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Group Title: Bulletin of the Florida Museum of Natural History
Title: Doves (Columbidae) and Cuckoos (Cuclidae) from the early Miocene of Florida /
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Title: Doves (Columbidae) and Cuckoos (Cuclidae) from the early Miocene of Florida /
Physical Description: p. 1-16 : ill. ; 28 cm.
Language: English
Creator: Steadman, David W
Publisher: Florida Museum of Natural History, University of Florida
Place of Publication: Gainesville, FL
Publication Date: 2008
Copyright Date: 2008
 Subjects
Subject: Birds, Fossil -- Florida   ( lcsh )
Paleontology -- Florida -- Miocene   ( lcsh )
Pigeons -- Florida   ( lcsh )
Cuckoos -- Florida   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Abstract: A new genus, Arenicolumba (Aves: Columbiformes: Columbidae), is proposed for the fossil species Columbina prattae Becker and Brodkorb, 1992. This small dove is known from many hundreds of fossils, representing all major post-cranial skeletal elements, from the early Miocene (ca. 18.5 Ma; Hemingfordian land mammal age) Thomas Farm local fauna, Gilchrist County, Florida. Except for the single humerus of Gerandia from the Aquitanian (early Miocene; ca. 22 Ma) of France, Arenicolumba prattae is the oldest known member of the family Columbidae. The nearest living relatives of arenicolumba appear to be the African genera Oena and Turtur. Thomasococcyx philohippus, new genus and species (Aves: Cuculiformes: Cuculidae), is also described from the Thomas Farm local fauna. Neotropical ground-dwelling cuckoos in the genera Morococcyx, Tapera, Geococcyx, and Dromoccyx may be the closest living relatives of Thomasococcyx. Along with Cursoricoccyx from the Hemingfordian of Colorado, Thomascoccyx philohippus provides evidence that the subfamily of New World ground-dwelling cuckoos (Neomorphinae) existed by the early Miocene.
Bibliography: Includes bibliographical references (P. 15-16).
General Note: Cover title.
General Note: Bulletin of the Florida Museum of Natural History ; vol. 48, no. 1, pp. 1-16
Statement of Responsibility: David W. Steadman.
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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: oclc - 248615735
issn - 0071-6154 ;

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'FLORIDA
__MUSEUM
OF NATURAL HISTORYTM


BULLETIN


DOVES (COLUMBIDAE) AND CUCKOOS (CUCLIDAE) FROM
THE EARLY MIOCENE OF FLORIDA

David W. Steadman


Vol. 48, No. 1, pp. 1-16


2008


UNIVERSITY OF FLORIDA


GAINESVILLE











DOVES (COLUMBIDAE) AND CUCKOOS (CUCULIDAE) FROM THE EARLY
MIOCENE OF FLORIDA






David W. Steadman'



ABSTRACT

A new genus, Arenicolumba (Aves: Columbiformes: Columbidae), is proposed for the fossil species Columbina prattae Becker and Brodkorb,
1992. This small dove is known from many hundreds of fossils, representing all major post-cranial skeletal elements, from the early Miocene
(ca. 18.5 Ma; Hemingfordian land mammal age) Thomas Farm local fauna, Gilchrist County, Florida. Except for the single humerus of Gerandia
from the Aquitanian (early Miocene; ca. 22 Ma) of France, Arenicolumba prattae is the oldest known member of the family Columbidae. The
nearest living relatives of Arenicolumba appear to be the African genera Oena and Turtur. Thomasococcyxphilohippus, new genus and species
(Aves: Cuculiformes: Cuculidae), is also described from the Thomas Farm local fauna. Neotropical ground-dwelling cuckoos in the genera
Morococcyx, Tapera, Geococcyx, and Dromococcyx may be the closest living relatives of Thomasococcyx. Along with Cursoricoccyx from the
Hemingfordian of Colorado, Thomasococcyx philohippus provides evidence that the subfamily of New World ground-dwelling cuckoos
(Neomorphinae) existed by the early Miocene.


Key Words: Florida; Thomas Farm; early Miocene fossils; Hemingfordian land mammal age; Columbidae; Cuculidae.







TABLE OF CONTENTS

Introdu action ................................................... ........................................... . . 2
M materials and M ethods..................................................................... .................2...
Sy stem atic P aleontology ................................................... ........................ 3
Order Columbiformes Family Columbridae....................................... 3
A renicolum ba n gen .......................................... ................. 3
Order Cuculiform es Family Cuculidae........................ ...................5...
Thomascoccyxphilohippus n. gen. & sp............................... 5
Cuculidae, gen. uncertain................................. ................. 10
D isc u s sio n .............................................................................................................. 1 0
C o lu m b ifo rm e s ............................................... ... ....................... ................ 10
C uculiform es ................................................................................ .................13
A cknow ledgem ents .......................................... ... ..... ................ .... ............ 14
L literature C ited ....................... .................................................. ................ 15












'Florida Museum of Natural History, Dickinson Hall, PO Box 117800, University of Florida, Gainesville, Florida 32611-7800
Steadman, D.W. 2008. Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida. Florida Museum. Nat. Hist. Bull. 48(1):1-










