNM 436771 (paratype), study skin, collected by Carriker on the Río Urrao, Antioquia, on 14 September 1951 . Both specimens were examined by M. R.H. on 23 October 2013 .</p><p>Catharus o. tenebris ssp. nov.: ANSP 185734 (holotype), study skin, adult female, collected and prepared by T. J. Davis in humid montane forest on the eastern slope of Cordillera de las Lagunillas, north bank of Río Isimanchi, about 6 km north-west of San Andrés, Zamora-Chinchipe, Ecuador (–4.7833°, –79.3333°, elev. 2250 m), on 6 November 1992. When it was prepared, the holotype weighed 35 g with no bursa and an ovary that measured 8 × 4 mm. Neither the ova nor the oviduct were enlarged, the skull was 50% pneumatized, and there were insect remains in the stomach. In 2022, the crown of ANSP 185734 was slightly darker than Sepia (219), transitioning to a darker version of Hair Brown (119A) on the back and rump. The undertail coverts were Dark Drab (119B) and the flanks were darker and cooler than Vandyke Brown (121). The breast was Greyish Horn Colour (91) with faint vertical striations. The throat was Drab-Grey (119D), contrasting with the breast, and a dark brown line connected the malar regions across the chin. No moult was noted. Iris colour was recorded as ‘grey’ and there is a pencilled note by M.B. Robbins (MBR): ‘ Catharus fuscater caniceps ? 2 other adults at this locality [KU 186025 and ANSP 186025, both males] had greyish irides; M.B. R. ‘93. [and] 1 in MECN [Museo Ecuatoriano de Ciencias Naturales], Quito’. The maxilla was ‘black’ when the specimen was prepared, and it remains so (Table 4). The mandible and gape were ‘dull orange’ in life, the eye-ring was ‘orange’, and the feet were ‘yellowish brown’.</p><p>Geographic range</p><p>Catharus o. opertaneus: In Colombia, occurs on the western slope of the Western Andes near Frontino and Urrao (Antioquia), at 1980 and 2730 m, respectively (Wetmore 1955), in the Central Andes near the cities of Pereira (Risaralda) and Manizales (Caldas) between 2400 and 2600 m (Beltrán and Kattan 2001), and at several other sites in Antioquia and Caldas (Cuervo et al. 2008) and Tolima (Molina Martinez 2014). In Ecuador, occurs at Yanyacu Biological Station near Cosanga, Napo province (Dyrcz et al. 2015, Greeney et al. 2015). If intervening populations between north-eastern Ecuador (Sucumbios) and northern Colombia (Antioquia) are of the same species, as we hypothesize, then two specimens collected by Carriker near Popayán, Cauca, Colombia, in 1959 and 1960 are probably C. opertaneus (WFVZ 16446, 16447).</p><p>Catharus o. tenebris ssp. nov.: Restricted to the Río Chinchipe watershed of northern Peru and south-eastern Ecuador, south of the Río Zamora.</p><p>Adult specimens examined</p><p>Catharus o. opertaneus (N = 2): Colombia (N = 2): Antioquia (one male, one female): Heda Potreros, 15 miles south-west of Frontino: USNM 427029 (male); Heda la Ilusion, Río Urrao: USNM 436771 (female) .</p><p>Catharus o. tenebris ssp.nov.(N = 4): Ecuador (N = 1): Zamora-Chinchipe (one female): Cordillera de las Lagunillas, 6 km north of San Andrés: ANSP 185734 (female). Peru (N = 3): Piura (two males, one female): Playón: LSUMZ 88629 (male); ‘Machete’, on the Zapalache-Carmen Trail: LSUMZ 97810 (male); ‘Batán’, on the Zapalache-Carmen Trail: LSUMZ 97809 (female).</p><p>Immature specimens examined</p><p>Catharus o. opertaneus (N = 1): Ecuador (N = 1): Napo (1 male): Puente del Rio Quijos: AMNH 180631 (male) .</p><p>Catharus o. tenebris ssp. nov. (N = 0).