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The fossil avifauna of the tar seeps Las Breas de San Felipe, Matanzas, Cuba (1 Viewer)

Melanie

Well-known member
Germany
The fossil avifauna of the tar seeps Las Breas de San Felipe, Matanzas, Cuba
WILLIAM SUÁREZ

Abstract

The Cuban fossil avifauna, prior to this study, included 30 extinct and extirpated valid taxa. In the present contribution, we review the fossil avifauna from Las Breas de San Felipe, and in so doing increase the diversity of Cuban fossil birds to 36 species with the description of a New World vulture, Coragyps seductus sp. nov., three accipitrids, Gigantohierax itchei sp. nov., Buteogallus royi sp. nov., and Buteo sanfelipensis sp. nov., a small caracara, Milvago diazfrancoi sp. nov., plus Buteogallus cf. B. fragilis (L. Miller, 1911), which is recorded for the first time in Cuba and the Antillean Subregion. Of the total of 34 bird species now registered from Las Breas de San Felipe, 21 (61.8 %) are extinct and 13 (38.2 %) correspond to species that still live in Cuba, other Antillean islands, or in the American continent. Raptors dominate the assemblage, with 26 (76.5 %) species. These are mostly from Accipitridae and Falconidae, and 19 (55.9 %) are diurnal and 7 (20.6 %) nocturnal. This abundance of raptors in Las Breas de San Felipe is similar to the composition from other asphalt deposits known from the American continent such as Rancho La Brea. The palaeoavifauna from this locality can be split, according to ecological preferences, into three groups or guilds. Radiocarbon (14C) dates indicate a late Pleistocene age for some of the recovered bird remains, including those of Antigone cubensis (Fischer & Stephan, 1971) comb. nov., Gymnogyps varonai (Arredondo, 1971) and Ornimegalonyx oteroi (Arredondo, 1958).

https://mapress.com/j/zt/article/view/zootaxa.4780.1.1
 
Systematic Palaeontology - 1

Finaly I have the paper and my conputer is working again so I can give you the Systematic Paleontology.

All species are from the Pleistocene.

Systematic Palaeontology

Class AVES Linnaeus
Order ANSERIFORMES Wagler
Family ANATIDAE Leach
Genus Dendrocygna Swainson
Dendrocygna arborea (Linnaeus, 1758)
Referred material. San Felipe I: Distal half of right tibiotarsus, MNHNCu 75.4743; left tarsometatarsus with fragmentary, distal end, MNHNCu 75.4744.
Description. These specimens are inseparable from the corresponding elements in the skeleton of the living species Dendrocygna arborea Linnaeus, 1758.
Comments. First record of the West Indian Whistling-Duck in Cuban fossil deposits. This species is currently a common permanent resident in certain places of Cuba, Isla de la Juventud, and some cays, where it frequents marshes, mangroves, lagoons and rice paddies

Order GRUIFORMES Bonaparte
Family GRUIDAE Vigors
Genus Antigone Reichenbach, 1853
Antigone cubensis (Fischer & Stephan, 1971) comb. nov.
Referred material. San Felipe I: Sternal half of right coracoid, MNHNCu 75.4760; distal half of left ulna, MNHNCu 75.4761; proximal fragment of right femur, MNHNCu 75.4762; right tibiotarsus without proximal end, MNHNCu 75.4765; distal half of right tibiotarsus, MNHNCu 75.4767; distal half of left tibiotarsus, MNHNCu 75.4763; distal third of left tibiotarsus, MNHNCu 75.4766; proximal ends of right tarsometatarsi, MNHNCu 75.4772-4773; proximal ends of left tarsometatarsi, MNHNCu 75.4774-4775; distal ends of left tarsometatarsi, MNHNCu 75.4764, 75.4768-4769. San Felipe II: Distal ends of right tarsometatarsi, MNHNCu 75.4770-4771.
Description. Fossils here referred to Antigone cubensis (Fischer & Stephan, 1971) comb. nov., do not show significant differences in size or qualitative characters when compared with specimens of the type series of that species.
Comments. This taxon was previously registered for asphalt deposits by Iturralde-Vinent et al. (2000). The osteology of this crane is well known since its original description (Fischer 1968; Fischer & Stephan 1971). The abundant material recovered in the type locality—Cueva de Pío Domingo, Sumidero, Minas de Matahambre Municipality, Pinar del Río Province—was confused and apparently identified at an early stage (e.g., Castellanos 1968) as belonging to the genus Ciconia Brisson, in the family Ciconiidae (Gray). It has been suggested that the Cuban species derives from an extinct lineage of large cranes (Olson & Rasmussen 2001:291) known in deposits of North America since the Pliocene (Yorktown Formation), including Florida (Emslie 1995, 1998), and related to the living Sarus Crane, Antigone antigone (Linnaeus, 1758). In congruence, I relocate here the Cuban Flightless Crane in the genus Antigone, represented in the Cuban archipelago today (Garrido & Kirckconell 2011) by the endemic subspecies of the Sandhill Crane, A. canadensis nesiotes

Order CHARADRIIFORMES Huxley
Family BURHINIDAE Mathews
Genus Burhinus Illiger
Burhinus bistriatus (Wagler, 1829)
Referred material. San Felipe I: Right humerus without distal end, MNHNCu 75.4783. San Felipe II: Distal end of right humerus, MNHNCu 75.4792; distal end of right tibiotarsus, MNHNCu 75.4798.
Description. These fossils are similar in characters to specimens referred to Burhinus bistriatus (Wagler, 1829), from other fossil localities in Cuba, and to the neontological material of that species with which it has been compared.
Comments. Burhinidae was previously cited for Las Breas de San Felipe, as Burhinus sp., by Iturralde-Vinent et al. (2000). The present material, although scarce, allows refinement of the identification to the specific level. Burhinus bistriatus is also known, and is abundant, in other Cuban deposits (Arredondo 1984; Suárez 2000b; Suárez unpubl. data). A fossil subspecies from the Bahamas, B. b. nanus Brodkorb, 1959b, seems to be also present in Cuba (Suárez unpubl. data), being smaller than the living one in Hispaniola (Brodkorb 1959b; Olson & Hilgartner 1982), B. b. dominicensis (Cory, 1883), or another known from some Cuban deposits (Suárez & Olson unpubl. data). To determine the subspecific status of all these fossils from Cuba, it is necessary to expand the existing series. Oswald & Steadman (2018) consider the mentioned Bahamian taxon as a valid species.
 
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Systematic Palaeontology - 2

Order CICONIIFORMES (Bonaparte)
Family CICONIIDAE (Gray)
Genus Ciconia Brisson
Ciconia sp.
Referred material. San Felipe I: Distal end of right tibiotarsus, MNHNCu 75.4599.
Description. This specimen differs, according to Suárez & Olson (2003a:151), from the same element in Mycteria americana Linnaeus, 1758, by a wide intercondylar groove (Fig. 4A; very narrow in the living species). The only known specimen of Ciconia sp., has a distal width of 15.4; smaller than in the tibiotarsus of C. maltha L. Miller,
1910 (18.0–21.5, N = 25 [Howard 1942]), and C. maguari (Gmelin, 1789) (17.1–19.6, N = 5 [Suárez & Olson 2003a: table 1]).
Comments. Known in Cuba only from asphalt deposits, where the fragment in discussion seems to represent a new species of stork, not yet described (Suárez & Olson 2003a), smaller than Ciconia maltha, and similar in size to the White Stork, C. ciconia (Linnaeus, 1758). The only other material in Cuba referable to this genus is a proximal third of tarsometatarsus and a distal end of tibiotarsus, from the thermal baths of Ciego Montero, Palmira Municipality,Cienfuegos Province, originally identified as Jabiru mycteria Lichtenstein, 1819, by Wetmore (1928:2), and reassigned to C. maltha by Howard (1942). This last species has not been registered in Cuba since that date. Agnolín (2009) considered that C. maltha is a posterior synonym of C. lydekkeri (Ameghino, 1891).

Genus Mycteria Linnaeus
Mycteria wetmorei Howard, 1935
Referred material. San Felipe I: distal end of right tibiotarsus, MNHNCu 75.4603; left tarsometatarsus, MNHNCu 75.4757; proximal end of left tarsometatarsus (juvenile), MNHNCu 75.4604; distal end of right tarsometatarsus, MNHNCu 75.4605. San Felipe II: Proximal end of right carpometacarpus, MNHNCu 75.4602.
Description. These fossils were referred to Mycteria (see Suárez & Olson 2003a:151–152) by the diagnostic characters of the genus present in M. americana Linnaeus, 1758. They coincide with the Wood Stork in most of the qualitative characters, but are larger than in the mentioned species (Suárez & Olson 2003a: table 1).
Comments. Mycteria wetmorei Howard, 1935, as in the case of the other species of the Leptoptilini and Mycteriini tribes mentioned, have previously been recorded in paleontological deposits of Cuba only from tar seeps (Suárez & Olson 2003a). This extinct species differs from M. americana by its larger size, and by presenting a less arched mandible (Howard 1935; Olson 1991). The Cuban specimens represent the southernmost record of this palaeospecies and the only one outside the continental mainland (Suárez & Olson 2003a). The left tarsometatarsus MNHNCu 75.4757, reported herein, expands the Cuban series for this taxon and constitutes the most complete skeletal element known to date of this bird in the archipelago.