INTRODUCTION
The Thomas Farm site (NAD 27; 29.860N, 82.830W,
elev. 11 m) is a highly fossiliferous, sediment-filled sink-
hole in Gilchrist County, Florida (Pratt 1990). The rich
mammalian fauna at Thomas Farm indicates a
Hemingfordian land mammal age (LMA) in the early
Miocene Epoch, approximately 18.5 million years old
(Hulbert 2001, MacFadden 2001). Avian fossils from
Thomas Farm include at least 29 species in 25 genera
and 18 families (DWS personal observation), although
only seven of these species have been described until
now. Five of the described species have been assigned
to extinct genera, namely the accipitrids Promilio
floridanus, P. epileus, and P brodkorbi, the cracid
Boreortalis laesslei, and the phasianid Rhegminornis
calobates (Wetmore 1943, 1958, Brodkorb 1954, Olson
& Farrand 1974, Steadman 1980, Becker 1987). An
apparent anhinga, originally described as a cormorant
Phalacrocorax subvolans by Brodkorb (1956), was
assigned to the living genus Anhinga but without details
by Becker (1987).
The seventh described species from Thomas Farm
is a small columbid, Columbina prattae Becker and
Brodkorb (1992). This dove is by far the most abundant
species of bird at Thomas Farm. Assigning Columbina
prattae to an extant genus sets this species apart from
the rest of the avifauna, with the exception of Anhinga
subvolans. In this paper I re-examine the osteology of
C. prattae, and conclude that this dove is like most other
birds (and all mammals; Hulbert 2001, MacFadden 2001)
from Thomas Farm in being generically distinct from
living species.
I also describe a new genus and species of cuckoo
(Cuculidae), a family previously unreported from Tho-
mas Farm and with a limited fossil record in North
America.

MATERIALS AND METHODS
The fossils from Thomas Farm are housed in the Divi-
sion of Vertebrate Paleontology, Florida Museum of
Natural History, University of Florida. Certain of the
fossils, formerly in the private collection of Pierce
Brodkorb, have the catalogue prefix UF/PB. Some of
the modem skeletons also were originally in the private
collection of Pierce Brodkorb; these specimens have a
PB catalogue number and a UF catalogue number as-
signed in the 1990s. Both the UF and PB numbers are
listed for these specimens in the figure legends. UF is
used as a prefix for both the fossils housed in the Verte-
brate Paleontology Collection and for modem skeletons
housed in the Ornithology Collection.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(1)

I compared the fossils with the following modem
UF skeletons, supplemented by specimens from the
University of Kansas Natural History Museum (KU),
and the Louisiana State University Museum of Zoology
(LSUMZ) Columbina passerina UF 40074, 43317,
C. talpacoti UF 41837, C. buckleyi UF 41837, C. picui
UF 38965, C. minute UF 38786, C. cruziana LSUMZ
48726, 81201, Claravispretiosa UF 38787, Scardafella
inca UF 40806, Metriopelia melanoptera UF 39346,
M ceciliae UF 38788, Geotrygon montana UF 33846,
G chrysia UF 40006, Zenaida macroura UF 40939, Z
aurita UF 40007, Z asiatica LSUMZ 89696, Leptotila
verreauxi UF 41097, L. jamaicensis UF 40027, and
Patagioenas [formerly Columba] leucocephala UF
42212. In addition to these 18 species from the Ameri-
cas, I also examined modem skeletons of seven Old
World species of Columbidae Oena capensis UF
38785, Turtur chalcospilos UF 38782, Streptopelia
capicola UF 38781, Columba guinea UF 38769,
Ducula pistrinaria UF 40186, Geopelia cuneata UF
32379, and Gallicolumba jobiensis UF 41460.
For the Cuculidae (sequence and subfamilial clas-
sification follow Payne 2005), I examined modem skel-
etons of these New World species: Crotophaginae -
Guira guira UF 38969, Crotophaga major KU 32390,
C. ani UF 40013, 42473, C. sulcirostris UF 33855,
40789; Neomorphinae Tapera naevia UF 38735,
Dromococcyx phasianellus KU 91388, D. pavoninus
LSUMZ 101257, Morococcyx ihp.'-i,, UF 16796,
26250, 26251, 38737, Geococcyx californianus UF
26238, G velox UF 38729, Neomorphus rufipennis
KU 86604, N. geoffroyi LSUMZ 106946; Cuculinae
Coccycua (Piaya) minute UF 26232, Piaya cayana
UF 40787, P melanogaster KU 88663, Coccyzus
americanus UF 40346, C. minor UF 40029, Coccyzus
(Hyetornis) pluvialis UF 26233, and Coccyzus
(Saurothera) merlini UF 42489. Skeletons of these Old
World species also were examined: Centropodinae -
Centropus superciliosus UF 33856; Cuculinae -
Zanclostomus (Phaenicophaeus) javanicus UF 42704,
Phaenicophaeus (Rhopodytes) viridirostris UF 26237,
Clamator jacobinus UF 38730, Eudynamys
scolopacea UF 41473, Chrysococcyx lucidus UF
39450, and Cuculus canorus UF 35289. For the
Musophagidae, I examined skeletons of Corythaeola
cristata UF 38728, Corythaixoides concolor UF 38719,
Crinifer piscator UF 38718, Musophaga rossae UF
38727, and Tauraco corythaix UF 38726.
Measurements were taken with digital calipers,
read to the nearest 0.01 mm. Specimens were exam-
ined with a Leica MZ8 stereomicroscope, and photo-








STEADMAN: Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida


graphed with a Sony DSC-R1 camera. Osteological ter-
minology is from Howard (1929) and Baumel et al.
(1993). Character polarities were not proposed because
of a lack of strong, consistent evidence of which fami-
lies would be appropriate as outgroups for the Columbidae
or Cuculidae.

