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The fossil birds of the late Miocene and early Pliocene of Florida

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Title:
The fossil birds of the late Miocene and early Pliocene of Florida
Creator:
Becker, Jonathan J., 1955-
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Language:
English
Physical Description:
ix, 245 leaves : ill. ; 28 cm.

Subjects

Subjects / Keywords:
Birds ( jstor )
Fauna ( jstor )
Femur ( jstor )
Fossils ( jstor )
Genera ( jstor )
Humerus ( jstor )
Mining ( jstor )
Museums ( jstor )
Species ( jstor )
Taxa ( jstor )
Birds, Fossil ( lcsh )
Dissertations, Academic -- Zoology -- UF
Paleontology -- Florida -- Pliocene ( lcsh )
Zoology thesis Ph. D
Manatee County ( local )
Genre:
bibliography ( marcgt )
theses ( marcgt )
non-fiction ( marcgt )

Notes

Thesis:
Thesis (Ph. D.)--University of Florida, 1985.
Bibliography:
Includes bibliographical references (leaves 229-244).
Additional Physical Form:
Also available online.
General Note:
Typescript.
General Note:
Vita.
Statement of Responsibility:
by Jonathan J. Becker.

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University of Florida
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THE FOSSIL BIRDS OF THE LATE MIOCENE AND EARLY PLIOCENE
OF FLORIDA





BY


JONATHAN J. BECKER


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY



UNIVERSITY OF FLORIDA


1985

































Copyright 1985

by

Jonathan J. Becker
















ACKNOWLEDGMENTS

I would first like to thank my advisor, Pierce Brodkorb, for his

support, guidance, and friendship during the course of this project. He

has provided much encouragement and council throughout my researches on

fossil birds, as have the members of my committee--Drs. Richard A.

Kiltie, S. David Webb, Elizabeth S. Wing, and Ronald G. Wolff. I thank

them for the many hours spent reading this manuscript and for their many

helpful comments pertaining to my research.

A number of friends and colleagues contributed to this study with

their comments, suggestions, and discussions. They include S. Emslie, R.

Hulbert, and A. Pratt. Gary S. Morgan has taken much time to discuss the

geology of Florida and systematics, evolution, and biochronology of

fossil and Recent mammals, in addition to many other aspects of

vertebrate paleontology. I especially thank him for his comments on

Chapter III. Storrs Olson has freely shared his considerable knowledge

and insights of avian systematics, evolution and anatomy. David Steadman

also provided many helpful comments. I also thank Cynthia West for her

support and aid in completing this dissertation.

I thank the many amateur fossil collectors who have generously

donated fossil birds from the included localites to the Florida State

Museum. They include Danny Bryant, George Hesslop, and Ron Love. Phil

Whisler, of Venice, Florida, originally discovered the SR-64 locality and

brought it to the attention of the staff of the Florida State Museum.

Rick Carter, of Lakeland, Florida, deserves special mention, for over the


iii








last three years, he has donated hundreds of specimens of fossil birds

from the Bone Valley Mining District to the Florida State Museum.

Without his generous contributions, the Bone Valley portion of this study

would have been impossible to complete. John Waldrop is also gratefully

acknowledged for providing information about the avian localities in the

Bone Valley.

I also thank the following individuals and institutions for making

fossil and/or Recent specimens available for study: A. Andors, G.

Barrowclough, C. Houlton, R. H. Tedford, F. Vuilleumier, American Museum

of Natural History; P. Brodkorb, Department of Zoology, University of

Florida; J. Chenval, C. Mourer, Universite Claude Bernard; J. Hardy, B.

J. MacFadden, G. S. Morgan, S. D. Webb, T. Webber, Florida State Museum;

R. Mengel, University of Kansas; R. Payne, University of Michigan; M.

Voorhies, University of Nebraska; H. James, S. L. Olson, D. Steadman,

United States National Museum. Helen James, Storrs L. Olson, and David

Steadman generously provided accommodations during a lengthy stay in

Washington, D.C.

Financial support received while at the University of Florida,

includes teaching and research assistantships from the Department of

Zoology, College of Liberal Arts and Sciences; teaching assistantships

from the Department of Physiological Sciences, College of Veterinary

Medicine; a curatorial assistantship from the Department of Vertebrate

Paleontology, Florida State Museum; and grants from the Frank M. Chapman

Memorial Fund, American Museum of Natural History, and from Sigma Xi

Grants-In-Aid of research. The Department of Zoology, College of Liberal

Arts and Sciences, and the Florida State Museum supplied all expendable







equipment. I gratefully acknowledge these departments and institutions

for their support.

Last, I thank my parents, Elwood W. and Nita E. Becker, for their

encouragement and support over the years. They have provided not only the

opportunity, but much of the impetus, that has allowed me to finish my

formal education.














TABLE OF CONTENTS


ACKNOWLEDGMENTS . . . . . . . . . . ...

ABSTRACT . . . . . . . . . . . . . .

CHAPTER

I. INTRODUCTION AND PREVIOUS WORK . . . . . . .

Introduction . . . . . . . . . ...
Limitations of Study . . . . . . . . ...
Previous Work . . . . . . . . . ....

II. METHODS . . . . ........ . . .

Measurements . . . . . ... .
Computer Software . .. . . . . . ....
Nomenclature . .. . . . . .
Systematics . ...... . . . .
Paleoecology . . . .. . .............
Biochronology and Faunal Dynamics . . . . . .
Specimens Examined . . . . . . . . . .
Abbreviations . . . . . . . . . . .

III. GEOLOGY . . . . . . . . . . . .

Biochronology . . . . . . . . . ...
Local Faunas . . . . . . . . .....
Eustatic Sea-level Changes . . . . . . . .

IV. SYSTEMATIC PALEONTOLOGY . . . . . . . . .


Order Podicipediformes .
Order Pelecaniformes .
Order Ciconiiformes .
Order Accipitriformes
Order Anseriformes . .
Order Galliformes . .
Order Ralliformes . .
Order Charadriiformes
Order Strigiformes . .
Order Passeriformes .


.......... . . 49
. . . . . . . . .. 65
. . . . . . . . 90
S. . . . . . . 116
. . . . . . . . 136
. . . . . . . . . 155
* . . . . . . . . 158
. . . . . . 175
S. . . . . . . . 191
. . . . . .. . .. 195


PAGE

iii

viii









V. PALEOECOLOGY . . .


Introduction . . .
Local Faunas . . .

VI. BIOCHRONOLOGY AND FAUNAL


Introduction . . .
Faunal Dynamics ...
Biochronology . . .

VII. SUMMARY . . ..


Systematics . . .
Paleoecology . . .
Biochronology . . .

LITERATURE CITED . . .


BIOGRAPHICAL SKETCH . . .


* * *

* * *
* ft ft *


DYNAMICS


* * *
* * *
* * *


* * *

* * *
* * *
* * *

* * *

* * ft ft


198


198
199

205

205
205
213


224

224
227
228


229

245















Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

THE FOSSIL BIRDS FROM THE LATE MIOCENE AND EARLY PLIOCENE
OF FLORIDA

By

Jonathan J. Becker

August 1985

Chairman: Pierce Brodkorb
Major Department: Zoology

This study examined the non-marine avifauna from ten late Miocene

and early Pliocene localities in Florida. These localities include the

Love Bone Bed, McGehee Farm, Mixson's Bone Bed, Bone Valley Mining

District, Withlacoochee River 4A, Manatee County Dam, SR-6h, Haile VB,

Haile VI, and Haile XIXA. Non-marine genera (number species, if more

than one) present include Rollandia, Tachybaptus, Podilymbus (2),

Podiceps, Pliodytes, Phalacrocorax (3), Anhinga (2), Ardea (2), Egretta

(2 or 3), Ardeola, Nycticorax, Mycteria, Ciconia (3), Eudocimus,

Plegadis, Threskiornithinae, genus indeterminate, Pliogyps, Pandion (2),

Haliaeetus, Buteo, Aquila, Accipitrid, genus indeterminate (3),

Dendrocygna, Branta, Anserinae, genus indeterminate (4), Tadorine, genus

indeterminate, Anas (2), Anatine, genus indeterminate (2), Aythya,

Oxyura, Meleagridinae, genus indeterminate, Meleagris, Grus (2),

Balearicinae, genus indeterminate, Aramornis, Rallus (3), Rallid,

undescribed genus, Phoenicopterus (2), Jacana, Limosa, "Calidris" (6+),


viii






?Actitis, ?Arenaria, ?Philomachus, Tytonid, undescribed genus, Bubo,

Passeriformes (2).

The largest avifaunas are from the Love Bone Bed local fauna (44

taxa present) and the Bone Valley local fauna (41 taxa present, 31 here

included). These two localities are the most diverse non-marine and

marine avifaunas, respectively, known in North America prior to the

Pleistocene.

An analysis of the faunal dynamics of the Neogene fossil birds from

North America shows the following results. (1) Localities which have

produced fossil birds are not uniformly distributed through time--74.4%

of the localities are from the last 41% of the Neogene. (2) By the

Barstovian, a majority of the living families which have a fossil record,

have appeared. (3) Generic diversity increases from 10 to 98 during the

Neogene. (4) The marine avifauna is essentially established at a

diversity of 20 to 25 genera by the Clarendonian. (5) The non-marine

avifauna increases continually throughout the Neogene. (6) Origination

rates for marine birds peak in the Clarendonian with 4.4 genera appearing

per million years. (7) Extinction rates for marine birds are

consistently low throughout the Neogene. (8) Origination rates for non-

marine birds show a 2- to 4-fold increase in alternate Land Mammal Ages.

(9) Turnover rates parallel the origination rates described above.














CHAPTER I
INTRODUCTION AND PREVIOUS WORK

Introduction

Florida has one of the richest records of fossil birds in the world.

This study examines the systematics of the non-marine fossil birds which

lived in Florida during the late Miocene and early Pliocene (9.0--4.5

million years before present) and the paleoecology of the localities

which produced them. Included is material from 10 local faunas--the

Love Bone Bed, McGehee Farm, Mixson Bone Bed, Bone Valley, Withlacoochee

River 4A, Manatee County Dam Site, SR-64, Haile VB, Haile VI, and Haile

XIXA. Only a few of the birds from McGehee Farm and the early

collections of birds from Bone Valley have been studied previously (Table

1.1). In addition, the biochronology and faunal dynamics of the entire

North American Neogene avifauna are investigated. Specifically, the

following questions are addressed:

Systematics

1. What species of fossil birds are present in Florida during the

late Miocene and early Pliocene?

2. What are their systematic and biogeographic relationships to

other fossil and Recent species?

Paleoecology

1. Can fossil birds be used to reconstruct the paleoenvironments of

the fossil localities examined in this study?











Biochronology and Faunal Dynamics

1. What is the temporal distribution of the fossil localities

producing birds in the Neogene of North America?

2. What is the temporal distribution of the North American Neogene

avifauna?

3. When, and at what rate, do the North American Neogene avian

families and genera appear and become extinct?

4. How do the marine and non-marine localities and avifaunas differ

in questions 1 to 3 above?

5. What avian species are biostratigraphically useful in the

Neogene of North America?

6. What degree of temporal resolution does avian biochronology

offer?


Limitations of Study.

There have been several limitations imposed on this study by the

current knowledge of avian systematics and paleontology, and to a lesser

degree by the lack of previous studies dealing with the paleoecology and

biochronology of birds. Birds are often considered ". . the best known

and most completely described class of animals . (Welty 1975:1h).

In reality, this statement applies only to the simple designation of

living forms and to the obvious aspects of their behavior and ecology,

but definitely not to their evolution, their superspecific relationships,

and to many aspects of internal morphology. Many living groups of birds

still lack modern systematic revisions based on internal morphology, or

in many, even a description of their internal morphology. Most modern

orders have never been shown to be monophyletic, although many are

doubtlessly so. Modern classifications of birds above the specific level












(e.g., American Ornithologists' Union, 1983) has changed very little in

the 90 years since Gadow's (1893) classification. Olson (1981a:193),

addressing this problem, states

Many ornithologists appear to believe that the higher-
level systematics of birds is a closed book, the sequence of
orders and families in their field guides being an immutable
constant that was determined long ago according to some
infallible principle. In reality, the present classification
of birds amounts to little more than superstition and bears
about as much relationship to a true phylogeny of the Class
Ayes as Greek mythology does to the theory of relativity. A
glance at the Gadow-Wetmore classification now in use shows
that there is still no concept in ornithology of what
constitutes a primitive bird.

Certainly correct phylogenies are impossible to develop without an

accurate knowledge of primitive character states, the distribution of

primitive and derived characters within the group being studied, and

meaningful outgroup comparisons.

Many fossil species are described from single, non-diagnostic

elements, making useful comparisons between, or among, species

impossible. But even diagnostic elements, when singly preserved, add

little to our understanding of the phylogeny and evolution of that taxon.

Other species are arbitrarily allied with the wrong family, the wrong

order, and even in some cases, with the wrong class of vertebrates (cited

in Brodkorb, 1978:211-228), many because of lack of proper comparisons.

Only recently have many of these errors been recognized, due in part to

an increased number of workers in the field, and from greater

availability and use of comparative skeletal material. Adequate skeletal

collections are still lacking for many common species (Zusi et al.,

1982).

It is outside the scope of this study to revise the many recent and

fossil genera and families which need such treatment. In such cases, I











try to note the systematic problems in each group, briefly review its

fossil record, and describe the fossil material from Florida. This

material should be reexamined as the state of systematics of these groups

improves and as additional Recent and fossil material becomes available.



Previous Work

The earliest published report of fossil birds from Florida is

Sellard's (1916) description of a supposed jabiru (Jabiru? weillsi

=Ciconia maltha) from Vero, closely followed by Shufeldt's (1917a, 1917b)

study of this local fauna. Wetmore (1931) reviewed the Pleistocene

avifauna of Florida and was also the first to report (19h3) on the

Tertiary birds from Thomas Farm and the Bone Valley Mining District.

Numerous studies have appeared since then, primarily by Brodkorb or his

students. A few other faunal and systematic studies have also included

avian material from Florida.

Table 1.1 lists many of the fossil localities in Florida which have

a notable record of fossil birds. Reference to Brodkorb's Catalogue of

Fossil Birds (1963-1978), where the avifaunas from many Florida

Pleistocene localities were first reported, is omitted to conserve space.










Table 1.1. Avian Fossil Localities of Florida.

Locality Reference (s)

MIOCENE TO EARLIEST PLIOCENE (Pre-Blancan)

Bone Valley, Polk Co. Becker, 1985a; Brodkorb, 1953b, 1953c,
1953d, 1953e, 1955a, 1970; Olson, 1981b;
Steadman, 1980; Wetmore, 1943; this study


Gainesville Creeks,
Alachua Co.

Haile VI, Alachua Co.

Haile XIXA, Alachua Co.

Love Bone Bed, Alachua Co.


Manatee Co.Dam Site
Manatee Co.

McGehee Farm, Alachua Co.


Mixson Bone Bed, Levy Co.

Seaboard Airline Railroad,
Leon Co.

SR-64, Manatee Co.

Thomas Farm, Gilchrist Co.



Withlacoochee River 4A,
Marion Co.

LATE PLIOCENE (Blancan)

Haile XVA, Alachua Co.

Santa Fe IB, Gilchrist Co.


Brodkorb, 1963b


Brodkorb, 1963a; this study

this study

Becker, 1985a, 1985b; Webb et al., 1981;
this study

Webb and Tessman, 1968; this study

Brodkorb, 1963a; Hirschfeld and Webb, 1968;
Olson 1976; this study

this study

Brodkorb, 1963b


this study

Brodkorb, 1954a, 1956a, 1963b; Cracraft,
1971; Olson and Farrand, 1974; Steadman,
1980; Wetmore, 1943, 1958

Becker, 1985a; this study




Campbell, 1976; Steadman, 1980

Brodkorb, 1963d










Table 1.1--continued.

Locality Reference (s)

EARLY PLEISTOCENE (Irvingtonian)

Coleman IIA, Sumter Co. Ritchie 1980; Steadman, 1980

Haile XVIA, Alachua Co. Steadman, 1980

Inglis IA, Citrus Co. Carr, 1981; Ritchie, 1980; Steadman, 1980

Santa Fe River IIA, Steadman, 1980
Gilchrist Co.

Williston, Levy Co. Holman, 1959, 1961; Steadman, 1980

LATE PLEISTOCENE (Rancholabrean)

Arredondo, Alachua Co. Brodkorb, 1959; Holman, 1961; Olson, 1974b,
1977b; Steadman, 1976, 1980; Storer, 1976b

Aucilla River IA, Steadman, 1980


Jefferson Co.

Bowman IA, Putnam Co.

Bradenton, Manatee Co.


Catalina Lake,
Pinellas Co.

Coleman III, Sumter Co.

Crystal Spring Run,
Pasco Co.

Davis Quarry, Citrus Co.

Econfina River, Taylor Co.

Eichelberger Cave,
Marion Co.

Florida Lime Company,
Marion Co.

Haile IA, Alachua Co.

Haile IIA, Alachua Co.

Haile VIIA, Alachua Co.


Steadman, 1980

Becker, 1984; Steadman, 1980; Wetmore,
1931

Storer, 1976b


Ritchie, 1980

Brodkorb, 1956b


Steadman, 1980

Steadman, 1980

Brodkorb, 1955b; Holman, 1961


Steadman, 1980


Brodkorb, 1953a, 1954b; Olson, 1974b, 1977b

Holman, 1961; Steadman, 1980

Steadman, 1980











Table 1.1--continued

Locality

Haile XIB, Alachua Co.

Hog Cave, Sarasota Co.

Hog Creek, Manatee Co.

Hornsby Springs,
Alachua Co.

Itchtucknee River,
Columbia Co.

Jenny Spring, Gilchrist Co.

Kendrick IA, Marion Co.

Lake Monroe, Volusia Co.

Mefford Cave I, Marion Co.

Melbourne, Brevard Co.

Monkey Jungle, Dade Co.

Oakhurst Quarry, Marion Co.

Orange Lake, Marion Co.

Reddick IB, Marion Co.



Rock Springs, Orange Co.


Sabertooth Cave, Citrus Co.

St. John's Lock, Putnam Co.

St. Mark's River,
Leon/Wakulla Co.

Santa Fe River IA,
Gilchrist Co.

Santa Fe River IVA,
Gilchrist Co.

Seminole Field,
Pinellas Co.


Reference (s)

Ligon, 1965; Olson, 1974b

Steadman, 1980; Wetmore, 1931

Wetmore, 1931

Storer, 1976b

Campbell, 1980; McCoy, 1963; Olson, 1974a,
1974b, 1977b; Storer, 1976b; Wetmore, 1931

Storer, 1976b

Steadman, 1980

Holman, 1961; Storer, 1976b

Steadman, 1980

Holman, 1961; Steadman, 1980; Wetmore, 1931

Ober, 1978

Holman, 1961; Steadman, 1980

Holman, 1961

Brodkorb, 1952, 1957, 1963e; Hamon, 1964;
Holman, 1961; Olson, 1974b, 1977b;
Steadman, 1976, 1980; Storer, 1976b

Storer, 1976b; Steadman, 1980; Woolfenden,
1959

Holman, 1961; Wetmore, 1931

Storer, 1976b

Steadman, 1980


Steadman, 1980


Steadman, 1980


Holman, 1961; Olson, 1974b; Steadman, 1980;
Wetmore, 1931











Table 1.1--continued

Locality

Steinhatchee River,
Taylor/Dixie Co.

Venice Rocks, Manatee Co.

Vero (Stratum 2),
Indian River Co.


Warren's Cave, Alachua Co.

Wekiva Run III, Levy Co.

West Palm Beach,
Palm Beach Co.

Withlacoochee River,
Citrus Co.

Zuber, Marion Co.

HOLOCENE

Cotton Midden, Volusia Co.

Castle Windy Midden,
Volusia Co.

Good's Shellpit,
Volusia Co.

Green Mound Midden,
Volusia Co.

Nichol's Hammock, Dade Co.

Silver Glenn Springs,
Lake Co.

Summer Haven Midden,
St. Johns Co.

Vero (Stratum 3),
Indian River Co.

Wacissa River,
Jefferson Co.


Reference (s)

Steadman, 1980


Wetmore, 1931

Holman, 1961; Sellards, 1916; Shufeldt,
1917; Steadman, 1980; Storer, 1976b; Weigel,
1962; Wetmore, 1931

Holman, 1961

Steadman, 1980

Becker, 1985c


Steadman, 1980


Holman, 1961



Hay, 1902; Neill et al., 1956

Weigel, 1958


Steadman, 1980


Hamon, 1959


Hirschfeld, 1968; Steadman, 1980

Neill et al., 1956; Steadman, 1980


Brodkorb, 1960


see Vero, Stratum 2, above


Steadman, 1980
















CHAPTER II
METHODS

Measurements

Measurements made in this study are listed below and are illustrated

in Figures 2.1 2.4. They have been selected from the literature

dealing with the osteology and identification of both Recent and fossil

species of birds. Previous authors have dealt with only one specific

group (Steadman, 1980; Howard, 1932b; Ono, 1980) or with Recent birds

commonly found in archeological sites (von den Driesch, 1979; Gilbert, et

al., 1981). The most applicable anatomical studies on fossil birds are

by Ballmann (1969a, 1969b). I have modified measurements from the above

studies to make them applicable to the range of morphologies encountered

in the fossil birds of this study. Measurements for the less diagnostic

elements and for the cranium were not included. Measurements presented

in the systematic section of this dissertation depend on the fossil

material available and the morphology of the species considered.

Anatomical terminology follows Baumel et al. (1979) and Howard

(1929, 1980). Many Latin terms have been anglicized for ease of

communication, but the original Latin is given parenthetically when the

term is first used (below). Terms for soft tissue anatomy come from

Feduccia (1975) and Van den Berge (1975).










Scapula

1. LENGTH.-Greatest length from the acromion to the caudal

extremity of the scapula (Extremitas caudalis scapulae).

2. W-NECK.--Least width neck of scapula (Collum scapulae).

3. W-PROX.--Proximal width from the ventral tip of the glenoid

facet (Facies articularis humeralis) to the dorsal margin of the

scapular head (Caput scapulae).

4. ACR-GLN.--Length from tip of acromion through ventral tip of

glenoid facet.

5. D-GLN.--Depth of glenoid facet.

Coracoid

1. HEAD-FAC.-Length from head (Processus acrocoracoideus) through

external end of sternal facet (Facies articularis sternalis).

2. HEAD-IDA.--Length from head through internal distal angle

(Angulus medialis).

3. HEAD-CS.--Length from head through scapular facet(Cotyla

scapularis).

4. D-HEAD.-Least depth of head.

5. W-SHAFT.--Width of midshaft.

6. D-SHAFT.--Depth of midshaft.

7. FAC-IDA.-Length from external end of sternal facet through

internal distal angle.

8. IDA-PL.--Length from internal distal angle to medial most edge

of sternocoracoidal process (Processus lateralis).

9. IDA-PP.--Length from internal distal angle to procoracoid

process (Processus procoracoideus).










10. L-GLEN.-Length of glenoid facet from the most cranial portion

of glenoid through the most caudal point of scapular facet.

11. IDA-FNS.--Length from internal distal angle through the most

sternal edge of coracoidal fenestra (Foramen nervous

supracoracoidei).

12. ANG-HEAD.--Angle formed between axis of the head, as seen in

proximal view, and the plane parallel to the dorsal surface

(Facies dorsalis).

Humerus

1. LENGTH.--Greatest length from the head of the humerus (Caput

humeri) through the midpoint of the lateral condyle (Condylus

ventralis).

2. W-SHAFT.-Transverse width of midshaft.

3. D-SHAFT.--Depth of midshaft.

4. W-PROX.--Transverse width of proximal end from the external

tuberosity (Tuberculum dorsale) to the most ventral face of the

bicipital crest (Crista bicipitalis).

5. D-PROX.--Depth of proximal end, from the bicipital surface

(Facies bicipitalis) to the internal tuberosity (Tuberculum

ventrale), measured at right angles to the long axis of the

shaft.

5a. D-HEAD.--Depth of head, measured parallel to the axis of the

head.

6. L-DELTOID.--Length of deltoid crest (Crista pectoralis),

measured from the external tuberosity to the most distal

extension of the deltoid crest.










7. W-DIST.--Transverse width of distal end from the entepicondylar

prominence (Epicondylus ventralis) to the ectepicondylar

prominence (Epicondylus dorsalis).

8. D-DIST.--Depth of distal end from cranial face of external

condyle (Condylus dorsalis) through ridge slightly media from

external tricipital groove (Sulcus scapulotricipitis), measured

at right angles to the long axis of the shaft.

9. D-ENTEP.--Depth of entepicondyle (Epicondylus ventralis) from

attachment of the pronator brevis (Tuberculum supracondylare

ventrale) through entepicondyle (Processus flexoris), measured at

right angles to the long axis of the shaft.

Ulna

1. V-LENGTH.--Greatest length from olecranon through tip of

internal condyle (Condylus ventralis).

2. W-SHAFT.--Transverse width of midshaft.

3. D-SHAFT.--Depth of midshaft.

4. W-PROX.--Greatest transverse width of proximal articular

surface.

5. D-LENGTH.--Length from tip of olecranon to tip of external

cotyla (Cotyla dorsalis).

6. D-PROX.--Depth of proximal end from cranial tip of internal

cotyla (Cotyla ventralis) to caudal margin (Margo caudalis) of

shaft of ulna, measured at right angles to the long axis of the

shaft.

7. ECON.--Length from external condyle (Condylus dorsalis) through

ventral face of distal end.










8. CPTB.--Length from carpal tuberosity (Tuberculum carpal e)

through lateral face of distal end.

9. ECON-CPTB.--Length from external condyle through carpal

tuberosity.

10. ECON-ICON.--Length from external condyle through internal

condyle.

Radius

1. LENGTH.--Greatest length from the radial head (Caput radii)

through distal end of the radius (Extremitas distale radii).

2. W-SHAFT.--Transverse width of midshaft.

3. D-SHAFT.--Depth of midshaft.

4. W-PROX.--Greatest transverse width of proximal end.

5. D-PROX.--Greatest depth of proximal end.

6. W-DIST.--Greatest transverse width of distal end.

7. D-DIST.--Greatest depth of distal end.

Carpometacarpus

1. LENGTH.--Greatest length from most proximal portion of the

carpal trochlea (Facies articularis radiocarpalis of trochlea

carpalis) through facet for digit III (Facies articularis

digitalis minor).

2. W-PROX.--Transverse width proximal end from ligamental

attachment of pisiform process (Processus pisiformes) to dorsal

surface (Facies dorsalis), measured at right angles to the long

axis of the shaft.

2a. W-CARPAL.--Transverse width carpal trochlea measured at the

proximal edge of the articular facet.











3. D-PROX.--Depth of proximal end from tip of process of

metacarpal I (Processus extensoris) through caudal part of carpal

trochlea (Facies articularis ulnocarpalis), measured at right

angles to the long axis of the shaft.

4. L-MCI.--Length metacarpal I (Os metacarpalis alulare) from

process of metacarpal I to pollical facet (Processus alularis).

5. D-SHAFT.--Depth of midshaft of metacarpal II ( Os metacarpale

majus).

6. W-SHAFT.--Transverse width of midshaft of metacarpal II.

7. D-DIST.--Greatest depth of distal end, measured across dorsal

edge of facet for digit II (Facies articularis digitalis major).

8. W-DIST.--Transverse width distal end from edge of facet for

digit II through facet for digit III.

Furculum

1. LENGTH.--Greatest length, measured from furcular process to

scapular tuberosity.

2. D-PROX.--Greatest diameter of coracoidal facet.

Femur

1. M-LENGTH.--Greatest length from head of femur (Caput femoris)

through medial condyle (Condylus medialis).

2. L-LENGTH.--Greatest length from trochanter (Trochanter femoris)

through lateral condyle (Condylus lateralis).

3. W-SHAFT.--Transverse width of midshaft.

4. D-SHAFT.--Depth of midshaft.

5. W-PROX.--Transverse width of proximal end, measured from the

head of femur through lateral aspect of trochanter, taken at

right angles to the long axis of the shaft.










6. D-HEAD.--Greatest depth of femoral head.

7. W-DIST.--Greatest transverse width of distal end.

8. W-M&LCON.--Transverse width of medial and lateral condyles from

(Crista tibiofibularis) to medial border of medial condyle.

9. W-LCON.--Transverse width of lateral condyle.

10. W-L&FCON.--Transverse width of lateral condyle and fibular

condyle.

11. D-FCON.--Greatest depth of fibular condyle.

12. D-LCON.--Greatest depth of lateral condyle.

13. D-MCON.--Greatest depth of medial condyle.

Tibiotarsus

1. L-LENGTH.--Greatest length from interarticular area (Area

interarticularis) on proximal articular surface through lateral

condyle (Condylus lateralis).

2. M-LENGTH.--Greatest length from the most proximal portion of

cnemial crest (Crista cnemialis cranialis) through medial

condyle (Condylus medialis). This includes the patella, if

fused, as in loons and grebes.

3. FIBULAR.--Length from interarticular area on proximal articular

surface to the most distal point of fibular crest (Crista

fibularis).

4. W-SHAFT.--Transverse width of midshaft.

5. D-SHAFT.--Depth of midshaft.

6. W-PROX-M.--Transverse width of proximal articular surface from

articular facet for fibular head (Facies articularis fibularis)

to medial border of proximal articular surface.










7. D-PROX.--Depth of proximal end from most caudal edge of medial

articular face (Facies articularis medialis) to the most cranial

point of the cranial cnemial crest.

8. W-PROX-L.--Transverse width of proximal end from medial border

of cranial cnemial crest to lateral border of lateral cnemial

crest (Crista cnemialis lateralis).

9. W-DIST-CR.--Transverse width of distal end, measured across

cranial portion of condyles.

10. W-DIST-CD.--Transverse width of distal end, measured across

caudal portion of condyles.

11. D-MCON.--Greatest depth of medial condyle.

12. D-LCON.--Greatest depth of lateral condyle.

13. D-ICON.-Depth of area intercondylaris.

Tarsometatarsus

1. LENGTH.--Greatest length from intercondylar eminence (Eminentia

intercondylaris) through trochlea for digit III (Trochlea

metatarsi III).

2. W-SHAFT.-Transverse width of midshaft.

3. D-SHAFT.--Depth of midshaft.

4. FLEXOR.--Intercondylar eminence to middle of tubercle for

tibialis anterior (Tuberositas m. tibialis cranialis).

5. W-PROX.--Greatest transverse width proximal articular surface,

measured across dorsal surface.

6. D-MCOT.--Greatest depth medial cotyla.

7. D-LCOT.--Greatest depth lateral cotyla.









8. D-PROX.--Depth from dorsal edge of proximal articular surface

to closest hypotarsal canal (Canalis hypotarsi), or the closest

hypotarsal groove (Sulcus hypotarsi), if no canals are present as

in the Accipitridae.

9. W-HYPOTS.--Greatest transverse width of hypotarsus.

9a. W-HYPOTS-C.--Transverse width of tuberosity on medial

hypotarsal crest (Crista medialis hypotarsi), in cormorants only.

10. L-HYPOTS.--Length of medial hypotarsal crest.

11. D-PROX-L.--Depth of proximal end, measured from dorsal edge of

the proximal articular surface through the lateral hypotarsal

crest (Crista lateralis hypotarsi).

lla. D-PROX-M.--Depth of proximal end, measured from dorsal edge

of the proximal articular surface through the medial hypotarsal

crest, if the lateral hypotarsal crest is reduced.

12. L-MTI.--Greatest length of metatarsal I facet (Fossa

metatarsi I).

13. D-D-SHAFT.--Depth of shaft at cranial edge of distal canal

(Foramen vasculare distale).

14. W-DIST.--Greatest transverse width of distal end (if trochlea

are of equal length).

15. TRIII-TRIV.--Greatest transverse width from trochlea III

through trochlea IV (if trochlea II is elevated).

16. TRII-TRIV.-Greatest transverse width between plantar portion

of trochlea II and plantar portion of trochlea IV.

17. W-TRII.--Greatest transverse width of trochlea II.

18. D-TRII.--Greatest depth of trochlea II.

19. W-TRIII.--Greatest transverse width of trochlea III.










20. D-TRIII.--Greatest depth of trochlea III.

21. W-TRIV.-Greatest transverse width of trochlea IV.

22. D-TRIV.--Greatest depth of trochlea IV.



Computer Software

Biomedical Statistical Software, P-Series (Dixon, 1981) was used to

analyze many of the measurements. Programs used included BMDP1D (simple

descriptive statistics), BMDP6D (bivariate plots), and BMDP2M (cluster

analysis). Computations were made at the Northeast Regional Data Center

(NERDC) at the University of Florida, Gainesville.


Nomenclature

Common names have not been used for species. Few nomenclatural

systems have a more unstable, inaccurate, or confusing set of terms than

the American Ornithologists' Union's (1983) list of common names for

birds. I agree with J. L. Peters (1934:ii)

S inventing common English names for birds that do not have
them is a waste of time. After all, the primary reason for a
scientific name is to have a name intelligible to scientists
the world over.

Systematic nomenclature generally follows the Checklist of Birds of

the World (Mayr and Cottrell, 1979; Peters, 1931-1951) for Recent species

and Brodkorb's Catalogue of Fossil Birds (1963-1978) for fossil species.

Departures are accompanied by full citation.

































Figure 2.1. Schematic diagrams illustrating measurements of the
humerus, coracoid, and scapula. A. Humerus, cranial view. B.
Humerus, ventral view. C. Coracoid, dorsal view. D. Coracoid,
lateral view. E. Humerus, distal end view. F. Scapula, lateral
view. G. Scapula, proximal end view. Figures are not drawn to
scale. Measurements defined in the text.









B
~C


4

D


G



































Figure 2.2. Schematic diagrams illustrating measurements of the
radius, ulna, carpometacarpus, and furculum. A. Radius, medial
view. B. Radius, cranial view. C. Ulna, cranial view. D.
Ulna, caudal view. E. Carpometacarpus, ventral view. F.
Carpometacarpus, proximal end view. G. Carpometacarpus, distal
end view. LH. Ulna, distal end view. I. Furculum, lateral view.
Figures are not drawn to scale. Measurements are defined in text.




A
1a


D

8
3

































Figure 2.3. Schematic diagrams illustrating measurements of the
femur and tibiotarsus. A. Tibiotarsus, caudal view. B.
Tibiotarsus, proximal end view. C. Tibiotarsus, distal end view.
D. Femur, cranial view. E. Femur, proximal end view. F. Femur,
distal end view. G. Femur, caudal view of distal end. Figures
are not drawn to scale. Measurements are defined in the text.











A













1 13 12


-1o0--I



































Figure 2.4. Schematic diagrams illustrating the measurements of
the tarsometatarsus. A. Dorsal view. B. Lateral view. C.
Plantar view of distal end. D. Distal end view. E. Proximal end
view. Figures are not drawn to scale. Measurements are defined in
text.





26



B

A N \~10
4

\w


3

2- 1


C

13 12



14-1


15
Is-- 7
F/19- E

20 67 :8~

S22 21 17 18 MED
IV1I
k- 16










Systematics

I have associated the skeletal elements of the fossil taxa in this

study using the following general criteria (Howard, 1932b):

1. resemblance to other species.

2. size and proportion.

3. relative abundance.

Systematic reasoning and taxonomic practice generally follow Mayr

(1981), when possible. Skeletal elements are first grouped into

similarity classes by means of measurements and qualitative characters.

Each identifiable taxon is placed within a geneology of previously known

species by the hierarchial distribution of shared-derived characters, and

the new species are then integrated into an evolutionary classification.


Paleoecology

Paleoecological methods are described by Shipman (1981). See the

paleoecology section for further comments on the paleoecological methods

used in the excavation of the localities included in this study.


Biochronology and Faunal Dynamics

A database for the examination of the biochronology and the faunal

dynamics of fossil birds of the Neogene of North America was developed

from the following sources: (1) A literature survey of all fossil

localities in North America from the late Arikareean through the Blancan

that have produced fossil birds. This published information was then

emended to reflect the current concepts of geological formations,

geological correlations, mammalian systematics, and the relative and

absolute dating of fossil localities. Verification of all fossil

identifications was made whenever possible. (2) Information from other









major unpublished localities was added to this database. This produced a

total of 133 localities from the Neogene of North America (86 published,

47 unpublished) with a record of fossil birds. (3) The occurrences of

fossil taxa were condensed to tabular form to reflect the actual,

documented range of taxa at the family, subfamily, and generic level. A

given taxonomic range reflects the summation of all lower taxonomic ranks

plus material which is only diagnostic to that given level. (4) From

this table, geological ranges were inferred parsimoniously. For example,

if a genus is known from the early Hemingfordian and the late

Clarendonian, I inferred that it also occurred in North America during

the interval between these endpoints. (5) From this table (#3 above),

biochronologically useful species were identified.

Indices of avian faunal dynamics were calculated following Marshall

et al. (1982): the duration (d) of each land mammal age is given in

millions of years, to the nearest tenth, and is based on all available

data; diversity (Si) represents the total number of genera known for each

land mammal age; originations (Oi) are the number of generic first

appearances in a given land mammal age; extinctions (Ei) are last

appearances of a genus in a given land mammal age; running means (Rm)

compensates for time intervals of unequal duration by subtracting the

average of originations (Oi) and extinctions (Ei) for a given age from

the diversity (Si) of that age, or Rm = Si (Oi + Ei)/2; origination

rates (Or) adjust for unequal time intervals by dividing the total number

of generic originations (Oi) occurring during a given land mammal age by

the duration (d) of that interval; similarly, the extinction rate, Er =

Ei/d; turnover rates (T) are the average number of genera that either

originate or go extinct during a given land mammal age, or T = (Or +









Er)/2; per-genus turnover rate is the turnover rate adjusted for average

diversity, calculated by dividing the total turnover rate (T) by the

total running mean (Pm). A sampling index was calculated as the number

of localities per Land Mammal Age divided by the duration of that

interval. Marshall et al. (1982) should be referenced for additional

qualifications of each statistic.


Specimens Examined

Fossil specimens included in this study are housed in the

collections of the Florida State Museum (UF), the collection of Pierce

Brodkorb (PB), and the Frick collections of the American Museum of

Natural History (F:AM). I have tried to include all known avian material

from the late Miocene through early Pliocene from Florida within these

collections. Non-diagnostic skeletal elements (vertebrae, phalanges,

etc.) were not considered. Fossil material accessioned into the UF

collections after 01 June 1984 (primarily from Bone Valley) was not

included in this study.

I relied on Recent comparative material in the collections of P.

Brodkorb, Florida State Museum, United States National Museum, American

Museum of Natural History, University of Michigan, and Royal Ontario

Museum.

Abbreviations

Table 2.1 lists the common abbreviations and acronyms used in this

dissertation. Anatomical abbreviations were given earlier.











Table 2.1. List of acronyms of institutions and abbreviations of terms
used in the text.

Acronym Institutions/Collections

AMNH American Museum of Natural History

F:AM Frick Collections, American Museum of Natural History

LACM Los Angeles County Museum of Natural History

MCZ Museum of Comparative Zoology, Harvard University

PB Collection of Pierce Brodkorb

ROM Royal Ontario Museum

UF Florida State Museum, University of Florida

UM University of Michigan

UMCP University of California. Museum of Paleontology.


UNSM

USNM

YPM


Berkeley

University of Nebraska State Museum

United States National Museum

Yale Peabody Museum


Abbreviation

BMDP

1. f.

M.

MA

max.

min.

En.

MYBP

N

NALMA


Terms

Biomedical Statistical Program, P-series

local fauna

musculus

megannum (or million years)

maximum

minimum

millimeter

million years before present

number (of specimens)

North American Land Mammal Age
















CHAPTER III
GEOLOGY

Biochronology

The Clarendonian and Hemphillian land mammal ages were first

proposed by Wood et al. (1941) based on the stage of evolution of mammals

from two localities in the panhandle of Texas. Since then, the concept

of each has changed, owing to the increasing knowledge of this time

period in North America. Two additional ages were proposed by Schultz et

al. (1970) for parts of the time intervals covered by the original

definitions: the Valentinian (for the late Barstovian to early

Clarendonian) and the Kimballian (originally proposed as late

Hemphillian; now considered to be early Hemphillian). Neither has been

accepted for use on a continent-wide basis. Rather, each has been

applied only to fossils from the type formations of the proposed ages

(Valentine and Kimball formations). Pertinent references include Schultz

et al. (1970), Tedford (1970), Tedford et al. (in press), Breyer (1981),

and Voorhies (1984).

The Clarendonian, in the restricted sense of Tedford et al. (in

press) is defined on the

earliest appearance of Barbourofelis and, later in the
interval, Platybelodon, Amebelodon, and Ischyrictis
(Hoplictis). Characterization earliest appearance of
Nimravides, Epicyon (in the Great Plains and Gulf Coast),
Griphippus [=Pseudhipparionl, Astrohippus, Nannippus (Gulf
Coast), Macrogenis, Synthetoceras, Hemiauchenia, Megatylopus,
Antilocaprinae (Plioceras and Proantilocapra), latest
occurrence of Eucastor, Brachypsalis, Ischyrocyon, Cynarctus,
Aelurodon, Tomarctus, Hypohippus, Megahippus, Merychippus,
Paratoceras, Miolabis, Protolabis, and probably Ustatochoerus.










In Florida, common late Clarendonian taxa include Barbourofelis,

Nimravides, Mylagaulus, Pseudhipparion, Amebelodon, advanced species of

Eucastor, Pediomeryx, and Aelurodon.

The early Hemphillian is defined (Tedford et al., in press) by the

earliest appearance of Arvicolinae (limited occurrence of
Microtoscoptes and Paramicrotoscoptes), Pliotomodon, and
Megalonychidae (Pliometanastes), limited occurrence late in
interval of Simocyon, Indarctos, Plionarctos, Lutravus, and
Eomellivora, and earliest appearance, late in interval, of
Mylodontidae (Thinobadistes), Machairodus and the Bovidae
(Neotragocerus ). Characterization earliest appearance of
Dipoides, Pliosaccomys, Pliotaxidea, Vulpes, 'Canis',
Osteoborus, and Cranioceras (Yumaceras), latest occurrence of
Amphicyonidae, Leptarctus, Sthenictis, Nimravides,
Barbourofelis, Epicyon, Pliohippus, Protohippus,
Cormohipparion, Prosthenops, Aepycamelus, Pseudoceras and
Plioceras.

In Florida, common early Hemphillan taxa include Calippus,

Pediomeryx (Yumaceras), Aepycamelus, primitive species of Osteoborus,

Cormohipparion, and Nannippus, and advanced species of Epicyon. The

early sloths Pliometanastes and Thinobadistes are present. The

Eurasiatic immigrants Indarctos and Machairodus appear in the later part

of the early Hemphillian.

The late Hemphillian is defined (Tedford et al., in press) by the

limited occurrence of Promimomys, 'Propliophenacomys',
Plesiogulo, Agriotherium, and Plionarctos and the earliest
appearance of Megalonyx, Ochotona, Megantereon, Felis,
Enhydriodon, and Cervidae. Characterization earliest
appearance of Taxidea, Borophagus, Rhynchotherium, Platygonus,
and Mylohyus, limited occurrence of Pediomeryx, latest
occurrence of Mylagaulidae, Osteoborus, Astrohippus,
Neohipparion, Dinohippus and Rhinocerotidae.

In Florida, common late Hemphillian taxa include advanced species of

Osteoborus, Neohipparion, Nannippus, Pseudhipparion, Teleoceras, and

Hipparion. Also present are Plesiogulo, Megalonyx, antilocaprids,

Agriotherium, Enhydriodon s.l., Megantereon, Machairodus, Rhynchotherium,


Pliomastodon, Dinohippus, Plionarctos, Felis, and cervids.












Local Faunas

Love Bone Bed

The Love Bone Bed is located near the town of Archer, Alachua

County, along State Road 241, in the NW 1/4, SW 1/4, NW 1/4, Sec. 9, T.

11 S., R. 18 E., Archer Quadrangle, U. S. Geologic Survey 7.5 minute

series topographical map, 1969. Excavation and collection of fossil

vertebrates by the Florida State Museum took place from its discovery in

1974 until the quarry was closed during the summer of 1981. This local

fauna originated from the Alachua Formation (Williams et al., 1977).

The Love Bone Bed is considered latest Clarendonian in age (Webb et

al., 1981). Studies published to date on the Love Bone Bed and its

vertebrate fauna include a general overview of the geology and

paleontology of the locality, including a preliminary faunal list (Webb

et al., 1981); studies on the turtles Pseudemys caelata, and Deirochelys

carri (Jackson, 1976, 1978); a description of the rodent Mylagaulus

elassos Baskin (1980); description of the carnivores Barbourofelis lovei

and Nimravides galiani Baskin (1981); descriptions of the procyonids

Arctonasua floridana and Paranasua biradica Baskin (1982); the

description of the ruminant Pediomeryx hamiltoni Webb (1983); and a

population study on the three-toed horse Neohipparion cf. N. leptode

(Hulbert, 1982). Studies on fossil birds include papers on the fossil

herons (Becker, 1985a), a description of a new species of osprey (Becker,

1985b), and one on the fossil anhinga (Becker, ms.). Additional studies

are in progress.









Mixson Bone Bed

The Mixson Bone Bed is located approximately 2 miles northeast of

Williston, Levy County, in the NE 1/4, SW 1/4, Sec. 29, T. 12 S., R. 19

E., Williston Quadrangle, U. S. Geological Survey 7.5 minute series

topographical map, 1969. This is the type locality of the Alachua

Formation (Dall and Harris, 1892). There have been many studies on this

site, including papers by Leidy and Lucas (1896), Sellards (1916), Hay

(1923), Simpson (1930), Webb (1964, 1969, in press), and Harrison and

Manning (1983). Additional references are listed in Ray (1957).

Genera in common with McGehee include Calippus, Pediomeryx

(Yumaceras), Aepycamelus, Osteoborus, and Aelurodon (Webb, 1969). The

early Hemphillian mylodontid sloth Thinobadistes is present. Two other

early Hemphillian index genera are absent from this local fauna, although

they are present in other Florida sites--Pliometanastes, and Indarctos.

The first appearance of Indarctos occurs late in the early Hemphillian

and its absence in this local fauna probably has temporal significance.

The absence of Pliometanastes is usually considered an ecological

sampling bias.


McGehee Farm

This locality is almost exactly three miles north of Newberry,

Alachua County, along State Highway 45, Sl/2, NW1/h, Sec. 22, T. 9 S., R.

17 E., Newberry Quadrangle, U. S. Geologic Survey 7.5 series

topographical map, 1968, in northcentral Florida. This locality was

first discovered in 1958 and was extensively collected by the University

of Florida, with support of the Frick Corporation. This local fauna

originates from the Alachua Formation, and the geology of this locality

is briefly discussed by Webb (1964) and Hirschfeld and Webb (1968). The










latter publication includes a preliminary list of the fossil vertebrates

from this local fauna. No faunal revision of this locality has been

undertaken, but several papers treating specific groups have appeared

(sloths: Hirschfeld and Webb, 1968; nylagaulids: Webb, 1966; canids:

Webb, 1969; and protoceratids: Patton and Taylor, 1973).

The faunal composition of this local fauna indicates an early

Hemphillian age (Hirschfeld and Webb, 1968; Webb, 1969; Marshall et al.,

1979), based primarily on the presence of Pliometanastes, the early

Hemphillian megalonychid ground sloth. Typical early Hemphillian

mammalian genera present include Calippus, Pediomeryx (Yumaceras),

Aepycamelus, Osteoborus, and Aelurodon (Webb, 1969).

The birds of this locality were first studied by Brodkorb (1963a)

who reported Phalacrocorax wetmorei, and described two new species-

Nycticorax fidens and Ereunetus rayi. Later, Olson (1976) described

Jacana farrandi from here.


Withlacoochee River hA

The Withlacoochee River 4A local fauna lies approximately 8 km.

southeast of Dunnellon (center of N1/2, NW1/4, Sec. 30, T. 17 S., R. 20

E., Stokes Ferry Quadrangle, U. S. Geologic Survey 7.5 minute series

topographical map, 1954, Marion County), in northcentral Florida. The

fossil vertebrates originate from a massive green clay filling of a

sinkhole in the late Eocene Inglis Formation of the Ocala Group (Webb,

1969, 1973, 1976). This deposit of green clay is being eroded by the

Withlachoochee River; the fossils were collected in approximately 20 feet

of water using scuba equipment. Deposition almost certainly occurred in

a pond environment near sea-level as shown by the fine-grained sediments










with articulated fish skeletons, the present elevation, and the marine

taxa present (Becker, 1985a; Berta and Morgan, in press).

Studies on the fossil mammals from here include Webb's (1969) paper

on Osteoborus orc, Hirschfeld and Webb's (1968) study on Pliometanastes

protistus, Webb's (1973) mention of antilocaprids, and Wolff's (1978)

study of the cranial anatomy of Indarctos. The concurrent range zones of

the first two taxa, and the presence of Indarctos, Machairodus, and

Pseudoceras indicate an age of late early Hemphillian.

Fossil birds from here include a very a small species of Egretta and

an indeterminate species of Buteo (Becker, 1985a). A preliminary faunal

list is included in this paper.


Haile VB

This locality is from the NE 1/4, Sec. 23, T. 9 S., R. 18 E.,

Newberry Quadrangle, U. S. Geologic Survey 7.5 minute series

topographical map, 1968. Auffenberg (1954) discusses the geology of this

locality and describes the abundant material of Gavialosuchus americanus

(Sellards) from this locality. A number of equids are known from this

site including Pliohippus, Cormohipparion, Calippus, and Nannippus.


Haile VI

This locality is in the N1/2, SW1/4, Sec. 24, T. 9 S., R. 17 E.,

Newberry Quadrangle, U. S. Geologic Survey 7.5 minute series

topographical map, 1968, Alachua County, Florida. It was collected by

the Florida Geologic Survey and Florida State Museum. Auffenberg (1963)

discussed the geology and concluded that this locality represents a

stream deposit. The mammalian fauna includes 'Hipparion', Pseudoceras,

and the type of Mylagaulus kinseyi Webb (1966). A new species of










sparrow, Palaeostruthus eurius, was described from here (Brodkorb,

1963a). Reptiles known from here include Deirochelys and Gavialosuchus.


Haile XIXA

This locality is 2.5 miles NE of Newberry, Alachua County, in the NE

1/4, Sec. 26, T. 9 S., R. 17 E., Newberry Quadrangle, U. S. Geologic

Survey 7.5 minute series topographical map, 1968. It was collected by

the Florida State Museum staff. Vertebrates are mainly aquatic,

including a large amount of skeletal material of Gavialosuchus. Fossil

mammals present include Epicyon, geolocids, Pediomeryx, equids, and

Aepycamelus.


Bone Valley Mining District

The name "Bone Valley" is applied to the phosphate mining district

of central Florida, mainly in Polk County, but also including portions of

adjacent Hillsborough, Hardee, and Manatee counties. The vertebrate

fauna was first described by Sellards (1916) and later more fully studied

by Simpson (1930). Berta and Morgan (in press) present an account of the

present status of vertebrate paleontology of this area. Much of the

following comes from their paper.

Most of the fossil vertebrates from the Bone Valley area were

obtained from extensive open-pit phosphate mines. In situ collections

are rare and make up approximately 5 10 % of the Florida State Museum

collections, part of the USGS and USNM collections, and virtually none of

the Harvard collections. Avian fossils fron these in situ sites are very

rare. Fossils are more commonly found eroding from spoil piles after an

area has been strip-mined. Except for the in situ collections and a few

intensively collected concentrations, the fossil vertebrates collected










from one mine, or a single dragline operating within one mine, are

considered as coming from one broad "locality." The exact geographic

position of these localities range from precisely known (1/h, 1/4, 1/4

section) to generally known (from one mine--i.e. somewhere within 1 to

10 sections). For a few amateur collections where even the mine is

unknown, the only designation can be the Bone Valley Mining District

(i.e. somewhere within 100 square miles), but the majority of specimens

are identified as coming from one mine. Table 3.1 lists the mines, mine

codes, describes their approximate location, and lists the stratigraphic

codes used.

The age of the Bone Valley fossil vertebrates was much debated from

the 1920s through the 1950s (discussed in Brodkorb 1955a), with the

proposed age ranging from the Miocene through the Pleistocene. Brodkorb

(1955a), using a Lyellian method of percent extinct species in the fauna

and the temporal ranges of three species, suggested that the age of the

Bone Valley avifauna was between the late Miocene and middle Pliocene,

probably early or middle Pliocene (i.e. Clarendonian or Hemphillian; =

late Miocene of current usage).

The majority of fossil land mammals are late Hemphillian in age and

compare well with others of similar age in North America. There is no

evidence to suggest that the fossil birds, which are found in association

with these land mammals, are of a different age. Recently several older

local faunas (Barstovian, Clarendonian, and early Hemphillian) have been

found (MacFadden, 1982; MacFadden and Webb, 1982; Webb and Crissinger,

1984; Berta and Galiano, 1984; and Tedford et al., in press). These

older occurrences are from the Phosphoria, Nichols, Silver City, and

Kingsford mines and to my knowledge have produced no fossil birds. There










are also numerous Pleistocene sites in the Payne Creek Mine (Steadman,

1984), Peace River Mine (=Pool Branch; Webb, 1974), and Nichols Mine.

The Pleistocene fossil birds from these sites can usually be separated

from geologically older specimens by their association with Pleistocene

fossil land mammals. The Pleistocene fossil birds are not considered in

this study.

There have been many recent studies on non-marine mammalian taxa

from Bone Valley (Baskin, 1982; Berta and Galiano, 1983; Berta and

Morgan, in press; Harrison, 1981; MacFadden and Waldrop, 1980; MacFadden

and Galiano, 1981; MacFadden, 1984; Webb, 1969, 1973, 1983; Webb and

Crissinger, 1984; Wright and Webb, 1984), but no single faunal study of

terrestrial mammals from this local fauna. Reference to the earlier

papers published on the Bone Valley Mining District and its vertebrate

fauna are given by Ray (1957).

Bone Valley taxa which are typical of the late Hemphillian age

include the Eurasian immigrant taxa Agriotherium, Plesiogulo,

'Enhydriodon', and Cervidae. Other taxa present are Rhynchotherium,

Hexameryx, and advanced species of Osteoborus, Gomphotherium,

Pseudhipparion, Nannippus, and Dinohippus (Berta and Morgan, in press).

Other authors (MacFadden and Galiano, 1981; Berta and Galiano, 1983;

Wright and Webb, 1984) suggest that the upper Bone Valley Formation is

very late Hemphillian because of the presence of taxa equally indicative

of an early Blancan age such as Felis rexroadensis, Meganteron hesperus,

Dinohippus mexicanus, Mylohyus elmorei, Antilocapra (Subantilocapra),

Hemiauchenia, Nasua, and Platygonus.

The birds from Bone Valley have been studied by Brodkorb (1953b,

1953c, 1953d, 1953e, 1955a, 1970). His monograph (1955a) dealt with the










birds then known, primarily the more numerous marine birds. Table 3.2

gives a partial list of birds now known from the Bone Valley Mining

District and notes the taxa (marine) not included here. The majority of

birds here studied have been collected in the last 15 years and mainly

include the rarer, non-marine members of this avifauna.


Manatee County Dam Site

This local fauna originates from a borrow pit south of the Manatee

River in Sec. 30, T. 34 S., R. 20 E., Verna Quadrangle, U. S. Geologic

Survey 7.5 minute series topographical map, 1944, Manatee County. Like

the nearby SR-64 local fauna discussed below, the Manatee County Dam Site

is essentially an outlier of the classic Bone Valley local fauna. All

three share a similar fauna, geology, and paleoecology. Webb and Tessman

(1968) describe this local fauna and report the presence of one bird,

Phalacrocorax wetmorei.


SR-64

This locality is located 6 miles east of 1-75 along State Road 64 in

Sec. 35, T. 34 S., R. 19 E., Manatee County, Lorraine Quadrangle, U. S.

Geological Survey 7.5 minute series topographical map, 1973, Florida. It

was discovered by Philip Whisler, of Venice, Florida, in 1983. The

majority of fossil vertebrates are in his private collection, except for

a few speciemens in the Florida State Museum collections. Numerous

fossil vertebrates are recorded from here, including several species of

bony fish, sharks, rays, and several different turtles. Mammals include

odobenids, phocids, Schizodelphis, Megalodelphis, tremarctine ursids, cf.

