AN ANNOTATED LIST OF LATE QUATERNARY EXTINCT BIRDS OF CUBA

Abstract

· Within the Antilles, Cuba has a peculiarly diverse fossil avifauna. However, information on this avifauna is scattered among the specialized literature. Here I provide an updated annotated taxonomic list of the fossil birds from Cuba. This list includes 35 taxa, of which 17 are endemic, 12 actually extirpated, and 6 are undefined species identified only to genus level. The list is richly diverse in raptors with varied adaptations, including giant owls with limited flight and four large barn-owls, all with anatomical adaptations that suggest pronounced ground-dwelling. The raptor list includes five hawks, five falcons, and three vultures. There are also records of an egret, a stork, a crane, a snipe, and a nighthawk. Most species seem to have become extinct in Cuba, probably during the Late Holocene.Resumen · Lista de la avifauna fósil de Cuba Cuba tiene una avifauna fósil peculiarmente diversa. No obstante, la información taxonómica al respecto se encuentra dispersa en la literatura especializada. Se presenta aquí una lista actualizada sobre la taxonomía de la avifauna fósil de Cuba, reconociéndose 35 taxones extintos, incluyendo 17 endémicos y 12 taxones localmente extinguidos o extirpados y 6 taxones identificados solo al nivel de género. Entre la fauna extinguida conocida prevalecen las aves rapaces, incluyendo búhos gigantes, lechuzas y, un teratornítido con adaptaciones que indican capacidades nulas o limitadas de vuelo. Además, hay cinco gavilanes, cinco halcones y tres buitres. También se registran una grulla, una cigüeña, una garza, un alcaraván y otras aves que se han extinguido localmente, posiblemente durante el Holoceno tardío o periodo colonial.

Full text

(2019) 30: 57–67
Johanset Orihuela
Department of Earth and Environment (Geosciences), Florida Internaonal University, Miami, Florida 33199, USA.
E-mail: paleonycteris@gmail.com
Abstract · Within the Anlles, Cuba has a peculiarly diverse fossil avifauna. However, informaon on this avifauna is scaered among the
specialized literature. Here I provide an updated annotated taxonomic list of the fossil birds from Cuba. This list includes 35 taxa, of which 17
are endemic, 12 actually exrpated, and 6 are undened species idened only to genus level. The list is richly diverse in raptors with varied
adaptaons, including giant owls with limited ight and four large barn-owls, all with anatomical adaptaons that suggest pronounced
ground-dwelling. The raptor list includes ve hawks, ve falcons, and three vultures. There are also records of an egret, a stork, a crane, a
snipe, and a nighthawk. Most species seem to have become exnct in Cuba, probably during the Late Holocene.
Resumen · Lista de la avifauna fósil de Cuba
Cuba ene una avifauna fósil peculiarmente diversa. No obstante, la información taxonómica al respecto se encuentra dispersa en la literatu-
ra especializada. Se presenta aquí una lista actualizada sobre la taxonomía de la avifauna fósil de Cuba, reconociéndose 35 taxones exntos,
incluyendo 17 endémicos y 12 taxones localmente exnguidos o exrpados y 6 taxones idencados solo al nivel de género. Entre la fauna
exnguida conocida prevalecen las aves rapaces, incluyendo búhos gigantes, lechuzas y, un teratornído con adaptaciones que indican capa-
cidades nulas o limitadas de vuelo. Además, hay cinco gavilanes, cinco halcones y tres buitres. También se registran una grulla, una cigüeña,
una garza, un alcaraván y otras aves que se han exnguido localmente, posiblemente durante el Holoceno tardío o periodo colonial.
Key words: Anlles · Exncon · Fossil birds · Insularity · Late Holocene · Late Pleistocene · Raptors · Teratorns
INTRODUCTION
Currently, the Cuban archipelago has a diverse avifauna comprising 371 recorded species within 20 orders and 60 families
(Garrido & Kirkconnell 2000). Of these, six genera and 21 species are endemic to the archipelago (Garrido & Kirkconnell 2000,
González 2012). Bird fossil and/or subfossil specimens indicate a diverse and unique exnct avifauna, whose remains are gen-
erally found in Late Quaternary deposits throughout the island. These avian assemblages are peculiarly diverse in endemic
birds of prey, including large-sized species with ground-dwelling anies, limited ight, and relavely large size. Oversized
owls of the genus Ornimegalonyx, for instance, are among the world’s largest strigids (Arredondo 1970, Garrido & Kirkconnell
2000).
Cuba’s unique exnct bird fauna was rst studied by Alexander Wetmore, who reported La Brea Stork (Ciconia maltha),
Cuban Macaw (Ara tricolor), and Black-chested Buzzard-Eagle (Geranoaetus melanoleucus) from fossils found in the thermal
baths, sinkhole deposits of Ciego Montero, in central Cuba (Wetmore 1928). Yet we owe the late Cuban paleontologist Oscar
Arredondo de la Mata for laying the foundaons in the study of the island’s paleofauna by publishing over 134 scienc papers
on fossil vertebrates between 1945 and 2001, on which he described 38 species, especially birds (Arredondo 2007: 152). His
work has been connued and complemented by the eorts of Jiménez (1997, 2001), Jiménez & Arrazcaeta (2008, 2015); Suár-
ez (2000a, 2000b, 2000c, 2004a, 2004b, 2004c), and Suárez & Olson (2001, 2003, 2007, 2009, 2015). These contributors have
not only revised Arredondo’s records, amending and revising specimens but have also discovered and described new ones that
expand the understanding of Cuban and Caribbean ancient avifaunas.
