Systematics and distribution of the living and fossil small barn owls of the West Ind (1 Viewer)

Systematics and distribution of the living and fossil small barn owls of the West Indies (Aves: Strigiformes: Tytonidae)
WILLIAM SUÁREZ, STORRS L. OLSON

Abstract

After reviewing the systematics and distribution of the living and fossil small West Indian taxa of Tytonidae (Tyto), we reached the following conclusions: (1) Strix tuidara J. E. Gray, 1827, type locality of Brazil, is the earliest available and correct name to be used in a binomen for New World mainland barn owls; (2) the North American mainland subspecies Tyto tuidara pratincola (Bonaparte, 1838), new combination, is resident in the Bahamas (“Tyto perlatus lucayanus” Riley, 1913, is a synonym), where it probably did not colonize until after the European introduction of Rattus Fischer, in Hispaniola (Dominican Republic and Haiti) where it became established in the 20th century, and subsequently in Puerto Rico; (3) Tyto furcata (Temminck, 1827) of Cuba, Jamaica and the Cayman Islands is a different species restricted to its insular distribution, with “T. alba niveicauda” Parkes & Phillips, 1978, of the Isle of Pines (now Isla de la Juventud) as a synonym; (4) the distinct species Tyto glaucops (Kaup, 1852), now endemic to Hispaniola, once occurred in Puerto Rico, as the fossil species “T. cavatica” Wetmore, 1920, is here shown to be a synonym; (5) the smallest taxon Tyto insularis (Pelzeln, 1872) of the southern Lesser Antilles is treated as a separate species, in which the nominate subspecies T. i. insularis (St. Vincent, Grenada, and the Grenadines) differs slightly but consistently in coloration from T. i. nigrescens (Lawrence, 1878) of Dominica; (6) another barn owl, Tyto maniola, new species, of this group of small tytonids from the West Indies inhabited Cuba during part of the Quaternary, and is here named and described.



Keywords

Aves, Tytonidae, Distribution, Fossil Birds, Systematics, Tyto, West Indies


www.biotaxa.org/Zootaxa/article/view/zootaxa.4830.3.4

Systematic Paleontology -1-

Systematic Paleontology

Class AVES Linnaeus
Order STRIGIFORMES (Wagler)
Family TYTONIDAE Ridgway
Genus Tyto Billberg
Tyto tuidara pratincola (Bonaparte), new combination
Strix pratincola Bonaparte, 1838: 7 (type locality: “northern parts” of America = North America).
Tyto perlatus lucayanus Riley, 1913: 153, synonym (type locality: New Providence Island, Bahamas).

Distribution in the West Indies. Resident in most of the islands of the Bahamas (Bond 1956; Buden 1987; White 1998). Accidental in Cuba (Garrido 1978; Parkes & Phillips 1978), Jamaica and Puerto Rico (Raffaele 1989; Raffaele et al. 1998; Jansen & Fuchs 2019) but has recently become established in the latter (Thorstrom & Gallardo 2017, Fig. 1). A nesting pair was located a few years ago in the municipality of Aguada, Puerto Rico (Sergio A. Colón, pers. comm. to W. Suárez, 2015; see Thorstrom & Gallardo 2017). A mounted, historical specimen identified as Tyto alba ssp. was obtained, supposedly from Puerto Rico, during the Baudin Expedition (1796–1798), but it may have come from elsewhere (see Jansen & Fuchs 2019). Bond (1963, 1980:5) reported this subspecies from Hispaniola (which it colonized ca. 1950), where by some time prior to 1980 it was said to be “widespread in open country, but appears to be most numerous in the extreme northwestern part of the Dominican Republic.” Keith et al. (2003) listed 28 specimens of this taxon in three museums taken in Haiti (8) and Dominican Republic (20) and felt that some specimens indicated vagrancy from North America in winter in addition to a resident population. Individuals of this barn owl have been observed several times in the Dominican Republic by one of us (Suárez unpubl. data). Steadman & Takano (2013) reported remains identified as Tyto alba (= T. t. pratincola, see op. cit., p. 940) from high elevation late Holocene cave deposits in Haiti (Massif de la Selle) dated at c. 1600–600 cal. BP. They therefore suggested that “fossils of Tyto alba confirm that this species, believed perhaps to be a recent colonist to Hispaniola…has a history there measured in millennia rather than decades”, meaning that it must have been resident in Hispaniola well before the arrival of Europeans in the New World. They did not address the possibility, however, that the fossil material from Haiti might pertain to the Antillean species Tyto furcata (see Discussion), which is very common in nearby Cuba (Garrido & García Montaña 1975; Garrido & Kirkconnell 2011), rather than to the recent colonizer.
Remarks. The putative Bahaman subspecies “Tyto perlatus lucayanus”, established by Riley (1913) on very weak characters, was recognized by Ridgway (1914) and Bond (1956), but on comparison with better material by Buden (1987) was deemed inseparable from the mainland subspecies pratincola. Our examination of the holotype of “T. p. lucayanus” and an additional specimen from New Providence Island fully corroborates Buden’s conclusion.