Agriotherium, felids, Nannippus minor, Neohipparion eurystyle,

Rhinocerotids, and Hexameryx. Based on the presence of Nannippus minor,










Neohipparion eurystyle, and Hexameryx, this locality is late Hemphillian

in age. As for the Manatee County Dam Site, this locality is here

considered an outlier of the classic Bone Valley Formation.


Eustatic Sea-Level Changes

Webb and others (Webb and Tessmann, 1968; MacFadden and Webb, 1982;

Webb, 1984) have proposed a model using the present elevations of fossil

localities with marine or estuarine taxa to reflect the fluctuations of

sea-levels during the late Miocene and early Pliocene in peninsular

Florida. This scheme is predicated on two assumptions--that the

Florida peninsula had been tectonically stable over the later Cenozoic

and that the sedimentary sequence reflects actual sea-level change. If

these assumptions are valid, then the present elevation of the localities

which contain marine or estuarine taxa should reflect the elevation of

the ocean at the time when these localities were deposited.

The following evidence argues against this model: (1) The relict

Pleistocene shorelines increase in elevation from a low point in southern

Georgia to a maximum elevation in northern peninsular Florida and

gradually lose elevation to the south. This indicates that the northern

part of peninsular Florida has not been stable, and has been

differentially uplifted. Opdyke et al. (1984) argue that this occurred

during the Pleistocene due to the subsurface solution of limestone and

the concomitant isostatic uplift and document an uplift in the magnitude

of 30-50 meters since this time. Most of the high-sea-level localities

are from the northern part of Florida in highly karsted areas.

(2) The elevational changes between most localities could easily be

accounted for by a slight regional dip to the beds. For example, a dip

of 1/20 of 1 degree would account for an elevational difference of 138











feet over 30 miles (the distance between the Manatee County Dam Site and

the "classic" Bone Valley exposures). This would also account for the

mixture of land and marine vertebrates in these sites, presuming the

tilting is post-depositional.

There is no question that there were eustatic sea-level changes

during this time period, such as the Messinian. But the elevations of

the Florida fossil vertebrate localities are of very doubtful value as

evidence. It should be noted that the rejection of the above hypothesis

does not prevent using the faunal composition of these localities to

determine their relative proximity to the ocean at the time of

deposition. They simply cannot be used as an absolute scale to measure

vertical sea-level changes.





































Figure 3.1. Correlation Chart of Included Local Faunas.










NORTH
AMERICA!
LAND
MAMMAL
AGES


z

-J
-J
*I
a-
2 >w

UJ


w
I-
-J


MANATEE CO. ISR-64
DAM L.F. L.F.


ITHLACOOCHEE
RIVER 4A L.F.





































Figure 3.2. Location of Included Local Faunas.






46

LOCAL FAUNA

ALACHUA 1 MCGEHEE
2 HAILE SITES
1 3 LOVE BONE BED
02 e 4 MIXSON BONE BED
4* 5 WITHLACOOCHEE
RIVER 4A
LEVY 6 BONE VALLEY
7 SR-64
I--IMARION
8 MANATEE CO. DAM










i5
0 CITRUS ----








N
'F
_

POLK




6



| ~7"


^ MANATEE \









Table 3.1. A partial list of Bone Valley mines, their mine codes,
approximate location, and the stratigraphic codes commonly used.


Mines

Chicora

District Grade

Estech

Ft. Green

Ft. Meade

Gardinier

Hooker's Prairie

Kingsford

New Palmetto

Nichols

Palmetto

Payne Creek

Peace River

Swift

Tiger Bay


Stratigraphic Codes

0

1

2

3

4

5


Codes

BVC

BVPC

BVS

BVFG

BVFM

BVG

BVHP

BVK

BVNP

BVN

BVP

BVPC

BVFM

BVS

BVTB


Township


see

see

32S

31S

32S

31S

31S

32S

30S

32S

32S

see

31S

31S


Range Sections


Payne Creek Below

Swift Below

23E

25E

24,24E

24E

23E

24E

23E

24E

24E

Ft. Meade Above

24E

24E


2,3,10-14,22-24

2,13

unknown

17,18,20,28-30

3

3

19,28,29

9,10,15,16,21,22

13,14,23,24,29-32



14

12


Explaination

No stratigraphic data

In place Hawthorn Fm. dolomitic

In place "lower Bone Valley Fm."

In place "upper Bone Valley Fm."

In place Pleistocene sediments

Soil zone (upper clay)










Table 3.2 Checklist of birds from the late Miocene and early Pliocene
Bone Valley Mining District. Asterisks denote marine taxa, which are not
included in this study. Taxa are based on previously published works and
on original identifications.

*Family Gaviidae Family Ciconiidae


*Gavia palaeodytes

*Gavia concinna

Family Podicipedidae

Podiceps sp.

Pliodytes lanquisti

*Family Diomedidae

*Diomedea anglica

Family Phalacrocoracidae

Phalacrocorax wetmorei

Phalacrocorax idahensis

*Family Sulidae

*Morus peninsularis

*Sula guano

*Sula phosphata

*Family Procellaridae

*Family Pelecanidae

*Pelecanus sp.

Family Plataleidae

Eudocimus sp.

Family Ardeidae


Ciconia sp.

Family Anatidae

Bucephala ossivallis

Family Pandionidae

Pandion sp.

Family Accipitridae

Haliaeetus sp.

Buteo sp.

Family Scolopacidae

Calidris pacis

Erolia penpusilla

Limosa ossivallis

Family Phoenicopteridae

Phoenicopterus floridanus

*Family Haematopidae

*Haematopus sulcatus

*Family Laridae

*Larus elmorei

*Family Alcidae

*Australca grandis


Ardea polkensis














CHAPTER IV
SYSTEMATIC PALEONTOLOGY

Table 4.1 lists the non-marine avian taxa now known to occur in

Florida during the late Miocene and early Pliocene. In the following

systematic section, I have tried to present osteological characters which

define the taxonomic groups in a hierarchical fashion. However,

considering the relatively small number of fossil and recent specimens

available for some species, and the restricted geographical area from

which many of the Recent species were collected, I would not be surprised

if some "diagnostic" characters do not hold when a larger number of

specimens are examined.


Order Podicipediformes (FUrbringer, 1888)

Family Podicipedidae (Bonaparte, 1831)

Tribe Podilymbini Storer, 1963

Characters. Separate canal through hypotarsus for the tendon of

insertion of M. flexor perforatus digiti II (Storer, 1963). Murray

(1967) gives additional osteological characters for the separation of the

different taxa of this family.


Genus Rollandia Bonaparte, 1856

Rollandia sp.

Material. Love Bone Bed local fauna, questionably referred; UF

29670, UF 29673, right coracoids; UF 25815, left coracoid.

Mixson Bone Bed local fauna; F:AM FLA-120-2183, complete right

tarsometatarsus.










McGehee Farm local fauna; UF 9488, shaft and distal end of right

humerus; UF 12468, right coracoid.

Description. Humerus (UF 9488) poorly preserved; larger and much

more robust than both species of Tachybaptus (dominicus and ruficollis).

Morphology of distal end similar to that of Rollandia rolland, but shaft

more robust. Measurements given in Table 4.2.

Coracoids from the Love Bone Bed are extremely worn and are here

assigned strictly on the basis of their size being near Rollandia (larger

than all species of Tachybaptus and smaller than the smallest males of

Podilymbus podiceps). Shaft similar to, or slightly more robust than,

that of Rollandia rolland. Proximal edge of ventral sternal articulation

flared (as in Rollandia rolland, less so in Tachybaptus). Coracoid from

McGehee Farm local fauna (UF 12488) similar to those from the Love Bone

Bed, except by having a proportionally longer shaft. Measurements are

given in Table 4.2.

Overall length of tarsometatarsus similar to Rollandia rolland

chilensis. Distinguished from Podilymbus by lacking the cranial

expansion of the proximal end of the tarsometatarsus and by having

trochlea II placed lower on the shaft. Distinguished from Podiceps

species by having the shaft less laterally compressed. Tarsometatarsus

differs from that of Rollandia rolland chilensis by having a more deeply

excavated lateral parahypotarsal fossa, a hypotarsus with a smaller

transverse width, a more distinct ridge extending distally from the

hypotarsus, and trochlea II slightly more narrow. Measurements given in

Table 4.3.

Remarks. This species appears to be slightly more robust than the

modern Rollandia rolland chilensis. The lack of a series of modern sexed










skeletons, necessary to determine the variability of the characters used

above, prevents the naming of this species as new.


Genus Tachybaptus Reichenbach, 1853

Tachybaptus sp. indet.

Material. Love Bone Bed local fauna; UF 25796, UF 25817, UF 25818,

UF 29671, complete left coracoids; UF 29668, UF 29669, UF 29672, humeral

ends left coracoids; UF 26006, UF 26014, UF 26017, UF 26019, UF 29664, UF

29665, UF 29666, UF 29669, complete right coracoids. UF 25773, complete

right femur. UF 29663, proximal end of right tibiotarsus (questionably

referred).

McGehee Farm local fauna; UF 67810, proximal end right tibiotarsus

(questionably referred).

Description. All coracoids are waterworn, abraded, or broken to

varying degrees, making detailed descriptions difficult. All coracoids

near that of Tachybaptus dominicus in size and overall shape. -Additional

description is not possible.

Femur also near Tachybaptus dominicus in size and general

morphology, but the femoral shaft slightly more gracile. Depression

cranial to the patellar sulcus is absent in fossil. Measurements are

given in Table 4.4.

Both tibiotarsal fragments are questionably referred solely on the

basis of size.

Remarks. The use of generic names follow Storer (1976a). This

material is not diagnostic enough to allow identification to the level of

species.










Genus Podilymbus Lesson, 1831

Podilymbus cf. P. podiceps

Material. Bone Valley Mining District, Gardinier Mine; UF 65678,

distal half of left tibiotarsus; Palmetto Mine; UF 21147, distal half of

left tibiotarsus.

Description. Tibiotarsus similar in size and general morphology to

that of Podilymbus podiceps. Distinguished from the tibiotarsus of

Podiceps by having a smooth medial border of the medial condyle (notched

in Podiceps) and a less distinct depression epicondylaris medialis (very

deep in Podiceps). Differs from that of Podilymbus podiceps as follows:

Shaft of UF 21147 slightly more robust than in males; tubercle slightly

proximal to medial attachment of supratendial bridge better developed;

deeper depression epicondylaris lateralis than in most specimens. Other

characters within range of variation of modern populations of Podilymbus

podiceps. Measurements are given in Table 4.5.


Podilymbus sp. A

Material. Mixson Bone Bed local fauna; F:AM FLA 66-1115, proximal

end of right tarsometatarsus; F:AM FLA 66-1116, proximal end of left

tarsometatarsus.

Remarks. Both specimens are juvenile, at a similar stage of

ossification, and have identical morphology; they may represent the same

individual. Tarsometatarsal morphology similar to that of Podilymbus

podiceps, but differs by being smaller, having a sharper intercondylar

knob, and a smooth cranio-dorsal border of the medial cotyla (notched in

P. podiceps). Both specimens are too small to correspond to that

expected of Podilymbus cf. P. podiceps from the Bone Valley.

Measurements given in Table 4.3.












Tribe Podicipedini Storer, 1963

Characters. Absence of a separate canal in the hypotarsus for the

tendon of insertion of the M. flexor perforatus digiti II (Storer, 1963).

See Murray (1967) for additional osteological characters.


Genus Podiceps Latham, 1787

Podiceps sp. indet.

Material. Bone Valley Mining District, Payne Creek Mine; UF 21205,

proximal end of left tarsometatarsus.

Remarks. Agrees with Podiceps in hypotarsal configuration (no

extra canal). Waterworn and abraded, and is not identifiable to species.

Similar in size to Podiceps nigricollis or P. o. occipitalis. (P. auritus

much larger).


Tribe indet.

Genus Pliodytes Brodkorb, 1953

Pliodytes lanquisti Brodkorb, 1953

Material. Bone Valley Mining District, Palmetto Mine; PB 299,

complete right coracoid holotypee).

Remarks. This species is known only from the holotype. Brodkorb

(1953e) states that it possesses characters in common with both

Podilymbus and Podiceps but also has its own unique characters. As the

tribes of grebes are defined on tarsometatarsal characters (Murray, 1967;

Storer, 1963), it is not possible to assign this genus to a tribe. As

more fossil material of grebes from the Bone Valley becomes available,

this species should be restudied to determine its generic validity and

relationships to other species.










Remarks on the Family Podicepidadae.

The family Podicipedidae includes 11 fossil and 19 living species.

The earliest certain grebe is Podiceps oligocaenus (Shufeldt), based on a

fragmented left femur (missing the proximal end, distal end badly

abraded) from the Arikareean John Day Formation, Oregon. It is

intermediate in size between the living Podiceps grisegena and P.

nigricollis. Although Wetmore (1937) considers it to be correctly

allocated to genus, next to nothing can be said about its relationships.

Podiceps pisanus (Portis) based on the distal end of a right

humerus, is from the Middle Pliocene (=late Miocene?) of Italy. This

species may also be present at the Hemphillian Lee Creek local fauna

(Olson, ms.). The only other late Miocene species of grebe now known is

Pliodytes lanquisti Brodkorb, discussed above.

There are five species of Pliocene (i.e. Blancan) grebes, all from

North America. Podiceps subparvus (L. Miller and Bowman), from the early

Blancan San Diego Formation, California, is based on a distal end of a

femur. It is approximately the same size as that of the living

Podilymbus podiceps and is now known from additional material. Murray

(1967) in his review of Pliocene grebes, described one new genus and four

new species. Pliolymbus baryosteus Murray, from the Fox Canyon local

fauna, Kansas, of Blancan age, is based on the cranial portion of a

sternum. Murray (1967) states that this is a small grebe with a robust

skeleton but does not suggest any possible relationships between this

species and other living or fossil species of grebes. Podiceps discors

Murray, also from Fox Canyon, is based on a left tarsometatarsus. It is

near the size of Podiceps nigricollis. Murray (1967) also tentatively

refers material from the Hagerman local fauna, the San Diego Formation,










California, and the Curtis Ranch, San Pedro Valley, Arizona, to this

species. Aechmophorus elasson, Murray, from the Blancan Hagerman local

fauna, was described on the distal end of an humerus and an associated

left ulna. It is similar to the living A. occidentalis. Podilymbus

majusculus Murray, also from the Hagerman local fauna, is based on a

nearly complete tarsometatarsus. It is larger than Podilymbus podiceps.

He also tentatively refers material from the Rexroad and Saw Rock Canyon

local faunas to this species.

Pleistocene species include Podiceps parvus (Shufeldt), based on a

lectotype right tarsometatarsus selected by Wetmore (1937) from the

Fossil Lake local fauna, Oregon. It is similar to the living P.

grisegena but is appreciably smaller (Howard, 1946). It is also know

from a well-core in the Tulane Formation of Kern County, California

(Wetmore, 1937). Podiceps dixi Brodkorb is known only from the proximal

end of a right carpometacarpus from Reddick, Florida. It was named after

the Dixie Lime Products Company which owned the quarry in which it was

found (Brodkorb, pers. comm., 1984; etymology omitted in Brodkorb,

1963e). It resembles the living Podiceps auritus, but is somewhat

larger (Brodkorb, 1963e). Podilymbus wetmorei Storer is based on a type

left tarsometatarsus, also from Reddick, Florida, and from a referred

tarsometatarsus and two femora from the Itchtucknee River, Florida. This

species is diagnosed as being the size of Podilymbus podiceps but more

robust. It is only known from these four elements.

The distribution of fossil grebes from the late Miocene through the

early Pliocene of Florida is shown in Table 4.1. Either Podilymbus cf.

P. podiceps or Podiceps sp. from Bone Valley could possibly be

conspecific with Pliodytes lanquisti. These species are only known from










a few specimens, none of which are directly comparable. Additional

material will eventually determine the validity of these assignments.

The occurrence of Tachybaptus in Florida is not surprising

considering the present range of Tachybaptus dominicus throughout the

Caribbean. Storer (1976a:124) suggests that T. dominicus has long been

separated from its Old World relatives (T. ruficollis subgroup).

Supporting this view is the lack of an extra canal in the hypotarsus of

T. dominicus (present in T. ruficollis subgroup). Storer (1976a)

considers this a derived character of T. dominicus.

The absence of small grebes the size of Tachybaptus from the Bone

Valley is probably due to a sampling bias toward large specimens or

possibly a general rarity of grebes due to the ecology of the area during

the deposition of the Bone Valley Formation.

The presence of Rollandia in the late Miocene of Florida suggests

that this genus, like Tachybaptus, had a far greater range in the past.

None of the fossil material of this family now known gives any

indication of the higher level systematic relationships of this group.

Postulated relationships include the Gaviidae, Hesperornithiformes,

Sphenisciformes (Cracraft, 1982), based on primarily foot-propelled

swimming adaptations; and the Rhinochetidae and Eurypygidae, on an

apparently unique configuration of the M. longus colli (Zusi and Storer,

1969).

Considering the distribution and diversity of the living species of

grebes (cf. Storer, 1963) it is probable that this group originated in

either North or South America. Supporting this view is the presence of

two genera unique to the Americas (Aechomorphus and Podilymbus) and the

diversity of the fossil record (ten out of eleven fossil species occur in










North America). This view is further strengthened by the absence of

grebes in the Early Tertiary European fossil localities which have

otherwise produced a rich aquatic avifauna (St.-Ge'rand-le-Puy; Cheneval,

1984; Phosphorites du Quercy; Mourer-Chauvir4, 1982). The paucity of

knowledge about the evolution of birds in the early Tertiary of South

America makes it premature to decide between a North or South American

origin, although this did not prevent Storer (1967) from suggesting a

South American origin of the family based solely on the diversity of

living species.










Table 4.1. Checklist of non-marine avian taxa discussed in the text.
Localities where each taxon occurs are given in parentheses -- Love Bone
Bed (LOV), McGehee Farm (MCG), Mixson Bone Bed (MIX), Bone Valley (BV),
Withlacoochee River hA (WITH 4A), Manatee County Dam (MD), SR-64, Haile
VB (H5B), Haile VI (H6), and Haile XIXA (H19A).

Class Ayes
Order Podicipediformes

Family Podicipedidae
Rollandia sp. (LOV, MIX, MCG)
Tachybaptus sp. (LOV, MCG)
Podilymbus cf P. podiceps (BV)
Podilymbus sp. A (MIX)
Podiceps sp. (BV)
Pliodytes lanquisti (BV)

Order Pelecaniformes

Family Phalacrocoracidae
Phalacrocorax sp. A (LOV, MCG, H19A)
Phalacrocorax wetmorei (BV, MD, SR-64)
Phalacrocorax cf. idahensis (BV)

Family Anhingidae
Anhinga grandis (LOV, MCG, H19A)
Anhinga sp. (BV)

Order Ciconiiformes

Family Ardeidae
Ardea polkensis (BV)
Areda sp. indet.(LOV)
Egretta sp. indet. (LOV, BV)
Egretta subfluvia (WITH hA)
Ardeola sp. (LOV)
Nycticorax fidens (MCG)

Family Ciconiidae
Mycteria sp. (LOV,MCG)
Ciconia sp. A (LOV)
Ciconia sp. B (MIX,BV)
cf. Ciconia sp. C (BV)


Family Plataleidae
Eudocimns sp. A. (BV)
Plegadis cf. P. pharangites (LOV)
Threskiornithinae, genus et species indet. (LOV)

Order Falconiformes (auct.)

Family Vulturidae
Pliogyps undescribed sp. (LOV)











Table 4.1--continued.

Family Pandionidae
Pandion lovensis (LOV)
Pandion sp. (BV)

Family Accipitridae
Haliaeetus (?) sp. (BV)
Buteo near B. Jamaciensis (WITH 4A)
Aquila sp. rBV
Accipitrid, genus indet. sp. A (LOV)
Accipitrid, genus indet. sp. B (BV)
Accipitrid, genus indet. sp. C (LOV)
Accipitrid, genus indet. (WITH 4A, BV)

Order Anseriformes

Family Anatidae
Dendrocygna sp. (LOV)
Branta sp. A (LOV)
Anserinae, genus indet. sp. B (LOV)
Anserinae, genus indet. sp. C (LOV, BV)
Anserinae, genus indet. sp. D (BV)
Tadorini, genus indet. sp. A (BV)
Anas undescribed sp. A (LOV, MCG)
Anas size near A. acuta (LOV, MCG)
Anatini, genus indet. sp. A (LOV)
Anatini, genus indet. sp. B (LOV)
Aythya sp. A (BV)
Oxyura cf. 0. dominica (BV)

Order Galliformes

Family Phasianidae
Meleagridinae, genus indet. (LOV)
Meleagris sp. (BV)


Order Gruiformes (auct.)

Family Gruidae
Grus sp. A (LOV)
Grus sp. B (LOV)
Balearicinae, genus indet. (BV)
Aramornis (cf.) (LOV)










Table 4.1--continued.


Family Rallidae
Rallus sp. A (LOV)
Rallus sp. B (BV)
Rallus (cf.) sp. C (LOV)
Undescribed genus (LOV, MCG)

Order Charadriiformes

Family Phoenicopteridae
Phoenicopterus floridanus (BV)
Phoenicopterus sp. A (LOV, MCG)

Family Jacanidae
Jacana farrandi (LOV, MCG)

Family Scolopacidae
Limosa ossivallis (BV)
Erolia penepusilla (BV)
Ereunetes rayi (MCG)
Calidris pacis (BV)
"Calidris" sp. indet. 1 (LOV, MCG, BV)
"Calidris" sp. indet. 2 (LOV)
"Calidris" sp. indet. 3 (MCG)
"Calidris" sp. indet. 4 (LOV)
??Actitis sp. indet. 5 (LOV)
??Arenaria sp. indet. 6 (LOV)
Genus indet. sp. indet. 7 (LOV)
Genus indet. sp. indet. 8 (LOV)
?Philomachus sp. (BV)


Order Strigiformes

Family Tytonidae
Undescribed genus (LOV)

Family Strigidae
Bubo sp. (BV)

Order Passeriformes

Suborder indet. sp. A (LOV)
Suborder indet. sp. B (LOV)

Family Fringillidae
Palaeostruthus eurius (H 6)









Table 4.2. Measurements of humeri and coracoids of the grebes Rollandia
rolland chiliensis (N = 6, 2 males, 1 female, 3 unsexed), Tachybaptus
dominicus (N = 7, 4 males, 3 females), and Rollandia sp. from McGehee
Farm local fauna. Data are mean + standard deviation and range. (*)
specimen damaged. Abbreviations defined in methods section.


Measurement


Humerus
W-SHAFT


D-SHAFT


W-DIST


Coracoid
HEAD-FAC


HEAD-IDA


HEAD-CS


D-HEAD


W-SHAFT


D-SHAFT


FAC-IDA


L-GLEN


R. r. chilensis


2.67 + 0.15
2.5 2.8

2.47 + 0.12
2.3-- 2.6

5.35 + 0.23
5.1 5.7


25.45 + 0.71
24.3 26.2

24.50 + 0.68
23.3 25.3

7.03 + 0.23
6.7f-- 7.4

2.17 + 0.10
2.0 2.3

2.30 + 0.18
2.1 2.6

1.65 + 0.10
1.5 1.8

8.53 + 0.38
8.2 9.1

4.55 + 0.16
4.4 5.8


T. dominicus


2.39 + 0.15
2.2 2.6

2.11 + 0.15
1.9 2.3

5.00 + 0.31
4.6-- 5.5


21.80 + 1.57
20.1 24.2

21.17 + 1.26
19.4 23.2

6.10 + 0.34
5.6-- 6.6

2.17 + 0.17
2.0 2.4

1.94 + 0.21
1.7 2.3

1.29 + 0.09
1.1 1.4

7.44 + 0.41
7.1 8.3

4.31 + 0.25
3.8 4.5


Rollandia sp.


2.8


2.6


5.9



*24.6


*23.3


7.6


2.1


5.3









Table 4.3. Measurements of the tarsometatarsi of the grebes Rollandia
rolland chilensis (N = 6, 2 males, 1 female, 3 unsexed), Podilymbus
podiceps (N = 14, 7 males, 7 females), Rollandia sp., and Podilymbus
sp. A. from the Mixson's Bone Bed. Data are mean + standard deviation


and range.


Abbreviations defined in the methods section.


Measurements

Tarsometatarsus
LENGTH


W-PROX


W-HYPOTS


TRIII-TRIV


TRII-TRIV


W-TRII


W-TRIII


D-TRIII


R. r. chilensis


35.58 + 1.30
33.2 36.7


7.00 + 0.36
6.4 7.5

3.65 + 0.20
3.3 3.8

5.20 + 0.19
5.0 -- 5.4

5.22 + 0.12
5.-1 5.4

1.55 + 0.19
1.3 1.8

2.52 + 0.15
2.3 2.7

3.78 + 0.21
3.4 3.9


P. podiceps


40.15 + 2.36
36.3 44.9

8.09 + 0.57
7.17- 8.9

4.61 + 0.28
4.1 5.1

6.57 + 0.49
5.87- 7.2

6.24 + 0.49
5.3 7.2

1.96 + 0.24
1.6 2.4

2.76 + 0.54
1.5 3.3

5.00 + 0.41
4.5 5.6


R. sp.


36.4


7.2;
7.5

4.0;
4.0


6.7


3.5


5.6


5.2


2.6


4.1









Table 4.4. Measurements of the femora of the grebes Rollandia rolland
chilensis (N = 6, 2 males, 1 female, 3 unsexed), Tachybaptus dominicus
(N = 7, 4 males, 3 females), and Tachybaptus sp. from the Love Bone Bed.
Data are mean + standard deviation and range. (*) Specimen
damaged. Abbreviations defined in the methods section.


Measurements


Femora
M-LENGTH


L-LENGTH


W-SHAFT


D-SHAFT


W-PROX


D-HEAD


W-DIST


W-M&LCON


D-LCON


D-MCON


R. r. chilensis


30.73 + 1.55
27.9 -32.0

32.73 + 1.64
29.7 33.5

2.80 + 0.15
2.6-- 3.0

3.18 + 0.24
2.8 3.4

7.65 + 0.39
7.1 8.3

3.30 + 0.24
3.0 3.7

8.08 + 0.44
7.5 8.4

5.97 + 0.41
5.4-- 6.4

6.03 + 0.24
5.7 6.4

4.40 + 0.24
4.1 4.7


T. dominicus


25.03 + 1.53
23.5 27.8

26.81 + 1.61
25.1 29.7

2.53 + 0.22
2.3 2.9

2.67 + 0.30
2.3 3.0

6.53 + 0.30
6.2 6.9

2.74 + 0.20
2.4 3.0

6.84 + 0.53
5.9 7.4

4.96 + 0.40
4.4-- 5.5

4.97 + 0.37
4.3 5.5

3.44 + 0.26
3.1 3.8


Tachybaptus sp.


26.1


27.5


2.6


2.6


6.8


2.8


*6.0


*5.1


3.5










Table 4.5. Measurements of the tibiotarsi of the grebes Podilymbus
podiceps (N = 14, 7 males, 7 females) and Podilymbus cf. P. podiceps from
the Bone Valley Mining District. Data are mean + standard deviation and
range. (*) Specimen damaged. Abbreviations are defined in the methods
section.


Measurements

Tibiotarsus
LENGTH


M-LENGTH


W-SHAFT


D-SHAFT


W-PROX-M


W-DIST-CR


W-DIST-CD


D-MCON


D-LCON


D-ICON


Podilymbus podiceps


68.69 + 4.31
61.7 77.0

80.17 + 5.27
72.4 90.2

4.50 + 0.41
3.8 5.3

3.29 + 0.27
2.8 3.8

7.00 + 0.44
6.5 7.9

7.10 + 0.65
6.1 7.9

6.04 + 0.37
5.2 6.5

6.86 + 0.44
6.1 7.5

6.76 + 0.49
5.9 7.4

4.29 + 0.33
3.7 4.9


P. cf. podiceps


4.6; 4.9


3.5; 3.7


7.2; *7.4


*6.2; *6.5


*6.9; 7.4


7.0; 7.3


4.4; 4.9









Order Pelecaniformes Sharpe,1891

Family Phalacrocoracidae (Bonaparte, 1853)

Genus Phalacrocorax Brisson, 1760

Remarks. The following morphological descriptions are based on the

comparisons of a sample of 5 males and 5 females each of Phalacrocorax

auritus auritus and P. auritus floridanus, and' all available fossil

material.


Phalacrocorax sp. A.

Material. Love Bone Bed local fauna; UF 25735, UF 29661, distal

ends left humeri; UF 29662, distal end right ulna; UF 25877, distal end

left tibiotarsus; UF 25861, distal end right tarsometatarsus (badly

worn, tentatively referred); UF 25933, distal end left tarsometatarsus.

McGehee Farm local fauna; UF 11569, complete left coracoid; UF

31779, sternal end left coracoid; UF 12351, distal end right humerus; UF

4107, proximal end right ulna; UF 9492, proximal end right ulna

(questionably referred); UF 31778, proximal end right carpometacarpus; UF

11105, distal end left carpometacarpus; UF 29746, complete left

tarsometatarsus; UF 31777, proximal end right tarsometatarsus. PB 7964,

proximal end left carpometacarpus.

Haile XIXA; UF 29774, proximal end left humerus; UF 47340, proximal

end right carpometacarpus.

Description. Coracoids from McGehee Farm differ from those of both

subspecies of Phalacrocorax auritus (auritus and floridanus) examined,

and from P. wetmorei by having a more elliptical facies articularis

clavicularis, a more robust shaft in relation to the length of coracoid,

the brachial tuberosity more undercut, the impression for the attachment











of the coraco-brachialis more distinct, and in medial view, the shaft

more rotated ventrally.

Distal end of the humeri of the fossil species is generally smaller

than that of females of P. a. floridanus, and the shaft more slender

(much smaller than P. a. auritus). Other characters are within the range

of variation of P. auritus. Differs from the humeri of P. wetmorei by

being smaller, having a more shallow fossa brachialis of a different

angle and a much wider attachment of the anterior articular ligament

(=--tuberculum supracondylare ventrale).

The two ulnae that are sufficiently perserved to make comparisons

(UF 29662, UF 4107) appear small, about the size of small females of P.

a. floridanus. Characters are within the range of variation of this

species.

Carpometacarpus larger than that of the largest male P. a. auritus.

Process of metacarpal I more nearly square than in that of P. auritus.

Shaft of metacarpal II more robust and angular; anterior carpal facet not

extending up the carpal trochlea as in P. auritus. Pollical facet with a

small papilla .

Tibiotarsus indistinguishable from that of small females of P. a.

floridanus.

Tarsometatarsus description based on UF 29746 (UF 25933 broken and

badly worn; UF 31777 missing distal half, but both agree with UF 29746 in

all discernable characters). Tarsometatarsus short, about equal to that

of females of P. a. floridanus. Transverse width of shaft very narrow.

Lateral face of shaft much more flattened than P. auritus, causing the

posterior intermuscular line to be on the lateral edge of the shaft.

Proximal end narrow, lateral calcaneal ridge more narrow and elongate










than in P. auritus. Posterior opening of the lateral proximal vascular

foramen is located lateral to the ridge extending down from the lateral

calcaneal ridge. This ridge not extending as far down the shaft as in P.

auritus.

Remarks. If the carpometacarpus above is correctly assigned to the

same species as is represented by the other skeletal elements, then this

cormorant is quite different in proportions than Phalacrocorax auritus

and related species such as Phalacrocorax wetmorei.


Phalacrocorax wetmorei Brodkorb, 1955

Material. This material is very well represented in the Bone Valley

Fauna. Only Florida State Museum and Florida State Geological specimens

are included in the following referred material section. Material

accessioned into the Florida State Museum collections after 26 April 1984

has not been included in the list of referred specimens; material

accessioned after 5 March 1984 has not been included in the tables of

measurements. Additional material from Bone Valley (type material) is

listed in Brodkorb (1955a).

Manatee County Dam Site.--UF 11916, distal end right humerus.

SR-64 local fauna.--UF 67805, complete left coracoid; UF 64143,

humeral end left coracoid; UF 64144, humeral end right coracoid; UF

64146, humeral end right scapula; UF 64145, partial sternum; UF 64147,

proximal end right tibiotarsus; UF 64148, UF 64149, distal ends left

tarsometatarsi.

Bone Valley Mining District, Brewster Mine.--UF 61987, humeral end

right coracoid; UF 61988, distal end right tarsometatarsus; UF 65691,

proximal end left ulna.









Bone Valley Mining District, Chicora Mine.--UF 29733, humeral end

left coracoid.

Bone Valley Mining District, Fort Green Mine.--UF 61958, associated

(?) partial skeleton; UF 58062, right quadrate; UF 60047, caudal portion

left mandible; UF 53912, caudal portion right mandible; UF 57248, sternal

fragment with coracoidal sulci; UF 52415, UF 53938, proximal ends right

scapulae; UF 53873, proximal end left scapula; UF 62025, complete left

coracoid; UF 52413, UF 57332, UF 55838, UF 57246, UF 58058, UF 61960, UF

65711, humeral ends right coracoids; UF 53913, UF 55872, UF 57331, UF

58059, UF 58339, UF 61961, UF 65655, humeral ends left coracoids; UF

52414, UF 61962, UF 65656, sternal ends right coracoids; UF 53934, UF

55831, UF 55875, UF 61963, sternal ends left coracoids; UF 55810, UF

55811, UF 58378, UF 61964, proximal end right humeri; UF 58304, UF 60048,

proximal ends left humeri; UF 53937, shaft left humerus; UF 52410, UF

53914, UF 60050, UF 65657, distal ends right humeri; UF 55865, UF 58060,

UF 58338, UF 58419, UF 58420, UF 60049, distal ends left humeri; UF

55867, UF 57242, UF 58061, UF 60051, UF 65712, proximal ends right ulnae;

UF 55866, UF 61965, proximal ends left ulnae; UF 57243, UF 57247, UF

57249, UF 60052, UF 61966, distal ends right ulnae; UF 52411, UF 55812,

UF 55871, UF 57334, UF 57335, UF 58340, distal ends left ulnae; UF 55813,

UF 55832, UF 55869, UF 60053, UF 65661, proximal ends right

carpometacarpi; UF 52412, UF 55833, UF 55834, UF 57333, UF 58418, UF

61967, UF 61968, UF 61969, UF 58407, distal ends right carpometacarpi; UF

61959, partial synsacrum; UF 55814, UF 60054, complete right femora; UF

57336, complete left femur; UF 57244, proximal end right femur; UF 53872,

UF 53872, UF 55873, UF 55874, UF 57337, UF 60055, UF 61970, UF 55835,

distal ends right femora; UF 57391, UF 61971, distal ends left femora; UF









55868, proximal end right tibiotarsus; UF 55836, UF 57245, UF 60056,

proximal end left tibiotarsus; UF 55804, UF 57250, distal ends right

tibiotarsi; UF 53935, UF 55863, UF 55864, UF 57357, UF 58305, UF 60057,

UF 60058, UF 65713, distal ends left tibiotarsi; UF 55860, nearly

complete left tarsometatarsus; UF 52416, UF 52417, UF 52418, UF 55837, UF

58067, proximal end right tarsometatarsi; UF 55870, UF 58421, UF 61972,

proximal ends left tarsometatarsi; UF 53889, UF 53936, UF 55815, UF

57251, UF 58306, UF 58341, distal ends right tarsometatarsi; UF 58342, UF

60059, UF 65658, distal end left tarsometatarsi.

Bone Valley Mining District, Gardiner Mine.--UF 58438, caudal

portion right mandible; UF 61998, right clavicle; UF 61999, proximal end

right scapula; UF 62000, proximal end left scapula, UF 65667, complete

left coracoid; UF 58278, UF 58279, UF 58439, UF 58440, UF 62001, UF

65669, humeral ends right coracoids; UF 58277, UF 58280, UF 58446, UF

58447, UF 58469, UF 62002, UF 62003, UF 62004, UF 65668, humeral ends

left coracoids; UF 58448, sternal end left coracoid; UF 58470, UF 62005,

proximal ends right humeri; UF 58441, UF 58471, distal ends right humeri;

UF 58449, UF 58472, UF 62006, distal ends left humeri; UF 58281, UF

58442, UF 62007, UF 65670, proximal end right ulnae; UF 62008; proximal

end left ulna; UF 57307, UF 58285, UF 58286, UF 65671, distal end right

ulnae; UF 58282, UF 58283, UF 58284, UF 65672, UF 65749, distal end left

ulnae; UF 58443, proximal end right carpometacarpus; UF 58287, proximal

end left carpometacarpus; UF 58303, UF 58282, distal ends left

carpometecarpi; UF 58450, complete left femur; UF 58451, proximal end

left femur; UF 58473, distal end right femur; UF 57311, proximal end left

tibiotarsus; UF 58290, UF 58444, UF 58445, UF 58474, UF 58475, distal

ends right tibiotarsi; UF 58289, UF 58452, UF 58453, UF 62009, UF 62010,









UF 62011, UF 62012, distal ends left tibiotarsi; UF 65673, complete right

tarsometarsus; UF 58478, UF 62013, proximal end right tarsometatarsi; UF

58291, UF 58292, proximal end left tarsometarsi; UF 58293, UF 58476, UF

58477, UF 65674, distal ends right tarsometatarsi; UF 57310, distal end

left tarsometatarsus.

Bone Valley Mining District, Palmetto Mine.--UF 21058, sternal

fragment with coracoidal sulci; UF 13225, humeral end left coracoid; UF

21143, shaft left coracoid; UF 21115, shaft right coracoid; UF 21146,

distal end right humerus; UF 29734, distal end left humerus; UF 13231, UF

29740, distal ends right ulnae; UF 21091, UF 21120, proximal ends right

carpometacarpi, UF 21068, UF 21072, proximal ends left carpometacarpi; UF

21131, distal end right carpometacarpus; UF 49090, complete right femur;

UF 12352, complete left femur; UF 29735, UF 49091, proximal ends left

tarsometatarsi; UF 12868, distal end right tarsometatarsus.

Bone Valley Mining District, New Palmetto Mine.-UF 49691, UF 49692,

vertebrae (questionally referred), UF 49693, complete right femur; UF

49694, complete left tarsometatarsus.

Bone Valley Mining District, North Palmetto Mine.--UF 49097, humeral

end left coracoid; UF 49098, humeral end right coracoid.

Bone Valley Mining District, Southwest Palmetto Mine.-UF 49093,

humeral end right coracoid.

Bone Valley Mining District, Hookers Prairie Mine.-UF 49690,

complete right femur.

Bone Valley Mining District, Kingsford Mine.--UF 21186, humeral end

left coracoid; UF 52971, distal end right humerus; UF 13212, distal end

left humerus; UF 21185, proximal end right tibiotarsus.









Bone Valley Mining District, Payne Creek Mine.--UF 29741, UF 57304,

humeral ends right coracoids; UF 21203, proximal end left

carpometacarpus; UF 29742, distal end right carpometacarpus.

Bone Valley Mining District, Swift Mine.-UF 55883, distal end right

tibiotarsus; UF 17687, distal end right tatsometatarsus.

Bone Valley Mining District, specific locality unknown.--UF 61549,

caudal portion left mandible; UF 61550, nearly complete right coracoid;

UF 61553, nearly complete left coracoid; UF 61554, UF 61555, humeral ends

left coracoids; UF 61551, humeral end right coracoid; UF 61552, sternal

end right coracoid; UF 61556, sternal end left coracoid; UF 61557, UF

61558, proximal end right humerus; UF 61559, UF 61560, proximal end left

humeri; UF 61561, distal end right humerus; UF 61562, UF 61563, UF 61564,

UF 61565, UF 61566, UF 61567, UF 61568, UF 61569, distal end left humeri;

UF 61570, UF 61571, proximal end left ulnae; UF 61572, UF 61573, UF

61574, distal ends right ulnae; UF 61575, distal end left ulna; UF 61578,

nearly complete left carpometacarpus; UF 61576, UF 61577, proximal ends

right carpometacarpi; UF 61579, proximal end left carpometacarpus; UF

61596, complete left femur; UF 61580, proximal end left tibiotarsus; UF

61581, UF 61582, UF 61583, UF 61584, distal ends right tibiotarsi; UF

61585, UF 61586, UF 61587, distal ends left tibiotarsi; UF 61588, nearly

complete left tarsometatarsus; UF 61589, UF 61590, UF 61591, UF 61592,

proximal end left tarsometatarsus; UF 61593, distal end right

tarsometatarsus; UF 61594, UF 61595, distal ends left tarsometatarsi.

Bone Valley Mining District, specific locality unknown (FGS

collection).--V 7311, proximal end left humers; V 7313, distal end left

humerus; V 7309, distal end left ulna; V 7310, proximal end left

carpometacarpus; V 7312, distal end left femur.










Description. Scapula within range of variation of that of

Phalacrocorax auritus.

Coracoids appear to be well within the range of variation of those of

P. auritus. The characters used by Brodkorb (1955a: 12) "anterior

intermuscular line situated farther laterad" applies only at the extreme

sternal end of the coracoid. This line does not swing media; instead it

curves little as it extends down the shaft. Differs from UF 11569 from

McGehee by characters cited above. Coracoids from SR-64 are

indistinguishable from those of Phalacrocorax wetmorei from Bone Valley.

The two characters of the humerus used by Brodkorb (1955a:12) "the

head of humerus shallower" and "condyles averaging less deep" do not hold

when a large series of specimens are measured (Table 4.7). Brodkorb's

statements that the pneumatic fossa is narrow (slightly) and deeper are

supported. This is especially apparent by having a small, but deep fossa

paralleling the crus ventrale fossae. In P. wetmorei the pneumatic fossa

is perforated by several pneumatic foramina but it is rarely perforated

in P. auritus. In the few specimens of P. auritus in which these

foramina are present, they are very minute. Ligamental furrow (=

ligamental sulcus) does not appear to be relatively longer when compared

against a series of both sexes and subspecies of P. auritus. The distal

end of the humerus of P. wetmorei tends to be narrower, with a more

elongated attachment for the anterior articular ligament (= tuberculum

supracondylare ventrale) ending proximally in a narrower crest than in

the modern specimens of P. auritus. Measurements of ulnae are given in

Table 4.7.

Carpometacarpi of P. wetmorei are about as robust as those of

females of P. f. floridanus. The process of metacarpal I is slightly










more produced. Fovea carpalis caudalis deeper in P. wetmorei than in P.

auritus.

The femora are similar, except that the popliteal fossa is generally

less excavated than in P. auritus. Brodkorb's statement that the femur

is longer and narrower than that of P. auritus is not supported by the

larger sample size now available.

Tibiotarsus with no obvious qualitative morphological differences,

but see Table 4.10 for a few minor quantitative differences.

Tarsometatarsus with a lateral face flat, similar to but not as

extreme as, that found on specimens from the Love Bone Bed local fauna.

Other characters similar. See Table 4.11 for measurements.

Remarks. See comments under Family Remarks (below) pertaining to P.

auritus and related species.


Phalacrocorax cf. P. idahensis

Material. Bone Valley Mining District, Palmetto Mine (locality 2 of

Brodkorb, 1955); PB 311, proximal end left ulna.

Remarks. This proximal ulna (PB 311) is larger than that of other

specimens of P. wetmorei presently known from Bone Valley. Since it was

first reported by Brodkorb (1955), it has been additionally damaged in

transport and is now barely diagnostic at the generic level. Unless

additional material becomes available, the status of this enigmatic

record in Florida will probably never be satisfactorily resolved.


Remarks on the Family Phalacrocoracidae.

The cormorants have an extensive fossil record. Brodkorb (1963c)

lists 23 paleospecies; subsequently eight more have been described, or

are in the process of being described. Most have been referred to the










genus Phalacrocorax (presently with 26 paleospecies), many without

extensive comparisons with other recent and fossil species to determine

the variability of the of the characters used. It would be desirable to

revise the paleospecies of Phalacrocorax and integrate this extensive

fossil record with the recent species to produce a phylogeny of the

family. There are large amounts of fossil material available, permitting

analysis of variation of several fossil species (e.g. P. wetmorei, P.

oweri, etc.), a recent descriptive osteological study (Ono, 1980), and

papers identifying osteological characteristics and proportions of the

various subgenera of cormorants (Howard, 1932a, 1965; Brodkorb and

Mourer-Chauvirb, 1984). However, such a revision is beyond the scope of

this dissertation.

Species listed by Brodkorb (1963c) that are not cormorants include

all species of the genus Graculavus (moved to Charadriiformes, Olson and

Parris, ms), Actiornis anglicus (not a cormorant, nor ibis, Olson,

1981b), and Phalacrocorax mediterraneus (Gruiformes, Family

Bathornithidae = Paracrax antiqua, Cracraft, 1971).

The earliest cormorants appear in the early Miocene. Phalacrocorax

subvolans Brodkorb from the mid-Hemingfordian Thomas Farm local fauna,

Florida, is known only from a proximal humerus. It is currently under

study (Becker, in prep.). Phalacrocorax marinavis Shufeldt from the

Arikareean John Day Formation, Oregon, is known from a humerus, ulna,

tarsometatarsus, and part of a femur. It is somewhat smaller than P.

carbo but is reported to be allied with this species. Phalacrocorax

miocaenus (Milne-Edwards) from the Aquitanian of Langy, Vaumas, St.-
I
Gerand-le-Puy, and Montaigu, France, is known from most skeletal

elements. It was moved to a new genus Nectornis and is said to share










characters with Anhinga (Cheneval, 1984). Phalacrocorax littoralis

(Milne-Edwards) from the Aquitanian of St.-Gerand-le-Puy, France, and

from Germany was based on a coracoid and a few other skeletal elements.

It seems to be related to P. aristotelis. Phalacrocorax anatolicus

Mourer-Chauvire' from the lower or middle Miocene (probably Helvetian) of

Bes-Konak, Turkey, was described from a coracoid and most of a forelimb.

It appears to be related to the fossil species P. littoralis, P.

miocaenus, and to the recent P. aristotelis.

Phalacrocorax leptopus Brodkorb from the Clarendonian and

Hemphillian localities of Juntura, Oregon, is based on a coracoid,

tarsometatarsus, and scapula. It is a small species and resembles the

fossil P. littoralis. It is in need of additional comparisons to

elucidate its relationships.

Phalacrocorax femoralis L. Miller from the Barstovian or

Clarendonian Calabasas local fauna, California, is based on most of a

skeleton, preserved in a slab of fine-grained shale. It is the size of

PL. penicillatus, but Miller (1929) asserts that this species does not

appear closely related to any living species. Phalacrocorax lautus

Kurochkin and Ganya from the upper Miocene of Moldavia, is based on the

proximal half of a right femur. It appears closest to the living P.

pygmaceus.

Phalacrocorax praecarbo Ammon was described from the upper Miocene

Brown Coal Formation, near WUrttemburg, Germany, on the humeral end of a

coracoid. Brodkorb (1980) has moved Ardea brunhuberi Ammon (figured in

Ammon, 1911), based on a proximal end of a carpometacarpus to this

species and emended the name to P. brunhuberi (Ammon, 1918; cited as 1911

in Brodkorb, 1980). Olson (ms) also moves Botaurites auritus Ammon,










based on a cervical vertebra to P. brunhuberi. Phalacrocorax intermedius

(Milne-Edwards) from the Oreleanian of Orleanais, France was described

from a proximal end of a humerus. Phalacrocorax brunhuberi may be

synonymous with P. intermedius; only slightly smaller size and slightly

younger age prevented Brodkorb (1980) from placing it in synonyqr with

this species.

Phalacrocorax ibericum Villalta, probably from the Lower Pontian of

Spain, is based on the distal end of a humerus. Villalta states that

this cormorant is smaller than the other Aquitanian cormorants of Europe

(P. littoralis, P. miocenaeus, and P. intermedius) and is close to the

living P. carbo.

Phalacrocorax goletensis Howard from the late Hemphillian or early

Blancan La Goletia local fauna, Michocan, Mexico, is known from a

coracoid (type) and a referred distal humerus. It is possibly ancestral

to P. olivaceus.

Five species, all said to be ancestral to P. auritus, have been

described from the late Hemphillian to Mid-Pleistocene of North America.

Phalacrocorax wetmorei Brodkorb, described from the late Hemphillian Bone

Valley District, Florida, is known from all major skeletal elements. See

additional remarks above. Phalacrocorax kennelli Howard from the Blancan

San Diego Formation, was described on a partial coracoid, a humerus, a

furculum, and vertebrae. It agrees in size with P. pelagicus and P.

penicillatus. In morphology, the fossil species resembles P. auritus or

P. pelagicus (Howard, 1949). Phalacrocorax idahensis (Marsh) from the

Hemphillian Castle Creek, Idaho, is based on a proximal carpometacarpus.

Murray (1970) referred additional material to and redescribed this

species from the (Blancan) Hagerman local fauna. Phalacrocorax macer










Brodkorb from the (Blancan) Hagerman local fauna, Idaho, was originally

described on a carpometacarpus. Murray (1970) also redescribed this

species based on additional material. Phalacrocorax macropus (Cope) from

the Mid-Pleistocene Fossil Lake local fauna, Oregon, was described on a

tarsometatarsus. Howard (1946) referred many other specimens to this

species.

I do not believe that all these species are valid and are correctly

assigned to the ancestral lineage of P. auritus. The morphological

differences among the fossil species are comparable to those among

subspecies of modern P. auritus. It is very possible that like modern

cormorants that show geographic size variations (Palmer, 1962), the

fossil species are simply conspecific geographical variants. If this can

be demonstrated, then each of these fossil species should be maintained

as subspecies of the senior synonym, Phalacrocorax macropus (Marsh).

Valenticarbo praetermissus Harrison from the late Pliocene to early

Pleistocene of Siwalks, India, is based on a 100-year-old plaster cast of

a proximal end of a tarsometatarsus, lacking the hypotarsus. It is very

doubtful that this genus is valid (Olson, ms.). I am unaware of the

relationship of this supposed species.

Pliocarbo longipes Tugarinov from the early Pliocene of the Ukraine

was described from a worn tarsometatarsus and a referred femur. Olson

(ms) notes that although the size and proportions of the tarsometatarsus

are different from typical cormorants, the illustrations are too poor for

even a positive familial verification.

Phalacrocorax destenfanii Regalia from the Mid-Pliocene (Ruscinian?)

of Orciano, Pisano, Italy, was described from most major skeletal

elements. Phalacrocorax mongoliensis Kurochkin from the upper Pliocene










of Mongolia is based on the distal epiphysis of a left femur.

Phalacrocorax reliquus Kurochkin from the middle Pliocene of western

Mongolia, is based on the distal epyphysis of a right humerus. It has

the same dimensions as P. pelagicus. The relationships of these

cormorants have not been determined.

Phalacrocorax rogersi Howard from the Pliocene Veronica Springs

Quarry, California, is known only from the type coracoid. It is a large

species and appears close to P. perspicillatus and P_. pelagicus.

Phalacrocorax owrei Brodkorb and Mourer-Chauvird, from the lower

Pleistocene of Olduvai Gorge, Tanzania, is known from nearly all skeletal

elements. It has been assigned to the subgenus Stictocarbo, although its

tarsometatarsus is rather similar to P. fuscicollis (subgenus

Phalacrocorax). Phalacrocorax tanzaniae Harrison and Walker from the

Pleistocene Bed II of Olduvai Gorge, Tanzania, was described from a

tarsometatarsus and appears close to P. carbo. Phalacrocorax pampeanus

Moreno and Mercerat from the upper Pleistocene Lujan local fauna,

Argentina, was described from the proximal end of a humerus. It is very

close to P. olivaceus and may be ancestral to this recent species

(Howard, 1965). Phalacrocorax gregorii and P. vetustus DeVis from the

upper Pleistocene localities near Lake Eyre, South Australia, were

described from many elements.














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Table 4.9. Measurements of the femora of the cormorants Phalacrocorax
auritus auritus (N = 14, 7 males, 7 females), Phalacrocorax auritus
floridanus (N = 18, 9 males, 9 females), and Phalacrocorax wetmorei,
from the Bone Valley Mining District. Data are mean + standard
deviation, (N), and range. Abbreviations are defined in the methods
section.


Measurements


Femur
M-LENGTH


L-LENGTH


W-SHAFT


D-SHAFT


W-PROX


D-HEAD


W-DIST


W-M&LCON


W-LCON


W-L&FCON


D-FCON


D-LCON


D-MCON


P. a. auritus


54.89 + 2.40
49.5 58.7

56.58 + 2.63
50.1 60.0

6.48 + 0.36
5.6 7.1

8.15 + 0.50
6.9 8.6

16.13 + 0.60
15.5 17.4

7.02 + 0.33
6.4 7.6

15.71 + 0.59
14.9 16.6

12.21 + 0.65
11.3 13.2

3.34 + 0.27
3.0 3.9

7.25 + 0.25
6.9 7.7

8.81 + 0.47
8.3 9.5

10.36 + 0.37
9.8 -11.1

9.05 + 0.34
8.4 9.7


P. a. floridanus


52.12 + 3.15
43.2 57.2

54.03 + 2.51
49.4 59.1

5.94 + 0.38
5.2 6.5

7.38 + 0.46
6.7 8.4

14.56 + 0.85
13.2 16.2

6.55 + 0.33
6.0 7.3

14.93 + 0.84
13.3 16.5

11.38 + 0.77
10.0 13.0

3.02 + 0.33
2.5 3.6


6.74
5.8


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- 7.4


8.40 + 0.50
7.5 9.2

9.78 + 0.57
8.7 10.5

8.59 + 0.48
7.6 9.4


P. wetmorei


55.46 + 1.58 (9)
53.3 59.2

57.58 + 1.45 (12)
55.7 61.0

6.49 + 0.29 (17)
5.9 7.0


8.04 + 0.41
7.3 -- 9.0


(17)


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14.5 16.7


6.76 + 0.28
6.3 7.2


(16)


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14.4-- 16.0

11.59 + 0.39 (14)
10.8 12.3


2.99 + 0.20
2.7 3.3

6.85 + 0.32
6.4 7.5

8.68 + 0.40
8.1 9.4


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9.6 10.8

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Family Anhingidae Ridgway, 1887

Remarks. Skeletal elements of anhingas discussed below may be

distinguished from the Phalacrocoracidae as follows: Humerus--by

characters given by Miller (1966) and Martin and Mengel (1975). Coracoid

--head rotated ventrad and media, to produce a distinct notch between

head and shaft, when observed from either a ventral or medial view (head

merges smoothly with shaft in the Phalacrocoracidae). Procoracid

expanded and concave cotyla scapularis present.


Genus Anhinga Brisson, 1760

Anhinga grandis Martin and Mengel, 1975

Material. Love Bone Bed local fauna; UF 25739, proximal end right

humerus; UF 25723, UF25725, distal ends right humeri; UF 26000, nearly

complete right coracoid; UF 25873, distal end right tibiotarsus.

McGehee Farm local fauna; UF 11107, distal end right humerus.

Remarks. Elements described in detail in Becker (in prep.). Distal

humeri compare exactly in all features with the type. Coracoid assigned

to this species, as it is of correct size.


Anhinga sp. unknown

Material. Bone Valley Mining District, no specific locality; UF

29781, proximal end left ulna; UF 29780, distal end left ulna.

Remarks. The ulnae are much larger than comparable elements of A.

anhinga and those expected for A. grandis. Parenthetically, the ulna

from Coleman III (UF 16664), referred to A. cf. grandis by Ritchie (1980),

is definitely not Anhinga grandis and is not identifiable to a species.

Along with the two specimens from Bone Valley, this Coleman III specimen

represents a much larger species of anhinga which existed approximately










4.5 million years later than did A. grandis. Unfortunately, this species

is only known from ulnae and not from more diagnostic elements.


Remarks on the Family Anhingidae.

The earliest record of the Anhingidae is Protoplotus beauforti

Lambrecht based on a skeletal impression, from the late Eocene of

Sumatra. It is presently being restudied and will probably be referred

to a new family (P. V. Rich--in litt., cited in Olson, ms).

Anhinga pannonica Lambrecht was described from a cervical vertebra

and carpometacarpus from the late Miocene of Tataros, Hungary. Rich

(1972) also assigned another cervical vertebra and a partial humerus from

the late Miocene of Tunisia to this species. The only other anhinga

known from the late Miocene, Anhinga grandis, originally described from

the Hemphillian Cambridge (Ft.- 40) locality, Nebraska, is discussed

above.