In 1984, Arredondo published a preliminary list which was later updated by Suárez (2004a). These lists have not been re-
vised for over a decade, and do not reect current taxonomic arrangements. Since Suarez’ 2004 list, several new species have
been described while other taxa have been synonymized or removed from the record. For instance, specimens assigned to the
ightless ibis Xenicibis, the vulture Sarcoramphus, and the giant hawk Titanohierax, previously reported for Cuba, have been
removed (Suárez 2001a, 2001b, 2004a). The exnct stork Ciconia maltha is now considered a junior synonym of Ciconia lydek-
keri (Ameghino 1891) (Agnolin 2009), and the enigmac snipe reported by Suárez (2004) was recently nominated as Gallinago
kakuki (Steadman & Takano 2016). During the late 1960s and early 1970s, new records based on specimens found in Cueva de
Pío Domingo, Pinar del Río, were described (Fischer 1968, 1977; Fischer & Stephan 1971a, 1971b). Most of these records were
AN ANNOTATED LIST OF LATE QUATERNARY EXTINCT BIRDS OF CUBA
Receipt 10 May 2018 · First decision 30 July 2018 · Acceptance 15 April 2019 · Online publicaon 28 April 2019
Communicated by Kaspar Delhey © Neotropical Ornithological Society
57

ORNITOLOGÍA NEOTROPICAL (2019) 30: 57–67
58
subsequently revised by Olson, including Rallus sumiderensis
and Fulica picapicensis, which were idened as the Zapata
Rail (Cyanolimnas cerverai) and the exnct rail Nesotrochis
picapicensis, respecvely (Olson 1974).
Several amended records and the most recent discover-
ies, however, have not been assimilated into the recent pale-
ontological and zooarchaeological literature, and are in need
of a compilaon that can be used by specialists and non-
specialists alike. As a summary, here I provide an updated
annotated taxonomic list of the exnct and exrpated bird
species known from the Late Quaternary record of Cuba,
which builds on the previous list of Suárez (2004a), Arredon-
do, and others. Addionally, the holotypes and new speci-
mens of several species are illustrated in detail for the rst
me. However, a record of extant birds found in archaeologi-
cal contexts is not included but will be provided elsewhere.
This list can be useful for studies about the evoluon of the
avian diversity in the Greater Anlles.
METHODS
The list includes only the exnct taxa currently known by
fossil or subfossil remains. Although many of these exnct
species are known from Latest Quaternary, likely pre-
Columbian (Amerindian) Late Holocene deposits, the majori-
ty of them do not have direct or associated radiocarbon ag-
es. The few radiocarbon dates associated or directly taken
from bird remains will be briey discussed to provide a
framework for the survival or me of exncon.
The inclusion of all exisng representave specimens of a
given taxon, state of preservaon in parcular remains, and
detailed stragraphic provenance lie outside the scope of
this list. In most cases, this informaon is not provided in
original descripons or records cited for Cuban exnct birds.
In the case of stragraphy, most remains are usually found in
cave deposits which lack discernible stragraphy or a stra-
graphic framework. Nevertheless, to unify criteria on each
species account, the following informaon will be provided:
synonyms (when available), holotype, lectotype or other
collecon number and repository reference, type locality or
where it was rst discovered in Cuba, status, and remarks as
observaons on idencaon or age. Widespread, non-
endemic exnct species will be indicated as globally exnct,
endemics as exnct endemics, and those extant elsewhere,
but locally exnct in Cuba will be denoted as exrpated. Ana-
tomical terminology follows the terms of Baumel & Witmer
(1993), and the Handbook of the Birds of the World (HBW)
for common names. Radiocarbon dates are given in radiocar-
bon years before the present (rcyrBP, present being AD
1950) with a standard error. A map of the most important
Cuban localies where avian remains were found is provided
in Figure 1. Photographs in Figures 2, 5, and 7, are used with
the permission and courtesy of L. W. Viñola; the others are
from the author.
Instuonal abbreviaons. Collecons and instuons that
serve as repositories of Cuban exnct bird remains meno-
ned in the text include: AC 33: ArqueoCentro, Villa Clara Pro-
vince, Cuba; AMNH: American Museum of Natural History,
New York City, USA.; AV: Colección de la Universidad de La
Habana, La Habana, Cuba; BMNH: Brish Museum of Natural
History, London, UK; CAZG: Colección Arqueozoológica del
Gabinete de Arqueología, Ocina del Historiador de La Haba-
na; CZACC: former Colección Zoológica de la Academia de
Ciencias de Cuba, La Habana, now at the Instuto de Ecolo-
gía y Sistemáca (IES), Mayabeque, Cuba; DPUH: Departa-
mento de Paleontología, Facultad de Biología, Universidad
de La Habana, La Habana; FLMNH-UF: Florida Museum of
Natural History, University of Florida, Gainesville, USA; LWV:
Lazaro W. Viñola collecon, La Habana, Cuba; MCZ: Museum
of Comparave Zoology, University of Harvard, Cambridge,
Massachuses, USA; MNHNCu: Museo Nacional de Historia
Natural, La Habana, Cuba; OA: Oscar Arredondo collecon,
La Habana, Cuba; PB: Pierce Brodkorb collecon, Florida Mu-
seum of Natural History, Gainesville, USA; USNM: United
States Naonal Museum, Smithsonian Instuon, Washing-
ton, DC, USA; WS: William Suárez collecon, housed at the
MNHNCu.
RESULTS
Pelecaniformes
Ardeidae
Tigrisoma mexicanum Swainson 1834
First material reported in Cuba: complete le tarsometa-
tarsus (AC 33), collected in 2004. Locality: Casimba en Los
Buentes, a sinkhole deposit in Mal Páez, near Sagua La Gran-
de, Villa Clara Province (Olson & Suárez 2008).
Status: The Bare-throated Tiger-heron is exrpated in Cuba,
but with a current wide geographical range in Central Ameri-
ca (Butchard & Symes 2018).
Remarks: This specimen was reported in associaon with Ara
tricolor, another Cuban exnct endemic (Olson & Suárez
(2008). This constuted the rst fossil record of the species
in the West Indies. In the original report, Olson & Suárez
(2008) considered that this specimen could represent an
endemic subspecies.