Mycteria americana Linnaeus, 1758
Referred material. San Felipe II: Scapular portion of left coracoid, MNHNCu 75.4600; proximal end of right carpometacarpus, MNHNCu 75.4601.
Description. This material (see Suárez & Olson 2003a: fig. 1) agrees with the morphology observed in the species of the genus Mycteria by having a coracoid with medial surface of the acrocoracoid flattened, rather than inflated; a carpometacarpus with long proximal symphysis, internal carpal trochlea less externally bent, shallow anterior carpal fossa, and metacarpal I very excavated on both sides. It agrees in characters and size (see Suárez & Olson 2003a,151, table 1) with M. americana and differs from M. wetmorei for being slightly smaller (see preceding species).
Comments. The Wood Stork is currently considered a rare permanent resident in coastal and mangrove lagoons of Cuba, Isla de la Juventud, and major cays of the north coast (Garrido & Kirkconnell 2011: 52). It has been erroneously recorded from different deposits in North America, confused with the fossil species Mycteria wetmorei (see Olson 1991). Fossil remains of M. americana are known in the northern hemisphere only from Las Breas de San Felipe, and given the characteristics of the Cuban material, this is the only reliable record for this species in this part of the world

Order INSERTAE SEDIS
Family †TERATORNITHIDAE L. Miller
Genus †Oscaravis Suárez & Olson
Oscaravis olsoni (Arredondo & Arredondo, 2002b)
Referred material. San Felipe I: Right cuneiform, MNHNCu 75.4663; distal half of fragmentary left tibiotarsus, MNHNCu 75.4659; distal end of right tibiotarsus without anterior portions of the external condyle, MNHNCu
75.4660; distal end of right tibiotarsus without condyles, MNHNCu 75.4858; distal end of right tarsometatarsus, MNHNCu 75.4662. San Felipe II: Distal end of fragmentary left femur, MNHNCu 75.4857; distal end of left tarsometatarsus, MNHNCu 75.4661.
Description. Cuneiform resembles Teratornis merriami L. Miller, 1909, in general morphology although it is massive, especially in its ventral ramus, with greater ulnar joint and dorsal ramus connected to the body of the bone at a more open angle than in T. merriami or Aiolornis incredibilis Campbell, Scott & Springer, 1999 (see also Howard 1952). For description of the femur, an element partially known in the deposits under study, see Arredondo & Arredondo (2002b), and Suárez & Olson (2009a). Tibiotarsus (Fig. 5A–C) with a broad and expanded shaft at its junction with the condyles, broad tendinal groove and foramen, reduced internal condyle, wide intercondylar fossa (proximad). Tarsometatarsus (Fig. 5D–F) with short trochlea metatarsi III, proportionally long trochleae metatarsorum II and IV; relatively wide intertrochlear spaces. For comparisons, measurements, and a more complete description of this material, see Suárez & Olson (2009a).
Comments. The extinct family Teratornithidae was recorded outside continental America as Teratornis sp., by Suárez & Arredondo (1997). Following this record, T. olsoni Arredondo & Arredondo, 2002b, was described from Cuba, based on part of the fossils then known. New material (including specimens treated here), and a first revision of the Cuban Teratorn allowed erection of the genus Oscaravis (Suárez & Olson 2009a). Oscaravis olsoni has a larger size compared with Taubatornis campbelli Olson & Alvarenga, 2002, but is smaller than the remaining taxa described in different genera for the family (Teratornis L. Miller 1909, Cathartornis L. Miller 1910, Argentavis Campbell & Tonni 1980, Aiolornis Campbell et al. 1999; see also Campbell & Stenger 2002). This teratorn provides evidence of the ability of some members of Teratornithidae for overwater dispersal, not dependent on a continuous land bridge for expansion to North America (Olson & Alvarenga 2002; Suárez & Olson 2009a). Orihuela (2019:53) erroneously stated that in Cuba was “un teratornítido con adaptaciones que indican capacidades nulas o limitadas de vuelo” [a teratornitid with adaptations that indicate no or limited flight capabilities]. In the anatomy of O. olsoni such adaptations do not exist (see Arredondo & Arredondo 2002b; Suárez & Olson 2009a).

Figure 5. Oscaravis olsoni: Left tibiotarsus (MNHNCu 75.4659) in anterior (A), distal (B), and medial (C) views; left tarsometatarsus
(MNHNCu 75.4661) in anterior (D) and posterior (E) views; right tarsometatarsus (MNHNCu 75.4662) in distal view (F). Scale = 2 cm.
 

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Systematic Palaeontology - 3

Order CATHARTIFORMES Coues
Family CATHARTIDAE Lafresnaye
Genus Coragyps Le Maout
Coragyps seductus sp. nov.
Holotype. Left tarsometatarsus, MNHNCu 75.4719. Collected in San Felipe II, by William Suárez and Stephen Díaz Franco, on February 24, 2001.
Measurements (mm) of holotype. Total length: 86.0; proximal width: 16.4; least width of the shaft at midpoint: 7.4; distal width: 16.1; depth of trochlea metatarsi II: 7.9+; width and depth of trochlea metatarsi III: 6.7–9.8; depth of trochlea metatarsi IV: 9.1 (see Table 4).
Diagnosis. Similar to Coragyps occidentalis (L. Miller, 1909) in robustness, but having tarsometatarsus slender, with noticeable bilateral compression at distal end and trochlea metatarsi III long, thin, and not deep.
Etymology. From Latin seductus, remote, secluded, in reference to the allopatric condition of the Cuban species, given its insularity.
Paratypes (topotypes). San Felipe II: Proximal half of left femur without trochanter, MNHNCu 75.4718;
proximal end of left tarsometatarsus, MNHNCu 75.4720.
Description. The specimens herein described are referred to Coragyps and split from the remaining genera of Cathartidae from Cuba, Cathartes Illiger and Gymnogyps Lesson (Suárez 2000a, 2001; Suárez & Emslie 2003), by having a tarsometatarsus with an elongated and relatively less robust shaft, distal end flaring gradually from the shaft, with trochleae compressed bilaterally. Other characters present in the tarsometatarsus that segregate it from Cathartes include those described by Carr (1981:35). Femur and tarsometatarsus (Fig. 6A–F) similar in robustness to those of the fossil species Coragyps occidentalis (Table 4), and slightly more robust than in the examined series of C. atratus Bechstein, 1793. The femur MNHNCu 75.4718, is fragmentary and coincident in morphology and general characters with both Coragyps species. The fragmentary condition of that specimen does not allow other
comparisons. Tarsometatarsus with slender shaft of moderate robustness (shorter and more robust in C. occidentalis; slender and gracile in C. atratus), distal end with noticeable bilateral compression (proximal end consistently thinner than distal end in C. occidentalis and C. atratus); trochlea metatarsi III long, thin, and not too deep (deeper in C. occidentalis; shorter, wider, and deeper in C. atratus); trochleae metatarsorum II and IV, relatively small, the former well projected posteriad (similar in C. occidentalis; relatively larger and not projected in C. atratus).
Comments. Fossils of Coragyps seductus sp. nov. are extremely rare in Cuba, and unknown in other Quaternary deposits in the Cuban archipelago. Tarsometatarsi of C. seductus sp. nov. do not agree in their characters with the equivalent element in skeletons of C. atratus examined (N =18) from North and South America, nor with series of this element in C. occidentalis (N = 42) from asphalt deposits of Rancho La Brea (see Howard 1968). The material described here represents the first record of this genus in fossil deposits of Cuba and the Greater Antilles. A distal fragment of a carpometacarpus (WS s/n), collected by the author in a Quaternary cave deposit at Cueva de Sandoval, Caimito Municipality, Artemisa Province (formerly La Habana Province), before the material described herein, may represent this new taxon, but it lacks sufficient diagnostic characters for a positive identification. As in the case of the Antillean caracaras (see Suárez & Olson 2001b: 507), members of Cathartidae in Cuba during the Pleistocene were represented by endemic species, all extinct today (Suárez 2001; Suárez & Olson unpubl. data). Coragyps has been registered from multiple fossiliferous localities in Florida (Brodkorb 1964), where it has been common since the late Pliocene (Emslie 1998:28). A similar case holds for the genus Gymnogyps Lesson and the endemic Cuban Condor (Suárez & Emslie 2003:36).

Figure 6. Coragyps seductus sp. nov.: Left tarsometatarsus (Holotype, MNHNCu 75.4719) in anterior (A), posterior (B) and distal (C) views; left tarsometatarsus (Paratype, MNHNCu 75.4720) in proximal (D) view; left femur (Paratype, MNHNCu 75.4718) in anterior (E) and posterior (F) views. Scale = 2 cm.
 

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Systematic Palaeontology - 4

Genus Cathartes Illiger
Cathartes sp.
Referred material. San Felipe I: Right tibiotarsus without proximal end, MNHNCu 75.4750; distal ends of right tibiotarsi, MNHNCu 75.4749, MNHNCu 75.4754; distal end of left tibiotarsus, MNHNCu 75.4748; proximal half of right tarsometatarsus, MNHNCu 75.4752; proximal ends of right tarsometatarsi, MNHNCu 75.4745, 75.4753; distal half of right tarsometatarsus, MNHNCu 75.4746; proximal half of left tarsometatarsus, MNHNCu 75.4747; distal half of left tarsometatarsus, MNHNCu 75.4751. San Felipe II: Incomplete left coracoid, MNHNCu 75.4755.
Comments. The material here referred to Cathartes sp., together with some specimens from cave deposits, represents a new species which will be compared, described, and named in a separate paper (Suárez & Olson in prep.). This constitutes the first record of this taxon for Las Breas de San Felipe. Fossils of this small vulture are also known from cave deposits located in the western part of the Cuban archipelago (Suárez 2000a, 2001; Suárez & Olson in prep.)