The validity of A. laticeps Devis from the late Pleistocenffe of

Australia is somewhat questionable (Brodkorb and Mourer-Chauvire, 1982;

Olson, ms.). Anhinga hadarensis Brodkorb and Mourer-Chauvire, 1982 from

the Upper Pliocene Kadar Hadar member of the Hadar Formation, Ethiopia is

also known from the Omo Basin, Ethiopia, and Olduvai Gorge, Tanzania. It

appears to be the immediate ancestor to A. rufa (Brodkorb and Mourer-

Chauvire, 1982). Two Pleistocene fossil species of anhingas have been

shown to be cormorants. Anhinga parva Devis from Australia was shown by

Miller (1966) to be the cormorant Phalacrocorax melanoleucos and A. nana

Newton and Gadow, from Mauritius was shown by Olson (1975a) to be another

cormorant, Phalacrocorax africanus.







Order Ciconiiformes Garrod, 1874 (Auct.)

Family Ardeidae Vigors, 1825.

Remarks. The following account briefly establishes the presence and

distribution of the late Miocene and early Pliocene herons in Florida for

the paleoecological and biochronological aspects of this study. More

detailed descriptions and systematic remarks may be found in Becker

(1985a). Systematic nomenclature follows Payne and Risley (1976).


Genus Ardea Linnaeus, 1758

Ardea polkensis Brodkorb, 1955

Material. Bone Valley Phosphate Mining District, Palmetto Mine; PB

380, proximal end of right tarsometatarsus (type), UF 21138, distal end

right tarsometatarsus; Payne Creek Mine; PB 7924, humeral end of right

coracoid.

Remarks. This heron is about the size of A. cinerea and is a rare

member of the Bone Valley avifauna. On the material now known for this

species, it is not possible to determine its relationship to other

members of the genus Ardea.


Ardea sp. indet.

Material. Love Bone Bed local fauna; UF 25939, distal end left

tarsometatarsus, missing trochlea IV.

Remarks. This specimen represents a species of Ardea about the size

of A. herodias occidentalis.


Genus Egretta T. Forster, 1817

Egretta subfluvia Becker, 1985

Material. Withlacoochee River 4A local fauna; UF 19001, right

tarsometatarsus lacking only trochlea IV and hypotarsus.

Remarks. This species is a small heron about the size of Egretta











ibis, and is only known from the holotype. The tarsometatarsus is

proportionally narrower than in other members of this genus.


Egretta sp. indet.

Material. Love Bone Bed local fauna; UF 25759, proximal end of left

carpometacarpus, UF 26082, distal end right ulna. Bone Valley Phosphate

Mining District, Payne Creek Mine; PB 7925, coracoid.

Remarks. These specimens fall within the size range of the living

E. rufescens. Because of the large time interval (4.5 MA) between the

Love Bone Bed and the Bone Valley, it is unlikely that these elements

represent the same species.


Genus Ardeola Bole, 1822

Ardeola sp. indet.

Material. Love Bone Bed local fauna; UF 25940, distal one-third

left tarsometatarsus.

Remarks. Small, similar in size to Ardeola striata. Taxonomic

assignment based entirely on size.


Subfamily Nycticoracinae Payne and Risley, 1976

Genus Nycticorax T. Forster, 1817

Nycticorax fidens Brodkorb, 1963

Material. McGehee Farm local fauna; UF 3285, complete left femur.

Remarks. See Brodkorb (1963a) for description and remarks.


Remarks on the Family Ardeidae.

Table 4.1 summarizes the distribution of the fossil herons from

Florida. The Love Bone Bed local fauna has produced three herons--a

small Ardeola a large Egretta, and a very large Ardea. It is most




Full Text
Figure 2.1. Schematic diagrams illustrating measurements of the
humerus, coracoid, and scapula. A. Humerus, cranial view. B.
Humerus, ventral view. C. Coracoid, dorsal view. D. Coracoid,
lateral view. E. Humerus, distal end view. F. Scapula, lateral
view. G. Scapula, proximal end view. Figures are not drawn to
scale. Measurements defined in the text.


27
Systematics
I have associated the skeletal elements of the fossil taxa in this
study using the following general criteria (Howard, 1932b):
1. resemblance to other species.
2. size and proportion.
3. relative abundance.
Systematic reasoning and taxonomic practice generally follow Mayr
(1981), when possible. Skeletal elements are first grouped into
similarity classes by means of measurements and qualitative characters.
Each identifiable taxon is placed within a geneology of previously known
species by the hierarchial distribution of shared-derived characters, and
the new species are then integrated into an evolutionary classification.
Paleoecology
Paleoecological methods are described by Shipman (1981). See the
paleoecology section for further comments on the paleoecological methods
used in the excavation of the localities included in this study.
Biochronology and Faunal Dynamics
A database for the examination of the biochronology and the faunal
dynamics of fossil birds of the Neogene of North America was developed
from the following sources: (l) A literature survey of all fossil
localities in North America from the late Arikareean through the Blancan
that have produced fossil birds. This published information was then
emended to reflect the current concepts of geological formations,
geological correlations, mammalian systematics, and the relative and
absolute dating of fossil localities. Verification of all fossil
identifications was made whenever possible. (2) Information from other


158
Order Ralliformes (Reichenbach, 1852)
Family Gruidae Vigors, 1825
Remarks. The following elements are so waterworn and abraded that I
cannot do more than tentatively refer them to the family Gruidae.
Material. Love Bone Bed local fauna; UF 25966, UF 25967, cervical
vertebrae; UF 26086, right digit II, phalanx I (wing); UF 26087, distal
end left radius; UF 25761, UF 25762, distal ends left carpometacarpi; UF
29723, UF 2589J+, dital ends left tibiotarsi; UF 25958, UF 25960, distal
shafts left tarsometatarsi; UF 25862, UF 25860, fragmentary distal ends
right tarsometatarsi; UF 26032, UF 26033, UF 2603^, UF 26036, UF 26037,
UF 26038, UF 26039, UF 260L0, pedal phalanges.
Generic diagnosis. The following diagnosis is based on an
examination of the following genera (number of species in parenthesis):
Grus (9), Bugeranus (l), Anthropoides (2), and Balerica (2).
No diagnostic characters noted on portion of the coracoid and
carpometacarpus preserved. Femur with proximal portion of the cranial
intermuscular line located medially, away from the crista trochanteris in
Balerica (extending from the crista trochanteris in other genera
examined). Only a few characters on the distal end of the tibiotarsus
will separate all species of Grus from all other species of Bugeranus,
Anthropoides, and Balerica. In Grus, the lateral surface of the lateral
condyle is expanded (less so in other genera), the internal ligamental
process is expanded (similar in Bugeranus, less so in Anthropoides, and
Balerica). Balerica is distinguished by having the distal end anterio-
posteriorly flattened, with the medial condyle rotated outward, producing
a broad, U-shaped anterior intercondylar sulcus. The proximal end of the
tarsometatarsus in Balerica with a short hypotarsus (longer in Grus,
Anthropoides (but short in A. virgo), and Bugeranus); Anthropoides with a


75
characters with Anhinga (Cheneval, 1984). Phalacrocorax littoralis
(Milne-Edwards) from the Aquitanian of St.-Ge'rand-le-Puy, France, and
from Germany was based on a coracoid and a few other skeletal elements.
It seems to be related to P. aristotelis. Phalacrocorax anatolicus
Mourer-Chauvire' from the lower or middle Miocene (probably Helvetian) of
Bes-Konak, Turkey, was described from a coracoid and most of a forelimb.
It appears to be related to the fossil species P. littoralis, P.
miocaenus, and to the recent P. aristotelis.
Phalacrocorax leptopus Brodkorb from the Clarendonian and
Hemphillian localities of Juntura, Oregon, is based on a coracoid,
tarsometatarsus, and scapula. It is a small species and resembles the
fossil P. littoralis. It is in need of additional comparisons to
elucidate its relationships.
Phalacrocorax femoralis L. Miller from the Barstovian or
Clarendonian Calabasas local fauna, California, is based on most of a
skeleton, preserved in a slab of fine-grained shale. It is the size of
P. penicillatus, but Miller (1929) asserts that this species does not
appear closely related to any living species. Phalacrocorax lautus
Kurochkin and Ganya from the upper Miocene of Moldavia, is based on the
proximal half of a right femur. It appears closest to the living P.
pygmaceus.
Phalacrocorax praecarbo Ammon was described from the upper Miocene
Brown Coal Formation, near WUrttemburg, Germany, on the humeral end of a
coracoid. Brodkorb (1980) has moved Ardea brunhuberi Ammon (figured in
Ammon, 1911), based on a proximal end of a carpometacarpus to this
species and emended the name to P. brunhuberi (Ammon, 1918; cited as 1911
in Brodkorb, 1980). Olson (ms) also moves Botaurites auritus Ammon,


69
55868, proximal end right tibiotarsus; UF 55836, UF 57245, UF 60056,
proximal end left tibiotarsus; UF 55804, UF 57250, distal ends right
tibiotarsi; UF 53935, UF 55863, UF 55864, UF 57357, UF 58305, UF 60057,
UF 60058, UF 65713, distal ends left tibiotarsi; UF 55860, nearly
confete left tarsometatarsus; UF 524l6, UF 52417, UF 524l8, UF 55837, UF
58067, proximal end right tarsometatarsi; UF 55870, UF 58421, UF 61972,
proximal ends left tarsometatarsi; UF 53889, UF 53936, UF 55815, UF
57251, UF 58306, UF 58341, distal ends right tarsometatarsi; UF 58342, UF
60059, UF 65658, distal end left tarsometatarsi.
Bone Valley Mining District, Gardiner Mine.UF 58438, caudal
portion right mandible; UF 61998, right clavicle; UF 61999, proximal end
right scapula; UF 62000, proximal end left scapula, UF 65667, complete
left coracoid; UF 58278, UF 58279, UF 58439, UF 58440, UF 62001, UF
65669, humeral ends right coracoids; UF 58277, UF 58280, UF 58446, UF
58447, UF 58469, UF 62002, UF 62003, UF 62004, UF 65668, humeral ends
left coracoids; UF 58448, sternal end left coracoid; UF 58470, UF 62005,
proximal ends right humeri; UF 58441, UF 58471, distal ends right humeri;
UF 58449, UF 58472, UF 62006, distal ends left humeri; UF 58281, UF
58442, UF 62007, UF 6567O, proximal end right ulnae; UF 62008; proximal
end left ulna; UF 57307, UF 58285, UF 58286, UF 65671, distal end right
ulnae; UF 58282, UF 58283, UF 58284, UF 65672, UF 65749, distal end left
ulnae; UF 58443, proximal end right carpometacarpus; UF 58287, proximal
end left carpometacarpus; UF 58303, UF 58282, distal ends left
carpometecarpi; UF 58450, complete left femur; UF 58451, proximal end
left femur; UF 58473, distal end right femur; UF 57311, proximal end left
tibiotarsus; UF 58290, UF 58444, UF 58445, UF 58474, UF 58475, distal
ends right tibiotarsi; UF 58289, UF 58452, UF 58453, UF 62009, UF 62010,


Figure 6.2. Graphic representation of avian faunal dynamic
parameters. A. Number of localities per million years per NALMA.
B. Running mean per NALMA. C. Extinction rate per NALMA. D.
Origination rate per NALMA. Abbreviations as in Table 6.1.


37
sparrow, Palaeostruthus eurius, was described from here (Brodkorb,
1963a). Reptiles known from here include Deirochelys and Gavialosuchus.
Haile XIXA
This locality is 2.5 miles HE of Newberry, Alachua County, in the NE
1/4, Sec. 26, T. 9 S., R. 17 E., Newberry Quadrangle, U. S. Geologic
Survey 75 minute series topographical map, 1968. It was collected by
the Florida State Museum staff. Vertebrates are mainly aquatic,
including a large amount of skeletal material of Gavialosuchus. Fossil
mammals present include Epicyon, geolocids, Pediomeryx, equids, and
Aepycamelus.
Bone Valley Mining District
The name "Bone Valley" is applied to the phosphate mining district
of central Florida, mainly in Polk County, but also including portions of
adjacent Hillsborough, Hardee, and Manatee counties. The vertebrate
fauna was first described by Sel lards (1916) and later more fully studied
by Simpson (1930). Berta and Morgan (in press) present an account of the
present status of vertebrate paleontology of this area. Much of the
following comes from their paper.
Most of the fossil vertebrates from the Bone Valley area were
obtained from extensive open-pit phosphate mines. In situ collections
are rare and make up approximately 5-10 % of the Florida State Museum
collections, part of the USGS and USNM collections, and virtually none of
the Harvard collections. Avian fossils iron these in situ sites are very
rare. Fossils are more commonly found eroding from spoil piles after an
area has been strip-mined. Except for the in situ collections and a few
intensively collected concentrations, the fossil vertebrates collected


Figure 3.1
Correlation Chart of Included Local Faunas


207
An index of relative sampling may be calculated to adjust for the
unequal lengths of time represented by these discrete NALMAs (North
American Land Mammal Ages) and to take into account the number of
sampling sites. The number of local faunas of a given NALMA are divided
by the duration of that NALMA (Table 6.2). This shows that the last
three NALMAs of the Neogene are better sampled than the first three by a
factor of 2 or 3.
When the local faunas are further divided into marine and non-marine
groups, even greater discrepancies in the representations of the NALMAs
are apparent. In general the non-marine local faunas outnumber the
marine local faunas by at least a factor of 3 to 1 (Table 6.2; 6.3; and
Figure 6.U).
Additional comparisons can be made between the number of published
versus unpublished localities. While these comparisons are biased by the
collections I have been able to examine, marine avifaunas have.been more
completely described than terrestrial ones (compare Table 6.3; 6.L).
The fossil avifaunas from nearly one-half of all terrestrial local faunas
in which birds are present are entirely unstudied and unreported in the
literature. As all major collections in the United States were not
examined, this is certainly an underestimate of the amount of work left
to be done and underscores the preliminary nature of this examination of
faunal dynamics.
The Neogene Avifauna
The combined sample of all families and genera will be considered
first, then reconsidered separately in marine and non-marine groups.


168
Table 4.25 Measurements of the tarsometatarsi of Aramornis longurio
(F:AM 6269, holotype) and Aramornis sp. from the Love Bone Bed local
fauna. Abbreviation are defined in methods section. (*) Specimen
abraded or broken.
Measurements Aramornis lonRurio
Aramornis sp.
Tarsometatarsus
W-TRII 4.2
4.4
D-TRII
*7.2
W-TRIII 5.7
*6.0
D-TRIII 7.2
*7-7
D-TRIII


106
Family Plataleidae Bonaparte, 1838
Subfamily Threskiornithinae (Richmond, 1917)
Genus Eudocimus Wagler, 1832
Eudocimus sp. A
Material. Bone Valley Mining District, Gardinier Mine; UF 60040,
distal end left tibiotarsus. Palmetto Mine, PB 77^+9, proximal end left
tarsometarsus.
Description. Tibiotarsus similar in size and general morphology to
large individuals (males) of Eudocimus albus or E. ruber. Tendinal
bridge as in E. ruber (shorter in E. albus), tubercle well-developed (as
in E. albus, less developed in E. ruber), extensor sulcus more excavated
and flattened than either E. ruber or E. albus, well-developed crest
curves obliquely toward lateral face from tubercle (crest well developed
and oriented parallel to the long axis of the shaft in both E. albus and
E. ruber). Lateral margins of lateral condyle not as developed as in E.
albus or E. ruber. Proximal border of the posterior articular surface
extends farther proximally on the external side of the fossil (border is
usually straight in E. albus and E. ruber; this condition similar in some
individuals of Plegadis spp., especially P. ridgwayi).
Tarsometatarsus similar in size and overall morphology to Eudocimus
albus and E. ruber. I can find no character on the fossil specimen that
is not within the range of variation of these two species.
Remarks. Olson (1981b), in his review of the fossil ibises reported
Eudocimus sp. from a distal end of a tarsometatarsus from Bone Valley. I
have not been able to locate a distal tarsometatarsus in either the
Brodkorb collection or in the FSM collections. I therefore conclude that


64
Table 4.5 Measurements of the tibiotarsi of the grebes Podilymbus
podiceps (N = 14, T males, 7 females) and Podilymbus cf. _P. podiceps from
the Bone Valley Mining District. Data are mean _+ standard deviation and
range. (*) Specimen damaged. Abbreviations are defined in the methods
section.
Measurements
Podilymbus podiceps
P. cf. podiceps
Tibiotarsus
LENGTH
68.69 + 4.31
61.7 77.0

M-LENGTH
80.17 + 5.27
72.4 90.2

W-SHAFT
4.50 + 0.4l
3.8 5.3
4.6; 4.9
D-SHAFT
3.29 + 0.27
2.8 3.8
3.5; 3.7
W-PROX-M
7.00 + 0.44
6.5 7.9

W-DIST-CR
7.10 + 0.65
6.1 7.9
7.2; *7.4
W-DIST-CD
6.04 + 0.37
5.2 6.5
*6.2; *6.5
D-MCON
6.86 + 0.44
6.1 7.5
*6.9; 7.4
D-LCON
6.76 + 0.49
5.9 7.4
7.0; 7.3
D-ICON
4.29 + 0.33
3.7 4.9
4.4; 4.9


BIOGRAPHICAL SKETCH
Jonathan J. Becker was born in Jerome, Idaho, on 16 December 1955
and was raised on a nearby farm. He graduated from the Jerome High
School in May 1974 and attended Idaho State University in Pocatello,
Idaho, from 1974 to 1980; receiving a B. S. in zoology with high honors
in May 1978 and a M. S. in zoology/biology in May 1980. Since August
1980 he has attended the University of Florida, graduating with a Ph. D.
in zoology in August 1985* He is currently a postdoctoral fellow at the
National Museum of Natural History, Smithsonian Institution.
He is a member of the American Association for the Advancement of
Science, American Ornithologists' Union, American Society of
Mammologists, Biological Society of Washington, Cooper Ornithological
Society, Florida Acadeny of Science, Sigma Xi, Society of Systematic
Zoology, Society of Vertebrate Paleontology, and Wilson Ornithological
Society.
His current research interests include the functional morphology of
birds and mammals and the evolution, systematics, and biochronology of
the Neogene birds of North America.
245


TO
UF 62011, UF 62012, distal ends left tibiotarsi; UF 65673, complete right
tarsometarsus; UF 58478, UF 62013, proximal end right tarsometatarsi; UF
58291, UF 58292, proximal end left tarsometarsi; UF 58293, UF 58476, UF
58U77 UF 6567U, distal ends right tarsometatarsi; UF 57310, distal end
left tarsometatarsus.
Bone Valley Mining District, Palmetto Mine.UF 21058, sternal
fragment with coracoidal sulci; UF 13225, humeral end left coracoid; UF
21143, shaft left coracoid; UF 21115, shaft right coracoid; UF 21146,
distal end right humerus; UF 29734, distal end left humerus; UF 13231, UF
29740, distal ends right ulnae; UF 21091, UF 21120, proximal ends right
carpometacarpi, UF 21068, UF 21072, proximal ends left carpometacarpi; UF
21131, distal end right carpometacarpus; UF 49090, complete right femur;
UF 12352, complete left femur; UF 29735, UF 49091, proximal ends left
tarsometatarsi; UF 12868, distal end right tarsometatarsus.
Bone Valley Mining District, New Palmetto Mine.UF 49691, UF 49692,
vertebrae (questionably referred), UF 49693, complete right femur; UF
49694, complete left tarsometatarsus.
Bone Valley Mining District, North Palmetto Mine.UF 49097, humeral
end left coracoid; UF 49098, humeral end right coracoid.
Bone Valley Mining District, Southwest Palmetto Mine.UF 49093,
humeral end right coracoid.
Bone Valley Mining District, Hookers Prairie Mine.UF 49690,
complete right femur.
Bone Valley Mining District, Kingsford Mine.UF 21186, humeral end
left coracoid; UF 52971, distal end right humerus; UF 13212, distal end
left humerus; UF 21185, proximal end right tibiotarsus.


230
Becker, J. J. 1985a. Fossil herons (Aves: Ardeidae) of the late
Miocene and early Pliocene of Florida. Journal of Vertebrate
Paleontology, 5:24-31.
Becker, J. J. 1985b. Pandion lovensis, a new species of Osprey from the
late Miocene of Florida. Proceedings of the Biological Society of
Washington, 98:314-320.
Becker, J. J. 1985c. A late Pleistocene (Wisconsinian) avifauna from
West Palm Beach, Florida- Bulletin of the British Ornithologists'
Club, 105:37-40.
Becker, J. J. ms. Birds of the Pliocene (Blancan) Oreana local fauna,
Owyhee County, Idaho.
Becker, J. J. ms. Neogene Avian Localities of North America.
Becker, J. J. ms. Fossil Anhingas (Aves, Pelecaniformes, Anhingidae) of
North America.
Berta,. A. and H. Galiano. 1983. Megantereon hesperus from the late
Hemphillian of Florida, with remarks on the phylogenetic
relationships of machairodonts (Mammalia, Felidae,
Machairodontinae). Journal of Paleontology, 57:892-899.
Berta, A. and H. Galiano. 1984. A Miocene amphicyonid (Mammalia:
Carnivora) from the Bone Valley Formation of Florida. Journal of
Vertebrate Paleontology, 4:122-125.
Berta, A. and G. S. Morgan, in press. A new sea otter (Carnivora:
Mustelidae) from the late Miocene and early Pliocene (Hemphillian)
of North America- Journal of Paleontology.
Blake, E. R. 1977. Manual of Neotropical Birds. Spheniscidae
(Penguins) to Laridae (Gulls and Allies). Vol. 1. University of
Chicago Press, Chicago. 674 pp.
Breyer, J. A. 1981. The Kimballian land-mammal age: mene, mene, tekel,
upharsin (Dan. 5:25). Journal of Paleontology, 55:1207-1216.
Brodkorb, P. 1952. A new rail from the Pleistocene of Florida. Wilson
Bulletin, 64:80-82.
Brodkorb, P. 1953a. Pleistocene birds from Haile, Florida. Wilson
Bulletin, 5:49-50.
Brodkorb, P. 1953b. A Pliocene flamingo from Florida. Natural History
Miscellanea, No. 124:1-4.
Brodkorb, P. 1953c. A Pliocene gull from Florida. Wilson Bulletin,
65:94-98.
Brodkorb, P. 1953d. Review of Pliocene loons. Condor, 55:211-214.


THE FOSSIL BIRDS OF THE' LATE MIOCENE AND EARLY PLIOCENE
OF FLORIDA
BY
JONATHAN J. BECKER
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1985

Copyright 1985
8y
Jonathan J
Becker

ACKNOWLEDGMENTS
I would first like to thank my advisor, Pierce Brodkorb, for his
support, guidance, and friendship during the course of this project. He
has provided much encouragement and councel throughout ny researches on
fossil birds, as have the members of ny committeeDrs. Richard A.
Kiltie, S. David Webb, Elizabeth S. Wing, and Ronald G. Wolff. I thank
them for the many hours spent reading this manuscript and for their many
helpful comments pertaining to my research.
A number of friends and colleagues contributed to this study with
their comments, suggestions, and discussions. They include S. Emslie, R.
Hulbert, and A. Pratt. Gary S. Morgan has taken much time to discuss the
geology of Florida and systematics, evolution, and biochronology of
fossil and Recent mammals, in addition to many other aspects of
vertebrate paleontology. I especially thank him for his comments on
Chapter III. Storrs Olson has freely shared his considerable knowledge
and insights of avian systematics, evolution and anatomy. David Steadman
also provided many helpful comments. I also thank Cynthia West for her
support and aid in completing this dissertation.
I thank the many amateur fossil collectors who have generously
donated fossil birds from the included localites to the Florida State
Museum. They include Danny Bryant, George Hess lop, and Ron Love. Phil
Whisler, of Venice, Florida, originally discovered the SR-6U locality and
brought it to the attention of the staff of the Florida State Museum.
Rick Carter, of Lakeland, Florida, deserves special mention, for over the
iii

last three years, he has donated hundreds of specimens of fossil birds
from the Bone Valley Mining District to the Florida State Museum.
Without his generous contributions, the Bone Valley portion of this study
would have been impossible to complete. John Waldrop is also gratefully
acknowledged for providing information about the avian localities in the
Bone Valley.
I also thank the following individuals and institutions for making
fossil and/or Recent specimens available for study: A. Andors, G.
Barrowclough, C. Houlton, R. H. Tedford, F. Vuilleumier, American Museum
of Natural History; P. Brodkorb, Department of Zoology, University of
Florida; J. Chenval, C. Mourer, Universite Claude Bernard; J. Hardy, B.
J. MacFadden, G. S. Morgan, S. D. Webb, T. Webber, Florida State Museum;
R. Mengel, University of Kansas; R. Payne, University of Michigan; M.
Voorhies, University of Nebraska; H. James, S. L. Olson, D. Steadman,
United States National Museum. Helen James, Storrs L. Olson, and David
Steadman generously provided accommodations during a lengthy stay in
Washington, D.C.
Financial support received while at the University of Florida,
includes teaching and research assistantships from the Department of
Zoology, College of Liberal Arts and Sciences; teaching assistantships
from the Department of Physiological Sciences, College of Veterinary
Medicine; a curatorial assistantship from the Department of Vertebrate
Paleontology, Florida State Museum; and grants from the Frank M. Chapman
Memorial Fund, American Museum of Natural History, and from Sigma Xi
Grants-In-Aid of research. The Department of Zoology, College of Liberal
Arts and Sciences, and the Florida State Museum supplied all expendable
iv

equipment. I gratefully acknowledge these departments and institutions
for their support.
Last, I thank my parents, Elwood W. and Nita E. Becker, for their
encourgement and support over the years. They have provided not only the
opportunity, but much of the impetus, that has allowed me to finish my
formal education.
v

TABLE OF CONTENTS
PAGE
ACKNOWLEDGMENTS iii
ABSTRACT viii
CHAPTER
I. INTRODUCTION AND PREVIOUS WORK 1
Introduction 1
Limitations of Study 2
Previous Work U
II. METHODS 9
Measurements 9
Computer Software 18
Nomenclature 18
Systematics 27
Paleoecology 27
Biochronology and Faunal Dynamics 27
Specimens Examined 29
Abbreviations 29
III. GEOLOGY 31
Biochronology 31
Local Faunas 33
Eustatic Sea-level Changes 1+1
IV. SYSTEMATIC PALEONTOLOGY 1+9
Order Podicipediformes 1+9
Order Pelecaniformes 65
Order Ciconiiformes 90
Order Accipitriformes Il6
Order Anseriformes 136
Order Galliformes 155
Order Ralliformes 158
Order Charadriiformes 175
Order Strigiformes 191
Order Passeriformes 195
vi

V. PALEOECOLOGY 198
Introduction 198
Local Faunas 199
VI. BIOCHRONOLOGY AND FAUNAL DYNAMICS 205
Introduction 205
Faunal Dynamics 205
Biochronology . 213
VII. SUMMARY 224
Systematics 224
Paleoecology 227
Biochronology 228
LITERATURE CITED 229
BIOGRAPHICAL SKETCH 245
vii

Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
THE FOSSIL BIRDS FROM THE LATE MIOCENE AND EARLY PLIOCENE
OF FLORIDA
By
Jonathan J. Becker
August 1985
Chairman: Pierce Brodkorb
Maj or Department: Zoology
This study examined the non-marine avifauna from ten late Miocene
and early Pliocene localities in Florida. These localities include the
Love Bone Bed, McGehee Farm, Mixson's Bone Bed, Bone Valley Mining
District, Withlacoochee River UA, Manatee County Dam, SR-6U, Haile VB,
Haile VI, and Haile XIXA. Non-marine genera (number species, if more
than one) present include Rollandia, Tachybaptus, Podilymbus (2),
Podiceps, Pliodytes, Phalacrocorax (3), Anhinga (2), Ardea (2), Egretta
(2 or 3), Ardeola, Nycticorax, Mycteria, Ciconia (3), Eudocimus,
Plegadis, Threskiornithinae, genus indeterminate, Pliogyps, Pandion (2),
Haliaeetus, Buteo, Aguila, Accipitrid, genus indeterminate (3),
Dendrocygna, Branta, Anserinae, genus indeterminate (U), Tadorine, genus
indeterminate, Anas (2), Anatine, genus indeterminate (2), Aythya,
Oxyura, Meleagridinae, genus indeterminate, Meleagris, Grus (2),
Balearicinae, genus indeterminate, Aramornis, Rallus (3), Rallid,
undescribed genus, Phoenicopterus (2), Jacana, Limosa, "Calidris" (6+),
viii

?Actitis, ?Arenaria, ?Philomachus, Tytonid, undescribed genus, Bubo,
Passeriformes (2).
The largest avifaunas are from the Love Bone Bed local fauna (44
taxa present) and the Bone Valley local fauna (4l taxa present, 31 here
included). These two localities are the most diverse non-marine and
marine avifaunas, respectively, known in North America prior to the
Pleistocene.
An analysis of the faunal dynamics of the Neogene fossil birds from
North America shows the following results, (l) Localities which have
produced fossil birds are not uniformly distributed through time74.4%
of the localities are from the last 4l% of the Neogene. (2) By the
Barstovian, a majority of the living families which have a fossil record,
have appeared. (3) Generic diversity increases from 10 to 98 during the
Neogene. (4) The marine avifauna is essentially established at a
diversity of 20 to 25 genera by the Clarendonian. (5) The non-marine
avifauna increases continually throughout the Neogene. (6) Origination
rates for marine birds peak in the Clarendonian with 4.4 genera appearing
per million years. (7) Extinction rates for marine birds are
consistently low throughout the Neogene. (8) Origination rates for non
marine birds show a 2- to 4-fold increase in alternate Land Mammal Ages.
(9) Turnover rates parallel the origination rates described above.
ix

CHAPTER I
INTRODUCTION AND PREVIOUS WORK
Introduction
Florida has one of the richest records of fossil birds in the world.
This study examines the systematics of the non-marine fossil birds which
lived in Florida during the late Miocene and early Pliocene (9.04.5
million years before present) and the paleoecology of the localities
which produced them. Included is material from 10 local faunasthe
Love Bone Bed, McGehee Farm, Mixson Bone Bed, Bone Valley, Withlacoochee
River 4a, Manatee County Dam Site, SR-64, Haile VB, Haile VI, and Haile
XIXA. Only a few of the birds from McGehee Farm and the early
collections of birds from Bone Valley have been studied previously (Table
l.l). In addition, the biochronology and faunal dynamics of the entire
North American Neogene avifauna are investigated. Specifically, the
following questions are addressed:
Systematics
1. What species of fossil birds are present in Florida during the
late Miocene and early Pliocene?
2. What are their systematic and biogeographic relationships to
other fossil and Recent species?
Paleoecology
1. Can fossil birds be used to reconstruct the paleoenvironments of
the fossil localities examined in this study?
1

2
Biochronology and Faunal Dynamics
1. What is the temporal distribution of the fossil localities
producing birds in the Neogene of North America?
2. What is the temporal distribution of the North American Neogene
avifauna?
3. When, and at what rate, do the North American Neogene avian
families and genera appear and become extinct?
4. How do the marine and non-marine localities and avifaunas differ
in questions 1 to 3 above?
5. What avian species are biostratigraphically useful in the
Neogene of North America?
6. What degree of temporal resolution does avian biochronology
offer?
Limitations of Study.
There have been several limitations imposed on this study by the
current knowledge of avian systematics and paleontology, and to a lesser
degree by the lack of previous studies dealing with the paleoecology and
biochronology of birds. Birds are often considered ". . the best known
and most completely described class of animals . (Welty 1975:14).
In reality, this statement applies only to the simple designation of
living forms and to the obvious aspects of their behavior and ecology,
but definitely not to their evolution, their superspecific relationships,
and to many aspects of internal morphology. Many living groups of birds
still lack modern systematic revisions based on internal morphology, or
in many, even a description of their internal morphology. Most modern
orders have never been shown to be monophyletic, although many are
doubtlessly so. Modern classifications of birds above the specific level

3
(e.g., American Ornithologists' Union, 1983) has changed very little in
the 90 years since Gadov's (1893) classification. Olson (l98la:193),
addressing this problem, states
Many ornithologists appear to believe that the higher-
level systematics of birds is a closed book, the sequence of
orders and families in their field guides being an immutable
constant that was determined long ago according to some
infallible principle. In reality, the present classification
of birds amounts to little more than superstition and bears
about as much relationship to a true phylogeny of the Class
Aves as Greek mythology does to the theory of relativity. A
glance at the Gadov-Wetmore classification now in use shows
that there is still no concept in ornithology of what
constitutes a primitive bird.
Certainly correct phytogenies are impossible to develop without an
accurate knowledge of primitive character states, the distribution of
primitive and derived characters within the group being studied, and
meaningful outgroup comparisons.
Many fossil species are described from single, non-diagnostic
elements, making useful comparisons between, or among, species
impossible. But even diagnostic elements, when singly preserved, add
little to our understanding of the phylogeny and evolution of that taxon.
Other species are arbitrarily allied with the wrong family, the wrong
order, and even in some cases, with the wrong class of vertebrates (cited
in Brodkorb, 1978:211-228), many because of lack of proper comparisons.
Only recently have many of these errors been recognized, due in part to
an increased number of workers in the field, and from greater
availability and use of comparative skeletal material. Adequate skeletal
collections are still lacking for many common species (Zusi et al.,
1982).
It is outside the scope of this study to revise the many recent and
fossil genera and families which need such treatment. In such cases, I

4
try to note the systematic problems in each group, briefly review its
fossil record, and describe the fossil material from Florida. This
material should be reexamined as the state of systematics of these groups
improves and as additional Recent and fossil material becomes available.
Previous Work
The earliest published report of fossil birds from Florida is
Sellard's (1916) description of a supposed jabir (Jabir? weillsi
=Ciconia maltha) from Vero, closely followed by Shufeldt's (1917a, 1917b)
study of this local fauna. Wetmore (1931) reviewed the Pleistocene
avifauna of Florida and was also the first to report (19^3) on the
Tertiary birds from Thomas Farm and the Bone Valley Mining District.
Numerous studies have appeared since then, primarily by Brodkorb or his
students. A few other faunal and systematic studies have also included
avian material from Florida.
Table 1.1 lists many of the fossil localities in Florida which have
a notable record of fossil birds. Reference to Brodkorb's Catalogue of
Fossil Birds (1963-1978), where the avifaunas from many Florida
Pleistocene localities were first reported, is omitted to conserve space.

5
Table 1.1. Avian Fossil Localities of Florida.
Locality
Reference (s)
MIOCENE TO EARLIEST PLIOCENE
(Pre-Blancan)
Bone Valley, Polk Co.
Becker, 1985a; Brodkorb, 1953b, 1953c,
1953d, 1953e, 1955a, 1970; Olson, 1981b;
Steadman, 1980; Wetmore, 19^3; this study
Gainesville Creeks,
Alachua Co.
Brodkorb, 1963b
Haile VI, Alachua Co.
Brodkorb, 1963a; this study
Haile XIXA, Alachua Co.
this study
Love Bone Bed, Alachua Co.
Becker, 1985a, 1985b; Webb et al., I98I;
this study
Manatee Co.Dam Site
Manatee Co.
Webb and Tessman, 1968; this study
McGehee Farm, Alachua Co.
Brodkorb, 1963a; Hirschfeld and Webb, 1968;
Olson 1976; this study
Mixson Bone Bed, Levy Co.
this study
Seaboard Airline Railroad,
Leon Co.
Brodkorb, 1963b
SR-64, Manatee Co.
this study
Thomas Farm, Gilchrist Co.
Brodkorb, 1954a, 1956a, 1963b; Cracraft,
1971; Olson and Farrand, 1974; Steadman,
1980; Wetmore, 1943, 1958
Withlacoochee River UA,
Marion Co.
Becker, 1985a; this study
LATE PLIOCENE (Blancan)
Haile XVA, Alachua Co.
Campbell, 1976; Steadman, 1980
Santa Fe IB, Gilchrist Co.
Brodkorb, 1963d

6
Table 1.1continued.
Locality Reference (s)
EARLY PLEISTOCENE (irvingtonian)
Coleman IIA, Sumter Co.
Ritchie 1980; Steadman, 1980
Haile XVIA, Alachua Co.
Steadman, 1980
Inglis IA, Citrus Co.
Carr, 1981; Ritchie, 1980; Steadman, 1980
Santa Fe River IIA,
Gilchrist Co.
Steadman, 1980
Williston, Levy Co.
Holman, 1959, 1961; Steadman, 1980
LATE PLEISTOCENE (Rancholabrean)
Arredondo, Alachua Co.
Brodkorb, 1959; Holman, 196l; Olson, 1974b,
1977b; Steadman, 1976, 1980; Storer, 1976b
Aucilla River IA,
Jefferson Co.
Steadman, 1980
Bowman IA, Putnam Co.
Steadman, 1980
Bradenton, Manatee Co.
Becker, 1984; Steadman, 1980; Wetmore,
1931
Catalina Lake,
Pinellas Co.
Storer, 1976b
Coleman III, Sumter Co.
Ritchie, 1980
Crystal Spring Run,
Pasco Co.
Brodkorb, 1956b
Davis Quarry, Citrus Co.
Steadman, 1980
Econfina River, Taylor Co.
Steadman, 1980
Eichelberger Cave,
Marion Co.
Brodkorb, 1955b; Holman, 1961
Florida Lime Company,
Marion Co.
Steadman, 1980
Haile IA, Alachua Co.
Brodkorb, 1953a, 1954b; Olson, 1974b, 1977b
Haile IIA, Alachua Co.
Holman, 1961; Steadman, 1980
Haile VIIA, Alachua Co.
Steadman, 1980

7
Table 1.1continued
Locality
Reference (s)
Haile XIB, Alachua Co.
Lign, 1965i Olson, 19T^b
Hog Cave, Sarasota Co.
Steadman, 1980; Wetmore, 1931
Hog Creek, Manatee Co.
Wetmore, 1931
Hornsby Springs,
Alachua Co.
Storer, 1976b
Itchtucknee River,
Columbia Co.
Campbell, 1980; McCoy, 1963; Olson, 197^+a,
1971+b, 1977b; Storer, 1976b; Wetmore, 1931
Jenny Spring, Gilchrist Co.
Storer, 1976b
Kendrick IA, Marion Co.
Steadman, I98O
Lake Monroe, Volusia Co.
Holman, 1961; Storer, 1976b
Mefford Cave I, Marion Co.
Steadman, 1980
Melbourne, Brevard Co.
Holman, 1961; Steadman, 1980; Wetmore, 1931
Monkey Jungle, Dade Co.
Ober, 1978
Oakhurst Quarry, Marion Co.
Holman, 1961; Steadman, 1980
Orange Lake, Marion Co.
Holman, 1961
Reddick IB, Marion Co.
Brodkorb, 1952, 1957, 1963e; Hamon, I96U;
Holman, 1961; Olson, 197^b, 1977b;
Steadman, 1976, 1980; Storer, 1976b
Rock Springs, Orange Co.
Storer, 1976b; Steadman, 1980; Woolfenden,
1959
Sabertooth Cave, Citrus Co.
Holman, 196l; Wetmore, 1931
St. John's Lock, Putnam Co.
Storer, 1976b
St. Mark's River,
Leon/Wakulla Co.
Steadman, 1980
Santa Fe River IA,
Gilchrist Co.
Steadman, 1980
Santa Fe River IVA,
Gilchrist Co.
Steadman, I98O
Seminole Field,
Pinellas Co.
Holman, I96I; Olson, 1974b; Steadman, 1980;
Wetmore, 1931

8
Table 1.1continued
Locality
Reference (s)
Steinhatchee River,
Taylor/Dixie Co.
Steadman, 1980
Venice Rocks, Manatee Co.
Wetmore, 1931
Vero (Stratum 2),
Indian River Co.
Holman, 1961; Sellards, 1916; Shufeldt,
1917; Steadman, 1980; Storer, 1976b; Weigel,
1962; Wetmore, 1931
Warren's Cave, Alachua Co.
Holman, 1961
Wekiva Run III, Levy Co.
Steadman, 1980
West Palm Beach,
Palm Beach Co.
Becker, 1985c
Withlacoochee River,
Citrus Co.
Steadman, 1980
Zuber, Marion Co.
Holman, 1961
HOLOCENE
Cotton Midden, Volusia Co.
Hay, 1902; Neill et al., 1956
Castle Windy Midden,
Volusia Co.
Weigel, 1958
Good's Shellpit,
Volusia Co.
Steadman, 1980
Green Mound Midden,
Volusia Co.
Hamon, 1959
Nichol's Hammock, Dade Co.
Hirschfeld, 1968; Steadman, I98O
Silver Glenn Springs,
Lake Co.
Neill et al., 1956; Steadman, 1980
Summer Haven Midden,
St. Johns Co.
Brodkorb, i960
Vero (Stratum 3),
Indian River Co.
see Vero, Stratum 2, above
Wacissa River
Jefferson Co.
Steadman, 1980

CHAPTER II
METHODS
Measurements
Measurements made in this study are listed below and are illustrated
in Figures 2.1 2.k. They have been selected from the literature
dealing with the osteology and identification of both Recent and fossil
species of birds. Previous authors have dealt with only one specific
group (Steadman, 1980; Howard, 1932b; Ono, 1980) or with Recent birds
commonly found in archeological sites (von den Driesch, 1979; Gilbert, et
al., 1981). The most applicable anatomical studies on fossil birds are
by Ballmann (1969a, 1969b). I have modified measurements from the above
studies to make them applicable to the range of morphologies encountered
in the fossil birds of this study. Measurements for the less diagnostic
elements and for the cranium were not included. Measurements presented
in the systematic section of this dissertation depend on the fossil
material available and the morphology of the species considered.
Anatomical terminology follows Baumel et al. (1979) and Howard
(1929, 1980). Many Latin terms have been anglicized for ease of
communication, but the original Latin is given parenthetically when the
term is first used (below). Terms for soft tissue anatony come from
Feduccia (1975) and Van den Berge (1975 )
9

10
Scapula
1. LENGTH.Greatest length from the acromion to the caudal
extremity of the scapula (Extremitas caudalis scapulae).
2. W-NECK.Least width neck of scapula (Collum scapulae).
3. W-PROX.Proximal width from the ventral tip of the glenoid
facet (Facies articularis humeralis) to the dorsal margin of the
scapular head (Caput scapulae).
4. ACR-GLN.Length from tip of acromion through ventral tip of
glenoid facet.
5. D-GLN.Depth of glenoid facet.
Coracoid
1. HEAD-FAC.Length from head (Processus acrocoracoideus) through
external end of sternal facet (Facies articularis sternalis).
2. HEAD-IDA.Length from head through internal distal angle
(Angulus medial is).
3. HEAD-CS.Length from head through scapular facet (Cotyla
scapularis).
4. D-HEAD.Least depth of head.
5. W-SHAFT.Width of midshaft.
6. D-SHAFT.Depth of midshaft.
7. FAC-IDA.Length from external end of sternal facet through
internal distal angle.
8. IDA-PL.Length from internal distal angle to medial most edge
of sternocoracoidal process (Processus lateralis).
9 IDA-PP.Length from internal distal angle to procoracoid
process (Processus procoracoideus).

11
10. L-GLEN.Length of glenoid facet from the most cranial portion
of glenoid through the most caudal point of scapular facet.
11. IDA-FNS.Length from internal distal angle through the most
sternal' edge of coracoidal fenestra (Foramen nervus
12.ANG-HEAD.Angle formed between axis of the head, as seen in
proximal view, and the plane parallel to the dorsal surface
(Facies dorsalis).
Humerus
1. LENGTH.Greatest length from the head of the humerus (Caput
humeri) through the midpoint of the lateral condyle (Condylus
ventralis).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. W-PROX.Transverse width of proximal end from the external
tuberosity (Tuberculum dorsale) to the most ventral face of the
bicipital crest (Crista bicipitalis).
5. D-PROX.Depth of proximal end, from the bicipital surface
(Facies bicipitalis) to the internal tuberosity (Tuberculum
ventrale), measured at right angles to the long axis of the
shaft.
5a. D-HEAD.Depth of head, measured parallel to the axis of the
head.
6. L-DELTOID.Length of deltoid crest (Crista pectoralis),
measured from the external tuberosity to the most distal
extension of the deltoid crest

12
7. W-DIST.Transverse width of distal end from the entepicondylar
prominence (Epicondylus ventralis) to the ectepicondylar
prominence (Epicondylus dorsalis).
8. D-DIST.Depth of distal end from cranial face of external
condyle (Condylus dorsalis) through ridge slightly mediad from
external tricipital groove (Sulcus scapulotricipitis), measured
at right angles to the long axis of the shaft.
9. D-ENTEP.Depth of entepicondyle (Epicondylus ventralis) from
attachment of the pronator brevis (Tuberculum supracondylare
ventrale) through entepicondyle (Processus flexoris), measured at
right angles to the long axis of the shaft.
Ulna
1. V-LENGTH.Greatest length from olecranon through tip of
internal condyle (Condylus ventralis).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. W-PROX.Greatest transverse width of proximal articular
surface.
5* D-LENGTH.Length from tip of olecranon to tip of external
cotyla (Cotyla dorsalis).
6. D-PROX.Depth of proximal end from cranial tip of internal
cotyla (Cotyla ventralis) to caudal margin (Margo caudalis) of
shaft of ulna, measured at right angles to the long axis of the
shaft.
7. ECON.Length from external condyle (Condylus dorsalis) through
ventral face of distal end.

13
8. CPTB.Length from carpal tuberosity (Tuberculum carpale)
through lateral face of distal end.
9. ECON-CPTB.Length from external condyle through carpal
tuberosity.
10. EC0N-IC0N.Length from external condyle through internal
condyle.
Radius
1. LENGTH.Greatest length from the radial head (Caput radii)
through distal end of the radius (Extremitas distale radii).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. W-PROX.Greatest transverse width of proximal end.
5. D-PROX.Greatest depth of proximal end.
6. W-DIST.Greatest transverse width of distal end.
7. D-DIST.Greatest depth of distal end.
Carpometacarpus
1. LENGTH.Greatest length from most proximal portion of the
carpal trochlea (Facies articularis radiocarpalis of trochlea
carpalis) through facet for digit III (Facies articularis
digitalis minor).
2. W-PROX.Transverse width proximal end from ligamental
attachment of pisiform process (Processus pisiformes) to dorsal
surface (Facies dorsalis), measured at right angles to the long
axis of the shaft.
2a. W-CARPAL.Transverse width carpal trochlea measured at the
proximal edge of the articular facet.

3.
. D-PROX.Depth of proximal end from tip of process of
metacarpal I (Processus extensoris) through caudal part of carpal
trochlea (Facies articularis ulnocarpalis), measured at right
angles to the long axis of the shaft.
4. L-MCI.Length metacarpal I (Os metacarpal is alulare) from
process of metacarpal I to pollical facet (Processus alularis).
5. D-SHAFT.Depth of midshaft of metacarpal II ( 0s_ metacarpale
ma.jus).
6. W-SHAFT.Transverse width of midshaft of metacarpal II.
7. D-DIST.Greatest depth of distal end, measured across dorsal
edge of facet for digit II (Facies articularis digitalis major).
8. W-DIST.Transverse width distal end from edge of facet for
digit II through facet for digit III.
Furculum
1. LENGTH.Greatest length, measured from furcular process to
scapular tuberosity.
2. D-PROX.Greatest diameter of coracoidal facet.
Femur
1. M-LENGTH.Greatest length from head of femur (Caput femoris)
through medial condyle (Condylus medialis).
2. L-LENGTH.Greatest length from trochanter (Trochanter femoris)
through lateral condyle (Condylus lateralis).
3. W-SHAFT.Transverse width of raidshaft.
4. D-SHAFT.Depth of midshaft.
5 W-PROX.Transverse width of proximal end, measured from the
head of femur through lateral aspect of trochanter, taken at
right angles to the long axis of the shaft.

15
6. D-HEAD.Greatest depth of femoral head.
7. W-DIST.Greatest transverse width of distal end.
8. W-M&LCON.Transverse width of medial and lateral condyles from
(Crista tibiofibularis) to medial border of medial condyle.
9. W-LCON.Transverse width of lateral condyle.
10. W-L&FCON.Transverse width of lateral condyle and fibular
condyle.
11. D-FCON.Greatest depth of fibular condyle.
12. D-LCON.Greatest depth of lateral condyle.
13. D-MCON.Greatest depth of medial condyle.
Tibiotarsus
1. L-LENGTH.Greatest length from interarticular area (Area
interarticularis) on proximal articular surface through lateral
condyle (Condylus lateralis).
2. M-LENGTH.Greatest length from the most proximal portion of
cnemial crest (Crista cnemialis cranialis) through medial
condyle (Condylus medialis). This includes the patella, if
fused, as in loons and grebes.
3. FIBULAR.Length from interarticular area on proximal articular
surface to the most distal point of fibular crest (Crista
fibularis).
4. W-SHAFT.Transverse width of midshaft.
5. D-SHAFT.Depth of midshaft.
6. W-PROX-M.Transverse width of proximal articular surface from
articular facet for fibular head (Facies articularis fibularis)
to medial border of proximal articular surface.

16
7. D-PROX.Depth of proximal end from most caudal edge of medial
articular face (Facies articularis medialis) to the most cranial
point of the cranial cnemial crest.
8. W-PROX-L.Transverse width of proximal end from medial border
of cranial cnemial crest to lateral border of lateral cnemial
crest (Crista cnemialis lateralis).
9. W-DIST-CR.Transverse width of distal end, measured across
cranial portion of condyles.
10. W-DIST-CD.Transverse width of distal end, measured across
caudal portion of condyles.
11. D-MCON.Greatest depth of medial condyle.
12. D-LCON.Greatest depth of lateral condyle.
13. D-ICON.Depth of area intercondylaris.
Tarsometatarsus
1. LENGTH.Greatest length from intercondylar eminence (Eminentia
intercondylaris) through trochlea for digit III (Trochlea
metatarsi III).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. FLEXOR.Intercondylar eminence to middle of tubercle for
tibialis anterior (Tuberositas m. tibialis cranialis).
5 W-PROX.Greatest transverse width proximal articular surface,
measured across dorsal surface.
6. D-MCOT.Greatest depth medial cotyla.
7. D-LCOT.Greatest depth lateral cotyla.

IT
8. D-PROX.Depth from dorsal edge of proximal articular surface
to closest bypotarsal canal (Canalis hypotarsi), or the closest
hypotarsal groove (Sulcus hypotarsi), if no canals are present as
in the Accipitridae.
9. W-HYPOTS.Greatest transverse width of hypotarsus.
9a- W-HYPOTS-C.Transverse width of tuberosity on medial
hypotarsal crest (Crista medialis hypotarsi), in cormorants only.
10. L-HYPOTS.Length of medial hypotarsal crest.
11. D-PROX-L.Depth of proximal end, measured from dorsal edge of
the proximal articular surface through the lateral hypotarsal
crest (Crista lateralis hypotarsi).
11a- D-PROX-M.Depth of proximal end, measured from dorsal edge
of the proximal articular surface through the medial hypotarsal
crest, if the lateral hypotarsal crest is reduced.
12. L-MTI.Greatest length of metatarsal I facet (Fossa
metatarsi I_).
13. D-D-SHAFT.Depth of shaft at cranial edge of distal canal
(Foramen vasculare distale).
14. W-DIST.Greatest transverse width of distal end (if trochlea
are of equal length).
15. TRIII-TRIV.Greatest transverse width from trochlea III
through trochlea IV (if trochlea II is elevated).
16. TRII-TRIV.Greatest transverse width between plantar portion
of trochlea II and plantar portion of trochlea IV.
IT. W-TRII.Greatest transverse width of trochlea II.
l8. D-TRII.Greatest depth of trochlea II.
19* W-TRIII.Greatest transverse width of trochlea III.

20.
18
D-TRIII.Greatest depth of trochlea III.
21. W-TRIV.Greatest transverse width of trochlea IV.
22. D-TRIV.Greatest depth of trochlea IV.
Computer Software
Biomedical Statistical Software, P-Series (Dixon, 198l) was used to
analyze many of the measurements. Programs used included BMDP1D (simple
descriptive statistics), BMDP6D (bivariate plots), and BMDP2M (cluster
analysis). Computations were made at the Northeast Regional Data Center
(NERDC) at the University of Florida, Gainesville.
Nomenclature
Common names have not been used for species. Few nomenclatural
systems have a more unstable, inaccurate, or confusing set of terms than
the American Ornithologists' Union's (1983) list of common names for
birds. I agree with J. L. Peters (l93^:ii)
. . inventing common English names for birds that do not have
them is a waste of time. After all, the primary reason for a
scientific name is to have a name intelligible to scientists
the world over.
Systematic nomenclature generally follows the Checklist of Birds of
the World (Mayr and Cottrell, 1979; Peters, 1931-1951) for Recent species
and Brodkorb's Catalogue of Fossil Birds (1963-1978) for fossil species.
Departures are accompanied by full citation.

Figure 2.1. Schematic diagrams illustrating measurements of the
humerus, coracoid, and scapula. A. Humerus, cranial view. B.
Humerus, ventral view. C. Coracoid, dorsal view. D. Coracoid,
lateral view. E. Humerus, distal end view. F. Scapula, lateral
view. G. Scapula, proximal end view. Figures are not drawn to
scale. Measurements defined in the text.

20

Figure 2.2. Schematic diagrams illustrating measurements of the
radius, ulna, carpometacarpus, and furculum. A. Radius, medial
view. B. Radius, cranial view. C. Ulna, cranial view. D.
Ulna, caudal view. E. Carpometacarpus, ventral view. F.
Carpometacarpus, proximal end view. G. Carpometacarpus, distal
end view. H. Ulna, distal end view. I. Furculum, lateral view.
Figures are not drawn to scale. Measurements are defined in text.

22

Figure 2.3. Schematic diagrams illustrating measurements of the
femur and tibiotarsus. A. Tibiotarsus, caudal view. B.
Tibiotarsus, proximal end view. C. Tibiotarsus, distal end view.
D. Femur, cranial view. E. Femur, proximal end view. F. Femur,
distal end view. G. Femur, caudal view of distal end. Figures
are not drawn to scale. Measurements are defined in the text.

24

Figure 2.U. Schematic diagrams illustrating the measurements of
the tarsometatarsus. A. Dorsal view. B. Lateral view. C.
Plantar view of distal end. D. Distal end view. E. Proximal end
view. Figures are not drawn to scale. Measurements are defined in
text.

26

27
Systematics
I have associated the skeletal elements of the fossil taxa in this
study using the following general criteria (Howard, 1932b):
1. resemblance to other species.
2. size and proportion.
3. relative abundance.
Systematic reasoning and taxonomic practice generally follow Mayr
(1981), when possible. Skeletal elements are first grouped into
similarity classes by means of measurements and qualitative characters.
Each identifiable taxon is placed within a geneology of previously known
species by the hierarchial distribution of shared-derived characters, and
the new species are then integrated into an evolutionary classification.
Paleoecology
Paleoecological methods are described by Shipman (1981). See the
paleoecology section for further comments on the paleoecological methods
used in the excavation of the localities included in this study.
Biochronology and Faunal Dynamics
A database for the examination of the biochronology and the faunal
dynamics of fossil birds of the Neogene of North America was developed
from the following sources: (l) A literature survey of all fossil
localities in North America from the late Arikareean through the Blancan
that have produced fossil birds. This published information was then
emended to reflect the current concepts of geological formations,
geological correlations, mammalian systematics, and the relative and
absolute dating of fossil localities. Verification of all fossil
identifications was made whenever possible. (2) Information from other

28
major unpublished localities was added to this database. This produced a
total of 133 localities from the Neogene of North America (86 published,
47 unpublished) with a record of fossil birds. (3) The occurrences of
fossil taxa were condensed to tabular form to reflect the actual,
documented range of taxa at the family, subfamily, and generic level. A
given taxonomic range reflects the summation of all lower taxonomic ranks
plus material which is only diagnostic to that given level. (4) From
this table, geological ranges were inferred parsimoniously. For example,
if a genus is known from the early Hemingfordian and the late
Clarendonian, I inferred that it also occurred in North America during
the interval between these endpoints. (5) From this table (#3 above),
biochronologically useful species were identified.
Indices of avian faunal dynamics were calculated following Marshall
et al. (1982): the duration (d) of each land mammal age is given in
millions of years, to the nearest tenth, and is based on all available
data; diversity (Si) represents the total number of genera known for each
land mammal age; originations (Oi) are the number of generic first
appearances in a given land mammal age; extinctions (Ei) are last
appearances of a genus in a given land mammal age; running means (Rm)
compensates for time intervals of unequal duration by subtracting the
average of originations (Oi) and extinctions (Ei) for a given age from
the diversity (Si) of that age, or Rm = Si (Oi + Ei)/2; origination
rates (Or) adjust for unequal time intervals by dividing the total number
of generic originations (Oi) occurring during a given land mammal age by
the duration (d) of that interval; similarly, the extinction rate, Er =
Ei/d; turnover rates (T) are the average number of genera that either
originate or go extinct during a given land mammal age, or T = (Or +

29
Er)/2; per-genus turnover rate is the turnover rate adjusted for average
diversity, calculated by dividing the total turnover rate (T) by the
total running mean (P.m). A sampling index was calculated as the number
of localities per Land Mammal Age divided by the duration of that
interval. Marshall et al. (1982) should be referenced for additional
qualifications of each statistic.
Specimens Examined
Fossil specimens included in this study are housed in the
collections of the Florida State Museum (UF), the collection of Pierce
Brodkorb (PB), and the Frick collections of the American Museum of
Natural History (F:AM). I have tried to include all known avian material
from the late Miocene through early Pliocene from Florida within these
collections. Non-diagnostic skeletal elements (vertebrae, phalanges,
etc.) were not considered. Fossil material accessioned into the UF
collections after 01 June 1984 (primarily from Bone Valley) was not
included in this study.
I relied on Recent comparative material in the collections of P.
Brodkorb, Florida State Museum, United States National Museum, American
Museum of Natural History, University of Michigan, and Royal Ontario
Museum.
Abbreviations
Table 2.1 lists the common abbreviations and acronyms used in this
dissertation. Anatomical abbreviations were given earlier.

