Integrative taxonomy reveals hidden diversity in the Catharus fuscater (Passeriformes: Turdidae) complex in Central and South America Author Halley, Maưhew R. Department of Biodiversity, Earth and Environmental Science, Drexel University, Philadelphia, Pennsylvania, USA, 19104 Author Catanach, Therese A. Ornithology Department, Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania, USA, 19103 Author Klicka, John Burke Museum of Natural History and Culture, Department of Biology, University of Washington, Seaưle, Washington, USA, 98105 Author Weckstein, Jason D. Department of Biodiversity, Earth and Environmental Science, Drexel University, Philadelphia, Pennsylvania, USA, 19104 & Ornithology Department, Academy of Natural Sciences of Drexel University, Philadelphia, Pennsylvania, USA, 19103 text Zoological Journal of the Linnean Society 2023 2023-07-07 199 1 228 262 http://dx.doi.org/10.1093/zoolinnean/zlad031 journal article 266146 10.1093/zoolinnean/zlad031 b9190aec-ed9f-4c04-8914-daa1913ef189 0024-4082 8326302 Catharus hellmayri Berlepsch 1902 Talamanca nightingale-thrush ( Fig. 13 ) 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 . 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 . Catharus hellmayri Sharpe 1902: 182 ; Davis 1972: 176 . Type material 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. ) . Geographic range Northern mountains of Costa Rica (Rincón de la Vieja, Miravalles, Tenorio) to west-central Panama (Parque Nacional Santa Fé, Veraguas ). Adult specimens examined 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). 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 . Immature specimens examined 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) . Audio recordings examined 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. 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. Diagnosis 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). 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 ). 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. Etymology The proposed English name references the Cordillera de Talamanca, which encompasses nearly the entire range of C. hellmayri .