Genus Gymnogyps Lesson
Gymnogyps varonai (Arredondo, 1971)
Referred material. San Felipe I: Right coracoid, MNHNCu 75.4596; left humerus without proximal end, MNHNCu 75.4821; distal half of right tibiotarsus, MNHNCu 75.4823; right tarsometatarsus without distal end, MNHNCu 75.4822; distal end of left tarsometatarsus, MNHNCu 75.4598; left cuneiform, MNHNCu 75.4664. San Felipe II: Upper mandible (premaxillary-maxillary-maxillopalatine), MNHNCu 75.4613; premaxillary fragments, MNHNCu 75.4594-4595; distal end of left radius, MNHNCu 75.4776; proximal right carpometacarpus, MNHNCu 75.4597.
Description. It differs (see descriptions and figures of this material in Suárez 2000a, Suárez & Emslie 2003) from other species of the genus by having a skull with strong and massive bill, robust occipital and opisthotic processes, large occipital condyle, a more rostral placement of nuchal crest, and a high cranial vault. These osteological modifications indicate a more developed, or powerful, cervical musculature than in continental species. Postcranial elements (Fig. 7A–J) are slightly larger, but similar in characters to the living species Gymnogyps californianus (Shaw, 1797).
Measurements. Coracoids.—total length: 98.9; least width at midpoint: 17.0; depth at midpoint of glenoid facet: 21.8. Humerus.—width and depth of shaft at midpoint: 19.0–15.4; distal width: 45.7. Carpometacarpus.— proximal width of metacarpal II: 12.0. Tibiotarsus.—width and depth of shaft: 14.4–11.1; distal width: 25.5; depth of external condyle: 22.8; depth of internal condyle: 23.4. Tarsometatarsus.—Proximal width: 28.1; depth of medial cotyla: 13.7; depth of lateral cotyla: 11.7; least width and depth of shaft: 14.7–8.5; distal width: 33.1; width of trochlea metatarsi III: 11.6.
Comments. This taxon is well known from multiple deposits in Cuba, and the first bird species identified in Las Breas de San Felipe (Suárez 2000a). The Cuban Condor was described as an endemic, monotypic genus and species, Antillovultur varonai Arredondo, 1971, on the basis of few fragmentary bones from Quaternary deposits at Cueva de Paredones, La Habana (now Artemisa) Province (Arredondo 1971). The species was subsequently referred to Gymnogyps (see Suárez 2000a) and redescribed (Suárez & Emslie 2003). Gymnogyps varonai was the largest vulturid in the Cuban archipelago during the Quaternary, where remains are very common in fossil localities (Suárez 2000a, 2001). This condor evolved in Cuba from a population of G. californianus or G. kofordi Emslie, 1998, from Florida and its presence in Cuba demonstrates the ability of condors to disperse over water (Tambussi & Noriega 1999).

Order ACCIPITRIFORMES Vieillot
Family ACCIPITRIDAE Vieillot
Genus Accipiter Brisson
Accipiter striatus Vieillot, 1808
Referred material. San Felipe I: Tarsometatarsus without proximal end, MNHNCu 75.4756.
Description. This specimen is identical to the equivalent element in A. striatus Vieillot, 1808, and coincides in its smaller size and delicateness with the Cuban subspecies A. s. fringilloides Vigors, 1827. Tarsometatarsi examined in skeletons of the continental subspecies A. s. velox Wilson, 1812, are larger (see Wetmore 1937: 428).
Comments. The Sharp-shinned Hawk is today a rare permanent resident in Cuban forests (Garrido & Kirkconnell 2011: 77). It has been previously registered from cave deposits in western Cuba (Suárez & Arredondo 1997). MNHNCu 75.4756 was originally identified as Accipiter sp. by Iturralde-Vinent et al. (2000: table 2).
 
Systematic Palaeontology - 5

Genus Buteogallus Lesson
Buteogallus borrasi (Arredondo, 1970)
Referred material. San Felipe I: Distal end of right tibiotarsus, MNHNCu 75.4678; distal ends of left tibiotarsi, MNHNCu 75.4665, MNHNCu 75.4693; left tarsometatarsus without trochleae, MNHNCu 75.4666; left tarsometatarsus lacking proximal articular region and trochleae metatarsorum II and III, MNHNCu 75.4667; proximal half of left tarsometatarsus without inner calcaneal ridge of the hypotarsus, MNHNCu 75.4686; proximal end of left tarsometatarsus without calcaneal ridges of the hypotarsus, MNHNCu 75.4692; distal halves of left tarsometatarsi, MNHNCu 75.4687-4688; fragmentary distal end of left tarsometatarsus, MNHNCu 75.4691; proximal end of right tarsometatarsus without inner calcaneal ridge of the hypotarsus, MNHNCu 75.4677; proximal segment of shaft of a right tarsometatarsus, MNHNCu 75.4669; distal end of right tarsometatarsus, MNHNCu 75.4670; distal ends of right tarsometatarsi without trochlea metatarsi III, MNHNCu 75.4689-4690; distal shaft of right tarsometatarsus, MNHNCu 75.4668; left metatarsal I, MNHNCu 75.4694; right metatarsal I, MNHNCu 75.4682; phalanx 1, left digit I, MNHNCu 75.4697; phalanx 1 without distal end, left digit I, MNHNCu 75.4671; phalanges 1, right digits I, MNHNCu 75.4695-4696; ungual phalanges, left digits I, MNHNCu 75.4672, 75.4684-4685; ungual phalanx, right digit I, MNHNCu 75.4683; ungual phalanges, left digits II, MNHNCu 75.4679-4680; ungual phalanges, right digits II, MNHNCu 75.4673-4676; ungual left, MNHNCu 75.4681, and right, MNHNCu 75.4707, phalanges of digit III.
Description. General morphology as in Buteogallus urubitinga (Gmelin, 1788), but about a 33% larger. For a more detailed description see Suárez (2004a), and Suárez & Olson (2007, 2009b).
Measurements. See Suárez & Olson (2007: tables 1–4); Table 6.
Comments. Buteogallus borrasi (Arredondo, 1970) was recorded for this locality by Iturralde-Vinent et al. (2000), and the material under study was described by Suárez & Olson (2007, 2009b). This is the most common accipitrid found in Quaternary deposits along the Cuban archipelago (Suárez 2004a:124; Suárez & Olson 2007: 296), and also in Las Breas de San Felipe. Wetmore (1928: 3–4, figs. 1–2), recorded Geranoaetus melanoleucus (Vieillot, 1819) for Cuba, on the basis of an incomplete left carpometacarpus (AMNH 6190)—plus an ungual phalanx without catalog number—from the thermal baths of Ciego Montero, Palmira Municipality, Cienfuegos Province, noting that it “is exactly similar to Geranoaëtus [sic.] melanoleucus and is identified as that species”. Given the carpometacarpus of B. borrasi is about the size of (G. melanoleucus in square brackets: Proximal width, 8.3–8.4 (8.3) 2 [7.3–8.5 (7.7) 8]; proximal depth, 21.1–22.1 (21.6) 2 [17.8–21.8 (20.1) 8]; depth of metacarpal II at midpoint, 5.7 [4.9–5.6 (5.1) 4].) and very similar in characters to the same element in G. melanoleucus (Suárez unpubl. data), it seems that AMNH 6190 represents the very common Borras’ Hawk, instead of the species recorded in 1928. Unfortunately, at the present stage, the specimens in AMNH have not been located.

Buteogallus cf. †B. fragilis (L. Miller, 1911)
Referred material. San Felipe II: Distal end of right tibiotarsus, MNHNCu 75.4735; distal segment of shaft of left tarsometatarsus, MNHNCu 75.4736.
Description. Smaller when compared with Buteogallus borrasi or “Amplibuteo” woodwardi. Specimen MNHNCu 75.4735 differs from the tibiotarsus of Geranoaetus melanoleucus, and agrees with those of B. fragilis (L. Miller, 1911), in its slightly smaller size, flattened supratendinal bridge (not inflated or domed) with more horizontal orientation, less deep and wider tendinal groove, and medial condyle much more projected mediad. This specimen is identical in small details and size to tibiotarsus RLB E4091, within the series of this bone examined (N = 33). The distal fragment of tarsometatarsus MNHNCu 75.4736, differs from the equivalent element in G. melanoleucus, by having the metatarsal facet in a lower (or distad) position and being slightly smaller, also as in B. fragilis. This specimen is very similar in size and characters to RLB D3604, within the series of tarsometatarsi examined (N = 70) of the latter species. For comparisons between B. fragilis and B. borrasi, see Suárez & Olson (2007: 293).
Measurements. (RLB specimens in parenthesis). Tibiotarsus.—distal width: 15.3+ (15.3–17.8 [16.7] 27). Tarsometatarsus.— least width of shaft: 7.9 (6.4–8.4 [7.3] 33).
Comments. Comparisons with other accipitrids (see Appendix 1) and with extensive series of B. fragilis from Rancho La Brea, California, show that the two Cuban specimens are referable to this taxon. This record is the first for Cuba and the Antilles, as well as the first occurrence of the species outside the continental mainland. This raptor is probably another member of the Cuban avifauna derived from populations established in Florida, as the taxon is considered to have been present in that peninsula since the late Pliocene (Emslie 1998). Another extinct hawk, close in general character to those present in B. fragilis—see Howard (1932) for descriptions, comparisons, and discussion of characters that place “Geranoaetus” fragilis under Urubitinga (= Buteogallus)—but much smaller and not referable to the species B. gundlachii (Cabanis, 1855), B. anthracinus (Deppe, 1830), or any of the other member of the genus, is described below.