30
Table 2.1. List of acronyms of institutions and abbreviations of terms
used in the text.
Acronym
Institutions/Collections
AMNH
American Museum of Natural History
F:AM
Frick Collections, American Museum of Natural History
LACM
Los Angeles County Museum of Natural History
MCZ
Museum of Comparative Zoology, Harvard University
PB
Collection of Pierce Brodkorb
ROM
Royal Ontario Museum
UF
Florida State Museum, University of Florida
UM
University of Michigan
UMCP
University of California, Museum of Paleontology,
Berkeley
UNSM
University of Nebraska State Museum
USNM
United States National Museum
YPM
Yale Peabody Museum
Abbreviation
Terms
BMDP
Biomedical Statistical Program, P-series
1. f.
local fauna
M.
muscuius
MA
megannum (or million years)
max.
maximum
min.
minimum
mm.
millimeter
MYBP
million years before present
N
number (of specimens)
NALMA
North American Land Mammal Age

CHAPTER III
GEOLOGY
Biochronology
The Clarendonian and Hemphillian land mammal ages were first
proposed by Wood et al. (19^1) based on the stage of evolution of mammals
from two localities in the panhandle of Texas. Since then, the concept
of each has changed, owing to the increasing knowledge of this time
period in North America. Two additional ages were proposed by Schultz et
al. (1970) for parts of the time intervals covered by the original
definitions: the Valentinian (for the late Barstovian to early
Clarendonian) and the Kimballian (originally proposed as late
Hemphillian; now considered to be early Hemphillian). Neither has been
accepted for use on a continent-wide basis. Rather, each has been
applied only to fossils from the type formations of the proposed ages
(Valentine and Kimball formations). Pertinent references include Schultz
et al. (1970), Tedford (1970), Tedford et al. (in press), Breyer (1981),
and Voorhies (I98U).
The Clarendonian, in the restricted sense of Tedford et al. (in
press) is defined on the
earliest appearance of Barbourofelis and, later in the
interval, Platybelodon, Amebelodon, and Ischyrictis
(Hoplictis).Characterization earliest appearance of
Nimravides, Epicyon (in the Great Plains and Gulf Coast),
Griphippus [=Pseudhipparion], Astrohippus, Nannippus (Gulf
Coast), Macrogenis, Synthetoceras, Hemiauchenia, Megatylopus,
Antilocaprinae (Plioceras and Proantilocapra), latest
occurrence of Eucastor, Brachypsalis, Ischyrocyon, Cynarctus,
Aelurodon, Tomarctus, Hypohippus, Megahippus, Merychippus,
Paratoceras, Miolabis, Protolabis, and probably Ustatochoerus.
31

32
In Florida, common late Clarendonian taxa include Barbourofelis,
Nimravides, Mylagaulus, Pseudhipparion, Amebelodon, advanced species of
Eucastor, Pediomeryx, and Aelurodon.
The early Hemphillian is defined (Tedford et al., in press) by the
earliest appearance of Arvicolinae (limited occurrence of
Microtoscoptes and Paramicrotoscoptes), Pliotomodon, and
Megalonychidae (Pliometanastes), limited occurrence late in
interval of Simocyon, Indarctos, Plionarctos, Lutravus, and
Eomellivora, and earliest appearance, late in interval, of
Mylodontidae (Thinobadistes), Machairodus and the Bovidae
(NeotragocerusT. Characterization earliest appearance of
Dipoides, Pliosaccomys, Pliotaxidea, Vulpes, 'Cams',
Osteoborus, and Cranioceras (YumacerasT!latest occurrence of
Amphicyonidae, Leptarctus, Sthenictis, Nimravides,
Barbourofelis, Epicyon, Pliohippus, Protohippus,
Cormohipparion, Prosthenops, Aepycamelus, Pseudoceras and
Plioceras.
In Florida, common early Hemphillan taxa include Calippus,
Pediomeryx (Yumaceras), Aepycamelus, primitive species of Osteoborus,
Cormohipparion, and Nannippus, and advanced species of Epicyon. The
early sloths Pliometanastes and Thinobadistes are present. The
Eurasiatic immigrants Indarctos and Machairodus appear in the later part
of the early Hemphillian.
The late Hemphillian is defined (Tedford et al., in press) by the
limited occurrence of Promimomys, 'Propliophenacomys',
Plesiogulo, Agriotherium, and Plionarctos and the earliest
appearance of Megalonyx, Ochotona, Megantereon, Felis,
Enhydriodon, and Cervidae. Characterization earliest
appearance of Taxidea, Borophagus, Rhynchotherium, Platygonus,
and Mylohyus, limited occurrence of Pediomeryx, latest
occurrence of Mylagaulidae, Osteoborus, Astrohippus,
Neohipparion, Dinohippus and Rhinocerotidae.
In Florida, common late Hemphillian taxa include advanced species of
Osteoborus, Neohipparion, Nannippus, Pseudhipparion, Teleoceras, and
Hipparion. Also present are Plesiogulo, Megalonyx, antilocaprids,
Agriotherium, Enhydriodon s.l., Megantereon, Machairodus, Rhynchotherium,
Pliornastodon, Dinohippus, Plionarctos, Felis, and cervids.

33
Local Faunas
Love Bone Bed
The Love Bone Bed is located near the town of Archer, Alachua
County, along State Road 24l, in the NW 1/4, SW 1/4, NW 1/4, Sec. 9, T.
11 S., R. 18 E., Archer Quadrangle, U. S. Geologic Survey 7.5 minute
series topographical map, 1969. Excavation and collection of fossil
vertebrates by the Florida State Museum took place from its discovery in
1974 until the quarry was closed during the summer of 1981. This local
fauna originated from the Alachua Formation (Williams et al., 1977).
The Love Bone Bed is considered latest Clarendonian in age (Webb et
al., 198l) Studies published to date on the Love Bone Bed and its
vertebrate fauna include a general overview of the geology and
paleontology of the locality, including a preliminary faunal list (Webb
et al., 1981); studies on the turtles Pseudemys caelata, and Deirochelys
carri (Jackson, 1976, 1978); a description of the rodent Mylagaulus
elassos Baskin (1980); description of the carnivores Barbourofelis lovei
and Nimravides galiani Baskin (1981); descriptions of the procyonids
Arctonasua floridana and Paranasua biradica Baskin (1982); the
description of the ruminant Pedimeryx hamiltoni Webb (1983); and a
population study on the three-toed horse Neohipparion cf. N. leptode
(Hulbert, 1982). Studies on fossil birds include papers on the fossil
herons (Becker, 1985a), a description of a new species of osprey (Becker,
1985b), and one on the fossil anhinga (Becker, ms.). Additional studies
are in progress.

34
Mixson Bone Bed
The Mixson Bone Bed is located approximately 2 miles northeast of
Williston, Levy County, in the NE 1/4, SW 1/4, Sec. 29, T. 12 S., R. 19
E., Williston Quadrangle, U. S. Geological Survey 7*5 minute series
topographical map, 1969 This is the type locality of the Alachua
Formation (Dali and Harris, 1892). There have been many studies on this
site, including papers by Leidy and Lucas (1896), Sellards (1916), Hay
(1923), Simpson (1930), Webb (1964, 1969, in press), and Harrison and
Manning (1983). Additional references are listed in Ray (1957).
Genera in common with McGehee include Calippus, Pediomeryx
(Yumaceras), Aepycamelus, Osteoborus, and Aelurodon (Webb, 1969) The
early Hemphillian rqylodontid sloth Thinobadistes is present. Two other
early Hemphillian index genera are absent from this local fauna, although
they are present in other Florida sitesPliometanastes, and Indarctos.
The first appearance of Indarctos occurs late in the early Hemphillian
and its absence in this local fauna probably has temporal significance.
The absence of Pliometanastes is usually considered an ecological
sampling bias.
McGehee Farm
This locality is almost exactly three miles north of Newberry,
Alachua County, along State Highway 45, Sl/2, NW1/4, Sec. 22, T. 9 S., R.
17 E., Newberry Quadrangle, U. S. Geologic Survey 7.5 series
topographical map, 1968, in northcentral Florida. This locality was
first discovered in 1958 and was extensively collected by the University
of Florida, with support of the Frick. Corporation. This local fauna
originates from the Alachua Formation, and the geology of this locality
is briefly discussed by Webb (1964) and Hirschfeld and Webb (1968). The

35
latter publication includes a preliminary list of the fossil vertebrates
from this local fauna. No faunal revision of this locality has been
undertaken, but several papers treating specific groups have appeared
(sloths: Hirschfeld and Webb, 1968; mylagaulids: Webb, 1966; canids:
Webb, 1969; and protoceratids: Patton and Taylor, 1973).
The faunal composition of this local fauna indicates an early
Hemphillian age (Hirschfeld and Webb, 1968; Webb, 1969; Marshall et al.,
1979), based primarily on the presence of Pliometanastes, the early
Hemphillian megalonychid ground sloth. Typical early Hemphillian
mammalian genera present include Calippus, Pedimeryx (Yumaceras),
Aepycamelus, Osteoborus, and Aelurodon (Webb, 1969).
The birds of this locality were first studied by Brodkorb (1963a)
who reported Phalacrocorax wetmorei, and described two new species
Nycticorax fidens and Ereunetus rayi. Later, Olson (1976) described
Jacana farrandi from here.
Withlacoochee River 4a
The Withlacoochee River 4a local fauna lies approximately 8 km.
southeast of Dunnellon (center of Nl/2, NW1/4, Sec. 30, T. 17 S., R. 20
E., Stokes Ferry Quadrangle, U. S. Geologic Survey 75 minute series
topographical map, 195*+, Marion County), in northcentral Florida. The
fossil vertebrates originate from a massive green clay filling of a
sinkhole in the late Eocene Inglis Formation of the Ocala Group (Webb,
1969, 1973, 1976). This deposit of green clay is being eroded by the
Withlachoochee River; the fossils were collected in approximately 20 feet
of water using scuba equipment. Deposition almost certainly occurred in
a pond environment near sea-level as shown by the fine-grained sediments

36
with articulated fish skeletons, the present elevation, and the marine
taxa present (Becker, 1985a; Berta and Morgan, in press).
Studies on the fossil mammals from here include Webb's (1969) paper
on Qsteoborus ore, Hirschfeld and Webb's (1968) study on Pliometanastes
protistus, Webb's (1973) mention of antilocaprids, and Wolff's (1978)
study of the cranial anatomy of Indarctos. The concurrent range zones of
the first two taxa, and the presence of Indarctos, Machairodus, and
Pseudoceras indicate an age of late early Hemphillian.
Fossil birds from here include a very a small species of Egretta and
an indeterminate species of Buteo (Becker, 1985a). A preliminary faunal
list is included in this paper.
Haile VB
This locality is from the NE 1/4, Sec. 23, T. 9 S., R. 18 E.,
Newberry Quadrangle, U. S. Geologic Survey 7*5 minute series
topographical map, 1968. Auffenberg (1954) discusses the geology of this
locality and describes the abundant material of Gavialosuchus americanus
(Sellards) from this locality. A number of equids are known from this
site including Pliohippus, Cormohipparion, Calippus, and Nannippus.
Haile VI
This locality is in the Nl/2, SW1/4, Sec. 24, T. 9 S., R. 17 E.,
Newberry Quadrangle, U. S. Geologic Survey 7*5 minute series
topographical map, 1968, Alachua County, Florida. It was collected by
the Florida Geologic Survey and Florida State Museum. Auffenberg (1963)
discussed the geology and concluded that this locality represents a
stream deposit. The mammalian fauna includes 'Hipparion', Pseudoceras,
and the type of Mylagaulus kinseyi Webb (1966). A new species of

37
sparrow, Palaeostruthus eurius, was described from here (Brodkorb,
1963a). Reptiles known from here include Deirochelys and Gavialosuchus.
Haile XIXA
This locality is 2.5 miles HE of Newberry, Alachua County, in the NE
1/4, Sec. 26, T. 9 S., R. 17 E., Newberry Quadrangle, U. S. Geologic
Survey 75 minute series topographical map, 1968. It was collected by
the Florida State Museum staff. Vertebrates are mainly aquatic,
including a large amount of skeletal material of Gavialosuchus. Fossil
mammals present include Epicyon, geolocids, Pediomeryx, equids, and
Aepycamelus.
Bone Valley Mining District
The name "Bone Valley" is applied to the phosphate mining district
of central Florida, mainly in Polk County, but also including portions of
adjacent Hillsborough, Hardee, and Manatee counties. The vertebrate
fauna was first described by Sel lards (1916) and later more fully studied
by Simpson (1930). Berta and Morgan (in press) present an account of the
present status of vertebrate paleontology of this area. Much of the
following comes from their paper.
Most of the fossil vertebrates from the Bone Valley area were
obtained from extensive open-pit phosphate mines. In situ collections
are rare and make up approximately 5-10 % of the Florida State Museum
collections, part of the USGS and USNM collections, and virtually none of
the Harvard collections. Avian fossils iron these in situ sites are very
rare. Fossils are more commonly found eroding from spoil piles after an
area has been strip-mined. Except for the in situ collections and a few
intensively collected concentrations, the fossil vertebrates collected

38
from one mine, or a single dragline operating within one mine, are
considered as coming from one broad "locality." The exact geographic
position of these localities range from precisely known (1/4, 1/4, 1/4
section) to generally known (from one minei.e. somewhere within 1 to
10 sections). For a few amateur collections where even the mine is
unknown, the only designation can be the Bone Valley Mining District
(i.e. somewhere within 100 square miles), but the majority of specimens
are identified as coming from one mine. Table 3.1 lists the mines, mine
codes, describes their approximate location, and lists the stratigraphic
codes used.
The age of the Bone Valley fossil vertebrates was much debated from
the 1920s through the 1950s (discussed in Brodkorb 1955a), with the
proposed age ranging from the Miocene through the Pleistocene. Brodkorb
(1955a), using a Lyellian method of percent extinct species in the fauna
and the temporal ranges of three species, suggested that the age of the
Bone Valley avifauna was between the late Miocene and middle Pliocene,
probably early or middle Pliocene (i.e. Clarendonian or Hemphillian; =
late Miocene of current usage).
The majority of fossil land mammals are late Hemphillian in age and
compare well with others of similar age in North America. There is no
evidence to suggest that the fossil birds, which are found in association
with these land mammals, are of a different age. Recently several older
local faunas (Barstovian, Clarendonian, and early Hemphillian) have been
found (MacFadden, 1982; MacFadden and Webb, 1982; Webb and Crissinger,
1984; Berta and Galiano, 1984; and Tedford et al., in press). These
older occurrences are from the Phosphoria, Nichols, Silver City, and
Kingsford mines and to ny knowledge have produced no fossil birds. There

39
are also numerous Pleistocene sites in the Payne Creek Mine (Steadman,
1984), Peace River Mine (=Pool Branch; Webb, 197*0, and Nichols Mine.
The Pleistocene fossil birds from these sites can usually be separated
from geologically older specimens by their association with Pleistocene
fossil land mammals. The Pleistocene fossil birds are not considered in
this study.
There have been many recent studies on non-marine mammalian taxa
from Bone Valley (Baskin, 1982; Berta and Galiano, 1983; Berta and
Morgan, in press; Harrison, 1981; MacFadden and Waldrop, 1980; MacFadden
and Galiano, 1981; MacFadden, 1984; Webb, 1969, 1973, 1983; Webb and
Crissinger, 1984; Wright and Webb, 1984), but no single faunal study of
terrestrial mammals from this local fauna. Reference to the earlier
papers published on the Bone Valley Mining District and its vertebrate
fauna are given by Ray (1957)
Bone Valley taxa which are typical of the late Hemphillian age
include the Eurasian immigrant taxa Agriotherium, Plesiogulo,
'Enhydriodon', and Cervidae. Other taxa present are Rhynchotherium,
Hexameryx, and advanced species of Qsteoborus, Gomphotherium,
Pseudhipparion, Nannippus, and Dinohippus (Berta and Morgan, in press).
Other authors (MacFadden and Galiano, 198l; Berta and Galiano, 1983;
Wright and Webb, 1984) suggest that the upper Bone Valley Formation is
very late Hemphillian because of the presence of taxa equally indicative
of an early Blancan age such as Felis rexroadensis, Meganteron hesperus,
Dinohippus mexicanus, Mylohyus elmorei, Antilocapra (Subantilocapra),
Hemiauchenia, Nasua, and Platygonus.
The birds from Bone Valley have been studied by Brodkorb (1953b,
1953c, 1953d, 1953e, 1955a, 1970). His monograph (1955a) dealt with the

birds then known, primarily the more numerous marine birds. Table 3.2
gives a partial list of birds now known from the Bone Valley Mining
District and notes the taxa (marine) not included here. The majority of
birds here studied have been collected in the last 15 years and mainly
include the rarer, non-marine members of this avifauna.
Manatee County Dam Site
This local fauna originates from a borrow pit south of the Manatee
River in Sec. 30, T. 3k S., R. 20 E., Verna Quadrangle, U. S. Geologic
Survey 7*5 minute series topographical map, 19kk, Manatee County. Like
the nearby SR-6k local fauna discussed below, the Manatee County Dam Site
is essentially an outlier of the classic Bone Valley local fauna. All
three share a similar fauna, geology, and paleoecology. Webb and Tessman
(1968) describe this local fauna and report the presence of one bird,
Phalacrocorax wetmorei.
SR-6k
This locality is located 6 miles east of 1-75 along State Road 6k in
Sec. 35, T. 3k S., R. 19 E., Manatee County, Lorraine Quadrangle, U. S.
Geological Survey 7.5 minute series topographical map, 1973, Florida. It
was discovered by Philip Whisler, of Venice, Florida, in 1983- The
majority of fossil vertebrates are in his private collection, except for
a few speciemens in the Florida State Museum collections. Numerous
fossil vertebrates are recorded from here, including several species of
bony fish, sharks, rays, and several different turtles. Mammals include
odobenids, phocids, Schizodelphis, Megalodelphis, tremarctine ursids, cf.
Agriotherium, felids, Nannippus minor, Neohipparion eurystyle,
Rhinocerotids, and Hexameryx. Based on the presence of Nannippus minor,

Neohipparion eurystyle, and Hexameryx, this locality is late Hemphillian
in age. As for the Manatee County Dam Site, this locality is here
considered an outlier of the classic Bone Valley Formation.
Eustatic Sea-Level Changes
Webb and others (Webb and Tessmann, 1968; MacFadden and Webb, 1982;
Webb, 1984) have proposed a model using the present elevations of fossil
localities with marine or estuarine taxa to reflect the fluctuations of
sea-levels during the late Miocene and early Pliocene in peninsular
Florida. This scheme is predicated on two assumptionsthat the
Florida peninsula had been tectonically stable over the later Cenozoic
and that the sedimentary sequence reflects actual sea-level change. If
these assumptions are valid, then the present elevation of the localities
which contain marine or estuarine taxa should reflect the elevation of
the ocean at the time when these localities were deposited.
The following evidence argues against this model: (l) The relict
Pleistocene shorelines increase in elevation from a low point in southern
Georgia to a maximum elevation in northern peninsular Florida and
gradually lose elevation to the south. This indicates that the northern
part of peninsular Florida has not been stable, and has been
differentially uplifted. Opdyke et al. (1984) argue that this occurred
during the Pleistocene due to the subsurface solution of limestone and
the concomitant isostatic uplift and document an uplift in the magnitude
of 30-50 meters since this time. Most of the high-sea-level localities
are from the northern part of Florida in highly karsted areas.
(2) The elevational changes between most localities could easily be
accounted for by a slight regional dip to the beds. For example, a dip
of 1/20 of 1 degree would account for an elevational difference of 138

1+2
feet over 30 miles (the distance between the Manatee County Dam Site and
the "classic" Bone Valley exposures). This would also account for the
mixture of land and marine vertebrates in these sites, presuming the
tilting is post-depositional.
There is no question that there were eustatic sea-level changes
during this time period, such as the Messinian. But the elevations of
the Florida fossil vertebrate localities are of very doubtful value as
evidence. It should be noted that the rejection of the above hypothesis
does not prevent using the faunal composition of these localities to
determine their relative proximity to the ocean at the time of
deposition. They simply cannot be used as an absolute scale to measure
vertical sea-level changes.

Figure 3.1
Correlation Chart of Included Local Faunas

44

Figure 3.2. Location of Included Local Faunas

46

Table 3.1. A partial list of Bone Valley mines, their mine codes,
approximate location, and the stratigraphic codes commonly used.
Mines
Codes
Township Range
Sections
Chicora
BVC
District Grade
BVPC
see
Payne Creek Below
Estech
BVS
see
Swift Below
Ft. Green
BVFG
32S
23E
2,3,10-14,22-24
Ft. Meade
BVFM
31S
25E
2,13
Gardinier
BVG
32 S
24,24E
unknown
Hooker's Prairie
BVHP
31S
24e
17,18,20,28-30
Kingsford
BVK
31S
23E
3
New Palmetto
BVNP
32S
24E
3
Nichols
BVN
30S
23E
19,28,29
Palmetto
BVP
32S
2UE
9,10,15,16,21,22
Payne Creek
BVPC
32S
24E
13,14,23,24,29-32
Peace River
BVFM
see
Ft. Meade Above
Swift
BVS
31S
24E
14
Tiger Bay
BVTB
31S
24e
12
Stratigraphic Codes
Explaination
0
No
stratigraphic data
1
In place
Hawthorn Fm. dolomitic
2
In place
"lower Bone Valley
Fm."
3
In place
"upper Bone Valley
Fm."
4
In place
Pleistocene sediments
5
Soil zone
: (upper clay)

48
from the late Miocene and early Pliocene
Asterisks denote marine taxa, which are not
are based on previously published works and
Table 3.2 Checklist of birds
Bone Valley Mining District,
included in this study. Taxa
on original identifications.
^Family Gaviidae
*Gavia palaeodytes
*Gavia concinna
Family Podicipedidae
Podiceps sp.
Pliodytes lanquisti
^Family Diomedidae
*Diomedea anglica
Family Phalacrocoracidae
Phalacrocorax wetmorei
Phalacrocorax idahensis
*Family Sulidae
*Morus peninsularis
*Sula guano
*Sula phosphata
^Family Procellaridae
^Family Pelecanidae
*Pelecanus sp.
Family Plataleidae
Eudocimus sp.
Family Ardeidae
Ardea polkensis
Family Ciconiidae
Ciconia sp.
Family Anatidae
Bucephala ossivallis
Family Pandionidae
Pandion sp.
Family Accipitridae
Haliaeetus sp.
Buteo sp.
Family Scolopacidae
Calidris pacis
Erolia penpusilla
Limosa ossivallis
Family Phoenicopteridae
Phoenicopterus floridanus
*Family Haematopidae
*Haematopus sulcatus
*Family Laridae
*Larus elmorei
^Family Alcidae
*Australea granis

CHAPTER IV
SYSTEMATIC PALEONTOLOGY
Table 4.1 lists the non-marine avian taxa now known to occur in
Florida during the late Miocene and early Pliocene. In the following
systematic section, I have tried to present osteological characters which
define the taxonomic groups in a hierarchical fashion. However,
considering the relatively small number of fossil and recent specimens
available for some species, and the restricted geographical area from
which many of the Recent species were collected, I would not be surprised
if some "diagnostic" characters do not hold when a larger number of
specimens are examined.
Order Podicipediformes (Ftirbringer, 1888)
Family Podicipedidae (Bonaparte, 1831)
Tribe Podilymbini Storer, 1963
Characters. Separate canal through hypotarsus for the tendon of
insertion of M. flexor perforatus digiti II (Storer, 1963). Murray
(1967) gives additional osteological characters for the separation of the
different taxa of this family.
Genus Rollandia Bonaparte, 1856
Rollandia sp.
Material. Love Bone Bed local fauna, questionably referred; UF
29670, UF 29673, right coracoids; UF 25815, left coracoid.
Mixson Bone Bed local fauna; F:AM FLA-120-2183, complete right
tarsometatarsus.
49

50
McGehee Farm local fauna; UF 9488, shaft and distal end of right
humerus; UF 12468, right coracoid.
Description. Humerus (UF 9488) poorly preserved; larger and much
more robust than both species of Tachybaptus (dominicus and ruficollis).
Morphology of distal end similar to that of Rollandia rolland, but shaft
more robust. Measurements given in Table 4.2.
Coracoids from the Love Bone Bed are extremely worn and are here
assigned strictly on the basis of their size being near Rollandia (larger
than all species of Tachybaptus and smaller than the smallest males of
Podilymbus podiceps). Shaft similar to, or slightly more robust than,
that of Rollandia rolland. Proximal edge of ventral sternal articulation
flared (as in Rollandia rolland, less so in Tachybaptus). Coracoid from
McGehee Farm local fauna (UF 12488) similar to those from the Love Bone
Bed, except by having a proportionally longer shaft. Measurements are
given in Table 4.2.
Overall length of tarsometatarsus similar to Rollandia rolland
chilensis. Distinguished from Podilymbus by lacking the cranial
expansion of the proximal end of the tarsometatarsus and by having
trochlea II placed lower on the shaft. Distinguished from Podiceps
species by having the shaft less laterally compressed. Tarsometatarsus
differs from that of Rollandia rolland chilensis by having a more deeply
excavated lateral parahypotarsal fossa, a hypotarsus with a smaller
transverse width, a more distinct ridge extending distally from the
hypotarsus, and trochlea II slightly more narrow. Measurements given in
Table 4.3.
Remarks. This species appears to be slightly more robust than the
modern Rollandia rolland chilensis. The lack of a series of modern sexed

51
skeletons, necessary to determine the variability of the characters used
above, prevents the naming of this species as new.
Genus Tachybaptus Reichenbach, 1853
Tachybaptus sp. indet.
Material. Love Bone Bed local fauna; UF 25796, UF 25817, UF 25818,
UF 29671, complete left coracoids; UF 29668, UF 29669, UF 29672, humeral
ends left coracoids; UF 26006, UF 260l4, UF 26017, UF 26019, UF 2966k, UF
29665, UF 29666, UF 29669, complete right coracoids. UF 25773, complete
right femur. UF 29663, proximal end of right tibiotarsus (questionably
referred).
McGehee Farm local fauna; UF 67810, proximal end right tibiotarsus
(questionably referred).
Description. All coracoids are waterworn, abraded, or broken to
varying degrees, making detailed descriptions difficult. All coracoids
near that of Tachybaptus dominicus in size and overall shape. ^Additional
description is not possible.
Femur also near Tachybaptus dominicus in size and general
morphology, but the femoral shaft slightly more gracile. Depression
cranial to the patellar sulcus is absent in fossil. Measurements are
given in Table 4.4.
Both tibiotarsal fragments are questionably referred solely on the
basis of size.
Remarks. The use of generic names follow Storer (1976a). This
material is not diagnostic enough to allow identification to the level of
species.

52
Genus Podilymbus Lesson, 1831
Podilymbus cf. P. podiceps
Material. Bone Valley Mining District, Gardinier Mine; UF 65678,
distal half of left tibiotarsus; Palmetto Mine; UF 21147, distal half of
left tibiotarsus.
Description. Tibiotarsus similar in size and general morphology to
that of Podilymbus podiceps. Distinguished from the tibiotarsus of
Podiceps by having a smooth medial border of the medial condyle (notched
in Podiceps) and a less distinct depressio epicondylaris medialis (very
deep in Podiceps). Differs from that of Podilymbus podiceps as follows:
Shaft of UF 21147 slightly more robust than in males; tubercle slightly
proximal to medial attachment of supratendial bridge better developed;
deeper depressio epicondylaris lateralis than in most specimens. Other
characters within range of variation of modern populations of Podilymbus
podiceps. Measurements are given in Table 4.5.
Podilymbus sp. A
Material. Mixson Bone Bed local fauna; F:AM FLA 66-1115, proximal
end of right tarsometatarsus; F:AM FLA 66-1116, proximal end of left
tarsometatarsus.
Remarks. Both specimens are juvenile, at a similar stage of
ossification, and have identical morphology; they may represent the same
individual. Tarsometatarsal morphology similar to that of Podilymbus
podiceps, but differs by being smaller, having a sharper intercondylar
knob, and a smooth cranio-dorsal border of the medial cotyla (notched in
P. podiceps). Both specimens are too small to correspond to that
expected of Podilymbus cf. P. podiceps from the Bone Valley.
Measurements given in Table 4.3.

53
Tribe Podicipedini Storer, 19^3
Characters. Absence of a separate canal in the hypotarsus for the
tendon of insertion of the M. flexor perforatus digiti II (Storer, 1963).
See Murray (1967) for additional osteological characters.
Genus Podiceps Latham, 1787
Podiceps sp. indet.
Material. Bone Valley Mining District, Payne Creek Mine; UF 21205,
proximal end of left tarsometatarsus.
Remarks. Agrees with Podiceps in hypotarsal configuration (no
extra canal). Waterworn and abraded, and is not identifiable to species.
Similar in size to Podiceps nigricollis or P. o. occipitalis. (£. auritus
much larger).
Tribe indet.
Genus Pliodytes Brodkorb, 1953
Pliodytes lanquisti Brodkorb, 1953
Material. Bone Valley Mining District, Palmetto Mine; PB 299,
complete right coracoid (holotype).
Remarks. This species is known only from the holotype. Brodkorb
(l953e) states that it possesses characters in common with both
Podilymbus and Podiceps but also has its own unique characters. As the
tribes of grebes are defined on tarsometatarsal characters (Murray, 1967;
Storer, 1963), it is not possible to assign this genus to a tribe. As
more fossil material of grebes from the Bone Valley becomes available,
this species should be restudied to determine its generic validity and
relationships to other species.

Remarks on the Family Podicepidadae.
The family Podicipedidae includes 11 fossil and 19 living species.
The earliest certain grebe is Podiceps oligocaenus (Shufeldt), based on a
fragmented left femur (missing the proximal end, distal end badly
abraded) from the Arikareean John Day Formation, Oregon. It is
intermediate in size between the living Podiceps grisegena and P.
nigricollis. Although Wetmore (1937) considers it to be correctly
allocated to genus, next to nothing can be said about its relationships.
Podiceps pisanus (Portis) based on the distal end of a right
humerus, is from the Middle Pliocene (=late Miocene?) of Italy. This
species may also be present at the Hemphillian Lee Creek local fauna
(Olson, ms.). The only other late Miocene species of grebe now known is
Pliodytes lanquisti Brodkorb, discussed above.
There are five species of Pliocene (i.e. Blancan) grebes, all from
North America. Podiceps subparvus (L. Miller and Bowman), from the early
Blancan San Diego Formation, California, is based on a distal end of a
femur. It is approximately the same size as that of the living
Podilymbus podiceps and is now known from additional material. Murray
(1967) in his review of Pliocene grebes, described one new genus and four
new species. Pliolymbus baryosteus Murray, from the Fox Canyon local
fauna, Kansas, of Blancan age, is based on the cranial portion of a
sternum. Murray (1967) states that this is a small grebe with a robust
skeleton but does not suggest any possible relationships between this
species and other living or fossil species of grebes. Podiceps discors
Murray, also from Fox Canyon, is based on a left tarsometatarsus. It is
near the size of Podiceps nigricollis. Murray (1967) also tentatively
refers material from the Hagerman local fauna, the San Diego Formation,

55
California, and the Curtis Ranch, San Pedro Valley, Arizona, to this
species. Aechmophorus elasson, Murray, from the Blancan Hagerman local
fauna, was described on the distal end of an humerus and an associated
left ulna. It is similar to the living A. occidentalis. Podilymbus
ma,i us cuius Murray, also from the Hagerman local fauna, is based on a
nearly complete tarsometatarsus. It is larger than Podilymbus podiceps.
He also tentatively refers material from the Rexroad and Saw Rock Canyon
local faunas to this species.
Pleistocene species include Podiceps parvus (Shufeldt), based on a
lectotype right tarsometatarsus selected by Wetmore (1937) from the
Fossil Lake local fauna, Oregon. It is similar to the living P.
grisegena but is appreciably smaller (Howard, 1946). It is also know
from a well-core in the Tulane Formation of Kern County, California
(Wetmore, 1937). Podiceps dixi Brodkorb is known only from the proximal
end of a right carpometacarpus from Reddick, Florida. It was named after
the Dixie Lime Products Company which owned the quarry in which it was
found (Brodkorb, pers. coram., 1984; etymology omitted in Brodkorb,
1963e). It resembles the living Podiceps auritus, but is somewhat
larger (Brodkorb, 1963e). Podilymbus wetmorei Storer is based on a type
left tarsometatarsus, also from Reddick, Florida, and from a referred
tarsometatarsus and two femora from the Itchtucknee River, Florida. This
species is diagnosed as being the size of Podilymbus podiceps but more
robust. It is only known from these four elements.
The distribution of fossil grebes from the late Miocene through the
early Pliocene of Florida is shown in Table 4.1. Either Podilymbus cf.
P. podiceps or Podiceps sp. from Bone Valley could possibly be
conspecific with Pliodytes lanquisti. These species are only known from

56
a few specimens, none of which are directly comparable. Additional
material will eventually determine the validity of these assignments.
The occurrence of Tachybaptus in Florida is not surprising
considering the present range of Tachybaptus dominicus throughout the
Caribbean. Storer (1976a:124) suggests that T. dominicus has long been
separated from its Old World relatives (T. ruficollis subgroup).
Supporting this view is the lack of an extra canal in the hypotarsus of
T. dominicus (present in T. ruficollis subgroup). Storer (1976a)
considers this a derived character of T. dominicus.
The absence of small grebes the size of Tachybaptus from the Bone
Valley is probably due to a sampling bias toward large specimens or
possibly a general rarity of grebes due to the ecology of the area during
the deposition of the Bone Valley Formation.
The presence of Rollandia in the late Miocene of Florida suggests
that this genus, like Tachybaptus, had a far greater range in the past.
None of the fossil material of this family now known gives any
indication of the higher level systematic relationships of this group.
Postulated relationships include the Gaviidae, Hesperornithiformes,
Sphenisciformes (Cracraft, 1982), based on primarily foot-propelled
swimming adaptations; and the Rhinochetidae and Eurypygidae, on an
apparently unique configuration of the M. longus colli (Zusi and Storer,
1969).
Considering the distribution and diversity of the living species of
grebes (cf. Storer, 1963) it is probable that this group originated in
either North or South America. Supporting this view is the presence of
two genera unique to the Americas (Aechomorphus and Podilymbus) and the
diversity of the fossil record (ten out of eleven fossil species occur in

57
North America). This view is further strengthened by the absence of
grebes in the Early Tertiary European fossil localities which have
otherwise produced a rich aquatic avifauna (St.-Gerand-le-Puy; Cheneval,
1984; Phosphorites du Quercy; Mourer-Chauvire, 1982). The paucity of
knowledge about the evolution of birds in the early Tertiary of South
America makes it premature to decide between a North or South American
origin, although this did not prevent Storer (1967) from suggesting a
South American origin of the family based solely on the diversity of
living species

58
Table 4.1. Checklist of non-marine avian taxa discussed in the text.
Localities where each taxon occurs are given in parentheses Love Bone
Bed (LOV), McGehee Farm (MCG), Mixson Bone Bed (MIX), Bone Valley (BV),
Withlacoochee River 4a (WITH 4a), Manatee County Dam (MD), SR-64, Haile
VB (H5B), Haile VI (h6), and Haile XIXA (H19A).
Class Aves
Order Podicipediformes
Family Podicipedidae
Rollandia sp. (LOV, MIX, MCG)
Tachybaptus sp. (LOV, MCG)
Podilymbus cf. P. podiceps (BV)
Podilymbus sp. A (MIX)
Podiceps sp. (BV)
Pliodytes lanquisti (BV)
Order Pelecaniformes
Family Phalacrocoracidae
Phalacrocorax sp. A (LOV, MCG, H19A)
Phalacrocorax wetmorei (BV, MD, SR-64)
Phalacrocorax cf. idahensis (BV)
Family Anhingidae
Anhinga granis (LOV, MCG, H19A)
Anhinga sp.(BV)
Order Ciconiiformes
Family Ardeidae
Ardea polkensis (BV)
Areda sp. indet.(LOV)
Egretta sp. indet. (LOV, BV)
Egretta subfluvia (WITH 4a)
Ardeola sp. (LOV)
Nycticorax fidens (MCG)
Family Ciconiidae
Mycteria sp. (LOV,MCG)
Ciconia sp. A (LOV)
Ciconia sp. B (MIX,BV)
cf. Ciconia sp. C (BV)
Family Plataleidae
Eudocimus sp. A. (BV)
Plegadis cf. P. pharangites (LOV)
Threskiornithinae, genus et species indet. (LOV)
Order Falconiformes (auct.)
Family Vulturidae
Pliogyps undescribed sp. (LOV)

59
Table 4.1continued.
Family Pandionidae
Pandion lovensis (LOV)
Pandion sp. (BV)
Family Accipitridae
Haliaeetus (?) sp. (BV)
Buteo near B. jamaciensis (WITH 4a)
Aguila sp. TbV)
Accipitrid, genus indet. sp. A (LOV)
Accipitrid, genus indet. sp. B (BV)
Accipitrid, genus indet. sp. C (LOV)
Accipitrid, genus indet. (WITH 4a, BV)
Order Anseriformes
Family Anatidae
Dendrocygna sp. (LOV)
Branta sp. A (LOV)
Anserinae, genus indet. sp. B (LOV)
Anserinae, genus indet. sp. C (LOV, BV)
Anserinae, genus indet. sp. D (BV)
Tadorini, genus indet. sp. A (BV)
Anas undescribed sp. A (LOV, MCG)
Anas size near A. acuta (LOV, MCG)
Anatini, genus indet. sp. A (LOV)
Anatini, genus indet. sp. B (LOV)
Aythya sp. A (BV)
Qxyura cf. 0_. dominica (BV)
Order Galliformes
Family Phasianidae
Meleagridinae, genus indet. (LOV)
Meleagris sp. (BV)
Order Gruiformes (auct.)
Family Gruidae
Grus sp. A (LOV)
Grus sp. B (LOV)
Balearicinae, genus indet. (BV)
Aramornis (cf.) (LOV)

Table 4.1continued.
Family Rallidae
Rallus sp. A (LOV)
Rallus sp. B (BV)
Rallus (cf.) sp. C (LOV)
Undescribed genus (LOV, MCG)
Order Charadriiformes
Family Phoenicopteridae
Phoenicopterus floridanus (BV)
Phoenicopterus sp. A (LOV, MCG)
Family Jacanidae
Jacana farrandi (LOV, MCG)
Family Scolopacidae
Limosa ossivallis (BV)
Erolia penepusilla (BV)
Ereunetes rayi (MCG)
Calidris pacis (BV)
"Calidris" sp. indet. 1 (LOV, MCG, BV)
"Calidris" sp. indet. 2 (LOV)
"Calidris" sp. indet. 3 (MCG)
"Calidris" sp. indet. 4 (LOV)
??Actitis sp. indet. 5 (LOV)
??Arenaria sp. indet. 6 (LOV)
Genus indet. sp. indet. 7 (LOV)
Genus indet. sp. indet. 8 (LOV)
?Philomachus sp. (BV)
Order Strigiformes
Family Tytonidae
Undescribed genus (LOV)
Family Strigidae
Bubo sp. (BV)
Order Passeriformes
Suborder indet. sp. A (LOV)
Suborder indet. sp. B (LOV)
Family Fringillidae
Palaeostruthus eurius (H 6)

6l
Table 4.2. Measurements of humeri and coracoids of the grebes Rollandia
rolland chiliensis (N =6, 2 males, 1 female, 3 unsexed), Tachybaptus
dominicus (N = 7, 4 males, 3 females), and Rollandia sp. from McGehee
Farm local fauna. Data are mean +_ standard deviation and range. (*)
specimen damaged. Abbreviations defined in methods section.
Measurement
R. r. chilensis
T. dominicus
Rollandia sp
Humerus
W-SHAFT
2.67 + 0.15
2.5 2.8
2.39 + 0.15
2.2 2.6
2.8
D-SHAFT
2.U7 + 0.12
2.3 2.6
2.11 + 0.15
1.9 2.3
2.6
W-DIST
5.35 + 0.23
5.1 5.7
5.00 + 0.31
1+.6-- 5.5
5.9
Coracoid
HEAD-FAC
25.1+5 + 0.71
24.3 26.2
21.80 + 1.57
20.1 24.2
*24.6
HEAD-IDA
21+.50 + 0.68
23.3 25.3
21.17 + 1.26
19-4 23.2
*23.3
HEAD-CS
7.03 + 0.23
6.7 7.1+
6.10 + 0.34
5.6"- 6.6
7.6
D-HEAD
2.17 + 0.10
2.17 + 0.17
__
2.0 2.3
2.0 2.4

W-SHAFT
2.30 + 0.18
2.1 2.6
1.94 + 0.21
1.7 2.3
2.1
D-SHAFT
1.65 + 0.10
1.5 1.8
1.29 + 0.09
1.1 1.4
1.7
FAC-IDA
8.53 + 0.38
8.2 9.1
7.44 + 0.4l
7.1 8.3

L-GLEN
1+.55 + 0.16
1+.1+ 5-8
4.31 + 0.25
3.8 4.5
5.3

62
Table 4.3. Measurements of the tarsometatarsi of the grebes Rollandia
rolland chilensis (N = 6, 2 males, 1 female, 3 unsexed), Podilymbus
podiceps (N = 14, T males, T females), Rollandia sp., and Podilymbus
sp. A. from the Mixson's Bone Bed. Data are mean + standard deviation
and range. Abbreviations defined
Measurements
R. r. chilensis
Tarsometatarsus
LENGTH
35.58 + 1.30
33.2 36.7
W-PROX
7.00 + 0.36
6.4 7.5
W-HYPOTS
3.65 + 0.20
3.3 3.8
TRIII-TRIV
5.20 + 0.19
5.0 5.4
TRII-TRIV
5.22 + 0.12
5.1 5*4
W-TRII
1.55 + 0.19
1.3 1.8
W-TRIII
2.52 + 0.15
2.3 2.7
D-TRIII
3.78 + 0.21
3.4 3.9
the methods section.
P. podiceps
R. sp.
P.
40.15 + 2.36
36.3 44.9
36.4

8.09 + 0.57
7.1 8.9
6.7
7.2;
7.5
4.6l + 0.28
4.1 5.1
3.5
4.0;
4.0
6.57 + 0.49
5.8 7.2
5.6

6.24 + 0.49
5-3 7.2
5.2

1.96 + 0.24
1.6 2.4
1.7

2.76 + 0.54
1.5 3.3
2.6

5.00 + 0.4i
4.5 5.6
4.1


63
Table 4.4. Measurements of the femora of the grebes Rollandia rolland
chilensis (N = 6, 2 males, 1 female, 3 unsexed), Tachybaptus dominicus
(N = 7, 4 males, 3 females), and Tachybaptus sp. from the Love Bone Bed.
Data are mean _+ standard deviation and range. (*) Specimen
damaged. Abbreviations defined in the methods section.
Measurements
R. r. chilensis
T. dominicus
Tachybaptus sp
Femora
M-LENGTH
30.73 + 1.55
27.9 -32.0
25.03 + 1.53
23.5 27-8
26.1
L-LENGTH
32.73 + 1.64
29-7 33.5
26.81 + 1.6l
25.1 29.7
27.5
W-SHAFT
2.80 + 0.15
2.6- 3.0
2.53 + 0.22
2.3 2.9
2.6
D-SHAFT
3.18 + 0.24
2.8 3.4
2.67 + 0.30
2.3 3.0
2.6
W-PROX
7.65 + 0.39
7.1 8.3
6.53 + 0.30
6.2 6.9
6.8
D-HEAD
3.30 + 0.24
3.0 3.7
2.74 + 0.20
2.4 3.0
2.8
W-DIST
8.08 + 0.44
7.5 8.4
6.84 + 0.53
5.9 7.4
*6.0
W-M&LCON
5.97 + 0.4l
5.4"- 6.4
4.96 + 0.40
4.4 5-5
*5.l~
D-LCON
6.03 + 0.24
5.7 6.4
4.97 + 0.37
4.3 5.5

D-MCON
4.40 + 0.24
4.1 4.7
3.44 + 0.26
3.1 3.8
3.5

64
Table 4.5 Measurements of the tibiotarsi of the grebes Podilymbus
podiceps (N = 14, T males, 7 females) and Podilymbus cf. _P. podiceps from
the Bone Valley Mining District. Data are mean _+ standard deviation and
range. (*) Specimen damaged. Abbreviations are defined in the methods
section.
Measurements
Podilymbus podiceps
P. cf. podiceps
Tibiotarsus
LENGTH
68.69 + 4.31
61.7 77.0

M-LENGTH
80.17 + 5.27
72.4 90.2

W-SHAFT
4.50 + 0.4l
3.8 5.3
4.6; 4.9
D-SHAFT
3.29 + 0.27
2.8 3.8
3.5; 3.7
W-PROX-M
7.00 + 0.44
6.5 7.9

W-DIST-CR
7.10 + 0.65
6.1 7.9
7.2; *7.4
W-DIST-CD
6.04 + 0.37
5.2 6.5
*6.2; *6.5
D-MCON
6.86 + 0.44
6.1 7.5
*6.9; 7.4
D-LCON
6.76 + 0.49
5.9 7.4
7.0; 7.3
D-ICON
4.29 + 0.33
3.7 4.9
4.4; 4.9

65
Order Pelecaniformes Sharpe,1891
Family Phalacrocoracidae (Bonaparte, 1853)
Genus Phalacrocorax Brisson, 1760
Remarks. The following morphological descriptions are based on the
comparisons of a sample of 5 males and 5 females each of Phalacrocorax
auritus auritus and P. auritus floridanus, and' all available fossil
material.
Phalacrocorax sp. A.
Material. Love Bone Bed local fauna; UF 25735, UF 29661, distal
ends left humeri; UF 29662, distal end right ulna; UF 25877, distal end
left tibiotarsus; UF 25861, distal end right tarsometatarsus (badly
worn, tentatively referred); UF 25933, distal end left tarsometatarsus.
McGehee Farm local fauna; UF II569, complete left coracoid; UF
31779, sternal end left coracoid; UF 12351, distal end right humerus; UF
Ul07, proximal end right ulna; UF 9^92, proximal end right ulna
(questionably referred); UF 31778, proximal end right carpometacarpus; UF
11105, distal end left carpometacarpus; UF 297^6, complete left
tarsometatarsus; UF 31777, proximal end right tarsometatarsus. PB 796U,
proximal end left carpometacarpus.
Haile XIXA; UF 2977^, proximal end left humerus; UF 473^0, proximal
end right carpometacarpus.
Description. Coracoids from McGehee Farm differ from those of both
subspecies of Phalacrocorax auritus (auritus and floridanus) examined,
and from P. wetmorei by having a more elliptical facies articularis
clavicularis, a more robust shaft in relation to the length of coracoid,
the brachial tuberosity more undercut, the impression for the attachment

66
of the coraco-brachialis more distinct, and in medial view, the shaft
more rotated ventrally.
Distal end of the humeri of the fossil species is generally smaller
than that of females of P. a. floridanus, and the shaft more slender
(much smaller than P. a. auritus). Other characters are within the range
of variation of £. auritus. Differs from the humeri of P. wetmorei by
being smaller, having a more shallow fossa brachialis of a different
angle and a much wider attachment of the anterior articular ligament
(=tuberculum supracondylare ventrale).
The two ulnae that are sufficiently perserved to make comparisons
(UF 29662, UF 4107) appear small, about the size of small females of P.
a. floridanus. Characters are within the range of variation of this
species.
Carpometacarpus larger than that of the largest male P. a. auritus.
Process of metacarpal I more nearly square than in that of P. auritus.
Shaft of metacarpal II more robust and angular; anterior carpal facet not
extending up the carpal trochlea as in P. auritus. Pollical facet with a
small papilla .
Tibiotarsus indistinguishable from that of small females of P. a.
floridanus.
Tarsometatarsus description based on UF 29746 (UF 25933 broken and
badly worn; UF 31777 missing distal half, but both agree with UF 29746 in
all discernable characters). Tarsometatarsus short, about equal to that
of females of P. a. floridanus. Transverse width of shaft very narrow.
Lateral face of shaft much more flattened than P. auritus, causing the
posterior intermuscular line to be on the lateral edge of the shaft.
Proximal end narrow, lateral calcaneal ridge more narrow and elongate

67
than in P. auritus. Posterior opening of the lateral proximal vascular
foramen is located lateral to the ridge extending down from the lateral
calcaneal ridge. This ridge not extending as far down the shaft as in P.
auritus.
Remarks. If the carpometacarpus above is correctly assigned to the
same species as is represented by the other skeletal elements, then this
cormorant is quite different in proportions than Phalacrocorax auritus
and related species such as Phalacrocorax wetmorei.
Phalacrocorax wetmorei Brodkorb, 1955
Material. This material is very well represented in the Bone Valley
Fauna. Only Florida State Museum and Florida State Geological specimens
are included in the following referred material section. Material
accessioned into the Florida State Museum collections after 26 April 1984
has not been included in the list of referred specimens; material
accessioned after 5 March 1984 has not been included in the tables of
measurements. Additional material from Bone Valley (type material) is
listed in Brodkorb (1955a).
Manatee County Dam Site.UF 11916, distal end right humerus.
SR-64 local fauna.UF 67805, complete left coracoid; UF 64l43,
humeral end left coracoid; UF 64l44, humeral end right coracoid; UF
64l46, humeral end right scapula; UF 64l45, partial sternum; UF 64l47,
proximal end right tibiotarsus; UF 64l48, UF 64l49, distal ends left
tarsometatarsi.
Bone Valley Mining District, Brewster Mine.UF 61987, humeral end
right coracoid; UF 61988, distal end right tarsometatarsus; UF 65691,
proximal end left ulna.

68
Bone Valley Mining District, Chicora Mine.UF 29733, humeral end
left coracoid.
Bone Valley Mining District, Fort Green Mine.UF 61958, associated
(?) partial skeleton; UF 58062, right quadrate; UF 60047, caudal portion
left mandible; UF 53912, caudal portion right mandible; UF 57248, sternal
fragment with coracoidal sulci; UF 52415, UF 53938, proximal ends right
scapulae; UF 53873, proximal end left scapula; UF 62025, complete left
coracoid; UF 52413, UF 57332, UF 55838, UF 57246, UF 58058, UF 61960, UF
65711, humeral ends right coracoids; UF 53913, UF 55872, UF 57331, UF
58059, UF 58339, UF 61961, UF 65655, humeral ends left coracoids; UF
524l4, UF 61962, UF 65656, sternal ends right coracoids; UF 53934, UF
55831, UF 55875, UF 61963, sternal ends left coracoids; UF 55810, UF
558II, UF 58378, UF 61964, proximal end right humeri; UF 58304, UF 60048,
proximal ends left humeri; UF 53937, shaft left humerus; UF 52410, UF
53914, UF 60050, UF 65657, distal ends right humeri; UF 55865, UF 58060,
UF 58338, UF 58419, UF 58420, UF 60049, distal ends left humeri; UF
55867, UF 57242, UF 58061, UF 60051, UF 65712, proximal ends right ulnae;
UF 55866, UF 61965, proximal ends left ulnae; UF 57243, UF 57247, UF
57249, UF 60052, UF 61966, distal ends right ulnae; UF 52411, UF 55812,
UF 55871, UF 57334, UF 57335, UF 58340, distal ends left ulnae; UF 55813,
UF 55832, UF 55869, UF 60053, UF 65661, proximal ends right
carpometacarpi; UF 52412, UF 55833, UF 55834, UF 57333, UF 584l8, UF
61967, UF 61968, UF 61969, UF 58407, distal ends right carpometacarpi; UF
81959, partial synsacrum; UF 55814, UF 60054, complete right femora; UF
57336, complete left femur; UF 57244, proximal end right femur; UF 53872,
UF 53872, UF 55873, UF 55874, UF 57337, UF 60055, UF 61970, UF 55835,
distal ends right femora; UF 57391, UF 61971, distal ends left femora; UF

69
55868, proximal end right tibiotarsus; UF 55836, UF 57245, UF 60056,
proximal end left tibiotarsus; UF 55804, UF 57250, distal ends right
tibiotarsi; UF 53935, UF 55863, UF 55864, UF 57357, UF 58305, UF 60057,
UF 60058, UF 65713, distal ends left tibiotarsi; UF 55860, nearly
confete left tarsometatarsus; UF 524l6, UF 52417, UF 524l8, UF 55837, UF
58067, proximal end right tarsometatarsi; UF 55870, UF 58421, UF 61972,
proximal ends left tarsometatarsi; UF 53889, UF 53936, UF 55815, UF
57251, UF 58306, UF 58341, distal ends right tarsometatarsi; UF 58342, UF
60059, UF 65658, distal end left tarsometatarsi.
Bone Valley Mining District, Gardiner Mine.UF 58438, caudal
portion right mandible; UF 61998, right clavicle; UF 61999, proximal end
right scapula; UF 62000, proximal end left scapula, UF 65667, complete
left coracoid; UF 58278, UF 58279, UF 58439, UF 58440, UF 62001, UF
65669, humeral ends right coracoids; UF 58277, UF 58280, UF 58446, UF
58447, UF 58469, UF 62002, UF 62003, UF 62004, UF 65668, humeral ends
left coracoids; UF 58448, sternal end left coracoid; UF 58470, UF 62005,
proximal ends right humeri; UF 58441, UF 58471, distal ends right humeri;
UF 58449, UF 58472, UF 62006, distal ends left humeri; UF 58281, UF
58442, UF 62007, UF 6567O, proximal end right ulnae; UF 62008; proximal
end left ulna; UF 57307, UF 58285, UF 58286, UF 65671, distal end right
ulnae; UF 58282, UF 58283, UF 58284, UF 65672, UF 65749, distal end left
ulnae; UF 58443, proximal end right carpometacarpus; UF 58287, proximal
end left carpometacarpus; UF 58303, UF 58282, distal ends left
carpometecarpi; UF 58450, complete left femur; UF 58451, proximal end
left femur; UF 58473, distal end right femur; UF 57311, proximal end left
tibiotarsus; UF 58290, UF 58444, UF 58445, UF 58474, UF 58475, distal
ends right tibiotarsi; UF 58289, UF 58452, UF 58453, UF 62009, UF 62010,

TO
UF 62011, UF 62012, distal ends left tibiotarsi; UF 65673, complete right
tarsometarsus; UF 58478, UF 62013, proximal end right tarsometatarsi; UF
58291, UF 58292, proximal end left tarsometarsi; UF 58293, UF 58476, UF
58U77 UF 6567U, distal ends right tarsometatarsi; UF 57310, distal end
left tarsometatarsus.
Bone Valley Mining District, Palmetto Mine.UF 21058, sternal
fragment with coracoidal sulci; UF 13225, humeral end left coracoid; UF
21143, shaft left coracoid; UF 21115, shaft right coracoid; UF 21146,
distal end right humerus; UF 29734, distal end left humerus; UF 13231, UF
29740, distal ends right ulnae; UF 21091, UF 21120, proximal ends right
carpometacarpi, UF 21068, UF 21072, proximal ends left carpometacarpi; UF
21131, distal end right carpometacarpus; UF 49090, complete right femur;
UF 12352, complete left femur; UF 29735, UF 49091, proximal ends left
tarsometatarsi; UF 12868, distal end right tarsometatarsus.
Bone Valley Mining District, New Palmetto Mine.UF 49691, UF 49692,
vertebrae (questionably referred), UF 49693, complete right femur; UF
49694, complete left tarsometatarsus.
Bone Valley Mining District, North Palmetto Mine.UF 49097, humeral
end left coracoid; UF 49098, humeral end right coracoid.
Bone Valley Mining District, Southwest Palmetto Mine.UF 49093,
humeral end right coracoid.
Bone Valley Mining District, Hookers Prairie Mine.UF 49690,
complete right femur.
Bone Valley Mining District, Kingsford Mine.UF 21186, humeral end
left coracoid; UF 52971, distal end right humerus; UF 13212, distal end
left humerus; UF 21185, proximal end right tibiotarsus.