</p><p>Audio recordings examined</p><p>Catharus o. opertaneus (N = 12): Colombia (N = 7): Antioquia: Reserva Natural Cañón Del Río Claro: ML 101421201 ; Caldas: Reserva Ecológica Río Blanco: ML 101421201, 177887161, XC 440235; Risaralda: Santuario de Fauna y Flora Otún Quimbaya: ML 127271981, XC 102252 ; Tolima: La Plata, Cañón del Combeima: ML 345974171 . Ecuador (N = 5): Napo: Cabañas San Isidro: ML 133453931; Cordillera Guacamayos: ML 142031581; Unknown locality: XC 255039; Volcán Sumaco:XC 443582 ; Sucumbíos: Lumbaquí, Gonzalo Pizarro: XC 529000 .</p><p>Catharus o. tenebris ssp. nov. (N = 4): Ecuador (N = 4): Zamora-Chinchipe: Old Loja-Zamora Rd.: ML 318532061; Parque Nacional Podocarpus: XC 460029; Reserva Tapichalaca: XC 250592, 250647 .</p><p>Diagnosis</p><p>Genetics: In the UCE tree, samples from the ranges of C. o. opertaneus and C. o. tenebris ssp. nov. formed reciprocally monophyletic clades, and together they formed a clade that was sister to the clade consisting of C. b. berlepschi + C. b. nebulus ssp. nov.. In the ND2 tree, C. o. opertaneus was sister to the Darién clade ( C. mirabilis + C. arcanus sp. nov.), with an estimated divergence of 2.5 Mya (95% HPD = 1.9–3.0), and C. o. tenebris ssp. nov. was sister to C. b. berlepschi, with a divergence estimate of 0.4 Mya (95% HPD = 0.2–0.5). ABGD and ASAP analyses of ND2 data both identified C. o. opertaneus and C. o. tenebris spp. nov. as unique genetic clusters (uncorrected p -distance = 0.07), supporting the formal taxonomic recognition of both.</p><p>Morphology: Catharus opertaneus is sexually monochromatic with dark ‘olive brown’ dorsal plumage, unlike any other taxon in the complex (Wetmore 1955; photos in: Beltrán and Kattan 2001, Dyrcz et al. 2015). It was the only taxon with no difference between the sexes in the colour of the maxilla (i.e. both completely black). M.R.H. examined the C. o. opertaneus type in 2015, but lacked specimens for the standardized colour comparison. In comparisons between C. o. tenebris ssp. nov. and the rest of the complex, the closest match in colour was the C. [f.] mentalis adult female, which nevertheless had a darker throat than C. o. tenebris ssp. nov. (Figs 15–17; Table 3). We were unable to determine whether C. o. opertaneus and C. o. tenebris ssp. nov. differ in plumage colour (Table 3), but note that skins of C. o. tenebris ssp. nov. had paler irides (e.g. ‘grey’ ANSP 185734 female, ‘grey brown’ LSUMZ 97809 female, ‘olive-brown’ LSUMZ 88629 male) and ‘orange’ orbital skin (ANSP 185734, LSUMZ 88629, 97809), whereas C. o. opertaneus is said to have ‘dark, cinnamon-brown’ irides and yellow orbital skin, in both northern (Colombian) and southern (Ecuadorean) populations (see photos in: Beltrán and Kattan 2001, Dyrcz et al. 2015).</p><p>Voice: Catharus opertaneus (sensu lato) was distinguished from all taxa, except C. b. nebulus ssp. nov. and C. [f.] mentalis, by its Type 3 (‘short/simple’) punctuation calls (Fig. 9). Too few recordings of song were available to assess the divergence of C. opertaneus from other taxa in the complex. Of the four triadic contours detected in C. o. opertaneus (ABC, ACB, CAB, CBA), only one (ABC) was shared with C. o. tenebris ssp. nov.. Of the four tetradic contours detected in C. o. opertaneus (ABCD, CDAB, CDBA, DCBA), only one (CDAB) was shared with C. o. tenebris ssp. nov.. An aberrant recording of C. o. opertaneus from Sucumbios, Ecuador (XC 529000) contained a unique triadic song type in which the upper two notes were rapidly trilled before descending to a lower note (i.e. a combination of contours BCA and CBA), unlike any other song in our dataset.