Buteogallus royi sp. nov.
Holotype. Left tarsometatarsus, MNHNCu 75.4909. Collected in San Felipe I, C area, on May 12, 2009, by William Suárez and Stephen Díaz Franco.
Measurements (mm) of holotype. Total length: 92.9; proximal width: 13.6+; proximal depth between calcaneal ridges of hypotarsus: 5.8; proximal width at level of tubercle for tibialis anticus: 10.5; least width and depth of shaft at midpoint: 6.7–8.1; least width and depth of shaft at proximal end of metatarsal facet: 7.4–5.5; distal width: 16.1+; width and depth of trochlea metatarsi III: 4.5–6.2; width of trochlea metatarsi II: 3.4; width of trochlea metatarsi IV: 3.3 (see Table 5).
Referred material. San Felipe II: Distal third of right ulna, MNHNCu 75.4737.
Etymology. Species dedicated to the memory of Dr. Roy E. Dickerson, discoverer of Las Breas de San Felipe, and the first to report fossil bird remains in Cuban tar seeps.
Diagnosis. Species similar in size to the living Buteogallus anthracinus and B. gundlachii, but with a tarsometatarsus longer and more robust, with deep and wide anterior metatarsal groove and tubercle for tibialis anticus located more proximad.
Description. Buteogallus royi sp. nov. differs from the extinct hawks B. fragilis, B. daguetti (L. Miller, 1911), B. terrestris (Campbell, 1979) and B. borrasi, as well as the living species B. solitarius (von Tschudi, 1844), B. coronatus (Vieillot, 1817), B. meridionalis (Latham, 1790) and B. urubitinga, due to its smaller size (Table 5). The tarsometatarsus present in B. aequinoctialis (Gmelin, 1788) is smaller. MNHNCu 75.4909 is comparable in size only to those of B. anthracinus and B. gundlachii (for size correlation of these living taxa see Olson 2006), but differs from these species by having the following combination of characters (Fig. 9–10A–C): shaft elongated and columnar (elongated but more gracile in B. anthracinus; consistently shorter and more gracile in B. gundlachii), reduced proximal end that is flaring bilaterally from a more proximal point on shaft (expanded, flaring gradually from a more distal point on shaft in B. anthracinus and B. gundlachii), broad and deep fossa infracotylaris dorsalis (reduced and less excavated in B. anthracinus and B. gundlachii), deep anterior metatarsal groove, very excavated and wide (less excavated and narrower in B. anthracinus and B. gundlachii), tubercle for tibialis anticus located proximad and laterad (distad and medially located in B. anthracinus and B. gundlachii), poorly developed internal metatarsal border (more developed and less flattened in B. anthracinus and B. gundlachii), trochlea metatarsi II more projected mediodistad (less projected in B. gundlachii; moderately projected in B. anthracinus), inner calcaneal ridge of the hypotarsus with base short, only slightly extended distally, and truncated by the medial proximal foramen (similar in some specimens of B. anthracinus; usually more distally extended in B. gundlachii), deep posterior metatarsal groove (less deep in B. anthracinus and B. gundlachii), metatarsal facet less expanded mediad (more medially expanded in B. anthracinus and B. gundlachii), and well developed crista plantaris lateralis (less developed in B. anthracinus and B. gundlachii).
Comments. The Roy’s Hawk, as the other extinct Buteogallus species present in this deposit, was a larger bird when compare with B. gundlachii, the only living species of the genus in Cuba (Wiley & Garrido 2005), which is common in mangroves, coastal swamps of the main island, Isla de la Juventud, and keys (Garrido & Kirkconnell 2011:79). The tarsometatarsus of B. royi sp. nov. is very different from that of B. anthracinus and B. gundlachii, being more similar to B. fragilis in some of its characters (see comparisons and description above; Fig. 9C). The latter species (and the new Cuban taxon described herein), is not referable to genera Buteo Lacépède, Rupornis Kaup, Parabuteo Ridgway, Geranoaetus, or Spizaetus Vieillot either, being similar in most of the characters to Buteogallus, in which it has been accommodated by Howard (1932), and where I maintain both species. The distal fragment of ulna (Fig. 10D–E), tentatively referred to B. royi sp. nov., is also similar in size and general characters to both B. anthracinus and B. gundlachii.

Figure 10. Buteogallus royi sp. nov.: Left tarsometatarsus (Holotype, MNHNCu 75.4909) in anterior (A), medial (B) and posterior (C) views; right ulna (MNHNCu 75.4737) in dorsal (D) and ventral (E) views. Buteo sanfelipensis sp. nov.: Left tarsometatarsus (Holotype, MNHNCu 75.4910) in anterior (F), medial (G) and posterior (H) views. Scale = 2 cm.
 

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Systematic Palaeontology - 6

Genus †Gigantohierax Arredondo & Arredondo
Type species.Gigantohierax suarezi Arredondo & Arredondo, 2002a
Emended diagnosis. Accipitrid with relatively small humerus; carpometacarpus short and robust; femur pneumatic, short and wide, well compressed anteroposteriorly; supratendinal bridge of the tibiotarsus with some horizontal orientation; tarsometatarsus columnar, with wide and robust shaft, proximal foramina and distal foramen large, tubercle for tibialis anticus located well proximad and placed in a deep anterior metatarsal groove, lateral border rounded (or convex) along its extension, especially at the proximal region, trochleae forming a poorly pronounced arc in distal view; phalanx I, digit I, heavy, large and wide; ungual phalanges only slightly arched.
Gigantohierax suarezi Arredondo & Arredondo, 2002a
Referred material. San Felipe I: Proximal halves of right, MNHNCu 75.4721, and left, MNHNCu 75.4722, humeri; fragmentary right carpometacarpus, MNHNCu 75.4724; left carpometacarpus without metacarpal III, MNHNCu 75.4723; right tarsometatarsus without proximal end and part of the medial border, MNHNCu 75.4728; distal half of left tarsometatarsus, MNHNCu 75.4729; distal end of left tarsometatarsus without trochlea metatarsi IV, MNHNCu 75.4730; ungual phalanges of left digit I, MNHNCu 75.4731, MNHNCu 75.4777; right, MNHNCu 75.4733, and left, MNHNCu 75.4732, metatarsal I.
Emended diagnosis. The largest species in the genus Gigantohierax.
Description. The specimens of Gigantohierax suarezi described here do not differ in size or qualitative characters from those recovered in other Cuban deposits. This material differs from all fossil and living accipitrids known in America by its huge size (Arredondo & Arredondo 2002a; Table 6; Suárez et al. in prep.), being similar in this respect to the Haast’s Eagle from New Zealand (see Holdaway 1990). It differs from other American taxa of Accipitridae examined (see Appendix 1) by having humerus (Fig. A–B) small in relation with the remaining elements of the skeleton, with shaft well arched. In palmar view the bicipital furrow is very shallow and expanded, but a small deep area exists distad to the head. Ligamental furrow inexistent, being indicated by a very small and round scar, located well proximad. Bicipital surface rounded (or convex), reduced, and bicipital crest angulated distally. Deltopectoral crest projected nearly perpendicular to the surface of the shaft. In anconal view, the proximal end is expanded. Head not flattened, very large and wide (or expanded mediolaterad). Capital groove wide but very shallow. The surface below (distad) to the head is very wide and flat. Attachment of supraspinatus being a very large, deep groove that gouges the shaft. As a result of this, the median crest is very prominent, and a well-developed ridge is characteristic in this portion of the shaft. In the proximal third, the medial side of the shaft is completely flat, and is more triangular in cross section at this point than in other eagles or hawks examined (see Appendix 1). Line of latissimus dorsi anterioris well developed, protruding posteriorly from shaft (in two specimens available to evaluate this character) at the level of the distal portion of the deltopectoral crest. Pneumatic foramen very large, its diameter much larger than the midshaft diameter. The external border of the shaft is acute distad (or below) to the bicipital crest, as a result of a compression (anteroposterior) that produce the posterolateral flat surface of the shaft at this point. Carpometacarpus (Fig.11C) short and heavy. This element is even larger and more robust than in Titanohierax gloveralleni Wetmore, 1937 (paratype, proximal end of carpometacarpus MCZ 2258) or “Amplibuteo” woodwardi and “A”. hibbardi (slightly more than the half of the length of the latter taxa). In lateral view, the metacarpal I is very wide at its base. The metacarpal III is thinner and located more medially, being much less extended laterad. The proximal symphysis is longer in internal view. In external (or lateral) view, there is a deep depression with a bony wall in the internal side, not present in T. gloveralleni. The internal face of the metacarpal I is markedly excavated. The space between metacarpals I and III is large. Metacarpals II and III well separated. Metacarpal II compressed anteroposteriorly. The trochleae are long. In internal view, the proximal symphysis is short and projected in an open angle. Tarsometatarsus (Fig.11D–G) elongated, larger than the femur (see measurements of this element in Arredondo & Arredondo 2002a: table 1). The proximal articulation and the medial proximal half of this element is unknown in the material from the tar seeps in study (but known from other Cuban localities). In anterior view, the shaft is long and columnar where both proximal and distal ends are proportionately small. Both anterior metatarsal borders are rounded, or convex, pronounced at both sides of the well excavated anterior metatarsal groove. Outer proximal foramen very large, close in size to the distal foramen (the inner is unknown in this material), perforating the bone from anterior to posterior face. Tubercle for tibialis anticus located central and high, slightly diagonal papilla separated from the anterolateral metatarsal border by a well grooved space. Distally the shaft is flat (by an anteroposterior compression), expanded above the trochleae. Outer extensor groove short and deep above (or proximad) to the distal foramen. Trochleae proportionately small in relation with the remaining bone, flaring well distad and abruptly from shaft (more evident in trochlea metatarsi II), and almost uniform in size. Notch between trochleae metatarsorum II and III slightly thinner. Trochlea metatarsi III short (depicting a nearly squared figure), slightly bended laterally, and with a well-defined groove. The outer ring of the latter is the distalmost point of the bone, and the trochlea metatarsi IV the shortest (less distad extended). In lateral view, the shaft is slightly arched and facies subcutanea lateralis ungrooved, being variable from a nearly flat to strongly convex surface. At distal end, the trochlea metatarsi IV is short and ovoid in shape, with posterior wing rounded, poorly projected posterodistad. At the base of the former trochlea, in the posterior half of the border of the shaft, a well-marked groove is present (at this level the bone is extremely rounded anteriorly). In posterior view, the outer proximal foramen is again about the same size of the distal foramen. Posterior metatarsal groove wide, moderately excavated proximad, very shallow distad. Crista plantaris medialis much less prominent distad than crista plantaris lateralis, although the former is better defined reaching distad the proximal end of the metatarsal facet. Metatarsal facet high (proximad), greatly excavated and expanded through its length, being deeper proximad. It is oriented mediad with prominent edges, being the border (medial border of the bone) very thin and acute. At the same level, but in the outer side, a wide and well-excavated (deeper distad) groove turns laterally and reaches the base of the trochlea metatarsi IV. The trochlea metatarsi III possess a moderate sized posterior articular surface, widely grooved, and located in an excavated, neck-like base. Trochlea metatarsi IV also well grooved, having an articular surface much more proximad than in the latter, as a result of its shortness. Trochlea metatarsi II wider at its base, which is relatively flat (or poorly excavated). The wing of this trochlea is located higher (proximal) than it’s distal contour. The distal end is expanded, produced by the flat and wide base of the trochlea metatarsi II (more noticeable than in anterior view). The internal and external intertrochlear notches are wider in posterior view than in anterior view, and both excavate (well proximad) the bone at both sides of the base (or neck) of the trochlea metatarsi III. In medial view, the metatarsal facet is facing medially, and as a result of this medial bending of the metatarsal facet onto the medial border, the crista plantaris medialis is quite visible in its distal portion (because it is less posteriorly located). The medial border along the metatarsal facet is just a thin, acute, and sharp edge, not having the flat surface found in most of the members of Accipitridae. In distal view, the trochleae are massive, aligned near the horizontal plane describing a very smooth arch. Trochleae metatarsorum II and IV with reduction of their posterolateral wings. Trochlea metatarsi III widely grooved through its articular surface. Trochlea metatarsi IV less grooved, only in the posterior half. Metatarsal I with wide digital condyle. The proximal phalanx (Fig. 11H) is relatively short, massive, and also wide. Ungual phalanx (Fig. 11I) with arc less pronounced (versus Arredondo & Arredondo 2002a:10) than in Aquila or Harpia, and close in this condition to the species of “Amplibuteo”.
Measurements. Humerus.—proximal width: 36.7; depth of head: 11.2; least width and depth of shaft at midpoint: 10.8–10.3. Carpometacarpus.—total length: 89.1; proximal width of metacarpal II: 8.0. For measurements of the tarsometatarsus, see Table 6.
Comments. This is the largest known species of Accipitridae in America, either extinct or living (Arredondo & Arredondo 2002a). It represents the diurnal size equivalent of Ornimegalonyx oteroi. In Cuba, its remains are found frequently in Quaternary cave deposits (Suárez pers. obs.). Gigantohierax is a valid genus, distinct from Titanohierax and other Buteonine hawks (Suárez et al. in prep.). Recently, Steadman et al. (2019) recorded a large eagle from Hispaniola (Haiti and the Dominican Republic), based on two fragmentary tibiotarsi and several phalanges recovered in cave deposits of the island. Examination of the photographs, measurements, and descriptions published there (op. cit., figs. 1–4), reveals that the supratendinal bridge in the largest of the two tibiotarsi (Museo Nacional de Historia Natural, Santo Domingo, MNHNSD FOS 24.001), is located more horizontally than in the smaller specimen, character that agrees, along with the huge size of the bone, with G. suarezi. The phalanges also coincide in their size with G. suarezi. The distal half of the left tibiotarsus mentioned above, are referred to the Cuban genus and identified herein as Gigantohierax sp., extending the ancient range of distribution of this genus to Hispaniola. The smaller tibiotarsus represents another species, one with a more vertical supratendinal bridge than in the former, and probably known from the West Indies. Therefore, in the material recorded for Hispaniola by Steadman et al. (2019), two large-sized accipitrids are represented, instead of one, as the authors concluded. Another species in the genus Gigantohierax from Cuba, also known from Las Breas de San Felipe, is represented by elements comparable to those known in other taxa of the Antillean Subregion and the continent.
 
