AN EARLY PLEISTOCENE AVIFAUNA
FROM INGLIS, FLORIDA
GAIL ELAINE SPEAKER CARR
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
I thank Pierce Brodkorb for his supervision and enthusiasm during
this study. Dr. Brodkorb has been more than a graduate advisor; he
has been an encouraging friend. I am indebted to David Webb, David
Johnston, and Thomas Emmel for sound advice and enlightening discussions.
Conversations with David Steadman and Thomas Ritchie were helpful
early in the study. Diana Matthiesen and Ronald Wolff aided me in fossil
photography. Stephen Carr kindly made the drawings.
My deepest thanks go to the members of my family who provided endless
amounts of financial and emotional support: my parents, Edwin and Donna
Speaker; my husband's parents, Archie and Marjorie Carr; and, above all,
my patient husband, Archie Carr, III.
TABLE OF CONTENTS
ACKNOWLEDGEMENTS .............................................. it
LIST OF TABLES ...................................... ...... .... iv
LIST OF FIGURES.... .............. ............................. viii
ABSTRACT ..................................................... x
INTRODUCTION.......................................... ........ 1
LIST OF SPECIES........................... ..................... 6
PALEOECOLOGY ............... ................. .......... ....... 129
COMPARISON WITH OTHER AVIFAUNAS............................... 131
AGE OF DEPOSIT.......................................... ...... 135
LITERATURE CITED................................................ 150
BIOGRAPHICAL SKETCH........................................... 162
LIST OF TABLES
1 List of the 47 non-passerine species from Inglis IA
with number of elements (Elem) and minimum number of
individuals (Ind)...................................... ... 3
2 Measurements of the ulna of the Late Miocene Gavia brodkorbi
(Howard, 1978), Recent G. stellata, Recent G. arctica
pacifica, Inglis G. arctica, Late Pliocene G. concinna
Recent G. immer, and Recent G. adamsii..................... 7
3 Measurements of the humerus and ulna of Recent Podiceps
auritus (2 males, 4 females, 4 ?) and Inglis P. dixi....... 11
4 Measurements of the ulna of Recent Podiceps dominicus
(1 male, 3 females) and Inglis P. dominicus............... 13
5 Measurements of the humerus, ulna, and tibiotarsus of
Recent Podilymbus podiceps (from Storer, 1976; 23 males
for humerus and tibiotarsus; 16 females for ulna) and
Inglis P. podiceps......................................... 15
6 Measurements of the humerus of Recent Eudocimus albus
(2 males, 4 females) and Inglis E. albus................... 17
7 Measurements of the sternum of Recent Ardea herodias
(1 male, 5 females) and Inglis Ardea sp.................... 19
8 Measurements of the carpometacarpus of Recent Botaurus
lentiginosus (6 males, 5 females) and Inglis B.
lentiginosus ............................................. 21
9 Measurements of the humerus of Aix sponsa (8 males,
5 females) and Inglis A. sponsa............................ 23
10 Measurements of the humerus, ulna, and carpometacarpus
of Anas discors (3 males, 3 females) and Inglis A. discors. 26
11 Measurements of the ulna of Aythya collaris (7 males, 2
females), A. valisineria (2 males, 6 females).............. 27
12 Measurements of the tarsometatarsus of Rancho La Brea
Anabernicula (Howard, 1964), Inglis A. gracilenta, A.
oregonensis (Howard, 1964), and Recent Tadorna ferruginea
(4 females) ................................................ 30
13 Proximal width of the tibiotarsus of modern Sarcoramphus
papa, Inglis cf. Gymnogyps californianus, Recent G.
californianus (Fisher, 1947), Pleistocene Teratornis
merriami (Fisher, 1945), and Pleistocene Vultur
(Campbell, 1973)........................................... 32
14 Measurements of the tarsometatarsus of mid-Pleistocene
Gymnogyps amplus from Reddick, late Pleistocene G.
amplus from Rancho La Brea (Fisher, 1947 and L. Miller,
1910), and Recent G. californianus (Fisher, 1947).......... 34
15 Measurements (mm.) of the coracoid, humerus, ulna, and
tibiotarsus of early Pleistocene Coragyps atratus from
Inglis, middle Pleistocene C. occidentalis from Reddick,
late Pleistocene C. o. occidentalis from Rancho La Brea
(Howard, 1968), late Pleistocene C. o. mexicanus from San
Josecito Cave, Mexico (Howard, 1968), Recent C. atratus
atratus from Florida and California (Howard, 1968), and
Recent C. stratus brasiliensis from Panama................. 36
16 Measurements of the quadrate and tarsometatarsus of Inglis
Cathartes aura and Recent C. aura (4 males, 5 females)..... 42
17 Measurements (mm.) of the carpometacarpus of Buteo
magnirostris, B. platypterus, B. nitidus, B. lineatus, B.
lagopus, B. swainsoni, B. jamaicensis, Inglis Buteo n. sp.,
and B. regalis............................................. 44
18 Measurements (mm.) of the tibiotarsus and tarsometatarsus
of Buteo lagopus, B. swainsoni, B. jamaicensis, Inglis
Buteo n. sp., and B. regalis............................... 47
19 Measurements of phalanx I of digit I and the ungual
phalanges of digits I, II, III, and IV of Recent Buteo
jamaicensis (2 males, 2 females) and Inglis Buteo n. sp.... 49
20 Measurements (mm.) of the scapula, tibiotarsus,
metatarsal I, phalanx I of digit III, ungual phalanx of
digit I, phalanx II of digit II, phalanx I of digit III,
phalanx III of digit III, phalanx I of digit IV, and ungual
phalanx of digit IV from Recent Aquila chrysaetos and
Aquila n. sp............................................... 55
21 Measurements of the carpometacarpus and femur from early
Pleistocene Aquila n. sp. from Inglis, middle Pleistocene
A. chrysaetos bonifacti from Saint-Estive, France (Mourer-
Chauvire, 1975), late Pleistocene A. chrysaetos from
Rancho La Brea (Howard, 1932), and Recent A. chrysaetos
from North America (Brodkorb Collection and Howard, 1932)
and from Europe (Mourer-Chauvire, 1975).................... 58
22 Femur measurements (mm.) of Aguila borrasi (Arredondo,
1976) and Aquila n. sp. .................................. 63
23 Measurements of the mandible and femur of Neophron
percnopterus (2 females), Necrosyrtes monachus (sex ?),
Neophrontops americanus (Howard, 1932), Palaeoborus
umbrosus (Cope, 1877), Inglis Neophrontops slaughter,
and Neogyps errans (Howard, 1932). measured from
photograph in Howard (1932)................................ 67
24 Measurements (mm.) of the tibiotarsus of Neophron
percnopterus (2 females), Necrosyrtes monachus (sex ?),
Arikarornis macdonaldi (Howard, 1966), Neophrontops
americanus (Feduccia, 1974), N. slaughter (Feduccia,
1974), Inglis N. slaughter, Palaeoborus umbrosus (Cope,
1877), and Neogyps errans (Howard, 1932)................... 71
25 Measurements of the tarsometatarsus of Neophron
perenopterus (2 females), Necrosyrtes monachus (sex ?),
Neophrontops americanus (Howard, 1963), Neophrontops
vallecitoensis (Howard, 1963), Inglis Neophrontops
slaughter, and Neogyps errans (Howard, 1966).............. 76
26 The fossil Gypaetinae of the New World (revised from
Feduccia, 1974............................................. 77
27 Measurements of the ulna and tibiotarsus of Recent
Accipiter cooperii (3 males, 3 females) and Inglis A.
28 Measurements of the carpometacarpus and tarsometatarsus
of Rallus limicola (5 males, 4 females), Inglis R.
longirostris, R. longirostris waynei (1 male, 6 female),
Inglis R. elegans, R. elegans elegans (2 male, 2 female),
Fulica americana americana (2 males, 6 females), Gallinula
chloropus cachinnans (5 males, 4 females), and Porphyrula
martinica (3 males, 3 females)............................. 84
29 Measurements of the carpometacarpus and tibiotarsus of
Rallus limicola (5 males, 4 females), Porzana carolina
(7 males, 4 females, 1 ?), and Inglis P. carolina.......... 87
30 Measurements of the ulna of Anthropoides virgo (1 male,
I female), Balearica pavonina (1 male, 2 females, 1 ?),
Grus grus grus (1 ?), Grus g. lilfordi (2 females), Grus
canadensis rowani (2 males, 1 female), Grus c. tabida
(3 males, 4 females), G. c. pratensis (4 males, 2 females),
Grus americana (1 ?), and Inglis Grus americana........... 89
31 Measurements of the phalanges of the Inglis Titanis
32 Measurements of the humerus, carpometacarpus. amd tarso-
metatarsus of Recent Capella gallinago delicate (1 male,
3 females, 1 ?) and Inglis C. gallinago.................... 96
33 Measurements of the coracoid and tarsometatarsus of Recent
Philohela minor (6 males, 8 females), Inglis P. minor, and
Recent Scolopax rusticola (1 male, 1 ?).................... 98
34 Measurements of the ulna and tarsometatarsus of Recent
Coccyzus erythropthalmus (1 male, 1 female), C. minor
(1 male, 1 ?), C. americanus (4 males, 1 female), and
Inglis C. americanus ...................................... 101
35 Measurements of the ulna and ulnare of Tyto alba (3
males, 2 females), Strix varia (1 male, 3 females, 3 ?),
Inglis Bubo virginianus, B. v. virginianus (6 males, 8
females), and Nyctea scandiaca (1 male).................... 104
36 Measurements of the femur of Strix varia (I male, 2
females, 2 ?), Inglis Bubo virginianus, and Bubo v.
virginianus (6 males, 8 females).......................... 105
37 Skeletal measurements (mm.) of Otus flammeolus, 0. asio,
Inglis 0. asio, Aegolius funereus richardsoni, and
Speotyto cunicularia floridana ............................ 109
38 Femur measurements of Recent Tyto alba (1 male,
2 females), Inglis T. alba, and T. pollens (Wetmore,
1937); tarsometatarsus measurements of T. cavatica
(Wetmore, 1920), Recent T. alba, and T. ostologa
(Wetmore, 1922) ............................................ 115
39 Measurements of the scapula, femur, tibiotarsus, and
tarsometatarsus of Glaucidium sp. 2 (tibiotarsus and
tarsomotatarsus only), G. minutissimus (scapula and femur
only), G. gnoma, G. brasilianum, and G. sp. 1.............. 118
40 Measurements of the humerus and tibiotarsus of Recent
Asio otus (2 males, 1 female), Inglis A. priscus, Recent
A. flammeus (1 male, 3 females), and Recent A. capensis
I male, 1 female) .......................................... 121
41 Comparison of the humerus (distal end) of three species of
42 Measurements of the coracold, humerus, ulna, femur,
tibiotarsus, and tarsometatarsus of Dendrocopos pubescens
(1 male), D. borealis (2 females), Sphyrapicus various (3
males), Melanerpes carolinus (3 males, 2 females), M.
erythrocephalus (2 males), and Colaptes auratus (1 male)... 125
43 Comparison of Inglis IA with some other North American
paleoavifaunas: Rexroad (Feduccia, 1975), lagerman
(Feduccia, 1975), Coleman IIA and III (Ritchie, 1980),
Arredondo (Brodkorb, 1959a), and Rancho La Brea (Howard,
44 Minimum number of individuals (Ind.) and percentage of
total for each order of birds from Inglis IA, Coleman IIA
(Ritchie, 1976), and Rancho La Brea (Howard, 1930)......... 133
LIST OF FIGURES
1 Titanis walleri, second cervical vertebra (axis) in dorsal
view, slightly smaller than natural size..................... 137
2 Titanis walleri, third cervical vertebra in dorsal view,
slightly smaller than natural size........................... 139
3 Titanis walleri, second cervical vertebra (axis)............. 141
4 Titanis walleri, third cervical vertebra...................... 143
5 Titanis walleri, left carpometacarpus in internal and
external views; Ixobrychus n. sp., distal ends of left
tarsometatarsus and left tibiotarsus in anterior and
posterior views.............................................. 145
6 Aquila n. sp., right carpometacarpus in internal and
external views; Buteo n. sp., left carpometacarpiis in
internal and external views.................................. 147
7 Neophrontops slaughter (coracoid, tibiotarsus,
tarsometatarsus) ........................................... 149
Abstract of Dissertation Presented to the Graduate Council of
the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
AN EARLY PLEISTOCENE AVIFAUNA
FROM INGLIS, FLORIDA
Gall Elaine Speaker Carr
Chairman: Pierce Brodkorb
Major Department: Zoology
Inglis IA in Citrus County, Florida, is a fossiliferous early
Pleistocene (Irvingtonian) sinkhole deposit in the Inglis member of
the late Eocene Ocala Limestone. The non-passerine species of the
paleoavifauna consists of over 2,710 specimens representing a minimum of
258 individuals of 12 orders, 18 families, 41 genera, and 47 species.