</p><p>Comments</p><p>Based on geography, we inferred that intervening populations in the Central Andes, between Sucumbios (ZMUC 119587) and Antioquia (ZMUC 134855), belong to the same clade ( C. opertaneus). We hypothesize that the placement of C. opertaneus in the ND2 tree, as sister to the Darién clade with relatively low support, may be evidence of ancient episodes of hybridization and mitochondrial capture between those taxa (see: Toews and Brelsford 2012). Greeney et al. (2015) documented cooperative breeding in C. o. opertaneus in the Ecuadorean population (i.e. five adults provisioning young in a single nest), the first report of this behaviour in a resident Neotropical Catharus species. Previously, this rare behaviour was known only in Bicknell’s thrush ( C. bicknelli, Goetz et al. 2003) and Veery ( C. fuscescens, Halley and Heckscher 2012, Halley 2014, Halley et al. 2016).</p><p>Etymology</p><p>The proposed English name references Antioquia, Colombia, where the types were collected. The scientific name C. o. tenebris ssp. nov. is derived from the feminine Latin noun tenebrae (darkness), referring to the dark forest habitat of the taxon .</p></div>	https://treatment.plazi.org/id/03A087D88361FFC5B05948A520C4FC2E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
03A087D88367FFC6B1BE4EB922E9FC12.text	03A087D88367FFC6B1BE4EB922E9FC12.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Catharus mentalis Sclater and Salvin 1879	<div><p>Catharus mentalis Sclater and Salvin 1879</p><p>Cochabamba nightingale-thrush</p><p>(Figs. 15, 16)</p><p>Catharus mentalis Sclater and Salvin 1876: 352, 1879: 591; Seebohm 1881: 285; Sharpe 1902: 182; Warren and Harrison 1971: 346.</p><p>Catharus fuscater mentalis Berlepsch 1902, Hellmayr 1934: 467; Bond and Meyer de Schauensee 1942, Bond 1956: 242; Ridgley and Tudor 1989: 109; Fjeldså and Krabbe 1990: 554; Clement 2000: 299; Collar 2005: 700 (in part); Halley 2020, Halley 2021.</p><p>Catharus fuscater Meyer de Schauensee 1966: 412 (in part); Meyer de Schauensee 1970: 342 (in part); Schulenberg et al. 2007(in part); Vidoz 2009, Remsen et al. 2023 (in part).</p><p>Type material</p><p>Monotypic species. NHMUK 1885.3.2.35 (syntype), study skin, from the Salvin–Godman Collection, collected by C. Buckley at ‘Suape’ in ‘prov. Yungas, Bolivia’ (= Suapi, La Paz, approximately –15.30˚, –67.31˚; see: Paynter 1992: 141), in 1876 ( Warren and Harrison 1971: 346) ; NHMUK 1885.3.2.36 (syntype), collected by C. Buckley in ‘Yungas’ (presumably at Suapi, La Paz) in 1876 ; NHMUK 1886.8.2.20 (syntype), collected by C. Buckley at ‘Suapi’, Bolivia. These specimens were not examined in this study. Label data were provided by A.L. Bond (in litt.) .</p><p>Geographic range</p><p>Occurs in southern Peru, east of the Río Apurímac, east to Santa Cruz, Bolivia (Vidoz 2009).</p><p>Adult specimens examined</p><p>Bolivia (N = 12): La Paz (five males, three females): Laguna Zongo: DMNH 67204, 67207 (males), DMNH 67201, 67206 (females); unspecified locality on the Río Aceramarca: AMNH 229261 (male); Sandillani: AMNH 138743 (male); Nequejahuruira: AMNH 229258 (male), AMNH 229257 (female) ; Cochabamba (two males, one female): Cerro Incachaca: CM 120346, FMNH 181675 (males), CM 120345 (female) .</p><p>Peru (N = 9): Cusco (five males, two females): La Esperanza, Paucartambo: FMNH 433742, 433738 (males), FMNH 433740 (female); Pillahuatta, Puacartambo: FMNH 430057 (male); Bosque San Luis: FMNH 299706 (female); Limacpunco: FMNH 222381 (male); San Pedro Village: FMNH 364458 (male); Puno (one male): Oconeque: ANSP 102367 (male) .</p><p>Immature specimens examined</p><p>Bolivia (N = 6): La Paz (five males, two females): Laguna Zongo: DMNH 67200, 67202, 67203, 67205 (males), DMNH 67198, 67199 (females); Acara: AMNH 229260 (male) .</p><p>Peru (N = 3): Cusco (one male, two females): Occobauiba Valley, Tocopoquen: USNM 273314 (female); Bosque San Luis: FMNH 299705 (male); Pensión Suecia, km 138.5 on Cusco-Shintuya Highway, Cosnipata Valley: FMNH 398318 (female) .</p><p>Audio recordings examined</p><p>Bolivia (N = 3): La Paz: Fuertecillo: XC 123041; Río Milluni near Titi Amaya, Inquisivi: XC 1998; Muñamachay: XC 73252.