Systematic Palaeontology - 7

Gigantohierax itchei sp. nov.
Holotype. Distal third of right tarsometatarsus without trochlea metatarsi IV, MNHNCu 75.4869. Collected by members of the Departamento de Geología y Paleontología, MNHNCu, in San Felipe II, during field expedition to the type locality in 1998.
Measurements (mm) of holotype. Total length as preserved: 69.5; least width and depth of shaft: 12.0–9.0; distal width from the wing of trochlea metatarsi II to the outer rim of trochlea metatarsi III: 23.5; width and depth of trochlea metatarsi III: 8.4–10.1 (see Table 6).
Paratype (topotype). San Felipe II: Proximal end of fragmentary right femur (MNHNCu 75.4725).
Measurements (mm) of paratype. Femur (G. suarezi in parentheses): depth of neck: 13.7 (19.0); width and depth at level of the distal end of the trochanteric ridge: 18.5 (23.9)–12.2 (16.2); length from distal point of the pneumatic foramen to the distal end of the trochanteric ridge: 19.5 (15.5).
Etymology. After my dear friend Irving “Itche” Himel, Toronto, Canada, for his sincere friendship and unconditional support during my paleontological studies in his country.
Diagnosis. The smallest species in the genus Gigantohierax.
Description. Specimens of Gigantohierax itchei sp. nov. are ~29% smaller than the equivalent elements in the skeleton of G. suarezi (mean 17.63%, range 9.27–26.84% in between-species of large Accipitridae, see Holdaway 1990). The femur (Fig. 11J–L) in anterior view has a large pneumatic depression (= proximal pneumatic foramen), located proximomediad to the distal pneumatic foramen (similar, greater pneumaticity and presence of perforations in the basal wall by some very small foramina in G. suarezi); small distal pneumatic foramen, apparently semitriangular, according to the preserved portions (larger, semitriangular in G. suarezi); the distance be ween the distal margin of the distal pneumatic foramen and the distal end of the trochanteric ridge is very large (smaller in G. suarezi). In posterior view, the proximal surface is irregular, being inflated (or bulky) in its center, as result of the high development of the obturator ridge (flattened and pneumatic surface, with little or no development of the obturator ridge in G. suarezi). The tarsometatarsus (Fig. 11M–P) has a shaft proportionally thinner (anterior and/or posterior view), less flattened or compressed (lateral and/or medial view) than in the largest species (more flattened and compressed, being wide in G. suarezi). In anterior view, it is deeply excavated by a distinctive, very deep and narrow anterior metatarsal groove, which extends well distad (groove much less excavated or extended distad, with anterior surface of the shaft more flattened or convex in G. suarezi). The distal foramen is large (proportionately smaller in G. suarezi), trochlea metatarsi III with very wide and grooved articular surface (relatively narrow in G. suarezi). In posterior view, the posterior metatarsal groove is slightly deeper, but less extended distally (superficial, ending more distad in G. suarezi); crista plantaris medialis better defined (rounded, less defined in G. suarezi); larger distal foramen (relatively smaller in G. suarezi); fossa supratrochlearis plantaris shallow (deep and expanded in G. suarezi); broadly furrowed trochlea metatarsi III (narrow groove in G. suarezi) with very narrow and laterally oriented inner ring (much wider and mediad in G. suarezi); internal side of this last trochlea very excavated medially (less excavated, more superficial in G. suarezi). In cross section, the shaft is squarer, furrowed by the deep anterior metatarsal groove (more triangular, less excavated in G. suarezi).
Comparisons with other Antillean and continental related fossil accipitrids. Gigantohierax itchei sp. nov. is a larger species compared with Buteogallus borrasi, “Amplibuteo” woodwardi, or “A” hibbardi, being close in size to Titanohierax gloveralleni of the Bahamas (Table 6). Gigantohierax itchei sp. nov. differs from all these species by having a wider femur (unknown element in T. gloveralleni), flaring greatly (anterior and/or posterior view) proximad; presence of distinctive concavity formed by the most proximal pneumatic foramen; second (more distad) pneumatic foramen small and semitriangular in shape. In posterior view, the proximal surface is flattened and expanded, not rounded. In lateral view, this element is greatly compressed in anteroposterior direction, being almost flat and the trochanteric ridge is narrow and long distad. Tarsometatarsus (Fig. 11M–P) also more compressed anteroposteriorly, being wide and flat, especially at the distal end; very deep anterior metatarsal groove that extends well down (distad) on shaft; distal foramen large; surface between this last foramen and the external border very wide and rounded (very thin and acute in T. gloveralleni; similar in the other species); trochleae flaring abruptly and distally on shaft; trochlea metatarsi III short, wide, and with a distinct wide groove (longer, thinner, with thin groove in T. gloveralleni; proportionately similar in the remaining species compared); in lateral view, the lateral border or surface is thinner (by the great anteroposterior compression) and rounded, especially at the distal end (wide, concave because it is deeply grooved, less rounded distally in B. borrasi; very wide and flat even distally, only slightly grooved in T. gloveralleni; intermediate in characters in “A”. woodwardi and “A”. hibbardi); internal (inner) side forming a thin, acute or sharp border, specially above the level of the metatarsal facet. In posterior view, the poste rior metatarsal groove is shallow, crista plantaris medialis and crista plantaris lateralis with poor development; metatarsal facet higher (proximal) on shaft; distal foramen ovoid and much larger; trochlea metatarsi III wide, widely grooved, with short articular surface and distinct long neck at base (wide and also widely grooved, long surface without neck in B. borrasi; much thinner and less grooved, much longer or extended posterior surface without neck in T. gloveralleni; intermediate in characters in “A”. woodwardi and “A”. hibbardi); in distal view, the later trochlea is wide but not deep, being squared in morphology (much thinner, very deep, rectangular in T. gloveralleni); outer rim very wide.
Comments. In the initial stages of the process of identification of the specimens described here as Gigantohierax itchei sp. nov. the author considered as the first hypothesis the logical possibility that the holotype of this species, due to its similar size, could represent Titanohierax gloveralleni, an extinct and large buteonine hawk unknown in Cuba so far, and described from the nearby Bahamas (Wetmore 1937; Olson & Hilgartner 1982). After direct comparisons with the type series of the Bahamian taxon, this hypothesis was quickly dismissed. Titanohierax is closer to the genus Geranoaetus (and Buteo Lacépède) than to Buteogallus (Olson & Hilgartner 1982; Suárez & Olson 2007), or Gigantohierax—which shares some characters with Buteogallus (see descriptions above)—, as seen in the anatomy of its tarsometatarsus, especially on both sides of the shaft and in the trochleae. After comparisons and explorations on the osteology of both species described in the extinct genus “Amplibuteo” Campbell (see Campbell 1979), it seems that those must be more accurately placed under Buteogallus (see Olson 2007; Suárez & Olson 2009b).