Thirty-four percent of the species are extinct. Five new species are
described. The families represented and the number of identified
species in each family are: Gaviidae, 1; Podicipedidae, 3; Ardeidae, 3;
Plataleidae, 1; Anatidae, 6; Vulturidae, 3; Accipitridae, 5; Falconidae,
2; Phasianidae, 2; Rallidae, 3; Gruidae, 1; Phorusrhacidae, 1; Scolo-
pacidae, 2; Columbidae, 1; Cuculidae, 1; Strigidae, 7;; Tytonidae, 1;
Inglis IA represents a community from a late glacial (Kansan)
habitat of high pine, xeric hammock, and savanna. The composition of
this avifauna is similar to that of a younger Irvingtonian site
from Coleman, Florida.
Florida's wealth of fossil material has kept workers in avian
paleontology busy for many years. A list of only the major publi-
cations on fossil avifaunas from this state includes Wetmore (1931
and 1943c), Brodkorb (1953b, 1955, 1959a, 1963d), McCoy (1963),
Ligon (1965), Ritchie (1980), and Campbell (1980). These studies
cover localities ranging in age from late Miocene to late Pleistocene.
The oldest Pleistocene localities in Florida from which avian
fossils have been studied are: one Blancan site, Santa Fe River I
(Brodkorb, 1963c); and three Irvingtonian sites, Inglis IA (this
paper) and the younger Coleman IIA and III (Ritchie, 1976 and 1980).
Therefore, the Inglis IA collection represents the earliest Florida
Pleistocene avifauna known.
Inglis IA is located in Section 8, R. 16 E., T. 17 S., in Citrus
County, Florida. The site was discovered at sea level on the north
bank of the Cross Florida Barge Canal by Jean Klein and Robert Martin
in 1967. The Florida State Museum collected fossils there between 1967
Inglis IA is a sinkhole deposit in the Inglis member of the late
Eocene Ocala Limestone. Klein (1971) and Meylan (1980) provide exten-
sive discussions of the geology of the deposit. The sediment filling
the sinkhole consisted of six layers. The lowest level which covered
the floor of the sink was the basal conglomerate composed of sand,
limestone, and bone fragments. Above this was the lower clay layer,
followed by the lower sand layer, upper clay layer, and then the upper
sand layer. The thin clay layers, less than 5 cm. deep, Indicate
periods of reduced energy of deposition which allowed the fine clay
particles to accumulate during periods of high water level in the sink.
The sandy levels varied between one and two meters in thickness, and
housed most of the fossils. The highest layer of the deposit was
composed of cemented sandstone plugging the neck of the sinkhole.
These alternating levels of sand and clay are indications of
fluctuations in the water table. Klein (1971) believes the time span
represented by this series of layers to be a short one based on the
following geological evidence: (1) the small size of the chamber; (2)
no collapse within the chamber; and (3) the small diameter of the
opening to the surface. Klein (1971) concludes that the Inglis IA
population sample is isochronous.
The ferungulates of Inglis IA were studied by Klein (1971), the
turkeys by Steadman (1975 and 1980), the quails by Ritchie (1976 and
1980), and the squamate reptiles by Meylan (1980). Webb (1974) lists
the mammals from Inglis IA.
The non-passerine avifauna from Inglis IA contains representa-
tives of 12 orders, 18 families, 41 genera, and 47 species. This
includes one extinct family, 4 extinct genera, and 16 extinct species.
Thirty-four percent of the non-passerine avifauna is extinct. Five
new species are described. There are 2,710 elements and a minimum
number of 258 individuals. Table 1 lists the non-passerine birds
identified from Inglis IA.
Table 1. List of the 47 non-passerine species from Inglis 1A, with
number of elements (Elem) and minimum number of individuals (Ind).
Extinct species are marked with an asterisk (*).
Gavia artica 1 1
Podilymus podiceps 6 2
Podiceps dominicus 1 1
*Podiceps dixi 3 1
Ardea sp. 1 1
Botaurus lentiginosus 1 1
Ixobrychus n. sp. 2 1
Eudocimus albus 1 1
*Anabernicula gracilenta 2 1
Aix sponsa 5 1
Anas platyrhynchos 2 1
Anas strepera 4 1
Anas discors 5 2
Aythya americana 1 1
Coragyps atratus 17 2
Cathartes aura 2 1
cf.Gymnogyps californianus 1 1
Buteo n. sp. 12 2
Aquila n. sp. 34 2
*Buteogallus fragilis I 1
*Neophrontops slaughter 7 1
Accipiter cooperil 6 2
Falco columbarius 2 1
Falco sparverius 14 2
*Colinus suilium 1100 115
*Meleagris anza 1230 50
Table 1. Continued.
Rallus longirostris 11 3
Rallus elegans 2 2
Porzana carolina 4 3
Grus americana 1 1
*Titanis walleri 11 2
Capella galinago 10 4
Philohela minor 12 2
Zenaidura macroura 40
Coccyzus americanus 25
Bubo virginianus 10
Otus asio 44
*Speotyto megalopeza 6
Glaucidium n. sp. I 12 3
Glaucidium n. sp. 2 3 2
*Asio priscus 3 1
Strix varia 4
Tyto alba 1
*Campephilus dalquesti 1 I
Colaptes auratus 10 3
Melanerpes erythrocephalus 38 7
cf. Sphyrapicus various 1 1
Totals 2,710 258
Twenty-three species are represented by 5 or more specimens,
17 by 10 or more specimens, and 7 by 15 or more specimens. Twelve
species are represented by only one specimen. The extinct turkey,
Meleagris anza, is the best represented species, having 1,230 fossils
from a minimum number of 50 individuals. Another extinct gallin-
aceous bird, the quail Colinus suilium, is a close second with 1,100
elements from a minimum number of 115 individuals.
The habitat requirements of the avifauna range from aquatic
conditions to drier forested areas and open savanna country. Species
requiring aquatic or marsh habitats include the loon, grebes (three
species), herons (three species), ibis, ducks (six species), and
rails (three species). The woodpeckers (four species) are abundant,
as is the cuckoo. Species with terrestrial requirements include the
quail, turkey, crane, phorusrhacid, and dove.
The ten species of diurnal raptors include three vulturids, five
accipitrids, and two falconids. The owls are represented by eight
species, including two new species of Glaucidium, the pygmy-owls.
The fossil birds from Inglis IA are housed in the Florida State
Museum at the University of Florida in Gainesville.
Most of the modern skeletons examined in this study were on loan
from the collection of Pierce Brodkorb. Several skeletons were
borrowed from the Florida State Museum and the Smithsonian
Avian osteological terms are those of Howard (1929). Osteo-
logical measurements were taken following those of Steadman (1975) and
All measurements are given in millimeters, with range (R), mean
(M), and number of specimens (N).
LIST OF SPECIES
Asterisks denote extinct taxa. Classification follows that of
Brodkorb (1963a, 1964a, 1967, 1971a).
Order Gaviiformes Wetmore and W. D. Miller
Family Gaviidae Allen
Subfamily Gavinae (Allen)
Genus Gavia Forster
Gavia arctica (Linnaeus)
Material: Proximal end of 1 left ulna.
Characters: Gavia arctica is distinguished from G. immer and G.
adamsii by having a smaller size, and from G. stellata by Wetmore's (1940)
Remarks: Two species of Gavia ace described from the Miocene. Gavia
portisi (Regalia, 1902) of the Middle Miocene of Italy is known only from
a cervical vertebra nearly as large as that of Recent G. immer. Howard
(1978) describes the small G. brodkorbi based on an ulna from the late
Miocene of California. Measurements of this ulna are listed in Table 2
along with those of some other species of Gavia.
Gavia concinna Wetmore (1940) of the Middle Pliocene of California
and the Upper Pliocene of Florida is known by the ulna and by several
Table 2. Measurements of the ulna of Late Miocene Cavia brodkorbi
(Howard, 1978), Recent C. stellata, Recent G. arctica pacifica, Inglis
C. artica, Late Pliocene C. concinna, Recent C. immer, and Recent
C. brodkorbi C. stellata C. arc. paclfica
Total R 81.0 100.4-114.1 104.6-121.2
Length M 113.30
N 1 11 7
Proximal R 9.4 9.3-11.2 9.4-11.9
Width M 10.76
N 1 11 7
Midshaft R 4.9 4.4-5.5 4.7-5.6
Width M 5.11
N 1 11 7
Midshaft R 5.4-6.2 5.6-7.3
Depth M 6.39
N 11 7
Distal R 7.3 11.7-12.5 11.4-16.2
Depth M 13.83
N 11 7
Table 2. Extended.
other elements. The ulna of this loon is intermediate in size between
those of Recent G. stellata and G. immer, and overlaps that of G. artica
pacifica (Table 2). The ulna of the Inglis Gavia is slightly closer in
measurements to that of G. concinna, but in the characters listed by Wetmore
(1940), it resembles that of Recent G. arctica pacifica.
Wetmore's (1943c) Gavia palaeodytes from the Middle Pliocene of Florida
is not known for the ulna. However, other elements show it to be a bird
close in size to Recent G. stellata.
Gavia howardae Brodkorb (1953b) is recorded from the Middle Pliocene of
California, is not known for the ulna, and is smaller than G. stellata, G.
palaeodytes, and G. concinna.
Gavia arctica is known from Pleistocene deposits in Denmark (H. Winge,
1903) and California (Howard, 1949). This is the first fossil record of
this species in Florida. In recent times G. arctica has been recorded in
Florida four times (Hopkins and Woolfenden, 1977).
Order Podicipediformes (Furbringer)
Family Podicipedidae (Bonaparte)
Genus Podiceps Latham
Podiceps *dixi Brodkorb
Material: Distal portion of 1 right humerus, distal end of 1 left
ulna, one anterior sternal fragment. The 3 specimens represent a minimum
number of 1 individual.
Characters: Podiceps and Podilymbus are separated using characters
listed by Murray (1967).
The sternum of Podiceps differs from that of Podilymbus in having
medial area of coracoidal sulcus smooth (ventral lip projecting anteriorly
beyond dorsal lip in Podilymbus).
Remarks: Brodkorb (1963b) describes Podiceps dixi based on the
proximal end of a carpometacarpus from the Middle Pleistocene of
Reddick, Florida. Brodkorb's (1963b) measurements show Podiceps dixi
to be larger than living P. auritus, and smaller than living P.
cristatus, P. grisegena, and P. parvus (Shufeldt) from the Lower and
Middle Pleistocene of California and Oregon. Podiceps pisanus (Portis)
of Italy and P. subparvus (L. Miller and Bowman) of California, both
from the Middle Pliocene, are smaller than P. dixi (Brodkorb, 1963b).
Podiceps dixi is larger than Pliolymbus baryosteus Murray from the
Upper Pliocene, which Murray (1967) describes as a small robust grebe.
An ulna referred by Murray (1967) to Pliolymbus baryosteus Is shorter
than that of Podiceps nigricollis, which is much smaller than the
Inglis ulna of P. dixi.
Podiceps dixi is also larger than P. discors Murray of the Upper
Pliocene of Kansas and Idaho. Podlceps discors may have had a size
range similar to that of P. auritus (Murray, 1967).
Pliodytes lanquisti Brodkorb (1953a) is based on a coracold from
the Lower Pliocene of the Bone Valley Formation. The coracold is longer
than that of Podiceps auritus, shorter than that of Podiceps parvus
(Shufeldt), and close in size to that of Podilymbus podiceps.
Measurements of the Inglis Podiceps dixi are given in Table 3.
Table 3. Measurements of the humerus and ulna of Recent Podiceps
auritus (2e, 4?, 4?) and Inglis P. dixi.
P. auritus P. dixi
Midshaft R 3.9-4.6 4.3
Width M 4.16
N 8 1
Midshaft R 3.1-3.7 3.7
Depth M 3.40
N 8 1
Distal R 3.8-4.5 4.8
Depth M 4.15
N 10 1
Podiceps dominicus (Linnaeus)
Material: One complete left ulna.