Tyto furcata (Temminck)
Strix furcata Temminck, 1827: text accompanying plate 432 (type locality: “au Mexique et aux Antilles; l’individu don’t nous donnons la figure a été envoyé de l’ile de Cuba”).
Tyto alba niveicauda Parkes & Phillips, 1978: 483, new synonymy (type locality: Los Indios, Isle of Pines, Cuba).

Distribution. Resident in the Cuban archipelago, Jamaica and the three Cayman Islands (Fig. 1). Erroneously considered as resident in the Bahamas by Steadman & Hilgartner (1999:79).
Remarks. Tyto furcata is one of the most common tytonids found in Quaternary cave deposits in Cuba, sometimes in association with other large extinct raptors (see Arredondo 1971, 1984; Suárez 2000a; Suárez & Olson 2015). Remains of T. furcata have been identified as well from asphalt deposits at Las Breas de San Felipe, Matanzas Province (Table 4), Cuba, from where some remains may fall well within the late Pleistocene. The Whitewinged Barn Owl originated in the Cuban archipelago from where it colonized Jamaica, the Caymans and probably other islands (cf. Hispaniola, see preceding species). This taxon is the main species responsible for Cuban deposits containing the remains of some extinct small vertebrates, where the nesophontid insectivore Nesophontes micrus G. M. Allen, 1917, and the echymid rodent Boromys torrei G. M. Allen, 1917, were the principal prey. After the total extinction of the former endemic mammals during the Holocene (Alcover et al. 1998; Silva Taboada et al. 2008), this barn owl probably was forced to prey more on species of bats, birds, immature specimens of the living hutias (Suárez, pers. observ.), and other small vertebrate and invertebrates, as it does today (see Johnston 1974; MacFarlane & Garrett 1989; Suárez 1998; Arredondo & Chirino 2002; Jiménez Vázquez et al. 2005; Hernández-Muñoz & Mancina 2011; López 2012; Morgan et al. 2019, among others). Following European colonization, Tyto furcata turned to the equivalent-sized introduced rodents of the genera Mus Clerck and Rattus Fischer (see literature above) as their principal prey in a new era for this species.

Systematic Paleontology -2-

Tyto glaucops (Kaup)
Strix glaucops Kaup, 1852: 118 (type locality: “Jamaica” = St. Domingo by inference from Sharpe 1875: 302 = “Haiti or Santo Domingo” of subsequent authors, e.g. Ridgway 1914: 612).
Strix dominicensis Cory, 1883: 95, synonym (type locality: Santo Domingo = Puerto Plata, Dominican Republic fide Wetmore & Swales 1931). Nec Strix dominicensis Gmelin, 1788: 296.
Tyto cavatica Wetmore, 1920: 80, new synonymy (type locality: “cave on the property of Don Gervacio Toraño, near Utuado, Porto Rico”).