</p><p>Peru (N = 4): Junín: Cordillera Vilcabamba, headwaters of Río Poyeni: ML 92162 ; Puno: Sina: ML 148105, 148129, 148130.</p><p>Diagnosis</p><p>Genetics: In the UCE tree, samples of C. mentalis formed a clade that was sister to a large South American clade composed of samples of C. o. opertaneus, C. o. tenebris ssp. nov., C. b. berlepschi, and C. b. nebulus ssp. nov.. In the ND2 tree, samples of C. mentalis also formed a clade, but its placement in the broader phylogeny of the complex was unresolved. The divergence of C. mentalis from C. f. fuscater was estimated at 2.4 Mya (95% HPD = 1.9–3.0), and C. mentalis was also highly divergent from C. b. nebulus ssp. nov., its northern geographic neighbour and the only population likely to establish secondary contact with it during glacial maxima (mean uncorrected p -distance = 0.07 ± &lt;0.01). ABGD and ASAP analyses both identified C. mentalis as an independent genetic cluster.</p><p>Morphology: Specimens of C. mentalis exhibited polychromatic plumage colour, with two distinct colour phenotypes in males, unlike other taxa in the complex (Fig. 8), except possibly C. f. fuscater and C. b. nebulus ssp. nov. (see above). Male specimens were clearly separable into ‘grey’ (FMNH 222381, FMNH 430057) and ‘brown’ (ANSP 102367, FMNH 181675, FMNH 299705, FMNH 364458) phenotypes. ‘Grey’ males were similar to C. f. sanctaemartae males, but with slightly browner (and less dark) breasts and throats and reduced black on the chin. ‘Brown’ males resembled C. mentalis females in plumage colour, and in having a black maxilla (a retained juvenile character).Females of C. mentalis were similar in colour to C. f. fuscater females (Tables 3, 4).</p><p>Voice: Catharus mentalis was distinguished from all taxa except C. o. tenebris ssp. nov., C. o. opertaneus, and C. b. nebulus ssp. nov., by its Type 3 (‘short/simple’) punctuation call structure. Its ‘ascending’ blurred call was of higher frequency, simpler in structure, and shorter in duration than the ‘check-shaped’ blurred calls of C. b. nebulus ssp. nov. and C. b. berlepschi (Fig. 11). Among these taxa, the punctuation calls of C. mentalis were the simplest in structure (Fig. 9). No tetradic song contours were detected in C. mentalis (Fig. 12). Our classification system for song contours was unable to score a remarkably slow-paced song (BAC) followed by a rapid trill (XC 123041) and a six-note contour that featured a typical BA contour followed by an extremely rapid CDAB contour (XC 1998). Although our sample of recordings is small, these two songs were unique in our dataset, suggesting that C. mentalis is vocally divergent from the rest of the complex. Of the three triadic contours (ACB, BAC, CBA) detected in C. mentalis, two (ACB, BAC) were detected in its northern neighbour (C. b. nebulus ssp. nov.).</p><p>Comments</p><p>To our knowledge, the plumage polychromatism of C. mentalis is novel in the C. fuscater complex (except possibly in C. f. [ fuscater] and C. b. nebulus ssp. nov., see above) and unprecedented in the genus Catharus . This pattern is illustrated by two specimens with enlarged testes (FMNH 433738, 433742), collected at the same site in November 2001, a ‘mossy forest’ at La Esperanza (2800 m. elevation), north-east of Paucartambo, Cusco, Peru (Fig. 8).</p></div>	https://treatment.plazi.org/id/03A087D88367FFC6B1BE4EB922E9FC12	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Halley, Maưhew R.;Catanach, Therese A.;Klicka, John;Weckstein, Jason D.	Halley, Maưhew R., Catanach, Therese A., Klicka, John, Weckstein, Jason D. (2023): Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America. Zoological Journal of the Linnean Society 199 (1): 228-262, DOI: 10.1093/zoolinnean/zlad031, URL: http://dx.doi.org/10.1093/zoolinnean/zlad031
