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Title: Pleistocene birds from New Providence Island, Bahamas
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00001556/00001
 Material Information
Title: Pleistocene birds from New Providence Island, Bahamas
Alternate Title: Bulletin of the Florida State Museum ; volume 4
Physical Description: 350-371 p. : illus., map. ;
Language: English
Creator: Brodkorb, Pierce, 1908-
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1959
Subject: Birds, Fossil   ( lcsh )
Paleontology -- Bahamas -- New Providence Island   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Bibliography: p. 369-371.
 Record Information
Bibliographic ID: UF00001556
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0864
notis - ALW8331
alephbibnum - 002363712
oclc - 05069405
lccn - a 59009672

Table of Contents
    Front Cover
        Page 345
        Page 346
        Page 347
        Page 348
        Page 349
        Page 350
        Page 351
        Page 352
        Page 353
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        Page 355
        Page 356
        Page 357
        Page 358
        Page 359
        Page 360
        Page 361
        Page 362
        Page 363
        Page 364
        Page 365
        Page 366
        Page 367
        Page 368
        Page 369
        Page 370
        Page 371
    Back Cover
        Page 372
        Page 373
Full Text
2. ~ /










BIOLOGICAL SCIENCES, are published at irregular intervals. Volumes contain
about 300 pages and are not necessarily completed in any one calendar year.


The publication of this volume of THE BUL-
LETIN was made possible by a grant from
the Graduate School, University of Florida.

All communications concerning purchase or exchange of the publication should
be addressed to the Curator of Biological Sciences, Florida State Museum, Scagle
Building, Gainesville, Florida. Manuscripts should be sent to the Editor of the
BULLETIN, Flint Hall, University of Florida, Gainesville, Florida.

Publication date
No. 1 23 April 1959
No. 2 27 April 1959 ........
No. 3 28 April 1959
No. 4 7 May 1959 .......
No. 5 14 May 1959 .............- .....
No. 6 15 May 1959 ............. ......
No. 7 20 May 1959 ._
No. 8 21 May 1959 ......-....-...-.. ..--.-- .
No. 9 22 May 1959.... ..
No. 10 3 June 1959
No. 11 4 June 1959


A Tabular Summary of the Biology of North American Mayfly
Nymphs (Ephemeroptera). LEWIS BEHNER . . . 1

Growth and Development of the Eastern Harvest Mouse,
Reithrodontomys humulis. JAMES N. LAYNE . . 61

A Checklist of the Chironomidae (Insecta) of Florida
(Diptera: Chironomidac). ELTZABETH C. BECK and
WILLIAM. M.. . . . . . 85

Variation in Lizards of the Leiocephalus cubensis Complex in
Cuba and the Isla de Pinos. ALBERT SCHWATZ . . 97

The Freshwater Decapod Crustaceans of the Apalachicola Drain-
age System in Florida, Southern Alabama, and Georgia.
HOnTON H. HOBBS, JR., and C. W. HART, JR. . ... 145

The Ostracods of the Genus Entocythere from the Lower Chatta-
hoochee Flint Basin: With a Review of the Occurrence of the
Genus in Florida, and Descriptions of Two New Species.
C. W HART, JR. . . . . . . . .193

Boarfishes of the Genus Antigonia of the Western Atlantic.
FREDERICK H. BERRY .... . . . .205

The Epaxial Musculature of Siren, Amphiuna, and Necturus
(Amphibia). WALTER AUFFENBERC . . . . .253

The Pleistocene Avifauna of Arredondo, Florida.
PIERCE BRODKORB . . . . . .269

The Atlantic Loggerhead Sea Turtle, Caretta caretta caretta (L.),
in America. DAVID K. CALDWELL, ARCHIE CARR, and others 293

Pleistocene Birds from New Providence Island, Bahamas.
PIERCE BRODKORB . . . . . . .349





Volume 4


Number II


Pierce Brodkorb





SYNOPSIS: Only 4 of 15 species of birds from a Pleistocene deposit on New
Providence still exist on the island. The other species either are extinct or occur
now farther south in the Bahamas and Greater Antilles. The extinct forms in in-
clude 2 raptors previously known only from Great Extina Island and 6 new spe-
uies: Caracara creightoni, Burhinus nanus, Glaucidium dickinsoni, Otus providen-
tiae, Bathoceleus hyphalns (new genus, Picidae), and Corvus wetimorei.
The fossil deposit is assigned to the pre-Pam lico portion ot the Wisconsin
glacial stage, when the sea had retreated to the 10-fathom mark to expose a
large land mass which reached within 10 miles of Cuba. At this time the avi-
fauna of the Bahamas appears to have been about 40 percent richer than at
present. The faunal tie to the Greater Antilles, particularly Cuba, was strong.
while the relationship to Florida was weaker than today.

Knowledge of the past bird life of the island of New Providence
has heretofore been lacking. Such information from elsewhere in the
Bahamas is meager, being limited to a paper on Pleistocene birds from
Great Exuma Island (Wetmore, 1937B) and to a report of bird re-
mains from an Indian midden of ceramic age on Crooked Island
(Wetmore, 1938). Although small, these collections are of great
interest. From Great Exuma the 13 species identified include 3 ex-
tinct raptors and 3 living forms presently confined to the Greater
Antilles. The 11 species reported from the pre-Columbian site on
Crooked Island also include several birds that no longer occur on
that island, although still extant elsewhere in the West Indies.
During the summer of 1958 Dr. J. C. Dickinson, Jr., of the Uni-
Sversity of Florida, discovered a Pleistocene vertebrate locality on
New Providence. With the assistance of Dr. Walter Auffenberg pre-
liminary excavations were carried out during August. Abundant re-
mains of large extinct rodents (Geocapromys) comprise the bulk of
the collection, but bats, birds, reptiles, and frogs are also present.
The avian material, which forms the subject of this report, contains
a high proportion of extinct species, besides others that no longer

The author is Professor of Biological Sciences at the University of Florida,
Gainesville. Manuscript received 8 May 1959-Eu.