Characters: The ulna of the Inglis Podiceps dominicus differs from
that of Recent P. dominicus in having (1) internal cotylar area flatter
(more excavated in modern form); (2) shaft more robust; and (3) external
condylar ridge concave at junction with shaft (convex in modern form).
Measurements of the fossil ulna are compared to those of Recent P.
dominicus in Table 4.
Remarks: Podiceps dominicus is known from the Pleistocene of Brazil
(0. Winge, 1887). This is the first fossil record of the Least Crebe for
Genus Podilymbus Lesson
Podilymbus podiceps (Linnaeus)
Material: One complete left humerus, proximal end of 1 left ulna,
distal end of 1 left ulna, proximal end of 1 right tibiotarsus, distal
ends of 2 right tibiotarsi. The 6 specimens represent a minimun number
of 2 individuals.
Characters: The humerus of Podilymbus differs from that of Podiceps
in having (1) larger bicipital crest; (2) deltoid crest extending farther
down shaft; and (3) brachial depression deeper.
The ulna in having (1) olecranon process broad; (2) intercotylar
region smooth (sharp ridge in Podiceps); (3) in internal view, edge of
internal cotyla smooth (notch at junction with olecranon in Podiceps);
Table 4. Measurements of
and Inglis P. dominicus.
the ulna of Recent Podiceps dominicus (lc, 3?)
(4) tendinal groove smaller; and (5) carpal tuberosity short and blunt
(long and sharp Podiceps).
The tibiotarsus in having (1) in external view, outer cnemial
crest forming a sharp angle on anterior side (curving in Podiceps);
(2) condyles much larger; and (3) distal width greater.
Remarks: Shufeldt (1913) describes Podilymbus magnus from the
Pleistocene of Fossil Lake, Oregon, as larger than P. podiceps. How-
ever, Wetmore (1937c) synonymizes P. magnus Shufeldt with P. podiceps.
Later, Brodkorb (1959a) revives P. magnus after measuring grebe fossils
from Arredondo, Florida. Measurements of Pleistocene grebes labelled
Podilymbus magnus from the Santa Fe River (in the Brodkorb Collection)
all fall within Storer's (1976) ranges for Recent P. podiceps. Finally,
Brodkorb (1963a, p.230) comments on Pleistocene Podilymbus podiceps,
Specimens from Fossil Lake and some of the Floridian
localities average large and are perhaps recognizable
as a temporal subspecies, P. podiceps magnus Shufeldt.
Storer (1976) compares measurements for 39 modern skeletons and
220 Late Pleistocene fossils of P. podiceps, and shows that there is
no significant size differences between the two groups. Measurements
of the Inglis fossils fit well with Storer's (1976) Pied-billed Grebe
measurements (Table 5).
Podilymbus majusculus Murray (1967) and P. parvus (Shufeldt, 1913),
both from the Upper Pliocene, are larger than P. podiceps. The Pleis-
tocene P. wetmorei Storer (1976) has thicker bones than the Pied-billed
Grebe. (Brodkorb, pers. com., suspects that P. wetmorei really ought to
be called P. podiceps magnus.) There appears to have been a decrease in
size in this genus since the Upper Pliocene.
Table 5. Measurements of the humerus, ulna, and tibiotarsus of Recent
Podilymbus podiceps (from Storer 1976; 23?' for humerus and tibiotarsus;
16Z for ulna) and Inglis P. podiceps.
Order Ardeiformes (Wagler)
Suborder Plataleae Newton
Family Plataleidae Bonaparte
Subfamily Threskiornithinae (Richmond)
Genus Eudocimus Wagler
Eudocimus albus (Linnaeus)
Material: Proximal end of 1 left humerus.
Characters: The humerus of Eudocimus albus differs from that of Ajaia
ajaja in having (1) smaller size; (2) deltoid crest more rounded; and (3)
bicipital crest more rounded.
The Inglis humerus is greatly abraded, making the usual osteological
measurements impossible. A few measurements are given in Table 6.
Remarks: Brodkorb (1963a) lists several Pleistocene sites in Florida
yielding E. albus fossils.
Xenicibis xympithecus Olson and Steadman (1977), a flightless ibis
from a Late Quarternary cave deposit on Jamaica, has a relatively small
Suborder Ardeae Wagler
Family Ardeidae Vigors
Genus Ardea Linaeus
Material: Anterior fragment of 1 sternum.
Characters: The sternum of Ardea n. sp. differs from those of
A. herodias and A. occidentalis in having (1) anterior crinal margin
with long, deep grooves on either side of midline ridge; (2) right
Table 6. Measurements of the humerus of Recent Eudocimus albus (2 d,4*)
and Inglis E. albus. Shaft measurements taken just distal to deltoid
albus E. albus
Head R 15.0-17.1 15.9
Length M 15.82
N 6 1
Head R 6.2-7.1 6.5
Width M 6.62
N 6 1
Shaft R 7.5-9.2 8.5
Width M 8.62
N 6 1
Shaft R 6.0-7.0 6.3
Depth M 6.43
N 6 1
corner of left coracoidal sulcus flat and without encircling ridge (area
depressed and ridge present in A. herodias and A. occidentalis); and (3)
carinal apex higher.
Measurements of the sternal fragment are given in Table 7.
Remarks: Ardea herodias has been recorded from several Pleistocene
sites in Florida (Brodkorb, 1963a).
Howard (1946) synonymized Ardea paloccidentalis Shufeldt with Botaurus
lentiginosus. Ardea polkensis Brodkorb (1955) of Lower Pliocene was
similar to, but smaller than, A. herodias.
Genus Ixobrychus Billberg
Ixobrychus n. sp.
Material: Distal end of i left cibiotarsus; distal end of 1 left
tarsometatarsus. The 2 specimens represent a minimum number of 1
individual. Both specimens are too worn for measurements.
Characters: Tibiotarsus having (1) size similar to that of Ixobrychus
involucris and I. sinensis (I. exilis smaller; I. minutus slightly larger;
I. cinnamomeus and I. sturmii much larger); (2) internal ligamental
prominence very flat, as in I. exilis (high and distinct in I. involucris,
I. sinensis and I. minutes); (3) tendinal bridge width as in I. sinensis
and I. exilis (bridge wider in I. minutus and I. involucris); and (4)
openings on either side of tendinal bridge large and rounded, as in I.
exilis and I. minutes (openings small and narrow in I. sinensis and I.
The tarsometatarsus (with trochleae crushed or lost) is slightly
larger than that of I. exilis (those of I. sinensis, I. involucris and
I. minutes all larger than the Inglis fossil). Ixobrychus cinnamomeus
Table 7. Measurements of the
(13, 5$) and Inglis Ardea sp.
sternum of Recent Ardea herodias
Length of right
Width of dorsal
Lip of right
Dorsal lips of
and I. sturmii tarcometatarsi are much larger that the Inglis tarso-
Remarks: Ritchie (1976) reports an Ixobrychus tibiotarsus from
Coleman IIA of the mid-Pleistocene of Florida, which is probably a new
species intermediate in size between I. exilis and Butorides virescens.
The Inglis tibiotarsus is the same size as the Coleman fossil and conforms
with the characters given by Ritchie (1976, 1980) The tarsometatarsus
is also a bit larger than that of I. exilis, and most likely belongs to
the same Ixobrychus species as the tibiotarsus.
Wetmore (1928a) reports I. exilis from the Pleistocene Ciego Montero
deposit of Cuba.
The Recent Least Bittern, I. exilis, ranges from southern Canada to
southern Peru (Blake, 1977), inhabiting fresh-water marshes with tall
grasses (Wetmore, 1965).
Genus Botaurus Stephens
Botaurus lentiginosus (Rackett)
Material: One left carpometacarpus, missing metacarpal III.
Remarks: Botaurus lentiginosus is known from several Pleistocene
localities in Florida (Brodkorb, 1963a).
The Upper Pleistocene Palaeophoyx columbians McCoy (1963) from the
Itchtucknee River, Florida, was restudied by Olson (1974a), who synonymized
it with Botaurus lentiginosus. Olson (1974a) comments that the Pleistocene
B. lentiginosus may have been a bit smaller than the modern form.
Measurements of the Inglis fossil are compared with those of Recent
B. lentiginosus in Table 8.
Table 8. Measurements of the carpometacarpus of Recent Botaurus
lentiginosus (6 e,5 ) and Inglis B. lentiginosus.
Order Anseriformes (Wagler)
Suborder Anseres Wagler
Family Anatidae Vigors
Subfamily Anatinae (Vigors)
Genus Aix Boie
Aix sponsa (Linnaeus)
Material: Proximal end of 1 right scapula, shaft of 1 right
coracoid, one furcula, I complete right humerus, distal end of 1
left carpometacarpus. The 5 specimens represent a minimum number of
Characters: The humerus agrees with those characters listed by
Ligon (1965) for distinguishing Aix from Spatula. Measurements of
the humerus of Aix sponsa appear in Table 9.
Remarks: No extinct species of Aix have been described.
Aix sponsa has been recorded for a number of Pleistocene
localities in Florida (Brodkorb, 1964a).
Genus Anas Linnaeus
Anas platyrhynchos Linnaeus
Material: Proximal end of 1 left humerus, fragment of 1 sternum.
Characters: Specimen agrees with characters of the humerus
listed by Woolfenden (1961) for Anas. Following Ligon (1965),
A. rubripes and A. fulvigula are combined with A. platyrhynchos.
The Inglis A. platyrhynchos differs from A. pachyscelus following
the characters in Wetmore's (1960) description.
Table 9. Measurements
Inglis A. sponsa.
of the humerus
of Aix snsa (8 e, 5*) and
Anas itchtucknee McCoy (1963) from the Upper Pleistocene of
Florida is slightly larger than Recent A. discors, and is therefore
much smaller the the Inglis A. platyrhynchos.
Janossy (1978) describes two species of Anas from the Lower
Pleistocene of the Carpathian Basin of Poland, Czechoslovakia, and
Austria. Anas submajor is the size of A. platyrhynchos, but
differs in the proportions of some long bones. Anas albae is a very
small duck, with a carpometacarpus shorter than that of A. crecca.
Campbell (1979) describes three species of Anas from the late
Pleistocene of the Talara Tar Seeps, northwestern Peru. Anas
talarae, A. amotape, and A. sanctaehelenae are all small ducks,
which Campbell compares with the four smallest living species of Anas:
A. leucophrys, A. brasiliensis, A. crecca, and A. cyanoptera.
Anas strepera Linnaeus
Material: Shaft of 1 right humerus, proximal end of 1 left ulna,
distal end of 1 right ulna, fragment of 1 left femur, distal end.
The 4 specimens represent a minimum number of 1 individual.
Characters: The medium-sized Anas strepera is distinguished from
A. acuta based on the characters of the ulna listed by Ligon (1965).
Remarks: Anas strepera is reported from the Itchtucknee River,
Florida (McCoy, 1963).
Anas discors (Linnaeus)
Material: One complete left humerus, distal shaft of 1 left
humerus, 1 complete left ulna, 1 right and 1 left carpometacarpus
(both lacking metacarpal III). The 5 specimens represent a minimum
number of 2 individuals.
Characters: Anas discors is distinguished from A. itchtucknee
based on the characters listed by McCoy (1963).
Measurements of the humerus, ulna, and carpometacarpus are given
in Table 10.
Remarks: The carpometacarpus of Anas albae Janossy (1978) has a
total length of 33.7 nun, which is smaller than the minimum length of
the carpometacarpus for A. discors (Table 10).
Campbell's (1979) Anas talarae and A. sanctaehelenae are smaller
than A. discors, and his A. amotape is about the same size.
However, the Inglis A. discors differs from the A. amotape following
Campbell's list of characters (1978).
Subfamily Aythyinae (Delacour and Mayr)
Genus Aythya Bole
Aythya americana (Eyton)
Material! One complete left ulna.
Characters: The ulna of Aythya americana is larger than those of
A. collaris, A. affinis, and A. fuligula, and is distinguished from
those of A. marila and A. valisineria by having (1) cotylae smaller;
(2) impression of brachialis anticus shallower; (3) carpal
tuberosity smaller; and (4) external condyle smaller.
Measurements of the fossil are compared with several species of
Aythya in Table 11.
