• TABLE OF CONTENTS
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 Title Page
 Acknowledgement
 Table of Contents
 List of Tables
 List of Figures
 Abstract
 Introduction
 List of species
 Paleoecology
 Comparison with other avifauna...
 Age of deposit
 Bibliography
 Biographical sketch














Group Title: early pleistocene avifauna from Inglis, Florida
Title: An early pleistocene avifauna from Inglis, Florida
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Permanent Link: http://ufdc.ufl.edu/UF00099233/00001
 Material Information
Title: An early pleistocene avifauna from Inglis, Florida
Physical Description: x, 162 leaves : ill. ; 28 cm.
Language: English
Creator: Carr, Gail Elaine Speaker, 1952-
Copyright Date: 1981
 Subjects
Subject: Birds, Fossil   ( lcsh )
Geology -- Florida -- Inglis   ( lcsh )
Geology, Stratigraphic -- Pleistocene   ( lcsh )
Zoology thesis Ph. D
Dissertations, Academic -- Zoology -- UF
Genre: bibliography   ( marcgt )
non-fiction   ( marcgt )
 Notes
Thesis: Thesis (Ph. D.)--University of Florida, 1981.
Bibliography: Bibliography: leaves 150-161.
General Note: Typescript.
General Note: Vita.
Statement of Responsibility: by Gail Elaine Speaker Carr.
 Record Information
Bibliographic ID: UF00099233
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: alephbibnum - 000294675
oclc - 07784437
notis - ABS1010

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Table of Contents
    Title Page
        Page i
    Acknowledgement
        Page ii
    Table of Contents
        Page iii
    List of Tables
        Page iv
        Page v
        Page vi
        Page vii
        Page viii
    List of Figures
        Page ix
    Abstract
        Page x
    Introduction
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
    List of species
        Page 6
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    Paleoecology
        Page 129
        Page 130
    Comparison with other avifaunas
        Page 131
        Page 132
        Page 133
        Page 134
    Age of deposit
        Page 135
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    Bibliography
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    Biographical sketch
        Page 162
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Full Text


















AN EARLY PLEISTOCENE AVIFAUNA
FROM INGLIS, FLORIDA







BY


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

















ACKNOWLEDGEMENTS


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




Page


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


Table Page

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.
cooperii.................................................... 79

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
walleri.................................................... 94

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


vii










41 Comparison of the humerus (distal end) of three species of
Campephilus................................................ 123

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,
1930)...................................................... 132

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


viii


















LIST OF FIGURES


Figure Page

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

By

Gall Elaine Speaker Carr

March, 1981



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;

Picidae, 4.

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.














INTRODUCTION


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

and 1974.

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 (*).


Elem Ind

Gaviiformes
Gavildae
Gavia artica 1 1

Podicipediformes
Podicipedidae
Podilymus podiceps 6 2
Podiceps dominicus 1 1
*Podiceps dixi 3 1

Ardeiformes
Ardeidae
Ardea sp. 1 1
Botaurus lentiginosus 1 1
Ixobrychus n. sp. 2 1
Plataleidae
Eudocimus albus 1 1

Anseriformes
Anatidae
*Anabernicula gracilenta 2 1
Aix sponsa 5 1
Anas platyrhynchos 2 1
Anas strepera 4 1
Anas discors 5 2
Aythya americana 1 1

Accipitriformes
Vulturidae
Coragyps atratus 17 2
Cathartes aura 2 1
cf.Gymnogyps californianus 1 1
Accipitridae
Buteo n. sp. 12 2
Aquila n. sp. 34 2
*Buteogallus fragilis I 1
*Neophrontops slaughter 7 1
Accipiter cooperil 6 2
Falconidae
Falco columbarius 2 1
Falco sparverius 14 2

Galliformes
Phasianidae
*Colinus suilium 1100 115
*Meleagris anza 1230 50


(Continued)









Table 1. Continued.