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exist on New Providence. As it adds materially to our knowledge
of the origin of the ornis of the Bahamas, it is to be hoped that further
investigations may soon be made at this important site.

The fossil locality is a sinkhole in oolitic limestone near the west-
crn end of New Providence. A large banana tree growing in the sink
led the collectors to designate it the Banana Hole. The sink is lo-
cated along the roadside between Lyford Cay and Clifton Pier. It is
just north of the old settlement of Clifton and lies about one-half
mile inland from Clifton Point. The surface elevation is about 30
feet above present sea level. The floor of the sink lies about 20 feet
below the surface and consists of reddish brown clay containing ver-
tebrate fossils. Excavation was made to a depth of only 1 foot.
When brought to the laboratory the bones were covered with
reddish matrix. Washing and microscopic inspection showed the
matrix to consist of nothing but clay, with no other minerals or ma-
rine invertebrates. The bones themselves, when cleaned, have a
huffy coloration and are more heavily mineralized than cave fossils
usually are. Many of them are abraded, but others show no erosion.
They are thought to have accumulated in the sink largely through
the feeding activities of raptorial birds.

Class AVES
Calohierax quadratus Wetmore
Plate I, fig. 1
Calohierax quadratus \VWetiore. 1937B, p. 429, fig. 1-3 (Pleistocene, Great Exunma
REFERRED MATERIAL. Distal portion of right tibiotarsus, UFC
DESCRIPTION. Agrees with Buteo Lacepbde and Buteogallus Les-
son and differs from Accipiter Brisson in having shaft of tibiotarsus
stout and flaring gently above internal condyle; internal rugosity of
oblique ligament situated near upper end of supratendinal bridge.
Differs from Buteo and Buteogallus in having raised edges of peroneal
groove forming ridges, not shelves; internal rugosity of oblique liga-
ment situated slightly lower on shaft.


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Plate I. Fig. 1 Calohierax quadratus Wetmore, tibiotarsus, UFC 3152, X 1.3.
I.ig. 2-4, Burhinlus naous n. sp., holotypc conraioid, UFC 3154, X I.3; tibiotarsaLs
UF'C 3155, X 1.3; tarsometatarsus. UFC 3156, X 1.1. Fig. 5-6, Tyto pollens
Wetinore, tibiotarsus, UFC 3195, X 0.9; tarsometatarsus, UIC 3196, X 1. Fig. 7,
Caracara creightoni n. sp., holotype carpometacarpus, UFC 3153, X 1.3.

Vol. 4


Size near that of female Buteo lineatus alleni, larger than Buteo
platypterus, smaller than Buteo jamaicensis and Buteogallus anthraci-
nus. Least width of shaft, 5.1 mm.; abrasion of the condyles prevents
taking other standard measurements.
DISTnuRTIoN. Calohierax was described on the basis of a frag-
mentary tarsometatarsus from Great Exuma, and the type has hereto-
fore remained unique.


Caracara creightoni, new species
Plate I, fig. 7
HOLOTYPE. Left carpometacarpus, lacking proximal end and shaft
of metacarpal III, UFC 3153. From Pleistocene at Banana Hole,
New Providence Island, Bahamas. Collected by J. C. Dickinson, Jr.,
and Walter Auffenberg, 28 August 1958.
DIAGNosIs. Differs from living Caracara cheriway (Jacquin) of
tropical and subtropical America in having carpometacarpus with
distal edge of pisiform process curved forward; a depression on shaft
distad to pisiform process; metacarpal III with its base nearly straight,
without medial angulation proximal to intermetacarpal tuberosity;
tuberosity of metacarpal II, in medial view, with its outline more
angular and less rounded, and its base more deeply excavated dor-
sally and laterally; subterminal tubercle on dorsal surface of shaft
of metacarpal II obsolete. Size somewhat smaller than in congeneric
species, living and fossil.
Length, from anterior edge of metacarpal I to distal condyle of
metacarpal II, 44.9 mm. (46.0-49.0 in C. cheriway); width of meta-
carpal II, at level of intermetacarpal tuberosity, 4.9 mm. (5.1-5.3 in
C. cheriway).
ETYMOLOGy. The new species from the Bahamas is dedicated to
Albert M. Creighton, of Boston and Nassau, in recognition of his sup-
port of the summer's field work.
In substituting the generic name Caracara Merrem for Polyborus
Vieillot, Hellmayr and Conover (1949) retained the masculine end-
ings of the several species formerly in Polyborus, and this action was
followed by the American Ornithologists' Union (1957). As the name
Caracara is of barbaric rather than classical origin, its gender is to
be determined by the original author. Merrem (1826) included four
species in Caracara. For three of these, C. accipitrina, C. aquilina,


and C. crotophaga, lie used adjectives with feminine endings. Only
the fourth specific name, Caracara plancus, is masculine, but the
word plancus is a noun (meaning a kind of eagle), and therefore its
gender is not effected by the gender of Caracara. As all the adjec-
tives Merrem used in Caracara are feminine, it is obvious that he con-
sidered his new generic term as of feminine gender, and the endings
of the specific adjectives currently placed in Caracara should agree.
DITnusBUTIoN. In the modern fauna this arid tropical and sub-
tropical genus occurs on the mainland from southern South America
to Arizona, Texas, and southern Florida. Its modern West Indian
range extends only to Cuba and the Isle of Pines, where the main-
land species, Caracara cheriway, occurs.
Caracara prelutosa (Howard, 1938) is known from the Pleistocene
of California and Florida, and from early Recent deposits in New
Mexico. It was represented in the Pleistocene of Nuevo Le6n by a
slightly differentiated race, Caracara prelatosa grinnelli (Howard.
1940). The only prehistoric West Indian representative hitherto known
is Caracara latbchrosa (Wetmore, 1920, 1922A), from a Quaternary
cave deposit on Puerto Rico.