Ciconiiformes
Ciconiidae
Ciconia lydekkeri Ameghino 1891
Synonyms: Jabiru mycteria Wetmore (1928)
Ciconia maltha (Howard 1942)
C. maltha (Miller 1910: 440). See Agnolin (2009)
Ciconia lydekkeri (as redescribed by Agnolin 2009)
First material reported in Cuba: Right tarsometatarsus and
le distal biotarsus (AMNH without a number) collected by
Barnum Brown and Carlos de la Torre between 1910 and
1918 (Suárez & Olson 2003a).
First fossil locality in Cuba: Ciego Montero thermal bath de-
posits, Cienfuegos Province (Wetmore 1928).
Holotype of Ciconia lydekkeri: Right distal tarsometatarsus
(BMNH 18879).
Status: Ciconia lydekkeri is a globally exnct species with a
pan-American Quaternary distribuon (Agnolin 2009).
Ciconia sp.
Type material: Right extreme distal biotarsus Museo Nacio-
nal de Historia Natural, La Habana, Cuba (MNHNCu P4599).
Type locality: Breas de San Felipe tar deposits, near Mar,
Matanzas Province (Suárez & Olson 2003a); locality 13
(Figure 1). The fauna at this locality has yielded Late Pleisto-

CUBAN EXTINCT BIRD LIST
59
cene to Late Holocene radiocarbon dates between 4960 ±
280 and 11,880 ± 420 rcyr BP (Jull et al. 2004).
Status: Undened.
Remarks: Species was described by Suárez & Olson (2003a:
151) as “smaller than C. maltha.”
Mycteria wetmorei Howard 1935
First material reported in Cuba: Right carpometacarpus
(MNHNCu P4602) and a right distal biotarsus (MNHNCu
P4603).
First report locality in Cuba: San Felipe tar deposits, near
Mar, Matanzas Province (Iturralde et al. 2000, Jull et al.
2004).
Status: Globally exnct, with former distribuon in North
America (Suárez & Olson 2003a). Remarks: Species occurred
in sympatry with Ciconia sp. at the San Felipe tar deposits
(Suárez & Olson 2003a).
Incertae sedis
Teratornithidae
Oscaravis olsoni (Arredondo & Arredondo 1999)
Synonyms: Teratornis olsoni Arredondo & Arredondo 1999:
310
Oscaravis olsoni (as redescribed by Suárez & Olson 2009:
106)
Holotype: near complete right femur (CZACC 400-649, now
at IES) (Figure 2).
Type locality: Cueva de Paredones, San Antonio de Los Ba-
ños, Artemisa Province.
Remarks: Specimens of this species are known from at least
ve cave sites distributed between the current Artemisa, La
Habana, and Mayabeque provinces (Figure 1).
Status: Exnct endemic.
Catharformes
Cathardae
Gymnogyps varonai (Arredondo 1971)
Synonyms: Anllovultur varonai Arredondo (1971: 310)
Gymnogyps (Suárez 2000a). See also Suárez & Emslie (2003)
Holotype: Le proximal tarsometatarsus (DPUH 1254).
Type locality: Cueva de Paredones, San Antonio de Los Ba-
ños, Artemisa Province.
Status: Exnct endemic.
Cathartes? sp. 1
Cathardae gen. et sp. indet.
Remarks: Suárez (2001c: 110) menoned the presence of
two undescribed and possibly dierent New World vultures
(Cathardae). One is probably referable to Cathartes but
diers from Cathartes aura (Suárez 2001). Another taxon of
the same family but gen. et sp. indet. could represent a
dierent genus altogether (Suárez 2001c: 110, L. W. Viñola
pers. comm.).
Suárez (2000) revised specimens previously idened as fos-
sil Cathartes aura from Mayabeque (Arredondo & Varona
1974) and concluded that these represented modern speci-
mens, thus deleng this taxon from the Cuban fossil record
(Jiménez & Arrazcaeta 2008). Cathartes aura is supposed to
have arrived in Cuba during the post-Colombian, so far re-
ported only from 17th-century archaeological contexts from
La Habana Vieja (Jiménez & Arrazcaeta 2008). However, a
premaxillary specimen of Cathartes aura was recently found
in a paleontological deposit at Cueva de la Caja, Mayabeque
province, where radiocarbon dated specimens yielded
Figure 1. Map of the Cuban archipelago depicng main localies were exnct avifauna has been found: (1) Cueva de Pío Domingo, Sumide-
ro, Pinar del Río Province; (2) Cueva El Abrón, Sierra de la Güira, Pinar del Río Province (Cueva del Mono Fósil is near); (3) Sierra de Caballos,
Isla de la Juventud Municipality : (4) Cueva de Sandoval, Caimito, Artemisa Province; (5) Cueva Lamas, Santa Fe, La Habana Province; (6)
Cueva de Paredones, San Antonio de los Baños, Artemisa Province; (7) Cueva del Túnel, La Salud, Mayabeque Province; (8) Cuevas Blancas,
Quivicán, Mayabeque Province; (9) Cueva de la Caja (or “Cueva de los Nesofontes”), Mayabeque Province; (10) Cueva de Bellamar, Matan-
zas, Matanzas Province; (11) Cueva del Quinto, Camarioca, Matanzas Province; (12) Cueva de Cantel, Calero, Matanzas Province; (13) Breas
de San Felipe (San Felipe tar pits), Mar, Matanzas Province; (14) Baños de Ciego Montero, Cienfuegos Province; (15) Hornos de Cal, Sanc
Spíritus Province; (16) Cueva del Indio, Daiquirí, Sanago de Cuba Province.

ORNITOLOGÍA NEOTROPICAL (2019) 30: 57–67
60
ages between 1960 ± 30 and 1290 ± 30 BP (Orihuela unpubl.
data). This suggests the presence of Cathartes aura well in
the pre-Columbian, Late Holocene of Cuba.
Status: Undened.
Accipitriformes
Accipitridae
Amplibuteo woodwardi (L. Miller 1911)
Synonyms: Amplibuteo sp. (Suárez & Arredondo 1997)
First material reported in Cuba: Incomplete associated skele-
ton, William Suárez collecon (WS 365), housed at MNHNCu.