SYSTEMATIC PALEONTOLOGY
Order Columbiformes (Latham 1790)
Family Columbidae (Illiger 1811)
Six of the seven characters (based on three skel-
etal elements) that Becker and Brodkorb (1992) used to
assign the species prattae to the genus Columbina are,
in fact, found in many living genera of Columbidae world-
wide. Their seventh character ["the dorsal surface of
trochlea IV extending dorsad (in distal view) past the
level of trochlea III"] does distinguish the tarsometatar-
sus of Columbina sensu lato (i.e., including Claravis,
Scardafella, and Metriopelia) from that of Geotrygon,
Zenaida, Leptotila, and Patagioenas but not from that
in Old World genera such as Turtur and Ducula.


In the diagnosis of the new genus, I will begin with
the tarsometatarsus since that is the holotype (UF
106594) of Columbina prattae as well as the skeletal
element with the largest number of diagnostic charac-
ters.
Arenicolumba n. gen.
Diagnosis.-A small columbid (Table 1) that dif-
fers from other genera (especially Columbina, Claravis,
Scardafella, Metriopelia, Geotrygon, Zenaida,
Leptotila, and Patagioenas) as follows. Tarsometatar-
sus (Fig. 1B, C): 1, in acrotarsial aspect, intercotylar
knob with flatter proximal surface (less rounded, less
pointed); 2, facet for hallux located more proximad than
in all genera except Patagioenas; 3, larger foramen
vasculare distale; 4, crista plantare lateralis more dis-
tinct than in all genera except Geotrygon; 5, in both
plantar and acrotarsial aspect, the articulating surface
of trochlea metatarsi III shorter than in all genera ex-
cept Patagioenas; 6, in distal aspect, trochlea metatarsi
III wide relative to its depth; 7, central groove in tro-
chlea metatarsi III deeper. Tibiotarsus (Fig. 2B): 8, inci-


Table 1. Measurements (in mm) ofArenicolumba prattae (Hemingfordian LMA, Thomas Farm, Florida) compared
with those of adult skeletons of two widespread, living North American species, Columbina passerina and Zenaida
macroura (F = female, M = male).

Coracoid: Tarsometatarsus:
depth of distal width
processus Tibiotarsus: (without medio-plantar
acrocoracoideus distal width process)

Arenicolumba prattae 2.47 3.39 4.09
Thomas Farm fossils 2.08-2.69 3.33-3.42 4.02-4.22
18 3 3

Columbina passerina F 2.05 2.83 3.28
Florida 1.86-2.13 2.65-2.97 3.12-3.50
12 12 12

Columbina passerina M 2.15 2.82 3.35
Florida 1.96-2.28 2.48-3.20 3.17-3.50
14 14 14

Zenaida macroura F 3.11 3.97 4.53
Florida 2.92-3.32 3.80-4.22 4.29-4.77
20 20 20

Zenaida macroura M 3.30 4.08 4.66
Florida 3.07-3.63 3.79-4.27 4.38-4.95
20 20 20








BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(1)


Figure 1. The tarsometatarsus of columbids in acrotarsial (upper row) and plantar (lower row) aspects. A. Columbina
passerina UF 40074. B. Arenicolumba prattae UF 62093. C. Arenicolumba prattae UF 219393. D. Oena capensis
UF 38785 [PB 39624]. E. Turtur chalcospilos UF 38782 [PB 36021]. Scale = 10 mm.


Figure 2. The tibiotarsus of columbids in dorsal (upper row) and ventral (lower row) aspects. A. Columbinapasserina
UF 40074. B. Arenicolumba prattae UF 96296. C. Oena capensis UF 38785 [PB 39624]. D. Turtur chalcospilos
UF 38782 [PB 36021]. Scale = 10 mm.








STEADMAN: Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida


sura intercondylaris relatively wide (condylus medialis
and condylus lateralis relatively narrow). Sternum (Fig.
3B): 9, rostrum stermi (manubrium) more elongate. Co-
racoid (Fig. 4B, C): 10, in dorsal aspect, processus
procoracoideus extends farther sternally. Humerus (Fig.
5B, C): 11, incisura capitis extends farther dorso-ven-
trally to wrap around the ventral margin of tuberculum
ventral and separate it more from corpus humeri.
Type species.--Arenicolumba prattae is the type
and only known species in the genus.
Etymology.-From the Latin arena, a feminine
word that means "sand, sandy place" (Brown 1956:678)
and the Latin columba, a feminine word meaning "dove,
pigeon" (Brown 1956: 278). The second a in arena be-
comes a linking i because it looks and sounds better this
way (Winston 1999: 163). The name Arenicolumba re-
fers to the sandy sediment at Thomas Farm (see Pratt
1990) in which fossils of this dove and other small verte-
brates are commonly found.
Remarks.-The 11 diagnostic characters of
Arenicolumba also distinguish it from the seven Old
World genera of Columbidae (see MATERIALS AND
METHODS) except for agreement with Oena in char-
acters 1, 3, 5, 9, and 10, with Turtur in characters 2, 3,
5-7, and 10, with Streptopelia in character 4, with
Columba in characters 4 and 10, with Ducula in char-
acters 2 and 4, with Geopelia in characters 4 and 10,
and with Gallicolumba in characters 4-6 and 10.
Note in the diagnosis that Arenicolumba differs in
all 11 diagnostic characters from all four New World
genera/subgenera (depending on classification) of small
columbids, namely Columbina, Claravis, Scardafella,
and Metriopelia. Agreement with Oena and Turtur in


five and six of these 11 characters, respectively, sug-
gests that Arenicolumba may be more closely related
to these genera than to any extant New World genus of
Columbidae. Today, the single species of Oena and the
five species of Turtur are confined to Africa (Gibbs et
al. 2001).