TI
Bone Valley Mining District, Payne Creek Mine.UF 29741, UF 57304,
humeral ends right coracoids; UF 21203, proximal end left
carpometacarpus; UF 29742, distal end right carpometacarpus.
Bone Valley Mining District, Swift Mine.UF 55883, distal end right
tibiotarsus; UF 17687, distal end right tatsometatarsus.
Bone Valley Mining District, specific locality unknown.UF 61549,
caudal portion left mandible; UF 61550, nearly complete right coracoid;
UF 61553, nearly complete left coracoid; UF 61554, UF 61555, humeral ends
left coracoids; UF 61551, humeral end right coracoid; UF 61552, sternal
end right coracoid; UF 61556, sternal end left coracoid; UF 61557, UF
61558, proximal end right humerus; UF 61559, UF 61560, proximal end left
humeri; UF 61561, distal end right humerus; UF 61562, UF 61563, UF 61564,
UF 61565, UF 61566, UF 61567, UF 61568, UF 61569, distal end left humeri;
UF 61570, UF 61571, proximal end left ulnae; UF 61572, UF 61573, UF
61574, distal ends right ulnae; UF 61575, distal end left ulna; UF 61578,
nearly complete left carpometacarpus; UF 61576, UF 61577, proximal ends
right carpometacarpi; UF 61579, proximal end left carpometacarpus; UF
61596, complete left femur; UF 61580, proximal end left tibiotarsus; UF
61581, UF 61582, UF 61583, UF 61584, distal ends right tibiotarsi; UF
61585, UF 61586, UF 61587, distal ends left tibiotarsi; UF 61588, nearly
complete left tarsometatarsus; UF 61589, UF 61590, UF 61591, UF 61592,
proximal end left tarsometatarsus; UF 61593, distal end right
tarsometatarsus; UF 61594, UF 61595, distal ends left tarsometatarsi.
Bone Valley Mining District, specific locality unknown (FGS
collection).V 7311, proximal end left humers; V 7313, distal end left
humerus; V 7309, distal end left ulna; V 7310, proximal end left
carpometacarpus; V 7312, distal end left femur.

72
Description Scapula within range of variation of that of
Phalacrocorax auritus.
Coracoids appear to be well within the range of variation of those of
P. auritus. The characters used by Brodkorb (1955a: 12) "anterior
intermuscular line situated farther laterad" applies only at the extreme
sternal end of the coracoid. This line does not swing mediad; instead it
curves little as it extends down the shaft. Differs from UF 11569 from
McGehee by characters cited above. Coracoids from SR-64 are
indistinguishable from those of Phalacrocorax wetmorei from Bone Valley.
The two characters of the humerus used by Brodkorb (l955a:12) "the
head of humerus shallower" and "condyles averaging less deep" do not hold
when a large series of specimens are measured (Table 4.7). Brodkorb's
statements that the pneumatic fossa is narrow (slightly) and deeper are
supported. This is especially apparent by having a small, but deep fossa
paralleling the crus ventrale fossae. In P. wetmorei the pneumatic fossa
is perforated by several pneumatic foramina but it is rarely perforated
in P. auritus. In the few specimens of P. auritus in which these
foramina are present, they are very minute. Ligamental furrow (=
ligamental sulcus) does not appear to be relatively longer when compared
against a series of both sexes and subspecies of P. auritus. The distal
end of the humerus of P. wetmorei tends to be narrower, with a more
elongated attachment for the anterior articular ligament (= tuberculum
supracondylare ventrale) ending proximally in a narrower crest than in
the modern specimens of P. auritus. Measurements of ulnae are given in
Table 4.7.
Carpometacarpi of P. wetmorei are about as robust as those of
females of P. f. floridanus. The process of metacarpal I is slightly

73
more produced. Fovea carpalis caudalis deeper in P. wetmorei than in P.
auritus.
The femora are similar, except that the popliteal fossa is generally
less excavated than in P. auritus. Brodkorb's statement that the femur
is longer and narrower than that of P. auritus is not supported by the
larger sample size now available.
Tibiotarsus with no obvious qualitative morphological differences,
but see Table 4.10 for a few minor quantitative differences.
Tarsometatarsus with a lateral face flat, similar to but not as
extreme as, that found on specimens from the Love Bone Bed local fauna.
Other characters similar. See Table 4.11 for measurements.
Remarks. See comments under Family Remarks (below) pertaining to P.
auritus and related species.
Phalacrocorax cf. P. idahensis
Material. Bone Valley Mining District, Palmetto Mine (locality 2 of
Brodkorb, 1955); PB 311, proximal end left ulna.
Remarks. This proximal ulna (PB 311) is larger than that of other
specimens of P. wetmorei presently known from Bone Valley. Since it was
first reported by Brodkorb (1955), it has been additionally damaged in
transport and is now barely diagnostic at the generic level. Unless
additional material becomes available, the status of this enigmatic
record in Florida will probably never be satisfactorily resolved.
Remarks on the Family Phalacrocoracidae.
The cormorants have an extensive fossil record. Brodkorb (1963c)
lists 23 paleospecies; subsequently eight more have been described, or
are in the process of being described. Most have been referred to the

74
genus Phalacrocorax (presently with 26 paleospecies), many without
extensive comparisons with other recent and fossil species to determine
the variability of the of the characters used. It would be desirable to
revise the paleospecies of Phalacrocorax and integrate this extensive
fossil record with the recent species to produce a phylogeny of the
family. There are large amounts of fossil material available, permitting
analysis of variation of several fossil species (e.g. P. wetmorei, P.
oweri, etc.), a recent descriptive osteological study (Ono, 1980), and
papers identifying osteological characteristics and proportions of the
various subgenera of cormorants (Howard, 1932a, 1965; Brodkorb and
Mourer-Chauvirfe, 1984). However, such a revision is beyond the scope of
this dissertation.
Species listed by Brodkorb (1963c) that are not cormorants include
all species of the genus Graculavus (moved to Charadriiformes, Olson and
Parris, ms), Actiornis anglicus (not a cormorant, nor ibis, Olson,
198lb), and Phalacrocorax mediterraneus (Gruiformes, Family
Bathornithidae = Paracrax antigua, Cracraft, 1971)
The earliest cormorants appear in the early Miocene. Phalacrocorax
subvolans Brodkorb from the mid-Hemingfordian Thomas Farm local fauna,
Florida, is known only from a proximal humerus. It is currently under
study (Becker, in prep.). Phalacrocorax marinavis Shufeldt from the
Arikareean John Day Formation, Oregon, is known from a humerus, ulna,
tarsometatarsus, and part of a femur. It is somewhat smaller than P.
carbo but is reported to be allied with this species. Phalacrocorax
miocaenus (Milne-Edwards) from the Aquitanian of Langy, Vaumas, St.-
Grand-le-Puy, and Montaigu, France, is known from most skeletal
elements. It was moved to a new genus Nectornis and is said to share

75
characters with Anhinga (Cheneval, 1984). Phalacrocorax littoralis
(Milne-Edwards) from the Aquitanian of St.-Ge'rand-le-Puy, France, and
from Germany was based on a coracoid and a few other skeletal elements.
It seems to be related to P. aristotelis. Phalacrocorax anatolicus
Mourer-Chauvire' from the lower or middle Miocene (probably Helvetian) of
Bes-Konak, Turkey, was described from a coracoid and most of a forelimb.
It appears to be related to the fossil species P. littoralis, P.
miocaenus, and to the recent P. aristotelis.
Phalacrocorax leptopus Brodkorb from the Clarendonian and
Hemphillian localities of Juntura, Oregon, is based on a coracoid,
tarsometatarsus, and scapula. It is a small species and resembles the
fossil P. littoralis. It is in need of additional comparisons to
elucidate its relationships.
Phalacrocorax femoralis L. Miller from the Barstovian or
Clarendonian Calabasas local fauna, California, is based on most of a
skeleton, preserved in a slab of fine-grained shale. It is the size of
P. penicillatus, but Miller (1929) asserts that this species does not
appear closely related to any living species. Phalacrocorax lautus
Kurochkin and Ganya from the upper Miocene of Moldavia, is based on the
proximal half of a right femur. It appears closest to the living P.
pygmaceus.
Phalacrocorax praecarbo Ammon was described from the upper Miocene
Brown Coal Formation, near WUrttemburg, Germany, on the humeral end of a
coracoid. Brodkorb (1980) has moved Ardea brunhuberi Ammon (figured in
Ammon, 1911), based on a proximal end of a carpometacarpus to this
species and emended the name to P. brunhuberi (Ammon, 1918; cited as 1911
in Brodkorb, 1980). Olson (ms) also moves Botaurites auritus Ammon,

76
based on a cervical vertebra to P. brunhuberi. Phalacrocorax intermedius
(Milne-Edwards) from the Oreleanian of Orleanais, France was described
from a proximal end of a humerus. Phalacrocorax brunhuberi may be
synonymous with P. intermedius; only slightly smaller size and slightly
younger age prevented Brodkorb (1980) from placing it in synonymy with
this species.
Phalacrocorax ibericum Villalta, probably from the Lower Pontian of
Spain, is based on the distal end of a humerus. Villalta states that
this cormorant is smaller than the other Aquitanian cormorants of Europe
(P. littoralis, P. miocenaeus, and P. intermedius) and is close to the
living P. carbo.
Phalacrocorax goletensis Howard from the late Hemphillian or early
Blancan La Goletia local fauna, Michocan, Mexico, is known from a
coracoid (type) and a referred distal humerus. It is possibly ancestral
to P. olivaceus.
Five species, all said to be ancestral to P. auritus, have been
described from the late Hemphillian to Mid-Pleistocene of North America.
Phalacrocorax wetmorei Brodkorb, described from the late Hemphillian Bone
Valley District, Florida, is known from all major skeletal elements. See
additional remarks above. Phalacrocorax kennelli Howard from the Blancan
San Diego Formation, was described on a partial coracoid, a humerus, a
furculum, and vertebrae. It agrees in size with P. pelagicus and P.
penicillatus. In morphology, the fossil species resembles P. auritus or
P. pelagicus (Howard, 19^9). Phalacrocorax idahensis (Marsh) from the
Hemphillian Castle Creek, Idaho, is based on a proximal carpometacarpus.
Murray (1970) referred additional material to and redescribed this
species from the (Blancan) Hagerman local fauna. Phalacrocorax macer

77
Brodkorb from the (Blancan) Hagerman local fauna, Idaho, was originally-
described on a carpometacarpus. Murray- (1970) also redescribed this
species based on additional material. Phalacrocorax macropus (Cope) from
the Mid-Pleistocene Fossil Lake local fauna, Oregon, was described on a
tarsometatarsus. Howard (19^6) referred many other specimens to this
species.
I do not believe that all these species are valid and are correctly
assigned to the ancestral lineage of P. auritus. The morphological
differences among the fossil species are comparable to those among
subspecies of modern P. auritus. It is very possible that like modern
cormorants that show geographic size variations (Palmer, 1962), the
fossil species are simply conspecific geographical variants. If this can
be demonstrated, then each of these fossil species should be maintained
as subspecies of the senior synonym, Phalacrocorax macropus (Marsh).
Valenticarbo praetermissus Harrison from the late Pliocene to early
Pleistocene of Siwalks, India, is based on a 100-year-old plaster cast of
a proximal end of a tarsometatarsus, lacking the hypotarsus. It is very
doubtful that this genus is valid (Olson, ms.). I am unaware of the
relationship of this supposed species.
Pliocarbo longipes Tugarinov from the early Pliocene of the Ukraine
was described from a worn tarsometatarsus and a referred femur. Olson
(ms) notes that although the size and proportions of the tarsometatarsus
are different from typical cormorants, the illustrations are too poor for
even a positive familial verification.
Phalacrocorax destenfanii Regalia from the Mid-Pliocene (Ruscinian?)
of Orciano, Pisano, Italy, was described from most major skeletal
elements. Phalacrocorax mongoliensis Kurochkin from the upper Pliocene

78
of Mongolia is based on the distal epiphysis of a left femur.
Phalacrocorax reliquus Kurochkin from the middle Pliocene of western
Mongolia, is based on the distal epyphysis of a right humerus. It has
the same dimensions as P. pelagicus. The relationships of these
cormorants have not been determined.
Phalacrocorax rogersi Howard from the Pliocene Veronica Springs
Quarry, California, is known only from the type coracoid. It is a large
species and appears close to P. perspicillatus and P. pelagicus.
Phalacrocorax owrei Brodkorb and Mourer-Chauvire' from the lower
Pleistocene of Olduvai Gorge, Tanzania, is known from nearly all skeletal
elements. It has been assigned to the subgenus Stictocarbo, although its
tarsometatarsus is rather similar to P. fuscicollis (subgenus
Phalacrocorax). Phalacrocorax tanzaniae Harrison and Walker from the
Pleistocene Bed II of Olduvai Gorge, Tanzania, was described from a
tarsometatarsus and appears close to P. carbo. Phalacrocorax pampeanus
Moreno and Mercerat from the upper Pleistocene Lujan local fauna,
Argentina, was described from the proximal end of a humerus. It is very
close to P. olivaceus and may be ancestral to this recent species
(Howard, 1965). Phalacrocorax gregorii and P. vetustus DeVis from the
upper Pleistocene localities near Lake Eyre, South Australia, were
described from many elements.

Table 4.6. Measurements of coracoids and scapulae of the
7 males, 7 females), Phalacrocorax auritus floridanus (N
and Phalacrocorax sp. from the McGehee Farm local fauna,
range. Abbreviations are defined in methods section.
Measurements
P. a. auritus
P. a. floridanus
Coracoid
HEAD-IDA
64.35 + 2.79
59-3 67.9
60.87 + 2.83
56.3 65.6
HEAD-CS
22.74 + 1.08
20.1 -23.8
21.16 + 1.14
19.1 23.7
D-HEAD
8.28 + 0.47
7.6 9.0
7.69 + 0.58
6.5 8.7
W-SHAFT
5.26 + 0.33
4.9 6.0
4.84 + 0.46
4.1 5-5
D-SHAFT
6.21 + 0.47
5.5 7.2
5.36 + 0.66
4.1 6.4
IDA-PP
48.25 + 2.6l
43.4 -51.4
46.04 + 2.07
42.5 49.2
L-GLEN
Scapula
12.38 + 0.46
11.5 13.3
11.68 + 0.57
10.6 12.6
1
W-NECK
6.05 + 0.4l
5.4 6.8
5.34 + 0.37
4.3 -6.0
ACR-GLN
16.46 + 0.98
14.7 18.1
15.26 + 0.94
13.4 17.3
cormorants Phalacrocorax auritus auritus (N = l4,
: 18, 9 males, 9 females), Phalacrocorax wetmorei,
Data are mean +_ standard deviation, (n) and
P. wetmorei
Phalacrocorax sp.
64.38 + 2.60 (4)
60.7 66.8
59.3
22.40 + 0.64 (4l)
21.1 23.8
21.8
8.02 + 0.50 (40)
7.4 9-2
7.0
5.26 + 0.38 (7)
4.8 5.8
5-5
5.16 + 0.30 (7)
4.7 5.6
5.8
48.53 + 2.32 (4)
45.1 50.2
43.6
12.29 + 0.46 (44)
11.3 13.2
12.0
*^J
vO
5.83 + 0.38 (12)
5.1 6.4

15.54 + 1.35 (8)
12.5 16.6


Table 4.7. Measurements of the humeri and ulnae of the cormorants Phalacrocorax auritus auritus (N = 14, 7
males, 7 females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), Phalacrocorax vetmorei
from the Bone Valley Mining District, Manatee Co. Dam (M), and Haile XIXA (H), and Phalacrocorax sp. from
McGehee Farm and the Love Bone Bed. Data are mean _+ standard deviation, (N), and range. Abbreviations are
defined in the methods section.
Measurements
P. a. auritus
P. a. floridanus
P. vetmorei
MCD/HXIXA
McGehee
LBB
Humerus
W-SHAFT
8.04 + 0.50
7.2 8.9
7.16 + 0.47
6.3 7.9
8.09 + 0.36 (18)
7.1 8.6
~
7.4
7.8
D-SHAFT
6.85 + 0.44
6.0 7.7
6.16 + 0.37
5.5 6.8
6.22 + 0.33 (l8)
5.4 7.0

6.2
6.3
W-PROX
22.91 + 1.02
21.0 24.6
21.52 + 1.34
19.7 24.1
23.19 + 0.88 (16)
21.5 24.8
20.4
(H)


D-PROX
7.47 + 0.25
7.1 8.0
6.80 + 0.50
5.9 7.7
6.97 + 0.25 (18)
6.3 7.3
6.4
(H)


D-HEAD
11.10 + 0.54
10.5 11.8
10.21 + 0.75
8.8 11.3
10.85 + 0.18 (10)
10.6~- 11.1
9.5
(H)


L-DELTOID
36.22 + 1.69
33.4 38.8
34.31 + 1.92
31.4 37.6
36.89 + 1.75 (11)
34.1 39-3
34.2
(H)


W-DIST
15.94 + 0.52
15.4 16.8
14.79 + 0.91
12.6 16.4
|
15.63 + 0.43 (37)
14.9 16.8
15.4
(M)
15.2
13.8;
14.9
D-DIST
10.88 + 0.59
10.0 11.9
10.15 + 0.68
8.9 11.8
10.29 + 0.36 (35)
9.4 11.1
10.2
(M)
9.9
9.5
9.5
D-ENTEP
7.25 + 0.25
7.2 7.9
7.10 + 0.45
6.3 8.1
7.18 + 0.25 (39)
6.7 7.7
7.3
(M)
6.8
6.4;
6.9

Table 4.7continued
Measurements
P. a. auritus
P. a. floridanus
Ulna
W-PROX
11.95 + 0.4l
11.4 12.7
11.11 + 0.69
9.8 12.4
D-LENGTH
15.74 + 0.83
14.5 17.2
14.19 + 1.12
12.3 l6.4
D-PROX
12.50 + O.58
11.5 13.5
11.31 + 0.90
9.8 13.0
ECON
8.26 + 0.36
7.8 8.9
7.62 + 0.48
6.9 8.5
CPTB
8.95 + 0.27
8.3 9.3
8.37 + 0.46
7.5 9.0
ECON-CPIB
11.15 + 0.50
10.3 11.9
10.38 + 0.49
9-7 11.3
ECON-ICON
6.29 + 0.27
5.8 6.7
5.83 + 0.23
5.4 6.3
i
P. wetmorei
MCD/HXIXA McGehee LBB
11.52 + 0.42 (27)
10.F 12.8
11.1
14.73 + 0.70 (22)
13.3 l6.0
14.1
11.61+0.43 (27) 11.2
10.5 12.3
8.04 + 0.26 (27) 8.0
7.5 8.6
8.46 + 0.37 (27) 8.4
7-7 9.0
10.70 + 0.35 (28) 10.6
9.7 11.3
6.47 + 0.15 (26) 6.2
6.2 6.7
00

Table 4.8. Measurements of carpometacarpi of the cormorants Phalacrocorax auratus auritus (N = 14, 7
males,? females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), Phalacrocorax vetmorei,
from the Bone Valley, and Phalacrocorax species from McGehee Farm and Haile XIXA. Data are mean +_ standard
deviation, (N), and range. Abbreviations defined in methods section.
Measurements
P. a. auritus
P. a. floridanus
P. vetmorei
Haile XIXA
McGehee
Carpometacarpus
LENGTH
69.OI + 2.49
63.8 73.3
66.51 + 2.74
62.6- 70.1
69.6 (1)
'
W-PKOX
7.79 + 0.25
7.4 8.1
7.31 + 0.36
6.7 8.0
6.93 + 0.36 (26)
5.9 7.6
6.8
8.3
W-CARPAL
6.29 + 0.30
5.9 6.9
6.00 + 0.27
5.5 6.4
6.o4 + 0.26 (28)
5.F 6.4
6.1
5.6; 7.2
D-PROX
13.30 + 0.40
12.7 14.1
12.54 + 0.51
11.5 13.3
13.68 + 0.39 (24)
13.1 14.2
12.6
12.9; 16.4
L-MCI
10.64 + 0.47
9.4 11.3
10.06 + 0.68
8.9 11.4
9.87 + 0.38 (23)
9.3 10.5
9.1
9.7; 11.5
D-SHAFT
3.54 + 0.19
3.3 3.9
3.18 + 0.17
2.9 3.4
3.69 + 0.34 (8)
3.3 4.3

4.6
W-SHAFT
4.69 + 0.23
4.3 5.2
4.34 + 0.21
4.0 4.8
4.68 + 0.12 (8)
4.^ 4.9

5.4
D-DIST
5.01 + 0.23
4.6 5.3
4.71 + 0.29
4.0 5.2
4.91 + 0.33 (8)
4.F 5.4

4.8
W-DIST
7.44 + 0.22
7.1 7-7
7.08 + 0.37
6.3 7.7
7.42 + 0.21 (9)
7.2 7.8

7.1

83
Table 4.9 Measurements of the femora of the cormorants Phalacrocorax
auritus auritus (N = 14, 7 males, 7 females), Phalacrocorax auritus
floridanus (N 18, 9 males, 9 females), and Phalacrocorax vetmorei,
from the Bone Valley Mining District. Data are mean +_ standard
deviation, (N), and range. Abbreviations are defined in the methods
section.
Measurements P. a. auritus P. a. floridanus P. wetmorei
Femur
M-LENGTH
54.89 + 2.40
49.5 58.7
52.12 + 3.15
43.2 57.2
55.46 + 1.58 (9)
53.3 59.2
L-LENGTH
56.58 + 2.63
50.1 60.0
54.03 + 2.51
49.4 59.1
57.58 + 1.45 (12)
55.7 61.0
W-SHAFT
6.48 + 0.36
5.6 7.1
5.94 + 0.38
5.2 6.5
6.49 + 0.29 (17)
5.9 7.0
D-SHAFT
8.15 + 0.50
6.9 8.6
7.38 + 0.46
6.7 8.4
8.04 + 0.4l (17)
7.3 9.0
W-PROX
16.13 + 0.60
15.5 17.4
14.56 + O.85
13.2 16.2
15.43 + 0.59 (21)
14.5 16.7
D-HEAD
7.02 + 0.33
6.4 7.6
6.55 + 0.33
6.0 7.3
6.76 + 0.28 (16)
6.3 7.2
W-DIST
15.71 + 0.59
14.9 16.6
14.93 + 0.84
13.3 16.5
15.11 + 0.47 (17)
l4.4 16.D
W-M&LCON
12.21 + O.65
11.3 13.2
11.38 + 0.77
10.0 13.0
11.59 + 0.39 (l4)
10.8 12.3
W-LCON
3.34 + 0.27
3.0 3.9
3.02 + 0.33
2.5 3.6
2.99 + 0.20 (13)
2.7 3.3
W-L&FCON
7.25 + 0.25
6.9 7.7
6.74 + 0.4l
5.8 7.4
6.85 + 0.32 (15)
6.4 7.5
D-FCON
8.81 + 0.47
8.3 9.5
8.40 + 0.50
7.5 9.2
8.68 + 0.40 (17)
8.1 9.4
D-LCON
10.36 + 0.37
9.8 11.1
9.78 + 0.57
8.7 10.5
10.14 + 0.37 (l4)
9.6 10.8
D-MCON
9.05 + 0.34
8.4 9.7
8.59 + 0.48
7.6 9-4
8.68 + 0.33 (18)
8.1 9.4

Table 4.10. Measurements of the tibiotarsi of the cormorants Phalacrocorax auritus aurltus (N = 14, 7
males, 7 females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), Phalacrocorax vetmorei,
from the Bone Valley Mining District, and Phalacrocorax species from the Love Bone Bed. Data are mean
standard deviation
, (N), and range.
Abbreviations are
defined in the methods
section.
Measurements
P. a. auritus
P. a. floridanus
P. vetmorei
Phalacrocorax sp.
Tibiotarsus
FIBULAR
43.18 + 1.99
38.5 45.8
41.26 + 1.94
36.7 43.9
37.7 (1)

W-SHAFT
6.92 + 0.22
6.6 7.4
6.63 + 0.28
6.0 7.1
7.02 + 0.25 (5)
6.6- 7.2

D-SHAFT
5.41 + 0.46
4.8 6.5
4.88 + 0.34
4.3 5.4
5.14 + 0.09 (5)
5.0 5.2

W-PROX-M
11.70 + 0.67
10.7 12.7
10.84 + 0.8I
8.9 12.2
11.70 + 0.80 (5)
10.5 12.7

D-PROX
17.09 + 0.75
16.2 18.4
15.87 + 0.79
i4.1 17.6
17.22 + 0.78 (5)
16.5 18.3

W-PROX-L
11.80 + 0.58
10.6 12.7
11.09 + 0.56
10.1 12.2
10.14 + 0.51 (5)
9.3 10.7

i
00
-o

Table 4.10continued
Measurements
P. a. auritus
P. a. floridanus
W-DIST-CR
11.64 + 0.36
11.06 + 0.50
11.1 12.3
10.2 11.9
W-DIST-CD
11.16 + 0.40
10.47 + 0.50
10.3 11.6
9.8 11.6
D-MCON
10.98 + 0.42
10.41 + 0.60
10.5 11.6
9.2 12.0
D-LCON
10.15 + 0.28
9.76 + 0.45
9.8 10.T
9.0 10.7
D-ICON
6.31 + 0.44
5.70 + 0.34
5.4 6.7
5.1 6.4
i
P. wetmorei
Phalacrocorax sp.
11.57 + 0.49 (25)
10.6- 12.4
11.1; 10.6
11.04 + 0.60 (21)
9.8 12.0
10.7
11.59 + o.4o (38)
10.5 12.3
10.6
10.14 + 0.35 (31)
9.6 10.8
9.4
6.43 + 0.26 (40)
6.1 7.0
6.1; 6.0
00
Ln

Table 4.11. Measurements of the tarsoraetatarsi of the cormorants Phalacrocorax auritus auritus (N = l4, 7
males, 7 females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), and Phalacrocorax
wetmorei, from the Bone Valley Mining District, and Phalacrocorax species from the Love Bone Bed and McGehee Farm
local faunas. Data are mean +_ standard deviation, (N), and range. Abbreviations are defined in. the methods
section.
Measurements
P. a. auritus
P. a. floridanus
P. wetmorei
McGehee
Farm
Love
Tarsometatarsus
LENGTH
62.01 + 2.42
55.2 65.0
59.50 + 2.41
54.7 63.6
64.62 + 3.68 (5)
60.3 69.2
57.7

W-SHAFT
5.94 + 0.37
5.3 6.6
5.83 + 0.44
5.1 6.9
6.17 + 0.38 (13)
5.8 7.0
5.5;
6.2

D-SHAFT
5.24 + 0.24
4.9 5.6
5.27 + 0.52
4.6 6.2
5.53 + 0.44 (13)
4.9 -6.2
5.1;
5.1

W-PROX
12.84 + 0.44
12.0 13.6
12.20 + 0.57
11.0 13.4
12.96 + 0.47 (22)
12.1 14.1
11.5;
, 11.7

D-MCOT
8.70 + 0.38
8.0 9.2
8.44 + 0.29
7.9 9.0
9.01 + 0.58 (21)
8.1 10.0
7.9;
8.1

D-LCOT
7.42 + 0.44
6.8 7.9
7.25 + 0.49
6.5 8.3
7.22 + 0.42 (20)
6.iT 8.3
6.7;
7.2

W-HYPOTS
5.61 + 0.21
5.3 6.0
5.52 + 0.27
5.0 5.9
|
6.38 + 0.65 (9)
5.6" 7.8
5.5;
6.1

L-HYPOTS
9.67 + 0.67
9.1 10.7
9.74 + 0.62
9.0 11.0
10.62 + 1.02 (11)
8.7 12.6
9.6;
9.7


Table 4.11
continued
Measurements
P. a. auritus
P. a. florida:
D-PROX-M
17.66 + 0.54
16.90 + 0.66
l6.6 18.5
15.5 18.3
D-D-SHAFT
4.59 + 0.34
4.27 + 0.26
4.2 5.3
3.6 4.7
W-DIST
14.64 + 0.36
14.11 + 0.6l
14.0 15.1
12.6 14.9
W-TRII
3.94 + 0.17
3.81 + 0.23
3.7 4.2
3.2 4.0
D-TRII
6.20 + 0.29
5.92 + 0.35
5.7 6.7
5.0 6.3
W-TRIII
4.99 + 0.21
4.78 + 0.38
4.6 5.4
4.0 5.9
D-TRIII
6.89 + 0.25
6.48 + 0.34
6.5 7.3
5.7 6.7
W-TRIV
4.02 + 0.22
3.86 + 0.21
3.6 4.3
3.5 4.2
D-TRIV
8.16 + 0.21
7.84 + 0.35
7.9 8.5
7.0 8.4
P. wetmorei
McGehee Farm
Love
17.54 + 0.72 (15)
l6."iT 19.0
16.1; 16.7

4.54 + 0.21 (30)
3.9 4.9
4.1
4.5
14.42 + 0.49 (25)
13.2 15.8
13.6

4.16 + 0.27 (18)
3.Z 4.5
3.9

6.07 + 0.19 (20)
5.7 6.4
5.6

5.21 + 0.28 (30)
4.3 5.8
4.8
4.9
6.89 + 0.29 (29)
6.1 7.6(7)
6.8
6.7
4.04 + 0.29 (23)
3.17 4.5
3.6
4.3
6.65 + 0.27 (25)
5.9 7.2 (?)
6.2
6.4
00

88
Family Anhingidae Ridgway, l88T
Remarks. Skeletal elements of anhingas discussed below may be
distinguished from the Phalacrocoracidae as follows: Humerusby
characters given by Miller (1966) and Martin and Mengel (19T5) Coracoid
head rotated ventrad and mediad, to produce a distinct notch between
head and shaft, when observed from either a ventral or medial view (head
merges smoothly with shaft in the Phalacrocoracidae). Procoracid
expanded and concave cotyla scapularis present.
Genus Anhinga Brisson, 1760
Anhinga grandis Martin and Mengel, 1975
Material. Love Bone Bed local fauna; UF 25739, proximal end right
humerus; UF 25723, UF25725, distal ends right humeri; UF 26000, nearly
complete right coracoid; UF 25873, distal end right tibiotarsus.
McGehee Farm local fauna; UF 11107, distal end right humerus.
Remarks. Elements described in detail in Becker (in prep*). Distal
humeri compare exactly in all features with the type. Coracoid assigned
to this species, as it is of correct size.
Anhinga sp. unknown
Material. Bone Valley Mining District, no specific locality; UF
29781, proximal end left ulna; UF 29780, distal end left ulna.
Remarks. The ulnae are much larger than comparable elements of A.
anhinga and those expected for A. grandis. Parenthetically, the ulna
from Coleman III (UF 1666*0, refered to A. cf. grandis by Ritchie (1980),
is definitely not Anhinga grandis and is not identifiable to a species.
Along with the two specimens from Bone Valley, this Coleman III specimen
represents a much larger species of anhinga which existed approximately

89
4.5 million years later than did A. grandis. Unfortunately, this species
is only known from ulnae and not from more diagnostic elements.
Remarks on the Family Anhingidae.
The earliest record of the Anhingidae is Protoplotus beauforti
Lambrecht based on a skeletal impression, from the late Eocene of
Sumatra. It is presently being restudied and will probably be referred
to a new family (P. V. Richin litt., cited in Olson, ms).
Anhinga pannonica Lambrecht was described from a cervical vertebra
and carpometacarpus from the late Miocene of Tataros, Hungary. Rich
(1972) also assigned another cervical vertebra and a partial humerus from
the late Miocene of Tunisia to this species. The only other anhinga
known from the late Miocene, Anhinga grandis, originally described from
the Hemphillian Cambridge (Ft.- 40) locality, Nebraska, is discussed
above.
The validity of A. laticeps Devis from the late Pleistocene of
Australia is somewhat questionable (Brodkorb and Mourer-Chauvir',, 1982;
Olson, ms.). Anhinga hadarensis Brodkorb and Mourer-Chauvire'", 1982 from
the Upper Pliocene Kadar Hadar member of the Hadar Formation, Ethiopia is
also known from the Omo Basin, Ethiopia, and Olduvai Gorge, Tanzania. It
appears to be the immediate ancestor to A. rufa (Brodkorb and Mourer-
Chauvire, 1982). Two Pleistocene fossil species of anhingas have been
shown to be cormorants. Anhinga parva Devis from Australia was shown by
Miller (1966) to be the cormorant Phalacrocorax melanoleucos and A. nana
Newton and Gadow, from Mauritius was shown by Olson (1975a) to be another
cormorant, Phalacrocorax africanus.

90
Order Ciconiiformes Garrod, 1874 (Auct.)
Family Ardeidae Visors, 1825.
Remarks. The following account briefly establishes the presence and
distribution of the late Miocene and early Pliocene herons in Florida for
the paleoecological and biochronological aspects of this study. More
detailed descriptions and systematic remarks may be found in Becker
(1985a). Systematic nomemclature follows Payne and Risley (1976).
Genus Ardea Linnaeus, 1758
Ardea polkensis Brodkorb, 1955
Material. Bone Valley Phosphate Mining District, Palmetto Mine; PB
380, proximal end of right tarsometatarsus (type), UF 21138, distal end
right tarsometatarsus; Payne Creek Mine; PB 7924, humeral end of right
coracoid.
Remarks. This heron is about the size of A. cinerea and is a rare
member of the Bone Valley avifauna. On the material now known for this
species, it is not possible to determine its relationship to other
members of the genus Ardea.
Ardea sp. indet.
Material. Love Bone Bed local fauna; UF 25939, distal end left
tarsometatarsus, missing trochlea IV.
Remarks. This specimen represents a species of Ardea about the size
of A. herodias occidentalis.
Genus Egretta T. Forster, 1817
Egretta subfluvia Becker, 1985
Material. Withlacoochee River 4a local fauna; UF 19001, right
tarsometatarsus lacking only trochlea IV and hypotarsus.
Remarks. This species is a small heron about the size of Egretta

91
ibis, and is only known from the holotype. The tarsometatarsus is
, proportionally narrower than in other members of this genus.
Egretta sp. indet.
Material. Love Bone Bed local fauna; UF 25759, proximal end of left
carpometacarpus, UF 26082, distal end right ulna. Bone Valley Phosphate
Mining District, Payne Creek Mine; PB 7925, coracoid.
Remarks. These specimens fall within the size range of the living
E. rufescens. Because of the large time interval (4.5 MA) between the
Love Bone Bed and the Bone Valley, it is unlikely that these elements
represent the same species.
Genus Ardeola Boie, 1822
Ardeola sp. indet.
Material. Love Bone Bed local fauna; UF 25940, distal one-third
left tarsometatarsus.
Remarks. Small, similar in size to Ardeola striata. Taxonomic
assignment based entirely on size.
Subfamily Nycticoracinae Payne and Risley, 1976
Genus Nycticorax T. Forster, l8l7
Nycticorax fidens Brodkorb, 1963
Material. McGehee Farm local fauna; UF 3285, complete left femur.
Remarks. See Brodkorb (1963a) for description and remarks.
Remarks on the Family Ardeidae.
Table 4.1 summarizes the distribution of the fossil herons from
Florida. The Love Bone Bed local fauna has produced three heronsa
small Ardeola, a large Egretta, and a very large Ardea. It is most

92
probable that these fossil herons were members of the same diurnal fish
eating guild as are their modern counterparts. It is not suprising that
three herons occur together. As many as 12 species of herons occur
sympatrically in Florida today. They coexist by partitioning resources
such as size and kind of prey, use of habitat (e.g. water depth), and
foraging behavior (Recher and Recher, 1980).
The two herons from Bone Valley could possibly be viewed as marine
specialistsone medium-sized specalizing on small prey and the other
specializing on larger prey, as the living Egretta rufescens and Ardea
herodias occidentalis do today.
Little can be said of the paleoecology of Egretta sp. from the
Withlacoochee River bA but perhaps it was similar to E. ibis in its
ecology, as it is similar to this species in its morphology.
Family Ciconiidae (Gray, l8U0)
Remarks. There are only a few characters on the skeletal elements
preserved here that can distinguish between the genera Mycteria, Ciconia,
and Jabir (sensu Kahl, 1972). Size is of no generic value as evidenced
by species of Ciconia overlapping with species of all other ciconiid
genera.
Characters on the distal end of the tibiotarsus include the internal
ligamental prominence (= medial epicondyle) well-developed in Jabir
(less so in Mycteria and Ciconia); distal end laterally compressed in
Mycteria (less so in Ciconia, except the atypical C. abdimii and C.
episcopus; somewhat compressed in Jabir); distal opening of the tendinal
canal more medially placed in Jabir than in Ciconia or Mycteria;
tubercle slightly elevated above the surface of a strong ridge connecting

93
the lateral condyle and tubercle in Mycteria (much more elevated in
Jabir and Ciconia, excluding C. abdimii and episcopus). Proximal
tarsometatarsus with sulcus ligamentosus sloping gently to the hypotarsus
in Mycteria (sharply sloping and usually deeply excavated in Ciconia and
Jabir). Tibiotarsi and tarsometatarsi of all species of ciconiids (l4
species total) were examined except those of Ciconia nigra,
Ephippiorhynchus senegalensis, and Leptoptilos crumeniferus.
Subfamily ftycteriinae American Ornithologists' Union, 1908
Genus Mycteria Linnaeus, 1758
Mycteria sp. A
Material. Love Bone Bed local fauna; UF 25990, proximal end right
tarsometatarsus (questionally referred).
McGehee Farm local fauna: UF 29^75, proximal end right
tarsometatarsus.
Description. UF 25990 with hypotarsus broken. Tentatively referred
to Mycteria by having the intercotylar prominence sharp and elevated, as
in modern species of Mycteria (prominence more rounded in Ciconia and
Jabir). UF 297^5 agrees with Mycteria by having the intercotylar
prominence highly raised and the sulcus ligamentosus sloping gradually
toward the hypotarsus (sharply notched in Ciconia). There are no
qualitative characters outside the range of variation of the modern
species Mycteria americana, except possibly having the hypotarsus
slightly lower on the shaft.
Remarks. As Table 4.12 shows, this fossil material is within the
range of a modern population of Mycteria americana. However, the
proportions are slightly different, with the proximal tarsometatarsus

proportionally slightly wider than expected (Figure 4.l). It is very
likely that this material represents a species separate from the living
one. Mycteria wetmorei Howard, 1935, from the Pleistocene of California,
was described on the basis of a lower mandible and is said to be larger
than the living Mycteria americana. As there is unstudied post-cranial
material of this fossil species in many U. S. museum collections, this
material should be examined and Mycteria wetmorei revised before
determining the exact systematic position of the fossil material from
Florida.
Subfamily Ciconiinae Gray, l8h0
Genus Ciconia Brisson, 1760
Ciconia sp. A.
Material. Love Bone Bed local fauna; UF 26102, UF 2967^, distal
ends left tibiotarsi; UF 25906, UF 25909, distal ends right tibiotarsi;
UF 259^6, distal end left tarsometatarsus. UF 29675, distal shaft right
tibiotarsus; UF 25900, distal end left tibiotarsus (tentatively
referred).
Description. Size similar to that of Ciconia ciconia. Distal end
of tibiotarsus agrees with those of Ciconia by having the anterior
intercondylar sulcus broad (narrow in Mycteria) and the distal end not
laterally compressed (laterally compressed in Mycteria). Process for the
ligamental attachment above the distal end of the external condyle is
intermediate between papilla-like (as in Jabir) and crest-like (as in
Ciconia). See Figure U.2 for comparisons with other species of Ciconia.
Distal end of tarsometatarsus agrees with Ciconia in having trochlea
II less rotated ventrally (Mycteria more ventrally rotated). UF 29675

95
and UF 25900 are referred here strictly on the basis of size. Both are
too broken and/or worn for further description.
Remarks. See remarks pertaining to Ciconia sp. B., below.
Ciconia sp. B.
Material. Mixson's Bone Bed; F:AM 120-2185, distal end right
tibiotarsus; F:AM 205-3008, distal end left tibiotarsus. Bone Valley
Mining District, Palmetto Mine; UF 21135, distal end right tibiotarsus;
UF 21063, distal end left tarsometatarsus missing trochlea IV
(tentatively referred).
Description. Distal tibiotarsus (F:AM 120-2195) larger than that of
£. mar guar i, similar in size to that of a small Jabir. Agrees with
Ciconia by having the process for the ligamentous attachment above the
distal end of the external condyle ridge-like (papilla-like in Jabir),
and by having the distal opening of the tendinal canal placed more toward
the edge of the bone (more toward the middle of the bone in Jabir).
Agrees with Jabir in having the distal end slightly laterally
compressed.
Distal tibiotarsus (UF 21135) similar to F:AM 120-2185, but with a
robust shaft and having the distal opening of the tendinal canal more
toward the middle of the bone. Very similar to some specimens of Jabir,
but can be distinguished from this genus by having the ridge from the
condyle to tubercle slightly notched, and the ligamentous attachment
above the distal end of the external condyle ridge-like (as in Ciconia).
See Figure 4.2 for comparisons with other species of Ciconia.
Tarsometatarsus fragment assigned on size.

96
Remarks. The presence of this species, species A. above, and
species C. below, shows that the genus Ciconia was much more diverse in
North America than was previously known.
cf. Ciconia sp. C
Material. Bone Valley Mining District, Swift Mine (=Estech); UF
52958, distal end left tibiotarsus. Palmetto Mine; UF 12470, distal end
left tarsometatarsus missing trochlea IV (tentatively referred).
Description. Distal end of tibiotarsus (UF 52958) extremely large,
in the size range of Jabir mycteria. Distal end slightly laterally
compressed (as in Jabir). As in Ciconia, the external ligamental
attachment is ridge-like and distal opening of tendinal bridge is toward
the edge of the bone (contra Jabir). See Figure 4.2 for comparisons
with other species of Ciconia.
Tarsometatarsus assigned here on the basis of size.
Remarks. Although referrable to the genus Ciconia on the basis of
the above characters, this specimen bears a striking resemblance to that
of Jabir mycteria. Were it not for Howard's (1942) study of Ciconia
maltha and Jabir nycteria, I would be tempted to suggest that C. maltha
and J. mycteria are congeneric. Ciconia sp. C. is probably closely
related to the clade which gave rise to Ciconia maltha.
Remarks on the Family Ciconiidae.
Living species of storks have been revised by Kahl (1971, 1972) who
synomized a number of raonotypic genera in Peters (1931). Wood (1983,
1984) has analyzed the phenetic relationships within the Ciconiidae.
Apart from the substantial criticisms which have be made on the use of
phenetics as a basis for classifications (Mayr, 1969; Hull, 1970;

97
Johnson, 1970) I would also note that Wood used an extremely small
sample size (2-6 individuals per species, many of which were unsexed),
which prevented him from adequately accounting for the considerable
sexual variation which exists in storks.
The fossil species of storks are in critical need of revision. All
fossil species now known were described before Kahl's revisions (1971,
1972) which reduced the number of recent genera from 11 to 6. Because of
the plethora of monotypic genera in this family at the time of the
description of the fossil species, workers have tended to underestimate
the amount of morphological differences within a single genus (sensu
Kahl). Because of this, it is now often impossible to determine
relationships between fossil and recent genera.
A number of species originally described as storks have since been
moved to other families or synomized with other species, or both. These
include Pelargopappus stehlini and P. trouessarti (= Amphiserpentarius
schlosseri; Chauvire, in litt., to Olson, ms), Amphipelargus majori
(Ergilornithidae; Harrison, 1981), Palaeopelargus nobilis,
Xenorhynchopsis tibialis, X* minor, and Xenorhynchus nanus (2 are
flamingos, Rich, 1976; 2 in need of restudy, Olson, ms), and Ibis milne-
edwardsi (= Miophasianus altus; Olson, 197*+b). Lists of the other fossil
storks may be found in Brodkorb (1963c). Most of these are in need of
restudy.
In North America, fossil species of storks include Propelargus
olsoni Brodkorb from the Seaboard Airline Railroad local fauna in
Tallahassee of Barstovian age (see comments on the generic status of
Propelargus in Olson, ms), Ciconia maltha Miller now known from the
Blancan through the Rancholabrean of Idaho, California, and Florida, and

98
Mycteria wetmorei Howard from the Rancholabrean of California and
Florida. These species are also poorly defined on their postcranial
skeleton and are in need of revision.
The fossil species of storks in the late Miocene and early Pliocene
of Florida show Mycteria to be long established (Clarendonian to Recent)
and Ciconia to be more diverse in the past. The relationships between
Ciconia maltha, Ciconia sp. C., and Jabir mycteria should be further
investigated.

Figure 4.1. Plot of the transverse width of the proximal end (W-PROX) versus depth of medial cotyla (D-MCOT)
of the tarsometatarsi of the Recent Mycteria americana and Mycteria sp. A from the Love Bone Bed (L) and
the McGehee Farm (M) local faunas. Sex of Recent individuals is indicated by an arrow (male) or cross
(female) on the symbols.
i

MCOT (mm)
1 1-
10-
l
Q
9-
13
j p
14 15
W-PROX (mm)
8
16
nr
17
100

Figure 4.2. Plot of transverse width of the distal end, across the
cranial surface (W-DIST-CR) versus the depth of the area
intercondylaris (D-ICON) of the tibiotarsi of the following
ciconiid species: (l) Ciconia abdimii, (2) C. episcopcus, (3) C.
nigra, (4) C. ciconia, (5) C. maguari, (6) C. maltha from the
Pleistocene of Florida, (7?~Jabir mycteria, (triangles) Ciconia
sp. A from the Love Bone Bedj (squares) Ciconia sp. B from the
Mixson and Bone Valley local faunas, and (open circle) Ciconia sp.
C from the Bone Valley local fauna.

102
i

103
Table 4.12. Measurements of the tarsometatarsi of the storks Mycteria
americana (N = 10, 5 males, 5 females) and Mycteria species A. from the
Love Bone Bed and the McGehee Farm local fauna. Data are mean +_ standard
deviation and range. Abbreviations are defined in the methods section.
(*) Specimen broken.
Measurements Mycteria americana Mycteria sp. A
Tarsometatarsus
W-PROX
15.19 + 0.90
13.9 16.9
17.0;
l6.1
D-MCOT
9.30 + 0.56
8.4 10.3
9-4;
8.6
D-LCOT
8.88 + 0.60
8.1 9.8
9.6;
9.5*
W-HYPOTS
9.66 + 0.62
8.5 10.5
9.5
D-PROX-L
16.57 + 0.94
15.3 17.8
17.2

Table 4.13. Measurements of the tibiotarsi of the Recent storks Jabir
mycteria (N= 13, 2 males, 5 females, 6 unsexed), Ciconia maguari (N = 5,
1 male, 1 female, 3 unsexed), and Ciconia ciconia (N= 6, 2 males, 4
unsexed). Data are mean +_ standard deviation and range. Abbreviations
are defined in the methods section.
Measurement
Ciconia ciconia
Ciconia
maguari
Jabir nycteria
Tibiotarsus
W-SHAFT
8.38 + 0.56
9.50+1
3.48
11.13
+ 0.91
7.6 -
9.0
8.8 :
10.0
9.6 -
- 12.7
D-SHAFT
7.22 + C
5.48
8.26 + (
D.30
10.37
+ 0.64
6.6 -
7.7
7.8 i
3.6
9-3
- 11.1
W-DIST-CR
14.98 +
0.47
17.72 +
0.66
19.21
+ 1.58
14.4 -
15.8
17.0 -
18.5
16.3
- 22.3
W-DIST-CD
11.87 +
0.95
14.56 +
0.35
16.35
+ 1.31
10.7 -
13.0
14.0 -
14.9
13.6
- 18.9
D-MCON
18.97 +
1.13
21.46 +
0.95
25.78
+ 1.76
17.9 -
20.4
20.6 -
22.8
23.4
- 28.7
D-LCON
18.65 +
0.55
21.14 +
0.69
24.37
+ 1.68
18.4 -
19.3
20.4 -
22.1
22.2
- 27.4
D-ICON
11.60 +
0.56
13.52 +
0.6l
16.31
+ 1.07
11.1 -
12.3
12.8 -
14.5
15.1
- 18.3

Table 4.l4. Measureemnts of the tibiotarsi of the fossil storks from Florida; Ciconia sp. A, from the Love
Bone Bed Local fauna, Ciconia sp. B. from the Bone Valley Mining District and Mixson local fauna, Ciconia
sp. C. from theBone Valley Mining District, and Ciconia maltha from various Pleistocene localities in
Florida. Data are mean +_ standard deviation and range. Abbreviations are defined in the methods section.
Compare measurements with those in Table 4.13.
Measurements
Ciconia sp. A
Ciconia sp. B
Ciconia sp. C
Ciconia maltha
Tibiotarsus
W-SHAFT
9.04 + 0.66 (5)
8.1 9*6
9.8; 10.2; 11.8*
12.3
12.60 + 1.09 (6)
11.7 12.7
D-SHAFT
7.80 + 0.53 (5)
7.1 8.5
8.4; 8.3; 9-1
11.3
10.01 + 0.75 (7)
9.0 11.1
W-DIST-CR
14.23 + 0.84 (3)
13.7 15.2
17.5; l6.6; 18.0
19.1
19.47 + 0.90 (6)
18.5 20.6
W-DIST-CD
*11.5
14.2; 13.7; 13.2
15.0
15.98 + 0.68 (5)
15.3 17.0
D-MCON
18.5; 18.9
22.8: 21.7; 22.5
26.4
24.80 + 1.57 (5)
22.9 26.9
D-LCON
l8.8; 18.9
22.6; 20.8; 22.4
25.5
24.62 + 1.67 (5)
23.0 26.7
D-ICON
11.48 + 0.28 (5)
11.2 11.9
13.9; 13.0; 13.8
16.5
15.51 + 0.99 (7)
14.5 17.1
i
105

106
Family Plataleidae Bonaparte, 1838
Subfamily Threskiornithinae (Richmond, 1917)
Genus Eudocimus Wagler, 1832
Eudocimus sp. A
Material. Bone Valley Mining District, Gardinier Mine; UF 60040,
distal end left tibiotarsus. Palmetto Mine, PB 77^+9, proximal end left
tarsometarsus.
Description. Tibiotarsus similar in size and general morphology to
large individuals (males) of Eudocimus albus or E. ruber. Tendinal
bridge as in E. ruber (shorter in E. albus), tubercle well-developed (as
in E. albus, less developed in E. ruber), extensor sulcus more excavated
and flattened than either E. ruber or E. albus, well-developed crest
curves obliquely toward lateral face from tubercle (crest well developed
and oriented parallel to the long axis of the shaft in both E. albus and
E. ruber). Lateral margins of lateral condyle not as developed as in E.
albus or E. ruber. Proximal border of the posterior articular surface
extends farther proximally on the external side of the fossil (border is
usually straight in E. albus and E. ruber; this condition similar in some
individuals of Plegadis spp., especially P. ridgwayi).
Tarsometatarsus similar in size and overall morphology to Eudocimus
albus and E. ruber. I can find no character on the fossil specimen that
is not within the range of variation of these two species.
Remarks. Olson (1981b), in his review of the fossil ibises reported
Eudocimus sp. from a distal end of a tarsometatarsus from Bone Valley. I
have not been able to locate a distal tarsometatarsus in either the
Brodkorb collection or in the FSM collections. I therefore conclude that

107
Olson erroneously reported PB 7749 as a distal end rather than a proximal
end.
Nothing prevents the two specimens listed above from representing
the same species. Despite the apparent difference in size, the Bone
Valley specimens (see Table 4.15) and USNM 181027 from Lee Creek could
easily represent the same species of Eudocimus. The geologic ages are
similar, both fall within the size variation of a single species (see
Figure 4.3, where both specimens fall within the size range of E. albus),
and both USNM 181027 and UF 60040 have the proximal border of the
posterior articular surface extending farther proximally on the external
side. The distribution of this last character in some specimens of
Plegadis species cannot be evaluated with the skeletal material presently
available.
Genus Plegadis Kaup, 1829
Plegadis cf. £. pharangites (A. H. Miller and Bowman, 1956)
Material. Love Bone Bed local fauna; UF 25870, distal end right
tibiotarsus.
Description. Tibiotarsus similar in size to a very small Plegadis
chihi or P. ridgwayi. Distinguished from these species by having a more
distinct tuberculum on the distal end, a more gracile shaft, lateral
condyle merging with the shaft less abruptly, and the surface cranial to
the tendinal bridge more excavated.
Remarks. The material from the Love Bone Bed is not directly
comparable to that of Plegadis pharangites. Skeletal elements of both P,
pharangites and P. cf. P. pharangites are approximately 10 to 12 percent
smaller than P. mexicana (=P. chihi). While I doubt that this material
from the late Clarendonian of Florida is conspecific with that from the

108
Blancan of Texas, I cannot demonstrate any qualitative differences and
have therefore referred this distal end of a tibiotarsus to P.
pharangites strictly on the basis of size.
Threskiornithinae, gen. et sp. indet.
Material. Love Bone Bed local fauna; UF 26003, humeral end right
coracoid.
Description. Humeral end coracoid with ventral portion of head
abraded and with procoracoid broken and missing. Size of a large
Plegadis falcinellus, but also within the range of Eudocimus albus or E.
ruber. The impression for the acrocoracohumeralis ligament is wider in
the fossil specimen than in either Plegadis or Eudocimus. In anterior
view, the shaft appears wider, resembling Eudocimus rather than Plegadis.
Remarks. I can find no consistent characters on the humeral end of
the coracoid which will discriminate with confidence between species of
Plegadis and Eudocimus. This specimen is probably too large to represent
the other ibis (Plegadis cf. P. pharangites) from the Love Bone Bed.
Remarks on the Family Plateleidae.
Olson (l98lb) has recently discussed the fossil record of ibises.
There has been no additional species described since the appearance of
his paper.
I have not been able to find consistent osteological characters
which can separate all specimens of Eudocimus ruber from E. albus.
Considering that most skeletal measurements of these two species overlap
extensively (compare in Table 4.15), and that these "species" interbred
freely when E. ruber was introduced into south Florida, these two taxa
should probably be regarded as two color morphs of the same species. As

109
Olson (1981b) notes, doing this would cast considerable doubt on the
validity of Eudocimus peruvianus Campbell. Campbell (1979) states that
E. ruber differs more from E. albus and E. peruvianus than the latter two
do from each other. It is very likely that when a larger series of
recent E. ruber and E. albus are examined and compared with E.
peruvianus, there will be no consistent ostelogical differences between
these three species.
Similarily, Plegadis falcinellus and P. chihi are very similar
osteologically and may possibly be conspecific (see Table 4.l6 for
measurements; Palmer, 1962; Mayr and Short, 1970 for comments).

Figure 4.3. Plot of transverse width of distal end across the
caudal surface (W-DIST-CD) versus depth of medial condyle (D-MCON)
of the tibiotarsi of the following species of ibis: (l) Eudocimus
albus, (2) E. ruber, (3) Plegadis falcinellis, (4) F. chihi, (3)
P. ridgwayi, () Eudocimus sp. from the Lee Creek 1. f., (b)
Eudocimus sp. A from the Bone Valley 1. f., and (C) Plegadis cf. P.
pharangites from the Love Bone Bed local fauna.