Elem Ind

Ralliformes
Rallidae
Rallus longirostris 11 3
Rallus elegans 2 2
Porzana carolina 4 3
Gruidae
Grus americana 1 1
Phorusrhacidae
*Titanis walleri 11 2

Charadriiformes
Scolopacidae
Capella galinago 10 4
Philohela minor 12 2

Columbiformes
Columbidae
Zenaidura macroura 40

Cuculiformes
Cuculidae
Coccyzus americanus 25

Strigiformes
Strigidae
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
Tytonidae
Tyto alba 1

Piciformes
Picidae
*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

Institution.

Avian osteological terms are those of Howard (1929). Osteo-

logical measurements were taken following those of Steadman (1975) and

Ritchie (1976).

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)

Arctic Loon

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)

characters.

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

6

















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. adamsil.


C. brodkorbi C. stellata C. arc. paclfica


Ulna

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.


Inglis
G. arctica


12.6

1

5.8

1

7.3

1


G. immer


G. adamsii


G. concinna


11.6

1

5.3

1

7.6

1

16.6

1


131.3-159.5

29

13.2-15.0

29

6.8-7.4

29

8.1-10.1

29

17.8-21.7

29


163.7

1

16.7

1

7.8

1

10.7

1

23.3

1


--------






9

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.






11











Table 3. Measurements of the humerus and ulna of Recent Podiceps
auritus (2e, 4?, 4?) and Inglis P. dixi.


P. auritus P. dixi

Humerus

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

Ulna

Distal R 3.8-4.5 4.8
Depth M 4.15
N 10 1








Podiceps dominicus (Linnaeus)

Least Grebe

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

Florida.



Genus Podilymbus Lesson

Podilymbus podiceps (Linnaeus)

Pied-billed Grebe

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?)


Podiceps
dominicus



45.5-48.8
47.33


Ulna


Total
Length


Proximal
Width


Midshaft
Width


Midshaft
Depth


Distal
Depth


Inglis
P. dominicus



45.5

1

4.2

1

2.0

1

2.5

1

2.9

I


4.0-4.1
4.07
3

1.7-2.1
1.85
4

2.2-2.6
2.40
4

2.7-3.0
2.83


_---_-_1_--_11_1-__ 1__1__1









(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.


Humerus

Total
Length


Proximal
Width


Least
Width of
Shaft


Distal
Width


Ulna


Distal
Depth


Podilymbus
podiceps



74.3-81.2
77.63
23

13.0-14.8
13.97
23


Inglis
P. podiceps


75.2

1


2.9-3.3
3.09
23

6.6-7.5
7.20
23



3.5-4.1
3.80
16



6.9-8.0
7.56
23


Tibiotarsus


Distal
Width










Order Ardeiformes (Wagler)

Suborder Plataleae Newton

Family Plataleidae Bonaparte

Subfamily Threskiornithinae (Richmond)

Genus Eudocimus Wagler

Eudocimus albus (Linnaeus)

White Ibis

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

humerus.



Suborder Ardeae Wagler

Family Ardeidae Vigors

Genus Ardea Linaeus

Ardea sp.

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
crest.


Eudocimus Inglis
albus E. albus

Humerus

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.

involucris).

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


Sternum


Length of right
Coracoidal sulcus


Width of dorsal
Lip of right
Coracoidal sulcus

Distance between
Dorsal lips of
Coracoidal sulcus


Inglis
Ardea sp.


Ardea
herodias



21.8-24.5
23.85
6


23.0

1


3.0-4.1
3.72
6

3.9-7.0
5.73
6










and I. sturmii tarcometatarsi are much larger that the Inglis tarso-

metatarsus.

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)

American Bittern

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.


Botaurus
lentiginosus


Carpometacarpus

Total
Length


Inglis
B. lent.