Table 10. Measurements of the humerus, ulna, and carpometacarpus of
Anas discors (3d, 3g.) and Inglis A. discors.
Table 11. Measurements of the ulna of Aythya collaris (7a,21-), A. affinis ( '4$), A. fuligula (4J,1,),
A. marila (5d,64), Inglis A. americana, A. americana (3d',10), and A. valisii ia (2?;,6g).
Aythya Aythya Aythya Aythya Inglis
collaris affinis fuligula marila Aythya
Remarks: The five known extinct species of Aythya are recorded
from the Upper Pleistocene of Australia (A. robusta, A. reclusa, and
A. effodiata) and the Upper Pliocene of Italy (A. aretina and
Subfamily Tadorninae Reichenbach
Genus *Anabernicula Ross
Anabernicula gracilenta Ross
Material: One right quadrate, 1 left tarsometatarsus (lacking
hypotarsus and inner trochlea). The 2 specimems represent a minimum
number of 1 individual.
Characters: Howard (1964a) lists characters of the tarsometa-
tarsus used to separate Anabernicula from Tadorna. Anabernicula
gracilenta is distinguished from A. oregonensis by having a slightly
smaller size. See Table 12.
Remarks: Four species of Anabernicula are known. Anabernicula
minuscule (Wetmore, 1924) from the lower Pleistocene of Arizona is
known only by the proximal half of the humerus. Anabernicula
oregonesis Howard (1964a) is recorded from the late Pleistocene of
Oregon, and is a slightly heavier bird than A. gracilenta.
Anabernicula gracilenta is recorded from the late Pleistocene of
California, Nevada, and New Mexico. Short (1970a) describes the stocky
A. robusta from the middle Pleistocene of Nebraska based on a
Howard (1963) describes another extinct genus of the subfamily
Tadorninae, Brantadorna downsi from the middle Pleistocene of
California. Brantadorna is distinguished from Anabernicula based on
characters of the humerus showing it to be more gooselike than
Table 12. Measurements of the tarsometatarsus of Rancho La Brea Anaber-
nicula gracilenta (Howard, 1964). Inglis A. gracilenta, A. oregonensis
Howard, (1964), and Recent Tadorna ferruginea (4 females).
A. grac. Inglis A. oreg. Tadorna
A. grac. ferruginea
Total R 56.0-63.3 61.6 59.2-62.8
Length H 58.9 61.12
N 5 1 4
Proximal R 9.9-11.7 11.7 11.2-12.0
Width M 10.7 11.62
N 9 1 4
Midshaft R 5.2 4.7-5.6
Width M 5.05
N 1 4
Midshaft R 4.9 4.3-5.2
Depth M 4.65
N 1 4
Width of R 4.5-5.3 5.7
Internal M 4.8
Condyle N 7 1
Depth of R 5.4-6.2 6.5
Internal M 5.7
Condyle N 7 1
Width R 11.3-13.0
Through M 12.25
Trochleae N 4
Order Accipitriformes (Viellot)
Suborder Sarcoramphi (Ridgway)
Family Vulturidae (Illiger)
Subfamily Vulturinae (Illiger)
Genus Gymnogyps Lesson
cf. Gymnogyps californianus (Shaw)
Material: Proximal end of one left tibiotarsus.
Characters: The cnemial crests of this fossil are worn, making
application of Fisher's (1945) characters for separating Gymnogyps
and Teraternis useless. Table 13 shows considerable overlap in the
proximal width of the Gymnogyps and Teratornis tibiotarsi. Howard's
(1974) detailed comparison of Breagyps, Vultur, and Gymnogyps
likewise cannot be used to identify this fossil, and she gives no
proximal width measurements for tibiotarsi. However, these genera
can be eliminated on the basis of their generally larger size with
respect to Gymnogyps. Modern Sarcoramphus papa is much smaller
than the fossil (Table 13).
Measurements in Table 13 show the Inglis bone to be at the mini-
mum size for Recent G. californianus. Measurements of the proximal
end of G. amplus have not been published. However, Fisher's (1947)
tibiotarsus measurements of total length, distal width, and fibular
crest length show the two species to be inseparable in size.
Remarks: The specimen is referred to the extant species Gymnogyps
californianus rather than to the extinct G. amplus L. Miller. Fisher
(1944) considers G. amplus to be the ancestral form of G. californianus,
and lumped all Pleistocene Gymnogyps into the species G. amplus. Loye
Table 13. Proximal width of the tibiotarsus of modern Sarcoramphus
papa, Inglis cf. Gymnogyps californianus, Recent G. californianus
(Fisher, 1947), Pleistocene Teratornis merriami (Fisher, 1945), and
Pleistocene Vultur (Campbell, 1973).
26.2 26.2-29.4 26.6-31.2 30.0-31.1
Miller (1957) does not agree with this move. Fisher's (1947) study
of Recent and fossil Gymnogyps reveals no qualitative or quantitative
differences in postcranial G. amplus or G. californianus bones (see
Table 14). Howard's (1974) skeletal measurements of G. californianus
show even more overlap in their size ranges than Fisher's (1947) paper
did. The only differences between these species are the characters of
the skull listed by Fisher (1944), but Wetmore (1959, p.12) remarks,
"even here the distinction is not clear-cut." Brodkorb (1964a) suggests
that G. amplus is a temporal subspecies of G. californianus. It seems
appropriate to here synonymize G. amplus with G. californianus.
Three species of condor are known from the Florida Pleistocene.
Gymnogyps amplus is recorded for the middle Pleistocene of Reddick by
Brodkorb (1957), and for the Upper Pleistocene of the Itchtucknee River
by McCoy (1963). Wetmore (1931.) lists G. californianus for the Upper
Pleistocene of Sarasota. Teratornis merriami is known from the Upper
Pleistocene of Seminole Field and Manatee County (Wetmore, 1931b).
The tibiotarsus is not represented for several fossil vulture
species. Pliogyps fisher Tordoff (1959) of the Upper Pliocene of Kan-
sas is known only from the tarsonetatarsus, and is smaller than Vultur,
Breagyps, and Cymnogyps. The Pliocene Vultur patruus (Lnnnberg, 1902)
of Bolivia is also known solely from the tarsometatarsus, as Is the
long and slender Cathartornis gracilis L. Miller (1910) of the Upper
Pleistocene of Rancho La Brea. Only the humerus is known of Sarco-
ramphus kernense (L. Miller, 1931) from California's middle Pliocene.
Antillovultur varonai Arredondo (1971 and 1976) from Cuba's late
Pleistocene is known from the tarsometatarsus and humerus, and is equal
in size to the Andean Condor, Vultur gryphus. The Antillovultur
tarsometatarsus is longer and more robust than that of Gymnogyps.
Table 14. Measurements of the tarsometatarsus of mid-Pleistocene
Gymnogyps amplus from Reddick, lage Pleistocene G. amplus from
Rancho La Brea (Fisher, 1947 and L. Miller, 1910), and Recent G.
californianus (Fisher, 1947).
G. amplus G. amplus G. calif.
Reddick Rancho La Brea Recent
Total R 126.9 113.0-134.0 120.0-131.4 113-117
Length M 123.00 123.70 114
N 1 100 14 6
Proximal R 29.3 26.5-31.7 26.5-30.1 25.5-28.0
Width M 28.50 28.01 26.6
N 1 86 14 6
Width R 30.8 29.5-34.5 30.7-33.2 28.3--30.2
Through M 32.3 32.09 29.40
Trochleae N 1 97 14 6
Genus Coragyps Geoffroy
Coragyps atratus (Bechstein)
Material: One axis, proximal end of 1 left scapula, proximal end of
1 left coracoid, 2 complete right coracoids, distal ends of 1 left and I
right humeri, proximal ends of 2 left and 1 right ulnae, distal ends of
2 left ulnae, 1 right ulnare, 1 right radiale, proximal end of I left
carpometacarpus, distal end of 1 right tibiotarsus, proximal end of 1
left tarsometatarsus. The 17 specimens represent a minimum number of 2
Characters: The tibiotarsus and tarsometatarsus conform to Cracraft
and Rich's (1972) diagnosis of the "Cathartidae" (Vulturidae). Coragyps
atratus is easy to distinguish from Cathartes aura, the Turkey Vulture,
in most elements by many characters.
The carpometacarpus proximall end) of Coragyps differs from that of
Cathartes in having (1) proximal process of metacarpal I narrow and
pointed (wide and rounded in Cathartes); (2) area between distal end of
tendinal groove and proximal edge of internal condyle narrow (wide in
Cathartes); and (3) prominence exterior to tendinal bridge low and blunt
(prominence sharp in Cathartes).
The tarsometatarsus proximall end) in having; (1) in proximal view,
posterior edge of hypotarsus straight (curved in Cathartes); and (2) in
anterior view, lacks prominence distal to external condyle.
Measurements are given in Table 15 comparing the Inglis fossils
with Coragyps occidentalis (L. Miller) from Reddick, C. occidentalis
occidentalis from Rancho La Brea (Howard, 1968), C. occidentalis mexi-
canus from San Josecito Cave (Howard, 1968), Recent C. atratus atratus
Table 15. Measurements (mm.) of the coracoid, humerus, ulna, and
tibiotarsus of early Pleistocene Coragyps atratus from Inglis,
middle Pleistocene C. occidentalis from Reddick, late Pleistocene C.
o. occidentalis from Rancho La Brea (Howard, 1968), late Pleistocene
C. o. mexicanus from San Josecito Cave, Mexico (Howard, 1968),
Recent C. atratus atratus from Florida and California (Howard, 1968)
and Recent C. atratus brasiliensis from Panama. *-surface worn.
R-range, M-mean, N-nmbcr.
Coragyps atratus C. occidentalis C. o. occi.
Inglis Reddick RLE
Length to int. R 59.2 64.4-65.4 60.3-69.4
Sternal facet M 64.90 64.5
N 1 2 70
Length to ext. R 65.5
Sternal facet M 1
Head to R 25.3-26.3 27.9-29.9
Scapular facet M 25.80 28.77
N 2 3
Midshaft R 7.4-8.3
Width M 7.85
Midshaft R 6.0-6.3
Depth M 6.15
Midshaft R 10.5
Width M -
Midshaft R 8.8
Depth M -
Distal R 23.7-24.2 26.1-29.1
Width M 23.95 27.4
N 2 50
Table 15. Extended.
C. o. mexicanus
San Josecito Cave
C. atratus atratus
C. a. brasiliensis
Table 15. Continued.
C. o. occi.
Table 15. Extended.
C. o. mexicanus
San Josecito Cave
C. atratus atratus
C. a. brasiliensis
from Florida and from Howard's (1968) paper (from the United States,
Mexico, and Guatemala), and Recent C. atratus brasiliensis from
Panama, (including Oaxaca, Mexico, and Guatemala specimens from Howard,
pers. com.). These measurements show an increase and then a decrease
in average size from early Pleistocene to Recent. However, there are
greatly overlapping ranges in the coracoid, humerus, ulna, and
Remarks: In Florida, C. occidentalis is listed for Reddick
(Brodkorb, 1957) and Haile (Ligon, 1965). Coragyps occidentalis is
a supposedly heavier-boned species than C. atratus, but Howard's
(1968, p.116) "Limb measurements of the extinct vulture, Coragyps
Qualitative distinctions from bones of the living
C. atratus are not evident in the fossil limb bones;
both species exhibit considerable variation. Differ-
ences lie in over-all size and proportions of the
However, Ligon's (1965) tibiotarsus measurements of C. occidentalis
are within the range of modern C. atratus.
Geographical variation in Recent Coragyps atratus has been dis-
cussed by Brodkorb (1944) and Wetmore (1962). The three subspecies
include the large C. atratus atratus, of the North American temperate
region, the small C. atratus brasiliensis of the tropical region, and
the large C. atratus foetens of the South American temperate region.
According to Howard (1968), there was a similar pattern of varia-
tion in the late Pleistocene. Coragyps o_ occidentalis was the larger
North American form, and C. o. mexicanus was the smaller tropical
There appears to have been a gradual increase, then decrease, in
Coragyps' size from early Pleistocene to Recent, plus geographical
variation in size at least in late Pleistocene to Recent. The size
fluctuation indicates climatic changes from warmer to cooler and back
to warm again in Recent tires.
In view of the lack of qualitative characters to separate C.
occidentalis and C. atratus, and the broad overlap in measurements, it
seems appropriate to synonymize C. occidentalis with C. atratus.