Figure 11. Gigantohierax suarezi: Left humerus (MNHNCu 75.4722) in anterior/cranial (A) and posterior/caudal (B) views; left carpometacarpus (MNHNCu 75.4723) in dorsal (C) view; right tarsometatarsus (MNHNCu 75.4728) in distal (D), anterior (E), medial (F) and posterior (G) views; phalanx I, left digit I, in dorsal (H) view; ungual phalanx, left digit I (MNHNCu 75.4731) in lateral (I) view. Gigantohierax itchei sp. nov.: Right femur (Paratype, MNHNCu 75.4725) in anterior (J), posterior (K) and lateral (L) views; right tarsometatarsus (Holotype, MNHNCu 75.4869) in anterior (M), medial (N), posterior (O) and distal (P) views. Scale = 2 cm.
 

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Systematic Palaeontology - 8

Genus Buteo Lacépède
Buteo lineatus (Gmelin, 1788)
Referred material. San Felipe II: Proximal end of right femur, MNHNCu 75.3231; distal halves of right MNHNCu 75.3232, and left, MNHNCu 75.3233 tibiotarsi; distal end of left tibiotarsus, MNHNCu 75.3234; distal halves of right MNHNCu 75.3235, and left, MNHNCu 75.3236 tarsometatarsi.
Description. Size, proportions and characters as in Buteo lineatus (see comparisons, description, measurements and illustrations of this material in Suárez & Olson 2003b).
Comments. This species was first recorded in the West Indies from cave deposits in the Bahamas (Olson 2000), after a review (Olson & Hilgartner 1982) of the fossil material of Calohierax quadratus Wetmore, 1937. The only record of this taxon in Cuba is from Las Breas de San Felipe (Suárez & Olson 2003b). Some elements of this species, especially if fragmentary, can be confounded with those of Accipiter gundlachi Lawrence, 1860, or A. cooperii (Bonaparte, 1828), or vice versa, by their roughly similar size, although most of the elements are diagnostic.

Buteo platypterus (Vieillot, 1823)
Referred material. San Felipe II: Distal half of left femur, MNHNCu 75.4740; distal end of right tibiotarsus, MNHNCu 75.4738; proximal end of left tarsometatarsus, MNHNCu 75.4739.
Description. This material represents the smaller species of Buteo in the deposit.
Comments. The Broad-winged Hawk was identified and recorded for Las Breas de San Felipe by Suárez & Olson (2003b:73), based on the material discussed here. Reported by Suárez & Arredondo (1997) from cave deposits in western Cuba. Buteo platypterus Vieillot,1823, is today a common resident in Cuba, and represents an endemic subspecies, B. p. cubanensis Burns, 1911, which lives in forests and pine forests (Garrido & García Montaña 1975; Garrido & Kirkconnell 2011). Individuals of the nominative subspecies are rare, some of them remaining on the island during the winter (Garrido & Kirkconnell 2011:79).

Buteo jamaicensis (Gmelin, 1788)
Referred material. San Felipe I: Distal end of left femur, MNHNCu 75.4741.
Description. This material coincides in its large size and is identical to the equivalent element in the skeleton of Buteo jamaicensis Gmelin, 1788.
Comments. Buteo jamaicensis is the largest species of its genus from these tar seeps. This hawk is the commonest accipitrid in Cuba today, which lives in forests located at low and middle height elevations, in addition to muddy forests (Garrido & Kirkconnell 2011:80).

Buteo sanfelipensis sp. nov.
Holotype. Left tarsometatarsus without trochlea metatarsi IV, MNHNCu 75.4910. Collected in San Felipe I, on May 14, 2009, by William Suárez and Stephen Díaz Franco.
Diagnosis. Tarsometatarsus of moderate length, close to B. swainsoni Bonaparte, 1838, or B. lagopus (Pontoppidan, 1763) in morphology, but with shorter shaft and relatively reduced distal end.
Etymology. The specific name sanfelipensis refers to the tar seeps type locality name, Las Breas de San Felipe.
Description. Tarsometatarsus (Fig. 10A–C) referable to Buteo by characters described by Campbell (1979: 74), of medium size when compared with the equivalent element in other resident (including extirpated) species of the genus in the Greater Antilles (Raffaele et al. 1998), where Buteo platypterus, B. ridgwayi (Cory, 1883), and B. lineatus are smaller species, but B. jamaicensis is larger. Similar to B. swainsoni or B. lagopus in size and general morphology, but more robust and short, with a relatively small distal end (slender and relatively thinner shaft in Buteo lineatus, B. nitidus [Latham, 1790], B. plagiatus Schlegel, 1862; B. ridgwayi, B. albonotatus Kaup, 1847; B. swainsoni, B. lagopus, B. platypterus and B. jamaicensis). Buteo brachyurus Vieillot, 1816, has a much shorter and stouter tarsometatarsus and B. regalis Gray, 1844, is slightly more shorter (proportionally), but it is much larger. Differs from the extinct species Buteo hoffstetteri Campbell, 1976, from the late Pleistocene of Ecuador, by having again a much shorter and robust shaft (see Campbell 1976).
Comments. The only living species of Buteo endemic to the Greater Antilles is B. ridgwayi, from Hispaniola (Raffaele et al. 1998, 2003; Keith et al. 2003; Latta et al. 2006), which is probably a derivative of B. lineatus (Olson 2000) from Cuba (Suárez & Olson 2003b). Buteo sanfelipensis sp. nov. seems to be its endemic equivalent in the Cuban archipelago. Of the remaining taxa known as residents in these islands, Buteo jamaicensis and B. platypterus, have been differentiated only at subspecific level, apparently by a more recent arrival to the Antillean Subregion. All Buteo species known from Cuba have been recorded at Las Breas de San Felipe (Table 10). Wetmore (1937) described the extinct hawk Calohierax quadratus from cave deposits in Great Exuma (= Little Exuma, Hecht 1955; Olson & Hilgartner 1982), Bahamas. This taxon is considered synonym of B. lineatus (Olson 2000; Olson & Hilgartner 1982), or as the valid species B. quadratus (Oswald & Steadman 2018). The following qualitative characters, apart from its larger size, distinguish the tarsometatarsus of Buteo sanfelipensis sp. nov. from that of B. quadratus (see Wetmore 1937:428–430; figs. 1–3): shaft with crista plantaris medialis and crista plantaris lateralis less parallel (flaring) proximad (longer, thinner and more uniform in width, parallel along the entire preserved length of the shaft in B. quadratus); metatarsal facet located higher (or proximad), and distance between its proximal end and the distal foramen, longer (lower and distance much shorter in B. quadratus); distal foramen large (this can be variable; much smaller and more proximad in B. quadratus). It is not impossible that remains of Buteo sanfelipensis sp. nov. can be found in Quaternary deposits of the Bahamas.

Figure 10. Buteogallus royi sp. nov.: Left tarsometatarsus (Holotype, MNHNCu 75.4909) in anterior (A), medial (B) and posterior (C) views; right ulna (MNHNCu 75.4737) in dorsal (D) and ventral (E) views. Buteo sanfelipensis sp. nov.: Left tarsometatarsus (Holotype, MNHNCu 75.4910) in anterior (F), medial (G) and posterior (H) views. Scale = 2 cm.
 

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Systematic Palaeontology - 9

Order STRIGIFORMES Wagler
Family TYTONIDAE Ridgway
Genus Tyto Billberg
Tyto furcata (Temminck, 1827)
Referred material. San Felipe I: Left tarsometatarsus, MNHNCu 75.4800.
Description. The tarsometatarsus reported here differs from all known large extinct tytonids from Cuba and agrees with the living Tyto furcata Temminck, 1827, by its smaller size and slender shaft (see Suárez & Olson 2015:543). Otherwise, it is larger when compared with Tyto sp. listed below.
Comments. The White-winged Barn Owl is a common permanent resident in Cuba (Garrido & Kirkconnell
2011: 153), where remains are known from Quaternary cave deposits (Arredondo 1972, 1975, 1976, 1982, 1984; Suárez 2000b; Jiménez et al. 2005).

Tyto sp.
Referred material. San Felipe I: Proximal end of right tarsometatarsus, MNHNCu 75.4656; proximal end of left tarsometatarsus, MNHNCu 75.4651; proximal end of left tarsometatarsus without inner calcaneal ridge, MNHNCu 75.4655; distal end of the left tarsometatarsus, MNHNCu 75.4657. San Felipe II: Right fragmentary carpometacarpus, MNHNCu 75.4654; distal end of right tarsometatarsus, MNHNCu 75.4652.
Comments. The material here referred to Tyto sp., together with additional specimens from cave deposits, represents a new species (Suárez & Díaz-Franco 2003:375; Suárez 2004b:156), which will be compared, described, and named in a separate paper (Suárez & Olson in press). This constitutes the first record of this taxon for asphalt deposits in Cuba. Fossils of this small barn owl are also known from cave deposits located west of the Cuban archipelago.

