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 los 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 orclenes Strigiformes y Based on discoveries of giant fossil raptorial birds Accipitriformes, de que dichos taxones consti- (Strigiformes and Accipitriformes) in Cuba and tuyeron el elemento faum'stico primoridal que con- other Antillean islands, the theory is put forth trolara con su accion predatora a roedores y des- here that these predators were the principal agents clentados 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 bubo 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. espedficas de todas estas grandes aves predatoras, incluyendose, distribucion geografica, tablas de medidas y una relacion de la fauna extinguida hallada asociada en las localidades tipicas. 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 Espeleologico, Martel 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 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY 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 Camagiiey, 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 rodents 1 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 Pfo 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 Rfo. 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 1) 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¬ 1 The mammalian classification used in this paper follows posed in 1958, established that the species belonged Varona (1974) and in some instances is at variance with in the family Strigidae, where he maintained it in Profess Aorredondo p’sreferences.—Ed. later publications (Brodkorb, 1969, 1971). Addi- NUMB E2R7 171 Table 1.—Associated fauna found in the type-localities of the large extinct species of Cuban birds of prey Species 172 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY 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. ostologa, and the other, Tyto gigantea, cies of Ornimegalonyx may exist among these re¬ being only, slightly smaller than T. riveroi, partly mains; one of these seems to be somewhat smaller contradicts the hypothesis that gigantism in Antil¬ than O. oteroi, while the other is larger. Consider¬ lean barn owls is attributable to insular evolution ing the enormous size of this owl, particularly of from smaller species that responded to the great its claws, it is quite conceivable that it could have abundance of food and the lack of competition made victims of juvenile edentates, notwithstand¬ from carnivorous mammals. The genus Tyto ing the fact that Ornimegalonyx appears to have evidently had already evolved giant species in Eu¬ been incapable of flight. rope, millions of years before the beginning of the In July 1954, explorations in the Cueva de Pleistocene. The following conclusions could there¬ Paredones in San Antonio de los Banos, revealed fore be drawn: (1) either the giant Antillean barn for the first time the fossil bones of an eagle larger owls evolved in parallel with those of Europe, ar¬ than any of the living species of the family Accipi- riving through convergence at species of approxi¬ tridae. Other bones of it were found a few years mately the same size, or (2) the Antillean forms later in the Cueva del Tunel, La Salud, Habana. are descended from Tertiary European forms that This eagle was named Aquila borrasi (Arredondo, established themselves in North America and 1970). According to its tarsometatarsus, it was very colonized the Antilles before or during the Pleisto¬ similar in size and morphology to the recently de¬ cene. Against this last suggestion is the absence on scribed species Garganoaetus freudenthali from the the American continent of giant species of Tyto. upper Miocene of Italy (Ballmann, 1973). Ornimegalonyx is truly exceptional for its ex¬ A fossil vulture from Cueva de Paredones was tremely large size. It appears to have evolved its recently described as a new genus and species, gigantism in Cuba from some remote smaller ances¬ Antillovultur varonai Arredondo (1971), and was tor. An affinity of Ornimegalonyx with any of the the size of an Andean Condor. Various bones found living genera of large continental owls is not in a cave in Habana and now under study, possibly clearly evident and its relationships may lie closer indicate another large species of vulture. to some extinct form rather than with any presently Two species of giant barn owls of the genus Tyto living. were discovered a little later (Arredondo, 1972a, 1972b). One of these, Tyto noeli, founded on The study of the origins, evolution, and paleo- abundant bones from two caves