Burhinus nanus, new species
Plate 1, figs. 2, 3, 4

HOLOTYPF. Left coracoid, lacking lower end, UFC 3154. From
Pleistocene at Banana Hole, New Providence Island, Bahamas. Col-
lected by J. C. Dickinson, Jr., and Walter Auffenberg, 28 August 1958.
DTAGNosis. Differs from living Burhinus domninicensis (Cory) of
Hispaniola in smaller size; much smaller than living Burhinus bistri-
atus (Wagler) of Central and South America (see table 1).
REFERRED MATEnIAL. Proximal portion of right tibiotarsus. UFC
3155; proximal portion of left tarsometatarsus, UFC 3156.
ETYr OLOGY. Latin, nanus, dwarf.
DISTRIBUTION. Although widespread in the Old World, the thick-
knees, family Burhinidae, occur in America at present only in arid
portions of the Neotropical mainland, from southern Mexico to Peru
and Brazil, and on the island of Hispaniola.

Vol. 4



Element B. nonus B. dominicensis (3) B. bistriatus (2)

Length to notch above in-
ternal sternal angle 22.5 24.9-25.3 26.1-28.8
Head to procoracoid foramen 13.5 14.1-15.1 14.5-15.9
Least w\idth of shaft 3.1 3.5- 3.6 .8- 3.9
Length to end of fibular ridge 22.6 24.3 24.9-26.7
Proximal width 8.2 9.7 10.2-11.2
Width through fibular ridge 5.1 6.1 6.9- 7.1
'T arsonllcttatarsus:
Proximal width 9.5 10.9-11.1
Depth through hypotarsus 8.8 9.6-10.0
Width of shaft at middle 3.4 3.6- 3.7



Columba squamosa Bonnaterre
REFEIRED MATERIAL. Anterior portion of sternum, UFC 3157;
left and right coracoids, UFC 3158-3159; 2 right ulnas, UFC 3160-
3161; right tarsometatarsus, UFC 3162.
DESCRIPTION. Bones from Columba squamosa differ from those
of Columba leucocephala in being larger and more robust, although
some measurements overlap. In addition the inner surface of the
sternal plate of C. squamosa has near its anterior end a large median
foramen, which is lacking in C. leucocephala. Most of the bones
referred above agree with modern skeletons, but the two ulnas are
even larger and heavier than in available comparative material. The
length of the single complete fossil ulna is 58.4 mm., against 54.1-
56.1 in modern C. squamosa. This could reflect a specific difference,
and therefore further material is desirable.
DISTRIBUTION. At the present time the Scaled Pigeon has a wide
distribution in the West Indies, being absent only from Jamaica and
the Bahamas. It was previously reported, however, from the Pleisto-
cene of Great Exuma in the Bahamas (Wetmore, 1937B) and has also


been found in prehistoric sites on Puerto Rico (Wetmore, 1922A,
1938), St. Croix (Wetmore, 1937A). and Martinique (Wetmore, 1952).

Columba leucocephala Linnacus
REFERRED MATERIAL. 3 left coracoids, UFC 3163-3165; 2 left hu-
meri, UFC 3166-3167; right ulna, UFC 3168; 4 left and 4 right carpo-
metacarpi, UFC 3169-3176; left digit IT, phalanx 1 of manus, UFC
3177; left and right femora, UFC 3178-3179; left tibiotarsus, UFC
3180; left and right tarsometatarsi, UFC 3181-3182.
DIs mRIBrioN. The White-crowned Pigeon is common today in
the Florida Keys, Bahamas, Greater Antilles, and northern Leeward
Islands. It has been recorded from prehistoric sites on Crooked Island
in the Bahamas (Wetmore, 1938), Puerto Rico (Wetmore, 1922A),
and St. Croix (Wetmore, 1937A), but this is the first Pleistocene record
of the species.
Zenaida aurita (Temminck)
REFERRED MATERIAL. Left coracoid, UFC 3183; right scapula,
UFC 3184; 2 left and 1 right humeri, UFC 3185-3187; 2 left ulnas, UFC
3188-3189; 3 left carpometacarpi, UFC 3190-3192.
DISTRIBUn ON. The Zenaida Dove occurs today throughout the
West Indies and on the coast of Yucatan. It has been recorded from
prehistoric sites on Puerto Rico (Wetmore, 1922A, 1938), St. Croix
(Wetmore, 1937A), and Martinique (Wetmore, 1952). but this is its
first Pleistocene occurrence.

Amazona leucocephala (Linnaeus)

REFERRED MATERIAL. Right tarsometatarsus. UFC 3193; distal
end of right radius, UFC 3194.
DISTRIBUTION. This parrot is limited at present to the Caymans,
Cuba, the Isle of Pines, and to Abaco, Acklin, and Great Inagua in
the Bahamas. Its former Bahaman range was more extensive, as it
occurred within recent years on Long and Fortune islands (Bond,
1956) and in pre-Columbian time on Crooked Island (Wetmore, 1938).
This is the first Pleistocene record.