The Inglis bones are the oldest known for this genus.
Genus Cathartes Illiger
Cathartes aura (Linnaeus)
Material: One left quadrate, proximal end of 1 left
tarsometatarsus. The 2 specimens represent a minimum number of 1
Characters: The identification of the tarsometatarsus is based on
the characters listed under the section on Coragyps atratus.
Measurements given in Table 16 compare these fossil elements with
those of nine modern specimens of Cathartes aura.
Remarks: Cathartes aura is known from many Pleistocene sites in
Florida (Brodkorb, 1964a). Inglis is the oldest fossil locality for
this genus and species.
Table 16. Measurements of the quadrate and tarsometatarsus of Inglis
Cathartes aura and Recent, C. aura (4 males, 5 females).
Suborder Accipitres (Vieillot)
Family Accipitridae (Vieillot)
Subfamily Buteoninae (Vigors)
Genus Buteo Lacepede
Buteo n. sp.
Holotype: One complete left carpometacarpus.
Paratypes: Proximal end of 1 left ulna, proximal end of 1 left
carpometacarpus, distal end of 1 left tibiotarsus, proximal end of 1
left tarsometatarsus, distal end of 1 left tarsometatarsus.
Referred material: One left digit II of phalanx of manus, 2 left
acetabular fragments, proximal end of 1 left tarsometatarsus, 1
phalanx I of digit I, I ungual phalanx each of digits I, II, III and
The 14 specimens represent a minimum number of 2 individuals.
Locality and age: Inglis IA, Citrus County, Florida. Early
Diagnosis: Carpometacarpus agrees with Buteo in the characters
listed by Campbell (1973, p. 98). It is larger than those of B.
magnirostris, B. platypterus, B. nitidus, B. lineatus, B. lagopus, and
B. swainsoni, and is slightly smaller than B. regalis. It agrees in
size with that of B. jamaicensis (Table 17).
The carpometacarpus differs from those of Buteo jamaicensis and
B. regalis in having (1) distal edge of pollical facet forming a broad
U-shape; (2) ridge between metacarpal I and external trochlea longer
and thinner; (3) internal ligamental fossa deeper; (4) internal carpal
trochlea shorter; (5) depression between pisiform process and
metacarpal I deeper; (6) tendinal groove wider; (7) in internal view,
wide area at distal end of metacarpal II more depressed.
Table 17. Measurements (mm) of the carpometacarpus of Buteo magnirostris,
B. platypterus, B. nitidus, B. lineatus, B. logopus, B. swainso'i, B.
jamaicensis, Inglis Buteo n. sp., and B. regalis.
(2 males, 2
females, 2 ?)
of Meta- M
carpal I N
(1 male, 3
Table 17. Extended.
(6 males, 6
females, 2 ?)
n. sp. regalis
Characters: The proximal end (76.8 mm. long) of the ulna differs
from those of Buteo jamaicensis and B. regalis in having (1) olecranon
process higher; (2) depression of internal side of olecranon deeper;
(3) intercotylar ridge higher; (4) tricipital attachment wider; (5)
bicipital attachment forming one elongated scar (forms 2 short scars
in B. jamaicensis and B. regalis); (6) impression of brachialis antics
bordered on internal side by a sharp ridge; (7) nutrient foremen much
larger. The proximal width of the Inglis Buteo n. sp. ulna is 16.9 mm.
The tibiotarsus differs from those of B. jamaicensis and B.
regalis in having (1) tendinal groove more medial (closer to external
condyle in B. jamaicensis and B. regalis); (2) anterior intercondylar
fossa shallower; (3) dismal edge of supratendinal bridge convex
(straight in B. jamaicensis and B. regalis). Measurements of the
tibiotarsus (Table 18) show it to be larger than those of B.
jamaicensis and B. regalis.
The tarsometatarsus in having (1) inner proximal foramen larger;
(2) in anterior view of proximal end, external edge of shaft nearly
straight (curves inward in B. jamaicensis and B. regalis); (3) area
proximal to tubercle for tibialis anticus relatively shallow, as in B.
regalis; (4) internal edge of shaft thicker proximally; (5) facet for
metatarsal I much larger and wider; (6) condyles larger (sternal
condyle missing in fossil). Tarsometatarsus measurements of several
Buteo are shown in Table 18. The referred piece of a tarsometatarsus
agrees with all the above characters except for number (4).
The carpometacarpus of Buteo n. sp. is closer in size to that of
B. jamaicensis than to that of B. regalis, while the tibiotarsus and
tarsometatarsus are slightly larger than those of B. regalis (Tables 17
The Inglis Buteo n. sp. is perhaps ancestral to B. jamaicensis,
as it shows more resemblance to that species in the characters listed
Measurements of the phalanges are shown in Table 19.
Remarks: Buteo typhoius Wetmore of the Upper and Lower Miocene
of Nebraska is known from the distal part of a tarsometatarsus and the
distal part of a tibiotarsus (Wetmore, 1923 and 1928b). The tibiotarsus
B. typhoius (from a juvenile individual) differs from that of Butteo n.
sp. in having (1) slightly smaller size, and (2) considerably different
shape and orientation of the tendinal bridge. The tarsometatarsus of
B. typhoius is larger than that of B. jamaicensis and Buteo n. sp.
The tarsometatarsus of Buteo contortus (Wetmore) from Nebraska's
Upper Miocene is much larger than that of Buteo n. sp., as is readily
seen from Wetmore's (1923) drawings and measurements.
The Lower Pliocene hawk from Nebraska, Buteo dananus (Marsh), was
described from the distal end of a tibiotarsus as being "nearly as
large as the Golden Eagle" (Marsh, 1871). Wetmore (1923) gives its
width across the condyles as just over 16 mm. The tibiotarsus of
Buteo n. sp. is distinguished from that of B. dananus, based on the
photograph of the type (Shufeldt, 1915), by having (1) proximal edge of
supratendinal bridge parallel with the long axis of the tibiotarsus
(edge makes an acute angle in B. dananus); (2) distal edge of supra-
tendinal bridge at a 450 angle to the long axis of the bone (edge is
almost perpendicular to long axis in B. dananus); and (3) distal
opening of tendinal groove triangular in shape (opening is nearly
spherical in B. dananus).
Table 19. Measurements of phalanx I of digit I and the ungual
phalanges of digits I, II, III, and IV of 1 Recent Buteo
jamaicensis (2 c,2 ?-) and Inglis Buteo n. sp.
Phalanx I of digit I
Ungual phalanx of digit I
of digit II
Ungual phalanx of digit III
of digit IV
Buteo n. sp.
Buteo conterminus (Wetmore), also of Nebraska's Lower Pliocene, is
even more massive than B. contortus in the size of the tarsometatarsus
(Wetmore, 1923), and is clearly not the same species as Buteo n. sp.
Two new genera of Pleistocene hawks from South America, Miraquila
terrestris Campbell and Amplibuteo hibbardi Campbell (1979) are
gigantic in comparison to Buteo n. sp. Campbell's (1979) Buteo sp. 1
and Buteo sp. 2 (not illustrated in his paper) are both smaller and
differ qualitatively from the Inglis Buteo n. sp.
Genus Aquila Brisson
Aquila n. sp.
Holotype: Complete right carpometacarpus.
Paratypes: Proximal end of I left humerus, proximal end of 2 left
ulnae, distal end of 1 left ulnae, proximal ends of 2 right and I left
radii, distal end of 1 left carpometacarpus, 1 right and 1 left pollex,
1 left digit [II of manus, 1 complete left femur, proximal end of 1
right femur, 1 complete right tibiotarsus, proximal end of 1 right
fibula, 1 right metatarsal, 1 subterminal phalanx of digit I, 1 ungual
phalanx of digit I, I phalanx II of digit II, 1 phalanx I of digit III,
1 phalanx III of digit III, 1 phalanx I of digit IV, and 1 ungual
phalanx of digit IV.
Referred material: Anterior end of 2 mandibles, posterior end of
1 left mandible, proximal end of 1 sternum, proximal end of 2 left
scapulae, midsection of 1 left coracoid, distal fragment of 1 right
humerus, distal end of 1 left radius, proximal fragment of 1 right
femur, and distal fragment of 1 right femur.
The 34 specimens represent a minimum number of 4 individuals.
Generic characters: Carpometacarpus agrees with that of Aquila
in having (1) in ventral view, carpal trochleae set far apart, with
depression for ulnare relatively flat, as in Spizaetus and Parabuteo
(trochleae closer together and depression for ulnare a rounded pit in
Haliaeetus, Heterospizias, and Buteo; very far apart in Accipiter;
very close in Buteogallus); (2) ligamental attachment of pisiform
process distinct, broad, and forming a deep pit anterior to pisiform
process, as in Accipiter, Spizaetus, and Heterospizias (smaller, less
distinct, and a shallow pit in Haliaeetus and Buteo; no pit in
Buteogallus and Parabuteo); (3) in internal view, tendinal groove
short and broad, and extending proximally from proximal end of
intermetacarpal space (groove long and narrow, coming to a distinct
point at proximal end in Haliaeetus and Accipiter; indistinct in
Buteogallus, Buteo, Spizaetus, Heterospizias, and Parabuteo); (4)
proximal fused area between metacarpal II and metacarpal III of medium
length, as in Accipiter, Buteogallus, Spizaetus, and Parabuteo (long
in Haliaeetus; very short in Buteo and Heterospizias); (5) tendinal
groove strongly curved around edge of shaft towards metacarpal I, as
in Accipiter, Buteogallus, Buteo, Spizaetus, Heterospizias, and
Parabuteo (groove more nearly parallel to shaft in Haliaeetus); distal
end of intermetacarpal space a broad U-shaped region, as in Buteo
(narrow U-shape in Haliaeetus, Buteogallus, Spizaetus, ieterospizias,
and Parabuteo; V-shape in Accipiter).
Humerus agrees with that of Aquila in having attachment of infra-
spinatus forming a long, deep groove (attachment flat and oval-shaped
with no such groove in Haliaeetus; short groove in Accipiter,
Buteogallus, Buteo, Spizaetus, Heterospizias, and Parabuteo).
Ulna agrees with that of Aquila in having (1) ridge separating
internal and external cotylae high (very low ridge in Haliaeetus and
Heterospizias; sharp prominence in Accipiter and Spizaetus; low ridge
in Buteogallus, Buteo, and Parabuteo); (2) bicipital scar more
distally placed than in Haliaeetus, Accipiter, Buteogallus, Buteo,
Spizaetus, Heterospizias, and Parabuteo.
Femur agrees with that of Aquila in having (1) anterior
intermuscular line extending much farther proximally than proximal
foramen, as in Accipiter, Buteogallus, Buteo, Spizaetus, Heterospizias
and Parabuteo (line stops at about level of foramen in Haliaeetus);
(2) posterior intermuscular line extending much farther proximally
than nutrient foramen, as in Accipiter, Buteo, Spizaetus ornatus, and
Heterospizias (stops at about level of foramen in Haliaeetus,
Buteogallus, Spizaetus tyrannus, and Parabuteo); (3) medial edge of
internal condyle rounded, as in Accipiter, Buteogallus, Buteo,
Spizaetus, Heterospizias, and Parabuteo (tapered in Haliaeetus);
external wing of fibular condyle thrust laterally, as in Accipiter,
Spizaetus, and Heterospizias (more compressed in Haliaeetus,
Buteogallus, Buteo, and Parabuteo); (5) external condyle large and
rounded, as in Accipiter, Buteogallus, Spizaetus, Heterospizias, and
Parabuteo (smaller and squared-off on proximal edge in Haliaeetus and
Buteo); (6) anterior view, ridges formed by internal and external
condyles extending proximad to same level, as in Buteogallus and
Spizaetus (ridge of external condyle extends much farther proximally
than ridge of internal condyle in Haliaeetus, Accipiter, Buteo,
Heterospizias, and Parabuteo).
Tibiotarsus agrees with that of Aquila in having (1) region
between proximal edge of supratendinal bridge and external condyle
wide, as in Spizaetus (narrow region in Haliaeetus, Accipiter, and
Buteo; area of medium width in Buteogallus, Heterospizias, and
Parabuteo); (2) in anterior view, proximal end of internal condyle
rounded, as in Accipiter, Buteogallus, Buteo, Heterospizias,
Parabuteo, and Spizaetus proximall end forms a point directed toward
supratendinal bridge in Haliaeetus); (3) in posterior view, internal
condyle longer and its internal ridge parallel with long axis of bone,
as in Accipiter, Buteogallus, Buteo, Heterospizias, Parabuteo, and
Spizaetus (internal condyle shorter and ridge angled toward center of
bone in Haliaeetus); (4) internal ligamental prominence low, as in
Spizaetus (high in Haliaeetus, Accipiter, Buteogallus, Buteo,
Heterospizias, and Parabuteo.