in Habana, was ecology of the giant raptorial birds of the Antilles similar in size to Tyto ostologa of Haiti. The other is of great interest and significance to our under¬ species, Tyto riveroi, based on the distal portion of standing of the environment and evolution of a tarsometatarsus from Cueva de Bellamar, Matan- many of the terrestrial vertebrates of those islands. zas, was truly gigantic, being larger than any of the It is hoped that this summary of what is known of fossil or living species of the genus. It is the one the Cuban birds will aid in that understanding. strigiform that most closely approaches the size of Abbreviations used are as follows: Academia de Ornimegalonyx oteroi, and like that species could Ciencias de Cuba (ACC), Departamento de Pale- also have captured small edentates. The eminent ontologia de la Universidad de la Habana paleornithologist Alexander Wetmore (1959) was (DPUH), Museo del Grupo de Exploraciones the first to report fossil remains of large barn owls from Cuba, but these were not named. 2 Cientificas “Pedro Borras Astorga’’ (GEC), Mu¬ seum of Comparative Zoology (MCZ), Museo " These specimens, a. humerus and a femur, are still at the Felipe Poey de la Academia de Ciencias de Cuba National Museum of Natural History. They are very much (MFP), Museo Montand de la Universidad de la larger than T. ostologa or T. noeli and most probably per¬ tain to the distinctive species T. riveroi. At a later date I Habana (MMEJH), personal collection of Oscar hope to be able to describe these fossils further, along with Arredondo (OA), Sociedad Espeleologica de Cuba abundan tunpublished materia lo fT .ostologa .—Ed. (SEC). NUMB E2R7 173 Order ACCIPITRIFORMES Family VULTURDDAE Genus Antillovultur Arredondo, 1971 Antillovultur varonai Arredondo, 1971 Holotype. —Proximal portion of left tarsometa- tarsus, DPUH 1254. Type-Locality.— Cueva de Paredones, San An¬ tonio de los Banos, Habana, Cuba. Age. —Late Pleistocene. Other Material.— GEC (unnumbered), distal portion of left humerus; OA 847, external trochlea of left tarsometatarsus; OA 848, body of seventh cervical vertebra. All specimens from the type- locality. Description. —The type (Figure 1 a,b) is a proxi¬ mal portion of a tarsometatarsus, 42.5 mm in length, lacking the proximal articulating surface, hypotarsus, and slightly more than half the distal portion of the bone. The estimated total length is 141 mm, or about equal to that of Vultur gryphus and longer and slightly more robust than in Gymnogyps californianus (Figure 2, Table 2). From these two species and Cathartes aura it differs in having the groove in the anterior face of the bone narrower and deeper and the internal tuber¬ cle for M. tibialis anticus more expanded. In ante¬ rior view, the surface of the shaft between the internal border and the groove is notably thick and rounded, whereas in Vultur, Cathartes, and Teratornis it is narrower and sharp-edged. The surface of the shaft delimited by the external bor¬ der and the groove is more slender than the internal ridge, contrary to the condition in the other genera mentioned. The shaft in medial view is proportionately more slender than in Vultur or Cathartes. These characters are considered to be of generic value. The distance between the proximal border of the larger tubercle for M. tibialis anticus and the proximal extremity of the anterior groove is greater than in Vultur or Teratornis and equal to that in Gymnogyps. Antillovultur has an addi¬ tional two proximal foraminae situated above the Figure 1.—Specimens of Antillovultur varonai, Cueva deParedones: a, holotype fragmentary proximal end of left usual two. Vultur similarly possesses another aper¬ tarsometatarsus (DPUH 1254) ,anterior view; b ,same ,pos¬ ture above the lateral proximal foramen but lacks terior view; c, left humerus lacking proximal end (GEC the medial proximal foramen of Antillovultur. unnumbered) p, alma rview (.Natura sl ize.) 174 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY Table 2 .