D-MCON (mm)
->l 00 (0 o
4^-j i J 1 1 1 L.
i

112
Table 4.15. Measurements of tibiotarsi, coracoids, and tarsometatarsi of
the ibises Eudocimus ruber (N =8, 2 males, 6 females), Eudocimus albus
(N = 12, 6 males, o females), and Eudocimus species from the Bone Valley
Mining District. Data are mean +_ standard deviation and range.
Abbreviations are defined in the methods section.
Measurements
Eudocimus ruber
Tibiotarsus
W-SHAFT
4.90
+
0.20
4.7
-
5.3
D-SHAFT
4.09
+
0.20
3.8
-
4.5
W-DIST-CR
9.09
+
0.64
8.5
-
10.5
W-DIST CD
6.74
+
0.30
6.4 -
- 1
r.3
D-MCON
9.69
+
0.42
9.3
-
10.6
D-LCON
9.14
+
0.50
8.6
-
10.2
D-ICON
6.04
+
0.27
5.8
-
6.6
Eudocimus albus Eudocimus sp.
5.02 + 0.40
4.5 5.6
5-6
4.53 + 0.53
3.9 5.8
4.7
9.74 + 0.74
8.5 10.5
10.7
7.07 + 0.4l
6.3 7-5
7.5
10.76 + 0.78
9.2 11.6
11.2
10.26 + 0.69
9.1 11.1
10.8
6.70 + 0.60
5.7 7.3
6.9

113
Table 4.15continued
Measurements Eudocimus ruber Eudocimus albus Eudocimus sp.
Coracoid
HEAD-CS
14.46 + 0.51
13.7 15.1
15.52 + 1.02
13.4 17.0
W-SHAFT
5.51 +0.31
5.2 6.1
5.93 + 0.4i
5.0 6.5

D-SHAFT
4.84 + 0.32
4.4 5.4
5.04 + 0.33
4.4 5.4

L-GLEN
9.50 + 0.35
8.9 9.9
10.50 + 0.44
9-8 11.1

Tarsometatarsus
W-PROX
10.76 + 0.83
10.1 12.6
11.47 + 0.98
10.2 12.8
10.3
D-MCOT
5.03 + 0.27
4.7 5.4
5.47 + 0.37
4.7 5.9
5.0
W-HYPOTS
5.24 + 0.37
4.8 5.8
5.65 + 0.54
4.9 6.5
5.5
D-PROX-L
9-33 + 0.49
10.33 + 0.73
9-5
8.8 10.4
9.1 11.3
_

Table 4.l6. Measurements of tibiotarsi, coracoids, and tarsoraetatarsi of the ibises Plegadis ridgwayi
(N = 2, unsexed), Plegadis chihi (N = 7, 2 males, 2 females, 3 unsexed; number may be less due to incomplete
specimens), Plegadis falcinellis (N =10, 7 males, 3 females), Plegadis species (tibiotarsus) and
Threskiornithinae, genus and species indet. (coracoid) from the Love Bone Bed local fauna. Data are
mean _+ standard deviation and range. Abbreviations are defined in the methods section. (*) specimen
damaged.
Measurements
P. ridgwayi
P. chihi
P. falcinellis
Plegadis sp.
Tibiotarsus
W-SHAFT
4.0;
4.3
4.33
+
0.47
4.52
+
0.37
3.7
3.8
-
4.9
3.9
-
5.0
D-SHAFT
3. 4;
3.6
3.80
+
0.35
4.05
+
0.34
3.2
3.5
-
4.3
3.6
-
4.7
W-DIST-CR
7.5;
7.9
8.45
+
0.66
8.63
+
O.56
7.2
7.6
-
9.1
7.8
-
9.7
W-DIST-CD
5.7;
5.5
5.93
+
0.32
6.28
+
0.43
*5.1
5-7
-
6.4
5.5
-
7.0
D-MCON
8.4;
8.4
9.20
+
0.69
9.93
+
O.58
*7-9
8.7
-
10.2
8.9
-
10.8
D-LCON
8.1;
8.3
9.18
+
0.75
9.56
+
0.71
*7.1
8.3
-
10.1
8.5
-
10.5
D-ICON
5.6;
5-5
6.34
+
0.54
6.39
+
0.43
4.9
5.8
-
'7.0
5.8
-
7.0
114

Table 4.l6continued
Measurements
P. ridfiwayi
P. chihi
Coracoid
HEAD-CS
12.6; 12.7
13.42 + 0.78
12.3 14.3
W-SHAFT
5-4; 5.+
5.37 + 0.50
4.6 5.9
D-SHAFT
3.7; 4.0
4.45 + 0.38
3.9 4.9
L-GLEN
8.5; 9.0
8.97 + 0.33
8.5 9.4
i
P. falcinellis
Threskiornithinae
Ren. et sp. indet,
13.99 + 0.86
12.2 15.0
16.2
5.52 + 0.54
4.6 6.3
6.1
4.54 + 0.38
4.0 5.1
5.0
9.54 + 0.69
8.1 10.3
11.4
115

116
Order Accipitriformes (Vieillott, l8l6) (Auct.)
Family Vulturidae (illiger, l8ll)
Remarks. Vulture tarsometatarsi and tibiotarsi, the only elements
considered here, are characterized by Cracraft and Rich (1972).
Genus Pliogyps Tordoff, 1959
Emended Generic Diagnosis. The tarsometatarsus of Pliogyps differs
from that of other living and fossil genera of vultures in having a
proportionally large trochlea for digit III, the proximal articular
surface wide and deep in comparison to the length of the bone, a
generally columnar form, with symmetrical lateral and medial flaring both
proximally and distally; shaft wide in comparison to length of bone;
hypotarsus merging distally with shaft by means of a broad, rounded ridge
(as in Vultur, Breagyps, Gymnogyps, and Geranogyps; narrow in Coragyps,
Cathartes, and Sarcoramphus). This last character may be strictly size
dependent and if so, not of value as a generic character.
Remarks. Two other proposed generic characters (Tordoff,- 1959)
appear to be variable within a species, and are of no generic value.
They are the shaft less deeply and extensively excavated anteriorly and
groove of trochlea for digit III ending anterioproximally in a shallow,
but distinct pit.
Pliogyps species
Referred Material. Love Bone Bed local fauna; UF 25719, fragment of
shaft of left humerus; UF 25886, distal end left tibiotarsus; UF 25952,
complete right tarsometatarsus missing a small portion of hypotarsus.
Diagnosis. Tarsometatarsus relatively more robust than any living
or fossil genus of vulturid except Pliogyps. Tarsometatarsus
distinguished from Pliogyps fisheri Tordoff 1959 by smaller size, by
having a narrow ridge extending from the hypotarsus farther down shaft

117
(caudal view), by having the sulcus extensorius more excavated and
extending farther down shaft (cranial view), and by having the shaft and
trochlea III proportionally less deep. Pliogyps sp. from the Love Bone
Bed has a proportionally greater power-arm ratio (defined below) than P.
fisheri.
Description. Tarsometatarsus proportionally different from all
other vultures except for Pliogyps fisheri (see Figures 4.4 4.6). The
proximal end wide and deep. In cranial view, the proximal vascular
foramina large and approximately equal in size. Papilla for the
attachment of M. tibialis cranialis large, rounded, and in two parts.
Sulcus extensoris extending down the shaft to distal foramen, with a
sharp lateral border. Distinct intermuscular line extending obliquely
through this sulcus (separating attachments for the extensor digitorum
brevis pars hallucis and extensor digitorum brevis pars adductor-extensor
digiti IV; Jollie 1976-1977:243). In caudal view, the tarsometatarsus
has a long ridge extending down the shaft from the hypotarsus,
terminating in an intermuscular line which then extends to the level of
the articular facet of metatarsal I. See Table 4.17 for measurements.
Distal end of tibiotarsus with a broad extensor sulcus. Slight
projection of bone on lateral surface (approximately 4 cm from distal
end) for attachment of the fibula. Intercondylar sulcus broad; with
external condyle merging evenly into it. In distal end view, the
intercondylar sulcus is not symmetrical, with the lateral border sloping
gradually, and the medial border sloping abruptly, up from the base of
the intercondylar sulcus (symmetrical or U-shaped in Coragyps,
Sarcoramphus, Breagyps, Gymnogyps; unsymmetrical or Pliogyps-like in
Cathartes). See Table 4.17 for measurements.

118
Humerus fragment tentatively referred.
Remarks. Table L.l8 lists indices for flexion of the intertarsal
joint (power-arm ratio of Jollie, 1976-1977; [(FLEXOR * LENGTH) X 100)
and for robustness of shaft [W-SHAFT - LENGTH) X 100]. These indices
show that Pliogyps sp. from the Love Bone Bed has an average flexion
ratio but a very broad tarsometatarsus. If this is considered with the
distinct, excavated muscle attachments discussed above, it is suggestive
of a powerful pelvic limb, more so than in most vulturids, reminiscent of
some accipitrids. Certainly the interpretation of this is very tentative
(see Fisher, 19^5; Becker, 1985b), but possibly Pliogyps sp. from the
Love Bone Bed was more rapacious than other living or fossil vultures, as
rapacious birds tend to have a higher flexor ratio than do non-rapacious
birds of equal size. As additional fossil material of this species
becomes available, this suggestion should be examined further.
Campbell and Tonni (1983) develop further ideas of Prange et al.
(1979) on the correlation between the cross-sectional area of the
tibiotarsus of a given species and its live weight. They empirically
determined the following regression
log Y = 2.51* log X 0.19906
where Y is the live body weight (gms) and X is the least shaft
circumference of the tibiotarsus (mm). The correlation coefficient for
this relationship is O.986, showing that the predictions of the live
weight should be very accurate. The least shaft circumference of
Pliogyps species (UF 25886) from the Love Bone Bed is 32 mm yielding a
predicted weight of 5*2 kg. Sarcoramphus papa, which has a
tarsometatarsus approximately as long as this species, weights between
3.0 and 3.75 kg (5 individuals, Brown and Amadon, 1968). This supports

119
Tordoff's (l959:3^1ff) conclusions that Pliogyps is relatively a very
heavy-bodied, short-legged bird.
The intergeneric relationships of living and fossil vultures in
general, and Pliogyps in particular, are very difficult to determine,
owing to a paucity of pre-Pleistocene fossil specimens required to
determine believable character states. Pliogyps shares tarsometatarsal
characters with Sarcoramphus (anterior fossa continuing down shaft to the
distal foramen, a similar size of the distal foramen and a similar shape
of the hypotarsal ridge). Characters in which Pliogyps differs from
Sarcoramphus include the degree of elevation of trochlea III (proximal
border merging smoothly with shaft (plantar surface) in all modern
skeleton of Sarcoramphus examined), and the amount of excavation of the
lateral parahypotarsal sulcus. Pliogyps also shares characters with
Vultur and Gymnogyps including the lateral side of the area proximal to
trochlea IV being inclined, and the excavation of the anterior fossa
extending to the distal foramen (although to a lesser degree than in
Sarcoramphus).
Remarks on the Family Vulturidae.
Lign (1967) and Rea (1983) discuss the relationships of the family
Vulturidae with other avian families. There have been many fossil
species described as vultures; these are listed in Brodkorb (1961+b).
Those which have been subsequently moved to other families and orders
include: Lithornis vulturinus Owen, volant paleognath (Olson, ms.);
Palaeogyps prodromus Wetmore and Meocathartes grallator Wetmore, Family
Bathornithidae (Olson, ms.). Olson (ms.) also considers several other
"vultures", not to be sufficently diagnostic to be maintained in this

120
family. They include: Eocathartes robustus Lambrecht, (too crushed);
Teracus littoralis Aymard, (incertae sedis; Olson 1978); and Phasmagyps
patritus Wetmore.
This' leaves Diatropornis ellioti (Milne-Edwards) and Pleiocathartes
europaeus Gail lard from the late Eocene and early Oligocene of the
phosphorites du Quercy as the oldest certain records of vultures in
Europe. Also known is Plesiocathartes? gaillardi from the early Miocene
of Spain and an unreported specimen of a large member of this family from
the early Oligocene of Mongolia (Kurochkin, in litt., to Olson, ms.).
In South America, the oldest current record of this family is
Dryornis pampeanus Moreno and Mercerat, from the Monte Hermoso Formation
in Argentina. This species was originally described as a species of
phorusrachid, but was later moved to the family Vulturidae (Brodkorb,
1967). Tonni (1980) states that it is close to the living Vultur. I
also note that Campbell (1979) synonymzed the fossil Vultur patruus
L&inberg, from the Pliocene of Tarija, Bolivia, with the living Vultur
gryphus.
In North America, the oldest records of vultures are Sarcoramphus
kernensis from the late Miocene (mid-HemphiIlian) of Kern River,
California, the species of Pliogyps from the Love Bone Bed local fauna,
discussed above, and an unreported species of vulturid from the mid-
Barstovian Sharkstooth Hill local fauna (Howard, in litt., 1984).
Sarcoramphus kernensis was originally described, and is still only known,
from a crushed distal end of a humerus. It was compared only with S.
papa (then Vultur papa). It should be re-examined and differentially
diagnosed to determine its correct generic position. There are several
described species of vultures from the Pliocene and Pleistocene of North

121
America which are listed in Brodkorb (1964b). Exactly how many of these
species are valid remains to be determined.
The earliest certain records for this family are therefore of late
Eocene and early Oligocene age in Europe and Asia; of mid to late Miocene
age in North America; and of Pliocene age in South America. The fossil
record would therefore suggest that this family had an Eurasian origin,
then invaded North America, and more recently South America.
An examination of the characters and proportions of the
tarsometatarsus and their variation in living and Neogene fossil species
suggests that the genera of vultures are oversplit. I agree with Mayr
and Short (1970) that Vultur Linnaeus and Gymnogyps Lesson (including G.
ampulus, G. howardae, and G. californianus) are congeneric. As Figure
4,4 and 4.5 show, the proportions of the tarsometatarsus of these species
differ little from each other. I would also tentatively include Breagyps
L. Miller and Howard and Geranogyps Campbell in the genus Vultur
Linnaeus, as they too are large vultures with similar tarsometatarsal
proportions.
However, I strongly disagree with Mayr and Short's (1970) opinion
that Pliogyps Tordoff should be included within the genus Vultur. The
tarsometatarsi of both species of Pliogyps differ greatly from Vultur in
proportions (Figure 4.4 and 4.5).
Therefore, on tarsometatarsal proportions, I would recognize the
following genera: Cathartes Illiger, Coragyps Geoffroy, Sarcoramphus
Dumeril, Pliogyps Tordoff, and Vultur Linnaeus, (including Gymnogyps,
Breagyps, and Geranogyps). If other skeletal elements show a similar
trend, I would urge adoption of these genera and their use as described
above.

122
Anti 1 lovultur Arredondo was described from a late Pleistocene cave
deposit in Cuba. The type is a 42.5 mm tarsometatarsal fragment which
lacks both the proximal end and the distal one-half of the bone. It is
also known from a referred distal portion of a humerus, a trochlea IV of
a tarsometatarsus, and a single cervical vertebra. The description
(Arredondo, 1976) does not clearly differentiate this material from that
of the genus Vultur.

Figure 4.4. Plot of greatest length (LENGTH) versus width of proximal end (W-PROX) of the tarsometatarsi
of the following species of vultures: (l) Vultur gryphus, (2) Sarcoramphus papa, (3) Gymnogyps
californianus, (4) Coragyps atratus atratus, (5) Cathartes aura, (A) Pliogyps fisheri, (B) Pliogyps
sp. from the Love Bone Bed 1^ fTj [c] Breagyps clarki, (D) Geranogyps reliquus~ (e) Gymnogyps howardae,
and (F) Gymnogyps amplus.
i

32
30
28
E 26
E
X
24
22-
oc
Q.
I 20-
£
18-
16
14
12
104-
60
\
5
70
80
i 1 1 1 1
90 100 110 120 130
LENGTH (mm)
140
124

Figure 4.5. Ratio diagram (after Simpson et al. i960) of measurements of the tarsometatarsi of the
following species of vultures: (A) Cathartes aura, (B) Coragyps atratus atratus, (C) Pliogyps sp. from
the Love Bone Bed 1. f., (D) Pliogyps fisheri, (E) Sarcoramphus papa, (f1 Gymnogyps californianus, (G)
Geranogyps reliquus, (H) Gymnogyps howardae, (i) Vultur gryphus, and (J) Breagyps clarki. Measurements,
defined in the methods section, are abbreviated as follows: (l) LENGTH, (2; W-PROX, (3) D-PROX-L, (4)
FLEXOR, (5) W-SHAFT, (6) D-SHAFT, (7) W-TRIII, (8) D-TRIII, (9) W-DIST.
1

.20 .15 .10 .05 0.0 + .05 .10 .15 .20
1 1 1 1 I I I L-
126

Figure 4.6. Pliogyps sp. A, B. UF 25886, distal end left
tibiotarsus. A. Caudal view. B. Cranial view. C, D. UF
right tarsometatarsus. C. Plantar view. D. Dorsal view.
A, B = 3 cm.; C, D = 5 cm.
25952,
Scale

128

129
Table 4.17. Measurements of the tibiotarsi and tarsometatarsi of the
vultures Coragyps atratus atratus (N = l6, 8 males, 8 females), Pliogyps
fisheri (holotype) from the Rexroad local fauna, and Pliogyps undescribed
species from the Love Bone Bed. Data are mean _+ standard deviation and
range. Abbreviations are described in the methods section.
Measurements Coragyps a. atratus Pliogyps fisheri Pliogyps sp.
Tibiotarsus
W-DIST-CR
12.79 + 0.4l
12.1 13.6

~
D-MCON
13.67 + 0.4l
13.2 l4.6


Tarsometatarsus
LENGTH
84.43 + 1.54
80.4 87.I
94.0
86.6
W-PROX
15.11 + 0.53
14.1 16.2
21.9
21.1
D-PROX
11.71 + 0.42
11.1 12.4


W-DIST
16.59 + 0.57
15.6 17.6
33.0

W-TRIII
6.43 + 0.21
6.0 6.7
9.6
9.2
D-TRIII
9-99 + 0.32
15.2
13.5

130
Table 4.18. Ratios of intertarsal flexion and tarsometatarsal robustness
of species of vultures. Flexion ratio is calculated by (FLEXOR + LENGTH)
X 100; Robustness ratio calculated by (W-SHAFT + LENGTH) X 100.
Increasing values are correlated with increasing force of flexion and
increasing robustness, respectively. (*) approximate.
RATIO
Species
LENGTH
FLEXOR
W-SHAFT
FLEXION
ROBUSTNESS
P. undesc. sp
86.6
14.9
10.8
17.2
12.4
P. fisheri
94
14.5
11.4
15.4
12.1
S. papa
*91
*16
*10.5
*16.5
*10.8
G. californica
*116
*22
*13.5
*19.0
*11.6
V. gryphus
*126
*22
*14
*17.4
*11.1
C. atratus
88.4
12.8
7.2
14.5
8.1
C. aura
65.9
11.3
7.5
17.1
11.4

131
Family Pandionidae (Sclater and Salvin, 1893)
Remarks. The following account briefly establishes the presence and
distribution of the late Miocene and early Pliocene ospreys in Florida
for the paleoecological and biochronological aspects of this study.
Detailed descriptions and systematic remarks may be found in Becker
(1985b).
Genus Pandion Savigny, 1809
Pandion lovensis Becker, 1985
Material. Love Bone Bed local fauna; UF 25950, nearly complete left
tarsometatarsus (holotype); UF 25766, distal half right femur; UF 2588U,
distal end of right tibiotarsus; UF 25928, complete left tibiotarsus; UF
25863, right tarsometatarsus lacking proximal end; UF 26055, UF 26056, UF
29660, ungual phalanges (paratypes).
Remarks. This species is more generalized, with longer and more
slender pelvic limb elements, than the living Pandion haliaetus. See
additional comments in Becker (1985b).
Pandion sp.
Material. Bone Valley Mining District, Palmetto Mine; UF 123^6,
ungual phalanx (claw).
Remarks. This specimen is not identifiable to species, but is
sufficently distinct to document a species of Pandion being present in
the Bone Valley Avifauna.
Remarks on the Family Pandionidae.
Warter (1976) has reviewed the fossil history of this family and
described the first fossil species of Pandion from the mid-Barstovian
Sharkstooth Hill local fauna, California. Recently, another fossil

132
species of Pandion was described from the late Clarendonian Love Bone Bed
local fauna and a phylogeny was proposed for this family (Becker, 1985b).
Family Accipitridae (Vieillot, l8l6)
Genus Haliaeetus Savigny, l809
? Haliaeetus sp.
Material. Bone Valley Mining District, Palmetto Mine; UF 21136,
distal end left humerus. Fort Green Mine; UF 55819, distal end left
tibiotarsus; UF 01956, distal end left tibiotarsus.
Description. Humerus similar in size and morphology to a small
Haliaeetus _1. leucocephalus. Provisionally assigned to the genus
Haliaeetus. Distinguished from Necrosyrtes and Neophron by having a
broader attachment for the anterior articular ligament. Distinguished
from Aguila, Necrosyrtes, and Neophron by having a narrow intercondylar
furrow (=incisura intercondylaris).
Tibiotarsi also the size of the modern Haliaeetus _1. leucocephalus.
There is little variation in size between the two tibiotarsi. UF 55819
has a slightly more slender shaft and a more horizontally placed tendinal
bridge.
Remarks. The distal tibiotarsus is rather undiagnostic in
accipitrids (Jollie, 1976-1977: 226-227), although it has been used in
the past to define species. This material is tentively assigned to
Haliaeetus on size, overall similarity, and the differences noted above.
The variablity of the characters used above has not been determined.
Additional analysis of the characters used by Rich (1980) to separate the
various subfamilies and genera of accipitrids is needed before this
material can be correctly placed within this family.

133
Genus Buteo Lacpde, 1799
Buteo near B. jamaciensis (Gmelin, 1788)
Material Withlacoochee River 4a local fauna; UF 67808, complete
left femur.
Description. Size of Buteo jamaciensis harlani. Contares well with
this subspecies in overall size, position of intermuscular lines.
Differs by having a more narrow patellar sulcus, a slightly more
constricted femoral head, greatest length slightly less, and slightly
more gracile.
Remarks. This specimen has been reported in Becker (1985a).
Genus Aguila Brisson, 1760
Aguila sp. A
Material. Bone Valley Mining District, Hookers Prarie Mine, UF
57299, distal end left tarsometatarsus with posterior wing of trochlea IV
broken off.
Description. Virtually indistinguishable from the range of
variation of that of the living Aguila chrysaetus in size and morphology.
Differs by having a more laterally compressed distal foramen and a
slightly deeper tendinal groove (= outer extensor groove of Howard,
1980).
Accipitrid, Genus indet., species A.
Material. Love Bone Bed local fauna, UF 29676, proximal end right
carpometacarpus.
Description. Similar in size to a large Haliaeetus 1eucocephalus or
a small Aguila chrysaetus. Carpometacarpus agrees with A. chrysaetus by
having a large process on metacarpal I (small in Haliaeetus) and shape of

13U
the distal portion of the carpal trochlea. Carpometacarpus agrees with
Haliaeetus by having a small anterior carpal facet; tendinal groove more
on the external surface and having a similar excavation on external side
of pollical facet.
Remarks. Material now available is not diagnostic to the generic
level. As additional material becomes available, this carpometacarpus
should be re-examined to determine its generic status. The time interval
between the Love Bone Bed and the Bone Valley local faunas make it
unlikely, although not impossible, that these specimens represent one of
the species of eagle from Bone Valley.
Accipitrid, Genus indet., species B.
Material. Bone Valley Mining District, Chicora Mine; PB 8100,
distal end right carpometacarpus.
Remarks. Carpometacarpus badly fractured and warped. It resembles
Haliaeetus but this material is not complete enough to discriminate
between Haliaeetus and species of Aguila of similar size.
Accipitrid, Genus indet., species C.
Material. Love Bone Bed local fauna; UF 25^91 distal end left
tarsometatarsus, lacking half of trochlea III and all of trochlea IV.
Remarks. Typical accipitrid tarsometatarsus. Size between males
and females of Buteo jamaciensis borealis; but also within the size range
of the Neotropical Spizaetus ornatus and of several African species of
the genera Lophaetus; Hieratus and Old World species of Aguila, including
A. rapax and wahlbergi. The incompleteness of the specimen and the lack
of diagnostic characters on the distal end of the tarsometatarsus
(Jollie, 1976-1977:266) prevents assignment of this specimen to a genus.

135
Owing to the long time interval between the Bone Valley local faunas and
the Love Bone Bed, it is unlikely that this specimen represents one of
the Bone Valley eagles.
Accipitrid, Genus indet.
Material. Withlacoochee River 4a local fauna; UF 67809, pedal
phalanx I, digit I; Bone Valley Mining District, District Grade (Agrico)
Mine; UF 57303, ungual phalanx (claw); Fort Green Mine, UF 61957, pedal
phalanx I, digit I (?); Palmetto Mine, UF 21123, UF 21125, ungual
phalanges (claws).
Remarks. All material is representive of a large accipitrid, but is
undiagnostic at the generic level.
Remarks on the Family Accipitridae.
The family Accipitridae had approximately 205 Recent species and 62
fossil species when Brodkorb published his Catalogue of Fossil Birds
(1964). Other species have been described since then. The systematics
of this family is based primarily on external characters (Brown and
Amadon, 1968), and probably does not accurately reflect the evolution of
this diverse group. Several recent studies (Jollie, 1976-1977; Rich,
1980) have described, in exhaustive detail, the morphology of this
family. But at the present, there is not a phylogeny of the family as a
whole based on internal morphology (but see comments in Olson, in press;
and Jollie, 1976-1977:309ff).
In Florida, there is also unreported accipitrid material from the
Hemingfordian Thomas Farm 1. f. and from several late Pleistocene
localities (Campbell, 1980; Carr, 1981).

Order Anseriformes (Wagler, 1831)
136
Family Anatidae Vigors, 1825
Remarks. There is a large amount of fossil material representing
anatids from nearly all local faunas included in this study, especially
from the Love Bone Bed local fauna. Unfortunately, much of it consists
of fragmentary, waterworn specimens or specimens of slight diagnostic
value (radii, ulnae, vertebrae, etc.). The correct taxonomic assignment
of these specimens is further complicated by the presence of both goose
like ducks (Anatinae: Tadornini) and duck-like geese (Anserinae:
Dendrocygnini) which precludes assigning specimens to subfamily on the
basis of size. Therefore, I do not assign material to a taxonomic rank
and list it under referred material unless it is clearly diagnostic. The
classification proposed by Woolfenden (1961) is followed.
Subfamily Anserinae Vigors, 1825
Tribe Dendrocygnini Reichenbach, "1850"
Genus Dendrocygna Swainson, 1831 _
Generic characters. The coracoids of Dendrocygna may be
distinguished from those of all other anseriform genera by the presence
of a pit-like depression on the ventro-lateral surface of the sternal
end. Other characters of the coracoid are listed in Woolfenden (1961).
The carpometacarpus of Dendrocygna may also be distinguished from all
other anseriform genera by its narrow and elongate proportions, the
metacarpal II incurved in dorsal view, the external rim of the carpal
trochlea only slightly notched, and a prominent neck present between the
carpal trochlea and metacarpal III (Woolfenden, 1961).

137
Dendrocygna sp.
Material. Love Bone Bed local fauna; UF 25992, UF 25997, UF 29765,
UF 29766, right coracoids; UF 29764, UF 29763, UF 25803, left coracoids;
UF 29762, UF 25774, UF 25839, humeral ends left coracoids (tenatively
referred); UF 25845, humeral end right coracoids (tentatively referred);
UF 25755, left carpometacarpus, missing metacarpal III, UF 25757,
prominent end left carpometacarpus.
Remarks. Coracoids and carpometacarpus (Fig. 4.7) typical of
Dendrocygna as described by Woolfenden (1961). I have been unable to
find either qualitative or quantitative characters which will distinguish
the fossil specimens from the Love Bone Bed from the few specimens of the
Recent species of Dendrocygna available, when the variation of modern
populations is taken into account. Measurements are given in Table 4.19.
Fossil species in this tribe include Dendrochen robusta A. H.
Miller, based on humeri from the early Hemingfordian Flint Hill local
fauna, South Dakota; and Dendrocygna eversa Wetmore, based on a proximal
humerus from the Blancan Benson local fauna, Arizona. Dendrocygna
validipinnis DeVis, from the Pleistocene of Australia was shown by Olson
(1977a) to be a junior synonym of Biziura.
Recently, Cheneval (1984) transfered three species previously
referred to Anas from St.-Gerand-le-Puy to the genus Dendrochen as D.
blanchardi, D. consobrina and D. natator. He also noted a possible
relationship between Romainvillia and Dendrochen.
Dendrocygna is predominately tropical, with the greatest diversity
of species occurring in the New World tropics and southeastern Asia. Two
species (D. viduata and D. autumnal is) occur in both the Old World and

138
New World tropics. D. autumnalis is widely distributed, highly
discontinuous, with no constant geographical variation (Friedmann, 1947).
Tribe Anserini Vigors, 1825
Genus Branta Scopoli, 1769
?Branta sp. A.
Material. Love Bone Bed local fauna; UF 29751, proximal end right
humerus; UF 29752, distal end right humerus; UF 25797, left coracoid; UF
26005, humeral end right coracoid; UF 25748, UF 25751, proximal ends
right carpometacarpi; UF 25890, UF 25891, distal ends left tibiotarsi; UF
25951, UF 29759, nearly complete left tarsometatarsi.
Remarks. Skeletal elements about the size of those of the Recent
Branta canadensis interior. Assignment to Branta is based on the
following characters: proximal humerus with the attachment of the M.
triceps externus with a distinct border; coracoid with furcular facet not
deeply undercut, with only a few pneumatic foramina present (Woolfenden
1961:49). This assignment is tentative, as similar character states are
approached in specimens of Anser species. Other elements were arbitarily
assigned on the basis of size.
Anserinae, Genus indet. sp. B.
Material. Love Bone Bed local fauna, UF 29753, left carpometacarpus
without metacarpal III; UF 25754, proximal end right carpometacarpus; UF
25758, proximal end left carpometacarpus; UF 29761, right coracoid; UF
25904, distal end right tibiotarsus; UF 25879, distal end left
tibiotarsus; UF 25929, UF 25947, distal ends left tarsometatarsi.

139
Remarks. Skeletal elements about the size of those of the living
males of Anser albifrons. Material badly waterworn, leaving no
diagnostic characters to permit assignment to genus.
Anserinae, Genus indet. sp. C. (or B.?)
Material. Love Bone Bed local fauna; UF 25750, proximal end right
carpometacarpus; UF 25763, distal end right carpometacarpus.
Bone Valley Mining District, near Brewster; PB 17^, proximal end
left carpometacarpus.
Remarks. The above skeletal elements are approximately the size of
those of Anser rossii. These elements may represent Anserinae, genus and
species indet. sp B. (above), depending upon how much size variation is
allowed within a single fossil species.
Anserinae, Genus indet. sp. D
Material. Bone Valley Mining District, specific locality unknown;
UF 61598, left coracoid.
Remarks. Coracoid much smaller than that of Anser rossii (or
species B or C above). Furcular facet deeply undercut as is typical of
the tribe Anserini (Woolfenden, 196l).
Subfamily Anatinae (Vigors, 1825)
Remarks. A large amount of material of anatines exists from nearly
all the localities included in this dissertation, but it is not of
sufficent diagnostic character to allow further identification. Rather
them arbitrarily assign specimens to size categories which would not
reflect the true species composition, I prefer to leave this material
undesignated pending further investigations of the anatid material from

other Neogene fossil localities in North America which have not been
included within this study.
Tribe Tadornini Reichenbach, "1850"
Remarks. The humerus of the tribe Tadornini may be distinguished by
the following combination of characters: capital shaft ridge prominent,
directed forward toward the external tuberosity. Characters that are
typical anatine in form are also present, such as having the external
head of the M. humerotriceps deeply undercutting the humeral head and
being continuous with the capital groove.
Genus and species indet.
Material. Bone Valley Mining District, Ft. Green Mine (# 13
dragline); UF 57253, proximal end right humerus.
Remarks. Typical Tadornine humerus about the size of that of a male
Chloephaga picta. This specimen could represent a large species in the
genus Anabernicula, but it is apparently larger than all species now
described in that genus. The specimen is not complete enough to warrant
assignment to genus.
Tribe Anatini Vigors,1825
Genus Anas Linnaeus, 1758
Anas undescribed sp. A.
Material. Love Bone Bed local fauna; UF 25738, complete left
humerus, UF 25720, left humerus missing distal end, UF 29750, proximal
end left humerus; UF 25731, 25736, proximal ends right humeri; UF 25837
UF 25853, UF 29770, UF 29761, complete right coracoids; UF 25805, UF
29768, complete left coracoids; UF 29801, UF 258L0, UF 258U1, humeral
ends right coracoids (tentativly referred); UF 25795, UF 25780, UF 29795,

humeral ends left coracoids (tentatively referred); UF 25756, left
carpometacarpus missing shaft of metacarpal III.
McGehee Farm local fauna; UF 8780, complete left coracoid; UF 12469,
right tarsometatarsus.
Description. The above taxonomic assignment is based primarily on
humeri. They are referrable to the Anatinae by lacking a prominent
capital shaft ridge, having the capital groove extending laterally across
the anconal surface and deeply undercutting the humeral head, and by
having a strongly developed attachment for the external head of the
triceps.
Within the subfamily Anatinae, the fossil humeri may be
distinguished from those of the tribe Tadornini by lacking a prominent
capital ridge shaft, by having the head unrotated, deltoid crest not
large and flaring and not extending distally. They may also be
distinguished from those of the tribe Cairinini by not having _a robust
shaft and the pneumatic fossa not restricted to a circular opening rimmed
with heavy bone; distinguished from those of the tribe Oxyurini by having
a very deep pneumatic fossa, extending well under head without numerous
foramina piercing the walls, by the entepicondyle not being reduced, and
by the scar for the attachment of the latissimus dorsi posterioris not
being in a line with the outer edge of the pectoral attachment;
distinguished from those of the tribe Mergini and the tribe Somaterini by
the internal tuberosity not being short and deep, and by the
entepicondylar prominence not being reduced.
The fossil humeri from the Love Bone Bed local fauna agree with the
humeri of both the Anatini and the Aythini by having the capital ridge
shaft obsolete and by having the pneumatic fossa ovaloid and unrimmed

142
with heavy bone. The fossil humeri particularly agree with the humeri of
the Anatini by having the impression of the brachialis not well defined
(Aythini with impression well-defined, with a distal medial rim sharp)
and by having the entepicondyle approximately equal in anconal height to
the ectepicondyle (Aythini with entepicondyle distinctly higher). In
three out of five fossil specimens, the pneumatic fossa is open and
contains struts (as in Anatini; in Aythini it is usually closed). Distal
end rotated medially as in Anas. Greatest morphological resemblence of
the humerus, coracoid, carpometacarpus, and tarsometatarsus is to the
genus Anas (Fig. 4.J).
Comparisons were made with the smallest living species of Anas, Anas
hottentata, to illustrate the morphological characters of this extremely
small fossil species from the late Miocene of Florida (see Table 4.20).
The humeri from the Love Bone Bed are smaller, less robust, but with
deltoid crest similar. The coracoids from the Love Bone Bed are smaller,
proportionally more stout, with the medial margin (in ventral view)
slightly inflated. Carpometacarpus from the Love Bone Bed is much
smaller, and more gracile. Tarsometatarsus from McGehee Farm is smaller,
more slender, with the lateral parahypotarsal sulcus more excavated.
Sulcus on anterior surface proximal to canal more defined.
Remarks. The above specimens agree with the genus Anas in all
characters except onewhether the humerus has an open or closed
pneumatic fossa. The two extremes of this character are one of the
defining characters which distinguish the tribes Anatini and Aythini
(Woolfenden, 1961: 12). This site is the entrance of the air sac system
into the shaft of the humerus and is highly adaptive as it allows for the
additional regulation of bouyancy in diving birds. It almost certainly

evolved more than once, as evidenced by its being open in some genera of
mergansers (always open in Mergus, almost always open in Lophodytes) and
closed in others (Mergellus) (Woolfenden, 1961).
Before the exact systematic position of this species can be
determined, additional comparisons are needed with small living and
fossil species of Anas. Of special interest is Anas luederitzensis
Lambrecht 1929, from the mid-Tertiary of southwest Africa. This species
is said to be distinguished from Anas querquedula and Anas cyanoptera by
having a pneumatic fossa not markedly perforated as in most anatids
(Howard, 1964).
?Anas, size near A. acuta
Material. Love Bone Bed local fauna; UF 26001, right coracoid; UF
25798, UF 25799, UF25800, UF 25801, left coracoids.
McGehee Farm local fauna; UF 9481, UF 31784; right coracoids; UF
9486, UF 9489, left coracoids.
Remarks. Typical Anas morphology. Skeletal elements slightly
larger than A. acuta; definately smaller than modern specimens of Anas
platyrhynchos.
Anatini, Genus indet., species A.
Material. Love Bone Bed local fuana; UF 29767, complete left
coracoid; UF 25808, UF 25854, humeral ends coracoids, tentatively
referred.
Remarks. Coracoids proportionally long and slender; slightly
smaller than females of A. crecca carolinensis. See remarks below under
"Genus indet., sp. B.".

144
Anatini, Genus indet., species B.
Material. Love Bone Bed local fauna; UF 25791, left coracoid; UF
25791, UF 25786, UF 25852, humeral ends coracoids, tentatively referred.
Remarks. Similar morphology as described above for genus and
species indet sp. A, but slightly larger than the coracoids of the males
of A. crecca carolinensis. This material probably represents the same
species as "A" above, but of the opposite sex.
Tribe Aythini (Delacour and Mayr, 1945)
Genus Aythya Boie, 1822
Aythya sp. A
Material. Bone Valley Mining District, Ft. Green Mine (#13
dragline); UF 49695, right carpometacarpus; UF 53945, proximal end right
carpometacarpus; Palmetto Mine; UF 21124, left coracoid, (tentative
referred); Ft. Green Mine (#6 dragline); UF 53866, left coracoid,
(tentatively referred); specific locality unknown, UF 61599, left
coracoid, (tentatively referred).
Remarks. Size similar to that of Aythya collaris. Carpometacarpus
referred to this tribe by lacking the distal swelling on the external rim
of the carpal trochlea (Woolfenden, 1961). Coracoids tenatively referred
here because of general overall similarity with Aythya in characters
(Woolfenden, 1961).
Tribe Mergini (Swainson, 1831)
Genus Bucephala Baird, 1858
Bucephala ossivallis Brodkorb, 1955
Material. Bone Valley Mining District, Palmetto Mine (= locality 2
of Brodkorb, 1955a); PB 172, humeral end left coracoid (holotype).

Remarks. Brodkorb (1955a) states that this coracoid agrees with
that of Bucephala clangula in general appearance and details of the
brachial tuberosity, but differs by being smaller and in details of the
procoracoid and triosseal canal.
Anatids are fairly rare members of the Bone Valley avifauna. As
additional material becomes available, this poorly known species should
be re-examined.
Tribe Oxyurini J. C. Phillips, 1926
Genus Oxyura Bonaparte, 1828
Oxyura cf. £. dominicus
Material. Bone Valley Mining District, Ft. Green Mine; UF 61950,
proximal ends left humerus.
Remarks. Size of a small Oxyura dominicus. Humeral head slightly
more undercut by the external head of the m. triceps in the fossi
specimen than in the series of recent skeletons of 0. dominicus. All
other characters within the range of variation of 0. dominicus.
Remarks on the Family Anatidae.
Howard (1964, 1973) has reviewed the extensive fossil record of the
Anatidae, so there is little need once again to review all taxa coverd by
Howard. New Miocene fossil species described since 1973 include
Cygnopterus alphonsi from the Aquitanian of St.-Gferand-le-Puy, France by
Cheneval (1984); and a Blancan goose from the Broadwater local fauna,
Anser thompsoni by Martin and Mengel (1980). Both of these species are
known from several elements.
Four new species have also been described from the late Pleistocene.
Anas schneideri was described from a single carpometacarpus from the

Rancholabrean Little Box Elder local fauna (Eraslie, 1985)* This species
is significantly smaller than the living A. crecca. Anas schneideri is
near the size of A. pullulans described from the mid to late Clarendonian
Black Butte local fauna, Oregon, but differs from this species by having
metacarpal I relatively higher (Emslie, 1985)* Campbell (1979) described
three new species of Anas (A. talarae, A. amotape, and A. sanctahelenae)
from the late Pleistocene of Bolivia and Ecuador. Differences between
these South American species of Anas and other living species of Anas
seem slight. Campbell (1979) also described, from the same localities, a
new genus and species (Nannonetta invisitata) of Tadorine.
Anatids are rare as fossils until the Neogene. Olson (ms) notes
that the major adaptive radiation of anatids took place in the Miocene;
for it is difficult to place fossils before this time in modern tribes
and genera.

Figure 4.7. Dendrocygna sp. A, B. Right coracoid, UF 29765*
A. Dorsal view. B. Ventral view. C, D. Left carpometacarpus, UF
25755 C. Ventral view. D. Dorsal view. Scale = 2.5 cm.

148

Figure 4.8. Anas sp. A, B. Left coracoid, UF 25805. A. Dorsal
view. B. Ventral view. C, D. Left carpometacarpus, UF 25756.
C. Ventral view. D. Dorsal view. E, F. Left humerus, UF 25738.
E. Cranial view. F. Caudal view. Scale (top) A-D = 2 cm.;
(bottom) E, F = 3 cm.

150

Table 4.19. Measurements of the coracoids and carpometacarpi of Dendrocygna viduata (N = 8, 4 males, 4
females), Dendrocygna arbrea (N = 4, 1 male, 3 females, maximun), Dendrocygna bicolor (N = 9 6 males, 3
females, maximun), Dendrocygna autumnalis (N = 8, 4 males, 4 females), and Dendrocygna species from the Love
Bone Bed local fauna. Data are mean +_ standard deviation (number) and range. Abbreviations are defined in
the methods section.
Measurement
D. viduata
D. arbrea
D. bicolor
D. autumnalis
Dendrocygna sp.
Coracoid
HEAD-IDA
40.79 + 1.74
38.6 44.3
44.98 + 1.97
42.9 46.8
42.14 + 3.97
37.8 52.1
42.46 + 1.6l
4o.l 44.8
44.93 + 1.97 (6)
42.1 47.9
HEAD-CS
14.75 + 0.88
l4.l 16.8
15.18 + 1.30
13.5 16.6
15.01 + 0.49
14.3 15.8
15.26 + 0.65
14.5 16.3
15.78 + 0.87 (9)
13.8 16.8
D-HEAD
3.79 + 0.16
3.5 4.0
4.00 + 0.28
3.8 4.0
3.44 + 0.17
3.2 3.7
3.80 + 0.30
3.4 4.3
4.10 + 0.39 (8)
3.8 5.0
W-SHAFT
3.69 + 0.07
3.6 3.8
4.18 + 0.17
4.0 4.4
4.01 + 0.42
3.3 4.5
4.10 + 0.21
3.8 4.5
4.46 + 0.32 (10)
4.1 5.0
D-SHAFT
3.60 + 0.29
3.2 4.0
3.68 + 0.17
3.5 3.9
3.77 + 0.17
3.6~- 4.4
3.86 + 0.19
3.6- 4.0
4.28 + 0.38 (10)
3.8 5.0
FAC-IDA
14.24 + 0.64
13.4 15.1
16.57 + 0.50
16.1 17.I
14.94 + 0.82
13.6 16.3
15.68 + 0.64
14.6 16.6
16.03 + 0.51 (4)
15.5 16.7
IDA-PP
30.05 + 1.20
28.9 32.7
33.53 + 1.53
32.2 35.0
29.54 + 1.17
27.0 30.7
31.26 + 1.45
29.4 33.4
32.86 + 1.13 (7)
31.8 35.2
i
151

Table 4.19continued
Measurement
D. viduata
D. arbrea
C arpometacarpus
LENGTH
52.95 + 2.06
51.5 57.7
59.70 + 2.77
56.5 61.3
W-CARPAL
4.79 + 0.20
4.5 5.1
5.57 + 0.06
5.5 5.6
D-PROX
10.60 + 0.26
10.2 11.0
11.70 + 0.20
11.5 11.9
D-SHAFT
3.00 + 0.18
2.8 3.3
3.43 + 0.06
3.4 3.5
W-SHAFT
3.35 + 0.12
3.2 3.5
3.97 + 0.12
3.9 4.1
W-DIST
6.08 + 0.31
5.5 6.5
6.60 + 0.30
6.3 6.9
i
D. bicolor D. autumnalis Dendrocygna sp.
51.93 + 1.51
49.5 54.1
54.63 + 2.64
50.0 58.5
55.0
4.54 + 0.23
4.1 4.8
4.90 + 0.19
4.6 5.1
5.2; 5.4
10.66 + 0.29
10.2 ll.O
11.30 + 0.51
10.7 12.2
11.3; 11.5
3.13 + 0.19
2.9 3.5
3.16 + 0.24
2.9 3.5
3.4; 3.6
3.38 + 0.21
3.1 3.7
3.70 + 0.26
3.3 4.1
4.0; 4.4
5.76 + 0.31
5.4 6.1
6.29 + 0.36
6.0 6.9
6.2
152

153
Table 4.20. Measurements of coracoids, humeri, carpometacarpi, and
tarsometatarsi of males of Anas hottentata and Anas sp. A from the Love
Bone Bed and McGehee local faunas. Data are mean _+ standard deviation
and range. Abbreviations defined in methods section.
Measurement
Anas hottentata
Anas sp. A
Coracoid
HEAD-IDA
29.7; 30.6; 29.7
25.85 + 0.72 (4)
24.8 26.4
HEAD-CS
9.7; 9.5; 9.7
8.93 + 0.43 (4)
8.3 9-2
W-SHAFT
3.2; 3.1; 3.0
3.06 + 0.23 (7)
2.8 3.4
D-SHAFT
2.2; 2.5; 2.6
2.56 + 0.50 (7)
2.1 3.6
IDA-PP
22.8; 24.0; 22.9
20.37 + 0.34 (6)
19.8 20.7
L-GLEN
5-8; 6.0; 6.1
6.10 + 0.23 (7)
5.B- 6.5
Humerus
LENGTH
54.6; 56.2; 54.2
50.8
W-SHAFT
4.1; 4.0; 4.3
3.63 + 0.31 (4)
3.2 3.9
D-SHAFT
3.5; 3.4; 3.7
3.03 + 0.49 (4)
2.Â¥ 3.5
W-PROX
11.5; 11.6; 11.6
12.2; 11.6
D-PROX
6.2; 6.5; 6.3
6.4; 5.3
D-HEAD
4.1; 4.4; 4.4
3.80 + 0.13 (4)
3.7 4.0
W-DIST
8.2; 8.3; 8.6
7.8
D-DIST
4.8; 4.8; 5.2
4.8
D-ENTEP
3.5; 3.5; 4.0
4.0

Table 4.20continued
Measurement Anas hottentata Anas sp.
Carpometacarpus
LENGTH
32.7; 32.0
27.9
W-CARPAL
3.3; 3.6
2.9
D-PROX
T.8; 7.8
6.7
L-MCI
4.9; 4.8
4.5
D-SHAFT
2.3; 2.3
2.1
W-SHAFT
2.4; 2.6
2.1
D-DIST
2.8; 3.0
2.2
W-DIST
4.3; 4.2
3.5
Tarsometatarsus
LENGTH
27.5; 27.9; 28.1
25.5
W-SHAFT
3.0; 3.1; 2.9
2.8
D-SHAFT
2.4; 2.5; 2.7
2.3
W-PROX
5.4; 5.8; 5.8
5.1

155
Order Galliformes (Temminck, 1820)
Family Phasianidae Vigors, 182$
Subfamily Meleagridinae (Gray, 1840)
Genus indet.
Material. Love Bone Bed local fauna; UF 25768, proximal half left
femur.
Remarks. Steadman (1980) reviewed and evaluated the previously used
taxonomic characters of the meleagridine femur. The femur of
Proagriocharis kimballensis is not known and that of Meleagris progenes
is damaged so that the few qualitative characters of value could not be
judged for these species. As there is a general increase in size through
time in this subfamily (Steadman, 1980:153), I attempted to assign this
specimen to a genus by comparing measurements of Recent and fossil
species (Table 4.21). These comparisons assume that similar measurements
(e.g. diameters of hind limb elements) do not vary between different
skeletal elements of a given species. Ratios of measurements in
Proagriocharis kimballensis to those of Meleagris gallopavo range from
about 0.57 to 0.71. In Meleagris progenes these ratios range from O.76
to 0.84. Ratios for UF 25768 range from 0.60 to 0.64 if the specimen is
assumed to be male (within the range of Proagriocharis kimballensis) or
from 0.80 to 0.83 if the specimen is assumed to be female (within the
range of Meleagris progenes). This specimen is therefore not assigned to
genus.
Genus Meleagris Linnaeus, 1758
cf. Meleagris sp.

156
Material. Bone Valley Mining District, Palmetto Mine; UF 21033,
distal end tibiotarsus.
Remarks. I have not been able to locate this specimen, but have
followed Steadman (I980:l4l) for this identification. There is additional
material of Meleagris in the UF collections from the Bone Valley Mining
District, but it is likely that it originated from Pleistocene deposits.
Remarks on the Family Phasianidae (Subfamily Meleagridinae).
See Steadman (1980) for an in-depth discussion of the osteology,
paleontology, systematics, and evolution of the subfamily Meleagridinae.

Table 4.21. Comparative measurements of the femora and tarsometatarsus of the turkeys Proagriocharis
kimballensis, Meleagris progenes and Meleagris gallopavo silvestris, and Meleagridinae, genus indet. from
the Love Bone local fauna. All measurements are means (Steadman, 1980) except those of the material from
the Love Bone Bed local fauna. Ratios were calculated by dividing the measurement in the fossil specimens
by that of M. silvestris.
Love
Bone Bed
P. kimballensis
M. progenes
M. g.
silvestri:
Measurement
Sex
Datum
Ratio
Datum
Ratio
Datum
Ratio
Datum
Ratio
Femur
W-PROX
M
20.9
0.603




34.65
1.00
F
20.9
0.795




26.28
1.00
D-HEAD
M
8.2
0.641
.
12.79
1.00
F
8.2
0.830




9.81
1.00
Tarsometatarsus
W-PROX
M


i4.0
0.572
18.7
0.764
24.49
1.00
F


12.95
O.665


19.46
1.00
W-SHAFT
M
7.45
0.798
9.34
1.00
F


5.05
0.690


7.32
1.00
D-SHAFT
M
...
__
5.05
0.836
6.04
1.00


3.5
0.709
3.9
O.789
4.94
1.00
1
157

158
Order Ralliformes (Reichenbach, 1852)
Family Gruidae Vigors, 1825
Remarks. The following elements are so waterworn and abraded that I
cannot do more than tentatively refer them to the family Gruidae.
Material. Love Bone Bed local fauna; UF 25966, UF 25967, cervical
vertebrae; UF 26086, right digit II, phalanx I (wing); UF 26087, distal
end left radius; UF 25761, UF 25762, distal ends left carpometacarpi; UF
29723, UF 2589J+, dital ends left tibiotarsi; UF 25958, UF 25960, distal
shafts left tarsometatarsi; UF 25862, UF 25860, fragmentary distal ends
right tarsometatarsi; UF 26032, UF 26033, UF 2603^, UF 26036, UF 26037,
UF 26038, UF 26039, UF 260L0, pedal phalanges.
Generic diagnosis. The following diagnosis is based on an
examination of the following genera (number of species in parenthesis):
Grus (9), Bugeranus (l), Anthropoides (2), and Balerica (2).
No diagnostic characters noted on portion of the coracoid and
carpometacarpus preserved. Femur with proximal portion of the cranial
intermuscular line located medially, away from the crista trochanteris in
Balerica (extending from the crista trochanteris in other genera
examined). Only a few characters on the distal end of the tibiotarsus
will separate all species of Grus from all other species of Bugeranus,
Anthropoides, and Balerica. In Grus, the lateral surface of the lateral
condyle is expanded (less so in other genera), the internal ligamental
process is expanded (similar in Bugeranus, less so in Anthropoides, and
Balerica). Balerica is distinguished by having the distal end anterio-
posteriorly flattened, with the medial condyle rotated outward, producing
a broad, U-shaped anterior intercondylar sulcus. The proximal end of the
tarsometatarsus in Balerica with a short hypotarsus (longer in Grus,
Anthropoides (but short in A. virgo), and Bugeranus); Anthropoides with a

159
transverse sulcus relatively narrow and deeply excavated (less so in
Grus, Balerica, and Bugeranus). Distal end of the tarsometatarsus in
Balerica with trochlea II higher on shaft than in that of other genera.
Subfamily Gruinae (Vigors, 1825)
Genus Grus Pallas, 1766
Grus sp. A.
Material. Love Bone Bed local fauna; UF 25752, proximal end right
carpometacarpus; UF 25903, distal end left tibiotarsus; UF 26092,
proximal end right tarsometatarsus.
Description. Tibiotarsal and tarsometatarsal characters as in the
genus Grus. Carpometacarpal fragment referred on basis of size. Smaller
than all species of living cranes except Grus canadensis. Differs from
Grus canadensis by having a less pronounced lateral expansion of the
lateral condyle, a less pronounced internal ligamental process, a more
robust shaft, a larger tendinal canal, and a broad triangular .crest
extending proximally from the lateral condyle.
Tarsometatarsus slightly larger that that of Grus canadensis;
carpometacarpus slightly smaller.
Remarks. The unexpected difference in size between the above
specimens could possibly be due to the elements representing two sexes,
two subspecies (analogous to the modern situation in Florida resident
population of Grus canadensis pratensis and the migratory Grus canadensis
tabida), or a single species of slightly different proportions. With
only the scanty material above, it is impossible to choose between these
possibilities.
The skeletal elements of the fossil species Grus nannodes from the
Hemphillian Edson local fauna, Kansas, is similar in size to the above

16o
specimens. Hovever, in the original description it was not demonstrated
that Grus nannodes actually belongs in the genus Grus. With the
prevalance of balearicinae-like cranes in the Tertiary of North America,
gruid fossil cannot be uncritically assigned to the genus Grus.
Therefore "Grus nannodes" should be re-examined before its current
taxonomic assignment is accepted.
Grus sp. B
Material. Love Bone Bed local fauna; UF 26002, sternal end right
coracoid; UF 29721, humeral end right coracoid; UF 25740, right scapula;
UF 25722, UF 25737, distal ends left humeri; UF 25749, right
carpometacarpus missing part of shaft of metacarpal III; UF 25753,
proximal end right carpometacarpus; UF 29720, proximal end right femur;
UF 25887 (tent, referred), UF 25893 (tent, referred), UF 25908, UF 25911,
distal ends left tibiotarsi; UF 25885, UF 25896 (tent, referred), UF
29722, distal ends right tibiotarsi; UF 25988, UF 25989, UF29725,
proximal ends left tarsometatarsi; UF 26093, UF 29724, proximal ends
right tarsometatarsi; UF 25857, distal end right tarsometatarsi; UF
25931, UF 25944, UF 25945, UF 29726, distal end left tarsometatarsi.
Remarks. Size similar to that of a large Grus americana or Grus
.japonensis. Coracoids, carpometacarpi, humeri, and femur referred here
on the basis of size, as are the several of the above waterworn
tibiotarsi.
Distal tibiotarsus with characters of the genus Grus, but also
similar to Grus (= Bugeranus) lsucogeranus. Larger than Grus canadensis,
G. grus, and G. monacha, slightly larger tha G. vipio and Grus rubicunda
(G. nigricollis not available). Similar in size to G. americana, G.

l6l
.laponensis, and G. antigone. These last three species have very similar
distal tibiotarsi and I have not been able to find consistent qualitative
characters on this element to separate these species from each other and
from the fossil specimens above. The proximal and distal ends of the
tarsometatarsus are also indistinguishable from those of G. americana, G.
Japonensis, and G. antigone. Two specimens (distal tibiotarsi; UF 25908,
UF 25911) are larger than all other fossil specimens and may represent
either sexual variation or a specific difference. The lack of an
adequate series of skeletons of Recent species prevents determination of
this question.
A large species of Grus is known from the Lee Creek local fauna
(Olson, ms). Grus sp. B. probably is closely related, or identical with
this species.
Subfamily Balearicinae (W. L. Sclater, 1924)
Balearicinae, Genus et. species indet. _
Material. Bone Valley Mining District, Nichols Mine; UF 24586,
distal end of right tibiotarsus.
Description. Tibiotarsus slightly smaller than that of a female
Anthropoides virgo, but having characters typical of the subfamily
Balearicinae. Comparisons with the tibiotarsus of an undescribed
balearicinae crane from the Hemphillian Long Island local fauna, Kansas,
(YPM 4662) shows UF 24586 to share a similar shape and position of the
condyles, although UF 24586 is much larger and is more anterior-
posteriorly compressed.
Remarks. Comparisons of this tibiotarsus with unpublished fossil
material in the Frick and USNM collections shows that this specimen is
distinct from that of both Probalearica and Aramornis. Additional

162
material of balearicine cranes is now under study by Feduccia. As much
of this material consists of complete, articulated skeletons, it seems
unneccessary to further describe this single, partial element. From the
material which I have seen, there are at least two species of
balearicinae cranes in North America at the end of the Hemphillian.
Based on the above specimen, the last occurrence now known of this
subfamily of cranes in North America is from the late Hemphillian Bone
Valley local fauna.
Genus Aramornis Wetmore, 1926
cf. Aramornis, sp. A.
Material. Love Bone Bed local fauna; UF 25949, distal end left
tarsometatarsus, missing trochlea IV.
Description/ Remarks. Tarsometatarsus compares well with the type
of Aramornis longurio (F:AM 6269) in relative proportions and positions
of the trochlae II and III, but differs by having a larger distal foramen
and a greater overall size. See Table 4.25 for measurements. Additional
material is needed to verify this generic assignment.
Remarks on the Family Gruidae.
Johnsgard (1983) considers the following species to be closely
related: Grus .japonensis and G. americana, Grus rubicundus and G.
antigone; Bugeranus carunculatus and B. leucogeranus; Antropoides virgo
and A. paradisea; and Balerica pavonina and B. regulorum. Additionally,
he considers Grus grus, G. monacha, G. canadensis, and G. vipio to form a
loose species cluster. Outside of these obvious groupings, he makes no
attempt to place them within a phylogeny. With the exception of Wood's
(19T9) phenetic study, little work has been done concerning the

163
relationships of the cranes above the species level. Wood's paper (1979)
unfortunately adds little new information; rather it addresses the
concordance between previously proposed classifications and the clusters
produced from multivariate analysis of different suites of characters.
The family Aramidae is here considered to be closely related to the
subfamily Balearicinae (Olson, ms). Fossil species of cranes (Gruidae +
Aramidae) are numerous throughout much of the Tertiary of North America
and Europe. They are listed in Brodkorb (1967), reviewed formally in
Cracraft (1973), and reviewed less formally by Olson (ms). Systematic
changes to Brodkorb (1967) may be found in the latter two publications.
My survey of mainly unpublished collections in the Frick
Collections, American Museum of Natural History, indicates there was a
great radiation of balearicinae cranes in North America during most of
the Tertiary (Becker, in prep; Olson, in press). Many fossil species
originally assigned to the Gruinae should be re-assigned to the
Balearicinae; and probably would have been long before now, were this
subfamily still extant in North America. The latest known occurrence of
this subfamily in North America is from the latest Hemphillian Bone
Valley local fauna.