Tarsometatarsus characters separating Aquila and Haliaeetus are
listed by L. Miller (1911).
Diagnosis: Carpometacarpus differs from that of Aquila
chrysaetos in having (1) anterior carpal fossa deeper; (2) large
foramen at external base of pollical facet (not present in A.
chrysaetos); (3) proximal, internal edge of metacarpal III has
distinct muscle scar that curved towards internal carpal trochlea
(scar less distinct and straighter in A. chrysaetos); (4) distal end
of scar described above comes to a prominence (smooth scar in A.
Specific characters: Aquila n. sp. differs from Recent Aquila
chrysaetos canadensis in having humerus with (1) much larger
tricipital fossa; (2) median crest forming a broad U-shaped curve
proximally (narrow curve in A. chrysaetos).
Ulna with (1) proximal radial depression deeper and broader; (2)
small prominence present just proximal to tendinal pit (not present in
Radius with (1) shaft heavier; (2) bicipital tubercle set farther
distally; (3) proximal width 13.3-13.8 mm. (x = 13.53, N = 3); (4)
proximal depth 9.3-9.7 mm. (x = 9.53, N = 3).
Femur with (1) in anterior view, two small, proximal foramina and
one large distal foramen to crochanteric ridge (only one large distal
foramen in A. chrysaetos); (2) large foramen internal to trochanteric
ridge elongated (more rounded in A. chrysaetos); (3) popliteal area
deeply depressed (flat in A. chrysaetos); (4) two foramina in center
of popliteal are a (set at distal end of popliteal area in A.
Tibiotarsus with (1) in anterior view, ridge of inner cnemial
crest does not extend as far down shaft; (2) intermuscular line nearly
absent (very prominent in A. chrysaetos).
Measurements of the scapula, metatarsal I, and some of the
phalanges are given in Table 20 for the Inglis Aquila n. sp. and for
Recent A. chrysaetos. Table 21 lists measurements of the carpometa-
carpus and femur of these species from several localities. These
measurements show the Inglis bird to be larger than any known Golden
Eagles of past or present.
Remarks: Wetmore (1937b) described two new genera of eagles from the
Pleistocene of Great Exuma Island in the Bahamas: Calohierax
Table 20. Measurements (mm.) of the scapula, tibiotarsus, metatarsal I,
phalanx I of digit I, and ungual phalanx of digit I, phalanx II of
digit II, phalanx I of digit III, phalanx III of digit III, phalanx I
of digit IV, and ungual phalanx of digit IV, from Recent Aquila
chrysaetos (3 8,1 $,1?) and Inglis Aquila n. sp.
Length from head
Ungual phalanx of digit I
Proximal width R
Proximal depth R
Aquila n. sp.
Table 20. Continued.
Phalanx II of digit II
Phalanx I of digit III
Distal width R
Phalanx III of digit III
Proximal width R
Aguila n. sp.
Table 20. Continued.
Phalanx I of digit IV
Ungual phalanx of digit IV
Aquila n. sp.
Table 21. Measurements of the carpometacarpus and femur from early
Pleistocene A. chrysaetos bonifacti from Saint-Esteve, France (Mourer-
Chauvire, 1975), late Pleistocene A. chrysaetos from Rancho LaBrea
(Howard, 1932), and Recent A. chrysaetos from North America (Brodkorb
Collection and Howard, 1932), and from Europe (Mourer-Chauvire', 1975).
Aquila n. sp.
A. c. bonifacti
Rancho La Brea
Height of R
Metacarpal I M
Width of M
Metacarpal II N
Depth of M
Metacarpal II N
Table 21. Extended.
Recent Aquila chrysaetos
Aqujiia n. sp. A. c. bonifacti
Rancho La Brea
Table 21. Extended.
Recent la chrysaetos
North America France
resembles Buteo in some characters and in size; the gigantic
Titanohierax resembles Recent Buteogallus urubitinga, but is nearly
twice as large. The Inglis Aquila n. sp. carpometacarpus appears to
be about 30% larger than the natural-size photograph of the
Titanohierax carpometacarpus. Wetmoregyps daggetti (L. Miller) from
the Upper Pleistocene of California and Mexico also bears a
resemblance to the genus Buteogallus (L. Miller, 1928).
Aquila borrasi Arrendondo of Cuba's late Pleistocene is known
from the tarsometatarsus, femur, and the ungual and subterminal
Table 22 gives femur measurements of A. borrasi and the Inglis
Aquila n. sp., which show A. borrasi to be a much larger species of
eagle. Visual comparison with the photograph in Arredondo's paper
(1970) confirms this. Aquila n. sp. more closely resembles A.
chrysaetos and differs from A. borrasi based on Arredondo's (1976)
characters of the femur for separating these two species.
Other fossil species of Aquila are known from the Upper Miocene
of France (A. delphinensis Gaillard and A. pennatoides Gaillard).
Fossil A. chrysaetos are known from many localities in Europe and
North America (Brodkorb, 1964a).
Mourer-Chauvire (1975) described Aquila chrysaetos bonifacti from
the Middle Pleistocene of Saint-Esteve, France, which she considers to
be the primitive form of today's Golden Eagle. No morphological
differences are found between these two forms, although statistical
analyses show allometric difference in the proportions of certain
bones. Also, the Saint-Esteve bird is almost always smaller than the
Recent A. chrysaetos. Mourer-Chauvire (1975) concludes that there was
Table 22. Femur measurements (mm.) of Aquila borrasi (Arredondo, 1976)
and Aquila n. sp. length as preserved is 140.0 mm.
Maximum width of pneumatic foramen
Length of pneumatic foramen
Least width of shaft
Aquila n. sp.
an increase in size of Golden Eagles throughout the course of the
Pleistocene in France.
Loye Miller (1911) remarks that the Rancho La Brea A. chrysaetos
has remained nearly unchanged to this day. Howard's (1947) study of
Rancho La Brea A. chrysaetos measurements indicate that Pleistocene
humeri and ulnae averaged longer, while tarsometatarsi averaged
shorter than Recent forms, although no proportional or structural
differences are evident.
Genus Buteogallus Lesson
Buteogallus *fragilis (L. Miller)
Extinct Fragile Eagle
Material: Proximal two-thirds of 1 left tarsometatarsus.
Characters: The fossil is placed in the genus Buteogallus and
separated from Geranoaetus, Harpyhaliatus, Spizaetus, Aquila, Buteo,
and Heterospizias based on characters number 3, 5, and (page 96) of
Tarsometatarsus larger, but resembling living Buteogallus
urubitinga in having (1) posterior metatarsal groove very deep for
most of the bone's length, and bordered by high ridges on internal and
external sides; (2) anterior proximal foramena set in a scooped-out
depression; (3) in posterior view, anterior end is cocked slightly
towards external side, with a "corner" formed on internal side at
level of attachment for tibialis antics; (4) edge of shaft external
to posterior metatarsal groove a distinct ridge, greatly flared
externally. Agrees with Buteogallus fragilis in characters and
measurements listed by L. Miller (1911). The type specimen
measurements are (1) proximal width 18.0 mm, (17.7 for Inglis fossil);
(2) length to center of papilla of tibialis anticus 18.6 mm (18.2 for
Remarks: This is the oldest fossil for the genus Buteogallus
which was previously known from the Upper Pleistocene of California
and Mexico (Howard and A. Miller, 1933).
Wetmore (1931a) records three fragmentary eagle ulnae from the
late Pleistocene of Seminole Field, Florida which he called
Geranoaetus sp. In his discussion section Wetmore mentions
Pleistocene G. fragilis (now Buteogallus fragilis), Pleistocene G.
grinelli), and Recent G. melanoleucas, the Black-chested Buzzard-Eagle
of South America. No characters or measurements are given. However,
Wetmore states that these ulnae agree in size with G. melanoleucas.
The elongated, narrow tarsometatarsus with attachment of the
tibialis anticus placed high on the bone indicated a relatively
weak-legged predator with poor lifting power.
The related Great Black Hawk of South America, Buteogallus
urubitinga, is described by Blake (1953, p.86) as, ". . decidedly
sluggish . soars at length . more abundant in the hot lowlands,
especially near water." Amadon (1949) also notes B. urubitinga living
near water, and remarks that it usually feeds upon reptiles,
amphibians, or crabs. Blake (1977) lists southern Mexico through
northern Argentina for this species' range.
Subfamily Gypaetinae (Vieillot)
Genus *Neophrontops L. Miller
Neophrontops slaughter Feduccia
Material: Left side of 1 mandible (without mandibular
symphysis), 1 left coracoid, distal end of 1 right ulna; proximal end
of 1 left radius; distal end of 1 left radius; 1 right femur,
proximal half of 1 left tibiotarsus, distal half of 1 right
tibiotarsus, proximal half of 1 right tarsometatarsus, distal half of
1 right tarsometatarsus.
Referred material: Shaft of 1 right radius.
The 11 specimens represent a minimum number of 2 individuals.
Characters: The mandible differs from those of recent Neophron
percnopterus ginginianus and Necrosyrtes monachus monachus in having
(1) larger size; (2) internal, posterior and exterior articular
processes relatively longer; (3) ramus longer and wider; (4) medial
lip of articular surface protruding with a broad, curved edge; (5)
internal articular process projecting more laterally (projects
dorsally in Neophron and Necrosyrtes.
Measurements of the mandible of several members of the subfamily
Gypaetinae are listed in Table 23. The mandible of Neogyps errans
L. Miller of the Upper Pleistocene resembles those of Neophron and
Necrosyrtes in characters (2) and (4) listed above, and is larger than
the Inglis bone. The fragment of mandible of Palaeoborus umbrosus is
larger and more robust than that of the Inglis Neophrontops slaughter,
while the mandible of Neophrontops americanus L. Miller is slightly
Table 23. Measurements of the mandible and femur of Neophron percnopterus (2*). Necrosyrtes
monachus (sex ?), Neophrontops americanus (Howard 1932), Palaeoborus umbrosus (Cope, 1877 ,
Inglis Neophrontops slaughter, and Neogyps errans (Howard, 1932). measured from photo-
graph in Howard (1932).
Neophron Necros Neophontops Palaeo
MANDIBLE percnop. monachus americanus umbrosus
Width, int. to
Depth of ramus
Width of Head
Table 23. Continued.
smaller than it. Neophrontops americanus is in close agreement with
N. slaughter in characters (2) and (4) above, based on Howard's
The left coracoid lacks the head, tip of procoracoid process, and
the sterno-coracoidal process. Howard's (1932) coracoid lengths for
112 N. americanus range from 49.5 to 56.2 mm. Howard divided the
distance from the sternal facet to the procoracoid by the total length
of the bone to get an average ratio of 65.7% for N. americanus (56.8%
for Neophron percnopterus). The distance from the sternal facet to the
procoracoid of the Inglis Neophrontops slaughter is 40.3 mm. The
total length could not be measured. However, a rough calculation
gives a total length of about 59.8 mm (40.3 divided by 65.7%).
The coracoid agrees with the characters listed by Brodkorb
(1961) for Neophrontops. Depth of sternal facet is 7.4 mm. (6.3 for N.
dakotensis); width of shaft at upper end of corracobrachial scar is
10.2 mm. (8.3 for N. dakotensis).
The ulna differs from that of Neophron percnopterus in having (1)
external condyle larger, and less rounded; (2) in internal view,
depression at proximal end of external condyle deeper; (3) carpal;
tuberosity less pronounced and blunter. Distal depth of ulna, 12.1 mm.
The radius has the bicipital tubercle larger, more elongated, and
set farther distally on shaft than in N. percnopterus. Distal width of
radius, 11.7 mm.