First fossil locality in Cuba: Cueva de Sandoval, Caimito, Arte-
misa Province (Suárez 2004b). Status: Globally exnct.
Remarks: This is an exnct taxon that inhabited North Ameri-
ca during the Quaternary. Its fossil remains are known from
the western United States and Florida and may be found in
other parts of the Caribbean, along with other yet un-
described accipitrids (Suárez 2004a, b: 124).
Buteo lineatus (Gmelin 1788)
First material reported in Cuba: Right proximal femur
(MNHNCu P4614), collected in 1988, at San Felipe tar pits,
locality II, Mar, Matanzas Province (Suárez & Olson 2003b).
Status: Exrpated. The Red-shouldered Hawk is an extant
taxon, with a wide distribuon in North and Mesoamerica
(Ferguson-Lee & Chrise 2005: 220).
Remarks: The Cuban remains were described as “more con-
sistently robust” than the comparave material of the Grey
Hawk (Buteo nidus), whose remains may also be expected
in West Indian contexts due to its wide distribuon (Suárez &
Olson 2003b). Fossil remains of F. lineatus have been also
reported from the Bahamas (Olson 2000).
Buteogallus borrasi (Arredondo 1970)
Synonyms: Aquila borrasi Arredondo (1970: 3)
Aquila sp. in Fischer (1977)
Titanohierax borrasi (Olson & Hilgartner 1982, Suárez 2000c)
Few specimens of this species were assigned to Sarco-
ramphus (? sp.) in Acevedo & Arredondo (1982), Arredondo
& Arredondo (1999), Garrido & Kirkconnell (2000), Iturralde
et al. (2000), and Suárez (2000c, 2001).
Holotype: Le tarsometatarsus (DPUH 1250), deposited at
Departamento de Paleontología, Facultad de Biología, Uni-
versidad de La Habana, La Habana.
Type locality: Cueva del Túnel, La Salud, Mayabeque Province
(locality 7 on Figure 1).
Remarks: Widespread species, known from eight cave depo-
sit localies, including Cueva Lamas, Cueva de Sandoval, Cue-
vas Blancas, and Breas de San Felipe (Figure 1). Buteogallus
borrasi was described as more robust and 33% larger than
Buteogallus urubinga (Suárez & Olson 2007).
Status: Exnct endemic, although Suárez and Olson insinuat-
ed that it may be present in other localies of the Greater
Anlles (Suárez & Olson 2007). Olson & Hilgartner (1982)
suggested that it should be included in the genus Titanohier-
ax. Recently, this taxon was transferred to the genus Bu-
teogallus, deleng Titanohierax from the Cuban record
(Suárez & Olson 2007).
Geranoaetus melanoleucus Swan 1922
First fossil material from Cuba: le carpometacarpus (AMNH
6190 in Wetmore 1928).
First fossil locality from Cuba: Ciego Montero, Cienfuegos
Province (Wetmore 1928).
Status: Exrpated. The Black-chested Buzzard Eagle has a
modern, wide distribuon in South America, especially the
southern cone, where it inhabits dry, mountainous scrubland
(Ferguson-Lee & Chrise 2005: 202).
Remarks: The species has not been reported since.
Gigantohierax suarezi Arredondo & Arredondo 1999
Synonym: Aguila borrasi (Arredondo, 1970: 3)
Holotype: Le femur (MNHNCu P574), discovered since the
late 1950s.
Type locality: Cueva de Sandoval, Caimito, Artemisa.
Status: Exnct endemic.
Remarks: Specimens of this species were originally included
in Arredondo’s descripon of Aquila borrasi, some of which
remains are now a synonym of Buteogallus borrasi.
Falconiformes
Falconidae
Caracara creightoni Brodkorb 1959
Synonyms: Milvago sp. of Suárez & Arredondo (1997) was
referred to this taxon by Suárez & Olson (2003c: 305). Cara-
cara plancus (= C. cheriway) in Jiménez (1997). See Suárez &
Olson (2003c: 306).
First fossil material from Cuba: Fragmentary skull in Oscar
Arredondo’s collecon (OA 3928). First fossil locality in Cuba:
Figure 2. Oscaravis olsoni holotype (complete right femur CZACC
400-649) from Cueva de los Paredones, Mayabeque; in (A) cranial
and (B) caudal views. Photograph used with the permission and
courtesy of L. W. Viñola. Scale bar=10 mm.

CUBAN EXTINCT BIRD LIST
61
Cueva Calero, Matanzas Province (Suárez & Arredondo 1997;
Suárez & Olson 2001b, 2003c).
Status: Globally exnct.
Remarks: Taxon with a wide distribuon in the Quaternary
that included Cuba and the Bahamas. Suárez & Olson (2001a)
hypothesized a recent arrival to Cuba, along with Cathartes.
In Cuba, it is known from at least six deposit localies in
northwestern and central provinces, suggesng also a wide-
spread distribuon within the main island. Archaeological
human remains at Cueva Calero have yielded Late Holocene
radiocarbon ages ranging between 1670 ± 60 and 1590 ± 70
rcyrs BP (Ulloa 2008, Roksandic et al. 2015). However,
whether said material occurred in the archaeological o pale-
ontological context of Calero’s deposit was not registered,
and thus the specimen’s direct age is unknown (Suárez &
Olson (2001b, 2003c). Remains of this species have recently
yielded a radiocarbon age of 2390 ± 30 rcyrs BP from a de-
posit in Great Abaco, Bahamas (Steadman & Franklin 2015:
Table 2).
Milvago carbo Suárez & Olson 2003
Holotype: Right tarsometatarsus (MNHNCu P4569).
Type locality: San Felipe tar deposits, Mar, Matanzas
Province (Iturralde et al. 2000).
Status: Exnct endemic.
Remarks: Occurred in probable sympatry with Caracara
creightoni at San Felipe (Suárez & Olson 2003).