Order CUCULIFORMES (Wagler 1830)
Family CUCULIDAE Vigors 1825
A set of 12 fossils representing five skeletal ele-
ments is referred to the Cuculidae because of this com-
bination of characters. Tarsometatarsus: two hypotarsal
canals; intercotylar knob prominent; in plantar aspect,
fused area ofcristae hypotarsi short; in proximal aspect,
cotyla medialis square; corpus tarsometatarsi relatively
uniform in width, and much wider than deep; trochlea
lateralis greatly elevated, dorso-ventrally expanded, and
antero-posteriorly compressed. Coracoid: facies
articularis sternalis developed on both dorsal and ventral
surfaces; facies articularis sternalis evenly developed
on dorsal surface. Scapula: facies articularis humeralis
well elevated from corpus scapulae; acromion blunt in
dorsal aspect; corpus scapulae of uniform depth through
much of its length. Humerus: distal end greatly expanded
ventrad, yielding an elongated processus flexorius and
epicondylus ventralis; fossa musculo brachialis distinct
and oval. Mandibular ramus: similar in thickness, curva-
ture, and overall proportions to that in Tapera and
Morococcyx.

Thomasococcyx philohippus n. gen. & sp.
Holotype.-Proximal end of right tarsometatarsus,
UF 96268 (Fig. 6A), Thomas Farm local fauna


Figure 3. The sternum of columbids in ventro-anterior aspect. A. Columbina passerina UF 40074. B. Arenicolumba
prattae UF 219359. C. Oena capensis UF 38785 [PB 39624]. D. Turtur chalcospilos UF 38782 [PB 36021].
Scale = 10 mm.








BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(1)


Figure 4. The coracoid of columbids in dorsal (upper row) and ventral (lower row) aspects. A. Columbina passerina
UF 40074. B. Arenicolumba prattae UF 223384. C. Arenicolumba prattae UF/PB 6503. D. Oena capensis UF
38785 [PB 39624]. E. Turtur chalcospilos UF 38782 [PB 36021]. Scale = 10 mm.


Figure 5. The humerus of columbids in palmar (upper row) and anconal (lower row) aspects. A. Columbinapasserina
UF 40074. B. Arenicolumba prattae UF 223471. C. Arenicolumba prattae UF 223408. D. Oena capensis UF
38785 [PB 39624]. E. Turtur chalcospilos UF 38782[PB 36021]. Scale = 10 mm.








STEADMAN: Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida


Figure 6. The proximal end of the tarsometatarsus of cuculids in acrotarsial (upper row), plantar (middle row), and
proximal (lower row) aspects. A. Thomasococcyx philohippus UF 96268 (Neomorphinae). B. Morococcyx
,.7i, 7,p.-'. UF 38737 [PB 29340] (Neomorphinae). C. Tapera naevia UF 38735 [PB 22915] (Neomorphinae).
D. Geococcyx velox UF 38729 [PB 17647] (Neomorphinae). E. Crotophaga sulcirostris UF 40789 (Crotophaginae).
F. Coccyzus americanus UF 40346 (Cuculinae). G. Clamatorjacobinus UF 38730 [PB 27715] (Cuculinae). Scale
= 10 mm.


(Hemingfordian LMA), Gilchrist County, Florida.
Paratypes.-Distal ends of tarsometatarsi UF
96268, 106406, 106595, 106735,223532 (Fig. 7A), man-
dibularrami UF 219354,219355, sternal end of coracoid
UF 106422 (Fig. 8A), coracoidal ends of scapulae UF
106451, 106638 (Fig. 9A), distal end of humerus UF
223499 (Fig. 10A), all from the Thomas Farm local fauna
(Hemingfordian LMA), Gilchrist County, Florida.
Diagnosis.-A medium-sized, presumably ground-
dwelling cuckoo defined by the following combination
of characters. Tarsometatarsus: cotyla lateralis more
anconally protrudent than in Cursoricoccyx; foramina
hypotarsi relatively larger than in Cursoricoccyx;
hypotarsus extends more media than in Cursoricoccyx;


sulcus extensorius relatively deeper than in
Cursoricoccyx, Geococcyx, Neomorphus, Coccyzus,
Clamator, Chysococcyx, or Cuculus; sulcus extensorius
relatively narrower than in Guira, Crotophaga,
Dromococcyx, Geococcyx, Neomorphus, Coccycua,
Piaya, Centropus, Phaenicophaeus, Clamator,
Eudynamys, Chrysococcyx, or Cuculus; combined
depth and width of sulcus extensorius most similar to
the condition in Tapera and Morococcyx; corpus tar-
sometatarsi narrower relative to its depth than in any
other genus, this difference being most pronounced com-
pared to specimens of Old World genera; foramen
vasculare distale larger than in all other genera. Cora-
coid: resembles that in Crotophaga, Morococcyx,








BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(1)


Figure 7. The distal end of the tarsometatarsus of cuculids in acrotarsial (upper row) and plantar (lower row) aspects.
A. Thomasococcyx philohippus UF 223532 (Neomorphinae). B. Morococcyx c, i.i'lip'-' UF 38737 [PB 29340]
(Neomorphinae). C. Tapera naevia UF 38735 [PB 22915] (Neomorphinae). D. Geococcyx velox UF 38729 [PB
17647] (Neomorphinae). E. Crotophaga sulcirostris UF 33855 (Crotophaginae). F. Coccyzus americanus UF
40346 (Cuculinae). G. Clamator jacobinus UF 38730 [PB 27715] (Cuculinae). Scale = 10 mm.



