The femur of the Inglis Neophrontops slaughter differs from that
of the living Necrosyrtes monachus and Neophron percnopterus in having
(1) much greater length; (2) trochanteric ridge long, as in Neophron;
(3) posterior intermuscular line running nearly entire length of shaft
and set closer to internal edge of shaft; (4) internal condyle thrust
farther posteriorly; (5) ridge proximal to popliteal area absent; (6)
politeal depression deeper distally; (7) pit above fibular condyle
deeply excavated; (8) proximal edge of fibular condyle with expanded
The Inglis femur differs from that of Palaeoborus umbrosus in
having (1) head rounder and set farther out from shaft; (2) popliteal
depression extending distally; (3) fibular condyle extending farther
The femur differs from that of Neogyps errans in having (1)
trochanteric ridge longer; (2) anterior intermuscular line more medial
in position; (3) fibular condyle extending farther externally.
The Inglis femur resembles that of Neophrontops americanus, but
differs in having greater length and condyles set farther apart (Table
The tibiotarsus is distinguished from other accipitrid
subfamilies and agrees with Gypaetinae in the shape of the inner
cnemial crest (Howard, 1932). Of the New World Gypaetinae, the
proximal end of the tibiotarsus is known only for Palaeoborus umbrosus
(Cope), Neogyps errans L. Miller, and Neophrontops americanus L.
The proximal piece of the tibiotarsus is distinguished at once
from that of the Lower Pliocene Palaeoborus umbrosus Cope by its
large, squared inner cnemial crest. Also, the Inglis tibiotarsus in
proximal view has a straight ridge over the cnemial crest as opposed
to a curved ridge for P. umbrosus as figured by Cope (1877).
Table 24. Measurements (mm.) of the tibiotarsus of Neophron
percnopterus (2 ), Necrosyrtes monachus (sex?), Arikarornis
macdonaldi(Howard, 1966), Neophrontops americanus (Feduccia,
Palaeoboru; umbrosus (Cope, 1877), and Neogyps errans (Howard, 1932).
Neophron Necros. Arikar. Neophrontops
percnop. monachus macdon americanus
Proximal R 13.3-14.3 15.4 13.3-15.2
Width M 13.80 14.42
N 2 1 5
Proximal R 18.3-18.5 20.0 17.0-18.6
Depth M 18.40 17.90
N 2 1 5
Distal R 14.0-14.4 15.1 13.2 13.2-15.0
Width M 14.20 14.30
N 2 1 1 6
Depth of R 9.7-10.5 12.3 9.3 10.2-11.3
Internal M 10.10 11.00
Condyle N 2 1 1 6
Depth of R 9.6-10.8 11.7 9.2
External M 10.20 -
Condyle N 2 1 1
Table 24. Extended.
The Inglis tibiotarsus differs from that of Neogyps errans in its
smaller size and its much more prominent inner cnemial crest.
The proximal end of the Inglis tibiotarsus closely resembles that
of Neophrontops americanus as figured by Howard (1932), but differs in
having (1) in internal view, inner cnemial crest with squared-off
corners both proximally and distally (in N. americanus corners are
pointed proximally and rounded distally); (2) in anterior view,
outermost edge of outer cnemial crest projecting distally; (3) shaft
Measurements of the tibiotarsus are summarized in Table 24.
The distal piece of tibiotarsus agrees with the following
characters listed by Howard (1966) for the Gypaetinae: (1) short,
broad supratendinal bridge, less vertical in position than in Aquila
or Buteo; (2) tendinal groove centrally placed on shaft above bridge;
(3) attachment of oblique ligament on slope of internal face of
groove; (4) anterior face of shaft external to groove well rounded.
The tibiotarsus is distinguished from the Lower Miocene
Arikarornis macdonaldi (Howard, 1966) in the unequal depth of the
internal and external condyles, the excentral position of the internal
ligamental prominence, and the more media posterior border of the
Palaeoborus rosatus and P. howardae are not known from the
tibiotarsus, however other skeletal elements of these species are
about the same size as those of Neogyps errans, which is larger than
the Inglis bird (Table 24).
The tibiotarsus differs from that of the late Pliocene Palaeoborus
umbrosus as figured by Cope (1877, p1.68) in having the tendinal groove
narrower, the proximal edge of the supratendinal bridge more vertical
in position, and the internal condyle projecting farther laterally.
The tibiotarsus differs from that of the Upper Pleistocene
Neogyps errans as figured by Howard (1932) in having (1) tendinal groove
narrower; (2) supratendinal bridge forming a 450 angle with the long
axis of the shaft (300 in Neogyps); (3) distal edge of supratendinal
bridge straighter; (4) opening below supratendinal bridge less rounded;
(5) internal condyle projecting farther laterally.
Five species of the extinct genus Neophrontops have been described,
but only N. slaughter Feduccia of the Upper Pliocene and N. americanus
L. Miller of the Upper Pleistocene are known from the distal end of the
tibiotarsus (Table 24).
The Inglis tibiotarsus agrees with Neophrontops slaughter in the
following characters listed by Feduccia (1974) which distinguish it from
N. americanus: (1) tendinal groove broader; (2) internal ligamental
prominence more pronounced; (3) shaft more robust; (4) external condyle
notched at junction with shaft.
The tarsometatarsus differs from Recent Neophron percnopterus gin--
ginianus, the Egyptian Vulture, by having (1) larger size; (2)
prominence between cotylae higher; (3) inner foramen much larger
(enlarged proximal foramina may indicate an immature individual); (4)
tubercle for tibialis anticus placed farther distally; (5) anterior
metatarsal groove deeper; (6) internal edge of shaft rounded at level
of anterior metatarsal groove; (7) in posterior view, outer proximal
foramen opens a bit farther distally than inner foramen foraminaa open
at same level in Neophron); (8) posterior metatarsal groove deeper,
wider; (9) calcaneal ridges of hypotarsus much closer together.
Measurements of the proximal piece of a tarsometatarsus are as
follows (those of two female Neophron percnopterus in parentheses):
proximal width, 17.5 (13.9-14.2); length to middle of tubercle for
tibialis anticus, 15.5 (12.6 for both); width at level of tubercle for
tibialis anticus, 5.8 (4.7-5.1).
Distal half of right tarsometatarsus differs from N. percnopterus
in having (1) internal ridge along posterior metatarsal groove
terminating before reaching metatarsal facet; (2) metatarsal facet
rounded proximally, with a narrow ridge projecting distally; (3)
distal foramen large and oval. The trochleae for digits II and IV
are damaged in this fossil. Table 25 gives measurements for the
distal end of the tarsometatarsus of several species of the family
Remarks: Neophrontops slaughter was previously known only from
the late Pliocene Hagerman local fauna in Idaho, which is dated at 3.5
million years B.P. and is older than the Inglis site.
Table 26 summarizes the New World fossil Gypaetinae.
Table 25. Measurements of the tarsometatarsus of Neophron percnopterus
(2 F), Necrosyrtes monachus (sex?), Neophrontops americanus (Howard,
1963), Neophrontops vallecitoensis (Howard, 1963), Inglis,
Neophrontops slaughter, and Neogyps errans (Howard, 1966).
Neophron Necros. Neophr. Neophr. Inglis Neogyps
percnop. monach. americ. valley. Neophr. errans
Distal R 14.9-16.8 18.1 17.5 19.6 17.4 21.4-24.1
width M 15.80 22.55
N 3 1 1 1 1 4
Depth of R 7.7-8.7 7.8 8.3 10.1 11.1-12.6
internal M 8.13 12.03
trochlea N 3 1 1 1 4
Depth of R 6.6-7.4 8.2 7.6 8.0 9.0-9.6
middle M 6.93 9.33
trochlea N 3 1 1 1 4
Depth of R 7.5-8.5 8.2 8.8 9.8 10.5-11.5
external M 8.05 11.13
trochlea N 3 1 1 1 4
Table 26. The fossil Gypaetinae of the New World (revised from Feduccia, 1974).
Arikarornis macdonaldi Howard 1966
Palaeoborus rosatus A. H. Miller
Palaeoborus howardae Wetmore 1936
Palaeoborus umbrosus (Cope 1874)
Neophrontops vetustus Wetmore 1943
Neophrontops dakotensis Compton 1935
Neophrontops slaughter Feduccia 1974
Neophrontops vallecitoensis Howard 1963
Neophrontops americanus L. Miller 1916
Neogyps errans L. Miller 1916
Lower & Middle
Skeletal elements known
Tibiotarsus, distal end
Ulna, distal end
Broken pieces of most
Humerus, distal end
Coracoid and humerus,
Tibiotarsus, distal end
All principal elements
All principal elements
Subfamily Accipitrinae (Vieillot)
Genus Accipiter Brisson
Accipiter cooperii (Bonaparte)
Material: One complete right ulna, distal ends of 1 right and
1 left ulnae, proximal ends of 1 right and 1 left tibiotarsi, distal
end of 1 right tibiotarsus. The 6 specimens represent a minimum
number of 2 individuals.
Characters: Accipiter cooperii is distinguished from A. gentilis
and A. striatus on the basis of size. Measurements of the fossils are
compared to those of Recent A. cooperii in Table 27.
Remarks: Accipiter cooperii has been recorded at the Reddick,
Arredondo, and Haile sites in Florida (Brodkorb, 1964a).
Family Falconidae Vigors
Subfamily Falconinae (Vigors)
Genus Falco Linnaeus
Falco sparverius Linnaeus
Material: Fragment of 1 premaxillary, proximal end of 1 left
coracoid, distal half of 1 right coracoid, proximal ends of 2 left
scapulae, 1 left humerus (lacking head), distal ends of 2 left humeri,
proximal half of 1 right ulna, posterior fragment of synsacral
vertebrae, 1 right femur, distal end of 1 left femur, proximal end of
1 left tibiotarsus, proximal end of 1 right tarsometatarsus. The 15
specimens represent a minimum number of 2 individuals.
Referred material: Shaft of 1 right ulna.
Table 27. Measurements of the ulna and tibiotarsus of Recent
Accipiter cooperii (39,3A) and Inglis A. cooperii.
cooperii A. cooperii
Distal R 6.2-7.2 6.2-7.1
Depth M 6.75 6.60
N 6 3
Width of R 8.0-9.6 8.8
Head M 8.78
N 6 1
Characters: These elements of Falco sparverius are distinguished
from those of Accipiter striatus as follows: (1) premaxillary having
ventral notch; (2) coracoid lacking dorsal lip bordering sternal lip;
(3) scapula having coracoidal articulation next to glenoid facet en-
larged; (4) humerus having brachial depression wider, groove between
ectepicondylar prominence and external condyle semi-circular, distal
end narrower, and attachment of anterior articular ligament larger;
(5) ulna having internal cotyla smaller and prominence for anterior
articular ligament set more distally on shaft; (6) femur having an-
terior foramen set closer to proximal end and fibular condyle less
prominent; (7) tibiotarsus having inner cnemial crest shorter, supra-
tendinal bridge with 3 openings (Brodkorb, 1960b), and internal
ligamental prominence low; (8) tarsometatarsus having shaft short and
broad, tubercle for tibialis anticus set towars internal side of
bone (centered on anterior side in Accipiter), hypotarsal ridge
extending far down shaft between proximal foramena (ridge absent in
Accipiter), and external trochlea much shorter than other two.
Falco sparverius is distinguished from other Recent species of
the genus Falco on the basis of size.
Remarks: Falco swarthi L. Miller (1927) from the Pleistocene of
McKittrick, California was a large species, exceeding F. peregrinus
and F. mexicanus in the size of the tarsometatarsus. From the
Pleistocene of Oregon, Howard (1946) described Falco oregonus based on
a carpometacarpus the same size as those of F. peregrinus and F.
mexicanus. Brodkorb's (1959a) Pleistocene Falco readei from
Arredondo, Florida has been restudied by Campbell (1979), who places
it in the genus Milvago.
Falco columbarius Linnaeus
Material: Anterior fragment of 1 mandible, distal end of 1 right
femur. The 2 specimens represent a minimum number of 1 individual.
Characters: The mandible of Falco columbarius differs from that
of F. sparverius in having a larger size.
The femur in having (1) in posterior view, prominence proximal to
internal condyle projecting farther internally; (2) prominence
proximal to popliteal region central on posterior side of shaft
(shifted externally in F. sparverius); and (3) in external view,
depression on fibular condyle crescent-shaped (roundish in F. spar-
verius). The distal width of the fossil femur is 6.8 mm.
Remarks: Campbell's (1979) Falco sp. is between F. peregrinus
and F. femoralis, and is therefore, much larger than the Inglis F.