Tyto cravesae Suárez & Olson, 2015
Referred material. San Felipe I: Distal end of the left tarsometatarsus, MNHNCu 75.4801.
Description. The specimen MNHNCu 75.4801 has been compared and described by Suárez & Olson (2015), where Tyto noeli Arredondo, 1972, is smaller, but T. pollens Wetmore, 1937, much larger. See additional comparisons in the original description of T. cravesae in Suárez & Olson (2015).
Measurements (Tyto noeli in parentheses, after Suárez & Olson 2015: table 2).—Distal width: 18.1 (14.4–17.3 [15.7] 12).
Comments. The distal end of the tarsometatarsus that represents this large barn owl in Las Breas de San Felipe was formerly recorded as Tyto sp. (Iturralde-Vinent et al. 2000: table 2), and subsequently designated as the paratype of T. cravesae (see Suárez & Olson 2015: 547). The assessment that T. cravesae was the rarest of the Cuban large barn owls (Orihuela 2019:62) and that in Cuba existed four large tytonids with marked anatomical adaptations for terrestrial locomotion (Orihuela 2019: 57), is wrong (see Suárez & Olson 2015: 533, 539–540).

Family STRIGIDAE Leach
Genus Margarobyas Olson & Suárez
Margarobyas lawrencii (Sclater & Salvin, 1868)
Referred material. San Felipe II: right ulna, MNHNCu 75.4809.
Description. This specimen is well preserved and it is indistinguishable from the homologous element in the skeleton of the monotypic Margarobyas lawrencii (Sclater & Salvin, 1868).
Comments. A common endemic species that inhabits forests of Cuba and Isla de la Juventud (Garrido & Kirkconnell 2011:154). Also very common in cave fossil deposits (Arredondo 1984; Jiménez Vázquez & Valdés 1995; Suárez unpubl. data).

Genus Pulsartrix Kaup
Pulsatrix arredondoi Brodkorb, 1969
Referred material. San Felipe II: Proximal half of the left tarsometatarsus, MNHNCu 75.4808.
Description. The tarsometatarsus MNHNCu 75.4808, despite being fragmentary, does not match that of any other Cuban owl in its size and robustness (Fig. 12C–D). It is referred to Pulsatrix arredondoi Brodkorb, 1969, by the shortness of its preserved shaft and by the marked anteroposterior curvature, diagnostic of this taxon (Brodkorb 1969; Arredondo & González Gotera 1982). The bone is also wide and recall the proportions in the much smaller tarsometatarsi present in species of Glaucidium Boie, but also Surnia Linnaeus.
Measurements (holotype of Pulsatrix arredondoi in parentheses, after Brodkorb 1969:113). Tarsometatarsus.— Proximal width: 12.0 (14.1); length of the base of the inner calcaneal ridge: 13.0 (13.6); depth of the outer edge: 4.4 (4.1).
Comments. This is the first record of the species for Las Breas de San Felipe. Pulsatrix arredondoi is known from caves deposits located in western and central Cuba (Arredondo 1984; Cuello 1988; Arredondo & Olson 1994; O. Jiménez Vázquez comm. pers. 2011).

Genus †Ornimegalonyx Arredondo
Ornimegalonyx oteroi Arredondo, 1958
Referred material. San Felipe I: Right tarsometatarsus, MNHNCu 75.4804. San Felipe II: Distal end of the left tarsometatarsus, MNHNCu 75.4807.
Description. This material does not differ in its gigantic size (Fig. 12E–H), or in characters described (see Arredondo 1958, 1976, 1982; Arredondo & Olson 1994) for the tarsometatarsus of Ornimegalonyx oteroi Arredondo, 1958. In addition, it compares well with specimens of this species examined from other Cuban fossiliferous localities (see Appendix 1).
Measurements. Tarsometatarsus.—Total length: 166.9; proximal width: 32.7; least shaft width: 14.5; 15.3; distal width: 31.6; 34.1; width of trochlea metatarsi III: 12.2.
Comments. Remains of the Cuban Giant Owl are well known from other localities in Cuba (Arredondo 1984) since the beginning of the second half of the last century (Arredondo 1958:10). The material now treated here allows refinement of the initial identification published by Iturralde-Vinent et al. (2000).

†Ornimegalonyx sp.
Referred material. San Felipe II: Proximal half of right femur, MNHNCu 75.4714; distal half of fragmentary left femur, MNHNCu 75.4715; proximal fragment of right tarsometatarsus, MNHNCu 75.4716; distal fragment of right tarsometatarsus, MNHNCu 75.4717.
Description. Differs from specimens of Ornimegalonyx oteroi present in this deposit, and from other fossil localities, by its smaller size. The elongated shaft of the tarsometatarsus in the genus Ornimegalonyx (see Arredondo & Olson 1994, fig. 1) separates it from Bubo osvaldoi Arredondo & Olson, 1994, a somewhat smaller Cuban strigid owl not identified so far from these tar seeps.
Comments. Remains of the genus Ornimegalonyx of similar size to the specimens recorded herein, are also known from eastern Cuba (Suárez unpubl. data).

Order FALCONIFORMES Sharpe
Family FALCONIDAE Leach
Genus Caracara Merrem
Caracara creightoni Brodkorb, 1959b
Referred material. San Felipe I: Premaxillary, MNHNCu 75.4742; shaft of right humerus, MNHNCu 75.4759; left humerus without proximal end, MNHNCu 75.4817; proximal half of left humerus, MNHNCu 75.4818; left carpometacarpus, MNHNCu 75.4819; distal halves of right tibiotarsi, MNHNCu 75.4852-4853; distal ends of right tibiotarsi, MNHNCu 75.4854-4856; shaft of left tibiotarsus, MNHNCu 75.4851; right tarsometatarsi, MNHNCu 75.4820, 75.4827-4828; proximal halves of right tarsometatarsi, MNHNCu 75.4844-4847; proximal ends of right tarsometatarsi, MNHNCu 75.4848-4850; right tarsometatarsi without proximal ends, MNHNCu 75.4829-4831; distal end of right tarsometatarsus, MNHNCu 75.4839; left tarsometatarsus without distal end, MNHNCu 75.4840; proximal ends of left tarsometatarsi, MNHNCu 75.4841-4843; distal halves of left tarsometatarsi, MNHNCu 75.4832-4835; distal ends of left tarsometatarsi, MNHNCu 75.4836-4838. San Felipe II: Fragmentary notarium, MNHNCu 75.4579; distal ends of right tibiotarsi, MNHNCu 75.4584-4585; distal ends of left tibiotarsi, MNHNCu 75.4580-4583; distal ends of right tarsometatarsi, MNHNCu 75.4586-4591; distal ends of left tarsometatarsi, MNHNCu 75.4592-4593.
Description. Caracara creightoni Brodkorb, 1959b, differs from C. cheriway (von Jacquin, 1784), C. plancus (J. F. Miller, 1777) and C. lutosa (Ridgway, 1876) in being of smaller size, but it is slightly large and more robust compared with the extinct C. seymouri Suárez & Olson, 2014, from South America. Qualitative characters include a large and high premaxillary, laterally compressed (ventral view), with very rounded dorsal arch (lateral view). Dorsally, culmen and bridge (between nasal openings) wide. Distance between nasal openings (which are wide and ovoid in Caracara, circular in species of Milvago) and tomium, deeper than in the other species. Ulna with large anterior articular ligament scar, tibiotarsus with long fibular crest and wide intercondylar groove. Notarium differs from species of Milvago due to the presence of vertebrae with less bilateral compression. Humerus smaller, with narrow capital groove, rather than wide, as in Milvago. Femur smaller, with shaft thinner at midpoint and well flared at proximal and distal ends. Femoral head less projected proximad, pneumatic foramen large, narrow and more vertical external condyle, and wide intercondylar groove. Tarsometatarsus shorter and relatively robust, with reduced trochlea metatarsi II and not rotated posteriad (large and posteriad rotated in Caracara cheriway, C. plancus and C. lutosa). These characters have been described by Suárez & Olson (2001b, 2003c).
Comments. This is the most common raptor in Las Breas de San Felipe. Caracara creightoni, was originally described from New Providence, in the Bahamas (Brodkorb 1959b; Olson & Hilgartner 1982; Steadman et al. 2007), recorded in Cuba (Suárez & Arredondo 1997), redescribed on the basis of Cuban material and reported for the asphalt deposits treated here (Suárez & Olson 2001b, 2003c). Fossils of the genus Caracara representing large species are known in the West Indies also from Puerto Rico and Mona Island (Wetmore 1920; Olson 2008), Cay man Islands (Morgan 1977, 1994; Olson & Hilgartner 1982), Jamaica (Olson 2008), and Hispaniola (Olson comm. pers. 2007), evidencing a greater distribution in the Antillean Subregion during the past (see summary of species in Suárez & Olson 2014). The Cayman Islands material, identified as C. creightoni (see Morgan 1977, 1994), represents a different species, larger in size (Suárez & Olson 2001b, 2003c; Olson 2008). Orihuela (2019:60) considered Milvago sp., registered only for cave deposits in Cuba by Suárez & Arredondo (1997), as a synonym of C. creightoni, mentioning Suárez & Olson (2003c:305) as proponents of this synonymy. This statement is false, since it does not exist in the cited literature, nor was it proposed by the authors.
 
Systematic Palaeontology - 10

Genus Milvago Spix
Milvago carbo Suárez & Olson, 2003c
Referred material. San Felipe II: Notarium, MNHNCu 75.4567; distal ends of left tibiotarsi, MNHNCu 75.4568, MNHNCu 75.4570-4571; right tarsometatarsus (holotype), MNHNCu 75.4569; proximal half of left tarsometatarsus without part of the inner and outer calcaneal ridges, MNHNCu 75.4572; shaft of left tarsometatarsus, MNHNCu 75.4573; proximal end of right tarsometatarsus, MNHNCu 75.4574; distal half of right tarsometatarsus, MNHNCu 75.4575-4576 (last one with abrasion); distal ends of left tarsometatarsi, MNHNCu 75.4577–4578 (this material is the type series of Milvago carbo Suárez & Olson, 2003c).
Description. This is the largest known species in the genus Milvago (Suárez & Olson 2003c: fig.1; table 1). The referred material is comparable in size to those of the preceding caracara, Caracara creightoni, but are gracile and the tarsometatarsus is very elongated, with wide trochlea metatarsi II at its base, rotated slightly backwards and with the posterior wing perpendicular to the main axis of the shaft (for discussion on osteological characters of the genus Milvago, see Campbell 1980; Emslie 1995; Suárez & Olson 2003c).
Comments. In the Greater Antilles, two extinct species described in the genus Milvago are known: a small one, M. alexandri Olson, 1976, from Quaternary cave deposits in Haiti, Hispaniola (Olson 1976), and M. carbo from Cuba, the largest representative of the genus. In South America, another fossil species, M. brodkorbi Campbell, 1979, is known from asphalt deposits in Talara, Peru (Campbell 1979).