164
Table 4.22. Measurements of the tibiotarsi and tarsometatarsi of Grus
canadensis tabida (N = 7, 4 males, 3 females), Grus canadensis canadensis
(N = 6, 3 males, 3 females), and Grus species A. from the Love Bone Bed
local fauna. Data are mean +_ standard deviation and range. Abbreviation
are defined in methods section.
Measurements
G. c. tabida
G. c. canadensis
Grus
Tibiotarsus
W-SHAFT
9.46 + 0.16
9.2 9.6
8.27 + 0.88
7.2 8.8
9.5
D-SHAFT
8.24 + 0.38
7.7 8.8
6.97 + 0.95
5.9 8.4
7.8
W-DIST-CR
18.73 + 0.84
17.7 20.1
17.33 + 2.09
15.2 20.3
18.8
W-DIST-CD
14.30 + 0.81
12.7 15.0
13.05 + 1.48
11.5 15.1
l4.i
D-MCON
18.73 + 0.76
17.5 19.7
17.52 + 2.14
15.1 20.6
17.9
D-LCON
18.39 + 0.60
17.3 19.1
16.80 + 1.86
14.7 19.6
17.0
D-ICON
10.29 + 0.38
9-7 10.9
9.32 + 0.86
8.3 10.4
9.7
Tarsometatarsus
W-PROX
20.83 + 0.99
19.8 22.2
19.23 + 1.93
17.3 21.6
22.5
D-PROX
13.37 + 0.33
12.9 13.8
11.70 + 1.01
10.4 13.1
12.7
D-PROX-L
19.57 + 0.80
18.5 20.5
17.82 + 1.59
l6.4 20.2
19.0

165
Table 4.23. Measurements of the humeri, tibiotarsi, and tarsometatarsi
of Grus americana (N = 9 maximun, 2 males, 2 females, 5 unsexed), Grus
japonensis (N = 6 maximun, all unsexed), and Grus sp. B. from the Love
Bone Bed local fauna. Data are mean +_ standard deviation and range.
Abbreviation are defined in methods section. (*) Specimen abraded or
broken.
Measurements
G. americana
Humerus
W-DIST
35.93
34.0
+ 1.84
- 38.5
D-DIST
20.56
18.8
+ 1.69
- 23.8
Tibiotarsus
W-SHAFT
11.19
10.3
+ 0.8l
- 13.0
D-SHAFT
9-T9
8.6 '
t- 0.83
- 11.3
W-DIST-CR
22.68
21.0
+ 1.00
- 24.1
W-DIST-CD
16.97
15.8
+ 0.71
- 18.1
D-MCON
21.72
21.2
+ 1.16
- 23.6
D-LCON
21.29
19.6
+ 1.33
- 23.1
D-ICON
12.64
11.6
+ 0.62
- 13.6
G. japonensis Grus sp. B.
38.73
+ 1.48
35.7
37.2
- 41.5
21.13
+ 0.94
18.0; *17.9
19.8
- 22.3
12.23
+ 0.34
10.7
11.8
- 12.8
10.33
1 +
0

-P"
V/l
10.7
9.7
- 11.0
24.83
+ 0.91
*23.7; 23.9;
23.9
- 26.4
23.1
18.52
+ 1.11
18.8; *15.2
17.2
- 20.4
24.93
+ 0.88
*24.1; *21.8
23.5
- 25.6
*23.2
23.33
+ 0.96
*22.8; 24.0;
22.2
- 24.5
*21.8; *21.5
13.08
+ 0.48
13.04 + 0.78 (:
12.5
- 13.6
12.3 14.2

166
Table 4.23continued
Measurements
G. americana
G. japonensis
Grus sp. B.
Tarsometatarsus
W-PROX
24.83 + 0.93
23.4 26.0
26.33 + 2.49
21.9 28.6
26.36 + 0.81 (5)
25.3 27.5
D-PROX
16.02 + 1.36
14.5 17.5
19.8
17.86 + 1.25 (5)
15.7 18.7
D-PROX-L
23.14 + 1.33
22.0 24.4
25.7
23.70 + 0.62 (5)
23.0 24.5
TRIII-TRIV
20.47 + 1.40
18.1 22.5
22.93 + 1.23
22.1 25.4
19.8; 21.8
TRII-TRIV
20.58 + 1.90
17.9 22.9
23.62 + 0.87
22.3 24.9
19.9
W-TRII
6.93 + O.58
6.5 8.0
7.95 + 0.29
7.6 8.4
7.48 + 0.35 (4)
7.3 8.0
D-TRII
13.60 + 0.45
12.8 14.0
15.15 + O.56
l4.4 15.9
*12.2; 13.3; l4.l
W-TRII
9.95 + 0.36
9.6 10.0
11.33 + 0.49
10.9 12.2
10.25 + 0.44 (4)
9.8 10.8
D-TRIII
12.33 + 0.56
11.5 13.0
13.98 + 0.59
13.0 14.5
13.3; *12.T; 13.4
W-TRIV
6.57 + 0.49
6.1 7.3
7.52 + 0.32
7.1 7-9
*6.0; 6.4; 6.6
D-TRIV
14.43 + 0.52
13.6 15.O
15.65 + 0.42
15.1 16.2
13.2; *13.4; 14.7

Table 4.24. Measurements of the tibiotarsi of Balerica pavonica (N = 8,
4 males, 4 females) and Balearicinae, genus and species indet. from the
Bone Valley Mining District. Data are mean + standard deviation and
range. Abbreviation are defined in
or broken.
methods section
Measurement
Balerica pavonica
Balearicinae
Tibiotarsus
W-SHAFT
9.16 + 0.49
8.2 9.8
8.3
D-SHAFT
7.98 + 0.44
7.4 8.6
6.6
W-DIST-CR
19.49 + 0.80
18.1 20.4
15.8
W-DIST-CD
13.10 + 0.79
12.2 l4.4
11.8
D-MCON
19.33 + 0.98
17.4 20.6
*14.0
D-LCON
18.04 + 0.66
17.5 19.0
14.1
D-ICON
10.75 + 0.45
10.1 11.5
8.3

168
Table 4.25 Measurements of the tarsometatarsi of Aramornis longurio
(F:AM 6269, holotype) and Aramornis sp. from the Love Bone Bed local
fauna. Abbreviation are defined in methods section. (*) Specimen
abraded or broken.
Measurements Aramornis lonRurio
Aramornis sp.
Tarsometatarsus
W-TRII 4.2
4.4
D-TRII
*7.2
W-TRIII 5.7
*6.0
D-TRIII 7.2
*7-7
D-TRIII

169
Family Rallidae Vigors, 1825
Family Characters. See references in family remarks section; also
Gilbert, et al. (1981).
Material. Love Bone Bed local fauna; UF 26015, UF 29714, UF 29709,
UF 25787, coracoid fragments.
Remarks. The above material appears to be rallid, but is not
identifiable to the generic level.
Genus Rallus Linnaeus, 1758
Rallus sp. A
Material. Love Bone Bed local fauna; UF 25936, distal end left
tarsometatarsus.
Remarks. Near Crex crex in size. Decidely larger than Rallus
limicola, but smaller than Rallus longirostris. I have not been able to
find qualitative characters in the distal ends of the tarsometatarsus to
distinguish between Rallus and Crex and have therefore arbitrally
assigned these specimens to Rallus.
Rallus sp. B
Material. Bone Valley Mining District, Palmetto Mine; UF 21060,
humeral end right coracoid. Payne Creek Mine; UF 21204, proximal end
left tarsometatarsus.
Remarks. Coracoid slightly smaller than that of females of Rallus
longirostris. Agrees with Rallus by having a small procoracoid process.
Tarsometatarsus near Crex crex in size, or intermediate between Rallus
limicola and Rallus longirostris. These two specimens could possibly
represent two different species, but owing to the slight difference in size
and the lack of additional specimens, I have listed them together as

170
representing an undetermined species of Rallus until tetter material is
available.
Rallus (cf.) sp. C.
Material. Love Bone Bed local fauna; UF 25732, distal end left
humerus; UF 26025, UF 26027, UF 26030, UF 29705, UF 29712, humeral ends
right coracoids; UF 29708, left coracoid; UF 26007, UF 26025, UF 29702,
UF 29703, UF 29704, UF 29710, UF 29711, UF 29713, UF 67807, humeral ends
left coracoids; UF 29707, distal ends right tarsometatarsus.
Remarks. All specimens badly water-worn, within the size range of
Rallus limicola or Porzana Carolina or slightly larger. I cannot
distinguish between these two genera on the basis of waterworn elements,
and have therefore arbitrarily assigned these specimens to Rallus on the
basis of size.
Undescribed Genus and Species
Material. Love Bone Bed local fauna; UF 25727, proximal end left
humerus; UF 25836, UF 25849, UF 29715, UF 29716, UF 29717, complete (or
nearly so) right coracoids; UF 29718, humeral end right coracoid; UF
29719, humeral end left coracoid; UF 25865, UF 25866, distal ends right
tarsometatarsi.
McGehee Farm local fauna; UF 9494, proximal end right humerus; UF
29748, nearly complete right coracoid.
Description. Coracoid with a greatly expanded procoracoid process,
extending relatively far down shaft; shaft fairly slender; dorsal surface
not deeply excavated; coracoid fenestra small; lateral process greatly
expanded (Fig. 4.9).

171
Humerus with shallow pneumatic foramen; wide and deep capital
groove. Crest-like margo caudalis extending from distal end of capital
groove (Fig. 4.9). Deltoid crest greatly expanded. Tarsometatarsi worn
and badly abraded and are assigned here mainly on the basis of size.
Distinguished from all genera examined (listed below) by having the
following combination of characters: coracoid with a large procoracoid,
flared medially, causing the triosseal canal to be open; dorsal surface
of coracoid not deeply excavated. Humerus with large deltoid crest and
well-developed capital shaft ridge.
Remarks. In trying to place this fossil material in a genus, I have
examined the skeletons of a large number of genera (number of species in
parentheses). For convience, nomenclature follows Peters (1934), but see
Olson (1973b) for phylogenetic relationships. Genera examined include:
Rallus (6), Alantisia (l), Ortygonax (2), Amaurolimnas (l), Ral lina (l),
Aramides (3), Gymnocrex (l), Gallirallus (2), Habroptila (l), Himantornis
(l), Canirallus (l), Crex (l), Limnocorax (l), Porzana (7), Laterallus
(4), Micropygia (l), Coturnicops (l), Neocrex (l), Sarothrura (l),
Poliolimnas (l), Porphyriops (l), Tribonyx (l), Amaurornis (l), Gallicrex
(l), Gall inula (2), Porphyrula (l), Porphyrio (3), and Flica (6).
None of the genera above match the characters of the fossil
material. There is a superficial resemblance of the coracoid (especially
the enlarged procoracoid process) to Canirallus, Gymnocrex, Aramides, and
Amaurolimnas. This appears to be due to shared primitive characters
between the fossil rail and these primitive living rails (Olson, 1973b).
Additional comparisons with other living and fossil rails are needed
to determine the systematic position of this fossil rail. Possibly
Flica infelix Brodkorb, known only from the distal end of a tibiotarsus,

172
from the late Miocene of Oregon, should he compared further with this
undescribed genus and species.
Remarks on the Family Rallidae.
The fossil record of rails has been recently reviewed (Feduccia,
1968; Cracraft, 1973; Olson, 197^+b, 1977b; Kurochkin, I98O; and references
therein). The generic or higher status of living rails has been reviewed
by Olson (1973b) and accounts have appeared for all living species
(Ripley, 1977).
The obvious lack of rails from many of the fossil localities in this
study is probably size related, whether due to inadequate sampling while
collecting, or bone destruction during deposition (or both).
Several fossil species already described should be re-examined as
additional material becomes available from Florida. The fossil species
Rallus phillipsi Wetmore, 1957, from the Wickieup 1. f. of Arizona was
described as intermediate in size between Rallus limicola and Rallus
longirostris. Olson (1977b) notes that when a larger series of recent
comparative skeletons are examined, Rallus phillipsi falls well within
the lower size range of the living Rallus longirostris. Rallus sp. A and
perhaps Rallus sp. B possibly has affinites with Ib_ phillipsi, but
without more material, this cannot be determined.
Rallus prenticei Wetmore 19^ from the Blancan of Kansas and Idaho,
was described as being somewhat larger and heavier than the living Rallus
limicola. Rallus sp. C may have affinities with this species. Better
material is again needed to identify this species with confidence.

Figure 4.9 A C. Rallid, undescribed genus. A, B. Right
coracoid, UF 29717. A. Ventral view. B. Dorsal view. C.
Proximal end right humerus, UF 9494, caudal view. D. Philomachus
sp., proximal end left humerus, UF 60062, caudal view. E.
Tytonid, undescribed genus, distal end right tibiotarsus, UF 25926,
cranial view, stereopair. Scale A D (top) = 1.5 cm; E (bottom) =
1.5 cm.

-T5S
174

175
Order Charadriiformes (Huxley, 1867)
Remarks. The phylogenetic position of the Phoenicopteridae has long
been debated by avian systematists. Flamingos have been placed with the
storks in the Order Ciconiidae, with the ducks in the Order Anseriformes,
or commonly in their own order, the Phoenicopteriformes. I follow the
recent work by Olson and Feduccia (1980) which establishes the
Phoenicopteridae as a family within the Order Charadriifromes, as shown
by their life history, behavior, nyology, pterylosis, natal down, oology,
parasites, biochemistry, osteology, and paleontology.
Family Phoenicopteridae Bonaparte, 1831
Remarks. Recent flamingos are usually divided into three genera,
based primarily on bill morphology and the presence (or absence) of a
very reduced hind toe. Phoenicopterus is usually considered to have the
most primitive feeding apparatus of the three, with Phoenicoparrus and
Phoeniconaias being more specalized. There are no characters -in the
post-cranial skeleton which will define these genera (Olson and Feduccia,
1980; personal observation). For the basis of comparison with fossil
species, I consider the living species to be congeneric. Olson and
Feduccia (1980) also consider the fossil genera Gervaisia Harrison and
Walker and Harrisonavis Kashin to be junior synonyms of Phoenicopterus
Linnaeus. Additionally, I consider Leakeyornis Rich and Walker to also
be a junior synonym of Phoenicopterus, as even the original authors of
Leakeyornis suggest (Rich and Walker, 1983: 104)! I follow Olson and
Feduccia (1980) and A. H. Miller (1944) in not recognizing the
Palaelodidae as a separate family. Both Palaelodus and Megapalaelodus
may be separated from Phoenicopterus by having a shorter tibiotarsus and
a shorter and more laterally compressed tarsometatarsus (Olson, ms).

176
These two genera are thought to be more specalized for swimming than
Phoenicopterus.
Genus Phoenicopterus Linnaeus, 1758
Remarks. The post-cranial skeletal elements of flamingos exhibit an
extremely large amount of sexual and individual variation. This
variability is especially evident in the tibiotarsus and tarsometatarsus,
with the males typically being much larger than females. The C. V's of
measurements from the Bone Valley material do not exceed those of a
Recent species, suggesting a single species is present here. At the Love
Bone Bed specimens range in size from smaller than that of the modern P.
minor to larger than that of the modern P. ruber, indicating that more
than one species is present. In trying to separate the Love Bone Bed
specimens into species, I found I could only separate the two extremes of
this continium. I have noted these extremes in the referred material as
"large" or "small".
Material. Love Bone Bed local fauna; UF 29678, humeral end left
coracoid; UF 26088, proximal end left radius; UF 29677, distal end right
ulna; UF 25769, proximal end left femur; UF 25957, distal portion of
shaft of tarsometatarsus; UF 259^3, UF 25930, distal fragments left
tarsometatarsus; UF 2031, UF 26035, UF 260k2, UF 260^3, pedal phalanges.
Remarks. The material listed above is considered to be either too
abraded or too undiagnostic for a more refined taxonomic assignment.
Phoenicopterus floridanus Brodkorb, 1953
Material. Bone Valley Mining District, Palmetto Mine (= Locality 2
of Brodkorb, 1955), referred by Brodkorb; PB lVf, distal end right
tibiotarsus (Holotype, not seen), PB 202, shaft right tibiotarsus (=

177
"paratype"), PB 146, PB 300, distal ends right tarsometatarsi (=
"paratype"). New material (referred by Becker); PB 139, proximal end
right carpometacarpus; PB 7980, distal end left tibiotarsus. Nichols
Mine; UF 24623, distal end right tibiotarsus (tentatively referred); Ft.
Green Mine (# 13 dragline), UF 52422, distal end right tibiotarsus; Payne
Creek Mine, UF 67811, distal end right tibiotarsus; No Specific Locality;
UF 29743, distal end right tibiotarsus; Palmetto Mine; UF 21164, distal
end right tarsometatarsus.
Description/ Remarks. For qualitative characters and remarks see
Brodkorb (1953b, 1955a) Measurements given in Table 4.26.
Phoenicopterus sp. A
Material. Love Bone Bed local fauna; UF 25905, UF 25907, UF 29685,
UF 29744, distal ends right tibiotarsi (large); UF 25882, UF 25883, UF
25892, UF 25898, UF 25910, UF 29684, distal ends left tibiotarsi (large);
UF 29686, distal end right tibiotarsus (small); UF 25889, UF 25899, UF
25927; distal ends left tibiotarsi (small); UF 25881, UF 25897, distal
ends right tibiotarsi (abraded); UF 25895, UF 29682, UF 29683, distal
ends left tibiotarsi (abraded); UF 29679, proximal end right
tarsometatarsus; UF 25859, UF 25864, distal ends right tarsometatarsi;
UF 25932, UF 25935, UF 29680, UF 29681, distal ends left tarsometatarsi.
McGehee Farm local fauna; UF 11103, distal end right
tarsometatarsus.
Description. Tibiotarsi with a pronounced bimodal size
distribution, less so in the tarsometatarsi. The group of small
tibiotarsi cannot be distinguished from the tibiotarsi of P. floridanus
when specimens are directly compared. The group of large tarsometatarsi

178
may be distinguished from that of P. floridanus by (l) larger size, (2)
proportionally wider distal end, (3) more robust shaft, (4) papilla just
medial to the proximal opening of the tendinal canal more expanded.
Comparisons of the distal end of a right tibiotarsus (USNM 242202)
from the Lee Creek local fauna with the large subset of tarsometatarsi
from the Love Bone Bed show it to be similar in all characters except for
the notch on the distal surface of the external condyle being slightly
more pronounced in distal end view. Distal opening of the tendinal canal
more transversely elongated and nutrient canal lateral to papilla is
larger and more pronounced in the Love Bone Bed specimens than in the Lee
Creek specimen in cranial view. Groove usually deeper and more
pronounced in the Love Bone Bed specimens than in the one from Lee Creek
in lateral view. In caudal view, the most cranial portion of the lateral
edge of the articular surface is expanded cranially and laterally in Lee
Creek specimens and not in specimens from the Love Bone Bed.
Distal ends of tarsometatarsi from the Love Bone Bed and McGehee may
be distinguished from the distal ends of tarsometatarsi of P. floridanus
by (l) averging larger and more robust (P. floridanus smaller and more
gracile) (2) caudal portion of the articular surface of trochlea III is
raised above shaft (P. floridanus blends relatively smoothly with shaft)
(3) Distal foramen elliptical and larger (P. floridanus smller, more
nearly circular) (4) Prominent nutrient foramen pierces the caudal
surface of the shaft cranial to the distal foramen (P. floridanus
nutrient foramina very small or absent).
Remarks. The flamingos from the Love Bone Bed show a distinct
bimodal distribution in the size classes of the tibiotarsi, with the
smaller group of tibiotarsi being very similar, if not identical with,

179
that of P. floridanus. Along with these small specimens are a group of
larger specimens which represent a much larger flamingo. If this
variation is considered to be sexual in origin, then there is more size
variation than has ever been observed in any single living or fossil
species of flamingo. This suggests that there are two species of
flamingos present at the Love Bone Bed, which overlap in size. The
similarity of the smaller flamingo from the Love Bone Bed to that of P.
floridanus from the Bone Valley (4 MA later in time) suggest that
flamingos have had a slow evolutionary rate. Pending further studies on
other Miocene flamingos, particularly P. stocki, I have left these
Florida specimens unassigned to species.
Remarks on the Family Phoenicopteridae.
Fossil species of Phoenicopterus are now known from the Aquitanian
in Europe and the late Miocene to the late Pleistocene of North America.
Phoenicopterus croizeti Gervais, was described from the Aquitanian of
France. It is known from abundant material and has recently been
restudied (Cheneval, 1984). Phoenicopterus stocki Miller, is based on a
distal tibiotarsus from the Hemphillian Yepomera local fauna. It is said
to have the morphological characters of the genus, but is of pigny size
(Miller, 1944: 77). Phoenicopterus floridanus Brodkorb, is discussed
above. Other Neogene species of Phoenicopterus include only P.
novaehollandiae A. Miller, from the late Oligocene or early Miocene of
Australia. Pleistocene species of Phoenicopterus include P. copei
Shufeldt, from Fossil Lake in Oregon and Manix Lake in California and £.
minutus from Manix Lake in California. All North American fossil species
should be revised and analysed with a larger database of skeletons to
account adequately for the large size variation present.

18o
Table 4.26. Measurements of the tibiotarsi and tarsometatarsi of the
fossil species Phoenicopterus floridanus from the Bone Valley Mining
District, Phoenicopterus sp. A~ (large), and Phoenicopterus sp. B.
(small) from the Love Bone Bed local fauna. Data are mean +_ standard
deviation (number) and range. Abbreviations defined in the methods
section.
Measurement P. floridanus P. sp. A (lg.) P. sp. A (sm.)
Tibiotarsus
W-DIST-CR
15.09
+ 0.4i
(4)
17.53
+ 0.70
(8)
15.0;
15.8
14.5
- 15.4
16.3
- 18.2
W-DIST-CD
10.39
+ 0.55
(4)
12.23
+ 0.33
(8)
9.6;
11.1
9.9
- 11.0
11.8
- 12.8
D-MCON
17.49
+ 0.81
(4)
20.33
+ 0.58
(6)
16.8;
19.5
16.4
- 18.3
19-3
- 20.8
D-LCON
17.51
+ 0.74
(5)
20.32
+ 0.63
(7)
16.8;
19.1
16.8
- 18.5
19.5
- 21.5
Tarsometatarsus
W-TRIII
TRIII-TRIV
7.83 + 0.64 (3)
7.1- 8.3
8.3; 9.4
8.08 + 0.60 (5)
7.1 8.6
14.25 + 0.24 (4)
14.1 14.6
Combined
in column
to the
left
13.4; 15.0
15.60 + 0.35 (3)
15.4 16.O
TRII-TRIV

Table 4.27. Measurements of the tibiotarsi and tarsometatarsi of the Recent Phoenicopterus jamesi (N = 9, 1
male, 2 females, 6 unsexed), Phoenicopterus chilensis (N = 8, 3 males 2 females, 3 unsexed), Phoenicopterus
ruber (N = 15, 6 males, 7 females, 2 unsexed), and Phoenicopterus minor (N = 7, 3 males, 4 females).Data
are mean _+ standard deviation (number) and range. Abbreviations defined in the methods section.
Measurement
P. ,1amesi
P. chilensis
P. ruber
P. minor
Tibiotarsus
W-DIST-CR
14.49 + 0.67
13.6 15.4
14.96 + 0.65
14.2 16.3
15.70 + 0.87
14.6 17.2
12.79 + 0.65
12.0 13.6
W-DIST-CD
10.21 + 0.39
9.6 10.8
10.26 + O.65
9-5 11.7
10.93 + 0.70
9.8 12.2
8.85 + 0.46
8.0 9.3
D-MCON
15.79 + 0.87
14.3 16.8
16.17 + 0.64
15.4 17.4
18.70 + 1.15
16.3 20.8
14.33 + 0.67
13.2 15.1
D-LCOT
15.29 + 0.69
13.9 16.3
16.29 + 0.53
15.5 17.1
18.80 + 1.19
l6.6 21.2
14.23 + 0.68
13.1 15.0
Tarsometatarsus
W-TRIII
7.09 + 0.44
6.3 7.7
7.55 + 0.38
7.2 8.4
7.85 + 0.50
7.2 8.8
6.31 + 0.40
5.7 6.8
TRIII-TRIV
II.92 + 0.49
11.2 12.5
12.47 + 0.37
11.9 13.1
13.91 + 0.86
12.6 15.9
11.15 + 0.65
10.3 12.0
TRII-TRIV
i4.oi + 0.64
13.2 15.0
14.13 + 0.97
13.0 15.9
14.99 + 0.80
13.7 16.5
12.22 + 0.77
11.6 13.5
i
oo

182
Family Jacanidae (Stejneger, 1885)
Characters. The Jacanidae may he distinguished from other families
in this order by having a tarsometatarsus with an extremely large distal
foramen with a deep tendinal groove leading into it and a deep pit
present on the medial surface of the inner trochlea (Olson, 1976).
Coracoid with procoracoid not perforated by a coracoidal fenestra
(perforated in all Charadriform families except the Rostratulidae,
Scolopacidae, Thinocoracidae, and Pedionomidae). Elongated tuberosity
(for attachment of part of the membrana sternocoracoclavicularis) present
on the procoracid and well developed in Jacanidae (less developed in the
Rostratulidae, small but distinct in Pedionomus; lacking in other
charadriforms).
Genus Jacana Brisson, 1760
Jacana farrandi Olson, 1976
Material. McGehee Farm local fauna (referred by Olson, 1976); UF
21219 (holotype), distal end of left tarsometatarsus, missing trochlea
IV; UF 11108 (paratype), left coracoid.
Love Bone Bed local fauna (referred by Becker); UF 25824, UF 29694,
UF 29696, UF 29698, humeral ends left coracoids; UF 260l6, UF 26026, UF
29690, UF 29691, humeral ends right coracoids; UF 29700, extreme proximal
end right humerus (tentatively referred); UF 25728, proximal end of left
humerus; UF 67806, distal end left tibiotarsus.
Description/ Remarks. Proximal end and shaft of humerus (UF 25728)
differs from Jacana spinosa by having shaft more gracile, pneumatic fossa
less deep, head of humerus not undercut by capital groove, and the small
papilla-like process not present. Deltoid crest broken. Coracoid
similar to the paratype coracoid described by Olson (1976). The

183
coracoids range in size from similar to the paratype of Jacana farrandi
to decidely larger. This supports Olson's assertion (1976:261-262) that
the paratype of J. farrandi (UF 11108) is from a male. Tibiotarsus
similar to Jacana spinosa except for a reduced lateral epicondyle. For
type description and additional remarks see Olson (1976).
Remarks on the Family Jacanidae.
Rhegminornis calobates was originally described by Wetmore (l9*+3b)
as a jacana from the early Hemingfordian Thomas Farm local fauna. Olson
and Farrand (197*0 and later Steadman (1980) showed that this species has
affinities with the Phasianidae (Meleagridinae). Olson (1976) later
described the only fossil species of this family now known, Jacana
farrandi (discussed above).
The Jacanidae have long been allied with the Rostratulidae of South
America. In these two families, the females are larger than the males
and the young are similar (downy, strongly marked dorsally with black-
edged stripes from the forehead to tail). The similar osteology of these
two families support their association. Strauch (1978) presented a
subordinal classification of the Charadriiformes and associated the
Jacanidae, Rostratulidae, Scolopacidae, Phalaropodidae, and Thinocoridae
in his suborder Scolopaci. Characters used to support this
classification included the absence of the maxillo-palatine strut A,
absence of the coracoidal fenestra, and presence of a ridge in the
capital groove of the humerus. Olson and Steadman (1981) showed that
Pedionomus (Pedionomidae) was not only a Charadriiform, but had
affinities with at least one of these families (Thinocoracidae). The
relationships of these families need additional study.

184
Table 4.28. Measurements of the humeri and tibiotarsi of Jacana spinosa
(N = 12, 6 males, 6 females) and Jacana farrandi. For measurements of
the type (tarsometatarsus) and paratype (coracoid) see Olson (1976).
Data are mean +_ standard deviation and range. Abbreviations are defined
in the methods section.
Measurements J. spinosa J. farrandi
males females
Humerus
W-SHAFT
2.67
+ 0.05
3.10
+ 0.13
3.4
2.6
- 2.7
3.0
- 3.3
D-SHAFT
2.25
+ 0.08
2.70
+ 0.09
2.9
2.1
- 2.3
2.6
- 2.8
W-PROX
8.25
+ 0.23
9.70
+ 0.52
9-6
7.9
- 8.4
9-0
- 10.6
D-PROX
4.68
+ 0.12
5.48
+ 0.18
5.9
4.5
- 4.8
5.2
- 5.7
D-HEAD
2.45
+ 0.08
2.82
+ 0.08
3.0
2.4
- 2.6
2.7
- 2.9
Tibiotarsus
W-SHAFT
2.33
+ 0.10
2.67
+ 0.14
2.8
2.2
- 2.5
2.5
- 2.8
D-SHAFT
1.93
+ 0.08
2.25
+ 0.14
2.6
1.8
- 2.0
2.1
- 2.4
W-DIST-CR
4.70
+ 0.11
5.35
+ 0.08
5.6
4.5
- 4.8
5.3
- 5.5
W-DIST-CD
3.52
+ 0.12
3.92
+ 0.04
3.9
3.4
- 3.7
3.9 4.0
D-MCON
5.28
+ 0.17
5.70
+ 0.20
5.8
5.1
- 5.5
5.4
- 5.9
D-LCON
4.90
+ 0.13
5.37
+ 0.10
5.6
4.7
- 5.0
5.2
- 5-5
D-ICON
3.32
+ 0.18
3.50
+ 0.l4
3.7
3.1
- 3.6
3.3
- 3.7

185
Family Scolopacidae Vigors, 1825
Remarks. Many of the specimens discussed below have been
arbitrarily assigned to the genus Calidris because of their similarity in
size and overall morphology to this genus. I have been unable to find
generic characters (i. e. non-size related) on the elements here
preserved which will confidently separate Calidris from other genera of
scolopacids. Possibly with study of additional series of all scolopacid
genera, characters could be isolated. But for the present, the following
generic assignments should be regarded as tentative.
Brodkorb (1955a, 1963a, 1967) described four species of scolopacids
from the late Miocene and early Pliocene of FloridaCalidris pacis,
Erolia penepusilla, and Limosa ossivallis from the Bone Valley Mining
District and Ereunetes rayi from the McGehee Farm local fauna. They are
listed below for the sake of completeness, but additional specimens
usually have not been assigned to them, pending further comparisons with
living and fossil species. All specimens are assigned at the species
level on the basis of size. This approach has the effect of
overestimating the diversity of scolopacids now known from in the late
Miocene and early Pliocene of Florida.
Genus Limosa Brisson, 1760
Limosa ossivallis Brodkorb, 1967
Material. Bone Valley Mining District, near Brewster (referred by
Brodkorb, 1955a); PB 526, distal end right tibiotarsus (holotype); PB 527
proximal end right tibiotarsus ("paratype"). Specimens referred by
Becker: Bone Valley Mining District, specific locality unknown; UF
60817, distal end left humerus; UF 61597, left coracoid; UF 6l600,

186
proximal end right tarsometatarsus; Palmetto Mine; UF 21092, distal end
right tibiotarsus.
Remarks. Assignment of additional specimens to Limosa ossivallis
very tentative as most of the skeletal elements are abraded. These
skeletal elements represent a scolopacid of the correct size for this
species (see Brodkorb, 1955a for measurements). The type material, and
the material referred above, should be compared with Limosa vanrossemi L.
Miller, which was described from the Lompoc local fauna of Mohinian age
(=late Miocene) of California. Miller (1925) states that it is closest
to the living Limosa fedoa and shows but slight divergence.
Genus Erolia Vieillot, l8l6
Erolia penepusilla Brodkorb, 1955
Material. Bone Valley Mining District, near Brewster; PB 6ll,
distal end left humerus (holotype).
Remarks. Brodkorb (1955a) found this species to be larger than Erolia
temninckii, E. ruficollis, E. minutilla, Ereunetes pusillus, and
Ereunetes mauri; smaller than Erolia bairdii and E. fusicollis. E.
penepusilla appears to be closest to E. minutilla.
Genus Ereunetes Illiger, l8ll
Ereunetes rayi Brodkorb, 1963
Material. McGehee Farm local fauna; UF 3978, humeral end left
coracoid (holotype).
Remarks. Brodkorb (1963a) found this species to be larger than that
of Ereunetes pusillius, E. mauri, and Erolia minutilla; but smaller than
Erolia bairdii and E. fuscicollis. Ereunetes rayi falls within the same

187
size class as Erolia penepusilla (above) and should be further compared
with this species.
Genus Calidris Merrem, 180^
Calidris pacis Brodkorb, 1963
Material. Bone Valley Mining District, near Brewster; PB 59*+
proximal end left humerus (Holotype).
Remarks. Brodkorb (1955a) states this species is almost identical
in size to Calidris canutus (see Brodkorb, 1955a:22 for measurements),
but differs significantly in the characters of the proximal end of the
humerus. Brodkorb (l955a:22) further suggests that this species may
require generic separation from Calidris when additional material is
known.
"Calidris" sp. 1
Material. Love Bone Bed local fauna; UF 25807, UF 29697, humeral
ends left coracoids; UF 26008, right coracoid; UF 29692, humeral end
right coracoid.
McGehee Farm local fauna; UF 3978, humeral end left coracoid; UF
9J+87, right coracoid.
Bone Valley Mining District, Ft. Green Mine (# 13 dragline); UF
539*+*+, right humerus lacking distal end.
Remarks. Skeletal elements similar in size to those of Calidris
minutilla, C. minuta, C. pusilla, and C. mauri. Slightly smaller than
those of C. subminuta females, C. ruficollis, £. bairdi, and C.
fusicollis; slightly larger than that of C. temnrinckii females.

188
"Calidris" sp. 2
Material. Love Bone Bed local fauna, UF 26018, right coracoid.
Remarks. Coracoid similar in size to that of Calidris melanotos
and morphologically indistinguishable.
"Calidris" sp. 3
Material. McGehee Farm local fauna; UF 31776, distal end left
tarsometatarsus.
Remarks. Distal tarsometatarsus larger than that of Calidris
minutilla and C. ruficollis; smaller than that of £. fusicollis.
Specimen very tentatively referred to genus.
Genus indet. sp. k
Material. Love Bone Bed local fauna, UF 25760, right
carpometacarpus missing minor metacarpus.
Remarks. Carpometacarpus similar to that of Calidris minutilla.
Genus Actitis Illiger, l8ll
??Actitis sp. indet. sp 5
Material. Love Bone Bed local fauna, UF 29701, distal end right
tibiotarsus.
Remarks. Tibiotarsus is about the size of, and very similar in
morphology to Actitis macularia. Species 3 above may belong here.
Assignment to Actitis very tentative.
Genus Arenaria Brisson, 1760
??Arenaria sp. indet. sp. 6
Material. Love Bone Bed local fauna, UF 258l4, UF 29693, humeral
ends left coracoids.

189
Remarks. The fossil coracoids are very similar in size and
morphology to the males of Arenaria melanocephala, but are slightly more
robust. Assignment to genus very tentative.
Genus indet. sp. 7
Material. Love Bone Bed local fauna, UF 25871, distal end left
tibiotarsus; UF 25858, distal end right tarsometatarsus.
Remarks. Fossil skeletal elements similar to those of females of
Limosa fedoa. Brief comparisons with Limosa ossivallis shows the
tibiotarsus from the Love Bone Bed local fauna to have a smaller
transverse width of both the shaft and the anterior portion of the distal
end and is of a slightly smaller size. Additional comparisons are needed
with Limosa vanrossemi from the Miocene of California.
Genus indet. sp. 8
Material. Love Bone Bed local fauna, UF 25810, UF 25822, UF 29695,
UF 29699, left coracoids, broken and abraded; UF 26009, UF 26013, UF
26028, UF 29867, UF 29688, UF 29689, right coracoids, broken and abraded.
Remarks. Coracoids similar in size to that of females of Tringa
flavipes, but slightly more slender. As all coracoids are broken or
abraded to varying degrees, assignment to genus is unwarranted.
Genus Philomachus Merrem, 1804
?Philomachus sp.
Material. Bone Valley Mining District, Ft. Green Mine; UF 60062,
proximal end left humerus.
Remarks. After an extended survey of scolpacid genera, I have found
this humerus (Figure 4.9) from Ft. Green to be most similar to that of
males of Philomachus pugnax. Because Philomachus is only a casual

190
visitor to North America today and has never been noted from Florida, I
hesitate to report it as a fossil until I make additional comparisons
with other recent and fossil species; and can distinguish the proximal
humerus of all genera of scolopacids with certainty by a differential
diagnosis.
Remarks on the Family Scolopacidae.
There are some 20 fossil species of scolopacids, many of which are
in need of revision. Reference to these may be found in Olson (ms) and
Brodkorb (1967). Living species are treated in Johnsgard (1981).

Order Strigiformes (Wagler, 1830)
191
Ordinal Characters. Tibiotarsus without tendinal bridge, with
ligamental tubercle, condyles prominent, nearly parallel, equal in size,
and circular in lateral view (the Psittacidae, the only other New World
family with a tibiotarsus lacking a tendinal bridge, does not have
condyles which are parallel, equal in size and circular in side view).
Family Tytonidae Ridgway, 1914
Characters. In cranial view, tendinal furrow not excavated (deeply
excavated in Strigidae); in caudal view, shaft merges with posterior
intercondylar sulcus evenly (depression present slightly craniad in
Strigidae).
Tytonidae, undescribed genus
Material. Love Bone Bed local fauna. UF 25926, distal end right
tibiotarsus.
Description. Tibiotarsus (Fig. 4.9) distinguished from all species
of Tyto examined (T. alba, glaucops, capensis, sanctialbani, ostologa,
pollens) and Phodilus badius by having a wider anterior intercondylar
sulcus, with the medial condyle sloping gradually into the anterior
intercondylar sulcus, and by having the area intercondylaris much broader
and more shallow (best seen in anterior view of distal end).
Measurements given in Table 4.29
Remarks. Although the above specimen is slightly abraded and is a
solitary specimen, there is no question that it represents a genus
distinct from the two living genera of this family. Additional
comparisons with owls from the early Tertiary are needed to determine the
exact systematic position of this taxon.

192
Remarks on the Family Tytonidae
There have been many fossil species of Tyto described (listed in
Brodkorb, 1971; Olson, in press); based mainly on size differences. The
morphology of the distal end of the tibiotarsus of these fossil species
(T. sanctialbani, T. ostologa, and T. pollens examined) is remarkably
uniform within this genus and is also very similar to that of Pholidus
badius.
There is only one fossil tytonid genus now known. Prosybris
Brodkorb is based on the type species P. antigua (Milne-Edwards, 1863)
from the Aquitanian of St.-Gerand-le-Puy, France. It was described on an
tarsometatarsus and is not directly comparable with the above specimen
from the Love Bone Bed local fauna. The tibiotarsus of Prosybris is
unknown.
A number of undescribed Oligocene owls are known. The above
specimen should be compared with these before allocating the Love Bone
Bed owl to a genus.
J

Table 4.29. Measurements of the tibiotarsi of the following tytonid owls: Tyto albus pratncola (N = l6, 8
males, 8 females), Tyto glaucops (N = 6, 2 males, 1 female, 3 unsexed), Tyto capensis (N = 1, unsexed),
Phodilus badius (N = 2, 1 male, 1 unsexed), and tytonid, undescribed species from the Love Bone Bed local
fauna. Data are mean _+ standard deviation and range. Abbreviations defined in the methods section.
Measurements
T. a. pratncola
T. glaucops
T. capensis
P. badius
LOV
W-DIST-CR
10.92 + 0.40
10.0 11.7
10.18 + 0.53
9.6 10.7
10.0
9.6; 9.2
11.7
D-ICON
4.83 + 0.26
4.5 5.5
4.47 + 0.48
4.0 5.3
4.5
3.5; 3.8
5.3
i
193

Family Strigidae Visors, 1825
Remarks. Femur referrable to the Strigidae by having a smooth
antero-dorsal condyle slope and having the postero-dorsal portion of the
external condyle joining the shaft abruptly (Ford, 1967).
Subfamily Buboninae Vigors, l8l5
Genus Bubo Burneril,l806
(cf.) Bubo sp.
Material. Bone Valley Mining District, Tiger Bay Mine. UF 29782,
distal end left femur.
Remarks. Distal end of the femur intermediate in size between that
of males and females of Bubo virginianus. Agrees with Bubo (and differs
from Strix and Asio) in placement of muscle scars, especially those above
the lateral condyle. Differs from Bubo virginianus and Nyctea scandica
by having the posterio-medial portion of the medial condyle merging
smoothly with the posterior intercondylar sulcus. In other Bubonini
examined, the posterior-medial portion of the medial condyle sharply
overhangs the popliteal fossa.
Remarks on the Family Strigidae.
There is several undescribed strigids awaiting a comprehensive
review. Olson (in press) briefly discusses much of this material.

195
Order Passeriformes (Linnaeus, 1758)
Ordinal Characters. Coracoids separate, lacking coracoidal
fenestra; process small; brachial tuberosity elongate and expanded toward
sternal end of element; head of coracoid usually small, somewhat pointed.
Family Fringillidae Vigors, 1825
Genus Palaeostruthus Wetmore, 1925
Palaeostruthus eurius Brodkorb, 1963
Material. Haile VI local fauna; PB 8502, distal portion of left
tarsometatarsus (holotype).
Remarks. See Brodkorb (1963a) and Steadman (1981) for discussion
and remarks pertaining to this species.
Family Indeterminable
Material. Love Bone Bed local fauna: UF 25806, left coracoid; UF
25T72, distal left femur; UF 29727, 29728, humeral ends right coracoids;
UF 29729, 29730, humeral ends left coracoids.
Remarks. The above specimens represent a minimum of two species
based on size. I have been unable to find characters on the humeral ends
of the coracoid to identify these specimens beyond the level of order.
Characters given by Hamon (1964) do not characterize the suborders of
passerines, although these characters appear adequate to discriminate
between the late Pleistocene and Recent taxa of North America which he
examined. The difficulties in identifying passerines are due to the
large number of genera and the general similarity of morphology within
the order (Frbringer, 1888). This is additionally complicated by the
poorly understood interrelationships between genera and families (Olson,
ms.).

196
Remarks on the Order Passeriformes
There have been very few fossil species of passerines described
(Brodkorb, 1978)- With a few exceptions, still fewer are believably
assigned to a genus or family. Most fossil species were compared to very
few other species and were not adequately diagnosed as a member of the
family in which they were placed.
Nine fossil genera of passerines have been described (Brodkorb,
1978). Miocitta Brodkorb is known only from the late Barstovian Kennesaw
local fauna, Colorado. Protocitta Brodkorb is from the Blancan of Texas
and Kansas and from the Pleistocene of Florida and Texas. Henocitta
Holman was described from the Pleistocene of Florida. All of the above
genera were described as medium to large sized jays.
Palaeoscinus Howard, from Mohanian of Tepsquet Creek, California,
was described from a slab and represents an extinct family of passerines.
Howard (1957) states this family has its affinities with the -
Pycnonotidae, Bombycillidae, Corvidae, and Cinclidae.
Necropsar Slater is based on a postcranial skeleton from the
Holocene of Rodriguez Island. It is placed in the family Sturnidae.
Genera of fossil icterids include Cremaster Brodkorb from the
Pleistocene of Florida, Pandanaris A. H. Miller from the Pleistocene of
Florida and California, and Pyelorhamphus A. H. Miller from the
Quaternary of New Mexico.
Palaeostruthus Wetmore was described as a late Clarendonian to early
Hemphillian genus of emberzid finch from Florida and Kansas. Steadman
(1981) synomized this genus with the living genus Ammodramus.
Additionally, Passerina (cf.) is reported from the Hemphillian Yepomera
local fauna (Steadman and McKittrick, 1982). These last two genera are

197
the only reports of nine-primaried oscines in the Tertiary (Steadman and
McKittrick, 1982).
No fossil passeriformes are recorded before the Miocene (Brodkorb,
1978; Olson and Feduccia, 1979) This could be explained simply as a
sampling bias, but considering the amount of detailed paleontological
field work done both in North America and in Europe, this absence seems
real, at least on these two continents. Prior the Miocene, a great
diversity of small non-passeriforms (mainly Coraciiformes) are known and
they evidently occupied many of the niches which are now filled by
passerines (Mourer-Chauvire^ 1982; Olson and Feduccia, 1979; Feduccia and
Olson, 1982). Feduccia and Olson (1982) noting the great radiation of
suboscines in South America, speculate that they were present in South
America for most of the Tertiary. They also argue that, in addition to
the suboscines, the entire order Passeriformes is of South American
origin.

CHAPTER V
PALEOECOLOGY
Introduction
In some situations, the avian specimens from one locality could be
used to reconstruct the fossil avian communities, and the fossil
environments sampled, using methods similar to those of mammalian
paleoecologists (see Shipman, 1981). Each locality could then be
quantitatively compared to other such fossil localities in North America
throughout the latter half of the Cenozoic. Two insurmountable problems,
sample size and collection technique, prevent such an approach being
applied to the localities included in this study.
Localities from which there are relatively few fossils are not
suited, or are severely limited in usefulness, for quantitative
paleoecological analyses because necessary data were never recorded and
because small forms are often poorly represented (Wolff, 1975). He
showed that when collected in a random manner, approximately 12,000 -
25,000 specimens are needed to represent all members of a mammalian
community, and that about 500 identifiable specimens are needed just to
represent the common members of a community. There is no reason to
I
believe that avian communities can be adequately represented by fewer
specimens, certainly when avian communities are as diverse and complex,
if not more so, than mammalian communities. Simply put, avian
communities cannot be reconstructed definitively from a meager handful of
haphazardly collected fossil bird bones. The problem of sample size
applies to all local faunas included in this study except for twothe
198

199
Love Bone Bed and Bone Valley. Although these two localities have large
samples of fossil birds, the techniques used to collect the fossil
vertebrates prevent a quantitative paleoecological analysis. Hence I am
limited to qualitative statements about the paleoecology of each of the
local faunas considered in this investigation. These statements are
based primarily on the birds present, the habitats used by their Recent
congeners (Blake, 1977; Palmer, 1962, 1975, 1976; Terres, 1980), and on a
general knowledge of the geology of the locality and its vertebrate
fauna. In general, the paleoecological reconstructions based on fossil
birds are similar to those based on other fossil groups.
Local Faunas
Love Bone Bed local fauna. The avifauna of the Love Bone Bed is
richly aquatic, with but a minor influence of more terrestrial species.
Taxa presently identified include 2 species of grebe, a species of
cormorant and one of anhinga, 3 species of heron, 2 species of~stork, 2
ibses, one vulture, one osprey, 2 accipitrids, 4 species of geese, 4
species of duck, a turkey, 3 cranes, 3 rails, 2 flamingos, a jacana, 7
species of shorebirds, a barn owl, and 2 species of passerine.
The more abundant species of those listed above include the grebe,
Tachybaptus sp.; a tree-duck, Dendrocygna sp.; a tiny species of teal,
Anas sp. A.; a larger duck, Anas near A. acuta; a crane, Grus sp. B.; two
species of rail, one a species of Rallus cf. sp. C., and the other an
undescribed genus; 2 flamingos in the genus Phoenicopterus; a jacana,
Jacana farrandi; and a indeterminate genus and species of scolopacid.
These abundant taxa suggest that more than one type of habitat has
been sampled. Modern species of grebe typically occur in freshwater,
from lakes to shallow ponds. Jacanas occur in freshwater marshes to

200
shores of rivers and often in ponds with floating vegetation. Species of
Dendrocygna may also be found in shallow ponds with floating vegetation,
but they have a greater tolerance to brackish water. Scolopacids may be
found in grassy marshes, mudflats, estuaries, and edges of ponds.
Flamingos are found in shallow water to mudflats. It would seem likely
therefore, that the environments around the Love Bone Bed during the time
of deposition would include freshwater ponds and streams, but probably
also with wet marshes, streams, estuaries, and mudflats nearby.
Although the avifauna from the Love Bone Bed is substantial, the
usefulness of this collection for qualitative paleoecological analysis is
limited by collection techniques. As in most fossil vertebrate
localities, this site was excavated to maximize the number of mammalian
specimens recovered per unit time spent digging. While sediment samples
were screenwashed for small specimens, it was not done in a systematic
fashion. By using these methods, many of the smaller specimens
apparently were never collected. In the original study, only two one-
quarter cubic meter samples were collected for paleoecological analysis
(Webb et al. 1981: 553ff). Each of these small samples was from a
different stratigraphic unit; vertebrate remains were reported by weight
only with no indication of number of specimens given, or of the weight of
the sediment.
It should also be noted that the MNI (minimum number of individuals)
of birds as given by Webb et al. 1981: 538) based on preliminary
identifications is not MNI but rather the estimated number of species of
birds. Neither MNI nor number of specimens are valid indicators of
abundance when samples are strongly biased toward larger specimens. It

201
seems likely that birds with large limb bones (flamingos, cranes) are
better represented in the collections than really existed either in the
deposit or in the fossil community preserved.
Mixson Bone Bed local fauna. Avian taxa now known from here include
two species of grebe (Rollandia sp. and Podilymbus sp. A); a large
stork, Ciconia sp. B; unstudied specimens of a large crane (?Grus sp. B),
and an anhinga, Anhinga grandis. These taxa suggest a shallow freshwater
environment with emergent vegetation surrounded by marshes or "wet
prairies". The lack of other avian species, and the lack of a greater
number of specimens should not be considered significant. This locality
was excavated primarily for the remains of large mammals and only a few
specimens of small vertebrates exist in collections.
McGehee Farm local fauna. Avian taxa known from the McGehee Farm 1.
f. include 2 species of grebe (Rollandia sp. and Tachybaptus sp.); an
undetermined species of cormorant; the anhinga, Anhinga grandis; the
night-heron, Nycticorax fidens; 2 species of duck, a small species of
teal and the other near Anas acuta in size; an undescribed genus of rail;
a flamingo, Phoenicopterus sp.; a jacana, Jacana farrandi; and three
shorebirds, probably all in the genus Calidris. None are abundant, but
the cormorant and the ducks are the most common. Based on the occurrence
of a similar set of avian taxa, the environments surrounding McGehee Farm
during deposition were probably very similar to those at the Love Bone
Bed.
Withlacoochee River kA local fauna. Only two avian species have
been recovered from this Hemphillian site. They consist of a single
femur representing an indeterminate species of Buteo about the size of

202
Buteo jamaicensis and the holotype of Egretta subfluvia, am egret about
the size of Egretta ibis. This locality probably represents a pond
environment with some marine influence (Becker, 1985a).
Haile VB local fauna. The meager avifauna from this local fauna
consists of several specimens of an indeterminate species of anatid. The
paleoenvironment of this site was highly aquatic as shown by the abundant
material of the crocodylian, Gavialosuchus.
Haile VI local fauna. Birds known from here include the holotype of
Palaeostruthus eurius and several specimens of an indeterminate species
of duck. Based on the birds, little can be said of the paleoecology of
this locality.
Haile XIXA local fauna. Avian taxa from here include a cormorant
and a few specimens of anatids. Further studies on the paleoecology of
this site await additional systematic studies on the vertebrates present.
Bone Valley Mining District. While large numbers of fossil birds
have been collected from here, it represents the most limited for
paleoecological analysis because of collection techniques. This mining
district, which is well over 100 square miles in extent and is being
systematically strip-mined, resulting in the moving of hundreds of
thousands of cubic yards of sediment. Many specimens of fossil birds
were collected from spoil areas in conjunction with the commerical
collection of the more numerous shark teeth and mammalian cranial
fragments. Because of these collection techniques, usually it is not
possible to quantify the relative abundance of fossil specimens of birds,
the volume of sediments from which they originated, the sedimentary

203
horizon from which they came, or their association with other fossil
vertebrates.
Like the Love Bone Bed, the Bone Valley avifauna is dominated by
aquatic birds. However, in sharp contrast, the most abundant taxa
present are marine. Birds from the Bone Valley Mining District included
in this study are 3 species of grebe, 2 species of cormorant, an anhinga,
2 herons, 2 storks, an ibis, an osprey, 4 species of hawks or eagles, a
goose, 4 species of duck, a turkey, a crane, a rail, a flamingo, 5
species of shorebird, and an owl. There is abundant material of
Phalacrocorax wetmorei, Aythya sp., and Phoenicopterus floridanus. Other
pelagic species, which were not included in this study, but are
abundantly represented, include several (3?) species of alcids, loons,
Larus elmorei, 2 species of Sula, and a species of Morus. Fossil
material still occurs in about the same proportions as was reported by
Brodkorb (1955a), even though the number of taxa in this avifauna have
increased. By far the most common taxon in this deposit is the
cormorant, Phalacrocorax wetmorei. The abundance of cormorants, sulids,
and gulls argues for a near-shore marine environment.
Brodkorb (1955) suggested that the large concentrations of seabirds,
especially cormorants, probably represented a breeding colony. For
future consideration of this hypothesis, I note that there is a very low
frequency of sub-adult specimens in the collections from Bone Valley. I
have been unable to find comparable data on Recent breeding colonies.
It has also been suggested (Brodkorb, 1955; and references therein)
that the large concentrations of fossil birds were responsible for the
formation of the phosphorite deposits. Current hypotheses (Riggs, 1984)
suggest that bacteria at the water-sediment interface are responsible for

20h
the primary formation of phosphate grains from the upwelling of nutrient-
rich waters. I suggest that the fossil birds in the Bone Valley Deposits
are present as a result of the abundant food supplies in these nutrient-
rich waters, rather than being the direct cause of it.
Manatee County Dam Site. The only avian taxon known from this local
fauna is one specimen representing Phalacrocorax cf. P. wetmorei. Based
on the available information pertaining to the geology, vertebrate fauna,
and geographic location, this locality is in all aspects essentially an
outlier of the Bone Valley local fauna discussed above. All comments
pertaining to the paleoecology of Bone Valley also apply to this local
fauna.
SR-6U. Fossil birds from this locality include the cormorant
Phalacrocorax wetmorei, loons, the flamingo, Phoenicopterus cf. P.
floridanus, and alcids. As with the Manatee County Dam Site above, all
available information about the geology, vertebrate fauna and geographic
location indicates this locality is an outlier of the Bone Valley local
fauna. See paleoecological comments under Bone Valley Mining District
above

CHAPTER VI
BIOCHRONOLOGY AND FAUNAL DYNAMICS
Introduction
The following analysis of the biochronology and faunal dynamics of
the Neogene (23 nybp to 1.8 mybp) avifauna of North America takes much of
its information from an accompanying project (Becker, ms.), which updates
and reviews the Neogene records of the North American avifauna. This
fossil record is still very incomplete in many areas, and its
intrepretation will doubtlessly change as new material becomes available
and previously described material is restudied. I should also note that
the current record of fossil birds suffers from decades of a typological
systematic approach, with many species, and even genera, being based more
on geography and on geologic age than on differences in morphology.
Faunal Dynamics
Remarks. Among the assumptions necessary to calculate faunal
dynamic parameters are the following:
(1) All taxa are correctly identified, are taxonomically valid, and
are correctly placed systematically.
(2) The stratigraphic context of the fossil specimens is known and the
specimen is correctly placed in a local fauna.
(3) The local fauna is correctly placed in geologic time.
(U) Taxa are correctly divided into marine and non-marine groups.
In one or another instance, each of these assumptions is surely
violated. Much work still remains to be done to verify some of the
205

206
supposed relationships of fossil birds suggested by previous authors and
to correct erroneous locality information. However, the value of such
investigations on fossil birds include:
(1) Demonstration of general trends in avian faunal dynamics in the
Neogene of North America. Major trends should still be evident,
even though they are based on incomplete or partially correct data.
(2) Taxa which do not parallel the general trend may be selected for
further investigations.
(3) Specific geographic areas and/or time intervals can be featured in
future work.
(U) Specific groups can be easily isolated, such as those of interest
from a zoogeographical standpoint.
Formulae used to calculate the parameters appearing in these table
are presented and defined in the methods section, and abbreviations are
given in Table 6.1. The separation of taxa into marine and non-marine
groups is based on the habitats used by the majority of living congeners.
The Local Faunas
The Neogene localities that have produced avian specimens are not
uniformly distributed in geologic time. Of 133 local faunas surveyed,
3.87 (5) are late Arikareean, 8.3 1 (ll) are Heraingfordian, 13.5 % (18)
are Barstovian, 22.6 % (30) are Clarendonian, 30.1 % (4o) are
HemphiIlian, and 21.8% (29) are Blancan in age. Or stated slightly
differently, 'J'h.hlo (99/130) of the localities examined are from the last
4l.O % (Clarendonian to Blancan; 8.7 MA / 21.2 MA) of the Neogene. If
only the published localities are used, instead of the total number of
localities, the percentages are roughly comparable, but the absolute
values are much lower (see Table 6.2).

