■JT Reprinted from: "Collected Papers in Avian Paleontology Honoring the 90th li *1 Birthday of Alexander Wetmore," Storrs L. Olson, editor, 21 May 1976. !# The Great Predatory Birds of the Pleistocene of Cuba Oscar Arredondo translated and amended by Storrs L. Olson ABSTRACT Recent paleontological investigations in Cuba Se expone la tesis, basada en Ios ultimos descu- have shown that the island was formerly inhabited brimientos realizados en Cuba y otras Antillas sobre by large populations of rodents and edentates. aves fosiles gigantes de los or denes Strigiformes y Based on discoveries of giant fossil raptorial birds Accipitriformes, de que dichos taxones consti- (Strigi formes and Accipitriformes) in Cuba and tuyeron el elemento faunistico primoridal que con- other Antillean islands, the theory is put forth trolara con su action predatora a roedores y des- here that these predators were the principal agents dentados que en estas islas existian, manteniendo in controlling the abundance of these native mam- asi el equilibrio biologico necesario. Se ofrecen mals. Details are given of the discovery in Cuba of datos concretos probatorios de la extremada abun- the gigantic owl Ornimegalonyx, two species of dancia en las Antillas de poblaciones de roedores y giant barn owls (Tyto), a giant species of eagle desdentados. Se dan detalles del descubrimiento en (Aquila borrasi), and a vulture (Antillovultur) Cuba del gigantesco buho Ornimegalonyx, de dos similar in size to the Andean Condor (Vultur especies gigantes de lechuzas del genero Tyto, de gryphus). The relationships and possible origins of una especie de aguila gigante y de un buitre simi- gigantism in these birds are discussed. Descriptions, lar en talla a Vultur gryphus. Se plantea una dis- geographic distribution, tables of measurements, cusion sobre el motivo que pudo originar el and a list of the extinct faunas found in each of the gigantismo en estas aves. Se ofrecen descripciones type-localities are offered for each species. especificas de todas estas grandes aves predatoras, incluyendose, distribution geografica, tablas de medidas y una relation de la fauna extinguida hallada asociada en las localidades tfpicas. Introduction thousands of mandibles of Capromys pleistocenicus can be extracted from a single small chamber, al- Recent studies of fossil material from numerous though most of these specimens are poorly pre- caves in Cuba have convincingly demonstrated the served. An examination of only a part of the fossil former extraordinary abundance of endemic mam- material taken from a small cave in Cayo Salinas, mals there. These include rodents of the genera a few miles east-southeast of Caibarien, Las Villas, Heteropsomys, Capromys (including Geocapro- yielded the remains of over 200 individuals of the mys), and Macrocapromys; ground sloths (Eden- edentate genus Mesocnus (Acevedo, Arredondo, tata) of the genera Cubanocnus, Miocnus, and Gonzalez, 1975). Further confirming the for- Mesocnus, and Megalocnus; and insectivores of the mer abundance of native mammals is the wide- genera Solenodon and Nesophontes, These genera spread occurrence in Cuba of remains of blood- were represented throughout the island by species eating vampire bats of the genus Desmodus very numerous in terms of individuals. From such (Koopman, 1958; Arredondo, 1958b; Woloszyn caves as Paredones and El Tunel in the province of and Mayo, 1974). These bats would necessarily Habana it is no exaggeration to say that tens of have required numerous, large, warm-blooded Oscar Arredondo, Grupo Espeleoldgico, Mattel de Cuba, mammals to sustain them. Similar abundant faunas Avenida 43, No. 5847, Apartado 4, Mariano 14, Habana, of large' rodents and ground sloths are also known Cuba. from cave deposits in Hispaniola and Puerto Rico. 169 170 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY Obviously, some natural regulator must have in cave deposits in Haiti was attributed to the acted to maintain a biological equilibrium between depredations of the extinct giant barn owl Tyto these prolific herbivorous mammals and the vegeta- ostologa, first described by Wetmore (1922). tion on which they fed. In the Antilles, however, Another giant barn owl, Tyto pollens Wetmore there are virtually no native carnivorous mammals. (1937), first reported from Great Exuma and later Exceptions are Cubacyon transversidens (Arre- from New Providence Island (Brodkorb, 1959), dondo and Varona, 1974), a canid known from a must have been an effective predator of Capromys single fragment of maxilla found in association with in the Bahamas, as no doubt were the extinct extinct vertebrates in a cave in the province of diurnal raptors (Accipitridae) of the genera Calo- Habana, and the extant populations of Procyon hierax and Titanohierax (Wetmore, 1937). The known from the Bahamas, Barbados, and Guade- presence of such large raptors elsewhere in the loupe, some or all of which may have been intro- Antilles suggested that the Cuban cave deposits duced to these islands by man (Varona, 1974; might