Distribution. Resident in Hispaniola; formerly present in Puerto Rico, where known only from Quaternary
fossil remains
Remarks. No resident species of Tyto has occurred in Puerto Rico in historical times until recently (see account for T. tuidara pratincola above). Fossils from Quaternary cave deposits on the island were described as an extinct species “Tyto cavatica” Wetmore (1920, 1922). The original material consisted of two proximal ends of tarsometatarsi and two distal ends of tibiotarsi, all from a single cave. These were considered to be closest to T. glaucops, but the species “T. cavatica” was distinguished on a supposed difference in the “talon” (inner calcaneal ridge) of the tarsometatarsus. Some subsequent literature (e.g. Steadman & Hilgartner 1999) conveys the idea, without reference to additional material, that “T. cavatica” was larger than T. glaucops, although Wetmore (op. cit.) made no such assertion, merely maintaining that it was smaller than T. alba perlata (= T. tuidara pratincola).
The main difference between “Tyto cavatica” and T. glaucops cited by Wetmore (1920:80) was that the main hypotarsal ridge is “larger and much longer” in the former. As he noted, however, this feature was not readily apparent in the holotype that he selected but was best observed in the paratypical tarsometatarsus that is now USNM 241263. This specimen is very obviously from a juvenile, however, so that the bone is still porous and evidently in the process of formation. Among more recently collected specimens from Puerto Rico is a complete tarsometatarsus of an adult (USNM 510230) in which the hypotarsus does not differ in size from that of T. glaucops. With regard to size, all of the newer fossil material from Puerto Rico falls exactly in the range of T. glaucops (Table 2–3) and we could detect no differences whatsoever between these two populations. Therefore, we consider “Tyto cavatica” Wetmore, 1920, to be a junior subjective synonym of T. glaucops (Kaup, 1852). The Baudin specimen supposedly from Puerto Rico does not represent Tyto glaucops (see Jansen & Fuchs 2019). Although the Puerto Rican extinct population of T. glaucops may represent a different subspecies from that of Hispaniola, no differences are known at present to justify use the name cavatica in a trinomen for its remains.

Tyto insularis insularis (Pelzeln)
Strix insularis Pelzeln, 1872: 23 (type locality: “Ins. St. Vincent (Capverden?)”, corrected to St. Vincent, West Indies by Hartert 1929).
Hybris nigrescens noctividus Barbour, 1912: 57, synonym (type locality: St. George’s, Grenada).

Distribution. Resident on the islands of St. Vincent, Grenada and the Grenadines, Lesser Antilles
Remarks. Barbour’s (1912) name noctividus for birds of Grenada was made in comparison only with birds from Dominica and without reference to Pelzeln’s name insularis, which at that time would have been thought to apply to birds of the Cape Verde Islands. Hartert (1929) emphatically synonymized noctividus with insularis, and we agree.

Tyto insularis nigrescens (Lawrence)
Strix flammea var. nigrescens Lawrence, 1878: 64 (type locality: Dominica).