Milvago sp. 1
Type locality: San Felipe tar pits, Mar, Matanzas Province
(Iturralde et al. 2000).
Status: Undened.
Remarks: The original indicaon of this species is from
Suárez & Arrendondo (1997). Suárez & Olson (2003: 302)
noted that although these specimens are too fragmentary
to be diagnosc, they are similar in size to Milvago chima-
chima.
Falco femoralis Temminck 1922
First fossil material from Cuba: Right carpometacarpus
(MNHNCu P4606), collected in 2001 at San Felipe I, Area C,
San Felipe tar pits, Mar, Matanzas Province (Suárez & Olson
2003b). Status: Exrpated.
Remarks: The Aplomado Falcon has an ample modern distri-
buon that includes the southern United States through to
southern South America (Ferguson-Lee & Chrise 2005).
Falco kurochkini Suárez & Olson 2001
Holotype: Le tarsometatarsus (MNHNCu P3229).
Type Locality: Cueva de Sandoval, Caimito, Artemisa Province
(Suárez 2000a, 2000b, 2000c, 2004b).
Status: Exnct endemic.
Gruiformes
Gruidae
Grus cubensis (Fischer & Stephan 1971)
Synonyms: Baeopteryx cubensis Fischer (1968)
Grus cubensis (Fischer & Stephan 1971)
Holotype: Skull (CZACC 1/67).
Type locality: Cueva de Pío Domingo, Sumidero, Pinar del
Río.
Status: Exnct endemic.
Rallidae
Nesotrochis picapicensis (Fischer & Stephan 1971) (Figure 3)
Synonyms: Fulica picapicensis by Fischer & Stephan (1971).
Nesotrochis picapicensis (Olson, 1974)
Holotype: Complete le humerus (AV 832/67).
Type locality: Pío Domingo, Sumidero, Pinar del Río Province.
Status: Exnct endemic.
Remarks: This taxon has been found in mid-late Holocene
deposits (mostly in archaeological sites dated between
1190±40 and 3331±17 rcyrs BP; Jiménez pers. Comm. 2018),
and post-Columbian cave deposits in Mayabeque (Jiménez
1997, 2001; Orihuela 2010). It is likely that this species be-
came exnct during the post-Columbian-colonial interval
(Jiménez & Arrazcaeta 2008, 2015).
Charadriiformes
Burhinidae
Burhinus sp. cf. B. bistriatus (Wagler 1829).
First fossil material from Cuba: Le distal humerus (OA
2958).
First fossil locality in Cuba: Cueva de Paredones, San Antonio
de Los Baños, Artemisa Province (Arredondo 1984; Suárez
2000b 2004b; Iturralde et al. 2000).
Status: Exrpated.
Remarks: This specimen has been idened as the Double-
striped Thick-knee Burhinus bistriatus by Suárez (2005). This
taxon has a current wide distribuon in North and Central
America, including parts of the Greater Anlles and The Ba-
hamas.
Figure 3. Nesotrochis picapicensis referred specimen: proximal end
of a right femur (MNHNCu 75.3231) from Cueva de los Nesofontes,
Loma del Palenque, Mayabeque Province; in medial (A) and cranial
(B) views. Scale bar = 10 mm.

ORNITOLOGÍA NEOTROPICAL (2019) 30: 57–67
62
Scolopacidae
Gallinago kakuki Steadman & Takano 2016
Synonyms: Capella sp. (Suárez 2004a, 2004c)
First fossil material from Cuba: Right humerus (MNHNCu
75.4709).
First fossil locality in Cuba: El Abrón Cave, Sierra de la Güira,
Pinar del Río (Suárez 2004b). See also Suárez & Díaz-Franco
(2003).
Gallinago kakuki holotype: Complete right humerus (UF
297382), collected in 1958–1960 from Banana Hole, Baha-
mas (Steadman & Takano 2016).
Status: Globally exnct.
Remarks: Widespread in the Greater Anlles, including Cay-
man, Bahamas, and likely within Cuba.
Psiaciformes
Psiacidae
Ara tricolor Bechstein 1811
Synonym: Ara cubensis of Wetherbee (1985).
First fossil material from Cuba: Right proximal carpometa-
carpus deposited at the AMNH without number.
First fossil locality in Cuba: The fossilized material was rst
discovered at Ciego Montero thermal bath deposits, Cien-
fuegos Province (Wetmore 1928: 4).
Status: Exnct endemic.
Remarks: Other fossil localies include the Casimba en Los
Buentes, where it was associated with remains of Tigrisoma
mexicanum (Olson & Suárez 2008). Exnct since the mid-late
19th century (Garrido & Kirkconnell 2000, Wiley & Kirwan
2013).
Strigiformes
Tytonidae
Tyto noeli (Arredondo 1972)
Synonyms: Tyto noelli (Arredondo 1972a) in Suárez & Olson
(2015)
Tyto neddi (Steadman & Hilgartner 1999: 76)
Tyto noeli (Suárez & Olson 2015: 541)
Holotype: Right tarsometatarsus (DPUH 1251), collected in
1968.
Type locality: Cueva del Túnel, La Salud, Mayabeque Provin-
ce.
Status: Globally exnct.
Remarks: Species with a wide Quaternary record that in-
cludes Jamaica and Barbuda (Suárez & Olson 2015). Also
with a widespread range in Cuba, where it is known so far
from at least ten localies. Occurs in sympatry with Tyto
furcata in several Cuban cave deposits (Arredondo 1972a,
Suárez 2002), and likely with Tyto cravesae at Cueva Cente-
lla, Matanzas (L. W. Viñola pers. comm.), where Tyto noeli
has been reported (Orihuela 2013) (Figure 4). Directly
radiocarbon dated material of this species from Cueva El
Abrón, Sierra de la Güira, Pinar del Río, yielded a Late Pleisto-
cene age of 17,406 ± 161 rcyr BP (Suárez & Díaz-Franco 2003:
373). At Drum Cave in Jamaica, remains of this species were
associated with radiocarbon dates between 3700 ± 150 and
6410 ± 110 rcyr BP (McFarlane et al. 2002).