—Measurements (mm) of the tarsometatarsus of Antillovultur varona icompared with other large New World vultures Character E*stimated. Cathartes aura has only two proximal foramina. A seventh cervical vertebra attributed to Antil¬ Teratornis differs in having three great united lovultur consists of the complete body lacking all of foramina arranged so as to form a kind of circle. the processes. It is similar to vertebrae of Vultur In Antillovultur the medial tubercle for M. tibi¬ in size and morphology. alis anticus is better developed than in either A distal portion of a left humerus (Figure 1c) Vultur, Gymnogyps, or Cathartes, and is situated is from a specimen very similar in size to Vultur. over a slight protruberance on the internal border This fragment has a length of 184 mm from the of the anterior groove. In posterior view the sur¬ distal end to a point a little beyond the protruber¬ face of the bone from the lateral foramen to the ance at the distal extremity of the deltoid crest. external border is flat and does not slope downward Taking this protruberance as a point of reference, as in Vultur. On the opposite side, the surface from the complete length of the bone can be estimated the medial foramen to the external border is very as 265 mm, which is about 20 mm less than in the depressed compared to that in Vultur. An external specimens of Vultur compared. The least width of trochlea from a left tarsometatarsus of this species the shaft is 20 mm (21.7 in Vultur ); the maximum is similar in size and shape to that of Vultur. distal width is 45.8 mm (54.3 in Vultur). The bone, although almost as large as that of Vultur, is more slender. The distal protruberance of the deltoid crest is at the same level as in Vultur. Antillovultur differs from Vultur in having the ectepicondylar prominence less pronounced, re¬ calling that of Cathartes. The internal and exter¬ nal condyles are slightly smaller than in Vultur but with the distal borders more prominent. The in¬ ternal condyle has a smooth and extensive depres¬ sion on the entepicondylar side, which is present in Cathartes but absent in Vultur. The attachment of the anterior articular ligament is as large as in Vultur, but situated closer to the internal condyle, the space between them being less than in Vultur and more similar to Cathartes. The foramen located in this space is isolated from the brachial depres¬ sion by a ridge that is absent in Vultur but some¬ abed what evident in Cathartes. The brachial depression Figure 2 .—Comparison of tarsometatars iof New World vul¬ is deeper and more pronounced than in Vultur, tures: a, Gymnogyps californianus; b, Vultur gryphus; c, particularly in the proximal region, but is notably Antillovultur varonai ;d ,Teratornis merriam i(based on Wet- less expanded than in either Vultur or Cathartes. more ,1931) .(Hal fnatura lsize.) The olecranal fossa is similar to that of Vultur and NUMB E2R7 175 less dilated than in Cathartes. The tricipital grooves larities to B. contortus in the general structure of are similar to those of Vultur but the external one the tarsometatarsus, it is larger and more robust is somewhat deeper and appears like that of than that species and appears more like A. chrysa¬ Cathartes. etos. According to the published figures, the re¬ cently described species Garganoaetus freudenthali Family ACCIPITRIDAE Ballmann (1973), from the upper Miocene ofItaly, is similar morphologically to B. borrasi, al¬ though its tarsometatarsus is slightly more robust. Genus Aquila Brisson Aquila borrasi was a gigantic form within its genus, the only other known fossil forms of which are Aquila borrasi Arredondo, 1970 Aquila delphinensis and A. pennatoides, described Holotype. —Left tarsometatarsus lacking troch- by Gaillard (1939) from tarsometatarsi from the leae, DPUH 1250. upper Miocene of France. Type-Locality.— Cueva del Tunel, La Salud, The femur of Aquila borrasi is larger and more Habana, Cuba. robust than that of any living eagle. Although the Other Localities.— Cueva de Paredones, San one known specimen is incomplete, its maximum Antonio de los Banos, Habana. Cueva de Pio length is estimated at about 155 mm, as opposed to Domingo, Sumidero, Pinar del Rio. 125 mm in Aquila chrysaetos, 114 mm in Haliae¬ Age.— Late Pleistocene. etus leucocephalus, and 96 mm in Spizaetus Other Material.— Right femur lacking con¬ ornatus. This is even larger than in the two im¬ dyles, SEC P-26; ungual phalanges, SEC P-31, mense living eagles Harpia harpyja (131 mm) P-32, P-35, P-1147, and ACC 1000a; subterminal phalanx, ACC 1000b; distal end of tarsometatarsus, SEC P-40. Description.