Figure 8. The coracoid of cuculids in dorsal (upper row), ventral (middle row), and sternal (lower row) aspects. A.
Thomasococcyx philohippus UF 106422 (Neomorphinae). B. Morococcyx i.wl,.:', pu UF 38737 [PB 29340]
(Neomorphinae). C. Tapera naevia UF 38735 [PB 22915] (Neomorphinae). D. Geococcyx velox UF 38729 [PB
17647] (Neomorphinae). E. Crotophaga sulcirostris UF 40789 (Crotophaginae). F. Coccyzus americanus UF
40346 (Cuculinae). G. Clamatorjacobinus UF 38730 [PB 27715] (Cuculinae). Scale = 10 mm.








STEADMAN: Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida


Figure 9. The scapula of cuculids in dorsal aspect. A. Thomasococcyx philohippus UF 106638 (Neomorphinae). B.
Cuculidae sp. UF/PB 6487. C. Morococcyx i, *hiv/ -',,s UF 38737 [PB 29340] (Neomorphinae). D. Tapera
naevia UF 38735 [PB 22915] (Neomorphinae). E. Geococcyx velox UF 38729 [PB 17647] (Neomorphinae). F.
Crotophaga sulcirostris UF 40789 (Crotophaginae). G Coccyzus americanus UF 40346 (Cuculinae). H. Clamator
jacobinus UF 38730 [PB 27715] (Cuculinae). Scale = 10 mm.


Figure 10. The humerus of cuculids in palmar (upper row) and anconal (lower row) aspects. A. Thomasococcyx
philohippus UF 223499 (Neomorphinae). B. Morococcyx c *.hi,.v/ -',,s UF 38737 [PB 29340] (Neomorphinae).
C. TaperanaeviaUF 38735 [PB 22915] (Neomorphinae). D. GeococcyxveloxUF 38729 [PB 17647] (Neomorphinae).
E. Crotophaga sulcirostris UF 40789 (Crotophaginae). F. Coccyzus americanus UF 40346 (Cuculinae). G Clamator
jacobinus UF 38730 [PB 27715] (Cuculinae). Scale = 10 mm.










Geococcyx, and Neomorphus more than in other gen-
era in the medial-lateral placement of, and great dorsal-
ventral depth of, the ventral shelf of facies articularis
sternalis (and resulting bulge of corpus coracoidei); agrees
with that in Tapera, Morococcyx, and Geococcyx, but
differs from that in other genera, in having a shallow but
sterno-humerally expanded dorsal component of facies
articularis sternalis. Scapula: relative to overall size of
the bone, facies articularis humeralis smaller than in all
other genera, and less elongated than in all New World
genera; acromion less elongate than in Guira,
Crotophaga, Tapera, Dromococcyx, and
Neomorphus; facies lateralis of corpus scapulae, be-
tween facies articularis clavicularis and facies articularis
humeralis, more concave than in all New World genera
except Guira and Neomorphus. Humerus: processus
flexorius smaller (less expanded posteriorly) than in all
genera except Dromococcyx and Geococcyx.
Etymology.-- Thomasococcyx is from Thomas,
referring to the Thomas Farm locality, and coccyx (from
the Greek kokkyx, m., cuckoo; Brown 1956:216, 241).
The Thomas family still cultivates much of the land sur-
rounding the fossil site. Thomasococcyx is masculine.
philohippus is from philos (Greek, m., dear one, friend;
Brown 1956:355) and hippos (Greek, m., horse; Brown
1956:423), an allusion to the three genera and species of
three-toed horses (see MacFadden 1992, 2001) that
shared the northern Florida landscape with this cursorial
cuckoo in the early Miocene. One of these horses,
Parahippus leonensis, is particularly common at Tho-
mas Farm (Hulbert 1984, 2001). The name philohippus
is a masculine noun in apposition.
Remarks.- Thomasococcyx philohippus is the
type and only known species in the genus. The tarsometa-
tarsus of Thomasococcyx is most similar qualitatively
to that in Tapera and Morococcyx among living genera
of cuckoos, whereas the coracoid and scapula are most
similar qualitatively to those in Morococcyx and
Geococcyx. The humerus of Thomasococcyx is most
similar qualitatively to that in Dromococcyx and
Geococcyx. Thus, Thomasococcyx is, as far as can be
determined now, a mosaic of living genera of
Neomorphinae.
The coracoid, scapula, humerus, and tarsometa-
tarsus of Thomasococcyx philohippus are all about
the size of the same elements in Morococcyx
i i.h,, Ii,.-,I, (Table 2), and therefore I interpret these
four skeletal elements to belong to the same extinct spe-
cies. I am somewhat less confident about referral of the
two mandibular rami (UF 219354, 219355) to
Thomasococcyx, given the great variation in bill depth
among living genera of cuckoos. Nevertheless, these


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(1)

two fossils respectively represent 8.5 mm (left) and 11.5
mm (right) lengths of the fused prearticular, articular,
and dentary of what is likely to have been a single man-
dibular ramus that was similar in size to that of
Morococcyx (as well as Tapera and Dromococcyx).