Vol. 4




Tyto pollens Wetmore
Plate T, figs. 5-6
Tyto pollens Wetmore, 1937B, p. 436, fig. 10-16 (Pleistocene, Great Exuma Island).
REFERRED MATERIAL. Distal portion of left tibiotarsus, UFC 3195;
left tarsometatarsus, UFC 3196; proximal portion of right tarsometa-
tarsus, apparently from another individual, UFC 3197; inner trochlea
of right tarsometatarsus, UFC 3198; outer trochlea of right tarsometa-
tarsus, UFC 3199.
DESCRIPTION. Measurements of these specimens are given in table
2. They indicate that while Tyto pollens was a very robust bird, with
the widths of the bones almost twice those of living Tyto alba pratin-
cola, the tarsometatarsus was relatively short and exceeded that of
the modern barn owl by only about 15 percent.


Element T. pollens T. ostologa T. alba pratincole (9)

Width through trochleac 16.7 9.9-11.0
Least width of shaft 7.8 4.8- 5.4
Length 93.5 75.4-81.9
Proximal width 17.7-18.2 17.5 10.0-11.1
Least width of shaft 8.6 4.5- 4.9
Width through trochleae 21.2 12.1-13.0
Width of outer trochlea 14.9, 14.9 13.5 7.8- 8.7
Width of inner trochlea 12.3-13.2 11.7 6.9- 7.5

DisTRnsUTOn. Previous knowledge of giant Pleistocene barn owls
is due to Wetmore (1922, 1937B, 1959), who reported Tyto ostologa
from Haiti, Tyto pollens from Great Exuma, and an unnamed form
from Cuba. These tremendous birds, which must have equaled or
exceeded in bulk any owl now living, were probably derived from
the same stock and apparently were contemporaries during the Pleis-
tocene. It is quite possible that they had differentiated from each
other only to the subspecific level.


The remains of these owls are associated with those of large
extinct rodents, which seem to have been their principal food.
Whether the giant stock ever extended to Puerto Rico is not
known. The Quaternary Tyto cavatica Wetmore (1920, 1922A) of
Puerto Rico was a small bird. From other parts of the world Pleisto-
cene barn owls have also been described, viz.: Tyto melitensis (Lydek-
ker, 1891) from Malta, Tyto sauzieri (Newton and Gadow, 1893) from
Mauritius, and Tyto newtoni (Rothschild, 1907) likewise from Mauri-
tius. These too were small and resembled modern barn owls in size.

Tyto alba (Scopoli)
REFERRED .MATERIAL. Premaxillary, UFC 3200; left tarsometatar-
sus, lacking both ends, UFC 3201. The premaxillary is from a young
but fully grown bird.
DISTRIBUTION. The Barn Owl is of almost cosmopolitan distribu-
tion and occurs today on New Providence, as well as others of the
Bahama Islands and through the West Indies. It has been recorded
from the Pleistocene of the United States, Mexico, Brazil, and Europe.
As the remains of 2 individuals were associated with the 2 indi-
viduals of the giant Tyto pollens, it appears that the species were
contemporaneous and sympatric.

Glaucidium dickinsoni, new species
Plate II, fig. 1
HOLOTYPE. Distal half of left tibiotarsus, UFC 3202. From Pleis-
tocene at Banana Hole, New Providence Island, Bahamas. Collected
by J. C. Dickinson, Jr., and Walter Auffenberg, 28 August 1958.
DIAGNOSIs. Agrees with modern Glancidium siiu (d'Orbigny) of
Cuba in having intercondylar sulcus wide; tendinal groove forming
a very deep fossa at its distal end, where encroaching under inter-
condylar bar. Differs in having intercondylar bar oblique rather than
transverse; peroneal groove reduced, not forming a shelf; internal
prominence for oblique ligament on anterior face of shaft, instead of

Plate II. Fig. 1, Glaucidium dickinsoni n. sp., holotype tibiotarsus, UFC
3202, X 1.7. Fig. 2-6, Otus providentiac n. sp., holotype tibiotarsus, UFC 3207,
X 1.7; coracoid, UFC 3203, X 2.2; earpounetacarpus, UFC 3205, X 1.9; tibiotarsus,
UFC 3206, X 1.9; ulna, UFC 3204, X 2.3. Fig. 7, Bathocleus hyphalos n. g.
and sp., holotype coracoid, UFC 3209, X 1.7.

Vol. 4









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protruding medially; shaft only slightly expanded distally. Size some-
what smaller. Distal width, 5.0 mm (5.5 in G. siju); width of shaft
through internal prominence for oblique ligament, 2.6 (3.2 in G.
siju); width of shaft above prominence, 2.4 (2.9 in G. siju).
ETYMOLocY. Named for the collector, J. C. Dickinson, Jr., Associ-
ate Professor of Biology at the University of Florida and Acting Di-
rector of the Florida State Museum.
DisTRmUTnON. This genus is represented in the modern fauna of
the West Indies only by G. siju of Cuba and the Isle of Pines, although
other species occur in the western United States, Central and South
America, and the Old World. No fossil species have previously
been described in this genus.

Otus providentiae, new species
Plate II, figs. 2-6
HOLOTYPE. Distal portion of left tibiotarsus, UFC 3207. From
Pleistocene at Banana Hole, New Providence Island, Bahamas. Col-
lected by J. C. Dickinson, Jr., and Walter Auffenberg, 28 August
DIAGNOSIS. Agrees with Otus Pennant and differs from Glaucid-
ium Boie in having intercondylar sulcus narrow; tendinal groove form-
ing a shallow fossa at its distal end, with a pit undercutting medial
end only of intercondylar bar.
Differs from modern Otus asio floridanus (Ridgway) of Florida in
having intercondylar bar straight, not undulating; tendinal pit under
medial end of bar deeper; shaft only slightly expanded distally, with
margins gently flaring, not convex.
Differs from modern Otus nudipes (Daudin) of Puerto Rico as from
Otus asio; in addition, peroneal groove obsolete, without raised edges;
tendinal fossa without a knob above external condyle; intercondylar
fossa only slightly excavating sides of condyles.
Size small. Distal width, 5.2 mm. (5.5-6.3 in floridanus, 6.0 in
nudipes); width of shaft through internal prominence for oblique lig-
ament, 2.8 (2.5-3.1 in floridanus, 3.4 in nudipes).
ETYiMOLOY. Latin, providentiae, of Providence.
REFERUr ) MATERIAL. Proximal three-fourths of left tibiotarsus of
another individual, UFC 3206; left coracoid, lacking upper end, UFC
3203; proximal portion of right ulna, UFC 3204; proximal portion of
left carpometacarpus, UFC 3205.