Tyto pollens (Wetmore 1937)
Synonym: Tyto riveroi Arredondo 1972b: 131, revised by
Suárez & Olson (2015: 539)
Holotype: Le femur (MZC 2262), collected in 1937.
Type locality of Tyto riveroi: Cueva de Bellamar, Matanzas
Bay, Matanzas Province (Arredondo 1972b).
Status: Globally exnct.
Remarks: Known from other Quaternary records in the Baha-
mas and within Cuba (Suárez & Olson 2015).
Tyto cravesae Suárez & Olson 2015
Synonym: Several specimens of this taxon were included in
the original type series of Tyto noeli in Oscar Arredondo’s
collecon (OA) (Suárez & Olson 2015).
Holotype: Associated postcranial long bone elements
(MNHNCu 75.590), likely represenng a single individual col-
lected in 1998.
Type locality: Salón del Pozo, Cueva de Paredones, San Anto-
nio de los Baños, Artemisa Province.
Status: Exnct endemic.
Remarks: This is the rarest of Cuban tytonid owls, known
so far from four localies in the provinces of Artemisa,
Mayabeque, and Matanzas, northwestern Cuba (Suárez &
Olson 2015). Associated with remains of Oscaravis olsoni and
Ornimegalonyx “minor”, Gymnogyps varonai and Puslatrix
arredondoi at the type locality.
Tyto sp. 1
A “small, undescribed species” was reported from Cueva El
Abrón, Sierra de la Güira, Pinar del Río (Suárez & Díaz-Franco
2003: 375).
Status: Undened.
Figure 4. Tyto noeli referred specimen (fragmentary right ulna
MNHNCu uncatalogued) from Cueva Centella, Matanzas Province;
in cranial (A), caudal (B) and dorsal (C) views. Scale bar = 10 mm.

CUBAN EXTINCT BIRD LIST
63
Strigidae
Bubo osvaldoi Arredondo & Olson 1994 (Figure 5)
Holotype: Right distal tarsometatarsus (MNHNCu 27.1).
Type locality: Cueva del Mono Fósil, Pinar del Río Province.
Status: Exnct endemic.
Remarks: Found in sites spanning from eastern and western
Cuba, suggesng it was widespread on the main island
(Arredondo & Arredondo 1999: 18).
Pulsatrix arredondoi Brodkorb 1969 (Figure 6)
Holotype: Le tarsometatarsus collected by Oscar Arredondo
in 1960, deposited in the Pierce Brodkorb collecon (PB
8420, sensu Brodkorb 1969).
Type locality: Cueva de Paredones, San Antonio de Los Ba-
ños, Artemisa Province. See also Arredondo & González
(1982) and Arredondo (1984).
Status: Exnct endemic.
Remarks: Previously known from three localies spanning
from Artemisia, Matanzas and San Spíritus, now with two
addional records in Mayabeque (Jiménez et al. in press).
One of the new records is from an archaeological assem-
blage at Cueva de los Muertos: a distal tarsometatarsus
(CAZG04), collected in May 2012 (Jiménez et al. in press).
This specimen (Figure 6) yielded a radiocarbon age of 1390 ±
30 rcyrs BP, which provides a last appearance date for this
taxon well into the Late Holocene (Jiménez et al. in press).
Ornimegalonyx
Remarks: All Ornimegalonyx species need revision. It is likely
that all Ornimegalonyx represent a single species. Their size
disparity could be due to sexual dimorphism, chrono-
temporal or/and individual variaon (Alegre 2002, Louchart
2005). This is a well-distributed genus within Cuba. As the
wide distribuon of other exrpated and exnct raptors sug-
gests, this genus likely inhabited other parts of the Greater
Anlles.
Ornimegalonyx oteroi (Arredondo 1958) (Figure 7)
Synonyms: Ornimegalonyx arredondoi (Arredondo 1958),
formerly redescribed by Brodkorb (1961)
Holotype: A le tarsometatarsus lectotype (MZC P-383E)
assigned by Brodkorb (1961: 634). Type locality: Cueva de Pío
Domingo, Sumidero, Pinar del Río Province.
Status: Exnct endemic.
Remarks: This hypodigm includes several specimens original-
ly assigned to Phorusrhacidae by Arredondo (1958) but in-
cluded in Strigidae by Brodkorb (1961). Also includes a speci-
men idened as Cathartes aura by Arredondo (1989: 9), but
amended by Suárez (2001: 110). Likely a widespread taxon.
Specimens are known from at least nine deposit localies
throughout the Cuban archipelago (Arredondo 1996).
Ornimegalonyx acevedoi Arredondo 1982
Holotype: Le tarsometatarsus (GEC without catalog num-
ber, see Suárez 2004a).
Type locality: Cueva del Quinto, Camarioca, Matanzas.
Status: Exnct endemic.
Ornimegalonyx minor Arredondo 1982
Synonyms: Ornimegalonyx sp. (Arredondo 1975)
O. “minor” (Suárez & Olson 2015)
Holotype: Right proximal femur (MCZ P-37).
Figure 5. Bubo osvaldoi holotype (fragmentary right tarsometa-
tarsus MNHNCu 27.1), from Cueva del Mono Fósil, Province Pinar
del Rio; in (A) dorsal and (B) plantar views. Photograph used with
the permission and courtesy of L. W. Viñola. Scale bar = 10 mm.
Figure 6. Pulsatrix arredondoi referred specimen, fragmentary right
tarsometatarsus (CAZGA04) from Cueva del Muerto, Mayabeque
Province; in (A) dorsal and (B) plantar views.

ORNITOLOGÍA NEOTROPICAL (2019) 30: 57–67
64
Type locality: Cueva de Paredones, San Antonio de los Baños,
Artemisa Province (Arredondo 1982, 1984).
Status: Exnct endemic.