Order Galliformes (Temminck)
Family Phasianidae Vigors
Subfamily Odontophorinae Gould
Genus Colinus Goldfuss
Colinus *suilium Brodkorb
Material: Nine skull parts, 16 fused thoracic vertebrae, 49
synsacra, 58 sternums, 16 furcula, 75 right and 97 left coracoids, 46
right and 48 left scapulae, 124 right and 98 left humeri, 33 right and
22 left ulnae, 19 radii, 5 ulnares, 29 right and 15 left carpometa-
carpi, 5 digital phalanges, 17 pelvic fragments, 70 right and 61 left
femora, 59 right and 56 left tibiotarsi, 1 fibula, 21 right and 36 left
tarsometatarsi, and 15 pedal phalanges. The 1100 specimens represent
a minimum of 115 individuals.
Remarks: Ritchie (1976) includes a detailed osteological
comparison between Colinus suilium from Inglis IA and from Coleman IIA
(a late Illinoian glacial stage site from Sumpter County, Florida).
Ritchie (1976) also compared measurements of the Colinus material from
those sites with those from 6 late Pleistocene sites and demonstrated
that C. suilium gradually increased in size throughout the
Subfamily Meleagrinae G. R. Gray
Genus Meleagris Linnaeus
Meleagris *anza (Howard)
Material: Steadman (1975) lists the Florida State Museum's
catalogue numbers for the Inglis Meleagris anza material in his
Appendix. The 1230 specimens represent a minumum number of 50
Remarks: Steadman's (1975) dissertation on the Plio-Pleistocene
evolution of turkeys was based heavily on his study of the huge
collection of Inglis Meleagris. Steadman simplified meleagrid
taxonomy by synonymizing (1) Proagriocharis Martin and Tate,
Agriocharis Chapman, and Parapavo L. Miller with Meleagris Linnaeus;
and (2) M. alta Marsh and M. tridens Wetmore with M. gallopavo. His
proposed phylogeny shows an increase in size from Late Pliocene to
Recent in Meleagris: M. kimballensis (Martin and Tate) M. progenes
(Brodkorb) M. leopoldi A. Miller and Bowman M. anza (Howard) M.
Order Ralliformes (Reichenbach)
Suborder Ralli (Reichenbach)
Family Rallidae Vigors
Subfamily Rallinae (Vigors)
Genus Rallus Linnaeus
Rallus longirostris Boddaert
Material% Proximal end of 1 right coracoid, distal end of 1
right coracoid, proximal end of 1 right ulna, 1 complete left
carpometacarpus, anterior fragment of 1 sternum, distal end of 1 left
tibiotarsus, shaft of 1 left tibiotarsus, proximal end of 1 left
tarsometatarsus, distal ends of 1 right and 2 left tarsometatarsi.
The 11 specimens represent a minimum number of 3 individuals.
Characters: Ligon (1965) lists characters useful in separating
Rallus, Gallinula, and Fulica. Rallus longirostris is distinguished
from R. limicola and R. elegans on the basis of its intermediate
Measurements of the carpometacarpus and tarsometatarsus of R.
longirostris are compared with those of some other species of rails
in Table 28.
Remarks: Rallus prenticei Wetmore (1944) of the Upper Pliocene
of Kansas is slightly larger than the Virginia Rail, R. prenticei, but
still smaller than R. longirostris.
In his study of the Pliocene rails of North America, Feduccia
(1968) places Porzana lacustris Brodkorb (1958) in the genus Rallus,
and compares it with the Recent R. sanguinolentus of South America.
Rallus lacustris (Brodkorb) is a little smaller than R. longirostris.
Table 28. Measurements of the carpometacarpus and tarsometatarsus of
Rallus limicola (5d?,4), Inglis R. longirostris, R. longirostris
waynei (ld',69), Inglis R. elegans (tarsus only), R. elegans elegans
(2d2,2), Fulica americana (2 d,29-), Gallinula chloropus cachinnana
(5 e,49), and Porphyrula martinica (3 c,3 -).
Rallus Inglis Rallus Inglis
limicola Rallus long. Rallus
long. waynei elegans
Total R 32.1-37.6 46.2-49.1 50.3-52.4
Length M 34.95 47.63 51.35
N 8 7 2
Proximal R 4.1-4.7 6.0 6.0-6.6 6.2-6.7
Width M 4.36 6.34 6.45
N 8 1 7 2
Midshaft R 2.0-2.4 2.7-3.2 3.3-3.5
Width M 2.16 2.91 3.40
N 8 7 2
Midshaft R 1.9-2.2 2.6-3.0 3.1-3.5
Depth M 2.08 2.77 3.30
N 8 7 2
Width R 4.4-4.9 6.2-6.6 6.3-7.1 7.0
Through M 4.60 6.43 6.71 7.00
Trochleae N 8 3 7 2
Total R 18.4-20.8 27.4 26.5-28.4
Length M 19.41 27.41
N 9 1 7
Proximal R 3.9-4.3 5.6 5.7-6.1
Width M 4.07 5.92
N 9 1 7
Distal R 2.5-2.9 4.0 3.8-4.4
Width M 2.69 4.02
N 9 1 7
Table 28. Extended.
Rallus Fulica Gallinula Porphyrula
elegans americana chloropus martinica
elegans americana cachinnans
53.1-63.8 50.5-61.7 52.9-62.6 58.8-64.3
59.27 54.07 55.82 60.78
4 8 9 5
6.5-7.6 7.8-8.9 6.9-8.2 6.8-7.5
7.17 8.27 7.56 7.13
4 8 9 6
3.1-3.6 3.3-4.4 3.4-4.6 4.1-4.9
3.37 3.66 3.75 4.54
4 8 9 5
3.0-3.7 2.9-4.2 3.1-3.7 3.0-3.7
3.37 3.48 3.44 3.44
4 8 9 5
6.7-8.2 7.6-9.1 7.3-8.7 6.9-7.5
7.50 8.40 7.95 7.25
4 8 9 6
30.4-35.4 37.6-41.7 31.0-36.0 29.1-31.3
32.72 38.82 33.72 30.48
4 8 9 6
6.0-7.5 7.0-7.6 6.8-7.5 6.3-6.6
6.87 7.27 7.14 6.50
4 8 9 6
4.1-5.0 4.7-5.3 4.5-5.3 4.2-4.7
4.42 5.02 4.83 4.55
4 8 9 6
Rallus elegans Audubon
Material: One complete right and 1 complete left tarsometatarsi.
The 2 specimens represent a minimum number of 2 individuals (due to a
difference in size).
Characters: Rallus elegans is distinguished from R. limicola, R.
longirostris, and the extinct species of Rallus on the basis of its
Measurements are given in Table 28.
Remarks: All three Recent species of North American Rallus have
been recorded previously from Pleistocene sites in Florida (Brodkorb,
Genus Porzana Vieillot
Porzana carolina (Linnaeus)
Material: One complete left carpometacarpus, distal ends of 2
right tibiotarsi, shaft of 1 right tibiotarsus. The 4 specimens
represent a minimum number of 3 individuals.
Characters: The genus Porzana is distinguished from Rallus based
on characters listed by Ligon (1965).
Measurements are given in Table 29.
Remarks: The Upper Pliocene Porzana insignis (Feduccia 1968) and
P. avita (Feduccia 1968) are smaller than P. carolina.
The Lower Pleistocene Porzana lacustris Brodkorb (1958) is trans-
ferred to the genus Rallus (Feduccia 1968).
Porzana auffenbergi Brodkorb (1954) is larger than P. carolina,
and P. guti (Brodkorb, 1952) is smaller. Both are from the Middle
Pleistocene of Florida.
Table 29. Measurements of the carpometacarpus and tibiotarsus of
Rallus limicola (5 males, 4 females), Porzana carolina (7 males,
4 females, 1 ?), and Inglis P. carolina.
Rallus Porzana Inglis
limicola carolina Porzana
Suborder Grues Bonapartes
Family Gruidae Vigors
Subfamily Gruinae (Vigors)
Genus Grus Pallas
Grus americana (Linnaeus)
Material: Proximal end of 1 right ulna, olecranon process
Characters: The ulna of Grus differs from that of Balearica in
having: (1) proximal radial depression deeper; (2) ridge internal to
impression of brachialis anticus more prominent; (3) prominence for
anterior articular ligament extending farther distally; and (4)
tricipital attachment extending distally past edge of external cotyla.
The ulna of Grus americana differs from that of Recent G. cana-
densis in having: (1) condyles larger; (2) impression of brachialis
anticus deeper; and (3) shaft heavier.
The proximal end of the ulna of the Inglis Grus americana
differs from that of Grus grus in having: (1) Impression of
brachialis anticus deeper with prominence for anterior articular
ligament overhanging: (2) impression for brachialis anticus narrower;
(3) ridge from bicipital attachment extending just slightly
posteriorly; (4) in internal view, scar for anterior articular
ligament wider proximally; (5) proximal radial depression deeper;
and (6) diameter of internal cotyla greater.
Measurements of the ulna of the Inglis Grus americana are
compared with those of several Recent species of cranes in Table 30.
Table 30. Measurements of the ulna of Anthropoides virgo (1 male, 1
female), Balearica pavonina (1 male, 2 females, 1 ?), Grus grus grus
(1 ?), Grus g. lilfordi (2 females), Grus canadensis rowani (2 males,
I female), Grus c. tabida (3 males, 4 females), Grus c. pratensis
(4 males, 2 females), Grus americana (1 ?), and Inglis Grus americana.
A. virgo B. pavonina Grus g. grus
G. g. lilfordi
G. c. row. G. c. tab. G. c. pra. G. amer.
Remarks: Probalearica crataegensis Brodkorb of Florida's Lower
Miocene resembles the living Balearica pavonina and was described from
a distal end of a tibiotarsus. This bird is much smaller than the
living Sandhill Crane.
Pliogrus germanicus Lambrecht and Pliogrus pentelici (Gaudry) are
recorded from the Lower Pliocene of Europe. Cracraft (1973) notes a
close similarity between Pliogrus and Grus, and suggests that the genus
Pliogrus may be poorly defined. The ulna of P. pentelici was mentioned
by Brodkorb (1967) but was not examined by Cracraft (1973).
Baeopteryx latipes Wetmore from the Lower Pleistocene of Bermuda
is known from several skeletal elements. The ulna of B. latipes is
smaller and apparently weaker than that of Grus canadensis.
Grus conferta A.H. Miller and Sibley (1942) is described from a
distal piece of tarsometatarsus from California's Upper Lower Pliocene,
and was in size similar to G. americana.
Grus nannodes Wetmore and Martin (1930) of the Middle Pliocene of
Kansas is based on a carpometacarpus lacking the proximal end. This
bird was small, being less than two-thirds the size of G. canadensis.
Two neospecies of Grus, G. canadensis (Linnaeus), the Sandhill
Crane, and G. americana, the Whooping Crane, are known from the
Pleistocene of North America, including several sites in Florida
Suborder Cariamae Furbringer
Family *Phorusrhacidae (Ameghino)
Subfamily *Phorusrhacinae (Ameghino)
Genus *Titanis Brodkorb
Titanis walleri Brodkorb
Material: One complete carpometacarpus, 2 complete cervical
vertebrae (C2 and C3), 1 vertebral fragment, distal shaft of 1 right
tibiotarsus, 1 metatarsal I fragment, 1 left phalanx I of digit III, 1
left phalanx II of digit III, 1 left phalanx III of digit III, 1 right
and 1 left phalanges I of digit IV. The 11 specimens represent a
minimum number of 2 individuals.
Characters: The second cervical vertebra (axis) differs from that
of Andalgalornis ferox Patterson and Kraglievich (1960, as drawn in
dorsal view on p. 42) in having: (1) prezygapophyses greatly reduced
to small prominences; (2) anterior end of neutral spine narrower and
posterior end rounder; (3) anapophyses enlarged and rounded dorsally;
and (4) anapophyses thinner and projecting farther laterally.
The third cervical vertebra differs in having: (1) edge of
centrum more deeply curved; (2) prezygapophyses rounded; (3) neural
spine set farther back posteriorly; and (4) anapophyses larger and
projecting farther laterally.
The following are measurements of C2 with those of C3 in
parentheses: anterior width of centrum, 44.2; anterior depth of
centrum, 31.2; diameter of neural canal, 8.2 (8.0); width through
diapophyses, 126.8 (138.7); width through hypapophysis, 33.0 (32.3);
length of centrum, 69.3 (65.1); height from hypapophysis through neural
spine, 105.0 (109.2). All measurements above are in millimeters.