Vol. 4


Proximal portion of tibiotarsus with a foramen on posterior side
of shaft between fibular ridge and head foramenn present in O. asio,
absent in 0. nudipes); flexor attachment distal to foramen forming
a pronounced, laterally deflected ridge (ridge small in O. asio, obso-
lete in 0. nudipes); size small, but shaft not narrowing near middle.
Proximal width, 4.8 mm. (4.7-5.1 in floridanus, 5.8 in nudipes); least
width of shaft, 2.4 (2.2-2.5 in floridanus, 2.7 in nudipes).
Coracoid agrees with that of Otus and differs from that of
Glaucidium in having anterior face of shaft perpendicular to procora-
coid, rather than inclined toward it; posterior surface of procoracoid
curved toward head, not flat; procoracoid foramen large and high,
near scapular facet. Resembles Otus nudipes rather than Otus asio
in having procoracoid narrow, rising toward head just medial to fora-
men and thus leaving a narrower groove along anterior face of bone.
Size small: length from scapular facet, 16.1 mm. (17.0-17.7 in flori-
danus, 17.3 in nudipes); sternal width, 6.0 (6.1-7.0 in floridanus, 7.3
in nudipes).
Ulna resembles that of 0. nudipes in having area for bicipital at-
tachment more rugose and closer to head than in 0. asio; differs
from both those species in having prominence for anterior articular
ligament rounded instead of angular. Proximal width, 4.9 mm. (4.8-
5.1 in floridanus, 5.5 in nudipes).
Carpometacarpus with intermetacarpal tuberosity smoothly round-
ed as in Otus (knoblike in Glaucidium). Resembles Otus asio in hav-
ing posterior carpal fossa elongate and shallow (round and deep in
Otus nudipes). Differs from both species of Otus in having process
of metacarpal I more upright, less bent back. Height through meta-
carpal I, 5.5 (5.0-5.7 in floridanus, 6.0 in nudipes).
Detailed comparison with Otus lawrencii (Sclater and Salvin) of
Cuba is not possible at this time, as skeletal material of that species
is unavailable. However, 0. lawrencii resembles 0. asio in size, and
thus the New Providence bird is smaller.
DISTRIBUTION. The genus has an almost world-wide distribution,
but the modem fauna of the West Indies contains only two species,
Otus nudipes (Daudin) of Puerto Rico and the Virgin Islands and Ottus
lawrencii (Sclater and Salvin) of Cuba and the Isle of Pines. Because
of their bare legs and lack of ear tufts, these two species arc often
placed in a separate genus, Gymnasio Bonaparte. In most characters
the New Providence screech owl is closer to true Otus than to Gym-
nasio. No fossil species of Otus were previously known.



Family PICmAE
Bathoceleus, new genus
TYPE OF GENUS. Bathoceleus hyphalus, new species.
DIAGNOSIS. Coracoid with internal margin of upper end of shaft
nearly straight (as in Xiphidiopicus; curved medially toward brachial
tuberosity in Colaptes, Nesoceleus, and Melanerpes); pneumatic foram-
ina not extending from region of brachial tuberosity onto upper end of
posterior face of shaft (resembling Nesoceleus and Melanerpes; area
pneumatic in Colaptes and Xiphidiopicus); a pneumatic foramen be-
tween scapular facet and procoracoid foramenn present in Colaptes,
Nesoceleus, and Xiphidiopicus; absent in Melanerpes); lower part of
shaft slightly compressed in medial view (as in Melanerpes and Xiphid-
iopicus; expanded in Colaptes and Nesoceleus); lower part of shaft
narrowly V-shaped in lateral view (concave on posterior face in allied
genera); scar of coracobrachialis longer and shallower than in allied
genera; a pneumatic foramen on posterior side above internal distal
angle (as in Nesoceleus; foramen absent in Colaptes, Melanerpes, and
Xiphidiopicus); anterior sternal facet curved (as in Xiphidiopicus;
straight in Colaptes, Nesoceleus, and Melanerpes).

Bathoceleus hyphalus, new species
Plate II, fig. 7
HOLOTYPE. Right coracoid, nearly complete, UFC 3209. From
Pleistocene at Banana Hole, New Providence Island, Bahamas. Col-
lected by J. C. Dickinson, Jr., and Walter Auffenberg, 28 August 1958.
DIAGNOSIS. Differs in generic characters from other West Indian
woodpeckers. Smaller than living Colaptes chrysocaulosus Gundlach
and Nesoceleus fernandinae (Vigors) of Cuba; larger than living Mel-
anerpes striatus (Miiller) of Hispaniola, Melanerpes superciliaris (Tem-
minck) of Cuba and the Bahamas, and Xiphidiopicus percussus (Tem-
minck) of Cuba. Length, 26.0+ mm.; least width of shaft, 2.0; width
of sternal facet, 5.6.
ETY OLOGY. Greek, bathos, deep, and keleos, masculine, wood-
pecker; Greek, huphalos, under the sea, in allusion to the former
larger extent of the Bahamas.
DisTRIUTrrio. The 3 previously described fossil woodpeckers are
from the Oligocene and Miocene of France.