— Tarsometatarsus (Figure 3) gen¬ erally similar to that of Aquila chrysaetos but notably longer, since even without the distal end it measures 97.7 mm. The estimated total length of the bone is 130 mm, or 34 mm longer than that of Aquila chrysaetos and larger than that of any liv¬ ing species of eagle (Table 3). The proximal artic¬ ular region is similar to that in Aquila but with the proximal foramina located only 5 mm from the internal cotyla, whereas in A. chrysaetos this dis¬ tance is nearly twice as great. That which remains of the base of the hypotarsus in the type indicates that this process was probably similar to that of A. chrysaetos. The tarsometatarsi of the fossil species Buteo typhoius and B. contortus, from the upper Miocene of Nebraska, and of B. conterminus, from the up¬ per Pliocene of Nebraska (Wetmore, 1923), are larger than those of Recent species of Aquila and Haliaeetus, but are more slender proximally, the articular region in proximal view being of a dif¬ ferent shape than in Aquila and also differing in the form and position of the middle trochlea. The tarsometatarsus of B. contortus measures 113 mm Figure 3. —Holotype left tarsometatarsus of Aquila borrasi in length, which is 17 mm less than in Aquila bor¬ (DPUH 1250), Cueva del Tunel. (Anterior view at natural rasi. Although the Cuban species has certain simi¬ size.) 176 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY Table 3 .—Measurements (mm) of the tarsometatarsus and claws of Aquila borrasi compared with other species of living eagles Character * Measurements from specimens in the USNM collections. It can be seen that the tarsometatarsus of Aquila borrasi, while decidedly longer ,is proportionately much more gracile than in either Pithecophaga or Harpia —Ed. and Pithecophaga jefferyi (130 mm). 3 Other meas¬ The ungual phalanges are very well developed urements (in mm) of this specimen are as fol¬ (Figure 4), being almost two times larger than lows: total length as preserved, 140, proximal those of A. chrysaetos. They resemble those of width 45.4, vertical diameter of head 16.3, width of Harpia harpyja in having the same degree of cur¬ neck 17.2, maximum width of pneumatic foramen vature. The ungual phalanx of digit IV is larger, 8.0, length of pneumatic foramen 14.0, least width while that of digit I is smaller than in Harpia of shaft 19.8. (Table 3). The shape of the articular region and The femur differs from that of allied genera by the ventral process of the first ungual phalanx of the lesser projection of the trochanter above the A. borrasi more closely resembles that of Aquila head and by the greater width between the anterior than Haliaeetus, Spizaetus, or Buteo. The ungual border of the head and the apex of the trochanter phalanges of digits II and IV are likewise similar (twice that of Aquila chrysaetos or Haliaeetus leu- to those of Aquila and differ from those of the cocephalus). The pneumatic opening is roughly other genera examined. The ungual phalanx of oval in shape but wider distally. It is located at the digit IV measures 33 mm through the ventral arc base of the trochanter as in Aquila but differs in and 35 mm through the dorsal arc. being not perfectly oval and in being oriented semi- obi iquely towards the external border of the tro¬ chanter. The foramina of the upper region of the Order STRIGIFORMES trochanter are larger and deeper than in the other species compared. The rugose intermuscular line Family STRIGIDAE on the anterior face of the bone angles below and near the pneumatic opening along the external Genus Ornimegalonyx Arredondo border of the shaft almost to its midpoint. In con¬ trast, this line in A. chrysaetos originates farther Ornimegalonyx oteroi Arredondo, 1958a above the upper border of the pneumatic opening Synonym.— Ornimegalonyx arredondoi Arre¬ and descends straight to the midpoint of the shaft. The head is massive and the neck is thick and dondo, 1958.Lectotype. —Left tarsometatarsus SEC P-383E. oriented slightly upwards. Lectotype designated by Brodkorb (1961); depos¬ 3 Measurements supplied from specimens in the collections ited in the Museum of Comparative Zoology, of the National Museum of Natural History, Smithsonian Harvard University. Institution.—Ed. Type-Locality. —Caverna de Pio Domingo, Si- NUMB E2R7 177 lonyx have come to light so far: the lectotype from Pio Domingo; one from Cueva de Quinto; two proximal halves from Paredones; a distal half from Cueva del Tunel and a proximal half from the same locality. Ultimately, two complete specimens from the same individual were found in a cave in the Sierra de Cubitas, Camaguey, along with other elements. The dimensions of these bones indicate the enor¬ mous size that this bird had in relation to all other known Strigiformes, living or fossil (Figures 5-9, Tables 4-6). The tarsometatarsus is almost double the length of that of Bubo bubo, or more than double if one considers the specimen from Cueva de Quinto (GEC unnumbered) (Figure 7 a) or that from Paredones. These bones are more than three times the size of the corresponding element of Nyctea scandiacci, more than four times