Catharus fuscater ( Lafresnaye 1845 )

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), pp. 228-262 : 248-250

Catharus fuscater ( Lafresnaye 1845)

Cordilleran nightingale-thrush

( Fig. 13 )

Myioturdus fuscater Lafresnaye 1845: 341 .

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).

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).

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.

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.

Type material

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.

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.

Geographic range

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).

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).

Adult specimens examined

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) .

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) .

Immature specimens examined

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) .

C. f. sanctaemartae (N = 0).

Audio recordings examined

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.

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.

Diagnosis

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 ± <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.

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).

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 ).

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.

Comments

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.

Figures

Figure 9. Taxonomic variation in the structure of ‘punctuation calls’, visualized in RavenPro 1.5 (Cornell University). We recognize three groups based on shared acoustic structure: (Type 1) pulsed/rippling:C. [f.] hellmayri, Undescribed 1, C. [f.] mirabilis; (Type 2) long/sinuous: C. [f.] sanctaemartae, C. [f.] fuscater, C. [f.] berplepschi; (Type 3) short/simple: Undescribed 2, C. [f.] opertaneus, Undescribed 3, C. [f.] mentalis.

Figure 10. Taxonomic variation in the structure of ‘contact calls’, visualized in RavenPro 1.5 (Cornell University). No recording of contact calls in C. [f.] mentalis was available.

Figure 11. Taxonomic variation in the structure of ‘blurred calls’, visualized in RavenPro 1.5 (Cornell University). The black line divides calls that modulate upward in frequency (left) and downward in frequency (right); both types were detected in C. [f.] berlepschi.

Figure 12. Heat map showing frequency distributions of song contours within each population. Dyadic, triadic, and tetradic song contour names follow Fig.5. Single-note songs are shown in the first column (A) and songs with five or six notes were combined into a single category, shown in the last column (FIVESIX). Open (white) circles denote song contours that were not detected in C. [f.] hellmayri, despite it having a much larger sample size than other taxa. Closed (black) circles denote contours that were only detected in one taxon.

Figure 13. Dorsal and ventral views of the adult plumages in C. hellmayri, C. arcanus sp. nov., C. mirabilis, and C. f.sanctaemartae. Photograph by M.R.H.