Vol. 4


Melanerpes superciliaris (Temminck)
REFERRED MATERIAL. Distal portion of right humerus, UFC 3208.
MEASUREMENTS. Distal width of humerus, 7.6 mm., identical in
size with a Recent specimen from Cuba and much larger than Mel-
anerepes striatus from Hispaniola.
DlsnRBUTION. The present distribution of this species is irregu-
lar. It occurs in the Bahamas on Grand Bahama and Abaco at the
northern end of the chain, on Watlings Island in the eastern part of
the Bahamas, and then reappears on Cuba, the Isle of Pines, and
Grand Cayman. Each of these islands has a distinct race, so it is
quite possible that more adequate material may prove the New
Providence bird to be separable. Wetmore (1937B) reported the
species from the Pleistocene of Great Exuma.

Corvus wetmorei, new species
Plate III, figs. 1-9
[?] Corvus nasicus Temminck, Wetmore, 1937B, p. 440 (Pleistocene, Great Exuina).
HOLOTYPE. Left humerus, lacking proximal end, UFC 3210. From
Pleistocene at Banana Hole, New Providence Island, Bahamas. Col-
lected by J. C. Dickinson, Jr., and Walter Auffenberg, 28 August 1958.
DIAGNosis. Resembles living Corvus lencognaphalus Daudin of
Cuba, Hispaniola, and Puerto Rico, but smaller (see table 3); entepi-
condyle small through reduction of area of origin of both heads of
flexor carpi ulnaris, resulting in its internal border being nearly
straight instead of produced internally, and further in being produced
distally for a shorter distance at a blunt rather than sharp angle (1250
instead of 100').
REFERRED MATERIAL. Proximal end of left humerus, possibly from
the same individual as the type, UFC 3211; left ulna, lacking proximal
end, UFC 3212; right carpometacarpus, with both ends broken, UFC
8213; 2 left femora, complete and proximal portion, UFC 3214-3215;
distal portion of left tibiotarsus, UFC 3216; proximal portion of left
tarsometatarsus, UFC 3217; unguis, UFC 3218.
ETYMOLOGY. Named for Alexander Wetmore, who is responsible
for all previous knowledge of fossil birds of the West Indies.



4 '

7. 9

Plate III. Fig. 1-9, Corvus wetmorei n. sp., proximal portion of humerus,
UFC 3211, X 1.1; holotype humerus, UFC 3210, X 1.2; ulna, UFC 3212, X 0.9;
carpometacarpus, UFC 3213, X 1.1; femora, UFC 3214-3215, X 1.1; tibiotarsus,
UFC 3216, X 1.3; unguis, UFC 3218, X 1; tarsometatarsus, UFC 3217, X 1.1.

DISTRBUTION. The crow bones from the Pleistocene of Great
Exuma (Wetmore, 1937B) and from the prehistoric site on Crooked
Island (Wetmore, 1938) should be restudied, since at that time skeletal
material of C. 1. nasicus was unavailable.
At present two size-classes of crows occur in the West Indies. The
larger species, Corvus leucognaphalus, is represented by two sub-
species. Corvus leucognaphalus nasicus inhabits Cuba, the Isle of
Pines, and Grand Caicos in the southern Bahamas. Corvus leucog-
naphalus leucognaphalus occurs on Hispaniola and Puerto Rico, and
it extended to St. Croix during pre-Columbian time (Wetmore, 1938).
The smaller size-class is represented in the modern fauna by
Corvus palmarum of Cuba and Hispaniola and by the aberrant Cor-

Vol. 4


ris jamaicensis of Jamaica. Corvus pumilus Wetmore (1920, 1922A)
was the Quaternary representative of the small crow on Puerto Rico.


Element C. wetmorei C. 1. nasicus (2) C. i. leucognaphalus (5)

Proximal width 16.1 16.9-18.7
Width of shaft 5.0 5.9- 6.3
Distal width 12.8 14.0 14.1-15.4
Depth through
condyles 7.6 8.8 9.0- 9.9
Mid-shaft width 3.8 3.9 4.3- 4.7
Posterior carpal
fossa to end of
space 27.0 26.4 28.4-29.8
Width of shaft of
nmtacarpal II 3.6 3.4 3.6- 4.0
Length 44.2 50.4-52.6
Width through
head 8.7-8.8 10.9-11.4
Least width
of shaft 3.3-.9 -- 4.3- 4.7
Distal width 9.3 plus -- 10.2-10.8
Distal width 7.8 8.5- 9.1 8.3- 9.5
Proximal width 8.7 10.1 8.9-10.1


Mimus gundlachii Cabanis

REFERRED MATERIAL. Proximal half of right humerus, UFC 3219;
distal portion of right tarsometatarsus, UFC 3220.
DESCRIPTION. In the humerus of Mimus the deltoid crest is angu-
lar, and the medial bar fuses to the side of the shaft rather than reach-


ing the floor of the tricipital fossa. Mimus gundlachii is larger than
Mimius polyglottos. The tarsometatarsus of M1. gundlachii has the
inner trochlea lengthened, so that it is decidedly longer than the outer
trochica and nearly as long as the middle one. In Af. polyglottos the
inner and outer trochleae are about equal in length, and both are
distinctly shorter than the middle one.
DISTRIBUTION. MimUs gundlachii has a relict distribution today,
in the Bahamas exceptt the northernmost islands), on cays off the
north coast of Cuba, and in southern Jamaica. It has been recorded
from the pre-Columbian site on Crooked Island (Wetmore, 1938), but
this is the first fossil record.
Other species of mockingbirds occur widely in the West Indies
and elsewhere in the Americas.