55.7-69.2
63.33
11

9.9-12.0
10.99


Proximal
Depth


Length of
Metacarpal I


Midshaft
Depth of
Metacarpal I

Midshaft
Width of
Metacarpal I


58.8.

1


6.9-8.6
7.99
11

2.4-3.5
2.99
11

3.4-4.3
3.90
11

6.5-8.2
7.39
11


Distal
Depth










Order Anseriformes (Wagler)

Suborder Anseres Wagler

Family Anatidae Vigors

Subfamily Anatinae (Vigors)

Genus Aix Boie

Aix sponsa (Linnaeus)

Wood Duck

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

1 individual.

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

Mallard

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


Aix
sponsa
males


Humerus

Total
Length


Proximal
Width


Midshaft
Width


Midshaft
Depth


Distal
Width


67.0-73.2
70.86
8

15.6-17.5
16.60
8

5.5-6.0
5.81
8

4.7-5.4
5.07
8

11.3-12.2
11.80
8


of Aix snsa (8 e, 5*) and


Aix
sponsa
females



69.0-73.9
70.76
5

15.7-16.6
15.92
5

5.5-5.9
5.68
5

4.6-5.2
4.96
5

11.5-12.7
11.86
5


Inglis
Aix
sponsa










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

Gadwall

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)

Blue-winged Teal

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)

Redhead

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.


Anas
discors


Humerus

Total
Length


Proximal
Width


Distal
Width


Ulna

Total
Length


Proximal
Width


Distal
Depth


61.5-64.1
63.34
5

13.2-13.9
13.55
6

9.3-10.4
9.83
6



51.0-54.1
52.96
6


Inglis
Anas
discors



61.8

1

13.8

1

9.8

1


6.0-6.8
6.45
6

5.3-5.8
5.51
6


Carpometacarpus

Total F
Length


Proximal R
Width P


36.7-39.5
37.88
6

8.6-9.2
8.93
6


37.9-38.3
38.10
2

9.2

1









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
americ.


Total
Length


Proximal R
Width M
N


Distal
Depth


63.4-68.0
65.62
9

7.2-7.6
7.41
9

5.9-6.5
6.11
9


66.3-71.3
68.75
8

7.2-7.8
7.48
8

5.8-6.3
6.06
8


69.1-72.9
70.68
5

7.5-7.8
7.64
5

6.0-6.1
6.04
5


69.5-79.5
76.46
11

7.6-8.7
8.26
11

6.1-7.1
6.61
11


Aythya Aythya
ameri. valisi.


76.2-83.9
79.17
13

8.0-8.7
8.33
13

6.4-7.5
6.90
13


79.0-84.1
81.90
8

8.3-9.0
8.67
8

6.8-7.5
7.11
8


Ulna






28



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

A. sepulta).










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

humerus.

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

Anabernicula.










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

Tarsometatarsus

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)

California Condor

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





32



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).


Sarcoramphus Gymnogyps
Inglis


R 18.8-19.8

M 19.3

N 2


Gymnogyps
Recent


Teratornis Vultur


26.2 26.2-29.4 26.6-31.2 30.0-31.1


27.3


29.0

15


30.6

2









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.





34



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
(Fisher) (Miller)

Tarsometatarsus

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)

Black Vulture

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

individuals.

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
Coracoid
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
N 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
N 2

Midshaft R 6.0-6.3
Depth M 6.15
N 2

Humerus
Midshaft R 10.5
Width M -
N 1

Midshaft R 8.8
Depth M -
N 1

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

58.5-65.3
62.0
38


C. atratus atratus
Florida California


57.9-64.0
61.71
9

63.1-68.7
66.51
9

24.4-26.8
25.66
9

7.0-7.9
7.59
9

5.4-6.4
6.09
9


10.2-11.4
10.87
9


56.6-62.6
59.87
6


C. a. brasiliensis
Panama

56.6-59.7
58.80
4


65.0-65.5
65.25
2

23.1-25.1
24.45
4

7.0-8.3
7.65
2

5.7-6.0
5.85
2


9.9-10.4
10.15
2


8.9-9.5
9.21
9


24.9-28.1
26.4
15


24.4-25.9
25.07
9


8.9-8.9
8.9
2


22.5-25.7
24.2
6


22.5-24.4
23.58
4











Table 15. Continued.