that of Asio otus, and eight and a half times the size of Glaucidium siju. Figure 4. —Phalanges of Aquila borras ifrom Cueva de Pare- In spite of its gigantic size, the tarsometatarsus of dones: a, ungual phalanx of digit I (SEC P-31); b, ungual Ornimegalonyx is proportionately less robust than phalanx of digit II (SEC P-32); c, subterminal phalanx (SEC that of Bubo, Nyctea, or Pulsatrix, the difference P-35 )(.Natura slize.) being due to the relative lengthening of the shaft in Ornimegalonyx. If the tarsometatarsus of Orni¬ megalonyx were reduced to the length of that of Asio otus, the two elements would be seen to be very similar in proportion, whereas if one magni¬ fied the tarsometatarsi of Bubo, Nyctea, or Pulsa¬ trix to the size of that of Ornimegalonyx, they would appear much stronger, wider, and more robust. In Ornimegalonyx the internal trochlea is pro¬ portionately shorter and wider than in Bubo, Nyctea, or Asio, being more similar to Pulsatrix. The middle trochlea is narrow and placed very close to the outer trochlea. The distal foramen is somewhat lower than in Bubo, Nyctea, or Asio, and the ossified bridge on the anteroproximal re¬ gion of the bone recalls that of Bubo and differs from Nyctea in that it is stronger and more circu¬ lar. The internal cotyla is similar to that in Bubo and Asio, but somewhat lower than in Nyctea. The wide, deep groove on the posterior face of the bone is more pronounced than in Bubo, Nyctea, or Asio. The tibiotarsi of the type individual are frac¬ tured into proximal and distal portions and shafts (Figure 5, Table 5). In a complete state they would have measured some 250 mm in length. The tibio- 178 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY Table 4. —Measurements (mm) of the tarsometatarsus of Ornimegalonyx oteroi compared with other owls E*stimated. tarsus from Cueva de Quinto has a length of 272 of Ornimegalonyx are less robust than in Bubo or mm, which is almost twice that of Bubo bubo or Nyctea, while in the femora the opposite occurs. Nyctea scandiaca. Its proximal width of 40 mm is Fragments of the sternum, as well as parts of the likewise twice that of those species. The tibiotarsus scapulae, ribs, vertebrae and carpometacarpi, were of the lectotype individual appears to have been associated with the type individual in Pio Do¬ straight, while that from Cueva de Quinto is mingo cave. The most important sternal fragments slightly curved and subtly twisted. Compared to are an anterior portion with the articulations for the modern genera examined, the tibiotarsus of the coracoids, and another fragment of the left Ornimegalonyx (Figure 6a) has the fibular articu¬ side containing four costal facets (the fifth having lation more pronounced, the rotular crest, and the been fractured off). The costal facets vary slightly cnemial crest more elevated, and the fossa proximal in size, the largest being 6X6 mm. Through these to the condyles on the anterior face deeper. fragments it has been possible to reconstruct the Eight femora of Ornimegalonyx have been sternum (Figure 8) as being wide, almost flat (both found so far. The left femur of the type individual dorsally and ventrally), with a vestigial keel, which is in the MCZ. Of the right, only a part of the indicates that the bird was hardly able to fly. Its proximal end remains in Cuba. The largest femur, estimated length is 120 mm (vs. 47 mm in Tyto represented by the proximal end only (Figure Id), alba), the estimated width 75 mm (30 mm in T. was found to the east of Sancti Spiritus and is de¬ alba)-, and the height at the keel some 30 mm posited in the Museo Montane of the University (25 mm in T. alba). The similarity of this last of Havana (MMUH 5072). Two incomplete fem¬ measurement in two species which otherwise differ ora from Paredones (formerly SEC P-37 and SEC so greatly in size is a further indication of the great P-38, but now in the MCZ) are smaller than those extent of the atrophy of the keel in Ornimegalonyx. of the type and for now are perhaps best referred The carpometacarpus of Ornimegalonyx is to as Ornimegalonyx sp. small in proportion to the enormous size of the The femur of Ornimegalonyx differs principally body. Its total length is estimated at about 90 mm from that of other owls in being much larger and whereas in Bubo bubo, a smaller volant species, it notably more robust (Table 6). That of the type is 85 mm. individual is almost one and a half times larger than the femur of Bubo bubo, while the large femur (MMUH 3072) from Sancti Spiritus is al¬ Ficur e5 —. Tibiotarsus t,arsometatarsus f,emur a, n dphalanges(SEC P-383E) from the same individual as the lectotype of most double the size of that of Nyctea scandiaca. Ornimegalonyx oteroi ,Caverna de Pio Domingo .The speci¬ It is curious that when reduced, the tarsometatarsi mens are covered with calcareous concretions .