be attributable to similar avian predators. Arredondo and Varona, 1974). A mandible found This was vividly confirmed by the discovery in in an Indian midden in Camaguey, Cuba, and at- Cuba of remains of five species of predatory birds tributed to Procyon lotor by the naturalist Andres of truly tremendous proportions. These birds Poey (Harrington, 1935), may similarly have occupied the niches which in continental areas are pertained to an introduced animal; the record is usually filled by various groups of carnivorous dubious in any case, since the specimen can no mammals. The absence of mammalian competitors, longer be found. Regardless, small procyonids combined with a superabundance of large prey, could not have been an influential factor in the are no doubt the principal factors contributing to control of the endemic Antillean rodents and the gigantism observed in these birds. The larger edentates. forms may have functioned particularly to keep The numerous and dense deposits of smaller the populations of edentates in check. vertebrates (Table 1) frequently found in Cuban caves abound with remains of the rodents1 Capro- mys pleistocenicus, Capromys nanus, Heteropso- Brief History of Certain Discoveries mys torrei, and H. offella; the insectivores Neso- On 2 January 1954, the remains of a gigantic phontes micrus and N. major; several genera of owl, the largest known, were discovered for the first bats; passeriform birds, particularly of the genera time in the depths of a large cavern known as Pio Mimocichla, Quiscalus and Dives; as well as non- Domingo Cave, located in the Sierra de Sumidero, passerines such as Crotophaga and Glaucidium. opposite Pica-Pica Valley in Pinar del Rio. These These remains are certainly attributable to the bones (Figure 5) were found in place, fastened to actions of medium-sized owls still existing in the the calcareous surface of the floor by travertine, island such as Tyto alba and Asio stygius, and per- just as were those of the various edentates en- haps to others now extinct, such as Pulsatrix countered in the immediate vicinity, This owl was arredondoi (Brodkorb, 1969). described under the name Ornimegalonyx oteroi The most outstanding accumulations of bones, (Arredondo, 1958a) and was at first erroneously however, are those of the larger mammals (Table regarded as a member of the Phorusrhacidae, an I) such as Capromys pilorides, C. columbianus, extinct family of flightless South American carniv- Macrocapromys acevedo, and occasional examples orous birds. Dr. Byran Patterson, who studied some of Cubanocnus gliriformis, as well as large birds of of these remains, afterwards informed me (pers. various orders. Such remains suggest that many comm.) of his belief that they actually pertained to larger predators were once active in Cuba. a great owl, two times larger in linear dimensions Much of the accumulation of vertebrate remains than Tyto ostologa of Haiti. Brodkorb (1961), recognizing the validity of the nomenclature pro- posed in 1958, established that the species belonged 'The mammalian classification used in this paper follows Varona (1974) and in some instances is at variance with in the family Strigidae, where he maintained it in Professor Arredondo's preferences.—Ed. later publications (Brodkorb, 1969, 1971). Addi- NUMBER 27 171 TABLE I.—Associated fauna found in the type-localities of the large extinct species of Cuban birds of prey Ornimegalonyx Tyto Tyto noeli and Antillovultur oteroi riveroi Aquila borrasi varonai Species Caverna de Pio Cueva de Domingo, Bellamar, Cueva del Tiinel, Cueva de Pinar del Rio Matanzas Habana Paredones, Habana AMPHIBIA Bufo sp X REPTILIA Iguanidae gen. and sp. indet. Epicrates cf. angulifer X X Geochelone cubensis X X X Crocodylus sp X X AVES Cathartes aura Antillovultur varonai X Vulturidae gen, and sp. indet. X Aquila borrasi ? X Grus cubensis ? Ornimegalonyx oteroi X X Ornimegalonyx sp X Pulsatrix arredondoi X Gymnoglaux sp X Tyto alba X Tyto noeli X Tyto riveroi MAMMALIA Solenodon cf. cubanus .... X X X Nesophontes micrus X X X Nesophontes major X Phyllonycteris poeyi X Natalus lepidus X Cubacyon tranversidens ... X Acratocnus sp X Miocnus antillensis X X X Cubanocnus gliriformis ■■ X X X X Mesocnus torrei X X X X Mesocnus browni X X Neomesocnus brevirostris X Megalocnus rodens X X X Megalocnus sp X Capromys pilorides X X X Capromys prehensilis X Capromys nanus X Capromys columbianus .... X X X Capromys pleistocenicus X X X Capromys sp. A X Capromys sp. B X Capromys sp, C X X Capromys sp. D X Macrocapromys acevedo ■■ X Heteropsomys torrei X X X Heteropsomys offella X X X X = present; ? = uncertain; not present. 172 SMITHSONIAN CONTRIBUTIONS TO PALEOBIOLOGY tional fossils of this giant owl were later found in The discovery of three species of tytonids in the various caves in the provinces of Habana, Matan- upper Miocene of Italy (Ballmann, 1973), one of zas, Las Villas, and Camaguey, as well as in the Isle which, Tyto robusta, is equal in size to T. noeli of Pines. It is possible that two unrecognized spe- and T.