Cuculidae, gen. uncertain
Material.-Scapula lacking distal end, UF/PB 6487
(Fig. 9).
Description.- This scapula compares closely in
overall size to that of Guira guira, Crotophaga ani,
Coccyzus (Saurothera) merlini, and Piaya cayana. It
is much too large to pertain to Thomasococcyx
philohippus (Table 2), from which it differs further by
having a much more dorsally flared facies articularis
clavicularis as in Tapera or Crotophaga. UF/PB 6478
is approximately the right size for Cursoricoccyx
gertrudae (Hemingfordian of Colorado; Martin and
Mengel 1984), based on extrapolation from the size of
the coracoid and carpometecarpus in the latter. More
material of this large cuckoo from Thomas Farm is
needed before it can be characterized further.

DISCUSSION
COLUMBIFORMES
Compared to many other non-passerine orders of
birds, the Columbiformes (traditionally but probably er-
roneously considered to consist of Pteroclididae +
Columbidae) has a poor fossil record in the Tertiary.
The Pteroclididae sandgrousee) is represented by sev-
eral Paleogene (late Eocene early Oligocene) fossils
from Europe that represent two extinct genera with no
apparent links to the Columbidae (Archaeoganga,
Leptoganga; Mourer-Chauvire 1992, 1993). I see no
characters in Arenicolumba that would link it more than
any other columbid to the Pteroclididae or any other fam-
ily.
The Columbidae has no Paleogene fossil evidence
at all, even among the very rich European and North
American avifaunas of the Eocene and Oligocene, where
the first evidence of many if not most living orders and
families of non-passerine birds is found (James 2005,
Mayr 2005). Nothing in the fossil record of columbids or
of birds in general supports the Cretaceous origin of
Columbiformes proposed by Pereira et al. (2007) from
mitochondrial and nuclear genetic sequences. Because
the Columbiformes almost certainly did not exist in the
Cretaceous, one need not invoke the breakup of
Gondwana as an essential part of their historic biogeog-
raphy. Nevertheless, the Columbiformes may have origi-
nated in the Southern hemisphere (Olson 1989), an idea
supported by their absence in the rich Paleogene fossil















Table 2. Measurements (in mm) of Thomasococcyxphilohippus, new genus and species (Hemingfordian LMA, Thomas Farm, Florida) compared to those of

Cursoricoccyx gertrudae (Hemingfordian LMA, Martin Canyon, Colorado), an indeterminate cuculid from Thomas Farm, and adult skeletal elements of

modem species of Cuculidae.



Scapula length
Species Mandible depth at Coracold Coracold length and width of Scapula width of Tarsometatarsus
junction of dentary and sternal width through scapular faces articularis corpus Humerus Carpometacarpus Tarsometatarsus Tarsometatarsus minimum depth of corpus
surangular and depth and glenoid facets humeralis scapulae distal width proximal width proximal width proximal depth tarsometatarsi


Thomasococcyx 2 15 (UF 219355)
philohippus, 2 18 (UF 219354)
new species



Cursoncoccyx ---
gertrudae
KUVP 25629-25631


4 03, 1 90
(UF 106422)


248, 1 91
(UF 106451)
2 47, 1 69
(UF 106638)


1 89
(UF 106451)
1 67
(UF 106638)


5 51
(KUVP 25630)


Cuculidae sp
Thomas Farm
UF/PB 6487


Gumra guira
UF 38969

Crotophaga major
KU 32390

Crotophaga am
UF 42473

Crotophaga
sulcirostris
UF 40789

Tapera naevia
UF 38735

Dromococcyx
phasianellus
KU 91388

Dromococcyx
pavomnus
LSUMZ 101257

Morococcyx
erythropygus
UF 38737

Geococcyx
cahformanus
UF 26238


5 84, 2 62



764, 220



571, 238



491, 1 94




3 92, 1 63



591, 223




4 03, 1 71




4 59, 1 54




8 78, 3 49


6 30
(UF 223499)


422 451
(UF 96268) (UF 96268)


1 14
(UF 106406)


6 03 5 68 5 57
(KUVP 25631) (KUVP 25629) (KUVP 25629)


3 18, 229




349, 228



3 83, 232



341, 251



2 59, 1 89




2 52, 1 48



3 04, 2 48




2 50, 1 71




2 34, 1 85




3 60, 2 70


2 12


725


4 29

















Table 2. (Cont.)


Species


Man
junction


Geococcyx velox
UF 38729

Neomorphus
rufipennms
KU 86604

Neomorphus
geoffroyi
LSUMZ 106946

Coccycua (Playa)
minute UF 26232

Playa cayana
UF 40787

Playa melanogaster
UF 40787

Coccyzus americanus
UF 40346

Coccyzus minor
UF 40029

Coccyzus (Hyetorms)
pluviahs UF 26233

Coccyzus (Saurothera)
merhm UF 42489

Centropus
supercilhosus
UF 33856

Zanclostomus
(Phaemncophaeus)
javamncus UF 42704

Clamator jacobinus
UF 38730


dible depth at
.n of dentary and
surangular



4 14



4 77


Coracold
sternal width
and depth



732, 2 18



779, 236


Coracold length
through scapular
and glenoid facets



5 09



5 96


8 73, 232




342, 148



556, 1 51



6 50, 1 80



466, 1 82



543, 200



717, 233



5 96, 1 81



6 62, 3 25




5 56, 1 70




427. 1 56


2 46


Scapula length
and width of
faces articularis
humeralis



3 86, 245



407, 254


467, 340


Scapula width of
corpus
scapulae



2 40



2 37


Humerus
distal width



8 43



9 19


Carpometacarpus
proximal width



6 07


Tarsometatarsus
Tarsometatarsus Tarsometatarsus minimum depth of corpus
proximal width proximal depth tarsometatarsi