Milvago diazfrancoi sp. nov.
Holotype. Right tarsometatarsus without trochleae metatarsorum II and IV, MNHNCu 75.4610. Collected in San Felipe I, on May 14, 2009, by William Suárez and Stephen Díaz Franco.
Measurements (mm) of holotype. Total length: 57.2; proximal width: 8.4; width of shaft at midpoint of tubercle for tibialis anticus: 6.9; least width of shaft: 3.1; width of trochlea metatarsi III: 2.9.
Diagnosis. The smallest or most delicate species in Milvago, with tarsometatarsus slightly more slender than in M. brodkorbi Campbell, 1979.
Etymology. Species dedicated to my colleague and friend Stephen Díaz Franco, specialist in Antillean fossil mammals and companion in all my expeditions to Las Breas de San Felipe, where together we collected most of the material treated here.
Paratypes (topotypes). San Felipe I: distal third of left tibiotarsus, MNHNCu 75.7021; distal half of right
tarsometatarsus, MNHNCu 75.7022; distal end of right tarsometatarsus, MNHNCu 75.4826; proximal half of right tarsometatarsus, MNHNCu 75.4825; distal half of left tarsometatarsus, MNHNCu 75.7023; distal end of left tarsometatarsus, MNHNCu 75.4824.
Description. This taxon is referable to the genus Milvago and not to Caracara, Daptrius Vieillot, Ibycter Vieillot, or Phalcoboenus d’Orbigny, by having (see Campbell 1979:94; Suárez & Olson 2003c:302) tarsometatarsus with short inner calcaneal ridge, outer calcaneal ridge with proximal orientation, shaft with prominent edges on both sides of the anterior metatarsal groove, trochleae bilaterally compressed, and trochlea metatarsi II broad at base. The tibiotarsus and tarsometatarsus (Fig. 14A–G) are considerably smaller than those present in Milvago carbo (see Suárez & Olson 2003c: fig. 1; table 1). Differs from M. chimachima Vieillot, 1816; M. alexandri, and M. brodkorbi, due to their smaller size (Table 9). The tarsometatarsus has an elongated shaft (shorter in M. chimachima , M. alexandri, and M. brodkorbi; slightly more longer in M. carbo), compressed bilaterally more abruptly at the distal level of the tubercle for the tibialis anticus (less compressed in M. chimachima, M. alexandri, M. brodkorbi, or M. carbo); the fossa supratrochlearis plantaris is expanded or large (smaller in M. chimachima, M. alexandri, M. brodkorbi, or M. carbo).
Comments. This is the second Cuban species, the third for the West Indies, and the fourth in America, described as extinct for the genus Milvago. The latter genus was initially reported for Cuba by Acevedo-González & Arredondo (1982:64, table 1), after an error of identification (Oscar Arredondo, pers. comm. 1997). The subsequent deletion of this taxon from the catalog of Cuban fossil birds (cf. Arredondo 1984) was not commented on. Subsequently, a proximal fragment of a tarsometatarsus (WS 977), from Cueva de Paredones, province of La Habana (now Artemisa), was registered by Suárez & Arredondo (1997:101) as Milvago sp. This constitutes the first valid record of the genus for Cuba. This fossil is not referable to M. diazfrancoi sp. nov., or M. carbo (see Suárez & Olson 2003c), and so the taxon that it represents is totally unknown until now in Las Breas de San Felipe (contra Orihuela 2019:61). Therefore, the alleged synonymy considered by Orihuela (2019:61), of the specimen WS 977 and the Milvago material recovered in these tar seeps is erroneous and unfounded. The species most biogeographically related to M. diazfrancoi sp. nov. is M. alexandri from Hispaniola, which seems to derive from M. chimachima (see Olson 1976), a taxon known from fossils (Pleistocene) in Florida (= Falco readei Brodkorb, 1959a; see Campbell 1980; Emslie 1998). Both have a much shorter tarsometatarsus compared with the species described here, in which this element resembles that of M. brodkorbi, but is more slender. Phalcoboenus chimango (Vieillot, 1816) has been traditionally placed in the genus Milvago (see Fuchs et al. 2012) and well compared with the known fossil species. Its tarsometatarsus is slightly even more slender than in M. diazfrancoi sp. nov. and about the same size (Table 9). Given the distribution of P. chimango (Canevari et al. 1991; White et al. 1994), and its generic position, it seems that all Antillean taxa are related in origins with M. chimachima, evolving with a more elongated tarsometatarsus than in the former, and more adapted to terrestrial locomotion, as in the case of P. chimango (see Mosto et al. 2013), under insular conditions and in sympatry with other endemic caracaras.

Figure 14. Milvago diazfrancoi sp. nov.: Right tarsometatarsus (Holotype, MNHNCu 75.4610) in anterior (A) and posterior (B) views; right tarsometatarsus (Paratype, MNHNCu 75.4825) in anterior view (C); left tarsometatarsus (Paratype, MNHNCu 75.7023) in posterior (D), and anterior (E) views; left tibiotarsus (Paratype, MNHNCu 75.7021) in distal (F) and anterior (G) views. Scale = 2 cm.
 

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Systematic Palaeontology - 11

Genus Falco Linnaeus
Falco sparverius Linnaeus, 1758
Referred material. San Felipe I: Distal half of left tibiotarsus, MNHNCu 75.4758.
Description. The known specimen does not differ from Falco sparverius sparverioides Vigors, 1827, being smaller and less robust when compared with the nominative form F. s. sparverius Linnaeus, 1758.
Comments. The American Kestrel is a common permanent resident in Cuba, Isla de la Juventud, and some cays, where it prefers habitats with open areas (Garrido & Kirkconnell 2011: 82). It has been previously registered for Cuban cave deposits (Jiménez Vázquez 1997; Suárez & Olson 2001a). As in Accipiter striatus fringilloides, remains of this kestrel of Cuba can be quickly distinguished from their continental equivalent by the smaller size.

Falco femoralis Temminck, 1822
Referred material. San Felipe I: Right carpometacarpus without minor metacarpal, MNHNCu 75. 4606; right carpometacarpus without distal end and minor metacarpal, MNHNCu 75.4607; distal end of left tibiotarsus, MNHNCu 75.4608; proximal end of left tarsometatarsus, MNHNCu 75.4609.
Description. These fossils were described by Suárez & Olson (2003b: fig. 1D,E; table 2) and coincide perfectly in size and characters with Falco femoralis Temminck, 1822. Falco sparverius, F. columbarius Linnaeus, 1758, and the fossil F. kurochkini Suárez & Olson, 2001a, all are smaller species, but the fossils are small in relation to the skeletal elements of F. peregrinus Tunstall, 1777.
Comments. The fossil material that represents this falcon in Las Breas de San Felipe, the only known from Cuba, was previously recorded by Suárez & Olson (2003b). The species lives on the continent, where it inhabits swampy places and plains (A.O.U. 1998). Its extinction in Cuba is difficult to explain, since this habitat has been present on this archipelago since the Pleistocene.

Order PASSERIFORMES Linnaeus
Family CORVIDAE Leach
Genus Corvus Linnaeus
Corvus palmarum Württemberg, 1835
Referred material. San Felipe II: left tarsometatarsus without proximal end, MNHNCu 75. 4816.
Description. Tarsometatarsus slender in comparison with the corresponding element in the skeleton of Corvus nasicus Temminck, 1826.
Measurements. Tarsometatarsus: Total length: 55.5; proximal width: 8.6; width and depth of shaft at midpoint: 3.4–4.2; distal width: 6.1.
Comments. The Palm Crow is a rare species in Cuba today, but locally common in some points of its current relictual distribution (Garrido & Kirkconnell 2011:191). Abundant material has been recovered in cave deposits of Artemisa Province (Suárez & Arredondo 1997:101; Suárez 2000b: 64, table 1), providing evidence of a greater former distribution within the Cuban archipelago. This species does not currently live in the Province of Matanzas, since it is limited to a few localities in open areas of the provinces of Pinar del Río (possibly extirpated population; Suárez per. obs. 2001) and Camagüey (Garrido & Kirkconnell 2011: 191).

Corvus nasicus Temminck, 1826
Referred material. San Felipe I: Proximal half of right ulna, MNHNCu 75.4812; left tarsometatarsus, MNHNCu 75.4810; distal half of right tarsometatarsus, MNHNCu 75.4811. San Felipe II: Proximal fragment of right ulna, MNHNCu 75.4813; distal fragment of left ulna, MNHNCu 75.4814; proximal fragment of left femur, MNHNCu 75.4815.
Description. The largest species of Corvus Linnaeus in this deposit. Tarsometatarsus shorter when compare with C. palmarum, and inseparable from Corvus nasicus Temminck, 1826.
Comments. The most common corvid in Las Breas de San Felipe, being one of the first known species in the locality (Iturralde-Vinent et al. 2000). The Cuban Crow is today a rare resident in Cuba, Isla de la Juventud, and in some large cays north of the Ciego de Ávila and Camagüey Provinces, found in forests, palm groves, pine forests, and around swampy areas, including those of Matanzas Province (Garrido & Kirkconnell 2011:192).
 
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