Coragyps atratus
Inglis


C. occidentalis
Reddick


*15.8-*16.1
15.95
2

12.3-13.0
12.65


Ulna
Proximal
Width


Distal
Depth


Tiblotarsus
Depth of
External
Condyle

Depth of
Internal
Condyle

Distal
Width


12.6-13.0
12.73
3

13.2-14.0
13.73
3

14.3
-


C. o. occi.
RLB

16.0-18.2
17.0
58


13.4-15.9
14.4
65









Table 15. Extended.

C. o. mexicanus
San Josecito Cave

15.6-17.7
16.6
20
















12.9-14.1
13.5
17


C. atratus atratus
Florida California

15.8-16.1 14.2-17.0
17.86 15.8
9 6

12.2-13.8
13.07
9


12.5-13.5
13.04
9

13.4-14.6
13.99
9

12.4-13.3
12.76
9


12.4-13.2
12.7
6


C. a. brasiliensis
Panama

14.2-17.0
15.70
4

11.8-13.7
12.75
2


12.2-12.5
12.35
2

13.5-13.9
13.70
2

12.3-12.8
12.53
4










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

tibiotarsus measurements.

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

occidentalis" states:

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
elements.

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

subspecies.










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)

Turkey Vulture

Material: One left quadrate, proximal end of 1 left

tarsometatarsus. The 2 specimens represent a minimum number of 1

individual.

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).


C. aura
Inglis


Quadrate


Length
(horizontal)


Depth


Tarsometatarsus

Proximal
Width


Midshaft
Width


Midshaft
Depth


15.7

1

7.0

1

5.0

1


C. aura
Recent


16.0-18.0
17.14
9

11.0-12.2
11.51
9



13.2-15.5
14.29
9

7.3-8.2
7.86
9

4.3-5.2
4.90
9







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

IV.

The 14 specimens represent a minimum number of 2 individuals.

Locality and age: Inglis IA, Citrus County, Florida. Early

Pleistocene.

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.


B.
magnirostris
(2 males, 2
females, 2 ?)


Total R
Length M
N

Proximal R
Height M
N

Height R
of Meta- M
carpal I N

Distal R
Width M
N


37.5-42.2
39.78
6

9.2-10.3
9.68
6

5.5-5.9
5.70
6

6.6-7.5
7.10
6


B.
platypterus
(1 male, 3
females)


44.5-46.5
45.6
4

10.8-11.3
10.95
4

6.7-7.0
6.83
4

7.2-8.0
7.63
4


B.
nitidus
(1 male,
4 females)

43.2-49.6
47.40
5

10.7-12.8
11.94
5

6.5-8.0
7.46
5

7.2-9.1
8.20
5


B.
lineatus
(6 males,
11 females)

47.0-55.2
51.31
17

10.8-13.7
11.85
17

6.5-8.2
7.34
17

8.0-9.7
8.79
17









Table 17. Extended.


B.
jamaicensis
(6 males, 6
females, 2 ?)

61.2-72.7,
65.52
14

14.7-17.8
16.08
14

9.2-11.9
10.56
14

10.9-13.2
11.81


B. B.
n. sp. regalis
(1 male,
2 females)


B.
lagopus
(1 male,
1 females)

62.9-64.6
63.75
2

14.3-14.4
14.35
2

7.2-9.2
8.20
2

10.5-10.6
10.55
2


B.
swainsoni
(2 males,
1 female)

60.8-66.4
63.20
3

13.5-15.2
14.27
3

8.6-10.5
9.27
3

9.9-11.3
10.53
3


69.3

1

16.9-18.4
17.65
2

10.4-10.6
10.50
2

13.3

1


72.4-79.4
75.90
2

16.1-18.6
17.35
2

11.1-12.1
11.60
2

12.2-13.8
13.00
2










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

and 18).