(Natura lsize.) NUMB E2R7 179 180 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY As with the other bones, the phalanges of Orni- digit I measures 40 mm. The proximal height of megalonyx stand apart from those of other owls by these phalanges ranges from 15-17 mm. their great size. The length of the ungual pha¬ It was not until the beginning of 1969 that frag¬ langes of digits II and III, measured through the ments of mandibles were found among the remains dorsal arc, range from 37 to 39 mm, while that of of the type individual of Ornimegalonyx oteroi. These consist of the two articular portions of both rami. These fragments permit for the first time a very approximate estimate of the size of the man¬ dible and ultimately of the whole skull (Figure 9a). These mandibles are very similar in overall morphology to those of the diminutive genus Glau- cidiam. They differ from Bubo bubo in that the internal angular process is greatly lengthened. The posterior angular process, in comparison to that of Bubo, is notably more robust, and the portions that remain of the surangular and angular are greatly thickened at the point of the fracture. Judging by comparison with recent owls, the length of the entire mandible of Ornimegalonyx oteroi would have been some 115 mm and the distance be¬ tween the external borders of the articulations was approximately 100 mm, or about two times larger than in Bubo bubo. A portion of a cranium of Ornimegalonyx was f Figure 7.—Some variation in the hindlimb o fOrnimegalonyx: a a, Ornimegalonyx cf. oteroi, left tarsometatarsus (GEC un¬ numbered), Cueva del Qtiinto; b, c, Ornimegalonyx oteroi, Figure 6 .—Hindlimb elements o fOrnimegalonyx otero ifrom left tarsometatarsus and right femur of the type individual Cueva de Paredones: a, right tibiotarsus lacking distal end (SEC. P-38E), Pio Domingo cave; d, Ornimegalonyx sp., (SEC P-28); b, posterior view of left tarsometatarsus lacking proximal end of left femur (MMUH 3072), Camera de los distal end (SEC P-39). (Natural size.) Hornos de Cal ,Sanct iSpiritus. NUMB E2R7 181 Tab 5le.—Measurements E*stimated. Table 6 .—Measurements (mm) of the femur of Ornimegalonyx otero icompared with other owls Es*timated. Figure 8.—Ventral and lateral views of the sternum of Ornimegalonyx oteroi, as reconstructed from two fragments from Pio Domingo and Paredones caves .(Natura lsize.) •• NUMB E2R7 183 found in Cueva de Paredones in 1959 (Figure larger than the males. Nevertheless, other bones 9b,c). For the time being it is best referred to only that are either larger or smaller than those of the as Ornimegalonyx sp., for it is apparent that it type individual and have distinct differences from does not correspond to the species O. oteroi. it, probably indicate additional species—the cran¬ Rather, it appears to belong to a smaller form, as ium and femora from Paredones and the femur also evidenced by the femora found in the same from Sancti Spiritus being examples. locality. The specimen consists of the posterior Ornimegalonyx had to have been the scourge portion of a cranium from the postorbital arc to and terror of most of the larger mammals of the the occiput and including the basisphenoid, the Pleistocene of Cuba and the claws and mandibles foramen magnum, and the occipital condyle. of this bird would have constituted a terrible com¬ Viewed from the front, the great thickness of the bination of superior destructive power. walls of the cranium are seen in the region of the break. Compared to Bubo bubo it is larger, and in Family TYTONIDAE ventral view it has the postorbital process better developed. The opisthotic process is rather promi¬ Genus Tyto Billberg nent and its extremity bends notably, hanging in the form of an ear. The basipterygoid processes are well developed. The foramen magnum is some¬ Tyto noeli Arredondo, 1972a what higher than wide, as opposed to Glaucidium Holotype.— Right tarsometatarsus, DPUH 1251. and Tyto in which it is wider than high. As in Type-Locality. —Cueva del Tunel, La Salud, Nyctea, the occipital condyle is very well developed Habana, Cuba. in relation to the foramen magnum, whereas it is Other Localities. —Cueva de Paredones, San proportionately much smaller in Glaucidium and Antonio de los Banos, Habana; Cueva del Indio, Tyto. Reparto El Globo, Calabazar, Habana; quarries From the actual and estimated measurements of near Sancti Spiritus, Las Villas. the various bones of Ornimegalonyx it can be es¬ Age. —Late Pleistocene. tablished that this great owl stood some 1100 mm Other Material.