Ornimegalonyx gigas Arredondo 1982
Synonym: Ornimegalonyx sp. Arredondo (1975: 142–143)
Holotype: Le proximal femur (MCZ 3032).
Type locality: Canteras de Hornos de Cal, San Spíritus.
Status: Exnct endemic.
Caprimulgiformes
Caprimulgidae
Siphonorhis daiquiri Olson 1985
Holotype: Right proximal humerus (USNM 336506).
Type locality: Cueva de Los Indios, near Daiquirí, Sanago de
Cuba (Figure 1, locality 16; see Anthony 1919 and Suárez
2000b).
Status: Exnct endemic.
Passeriformes
Rhinocrypdae
Scytalopus sp.
First fossil material from Cuba: A right proximal humerus
(USNM 33605).
First fossil locality in Cuba: Sierra de Caballos, Isle of Pines,
and Cueva de los Fósiles, Camagüey (Olson & Kurochkin
1987).
Status: Exrpated.
Remarks: Possible widespread in the island as suggested
by the widely separated record localies, one on the Isle
of Pines, western Cuba, and other in Camagüey, central
Cuba.
Icteridae
Dolichonyx kruegeri Fischer & Stephan 1971
Holotype: Right complete humerus (AV 877/67).
Type locality: Pío Domingo, Sumidero, Pinar del Río Province.
Status: Likely exrpated or accidental record.
Remarks: Forgoen record since the original descripon,
which may likely represent a misidened specimen of the
Bobolink (D. oryzivorus), who is an uncommon transient spe-
cies in Cuba (Garrido & Kirkconnell 2000: 218).
DISCUSSION
The Cuban avian fossil record indicates a high level of ende-
mism and diversity. The list reveals a total of 35 exnct spe-
cies, of which 17 are endemic taxa, and 12 are currently ex-
rpated forms. The status of the remaining six taxa (17%) is
sll uncertain or undened. These disappearances comprise
~ 9 % out of the total >370 bird species registered for the
Cuban archipelago (Garrido & Kirkconnell 2000).
The known assortment of Cuban exnct birds is remarka-
ble in its diversity of diurnal and nocturnal raptors, of which
most are endemic and notably oversized with ground-
dwelling adaptaons, represenng 65.7 % of the exnct
know fauna. The present avian assemblages include ve ac-
cipitrids (13%), six falconids (17.1 %), comprised of two large
hawks, plus two caracaras and two smaller falcons. The fossil
record is parcularly diverse in Strigiformes, which comprise
24% total of the currently know avifauna, or 13% if only one
Ornimegalonyx is counted. These include the six large, ight-
limited, endemic strigids of the genus Ornimegalonyx, plus
two endemic owl species and four exnct Tyto. Addionally,
there is a large condor and two yet undescribed vultures,
which comprise eight percent of the exnct avifauna. Per-
haps the most enigmac of these is the rare Oscaravis, an
exnct endemic teratorn bird; a probable relic of a more
ancient fauna with a South American anity (Suárez & Olson
2009). Many of the exrpated and globally exnct seem to
have been geographically widespread within the Greater
Anlles. In this sense, and taking into consideraon several
of the rich cave deposits being researched in the Bahamas
and Hispaniola, it would not be surprising to nd several Cu-
ban species in its assemblages, even few of those that are
today considered Cuban endemics, as our knowledge of their
geographical distribuon improves.
Cuba’s exnct avifauna is generally assumed to be Late
Quaternary in age (Suárez 2004a), although most species and
their contexts lack direct radiocarbon dates. For most of the
20th century, many of the species records were arbitrarily
considered of Late Pleistocene age, as was the case with oth-
er Cuban exnct vertebrates (e.g., Anthony 1919, Wetmore
1928, Acevedo et al. 1975; Arredondo 1958, 1970, 1972a,
1972b). However, recent direct radiocarbon dang has
shown that several of the Cuban exnct vertebrates have
late appearance dates that extend well into the Amerindian
pre-Columbian Late Holocene (MacPhee et al. 1999, Jull et al.
1994, Jiménez et al. 2005, Orihuela 2010, Orihuela & Tejedor
2012).
Of all the exnct Cuban birds, only two have been dated
directly by radiocarbon methods: Tyto noeli, which yielded a
Late Pleistocene date (17,406 ± 161 rcyr BP) from a cave de-
posit at Cueva El Abrón, in Pinar del Río Province (Suárez &
Diaz-Franco 2003; Tyrberg 2009), and Puslatrix arredondoi
from Cueva de los Muertos (Figure 6), which yielded an age
of 1390 ± 30 rcyr BP (Jiménez et al. in press). Others, such as
the ightless rail Nesotrochis picapicensis have been found
associated to radiocarbon dated Amerindian pre- and post-
Figure 7. Ornimegalonyx oteroi referred specimen (fragmentary le
tarsometatarsus LWV 890) from Cueva Beruvides, Matanzas Provin-
ce; in (A) dorsal, (B) plantar and (C) lateral views. Specimen record
used with permission and courtesy of L. W. Viñola. Scale bar = 10
mm.

CUBAN EXTINCT BIRD LIST
65
Arredondo, O (1958) Aves gigantes de nuestro pasado prehistórico. El
Cartero Cubano 17: 10–12.
Arredondo, O (1970) Nueva especie de ave pleistocenica del orden
Accipitriformes (Accipitridae) y nuevo género para las Anllas.
Ciencias Biológicas 4: 1–19.
Arredondo, O (1971) Nueva especie de ave fósil (Accipitriformes: Vultu-
ridae) del Pleistoceno de Cuba. Memorias de la Sociedad de Ciencias
Naturales, La Salle 31: 309–323.
Arredondo, O (1972a) Nueva especie de ave fósil (Strigiformes: Tytoni-
dae) del Pleistoceno superior de Cuba. Bolen de la Sociedad Vene-
zolana de Ciencias Naturales 29: 122–123.