The Inglis Buteo n. sp. is perhaps ancestral to B. jamaicensis,

as it shows more resemblance to that species in the characters listed

above.

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
Total R
Length M
N


Proximal
Width


Distal
Width


Ungual phalanx of digit I
Proximal R
Width M
N


Proximal
Depth


Ungual phalanx
Proximal
Width


Proximal
Depth


R
M
N

of digit II
R
M
N

R
M
N


Ungual phalanx of digit III
Proximal R
Width M
N


Proximal
Depth


Ungual phalanx
Proximal
Width


Proximal
Depth


of digit IV
R
M
N


B. jamaicensis
22.2-27.1
24.13
4

10.9-12.7
11.68
4


5.4-6.3
5.78
4


6.5-8.2
7.20
4

12.0-14.0
12.95
4


6.0-7.3
6.45
4

11.1-13.2
12.10
4


4.9-5.7
5.15
4

8.1-9.9
8.95
4


4.0-4.7
4.28
4

7.5-9.0
8.15
4


Buteo n. sp.
24.7

1

11.8

1

6.0

1


7.7-8.0
7.85
2

14.1-14.6
14.35
2









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.

chrysaetos).

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

A. chrysaetos).

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.

chrysaetos).

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.


Scapula

Proximal
Width


A. chrysaetos

23.7-27.1
25.85
4


Length of
Articular surface

Tibiotarsus

Length from head



Midshaft width



Midshaft depth


Metatarsal I

Length



Width of
Digital condyle

Phalanx I

Length



Proximal width


Ungual phalanx of digit I

Proximal width R
M
N

Proximal depth R


14.3-17.3
15.72
5


151.7-168.2
161.77
3

10.2-12.0
11.33
3

8.8-9.7
9.37
3


22.5-27.6
25.10
3

13.1-15.7
14.83
3


35.5-41.0
38.83
3

16.0-19.1
17.97
3


9.8-11.8
10.93
3

16.8-21.0
19.10
3


Aquila n. sp.

32.3-32.9
32.60
2

18.7-20.7
19.70
2


181.2

1

11.8

1

9.8

1


26.0

1

17.0

1


40.0

1


11.3

1

20.5

1









Table 20. Continued.


Phalanx II of digit II

Total R
Length M
N

Proximal R
Width M
N

Distal R
Width M
N

Phalanx I of digit III

Total R
Length M
N

Proximal R
Width M
N

Distal width R
Width M
N

Phalanx III of digit III

Total R
Length M
N

Proximal width R
Width M
N


Distal
Width


A. chrysaetos

29.2-34.5
32.30
3

10.5-12.1
11.46
3

8.3-10.2
9.46
3



24.1-27.8
26.30
3

9.6-12.0
11.07
3

7.2-8.2
7.73
3



24.3-28.0
26.73
3

7.7-8.2
8.03
3

6.4-7.5
7.10
3


Aguila n. sp.

33.3

I

10.8

1

9.1

1



32.0

1

14.3

1

9.2

I



28.2

1

7.9

1

7.1

1









Table 20. Continued.


Phalanx I of digit IV

Total R
Length M
N

Proximal R
Width M
NI

Distal R
Width M
N

Ungual phalanx of digit IV

Proximal R
Width M
N

Proximal R
Depth M
N


A. chrysaetos


12.9-15.2
14.30
3

7.5-9.3
8.36
3

5.7-7.1
6.40
3



6.6-7.3
6.60
3

10.0-12.2
11.26
3


Aquila n. sp.


