Systematics and distribution of the giant fossil barn owls of the West Indies (Aves: Strigiformes: Tytonidae) Author Suárez, William Author Olson, Storrs L. text Zootaxa 2015 4020 3 533 553 journal article 10.11646/zootaxa.4020.3.7 0485814f-c455-410f-883d-d84e904669e3 1175-5326 240356 E4BC27F4-67DE-4598-BB06-627C9332D101 Tyto noeli Arredondo Noel’s Giant Barn Owl; Lechuza Gigante de Noel ( Figs. 2 A,B,C; 3B; 4A; 5A,B,C,D,G; 6A,E,F; 7A,B,C) Tyto noeli Arredondo, 1972a : 416 (part). ( type locality: “Cueva del Túnel, Término Municipal de La Salud, La Habana [now Mayabeque]” Province, Cuba ). Tyto noelli Arredondo, 1972a : 428 , table 5 (lapsus). Tyto neddi Steadman & Hilgartner, 1999 : 76 , new synonymy (type-locality: "Rat Pocket, Gun Shop Cliff, Two Foot Bay, Barbuda "). FIGURE 4. Right femora of giant species of Tyto from the West Indies in anterior view (A–D) and internal view (E–G): A, T . noeli (large) WS 894; B, T. ostologa USNM 543283; C, T. cravesae , new species (holotype) MNHNCu 75.590; D, T. cravesae , new species (paratype, proximal segment) MNHNCu 75. 594, image reversed to facilitate comparisons; E, T. cravesae , new species (paratype, distal segment) MNHNCu 75. 594, image reversed to facilitate comparisons; F, T. ostologa USNM 543283; G, T. cravesae , new species (holotype) MNHNCu 75.590. Scale =1cm. Holotype . Right tarsometatarsus DPUH 1251. Collected March 3, 1968 , by Noel González Gotera and Oscar Arredondo. Emended diagnosis. The smallest and least robust of the fossil giant barn owls from the West Indies. Material examined. Cueva de Paredones, Ceiba del Agua, Municipality of Caimito, Artemisa (formerly La Habana) Province, Cuba . — Coracoid: nearly complete left (OA 3212, lacking sterno-coracoidal process and procoracoid), nearly complete right ( paratype OA 839, lacking sterno-coracoidal process). Humerus: distal end of left (WS 885, juvenile), proximal half of right (OA 3218). Ulna: incomplete left (WS 899, lacking distal end), distal half of left (OA 3016–3017), proximal half of right (OA 3210), proximal right ( CZACC 400–624, lacking olecranon), distal half of right (OA 3022), shaft of right (WS 915). Femur: complete left (WS 913–914, both fragmentary), distal half of left (OA 3214), distal left ( paratype OA 834, abraded), complete right (OA 3211). Tibiotarsus: distal half of left (OA 3213), distal left ( CZACC unnumbered, juvenile), proximal half of right ( paratype OA 827). Tarsometatarsus: complete left (WS 18A), distal half of left (OA 3112), distal left (WS 916, juvenile), incomplete right (OA 829, OA 3113; both lacking distal end), proximal shaft of right (OA 833), proximal right (OA unnumbered), distal right (OA 3119; WS 882, lacking wing of the trochlea for digit IV). Cueva de Sandoval, Vereda Nueva, Municipality of Caimito, Artemisa (formerly La Habana) Province, Cuba . — Coracoid: complete left (WS 369), complete right (WS 368, WS 851), sternal half of right (WS 909). Humerus: incomplete left (WS 980, lacking distal end, portions of deltopectoral crest and bicipital surface), proximal left (WS 1112, fragmentary), proximal shaft of left (WS 348), distal half of left (WS 983), shaft of left (WS 343, juvenile), distal shaft of left (WS 863, juvenile), distal left (WS 1108), proximal third of right ( CZACC unnumbered), distal half of right (WS 342), distal portion of shaft of right (WS 371–372). Ulna: proximal half of left (WS 370), proximal left (WS 906–907, lacking olecranon), proximal right (WS 905, lacking most of the articular surface), distal right (WS 950, WS 1111). Femur: incomplete left (WS 373, lacking proximal end; CZACC unnumbered, lacking proximal third), proximal half of left (WS 1032), incomplete right (WS 894–895, lacking head; CZACC unnumbered, lacking head and internal condyle), proximal half of right ( CZACC unnumbered), proximal right (WS 781; WS 841, juvenile), distal right (WS 1106, fragmentary). Tibiotarsus: incomplete left (WS 839, lacking proximal end, juvenile), distal half of left (WS 345), distal left (WS 902; WS 984, lacking internal condyle), incomplete right (WS 344, lacking distal end), distal right (WS 901; WS 903, lacking anterior portions of the lateral condyle—904, juvenile). Tarsometatarsus: complete left (WS 897E) and complete right (WS 898E) of the same individual, proximal left (WS 911, fragmentary), proximal right (WS 910, lacking inner calcaneal ridge), distal half of right (WS 846, lacking trochleae for digits II and III ; CZACC unnumbered), distal right (WS 912, lacking trochlea for digit IV, juvenile). Cueva del Indio, Calabazar, Municipality of Boyeros, Ciudad de La Habana Province, Cuba . — Femur: incomplete right (OA 1027, lacking condyles). Cueva del Túnel, La Salud, Municipality of Quivicán, Mayabeque (formerly La Habana) Province, Cuba . — Humerus: distal shaft of left (OA 814), proximal right (WS 295-296, both fragmentary), distal shaft of right (WS 291, juvenile). Ulna: proximal left (OA 807), distal left ( paratype OA 816), distal right ( paratype OA 815). Femur: incomplete left (OA 819, lacking both ends), incomplete right ( paratype OA 818, with fragmentary condyles). Tibiotarsus: proximal left (OA 809, lacking inner cnemial crest), distal half of left (WS 136), distal left ( paratype OA 812), distal shaft of left (OA 810), proximal half of right (WS 213, lacking outer cotyla), proximal right (WS 214), distal half of right (OA 3118). Tarsometatarsus: complete right ( holotype DPUH 1251), medial segment of right (OA 813, OA 821), distal right (OA 817). Cueva de Insunza, La Salud, Municipality of Quivicán, Mayabeque (formerly La Habana) Province, Cuba . — Humerus: distal shaft of right (WS 268). Tarsometatarsus: nearly complete left (WS 15, lacking fragments of shaft posteriorly). Cueva del Chicharrón, La Salud, Municipality of Quivicán, Mayabeque (formerly La Habana) Province, Cuba . — Carpometacarpus: incomplete left ( CZACC unnumbered, lacking metacarpal III ). Cuevas Blancas, Aguacate, Municipality of Quivicán, Mayabeque (formerly La Habana) Province, Cuba . — Carpometacarpus: proximal left ( CZACC unnumbered, CZACC unnumbered). Tibiotarsus: distal right ( CZACC unnumbered). Tarsometatarsus: proximal half of left ( CZACC unnumbered, lacking inner calcaneal ridge), distal left ( CZACC unnumbered). Cantera cerca de Loma de Mosas, 5 km Northeast of the city of Sancti Spirítus, Sancti Spirítus Province, Cuba . — Femur: proximal fragment of right (OA 3120). Cueva de la Jutía, Loma de los Rubíes, Municipality of Florencia, Ciego de Ávila Province, Cuba . — Tarsometatarsus: distal left ( CZACC unnumbered). Skeleton (Goat Bone) Cave, Clarendon Parish, Jamaica . — Coracoid: left ( USNM 535720). Drum Cave, Clarendon Parish, Jamaica . — Tarsometatarsus: distal half of right ( USNM 615828, lacking trochleae for digits II and III ), distal left ( USNM 615829, lacking trochlea for digit IV). Rat Pocket, Gun Shop Cliff, Two Foot Bay , Barbuda . — Coracoid: humeral end of left ( paratype of T . neddi USNM 359245). Femur: distal right ( holotype of T . neddi USNM 359240, fragmentary). Additional type material of T . neddi , not listed herein, includes a few pedal and ungual phalanges (see Steadman & Hilgartner 1999 ). Distribution. Cuba , from the province of Pinar del Río (see Suárez & Diáz-Franco 2003 , Suárez 2004b ) to Ciego de Ávila; also south-central Jamaica and Barbuda (see Fig. 1 ). Remarks. Tyto noeli is the commonest extinct large barn owl in the Quaternary of Cuba , its type series being the largest ( Arredondo 1972a ) for any fossil tytonid described from the West Indies. This taxon was originally known from two Quaternary cave sites in the former La Habana Province (now split in two provinces: Artemisa to the west, and Mayabeque to the east). Although Arredondo (1972a: 416) commented on the abundance of postcranial elements of T . noeli in his collection (at least 23 specimens ), in the original description only ten were used, plus three others that appeared only in tables and figures (for a total of 13; note that the legends to figures 2 and 4 are reversed in the original description). At that time, Arredondo (1972a) could rely only on the little published information on T. ostologa and T. pollens for comparison with T. noeli and hence was under the impression that T. noeli was about the same size, or even larger than those two species, which is not the case, however (for a correct size correlation see Olson & Hilgartner 1982 : 36; Steadman & Hilgartner 1999 : 79). Confusion arose because the type series was a composite which included a large and robust tarsometatarsus (and other specimens) lacking the distal end (OA 828), referred herein to a new species described below. In Table 2 of the original description Arredondo estimated the total length of this specimen at 100 millimeters, much larger even than T . pollens , which was incorrect. Arredondo did not consider that the holotypical tarsometatarsus of T . noeli had a shorter shaft when compared with the equivalent element in T . furcata , the species used by him as a model for his hypothetical reconstructions (O. Arredondo pers. comm. to W. Suárez in 1999). Overestimation is evident also in the length of the tibiotarsus reconstructed, based on fragments from different individuals (see Arredondo 1972a , figs. 4, 3). A relatively shorter shaft of the tarsometatarsus, compared with T . alba or with T . furcata , is a shared character of all the giant Tyto species from the West Indies (see Wetmore & Swales 1931 : 237; Brodkorb 1959 : 357). That mistake is what led Arredondo to consider that T. noeli , T. ostologa , and T. pollens were about the same size in all his subsequent publications ( Arredondo 1972b ; Acevedo-González et al. 1975 ; Arredondo 1976 , 1982 , 1984 ). Furthermore, and as result of that confusion, some fossils described herein as a new species, larger than T . noeli , provided the basis for speculations about the presence of T . riveroi in cave deposits in the former La Habana Province (see Suárez & Arredondo 1997 : 101). A few fragments (see Material examined; Figs. 2 A, 7B) from two caves in the Portland Ridge area of southern Jamaica collected in 1995 establish that T. noeli also occurred on that island. These bones probably date from well into the Holocene as surface material from Drum Cave had a radiocarbon age of 3700±150 yr and the lower level of Skeleton Cave dated to 6410±110 yr (McFarlane et al. 2002). In contrast, the only bones of this tytonid yet dated directly are from the Cueva El Abrón deposit in Pinar del Río Province, Cuba , of late Pleistocene (17, 406 ±161yr) age ( Suárez & Diáz-Franco 2003 ). We also compared the holotypical fragmentary distal end of a femur (USNM 359240) and the paratypical fragment of a coracoid (USNM 359245) of Tyto neddi from the island of Barbuda ( Steadman & Hilgartner 1999 ) with equivalent material of T. noeli from Cuba and could find no quantitative or qualitative characters by which the two could be separated ( Fig.5 A, C; Table 3 ). These two species had never been compared directly before. Although it is possible that more material of this owl from the Lesser Antilles might reveal some differences from T. noeli of Cuba and Jamaica , there is no basis for making a distinction at present. The characters proposed by Steadman & Hilgartner (1999: 77) in the diagnosis of Tyto neddi are found in our series of T. noeli . Therefore we provisionally consider Tyto neddi Steadman & Hilgartner (1999) to be a junior subjective synonym of T. noeli Arredondo (1972a) . TABLE 3. Measurements (mm) of elements of Tyto noeli from Cuba, Jamaica, and Barbuda.

Measurement	Cuba Range (mean) n	Jamaica Range (mean) n	Barbuda Range (mean) n
Coracoid Length of glenoid facet Width of shaft at narrowest point Depth of shaft at narrowest point	11.0–11.9 (11.4) 3 5.3–5.9 (5.5) 4 3.4–3.7 (3.6) 4	10.9 5.4 3.6	11.5 — 3.4
Femur Distal width Depth of internal condyle	13.4–15.4 (14.4) 10 8.6–10.0 (9.3) 10	— —	15.3 9.9
Tarsometatarsus Shaft width at midpoint Shaft depth at midpoint Width through trochleae for digits II and III Depth of trochlea for digit II	5.3–6.7 (6.1) 16 4.9–6.7 (5.4) 16 11.2–12.5 (11.9) 4 8.4–10.7(9.5) 8	5.9 4.8+ 10.8+ 9.3	— — — —
+ Abraded.

FIGURE 5. Distal ends of right femora in anterior (A, B) and posterior (C, D) views and distal ends of left tibiotarsi in anterior view (E–G) of giant species of Tyto from the West Indies: A and C, T . noeli (Barbuda, holotype of T . neddi ) USNM 359240; B and D, T . noeli (Cuba) WS 894; E, T. cravesae , new species (paratype) OA 831; F, T. cravesae , new species (paratype) MNHNCu 75.593; G, T. noeli (large) WS 345. Scale =1cm. The series of Tyto noeli we examined increases our knowledge of the size range for this species, providing evidence of marked sexual dimorphism, as can be expected in a large nocturnal raptor, in agreement with material of the other species described from the West Indies ( Olson & Hilgartner 1982 ; Steadman & Hilgartner 1999 ). From Barbuda , two different-sized species of Tyto were recorded by Steadman & Hilgartner (1999) , the smaller being about the size of the living Tyto alba and identified as Tyto sp. Pavia (2004: 635) interpreted these differences in size as sexual dimorphism of a single species of Tyto that occurred in that island. The series of large Tytonidae from the West Indies we examined does not support Pavia’s point of view, and we agree with Steadman & Hilgartner (1999) in recognizing two species from Barbuda . Similarly, remains of Tyto furcata and T. noeli can also be found in association in Cuban cave deposits ( Arredondo 1972a , 1975, 1976, 1982, 1984; Suárez 2000). Specimens of Tyto noeli are consistently larger and more robust than those of T . furcata ( Arredondo 1972a ) , and smaller and more gracile than in T. ostologa , T. pollens ( Table 1–2 ; see Olson & Hilgartner 1982 ; Steadman & Hilgartner 1999 ), or in the new species described below.