The present avifauna of the Bahamas contains 93 species of breed-
ing birds, besides winter visitants, transients, and stragglers (data
summarized from Bond, 1956). Endemism is mainly at the subspecific
level, as only 4 species of birds (Philodice evelynae, Callichelidon
cyaneoviridis, Vireo crassirostris, and Geothlypis rostrata) are confined
to the Bahamas as breeders, and only one of these (Callichelidon) has
differentiated generically.
The relationships of the present Bahaman avifauna are primarily
to the Greater Antilles and secondarily to Florida. Of the 93 resident
species no less than 77 percent breed also on Cuba, and 74 percent on
Hispaniola. While the Bahamas share 61 percent of their breeding
birds with Florida, these are mainly species of wide distribution, and
only 5 species (Pandion haliaetus, Dendrocopos villosus, Sitta pusilla,
Polioptila carulea, and Dendroica dominica) breed on the Bahamas
and in Florida but not on Cuba or Hispaniola.

The adjacent areas have richer avifaunas than the Bahamas. Flor-
ida has 141 breeding species of birds (Howell, 1982), and its avifauna
is thus 50 percent larger than that of the Bahamas. Cuba has 133
breeding species and Hispaniola 123 (Bond, 1956), and their avifaunas
are thus respectively 40 and 30 percent larger than that of the Ba-
hamas. Physical factors which might account for these differences
include separation from the mainland, absence of mountains, and
a small land mass.


The Bahamas share with Cuba and Hispaniola a water barrier
separating them from the mainland. Although the Bahamas in gen-
eral are closer to the mainland than is Cuba, yet they have fewer
resident species in common with Florida. for only 57 Florida species
occur in the Bahamas, against 61 Florida species that reach Cuba.
As the water barrier should have operated more effectively against
Cuba than against the Bahamas, some other reason must be found
to explain the paucity of the Bahaman avifauna.
The absence of mountains in the Bahamas undoubtedly reduces
the number of habitats available in comparison with Hispaniola, and
to a lesser extent with Cuba. Yet the highest elevation in Florida is
324 feet, less than that of Cat Island in the Bahamas, which reaches
about 400 feet, and the greater part of Cuba lies below the 500-foot
contour. Thus differences in elevation alone appear insufficient to
account for the poverty of the present Bahaman fauna.
The small land area of the Bahamas forms the most striking physi-
cal difference between that group and all three of the adjacent re-
gions. Although the Bahamas stretch over a distance comparable
to Cuba or Florida and greater than Hispaniola, their actual land
area of 4404 square miles is only 8 percent that of Florida, 10 percent
that of Cuba, and 14 percent that of Hispaniola. As this total land
area is further broken up into numerous small islands, it becomes ap-
parent that the absence of a large land mass is a major factor in re-
stricting the size of the avifauna.

Of the 15 species identified from the New Providence deposit.
only 4 still occur on that island, namely: Columba leucocephala,
Zenaida aurita, Tyto alba, and Mimuis gundlachii. Only 2 additional
species, Amazona leucocephala and Melanerpes superciliaris, are
found at present elsewhere in the Bahamas. While only 2 of these
6 species extend to Florida, every one of them is represented in the
present avifauna of Cuba, and 4 of them also occur on Hispaniola.
Columba squamosa still occurs on Cuba and Hispaniola, but is
now absent from the Bahamas.
The remaining 8 species are extinct. As with the extant birds,
their relationships are primarily with Cuba and secondarily with His-
paniola. Calohierax quadratus and Bathoceleus hyphalus had differ-
entiated to the generic level, and their relationships are doubtful.
Caracara creightoni has a generic representative on modern Cuba, had


another in the Quaternary of Puerto Rico, and others in the Pleisto-
cene and Recent faunas of Florida. Burhinus nanus is related to a
species on Hispaniola, the only place in the West Indies where the
family now occurs. Tyto pollens is allied to the Pleistocene barn owls
of Cuba and Hispaniola. Glaucidium dickinsoni was the Pleistocene
representative of a modern owl on Cuba. Otus providentiae has
generic representatives in the present fauna of Florida, Cuba, and
Puerto Rico. Corvus wetmorei is related to crows now living on Cuba,
Hispaniola, and Puerto Rico.
Thus of the 15 species identified from the Pleistocene of New
Providence, 80 percent are related to Cuban forms, 53 percent are
related to Hispaniolan forms, but only 27 percent occur in or were
derived from Florida. Moreover, all 4 species that are related to the
Florida ornis are likewise represented on Cuba, and 2 of them on
If the birds from the Pleistocene of Great Exuma are added to
those from New Providence, the combined list of 21 species of resi-
dent birds does not change the picture materially. The affinities of
71 percent of them are with Cuba, 62 percent with Hispaniola, but
only 24 percent with Florida. All the species in the Florida group
are likewise represented in the Cuban or Hispaniolan groups.
Thus during the Pleistocene the faunal tie of the Bahamas to the
Greater Antilles, particularly Cuba, was strong, while the relationship
to Florida was weaker than today.

Nine of the 21 resident Pleistocene birds identified from the Ba-
hamas are extinct. A 43 percent extinction rate is higher than re-
ported for other Pleistocene localities (Brodkorb, 1955). All the ex-
tinct species represent genera or ecological types absent from the
islands today. It is thus logical to conclude that the Pleistocene avi-
fauna of the Bahamas was about 40 percent richer than the present
depauperate fauna and comparable in numbers to the present avi-
faunas of Cuba or Hispaniola.