18.3

1

9.8

1

8.9

1



6.7

1

12.9

1









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.
Inglis


A. c. bonifacti
France


A. chrysaetos
Rancho La Brea


Carpometacarpus


Total
Length


Proximal
Width


Height of R
Metacarpal I M
M

Midshaft R
Width of M
Metacarpal II N

Midshaft R
Depth of M
Metacarpal II N

Distal R
Width M


Femur

Total
Length


Proximal
Width


Head
Width


Midshaft
Width


120.0

1

30.2

1


102.10
2


26.46
3


92.5-112.9


114.0-135.8


11.1

1

23.2

1



139.8

1

30.1

1

12.0

1

14.1

1


123.54
5


26.30
6


12.30
4









Table 21. Extended.


Recent Aquila chrysaetos
North America
Brodkorb Howard


94.2-113.2
105.00
3


96.6-105.5
102.20
3


24.5
26.43
3


15.0-17.5
16.57
3

6.0
6.90
3

8.0
8.77
3

18.6-21.5
20.43
3



118.7-133.8
128.73
3

26.7-33.0
30.47
3

10.1-12.0
11.33
3

12.0-14.0
13.10
3


118.6-129.2
123.46
3


France




14
14


25.77
14


127.55
20


27.98
20







12.82
20


~














Aqujiia n. sp. A. c. bonifacti
Inglis France


A. chrysaetos
Rancho La Brea


Table 21.





Femur

Midsliaft
Depth


Distal
Width


Depth of
Internal
Condyle

Depth of
External
Condyle


Continued.


20.92
4










Table 21. Extended.


Recent la chrysaetos
North America France
Brodkorb Howard

12.0-14.0
13.27
3

26.5-32.3
30.20 28.75
3 20

17.9-23.4
21.33
3

20.0-24.6
22.97
3









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

phalanges.

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






63




Table 22. Femur measurements (mm.) of Aquila borrasi (Arredondo, 1976)
and Aquila n. sp. length as preserved is 140.0 mm.


Total length

Proximal width

Head width

Maximum width of pneumatic foramen

Length of pneumatic foramen

Least width of shaft


A. borrasi

*155.

45.4

16.3

8.0

14.0

19.8


Aquila n. sp.

139.8

30.1

12.0

3.3

7.9

13.7









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

Campbell (1973).

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

Inglis fossil).

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
ext. articular
process

Depth of ramus
through suran-
gular process

FEMUR

Total length


Proximal
width


14.8-15.0
14.90
2

7.7-8.2
7.95
2



67.3-75.5
71.40
2

18.0-19.9
18.95
2


15.2


Inglis Neogyps
erransd


18.6 *22.3


10.1


80.3

1

20.8

1


71.7-83.4
79.80
5

14.8-15.8
15.48
4


93.5 103.4-113.4
- 107.58
1 4


21.0

1


18.5-20.8
19.20
4


Width of Head


7.0-7.6 7.8
7.30











Table 23. Continued.


Midshaft
width


Midshaft
depth


Distal width


Depth of
internal
condyle

Depth of
external
condyle


7.8-8.3
8.05
2

7.5-8.2
7.85
2

17.3-18.3
17.80
2

11.0-13.0
12.00
2

12.8-13.8
13.30
2


19.2

1

12.3

1

14.2


23.6

1

14.9


22.0

1

14.0

1

20.0


--~-









smaller than it. Neophrontops americanus is in close agreement with

N. slaughter in characters (2) and (4) above, based on Howard's

(1932) photograph.

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

prominence.

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

distally.

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

23).

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.

Miller.

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

Tibiotarsus
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.


Neophrontops
slaughter


Inglis N.
slaughter


16.1

1

19.3

1


11.4

1


Palaeoborus
umbrosus


20.0

1

17.0

1

16.0

1

13.0

1

12.0

1


Neogyps
errans


18.8-21.2
20.34
7

22.5-24.2
23.43
7

18.0-20.8
19.40
3

12.8-14.5
13.67
3

11.8-13.4
12.63
3









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

more robust.