— Cueva del Tunel: OA 818, high in life. Although the general aspect of its skel¬ right femur; OA 812, distal portion of left tibio- eton is similar to that of living owls, it is distin¬ tarsus; OA 804, distal portion of right humerus; guished from them by the long and robust hind- OA 806, proximal fragment of right humerus; OA limbs, provided with long, heavy toes armed with 822, shaft of right tibiotarsus; OA 815, distal por¬ the most powerful claws possessed by any strigiform tion of right ulna. Cueva de Paredones: OA 828, bird. Although the sternum is larger than in any proximal portion of right tarsometatarsus; OA 827, living owl, it is actually small in proportion to the proximal portion of right tibiotarsus; OA 839, rest of the bones of the skeleton. Its semiflat struc¬ right coracoid. Cueva del Indio: OA 1027, right ture and reduced keel show that Ornimegalonyx femur. was little or not at all capable of flight. In accord¬ Description.— Similar to the living species Tyto ance with this, the bones of the wing are poorly alba in its general skeletal configuration, but much developed, particularly the carpometacarpus. larger (Figures 10, 11, Tables 7 and 8), equaling in Some of the differences in morphology and size size the extinct species Tyto ostologa of Haiti and that are observed between individuals of Orni¬ T. pollens of the Bahamas. The tarsometatarsus megalonyx are probably attributable to sexual di¬ was between 90 and 100 mm long and was similar morphism, since in other owls the females are to that of T. pollens, but more slender, even in specimens that are longer than in T. pollens. The Figure 9. —Skulls of Ornimegalonyx: a ,hypothetica lrecon¬ tibiotarsus is likewise similar to that of T pollens struction of the skull of Ornimegalonyx oteroi to show the but is less robust. This slenderness of the hindlimb size as extrapolated from the two mandibular articulations is the most notable difference between the two associated with the type; b, c, anterior and ventral views of the cranium (GEC unnumbered) of Ornimegalonyx sp .from species. Cueva de Paredones (the left quadrate is incorrectly articu¬ In the femur, humerus, ulna, coracoid, and lated). (All figures natural size.) claws, the only pronounced difference from Tyto 184 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY Figure 10.—Paratypes of Tyto noeli from Cueva del Tunel and Cueva de Paradones compared with a Recent specimen of Tyto alba furcata (on the left in each pair): a, fragments of right humerus (OA 804 and 826); b, fragments of right ulna (OA 806 and 815); c, fragments of tibiotarsus (OA 827, 812, and 822); d, right femur (OA 818); e, right coracoid (OA 839). (Natural size.) alba is in size. Likewise, a fragment of the anterior may possibly be differentiated from T ostologa portion of a sternum of T. noeli from quarries near only at the subspecific level. The same could be Sancti Spiritus has the same conformation as that suggested for T. noeli. From the upper Miocene of of T. alba but is larger. One might expect to find Italy, a new species of giant barn owl, Tyto robusta greater distinctions in the skull, but so far only Ballmann (1973), has been described that is near fragments of the skull of T. noeli have been found. the size of T. noeli. The Cuban species is somewhat Brodkorb (1959:357) suggested that T. pollens larger and heavier, however. NUMBE 2R7 185 Table 7. —Measurements of limb bones of Tyto noeli compared with Tyto alba Character E*stimated. Table 8. —Measurements (mm) of the tarsometatarsi of the three Cuban species of Tyto Character E*stimated. Tyto riveroi Arredondo, 1972b aspect of that of Tyto alba and still more similar to that of T. noeli. Except for size, significant morph¬ Holotype. —Distal portion of a left tarsoraeta- tarsus, DPUH 1252. ological differences from the above species are al¬ Type-Locality. —Cueva de Bellamar, Mantanzas, most absent; however, the measurements of the Cuba. type of T. riveroi notably exceed the limits of Age.