The absence of a large land mass is thought to be of major im-
portance in restricting the size of the present depauperate avifauna
of the Bahamas. The richer Pleistocene avifauna would seem to re-
quire a larger land mass than presently exists in the Bahamas, and

Vol. 4


this would also seem to be needed to support giant raptorial birds,
such as Titanohierax gloveralleni and Tyto pollens, and the plentiful
large rodents on which they fed.
There were periodic rises in sea level during the interglacial stages
of the Pleistocene and drops from present level during the glacial
stages (Cooke, 1932, 1939, 1945; Flint, 1942, 1947; MacNeil, 1950).
The Bahamas have been a stable, reef-building area since the Creta-
ceous (Eardley, 1951). Therefore in the absence of evidence for
movement of the land, the requirement of a larger land mass can be
met only during one of the glacial stages.
During the Sangamon (third) interglacial stage the sea rose to a
height of 90-100 feet above its present level (authors cited above).
Such a rise would inundate the fossil locality on New Providence, but
no marine materials were recognized. It is thus concluded that the
sink hole at the fossil locality postdates Sangamon time.
According to Cooke, the sea dropped to 60 feet below its present
level during the early portion of the Wisconsin (fourth) glacial stage
and preceding the mid-Wisconsin (Pamlico) temporary rise in sea
level. Such a recession of the sea to the 10-fathom mark would ex-
pose the Great Bahama Bank, uniting it in one large island with
Bimini, Andros, New Providence, Great Exuma, and Long Island
(see map). This large island lay within 10 miles of Cuba. It was
thus easily accessible to Cuban species, and other islands stretching
toward the southeast provided an indirect connection to Hispaniola.
Therefore, in view of the geological and faunal evidence, the fossil
deposit is thought to represent early Wisconsin (pre-Pamlico) time.

For the use of comparative material I am indebted to Dr. Dean
Amadon of the American Museum of Natural History, Dr. Julian J.
Baumel of Creighton University. and Drs. Herbert Friedmann and
Alexander Wetmore of the United States National Museum. Clayton
Ray of Harvard University supplied casts of the types of birds de-
scribed from Great Exuma. The photographs were taken by J. Hill
Hamon of the University of Florida.

American Ornithologists' Union
1957. Check-list of North American birds. Fifth edition. Lord Baltimore
Press, Baltimore, xiii + 691.


Bond, James
1956. Check-list of birds of the West Indies. Fourth edition. Academy of
Natural Sciences of Philadelphia, Lancaster, Pa., ix + 214.

Brodkorb, Pierce
1955. The avifauna of the Bone Valley formation. Florida Geol. Surv., Re-
port Invest., no. 14: 1-57, 11 pl.

Cooke, C. 'Wythe
1932. Tentative correlation of American glacial chronology with the marine
time scale. Jour. Wash. Acad. Sci., 22: 310-312.
1939. Scenery of Florida interpreted by a geologist. Bull. Florida Geol. Surv.,
17: 1-118, 58 figs.
1945. Geology of Florida. Bull. Florida Geol. Surv., 29: 1-339, 47 figs., map.

Eardley, A. J.
1951. Structural geology of North America. Harper & Bros., New York,
624, illus.

Flint, R. F.
1942. Atlantic coastal "terraces." Jour. Wash. Acad. Sci., 32: 235-237.
1947. Glacial geology and the Pleistocene epoch. John Wiley & Sons, New
York, xviii + 589, illus.

Hellmayr, Charles E., and Boardman Conover
1949. Catalogue of birds of the Americas. Field Mus. Nat. Listt, zool. ser.,
13, part 1, no. 4: i-vi, 1-358.

Howard, Hildegarde
1938. The Rancho La Brea caracara: a new species. Carnegie Inst. Wash.
Publ. 487: 217-240, 3 pls.
1940. A new race of caracara from the Pleistocene of Mexico. Condor, 42:

Howell, Arthur H.
1932. Florida bird life. Coward-McCann, Inc., New York, xxiv + 579,
58 pl., 72 text-figs.

Lydekkcr, Richard
1891. Catalogue of the fossil birds in the British Museum. Taylor and Fran-
cis, London, xxvii + 368, 75 figs.

MacNeil, F. Stearns
1950. Pleistocene shore lines in Florida and Georgia. U. S. Geol. Surv. Prof.
Paper 221-F: 95-107, pl. 20-25, map.

Merrem, Blasius
1826. Caracara. In Ersch and Gruber, Allgemeine EncyclopAdie der Wissen-
schaft und Kiinste, 15: 159.

Vol. 4


Newton, Edward, and Hans Gadow
1893. On additional bones of the dodo and other extinct birds of Mauritius,
obtained by Mr. Th6odorc de Sauzier. Trans. Zool. Soc. London, 13:
282-288, pl. 33.
Rothschild, Walter
1907. Extinct birds. Hutchinson & Co., London, xxix + 244, 49 pl.
Wetmore, Alexander
1920. Five new species of birds from cave deposits in Porto Rico. Proc.
Biol. Soc. Wash., 33: 77-81.
1922a. Bird remains from the caves of Purto Rico. Bull. Am. Mus. Nat. Hist.,
46: 297-333, 25 figs.
1922b. Remains of birds from caves in the Republic of Haiti. Smiths. Misc.
Coll., 74, no. 41: 1-4, 2 figs.
1937a. Ancient records of birds from the island of St. Croix with observations
on extinct and living birds of Puerto Rico. Jour. Agr. Univ. Puerto Rico,
21:5-16, pl. 1.
1937b. Bird remains from cave deposits on Great Exmna Island in the Bahamas.
Bull. Mus. Comp. Zool., 80: 427-441, 1 pl., 16 figs.
1938. Bird remains from the West Indies. Auk, 55: 51-55.
1952. A record for the black-capped petrel, Pterodroma hasiata, in Martinique.
Auk, 69: 460.
1959. Birds of the Pleistocene in North America. Smiths. Misc. Coll., 138,
no. 4: 1-24.

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