707 668 213



8 44 849 2 31


2 83 1054


2 87, 1 94




241. 1 86








STEADMAN: Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida


record of Europe and North America.
The abundant specimens ofArenicolumbaprattae
from Thomas Farm represent the New World's earliest
record of Columbidae. The only earlier fossil assigned
to the Columbidae is the single humerus of Gerandia
calcaria from the early Miocene (Aquitanian; ca. 22
ma) of Saint Gerand-le-Puy, France (Lambrecht 1933,
Brodkorb 1971, Olson 1985). Continental fossils of
columbids that are younger than Arenicolumba, mainly
from the northern hemisphere, seem to represent mod-
em genera (Olson 1985, Becker and Brodkorb 1992).
Columbids do have an extensive Quaternary fossil
record, especially on tropical oceanic islands where con-
siderable anthropogenic extinction of species and even
genera is documented (Steadman 1992, 2006, Pregill et
al. 1994).
The family Columbidae is broadly distributed in tem-
perate and tropical regions today. Its greatest generic
and species-level diversity is in the Australian-Papuan-
Oceanic region (Steadman 2006), where the substantial
morphological and ecological diversity of pigeons and
doves stands in marked contrast to the more uniform
morphology and ecology of New World columbids.
In eight genetically-based cladograms of living
columbid genera by Pereira et al. (2007), Oena and
Turtur (the two living genera most similar to the extinct
Arenicolumba) are rather derived although often with
low posterior probabilities or bootstrap support. Should
the phylogeny hypothesized by Pereira et al. (2007) be
valid, it would suggest that Arenicolumba is not a basal
columbid. In having its proposed closest living relatives
confined to Africa, Arenicolumba resembles the primi-
tive crane Balearica (Gruidae), which occurs in Africa
today but is common in the North American Neogene
(Olson 1985, Feduccia and Voorhies 1992).
Of the nine osteological characters regarded by
Livezey & Zusi (2006) to be diagnostic or supportive for
defining the Columbidae, only seven (their characters
number 1307, 1356, 1369,1417,1723,2036,2119) could
potentially be evaluated in Arenicolumba. I found only
one of these characters (number 1723) to be valid and
diagnostic of columbids; the remaining six characters
are either highly generalized (apply to numerous fami-
lies of birds and are unlikely to represent homologies) or
cannot be interpreted.
At the time of its description in 1992, at least 375
specimens of Arenicolumba prattae had been recov-
ered. Subsequent field and laboratory work at Thomas
Farm has increased this number to more than 700, which
make up 75% to 80% of the site's identifiable bird fos-
sils. It is noteworthy that, among these numerous speci-
mens, only the ulnare, manus phalanges, and pedal pha-


langes are ever complete. The consistent breakage of
other skeletal elements supports the thesis of Pratt (1989)
that most breakage of bones in doves (and other small
vertebrates) within the Thomas Farm sinkhole was due
to damage by avian and mammalian predators at or just
before the time of deposition.

CUCULIFORMES
Traditionally (i.e., Mayr & Amadon 1951, Wetmore
1960), the Cuculiformes was considered to comprise two
families, the Cuculidae and Musophagidae. Endemic to
Africa today, musophagids (turacos and plantain-eaters)
are known as fossils from the Oligocene of Africa and
the Oligocene and Miocene of Europe; Ballmann 1970,
Feduccia 1996, Mayr 2005). The strictly cuculid char-
acters of the five skeletal elements available for
Thomasococcyx philohippus (tarsometatarsus, cora-
coid, scapula, humerus, and mandible) support the hy-
pothesis that the Musophagidae is only distantly related
to the Cuculidae and does not belong in the Cuculiformes.
This agrees with the proposals of Sibley & Ahlquist
(1990), Mindell et al. (1997), Sorenson et al. (2003), Mayr
& Ericson (2004), and perhaps Cracraft et al. (2004). I
should note, however, that the early Eocene Foro
panarium (known from a complete, associated skel-
eton, Green River Formation, Wyoming) was placed in
an extinct family of Cuculiformes with characters sug-
gestive of musophagids, cuculids, and the enigmatic
Opisthocomus hoazin (Olson 1992). Furthermore,
Hughes (2000) classified the Opisthocomidae (consist-
ing of the living Opisthocomus hoazin and the middle
Miocene fossil Hoazinoides magdalenae from Colom-
bia; A. H. Miller 1953) and Musophagidae together to
make up the Opisthocomiformes, this order being sister
to the Cuculiformes, consisting only of the Cuculidae.
Whether the Cuculidae originated in the New World
versus Old World, or Northern Hemisphere versus
Southern Hemisphere, awaits the discovery of more Pa-
leogene fossils. The Tertiary fossil record of cuckoos is
poor. Nearly all of the supposed Eocene and Oligocene
cuckoos from Europe are either not referable to the
Cuculidae or are of questionable identity; Dynamopterus
velox from the Eo-Oligocene Quercy fauna of France
may be a cuckoo but is based only on a humerus (Olson
& Feduccia 1979, Martin & Mengel 1984, Olson 1985,
Mayr 2005, Mayr & Mourer-Chauvire 2005).
In North America, Eocuculus cherpinae is based
on a partial associated skeleton from the late Eocene
(early Chadronian LMA; ca. 37-36 Ma) of Colorado
(Chandler 1999). Eocuculus was much smaller than
Thomasococcyx and had relatively short legs suggest-
ing that it was arboreal rather than ground-dwelling.