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

internal condyle.

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

Gypaetinae.

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
slaugh.

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).


Species

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


Age

Lower Miocene


Lower Miocene

Middle Miocene


Lower Miocene


Middle Miocene

Lower & Middle
Pliocene

Upper Pliocene

Middle Pleistocene


Upper Pleistocene

Upper Pleistocene


Skeletal elements known

Tibiotarsus, distal end


Ulna, distal end

Tarsometatarsus, distal
end

Broken pieces of most
elements

Humerus, distal end

Coracoid and humerus,
distal ends

Tibiotarsus, distal end

Tarsometatarsus,distal
end

All principal elements

All principal elements









Subfamily Accipitrinae (Vieillot)

Genus Accipiter Brisson

Accipiter cooperii (Bonaparte)

Cooper's Hawk

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

Sparrow Hawk

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.





79



Table 27. Measurements of the ulna and tibiotarsus of Recent
Accipiter cooperii (39,3A) and Inglis A. cooperii.


Accipter Inglis
cooperii A. cooperii

Ulna

Distal R 6.2-7.2 6.2-7.1
Depth M 6.75 6.60
N 6 3

Tibiotarsus

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

Pigeon Hawk

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.

columbarius.



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

Pleistocene.



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

individuals.

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.

gallopavo Linnaeus.









Order Ralliformes (Reichenbach)

Suborder Ralli (Reichenbach)

Family Rallidae Vigors

Subfamily Rallinae (Vigors)

Genus Rallus Linnaeus

Rallus longirostris Boddaert

Clapper Rail

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

size.

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

Tarsometatarsus

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

Carpometacarpus

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

King Rail

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

superior size.

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,

1967).



Genus Porzana Vieillot

Porzana carolina (Linnaeus)

Sora

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
carolina


Carpometacarpus

Total
Length


Proximal
Width


Distal
Width


Tibiotarsus

Distal
Width


Depth of
Internal
Condyle

Depth of
External
Condyle


18.4-20.8
19.41
9

3.9-4.3
4.07
9

2.5-2.9
2.69
9


3.9-4.4
4.17
9

4.2-4.7
4.47
9

4.0-4.6
4.33
9


18.6-21.3
20.22
12

3.9-4.4
4.22
12

2.4-2.8
2.68
12



3.5-4.3
4.08
12

4.0-4.7
4.35
12

3.8-4.5
4.23
12


4.4-4.6
4.50
2

4.4-4.9
4.65
2

4.4-4.8
4.60
2










Suborder Grues Bonapartes

Family Gruidae Vigors

Subfamily Gruinae (Vigors)

Genus Grus Pallas

Grus americana (Linnaeus)

Whooping Crane

Material: Proximal end of 1 right ulna, olecranon process

missing.

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


Ulna

Proximal R
Width M
N


Width at
Bicipital
Scar

Depth at
Bicipital
Scar


Ulna

Proximal R
Width M
N


Width at
Bicipital
Scar

Depth at
Bicipital
Scar


G. c. row. G. c. tab. G. c. pra. G. amer.


19.3-21.6
20.57
3

17.5-18.3
17.90
3

12.3-13.5
13.10
3


19.4-21.0
20.26
7

15.4-18.1
17.33
7

12.2-13.7
13.13
7


19.0-21.3
20.05
6

16.3-17.8
17.20
6

12.1-13.5
12.65
6


21.0

1

17.3

1

14.0

1


17.7-18.4
18.05
2

15.1-15.9
15.50
2

12.5-13.2
12.85
2


17.7-20.7
18.68
4

15.3-18.3
16.25
4

11.0-14.3
12.65
4


24.1

1

21.0

1

15.0

1


20.5-20.8
20.65
2

17.9-18.2
18.05
2

12.3-13.0
12.65
2


Inglis
G. amer.

23.1

1

19.2

1

14.0

1










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

(Brodkorb, 1967).








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.




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