Distribution. Resident on the island of Dominica, Lesser Antilles

Systematic Paleontology -3-

Tyto maniola, new species

The fossil material described and named herein is referable to Tytonidae and to the genus Tyto (see Figs. 3, 6, 7), and differs from Antillean Strigidae of similar size (see Materials and Methods) by the following combination of osteological characters (see also Mourer-Chauviré 1987; Arredondo & Olson 1994; Steadman & Hilgartner 1999; Pavia 2004; Suárez & Olson 2015, among others): Humerus with bicipital furrow very deep and impression of brachialis anticus much deeper. Ulna with prominence for anterior articular ligament more projected; impression of brachialis anticus deeper and more distally extended; bicipital attachment reduced; internal condyle better developed, and the sulcus intercondylaris much more extended into the internal side. Carpometacarpus with a deep fossa distad to the pisiform process (forming two marked ridges at the base of the latter); metacarpal II with internal border acute (less rounded); metacarpal III longer and well extended proximad. Tarsometatarsus with intercotylar prominence placed more anteriorly in relation to the position of anterior edges of both internal and external cotylae; ossified proximal supratendinal bridge absent (see also Brodkorb 1969:113); shaft convex anteriorly at the distal end, with lateral surface expanded (deeper); trochlea IV massive; intertrochlear notch between trochleae II and III reduced.
Holotype. Proximal half of left tarsometatarsus (MNHNCu 75.4651), collected by members of the Departamento de Geología y Paleontología (MNHNCu), during 25–28 November 1998.
Type locality. Las Breas de San Felipe (San Felipe II), Matanzas Province, Municipality of Martí, San Felipe Valley, 5.5 kms west of the town of Martí (ca. 22º57’N; 80º58’W; sheet Martí 4084-IV, 1:50,000 map, X502.850, Y347.100). Las Breas de San Felipe are the only known tar seeps in the Greater Antilles and contain two productive fossil sub-localities known as San Felipe I and San Felipe II, respectively (for detailed description and related fauna see Iturralde-Vinent et al. 2000; Suárez 2000a; Suárez & Emslie 2003; Suárez & Olson 2003a,b,c, 2007, 2009, 2015; Silva Taboada et al. 2008; Table 4). This is the type locality of the extinct Cuban Caracara Milvago carbo Suárez & Olson (2003c) and other Cuban extinct birds (see Suárez 2020).
Chronology. Late Pleistocene. Accelerator-mass spectrometer (AMS) radiocarbon (14C) dating of hindlimb material (directly associated with the holotype at San Felipe II) of Ornimegalonyx oteroi Arredondo, 1958, Gymnogyps varonai (Arredondo, 1971), and Antigone cubensis (Fischer & Stephan, 1971) (NSF laboratory numbers: AA 50562–50564, respectively; Arizona AMS facility, University of Arizona) range from 22,000±2,600 to >41,000 yr BP (Suárez 2020). In addition, radiocarbon dates from Las Breas de San Felipe based on humeral material of the extinct Cuban sloth Parocnus browni Matthew, 1931, yield ages ranging from 4,960±280 to 11,880±420 yr BP (see Jull et al. 2004; Steadman et al. 2005).
Paratypes. San Felipe I.—Tarsometatarsus: proximal left without inner calcaneal ridge (MNHNCu 75.4655), proximal right (MNHNCu 75.4656), distal left (MNHNCu 75.4657). San Felipe II.—Carpometacarpus: right fragmentary (MNHNCu 75.4654). Tarsometatarsus: distal end of right (MNHNCu 75.4652). All collected by William Suárez and Stephen Díaz Franco, 22–25 February 2001.
Cueva El Abrón, western part of Sierra de La Güira, municipality of La Palma, Pinar del Río Province, Cuba.—
Humerus: complete right (juvenile) WS 0.435. Collected February 1997 by William Suárez, Jesús Martínez and
Gabino de La Rosa.
Cueva de Paredones, Ceiba del Agua, Municipality of Caimito, Artemisa (formerly La Habana) Province,
Cuba.—Ulna: nearly complete right ulna WS 0.436. Collected March 1997 by William Suárez.
Etymology. Diminutive of L. mania, a goblin or specter to frighten children, from the small size of the species and the dread with which many people regard owls.
Diagnosis. Much smaller than Tyto furcata or T. tuidara pratincola, slightly smaller than T. glaucops, and larger than T. insularis, T. bargei, or T. punctatissima (see Tables 2–3). Differs from all of these taxa by having the tarsometatarsus with anterior metatarsal groove deeper proximally and the intercotylar prominence tending to be less pointed.
Remarks. This new species was discovered by WS during the 1990s on the basis of the only two cave specimens (humerus and ulna) described above. The ulna specimen from Cueva de Paredones was so fresh in appearance that it motivated exhaustive exploration by WS for several years, in attempts to locate a living bird. Further exploration in suitable regions is still needed to adequately test this possibility. This new tytonid is rare and remains scarce in the fossil record of Cuba, in contrast to the abundance of Tyto furcata and the much larger extinct species, T. noeli Arredondo, 1972 (see Suárez & Olson 2015). Deposits of accumulated pellets of T. maniola, new species, are virtually unknown in the Cuban archipelago. Smaller mammal species such as Nesophontes micrus and Boromys torrei,together with other supplementary vertebrates and invertebrates of adequate size, were the best candidates to form the bulk of its prey in Cuba. The known specimen from Cueva El Abrón was collected at about 1m depth. This level was later determined to belong to “layer VII” (from 1.00 to 1.60 m depth, brownish-yellow in color), after this important owl pellet deposit was revisited and better excavated by personnel of the MNHNCu. A new species of bat was discovered (Suárez & Díaz-Franco 2003) but no additional elements of Tyto maniola, new species. Associated there with the abundant material of the above-mentioned mammals were bird species not found today (extirpated) in Pinar del Río Province (see Garrido & Kirkconnell 2011), including the Cuban Parakeet Psittacara euops (Wagler, 1832), the Zapata Wren Ferminia cerverai Barbour, 1926 (for additional bird species recorded there see Suárez & Díaz-Franco 2003; Suárez 2004), and a large species of swift. Bones of Zapata Wren are common in some Quaternary cave deposits in western Cuba (Suárez unpubl. data), but the species had been confounded with a tapaculo (=?Scytalopus sp.), as result of their very similar morphology (see Olson & Kurochkin 1987; Jiménez Vázquez et al. 2005).
Possible competition with other nocturnal raptors, including T. furcata and Pulsatrix arredondoi Brodkorb, 1969, deserves further study. Fossils of T. maniola, new species, were found again years later by WS during excavations and study of fossil birds in asphalt deposits at Las Breas de San Felipe, Matanzas Province, western Cuba.

Fred

Holotype proximal left tarsometatarsus (MNHNCu 75.4651) of Tyto maniola, new species (Cuba), in anterior (A), posterior (B), superior (C), lateral (D) and medial (E) views. Scale = 1 cm.