— Late Pleistocene. either (Figure 11, Table 8). The estimated total Description. —Tarsometatarsus with the general length of this bone would be approximately 125 186 SMITHSONIA NCONTRIBUTION TS OPALEOBIOLOGY mm. The following slight morphological distinc¬ Tyto gigantea, recently described from the upper tions are also noted: greater separation of the in¬ Miocene of Gargano, Italy (Ballmann, 1973), was ternal and external trochleae from the middle an enormous barn owl, equal in size to T. riveroi. trochlea, the intertrochlear spaces being narrower According to the published figures, the Italian in T alba and T noeli; shaft proportionately species has the distal foramen somewhat more ele¬ wider and thicker. Compared with T. pollens, the vated and the middle trochlea lower and more same slight differences are apparent. elongate than in the Antillean species. Figure 11.—Comparison of the tarsometatarsi of the three Cuban species of Tyto: a, Recent Tyto alba furcata; b, Tyto noeli, holotype (DPUH 1251), Cueva del Tunel; c-e, Tyto riveroi, holotype (DPUH 1252) ,Cueva de Bellamar ,anterior ,posterior ,and latera lviews .(Natura lsize.) Literature Cited Acevedo ,M. ,O. Arredondo ,and N. Gonzdlez 1971 .Nuevo genero y especie de ave fdsi l(Accipitriformes: 1975. La Cueva clel Tunel. Volume 1, 74 pages, 17 fig¬ Vulturidae) del Pleistoceno de Cuba. Memoria de ures, 2 maps, 6 tables. Havana: Editorial Pueblo y la Sociedad de Ciencias Naturales La Salle ,31(90): Educacidn .Instituto de lLibro. 309-323 5f,igures. Arredond Oo,. 1958a. Aves gigantcs de nucstro pasado prehistdrico. El 1972a .Nueva especie de ave fdsi l(Strigiformes :Tytonidae) Cartero Cubano 1, 7(7) :10-12 6, figures. del Pleistoceno superior de Cuba. Boletin de la 1958b. E lVampiro Cubano. Scout (Havana) ,pages 6-7 ,10, Socieda dVenezolan ad eCiencia sNaturales 2, 9(122— f6igures. 123):415-43 1f5,igures. 1970. Nueva especie de ave pleistocdnica del orden Ac- 1972b .Especie nueva de lechuza (Strigiformes :Tytonidae) cipitriformes (Accipitridae) y nucvo genero para del Pleistoceno cubano. Boletin de la Sociedad las Antillas. Ciencias, series 4, Ciencias Bioldgicas, Venezolana de Ciencias Naturales, 30(124-125): 8:1-19 1, 0figures. 129-140 4f,igures. NUMBE 2R7 187 Arredondo, O., and L. S. Varona XXXII, Cultural. 1974. Nuevos g^nero y especie de mamffero (Carnivora: Koopman ,K .F. Canidae) del Cuaternario de Cuba. Poeyana, 131: 1958. A Fossil Vampire Bat from Cuba. Breviora, 90: 1-12 ,3 figures. 1-4 ,1 plate. Ballman nP,. Varona ,Luis S. 1973. Fossile Vogel aus dem Neogen der Halbinsel Gar- 1974. Catalogo de los mamiferos vivientes y extinguidos gano (Italien) .Scripta Geologica ,17:1-75 ,16 figures, de las Antillas. 139 pages. Havana: Academia de p7lates. Ciencias de Cuba. Brodkor bP,. Wetmor eA,. 1959. Pleistocene Birds from New Providence Island, 1922. Remains of Birds from Caves in the Republic of Bahamas. Bulletin of the Florida State Museum, Haiti. Smithsonian Miscellaneous Collections, 74 Biologic Saclience 4s(,11):349—371. (41) :1-4 ,2 figures. 1961. Recently Described Birds and Mammals from 1923. Avian Fossils from the Miocene and Pliocene of Cuban Caves .Journa lof Paleontology ,35(3):633— Nebraska. Bulletin of the American Museum of 635. Natura lHistory ,48:483-507 ,20 figures. 1969. An Extinct Pleistocene Owl from Cuba. Quarterly 1931. The Avifauna of the Pleistocene in Florida. Smith¬ Journal of the Florida Academy of Sciences, 31(2): sonian Miscellaneous Collections, 85(2): 1-41, 16 112-114 1 ,figure. figure s6p, lates. 1971. Catalogue of Fossil Birds, Part 4 (Columbiformes 1937. Bird Remains from Cave Deposits on Great Exuma through Piciformes). Bulletin of the Florida State Island in the Bahamas. Bulletin of the Museum of Museum B, iologica Sl ciences 1, 5(4) 1: 63-266. Comparative Zoology ,80(12):427-441 ,16 figures ,1 Gaillard C, . plate. 1939 .Contribution a 1’^tude des oiseaux fossiles .Archives 1959. Birds of the Pleistocene in North America. Smith¬ du Museum d’Histoire Naturelle de Lyon ,15(mem- sonia nMiscellaneou sCollections 1, 38(4 )1: -24. oire 2) :1-100 ,34 figures. Wotoszyn, B. W., and N. A. Mayo Harrington ,M .R. 1974. Postglacial Remains of a Vampire Bat (Chiroptera: 1935. Cuba antes de Colon. 290 pages. 111 figures, 108 Desmodus )from Cuba A. ct aZoologic aCracoviensia, plates. Havana: Coleccidn de Libros Cubanos, 19(13):253-265 ,5 figures ,1 plate. Arredondo, Oscar. and Olson, Storrs L. 1976. "The Great Predatory Birds of the Pleistocene of Cuba." Collected papers in avian paleontology honoring the 90th birthday of Alexander Wetmore 27, 169–187. View This Item Online: https://www.biodiversitylibrary.org/item/267468 Permalink: https://www.biodiversitylibrary.org/partpdf/352137 Holding Institution Smithsonian Libraries Sponsored by Smithsonian Institution Copyright & Reuse Copyright Status: In copyright. Digitized with the permission of the rights holder. 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