Chandler (1999) classified Eocuculus not only in
the Cuculidae but in the subfamily Cuculinae, with its
closest affinities to Cuculus (Old World) among living
genera. Mayr (2006) reported Eocuculus from the early
Oligocene (ca. 30 Ma) of France, excluded it from crown
group Cuculidae, and was justifiably tentative about re-
ferring it to the Cuculidae. Neococcyx mccorquodalei
is based only on the distal end of humerus from the early
Oligocene (late Chadronian LMA; ca. 34 Ma; Prothero
& Emry 2004) of Saskatchewan (Weigel 1963). I agree
with Olson (1985) thatNeococcyx probably is a cuckoo,
although the limited material precludes any precise hy-
pothesis of relationship.
Moving to the Neogene, roughly contemporane-
ous with Thomasococcyx is Cursoricoccyx geraldinae,
based on a humeral end of coracoid, a nearly complete
carpometacarpus, and a proximal end of tarsometatar-
sus from the early Miocene (Hemingfordian LMA)
Martin Canyon local fauna of Colorado (Martin &
Mengel 1984). While I agree with Martin and Mengel
(1984) that it belongs in the Neomorphinae, C.
geraldinae is much larger than T philohippus (Table
2) and differs further in four characters of the tarsometa-
tarsus (the only directly comparable element in the two
species).
Finally, Olson (1985:111) mentioned the distal end
ofa tibiotarsus from "a large cuckoo of uncertain affini-
ties" from the early Pliocene (late Hemphillian LMA)
Lee Creek local fauna, North Carolina. This specimen
(USNM 256228) was tentatively referred to the Galli-
formes (family, genus, and species indeterminate) by
Olson & Rasmussen (2001), who also noted that this
fossil may derive from the early to middle Miocene
Pungo River Formation (Barstovian LMA) rather than
the Hemphillian Yorktown Formation.
The interpretation of Thomasococcyx philohippus
as a ground-dwelling species is compatible with the os-
teological characters that ally it more closely to living
New World genera of ground-dwelling cuckoos
(Neomorphinae, which I regard as monophyletic and
more closely allied to the Crotophaginae than to the
Cuculinae) than to any of the living Old World genera of
Cuculidae. Following the classification of Payne (2005),
which is based on morphology, DNA, and breeding ecol-
ogy, my assignment of Thomasococcyx to the
Neomorphinae suggests that the split between the New
World ground-dwelling cuckoos and other cuculid sub-
families already had occurred by the early Miocene. At
this time at least two genera of Neomorphinae
(Cursoricoccyx, Thomasococcyx) existed in North
America. Today, Geococcyx is the only neomorphine
cuckoo north of ca. 240N, the remaining four genera


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 48(1)

(Tapera, Dromococcyx, Morococcyx, Neomorphus)
being strictly neotropical. I note here that Hughes (2000)
placed Morococcyx, Geococcyx, and Neomorphus in
the Neomorphinae, but Tapera and Dromococcyx in
the Cuculinae. I disagree, as does Payne (2005).
It is beyond the scope of this paper to assess the
novel hypothesis of Mayr & Ericson (2004) that the
Mesitomithidae (endemic to Madagascar; traditionally
placed in the Gruiformes) has a sister group relationship
to the Cuculidae. Of the 80 osteological characters used
by Mayr & Ericson (2004) to propose a close relation-
ship between the Cuculidae and Mesitomithidae, only
nine (their characters numbered 40, 54, 55, 73-78) can
be evaluated in the available material of Thomasococcyx
philohippus. In each case, the character state of T
philohippus agrees with that given for modem skel-
etons of both Cuculidae and Mesitomithidae by Mayr &
Ericson (2004), except that Thomasococcyx agrees with
Cuculidae to the exclusion of Mesitomithidae in charac-
ter 78.
Of the seven osteological characters that Livezey
& Zusi (2006) found to be diagnostic or supportive for
the Cuculidae (their characters number 1007, 1336, 1572,
1614, 1651, 1658, 1866), none can be evaluated in
Thomasococcyx. Nevertheless, I found six of these
seven characters to be generalized (applicable to nu-
merous families of birds; unlikely to represent homolo-
gies) and the other one (1651) impossible to interpret.

ACKNOWLEDGEMENTS
This research was supported by the National Science
Foundation (Assembling the Tree of Life grant DEB-
0228682) and the Toomey Foundation. I thank Richard
Hulbert, Bruce MacFadden, Arthur Poyer, and Erika
Simons for working with me in the field, generously shar-
ing their extensive knowledge of the Thomas Farm site,
and facilitating access to the Thomas Farm bird fossils.
I also thank the many volunteers who have worked so
hard at Thomas Farm through the years, with special
recognition to Micah Adams, Norma Jean Aker, Grant
Boardman, David Cale, Tabitha Cale, Wiley Dykes,
Carolyn Eastwood, Patricia Geisler, Karen Goodheart,
Bill Lee, Judy Lundquist, Mary Lynch, Kari Stopp, Bar-
bara Toomey, George Weems, and Marcia Wright. The
photographs were skillfully taken by Sean Roberts. For
comments that improved the manuscript, I thank Rich-
ard Franz, Richard Hulbert, Bruce MacFadden, and Storrs
Olson. I am grateful to Andrew Kratter, Mark Robbins,
Jeff Sailer, Terry Taylor, Tom Webber, and the late Pierce
Brodkorb for preparing high-quality modem skeletons
of columbids and cuckoos. For access to specimens
under their care, I thank Larry Martin (KUVP), Town








STEADMAN: Doves (Columbidae) and Cuckoos (Cuculidae) from the Early Miocene of Florida


Peterson (KU), and Fred Sheldon (LSUMNS).

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