Arredondo, O (1972b) Especie nueva de lechuza (Strigiformes: Tytoni-
dae) del Pleistoceno cubano. Bolen de la Sociedad Venezolana de
Ciencias Naturales 30: 124–125.
Arredondo, O (1975) Distribución geográca y descripción de algunos
huesos de Ornimegalonyx oteroi Arredondo 1958 (Strigiformes:
Strigidae) del Pleistoceno superior de Cuba. Memorias de la Socie-
dad de Ciencias Naturales La Salle 35: 135–190.
Arredondo, O (1976) The great predatory birds of the Pleistocene of
Cuba. Smithsonian Contribuon to Paleobiology 27: 169–188.
Arredondo, O (1982) Los Strigiformes fósiles del Pleistoceno cubano.
Bolen de la Sociedad Venezolana de Ciencias Naturales 140: 33–
55.
Arredondo, O (1984) Sinopsis de las aves halladas en depósitos fosilí-
feros pleisto-holocénicos de Cuba. Reporte de Invesgación del
Instuto de Zoología 17: 1–35.
Arredondo, O & C Arredondo (1999a) Nuevos género y especie de ave
fósil (Falconiformes: Accipitridae) del Cuaternario de Cuba. Poeyana
470–475: 9–14.
Arredondo, O & C Arredondo (1999b) Nueva especie de ave
(Falconiformes: Teratornithidae) del Pleistoceno de Cuba. Poeyana
470/475: 15–21.
Arredondo, O & N González (1982) Nuevo hallazgo de Pulsatrix arredon-
doi Brodkorb (Aves: Strigidae) del Pleistoceno de Cuba. Miscelánea
Zoológica 16: 1–2.
Arredondo, O & S L Olson (1994) A new species of owl of the genus
Bubo from the Pleistocene of Cuba (Aves: Strigiformes). Proceedings
of the Biological Society of Washington 107: 436–444.
Baumel, J & L Witmer (1993) Osteologia. Pp 45–132 in Baumel, J, A King,
J Breazile, H Evans & J Vanden Berge (eds). Handbook of avian anat-
omy: Nomina Anatomica Avium. Publicaon of the Nuall Ornithol-
ogy Club, Boston, Massachusses, USA.
Borroto-Páez, R & CA Mancina (2017) Biodiversity and conservaon of
Cuban mammals: past, present, and invasive species. Journal of
Mammalogy 98: 964–985.
Brodkorb, P (1961) Recently described birds and mammals from Cuban
caves. Journal of Paleontology 35: 633–635.
Brodkorb, P (1969) An exnct Pleistocene owl from Cuba. Quarterly
Journal of the Florida Academy of Sciences 31: 112–114.
Butchart, S & A Symes (2018) BirdLife Internaonal. 2016. Tigrisoma
mexicanum. The IUCN Red List of Threatened Species 2016:
e.T22697259A93605056. Downloaded on 23 March 2018 from
hp://dx.doi.org/10.2305/IUCN.UK.2016-3.RLTS.T22697259A9360-
5056.en.
Cooke, SB, LM Dávalos, AM Mychajliw, ST Turvey & NS Upham (2017)
Anthropogenic exncon dominates Holocene declines of West
Indian mammals. Annual Review of Ecology, Evoluon, and System-
acs 48: 301–327.
Ferguson-Lees, J & DA Chrise (2005) Raptors of the world. Princeton
Field Guides. Princeton Univ. Press, Princeton, New Jersey, USA.
Fischer, K (1968) Ein ugunfähiger Kranich aus dem Pleistozän von
Cuba. Monatsschri für Ornithologie und Vivarienkunde 15: 270–
271.
Fischer, K (1977) Quartäre Mikromammalia Cubas, vorwiegend aus der
Höhle San Jose de la Lamas, Santa Fe, Provinz Habana. Zeitschri für
Geologische Wissenscha 5: 213–255.
Columbian assemblages (dated between 1190 and 3331 BP)
in latest Holocene cave deposits of western Cuba (Jiménez
1997, 2001; Jiménez & Arrazcaeta 2008, 2015; Orihuela
2010, unpubl. data). The presence of many of these taxa, in
associaon with other small vertebrates in Cuban Amerin-
dian and post Columbian Holocene deposits, not only indi-
cate a very late survival past the climac uctuaons of
the Quaternary, including its Last Glacial Maximum, but also
human coexistence for several thousands of years before
nal exncon that implicate complex responses as causes
for their exncon or long lag periods before nal exnct-
on or exrpaon (MacPhee et al. 1999, Turvey 2009,
Orihuela & Tejedor 2012, unpubl. data). In the case of the
Cuban exnct endemics, the causes suggest the acon of
combined threats involving both background climate change
and direct human environmental degradaon, the introduc-
on of exoc species, deforestaon, and some cases over-
hunng, all especially during the last 500 years (Borroto-Páez
& Mancina 2017, Cooke et al. 2017). Although the
ming and likely causes leading to exncon are becoming
beer understood for mammals (Borroto-Páez & Mancina
2017), much remains to be unraveled in the case of birds,
many which sll require taxonomic denion or lack last
occurrence data. Further discovery, analysis, and direct da-
ng of addional fossils will likely increase the list of Cuba’s
exnct avifauna, shed light on its diversity and the causes
and ming of their exncon.
ACKNOWLEDGMENTS
I thank Lázaro W. Viñola, Adrián Tejedor, and Osvaldo Jimé-
nez Vázquez for revising several dras of this manuscript,
providing pernent literature, and clarifying several points
on the actual state of Cuba’s exnct birds. L. W. Viñola kindly
provided several photographs of holotype specimens to be
used here. Addional thanks are due to three referees, espe-
cially Federico Agnolin and Marcos Cenizo, whose sugges-
ons and correcons largely improved this work. Most espe-
cially, I thank Osvaldo Jiménez, Storrs Olson, William Suárez,
and Dave Steadman for their unfaltering dedicaon to Carib-
bean paleornithology. Their work has been an inspiraon to
many of us.
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