207
An index of relative sampling may be calculated to adjust for the
unequal lengths of time represented by these discrete NALMAs (North
American Land Mammal Ages) and to take into account the number of
sampling sites. The number of local faunas of a given NALMA are divided
by the duration of that NALMA (Table 6.2). This shows that the last
three NALMAs of the Neogene are better sampled than the first three by a
factor of 2 or 3.
When the local faunas are further divided into marine and non-marine
groups, even greater discrepancies in the representations of the NALMAs
are apparent. In general the non-marine local faunas outnumber the
marine local faunas by at least a factor of 3 to 1 (Table 6.2; 6.3; and
Figure 6.U).
Additional comparisons can be made between the number of published
versus unpublished localities. While these comparisons are biased by the
collections I have been able to examine, marine avifaunas have.been more
completely described than terrestrial ones (compare Table 6.3; 6.L).
The fossil avifaunas from nearly one-half of all terrestrial local faunas
in which birds are present are entirely unstudied and unreported in the
literature. As all major collections in the United States were not
examined, this is certainly an underestimate of the amount of work left
to be done and underscores the preliminary nature of this examination of
faunal dynamics.
The Neogene Avifauna
The combined sample of all families and genera will be considered
first, then reconsidered separately in marine and non-marine groups.

208
Families. The easiest statistic to calculate is the number of
families present in a given NALMA. In the late Arikareean 8 familes are
present of which 5 (63%) survive to the present. The number of families
rapidly increase to 18 (IT surviving; 94^) in the Hemingfordian, 31 (30
surviving; 97%) in the Barstovian, 37 (36 surviving; 97%) in the
Clarendonian, 40 (39 surviving; 98%) in the Hemphillian, and 4l (40
surviving; 98%) in the Blancan. By the Barstovian a majority of the
living families with a fossil record have appeared (Table 6.1).
Genera. The diversity (Si) of the genera in the Neogene of North
America begins with 10 (2 surviving; 20%) in the late Arikareean, 28 (ll
surviving; 39%) in the Hemingfordian, 38 (22 surviving; 56%) in the
Barstovian, 6l (43 surviving; 70%) in the Clarendonian, 78 (65 surviving;
83%) in Hemphillian, and increases to 98 (88 surviving; 90%) in the
Blancan.
Originations (Oi) and Extinctions (Ei) are given in Table 6.2.
These parameters are the simple counts of the first and last appearance,
respectively. Nine genera first appear in the late Arikareean. This
increases to about 20 in both the Hemingfordian and Barstovian and then
increases again to about 30 in the Clarendonian, Hemphillian, and
Blancan. Five extinctions occur in the late Arikareean, then 8
extinctions occur in each of the succesive NALMAs, with the exception of
the Clarendonian, which records 12 extinctions.
The raw counts of origination and extinctions are adjusted for the
unequal time interval in each of the NALMAs by dividing each count by the
duration of the given NALMA to produce Origination rates (Or) and
Extinction rates (Er) in Table 6.2.

209
Origination rates show a peak during the Hemingfordian, a very large
increase in the Clarendonian, a decrease in the Hemphillian, although not
to pre-Clarendonian level, then an additional increase in the Blancan.
Extinction rates tend to he much lower than origination rates, hut show a
roughly parallel trend. Again, a high value is noted in the
Clarendonian.
Turnover rates (T), or the numerical average of origination rates
and extinction rates, predictably parallel the trends in origination and
extinction rates described above. Running means (Rm), or the diversity
minus the average of originations and extinctions of a given land mammal
age, increase at a decreasing rate throughout the Neogene.
Turnover rates per genus (T/Rm) show a high value in the late
Arikareean, decrease through the Barstovian, then peak in the
Clarendonian, to decrease and show a low rate through to the end of the
Neogene.
Marine versus Non-marine
A more instructive approach is to divide the North American avifauna
into marine and non-marine subgroups and then compare and contrast these
two broad divisions (Table 6.3; 6.U and Figure 6.1; 6.2). The following
observations result.
(l) The North American marine avifauna is essentially established at a
diversity of 20 to 25 genera by the Clarendonian. The North
American non-marine avifauna is roughly stable at a diversity of 3
genera during the late Arikareean and Hemingfordian. From this
NALMA through the remainer of the Neogene, diversity increases from
13 to 26 genera by the Blancan.

210
(2) The absolute number of genera which become extinct is roughly stable
over most of the Neogene with about 2 generic extinctions per
million years in the marine avifauna and about 7 generic extinctions
per million years in the non-marine avifauna.
(3) In the marine avifauna, the rate of origination peaks during the
Clarendonian with 4.4 new genera appearing per million years.
(4) In the non-marine avifauna, there are peaks in the generic
origination rates during the Hemingfordian, Clarendonian and Blancan
Land Mammal Ages, with 6, 8, and 10 genera appearing per million
years, respectively. These peaks rise above a background rate of 2
to 4 genera appearing per million years in the intervening NALMAs,
or roughly a cyclic 2- to 4-fold change.
(5) Extinction rates are typically low in the marine avifauna throughout
the Neogene. With the exception of the Clarendonian, which shows an
extinction rate of 2 genera per million years, all other NALMAs have
an extinction rate much less than 1.0 extinction per million years.
In the non-marine avifauna, extinction rates average from 1.0 to 2.8
generic extinctions per million years.
(6) Turnover rates are consistently low in the Marine avifauna, with the
highest turnover rate being in the Clarendonian. In non-marine
avifauna, turnover peaks appear in alternating mammal ages (in the
Hemingfordian, Clarendonian, and Blancan). These peaks in turnover
represent a 2- to 3-fold increase over the rates in the intervening
NALMAs. Fluctuations in the turnover rates are governed more by
changes in origination rate than by changes in extinction rates.
(7) The per-genus turnover rate (T/Rm) in marine avifauna is high in the
late Arikareean (1.68) and then drops to approximately 0.20 through

211
the Clarendonian. In the Hemphillian and the Blancan it decreases
again to less than 0.1. In the non-marine avifauna the turnover
rate is 0.6 and 0.4 in the late Arikareean and Hemingfordian
respectively, and a rate of 0.2 in the Clarendonian. Other mammal
ages have a rate of 0.1 or less. The extremely high per-genus
turnover rate in the Arikareean may be an artifact of the low
sample size in this NALMA.
Discussion
Several factors could produce the results presented above, including
real changes in fossil avifaunas, but also including invalid assumptions
or errors in sampling. Certainly an obvious cause for some of the
changes in faunal dynamics is the unequal distribution of localities
within each mammal age. The absence of extinctions in the Hemingfordian
and Barstovian is probably an artifact resulting from an impoverished
prior record of taxa. The Clarendonian is the first NALMA in ihe Neogene
with a large number of both marine and non-marine localities. This
accounts at least for part of the peak in Or (marine) and the resultant
peak in turnover.
The mammalian fossil record in the Neogene has long been known to
consist of sequences of relatively long-lived "chronofaunas" whose
boundaries do not coincide with the boundaries of the NALMAs. It is
equally possible that birds exhibit a similar pattern of long periods of
faunal stasis with relatively rapid periods of turnover at the ends of
these periods. There is no a priori reason that these "avian" boundaries
would correspond with the "mammalian" boundary of the NALMA, or even with
the boundary of the mammalian chronofauna. Further work is needed to
reconstruct the database of Neogene localities from one keyed into

212
relative time (NALMAs) to one keyed to absolute time. This database
could then be clustered into "natural" groups of geological ranges of
species to discriminate between the lack of localities and real turnover.
Vuilleumier (198*0 examined the faunal turnover of the South
American avifauna and has recently (in press) expanded his investigations
to include the late Neogene North American fossil record of birds. It is
difficult to make comparisons between his study and the results presented
above because of the time scales used. The faunal parameters which
Vuilleumier (1984, in press) presents, are based on a division of
geologic time into Epochs, which have vastly uneven durations (Miocene,
IT MA; Pliocene, 2.5 MA). Many of the finer scale events presented above
would not be apparent in Vuilleumier's parameters.
The faunal dynamics of mammals have been extensively investigated
(Webb, 1976, Marshall, et al., 1982). Table 6.5 compares the most recent
information on faunal dynamics of mammals (Marshall, et al., 1982) with
those for birds. Generic extinction rates, generic origination rates,
and turnover rates are substantially higher for mammals than for birds.
One possible explanation is that mammals are evolving at a much
faster rate than are birds. However, considering the great diversity of
the Recent avifauna and the comparable amount of morphological difference
between equivalent taxonomic ranks of birds and mammals (Wyles et al.,
1983), this explanation seems at best, only partial.
A more likely explanation is that the differences in faunal dynamic
parameters between birds and mammals are due to the use of different
types of taxonomic characters. In systematics of fossil avian species,
the post-cranial skeleton is used almost exclusively. Conversely, in
fossil mammals, dental morphology is usually used. As the post-cranial

213
skeleton is likely to be much more conservative as a systematic character
than tooth morphology, a slower turnover rate in birds would not be
unexpected.
Biochronology
The following preliminary list of taxa represents the first attempt
at using fossil birds as biochronologically useful taxa. This list is
based on the current record of avian genera from the Neogene of North
America, with taxa of questionable validity being omitted. The following
list will doubtlessly change and increase as our knowledge of avian
evolution in the Neogene becomes more complete.
LATE ARIKAREEAN. First Appearance: Morus.
Last Appearance: Plotopteridae.
HEMINGFORDIAN. First Appearance: Puffinus, Anhinga, Sula,
Dendrocygnini, Anatini, Palaeoborus, Neophrontops, Falco, Rallidae,
Megapaloelodus, Burhinus, Strigidae. Last Appearance: None identified.
BARSTOVIAN. First Appearance: Gavia, Podicipedidae, Diomedea,
Fulmarus, Phalacrocorax, Microsula, Qsteodontornis, Ardea, Anserini,
Mergini, Vulturidae, Pandion, Rallus, Laridae, Stercorariidae, Alcinae,
Corvidae. Last Appearance: None identified.
CLARENDONIAN. First Appearance: Rollandia, Tachybaptus, Oceanodroma,
Miosula, Plegadis, Egretta, Ardeola, Mycteria, Ciconia, Pliogyps, Flica,
Grus, Limosa, Phoenicopterus, Jacana, Alca, Cepphus, Cerorhinca, Uria,
Aethia, Praemancalla, Tytonidae. Last Appearance: Qsteodontornis,
Microsula

21k
HEMPHILLIAN. First Appearance: Podiceps, Podilymbus, Pelecanus,
Eudocimus, Nycticorax, Cygnini, Bucephala, Oxyura, Haematopus, Calidris,
Larus, Pinguinus, Manca 11a. Last Appearance: Pelagornithidae,
Balearicinae, Premancalla, Megapaloelodus.
BLANCAN. First Appearance: Aechmophorus, Botaurus, Colinus, Meleagris,
Ptychoramphus, Titanis, Sterna. Last Appearance: Miosula, Pliogyps,
Mancalla.

Table 6.1. Number of families and genera present in each North American Land Mammal Age and the percentage
of these which are still living. See text for discussion of the division of families and genera into marine
and non-marine groups. Abbreviations: L. ARIK. late Arikareean (23.0 20.0 nybp), HEMING.
Hemingfordian (20.0 16.5 mybp), BARST. Barstovian (16.5 11.5 mybp), CLAR. Clarendonian (11.5 9*0
nybp), HEMP. Hemphillian
(9*0 4.5 mybp)
, BLAN. Blancan (4.5
- 1.8 mybp).
North American
genera
which lack a fossil record ¡
are omitted from
this table.
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Number families present
Marine
3
3
9
10
12
11
Non-marine
5
15
22
27
28
30
Total
8
18
31
37
40
4i
Number (%) living families
Marine
1 (33)
2 (66)
8 (89)
9 (90)
11 (92)
11 (100)
Non-marine
4 (80)
15 (100)
22 (100)
27 (100)
28 (100)
29 (97)
Total
5 (63)
17 (94)
30 (97)
36 (97)
39 (98)
40 (98)
Number genera present
Marine
3
3
12
21
26
26
Non-marine
7
25
26
40
52
72
Total
10
28
38
6l
78
98
Number {%) living genera
Marine
1 (33)
1 3 (100)
9 (75)
14 (67)
22 (85)
24 (92)
Non-marine
1 (14)
8 (32)
13 (50)
29 (73)
43 (83)
64 (89)
Total
2 (20)
11 (39)
22 (58)
43 (70)
65 (83)
88 (90)
215

Table 6.2 Faunal dynamics
of the North American Neogene
avifauna.
Abbreviations as in
Table 6.1
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Duration (MA)
3
3.5
5
2.5
4.5
2.7
Localities (Published)
5 (5)
11 (6)
18 (12)
30 (22)
4o (25)
29 (16)
Sampling Index
1.67
1.71
2.40
8.80
5.56
5.93
Number of genera (Si)
10
28
38
6l
78
98
Originations (No.)
9
23
18
31
29
29
Extinctions (No.)
5
8
8
12
9
7
Running mean (Rm)
3.00
12.50
25.00
39.50
59.00
80.00
Origination Rate
3.00
6.57
3.60
12.40
6.44
10.74
Extinction Rate
1.67
2.29
1.60
4.80
2.00
2.59
Turnover Rate (T)
2.34
4.43
2.60
8.60
4.22
6.67
T/Rm
0.78
0.35
0.10
0.22
0.07
0.08
T/Si
0.23
0.16
0.07
0.14
0.05
0.07
I
216

Table 6.3 Faunal dynamics of the marine Neogene birds from North America. Abbreviations as in Table 6.1
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Duration (MA)
3
3.5
5
2.5
4.5
2.7
Localities (Published)
3 (3)
1 (1)
3 (3)
8 (8)
11 do)
1(1)
Sampling Index
1.00
0.29
0.60
3.20
2.22
0.37
Number of genera (Si)
3
3
12
21
26
26
Originations (No.)
3
2
9
11
10
2
Extinctions (No.)
2
0
2
5
2
2
Running mean (Rm)
0.50
2.00
6.50
13.00
20.00
24.00
Origination Rate
1.00
0.86
1.80
4.40
2.22
0.74
Extinction Rate
0.67
0.00
0.40
2.00
0.44
0.74
Turnover Rate (T)
0.84
0.43
1.10
3.20
1.33
0.74
T/Rm
1.68
0.21
0.17
0.25
0.07
0.03
T/Si
0.28
0.l4
0.09
0.15
0.05
0.03
i
217

Table 6.4 Faunal dynamics of non-marine Neogene birds of North America. Abbreviations as in Table 6.1.
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Duration (MA)
3
3.5
5
2.5
4.5
2.7
Localities (Published)
2 (2)
10 (5)
15 (9)
22 (14)
29 (15)
28 (15)
Sampling Index
0.67
1.43
1.80
5.60
3.33
5.56
Number of genera (Si)
7
25
26
4o
52
72
Originations (No.)
6
21
9
20
19
27
Extinctions (No.)
3
8
6
7
7
5
Running mean (Rm)
2.50
10.50
18.50
26.50
39-00
56.00
Origination Rate
2.00
6.00
1.80
8.00
4.22
10.00
Extinction Rate
1.00
2.29
1.20
2.80
1.56
1.85
Turnover Rate (T)
1.50
U.15
1.50
5.40
2.89
5-93
T/Rm
0.60
0.40
0.08
0.20
0.07
0.11
T/Si
0.21
0.17
0.06
0.14
0.06
0.08
i
218

Table 6.5 Comparisons of avian and mammalian faunal dynamics. Avian parameters include only non-marine
faunas, from Table 6.4. Mammalian parameters are from Marshall et al. (1982). The mammalian parameters for
the Hemphillian and Blancan are recalculated to account for the use of a different duration of these NALMAs.
AVIAN
CLAR.
HEMP.
Running mean (Rm)
26.50
39.00
Origination rate (Or)
8.00
4.22
Extinction rate (Er)
2.80
I.56
Turnover rate (T)
5.40
2.89
Per-genus Turnover (T/Rm)
0.20
0.07
T/Si
0.14
0.06
BLANC.
CLAR.
MAMMALIAN
HEMP.
BLANC.
56.00
52.00
50.00
52.00
10.00
17.20
16.67
20.00
1.85
14.80
18.00
l4.8l
5-93
16.00
17.34
17.41
0.11
0.31
0.35
0.33
0.08
0.17
0.14
0.18
1
219

Figure 6.1. Distribution of avian genera and localities through
geologic time. Abbreviations as in Table 6.1.

NUMBER OF GENERA
221
GENERA LOCALITIES
L ARIK HEMING BARST 1 CLAR 1 HEMP 1 BLANC
NUMBER OF LOCALITIES

Figure 6.2. Graphic representation of avian faunal dynamic
parameters. A. Number of localities per million years per NALMA.
B. Running mean per NALMA. C. Extinction rate per NALMA. D.
Origination rate per NALMA. Abbreviations as in Table 6.1.

8
4
O
60
40
20
0
2
0
8
4
0
223
NON-MARiNE
MARINE
A
^ m m
L ARIK HEMING BARST

CHAPTER VII
SUMMARY
This study has examined three aspects of avian paleontology
systematics, paleoecology, and biochronology and faunal dynamics.
Systematics
It has first focused on the systematics of the non-marine fossil
birds from the late Miocene and early Pliocene of Florida. 78 taxa have
been identified in this study. Grebes (Family Podicipedidae) are
represented by 6 taxa. A species of Tachybaptus is abundant in the Love
Bone Bed local fauna and is also present from McGehee Farm. Rollandia is
known from a few specimens from the Love Bone Bed, Mixson, and McGehee
Farm. The skeleton of this species is slightly more robust than that of
the living Rollandia rolland chilensis. A small species of Podilymbus is
known from Mixson. The Bone Valley Mining District has produced
specimens of 3 grebesPodilymbus cf. JP. podiceps, Podiceps sp. and
Pliodytes lanquisti. All are rare members of the Bone Valley avifauna.
Pelecaniformes were represented by 2 or possibly 3 species of
cormorants (Family Phalacrocoracidae) and 1 or possibly 2 species of
anhinga (Family Anhingidae). The cormorant Phalacrocorax wetmorei is one
of the best represented Neogene fossil species, with well over 500
specimens known. Almost every skeletal element is known.
Phalacrocorax sp. A. is known from the Love Bone Bed, McGehee and
probably Haile XIXA. The anhinga, Anhinga grandis, is known from the
22 k

Love Bone Bed and from McGehee Farm. It is larger than the living New
World anhinga and is fairly completely known.
Six species of heron (family Ardeidae) are present. These include 2
species of Ardea, at least 2 species of Egretta, a species of Ardeola,
and a single species of Nycticorax. Herons are rare members of fossil
avifaunas.
Four storks are present as are 3 species of ibis. Storks (Family
Ciconiidae) include a species of Mycteria from the Love Bone Bed and
McGehee and 3 species of Ciconia, distributed between the Love Bone Bed,
Mixson Bone Bed, and Bone Valley. This shows that storks were more
diverse in the late Miocene and early Pliocene of North America than they
are today. Ibises (Family Plataleidae) include a species of Eudocimus
from the Bone Valley Mining District, and a specimen of Plegadis cf. P.
pharangites and one representing a large species of indeterminate ibis
from the Love Bone Bed.
Nine taxa of accipitriform birds are present, including a species of
New World Vulture (Family Vulturidae), 1 or possibly 2 species of Osprey
(Family Pandionidae), and 7 species of hawk or eagle (Family
Accipitridae). Of note from the Love Bone Bed is an undescribed species
of vulture, Pliogyps, and the most primitive species of osprey, Pandion
lovensis, now known. An indeterminate species of Pandion, ?Haliaeetus
sp., Aguila sp. A., and an indeterminate genus are all known from the
Bone Valley Mining District. A species of Buteo near the size of Buteo
jamaciensis occurs in the Withlacoochee River 4a local fauna.
Waterfowl (Family Anatidae) are common, with 5 species of geese and
8 species of ducks being present. Of note is a well-represented species
of Dendrocygna from the Love Bone Bed which cannot be distinguished from

226
the living species of this genus. A very small species of teal, Anas sp.
A., is also abundant in the Love Bone Bed local fauna. This species may
represent the smallest species of the genus Anas now known. Other
species of ducks and geese are common in these localities. A tadorine
duck is known from one specimen from the Bone Valley Mining District.
Gallinaceous birds are poorly represented. Two species of turkeys
(Family Phasianidae), each known from a single specimen, are known. One
is referable to the genus Meleagris and the other is not referable to a
genus.
Gruiform birds are also common with 4 species of crane (Family
Gruidae) and b species of rail (Family Rallidae) being known. Of note is
an undescribed species of primitive rail, which is known from abundant
material from the Love Bone Bed and from a few specimens from the McGehee
Farm local fauna. A specimen of a balearicine crane from the Bone Valley
Mining District represents the last occurrence of this subfamily of
cranes in North America. Today this subfamily is only known from Africa
south of the Sahara. Cranes are also shown to be much more diverse in
North America in the Miocene than in Pleistocene or Recent avifaunas.
Two species of flamingos (Family Phoenicopteridae; genus
Phoenicopterus) are known. They are present in the Love Bone Bed local
fauna, the McGehee Farm local fauna and from the Bone Valley. At least 2
different species of flamingo survived in North America until the
Pleistocene.
A jacana, or lily-trotter (Family Jacanidae), Jacana farrandi, is
known from the Love Bone Bed and from McGehee Farm. This distinctive
genus is usually not found north of Mexico today.

227
A number of shorebirds (Family Scolopacidae) are known. Most are
small and are tentatively referred to the genus Calidris.
Two owls are known. One is an undescribed genus (Family Tytonidae)
which appears to be related to the barn owl and the grass owl of south
east Asia. Another (Family Strigidae), probably in the genus Bubo, is
known from Bone Valley.
Three perching birds (Order Passeriformes) are present. All are
poorly known.
The most diverse localities are the Love Bone Bed local fauna with
approximately 44 taxa and the Bone Valley Mining District with
approximately 4l taxa, 31 of which were studied in this paper. These
localities are the most diverse non-marine and marine avifaunas,
respectively, known in North America prior to the Pleistocene. Other
localities are much less diverse. Approximately l4 taxa are represented
at McGehee Farm, while the other localities included in this study
typically have fewer than 3 or 4 taxa present.
Paleoecology
The second aspect of this study focused on paleoecology. Birds from
two environments dominate the late Miocene and early Pliocene localities
in Florida. Birds from the Bone Valley Mining District, SR-64, and
Manatee County Dam are interpreted as being from near-shore marine
environments. Abundantly represented taxa from other localities, such as
the Love Bone Bed and McGehee Farm, are interpreted as representing
freshwater ponds and streams, with marshes, mudflats, and estuaries not
far removed. More terrestrial localities are either very poorly
represented or lacking for this time period in Florida.

228
Biochronology and Faunal Dynamics
The final aspect of this dissertation focused on the biochronology
and faunal dynamics of the Neogene North American fossil record of birds.
From the examination of 133 Neogene localities which have produced fossil
birds, I show that the fossil localities which have produced fossil birds
are not uniformly distributed in time, with nearly 75% of the localities
occurring in the last 4l% of the Neogene. A majority of the living
families of North American birds which have a fossil record appear by the
Barstovian. Generic diversity increases throughout the Neogene. The
marine avifauna is essentially established by the Clarendonian at a level
of 20 25 genera, while the non-marine avifauna increases continually.
Generic origination rates peak for marine birds in the Clarendonian, but
extinction rates remain low throughout the Neogene. Generic origination
rates for non-marine birds show a cyclic nature every alternate Land
Mammal Age.
A preliminary list of biochronologically important genera of birds
is presented. As our knowledge of the North American Neogene avifauna
expands and becomes more complete, this list can be revised and updated.

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243
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wide inventory of avian anatomical specimens. Auk, 99:740-757.

BIOGRAPHICAL SKETCH
Jonathan J. Becker was born in Jerome, Idaho, on 16 December 1955
and was raised on a nearby farm. He graduated from the Jerome High
School in May 1974 and attended Idaho State University in Pocatello,
Idaho, from 1974 to 1980; receiving a B. S. in zoology with high honors
in May 1978 and a M. S. in zoology/biology in May 1980. Since August
1980 he has attended the University of Florida, graduating with a Ph. D.
in zoology in August 1985* He is currently a postdoctoral fellow at the
National Museum of Natural History, Smithsonian Institution.
He is a member of the American Association for the Advancement of
Science, American Ornithologists' Union, American Society of
Mammologists, Biological Society of Washington, Cooper Ornithological
Society, Florida Acadeny of Science, Sigma Xi, Society of Systematic
Zoology, Society of Vertebrate Paleontology, and Wilson Ornithological
Society.
His current research interests include the functional morphology of
birds and mammals and the evolution, systematics, and biochronology of
the Neogene birds of North America.
245

I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
c / 1
Pierce Brodkorb, Chairman
Professor of Zoology
I certify that I have read this study and that in ny opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
udmi
Richard A. Kilti
Assistant Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
S. David Webb
Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as^su-dssBrt^.tion f£>ir~^i^degree of Doctor
of Philosophy.
Ronald G. Wolff
Associate Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
<7
f
Elizabeth
Professor
S. Wing
of Anthropology
This dissertation was submitted to the Graduate Faculty of the Department
of Zoology in the College of Liberal Arts and Sciences and to the
Graduate School, and was accepted for partial fulfillment of the
requirements of the Doctor of Philosophy.
August 1985
Dean for Graduate Studies and Research



83
Table 4.9 Measurements of the femora of the cormorants Phalacrocorax
auritus auritus (N = 14, 7 males, 7 females), Phalacrocorax auritus
floridanus (N 18, 9 males, 9 females), and Phalacrocorax vetmorei,
from the Bone Valley Mining District. Data are mean +_ standard
deviation, (N), and range. Abbreviations are defined in the methods
section.
Measurements P. a. auritus P. a. floridanus P. wetmorei
Femur
M-LENGTH
54.89 + 2.40
49.5 58.7
52.12 + 3.15
43.2 57.2
55.46 + 1.58 (9)
53.3 59.2
L-LENGTH
56.58 + 2.63
50.1 60.0
54.03 + 2.51
49.4 59.1
57.58 + 1.45 (12)
55.7 61.0
W-SHAFT
6.48 + 0.36
5.6 7.1
5.94 + 0.38
5.2 6.5
6.49 + 0.29 (17)
5.9 7.0
D-SHAFT
8.15 + 0.50
6.9 8.6
7.38 + 0.46
6.7 8.4
8.04 + 0.4l (17)
7.3 9.0
W-PROX
16.13 + 0.60
15.5 17.4
14.56 + O.85
13.2 16.2
15.43 + 0.59 (21)
14.5 16.7
D-HEAD
7.02 + 0.33
6.4 7.6
6.55 + 0.33
6.0 7.3
6.76 + 0.28 (16)
6.3 7.2
W-DIST
15.71 + 0.59
14.9 16.6
14.93 + 0.84
13.3 16.5
15.11 + 0.47 (17)
l4.4 16.D
W-M&LCON
12.21 + O.65
11.3 13.2
11.38 + 0.77
10.0 13.0
11.59 + 0.39 (l4)
10.8 12.3
W-LCON
3.34 + 0.27
3.0 3.9
3.02 + 0.33
2.5 3.6
2.99 + 0.20 (13)
2.7 3.3
W-L&FCON
7.25 + 0.25
6.9 7.7
6.74 + 0.4l
5.8 7.4
6.85 + 0.32 (15)
6.4 7.5
D-FCON
8.81 + 0.47
8.3 9.5
8.40 + 0.50
7.5 9.2
8.68 + 0.40 (17)
8.1 9.4
D-LCON
10.36 + 0.37
9.8 11.1
9.78 + 0.57
8.7 10.5
10.14 + 0.37 (l4)
9.6 10.8
D-MCON
9.05 + 0.34
8.4 9.7
8.59 + 0.48
7.6 9-4
8.68 + 0.33 (18)
8.1 9.4


8
4
O
60
40
20
0
2
0
8
4
0
223
NON-MARiNE
MARINE
A
^ m m
L ARIK HEMING BARST


201
seems likely that birds with large limb bones (flamingos, cranes) are
better represented in the collections than really existed either in the
deposit or in the fossil community preserved.
Mixson Bone Bed local fauna. Avian taxa now known from here include
two species of grebe (Rollandia sp. and Podilymbus sp. A); a large
stork, Ciconia sp. B; unstudied specimens of a large crane (?Grus sp. B),
and an anhinga, Anhinga grandis. These taxa suggest a shallow freshwater
environment with emergent vegetation surrounded by marshes or "wet
prairies". The lack of other avian species, and the lack of a greater
number of specimens should not be considered significant. This locality
was excavated primarily for the remains of large mammals and only a few
specimens of small vertebrates exist in collections.
McGehee Farm local fauna. Avian taxa known from the McGehee Farm 1.
f. include 2 species of grebe (Rollandia sp. and Tachybaptus sp.); an
undetermined species of cormorant; the anhinga, Anhinga grandis; the
night-heron, Nycticorax fidens; 2 species of duck, a small species of
teal and the other near Anas acuta in size; an undescribed genus of rail;
a flamingo, Phoenicopterus sp.; a jacana, Jacana farrandi; and three
shorebirds, probably all in the genus Calidris. None are abundant, but
the cormorant and the ducks are the most common. Based on the occurrence
of a similar set of avian taxa, the environments surrounding McGehee Farm
during deposition were probably very similar to those at the Love Bone
Bed.
Withlacoochee River kA local fauna. Only two avian species have
been recovered from this Hemphillian site. They consist of a single
femur representing an indeterminate species of Buteo about the size of


CHAPTER I
INTRODUCTION AND PREVIOUS WORK
Introduction
Florida has one of the richest records of fossil birds in the world.
This study examines the systematics of the non-marine fossil birds which
lived in Florida during the late Miocene and early Pliocene (9.04.5
million years before present) and the paleoecology of the localities
which produced them. Included is material from 10 local faunasthe
Love Bone Bed, McGehee Farm, Mixson Bone Bed, Bone Valley, Withlacoochee
River 4a, Manatee County Dam Site, SR-64, Haile VB, Haile VI, and Haile
XIXA. Only a few of the birds from McGehee Farm and the early
collections of birds from Bone Valley have been studied previously (Table
l.l). In addition, the biochronology and faunal dynamics of the entire
North American Neogene avifauna are investigated. Specifically, the
following questions are addressed:
Systematics
1. What species of fossil birds are present in Florida during the
late Miocene and early Pliocene?
2. What are their systematic and biogeographic relationships to
other fossil and Recent species?
Paleoecology
1. Can fossil birds be used to reconstruct the paleoenvironments of
the fossil localities examined in this study?
1


77
Brodkorb from the (Blancan) Hagerman local fauna, Idaho, was originally-
described on a carpometacarpus. Murray- (1970) also redescribed this
species based on additional material. Phalacrocorax macropus (Cope) from
the Mid-Pleistocene Fossil Lake local fauna, Oregon, was described on a
tarsometatarsus. Howard (19^6) referred many other specimens to this
species.
I do not believe that all these species are valid and are correctly
assigned to the ancestral lineage of P. auritus. The morphological
differences among the fossil species are comparable to those among
subspecies of modern P. auritus. It is very possible that like modern
cormorants that show geographic size variations (Palmer, 1962), the
fossil species are simply conspecific geographical variants. If this can
be demonstrated, then each of these fossil species should be maintained
as subspecies of the senior synonym, Phalacrocorax macropus (Marsh).
Valenticarbo praetermissus Harrison from the late Pliocene to early
Pleistocene of Siwalks, India, is based on a 100-year-old plaster cast of
a proximal end of a tarsometatarsus, lacking the hypotarsus. It is very
doubtful that this genus is valid (Olson, ms.). I am unaware of the
relationship of this supposed species.
Pliocarbo longipes Tugarinov from the early Pliocene of the Ukraine
was described from a worn tarsometatarsus and a referred femur. Olson
(ms) notes that although the size and proportions of the tarsometatarsus
are different from typical cormorants, the illustrations are too poor for
even a positive familial verification.
Phalacrocorax destenfanii Regalia from the Mid-Pliocene (Ruscinian?)
of Orciano, Pisano, Italy, was described from most major skeletal
elements. Phalacrocorax mongoliensis Kurochkin from the upper Pliocene


176
These two genera are thought to be more specalized for swimming than
Phoenicopterus.
Genus Phoenicopterus Linnaeus, 1758
Remarks. The post-cranial skeletal elements of flamingos exhibit an
extremely large amount of sexual and individual variation. This
variability is especially evident in the tibiotarsus and tarsometatarsus,
with the males typically being much larger than females. The C. V's of
measurements from the Bone Valley material do not exceed those of a
Recent species, suggesting a single species is present here. At the Love
Bone Bed specimens range in size from smaller than that of the modern P.
minor to larger than that of the modern P. ruber, indicating that more
than one species is present. In trying to separate the Love Bone Bed
specimens into species, I found I could only separate the two extremes of
this continium. I have noted these extremes in the referred material as
"large" or "small".
Material. Love Bone Bed local fauna; UF 29678, humeral end left
coracoid; UF 26088, proximal end left radius; UF 29677, distal end right
ulna; UF 25769, proximal end left femur; UF 25957, distal portion of
shaft of tarsometatarsus; UF 259^3, UF 25930, distal fragments left
tarsometatarsus; UF 2031, UF 26035, UF 260k2, UF 260^3, pedal phalanges.
Remarks. The material listed above is considered to be either too
abraded or too undiagnostic for a more refined taxonomic assignment.
Phoenicopterus floridanus Brodkorb, 1953
Material. Bone Valley Mining District, Palmetto Mine (= Locality 2
of Brodkorb, 1955), referred by Brodkorb; PB lVf, distal end right
tibiotarsus (Holotype, not seen), PB 202, shaft right tibiotarsus (=


72
Description Scapula within range of variation of that of
Phalacrocorax auritus.
Coracoids appear to be well within the range of variation of those of
P. auritus. The characters used by Brodkorb (1955a: 12) "anterior
intermuscular line situated farther laterad" applies only at the extreme
sternal end of the coracoid. This line does not swing mediad; instead it
curves little as it extends down the shaft. Differs from UF 11569 from
McGehee by characters cited above. Coracoids from SR-64 are
indistinguishable from those of Phalacrocorax wetmorei from Bone Valley.
The two characters of the humerus used by Brodkorb (l955a:12) "the
head of humerus shallower" and "condyles averaging less deep" do not hold
when a large series of specimens are measured (Table 4.7). Brodkorb's
statements that the pneumatic fossa is narrow (slightly) and deeper are
supported. This is especially apparent by having a small, but deep fossa
paralleling the crus ventrale fossae. In P. wetmorei the pneumatic fossa
is perforated by several pneumatic foramina but it is rarely perforated
in P. auritus. In the few specimens of P. auritus in which these
foramina are present, they are very minute. Ligamental furrow (=
ligamental sulcus) does not appear to be relatively longer when compared
against a series of both sexes and subspecies of P. auritus. The distal
end of the humerus of P. wetmorei tends to be narrower, with a more
elongated attachment for the anterior articular ligament (= tuberculum
supracondylare ventrale) ending proximally in a narrower crest than in
the modern specimens of P. auritus. Measurements of ulnae are given in
Table 4.7.
Carpometacarpi of P. wetmorei are about as robust as those of
females of P. f. floridanus. The process of metacarpal I is slightly


Table 4.21. Comparative measurements of the femora and tarsometatarsus of the turkeys Proagriocharis
kimballensis, Meleagris progenes and Meleagris gallopavo silvestris, and Meleagridinae, genus indet. from
the Love Bone local fauna. All measurements are means (Steadman, 1980) except those of the material from
the Love Bone Bed local fauna. Ratios were calculated by dividing the measurement in the fossil specimens
by that of M. silvestris.
Love
Bone Bed
P. kimballensis
M. progenes
M. g.
silvestri:
Measurement
Sex
Datum
Ratio
Datum
Ratio
Datum
Ratio
Datum
Ratio
Femur
W-PROX
M
20.9
0.603




34.65
1.00
F
20.9
0.795




26.28
1.00
D-HEAD
M
8.2
0.641
.
12.79
1.00
F
8.2
0.830




9.81
1.00
Tarsometatarsus
W-PROX
M


i4.0
0.572
18.7
0.764
24.49
1.00
F


12.95
O.665


19.46
1.00
W-SHAFT
M
7.45
0.798
9.34
1.00
F


5.05
0.690


7.32
1.00
D-SHAFT
M
...
__
5.05
0.836
6.04
1.00


3.5
0.709
3.9
O.789
4.94
1.00
1
157


108
Blancan of Texas, I cannot demonstrate any qualitative differences and
have therefore referred this distal end of a tibiotarsus to P.
pharangites strictly on the basis of size.
Threskiornithinae, gen. et sp. indet.
Material. Love Bone Bed local fauna; UF 26003, humeral end right
coracoid.
Description. Humeral end coracoid with ventral portion of head
abraded and with procoracoid broken and missing. Size of a large
Plegadis falcinellus, but also within the range of Eudocimus albus or E.
ruber. The impression for the acrocoracohumeralis ligament is wider in
the fossil specimen than in either Plegadis or Eudocimus. In anterior
view, the shaft appears wider, resembling Eudocimus rather than Plegadis.
Remarks. I can find no consistent characters on the humeral end of
the coracoid which will discriminate with confidence between species of
Plegadis and Eudocimus. This specimen is probably too large to represent
the other ibis (Plegadis cf. P. pharangites) from the Love Bone Bed.
Remarks on the Family Plateleidae.
Olson (l98lb) has recently discussed the fossil record of ibises.
There has been no additional species described since the appearance of
his paper.
I have not been able to find consistent osteological characters
which can separate all specimens of Eudocimus ruber from E. albus.
Considering that most skeletal measurements of these two species overlap
extensively (compare in Table 4.15), and that these "species" interbred
freely when E. ruber was introduced into south Florida, these two taxa
should probably be regarded as two color morphs of the same species. As


195
Order Passeriformes (Linnaeus, 1758)
Ordinal Characters. Coracoids separate, lacking coracoidal
fenestra; process small; brachial tuberosity elongate and expanded toward
sternal end of element; head of coracoid usually small, somewhat pointed.
Family Fringillidae Vigors, 1825
Genus Palaeostruthus Wetmore, 1925
Palaeostruthus eurius Brodkorb, 1963
Material. Haile VI local fauna; PB 8502, distal portion of left
tarsometatarsus (holotype).
Remarks. See Brodkorb (1963a) and Steadman (1981) for discussion
and remarks pertaining to this species.
Family Indeterminable
Material. Love Bone Bed local fauna: UF 25806, left coracoid; UF
25T72, distal left femur; UF 29727, 29728, humeral ends right coracoids;
UF 29729, 29730, humeral ends left coracoids.
Remarks. The above specimens represent a minimum of two species
based on size. I have been unable to find characters on the humeral ends
of the coracoid to identify these specimens beyond the level of order.
Characters given by Hamon (1964) do not characterize the suborders of
passerines, although these characters appear adequate to discriminate
between the late Pleistocene and Recent taxa of North America which he
examined. The difficulties in identifying passerines are due to the
large number of genera and the general similarity of morphology within
the order (Frbringer, 1888). This is additionally complicated by the
poorly understood interrelationships between genera and families (Olson,
ms.).


Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy
THE FOSSIL BIRDS FROM THE LATE MIOCENE AND EARLY PLIOCENE
OF FLORIDA
By
Jonathan J. Becker
August 1985
Chairman: Pierce Brodkorb
Maj or Department: Zoology
This study examined the non-marine avifauna from ten late Miocene
and early Pliocene localities in Florida. These localities include the
Love Bone Bed, McGehee Farm, Mixson's Bone Bed, Bone Valley Mining
District, Withlacoochee River UA, Manatee County Dam, SR-6U, Haile VB,
Haile VI, and Haile XIXA. Non-marine genera (number species, if more
than one) present include Rollandia, Tachybaptus, Podilymbus (2),
Podiceps, Pliodytes, Phalacrocorax (3), Anhinga (2), Ardea (2), Egretta
(2 or 3), Ardeola, Nycticorax, Mycteria, Ciconia (3), Eudocimus,
Plegadis, Threskiornithinae, genus indeterminate, Pliogyps, Pandion (2),
Haliaeetus, Buteo, Aguila, Accipitrid, genus indeterminate (3),
Dendrocygna, Branta, Anserinae, genus indeterminate (U), Tadorine, genus
indeterminate, Anas (2), Anatine, genus indeterminate (2), Aythya,
Oxyura, Meleagridinae, genus indeterminate, Meleagris, Grus (2),
Balearicinae, genus indeterminate, Aramornis, Rallus (3), Rallid,
undescribed genus, Phoenicopterus (2), Jacana, Limosa, "Calidris" (6+),
viii


155
Order Galliformes (Temminck, 1820)
Family Phasianidae Vigors, 182$
Subfamily Meleagridinae (Gray, 1840)
Genus indet.
Material. Love Bone Bed local fauna; UF 25768, proximal half left
femur.
Remarks. Steadman (1980) reviewed and evaluated the previously used
taxonomic characters of the meleagridine femur. The femur of
Proagriocharis kimballensis is not known and that of Meleagris progenes
is damaged so that the few qualitative characters of value could not be
judged for these species. As there is a general increase in size through
time in this subfamily (Steadman, 1980:153), I attempted to assign this
specimen to a genus by comparing measurements of Recent and fossil
species (Table 4.21). These comparisons assume that similar measurements
(e.g. diameters of hind limb elements) do not vary between different
skeletal elements of a given species. Ratios of measurements in
Proagriocharis kimballensis to those of Meleagris gallopavo range from
about 0.57 to 0.71. In Meleagris progenes these ratios range from O.76
to 0.84. Ratios for UF 25768 range from 0.60 to 0.64 if the specimen is
assumed to be male (within the range of Proagriocharis kimballensis) or
from 0.80 to 0.83 if the specimen is assumed to be female (within the
range of Meleagris progenes). This specimen is therefore not assigned to
genus.
Genus Meleagris Linnaeus, 1758
cf. Meleagris sp.


159
transverse sulcus relatively narrow and deeply excavated (less so in
Grus, Balerica, and Bugeranus). Distal end of the tarsometatarsus in
Balerica with trochlea II higher on shaft than in that of other genera.
Subfamily Gruinae (Vigors, 1825)
Genus Grus Pallas, 1766
Grus sp. A.
Material. Love Bone Bed local fauna; UF 25752, proximal end right
carpometacarpus; UF 25903, distal end left tibiotarsus; UF 26092,
proximal end right tarsometatarsus.
Description. Tibiotarsal and tarsometatarsal characters as in the
genus Grus. Carpometacarpal fragment referred on basis of size. Smaller
than all species of living cranes except Grus canadensis. Differs from
Grus canadensis by having a less pronounced lateral expansion of the
lateral condyle, a less pronounced internal ligamental process, a more
robust shaft, a larger tendinal canal, and a broad triangular .crest
extending proximally from the lateral condyle.
Tarsometatarsus slightly larger that that of Grus canadensis;
carpometacarpus slightly smaller.
Remarks. The unexpected difference in size between the above
specimens could possibly be due to the elements representing two sexes,
two subspecies (analogous to the modern situation in Florida resident
population of Grus canadensis pratensis and the migratory Grus canadensis
tabida), or a single species of slightly different proportions. With
only the scanty material above, it is impossible to choose between these
possibilities.
The skeletal elements of the fossil species Grus nannodes from the
Hemphillian Edson local fauna, Kansas, is similar in size to the above


Table 4.29. Measurements of the tibiotarsi of the following tytonid owls: Tyto albus pratncola (N = l6, 8
males, 8 females), Tyto glaucops (N = 6, 2 males, 1 female, 3 unsexed), Tyto capensis (N = 1, unsexed),
Phodilus badius (N = 2, 1 male, 1 unsexed), and tytonid, undescribed species from the Love Bone Bed local
fauna. Data are mean _+ standard deviation and range. Abbreviations defined in the methods section.
Measurements
T. a. pratncola
T. glaucops
T. capensis
P. badius
LOV
W-DIST-CR
10.92 + 0.40
10.0 11.7
10.18 + 0.53
9.6 10.7
10.0
9.6; 9.2
11.7
D-ICON
4.83 + 0.26
4.5 5.5
4.47 + 0.48
4.0 5.3
4.5
3.5; 3.8
5.3
i
193


175
Order Charadriiformes (Huxley, 1867)
Remarks. The phylogenetic position of the Phoenicopteridae has long
been debated by avian systematists. Flamingos have been placed with the
storks in the Order Ciconiidae, with the ducks in the Order Anseriformes,
or commonly in their own order, the Phoenicopteriformes. I follow the
recent work by Olson and Feduccia (1980) which establishes the
Phoenicopteridae as a family within the Order Charadriifromes, as shown
by their life history, behavior, nyology, pterylosis, natal down, oology,
parasites, biochemistry, osteology, and paleontology.
Family Phoenicopteridae Bonaparte, 1831
Remarks. Recent flamingos are usually divided into three genera,
based primarily on bill morphology and the presence (or absence) of a
very reduced hind toe. Phoenicopterus is usually considered to have the
most primitive feeding apparatus of the three, with Phoenicoparrus and
Phoeniconaias being more specalized. There are no characters -in the
post-cranial skeleton which will define these genera (Olson and Feduccia,
1980; personal observation). For the basis of comparison with fossil
species, I consider the living species to be congeneric. Olson and
Feduccia (1980) also consider the fossil genera Gervaisia Harrison and
Walker and Harrisonavis Kashin to be junior synonyms of Phoenicopterus
Linnaeus. Additionally, I consider Leakeyornis Rich and Walker to also
be a junior synonym of Phoenicopterus, as even the original authors of
Leakeyornis suggest (Rich and Walker, 1983: 104)! I follow Olson and
Feduccia (1980) and A. H. Miller (1944) in not recognizing the
Palaelodidae as a separate family. Both Palaelodus and Megapalaelodus
may be separated from Phoenicopterus by having a shorter tibiotarsus and
a shorter and more laterally compressed tarsometatarsus (Olson, ms).


Figure 2.U. Schematic diagrams illustrating the measurements of
the tarsometatarsus. A. Dorsal view. B. Lateral view. C.
Plantar view of distal end. D. Distal end view. E. Proximal end
view. Figures are not drawn to scale. Measurements are defined in
text.


CHAPTER III
GEOLOGY
Biochronology
The Clarendonian and Hemphillian land mammal ages were first
proposed by Wood et al. (19^1) based on the stage of evolution of mammals
from two localities in the panhandle of Texas. Since then, the concept
of each has changed, owing to the increasing knowledge of this time
period in North America. Two additional ages were proposed by Schultz et
al. (1970) for parts of the time intervals covered by the original
definitions: the Valentinian (for the late Barstovian to early
Clarendonian) and the Kimballian (originally proposed as late
Hemphillian; now considered to be early Hemphillian). Neither has been
accepted for use on a continent-wide basis. Rather, each has been
applied only to fossils from the type formations of the proposed ages
(Valentine and Kimball formations). Pertinent references include Schultz
et al. (1970), Tedford (1970), Tedford et al. (in press), Breyer (1981),
and Voorhies (I98U).
The Clarendonian, in the restricted sense of Tedford et al. (in
press) is defined on the
earliest appearance of Barbourofelis and, later in the
interval, Platybelodon, Amebelodon, and Ischyrictis
(Hoplictis).Characterization earliest appearance of
Nimravides, Epicyon (in the Great Plains and Gulf Coast),
Griphippus [=Pseudhipparion], Astrohippus, Nannippus (Gulf
Coast), Macrogenis, Synthetoceras, Hemiauchenia, Megatylopus,
Antilocaprinae (Plioceras and Proantilocapra), latest
occurrence of Eucastor, Brachypsalis, Ischyrocyon, Cynarctus,
Aelurodon, Tomarctus, Hypohippus, Megahippus, Merychippus,
Paratoceras, Miolabis, Protolabis, and probably Ustatochoerus.
31


humeral ends left coracoids (tentatively referred); UF 25756, left
carpometacarpus missing shaft of metacarpal III.
McGehee Farm local fauna; UF 8780, complete left coracoid; UF 12469,
right tarsometatarsus.
Description. The above taxonomic assignment is based primarily on
humeri. They are referrable to the Anatinae by lacking a prominent
capital shaft ridge, having the capital groove extending laterally across
the anconal surface and deeply undercutting the humeral head, and by
having a strongly developed attachment for the external head of the
triceps.
Within the subfamily Anatinae, the fossil humeri may be
distinguished from those of the tribe Tadornini by lacking a prominent
capital ridge shaft, by having the head unrotated, deltoid crest not
large and flaring and not extending distally. They may also be
distinguished from those of the tribe Cairinini by not having _a robust
shaft and the pneumatic fossa not restricted to a circular opening rimmed
with heavy bone; distinguished from those of the tribe Oxyurini by having
a very deep pneumatic fossa, extending well under head without numerous
foramina piercing the walls, by the entepicondyle not being reduced, and
by the scar for the attachment of the latissimus dorsi posterioris not
being in a line with the outer edge of the pectoral attachment;
distinguished from those of the tribe Mergini and the tribe Somaterini by
the internal tuberosity not being short and deep, and by the
entepicondylar prominence not being reduced.
The fossil humeri from the Love Bone Bed local fauna agree with the
humeri of both the Anatini and the Aythini by having the capital ridge
shaft obsolete and by having the pneumatic fossa ovaloid and unrimmed


150


13U
the distal portion of the carpal trochlea. Carpometacarpus agrees with
Haliaeetus by having a small anterior carpal facet; tendinal groove more
on the external surface and having a similar excavation on external side
of pollical facet.
Remarks. Material now available is not diagnostic to the generic
level. As additional material becomes available, this carpometacarpus
should be re-examined to determine its generic status. The time interval
between the Love Bone Bed and the Bone Valley local faunas make it
unlikely, although not impossible, that these specimens represent one of
the species of eagle from Bone Valley.
Accipitrid, Genus indet., species B.
Material. Bone Valley Mining District, Chicora Mine; PB 8100,
distal end right carpometacarpus.
Remarks. Carpometacarpus badly fractured and warped. It resembles
Haliaeetus but this material is not complete enough to discriminate
between Haliaeetus and species of Aguila of similar size.
Accipitrid, Genus indet., species C.
Material. Love Bone Bed local fauna; UF 25^91 distal end left
tarsometatarsus, lacking half of trochlea III and all of trochlea IV.
Remarks. Typical accipitrid tarsometatarsus. Size between males
and females of Buteo jamaciensis borealis; but also within the size range
of the Neotropical Spizaetus ornatus and of several African species of
the genera Lophaetus; Hieratus and Old World species of Aguila, including
A. rapax and wahlbergi. The incompleteness of the specimen and the lack
of diagnostic characters on the distal end of the tarsometatarsus
(Jollie, 1976-1977:266) prevents assignment of this specimen to a genus.


other Neogene fossil localities in North America which have not been
included within this study.
Tribe Tadornini Reichenbach, "1850"
Remarks. The humerus of the tribe Tadornini may be distinguished by
the following combination of characters: capital shaft ridge prominent,
directed forward toward the external tuberosity. Characters that are
typical anatine in form are also present, such as having the external
head of the M. humerotriceps deeply undercutting the humeral head and
being continuous with the capital groove.
Genus and species indet.
Material. Bone Valley Mining District, Ft. Green Mine (# 13
dragline); UF 57253, proximal end right humerus.
Remarks. Typical Tadornine humerus about the size of that of a male
Chloephaga picta. This specimen could represent a large species in the
genus Anabernicula, but it is apparently larger than all species now
described in that genus. The specimen is not complete enough to warrant
assignment to genus.
Tribe Anatini Vigors,1825
Genus Anas Linnaeus, 1758
Anas undescribed sp. A.
Material. Love Bone Bed local fauna; UF 25738, complete left
humerus, UF 25720, left humerus missing distal end, UF 29750, proximal
end left humerus; UF 25731, 25736, proximal ends right humeri; UF 25837
UF 25853, UF 29770, UF 29761, complete right coracoids; UF 25805, UF
29768, complete left coracoids; UF 29801, UF 258L0, UF 258U1, humeral
ends right coracoids (tentativly referred); UF 25795, UF 25780, UF 29795,


7
Table 1.1continued
Locality
Reference (s)
Haile XIB, Alachua Co.
Lign, 1965i Olson, 19T^b
Hog Cave, Sarasota Co.
Steadman, 1980; Wetmore, 1931
Hog Creek, Manatee Co.
Wetmore, 1931
Hornsby Springs,
Alachua Co.
Storer, 1976b
Itchtucknee River,
Columbia Co.
Campbell, 1980; McCoy, 1963; Olson, 197^+a,
1971+b, 1977b; Storer, 1976b; Wetmore, 1931
Jenny Spring, Gilchrist Co.
Storer, 1976b
Kendrick IA, Marion Co.
Steadman, I98O
Lake Monroe, Volusia Co.
Holman, 1961; Storer, 1976b
Mefford Cave I, Marion Co.
Steadman, 1980
Melbourne, Brevard Co.
Holman, 1961; Steadman, 1980; Wetmore, 1931
Monkey Jungle, Dade Co.
Ober, 1978
Oakhurst Quarry, Marion Co.
Holman, 1961; Steadman, 1980
Orange Lake, Marion Co.
Holman, 1961
Reddick IB, Marion Co.
Brodkorb, 1952, 1957, 1963e; Hamon, I96U;
Holman, 1961; Olson, 197^b, 1977b;
Steadman, 1976, 1980; Storer, 1976b
Rock Springs, Orange Co.
Storer, 1976b; Steadman, 1980; Woolfenden,
1959
Sabertooth Cave, Citrus Co.
Holman, 196l; Wetmore, 1931
St. John's Lock, Putnam Co.
Storer, 1976b
St. Mark's River,
Leon/Wakulla Co.
Steadman, 1980
Santa Fe River IA,
Gilchrist Co.
Steadman, 1980
Santa Fe River IVA,
Gilchrist Co.
Steadman, I98O
Seminole Field,
Pinellas Co.
Holman, I96I; Olson, 1974b; Steadman, 1980;
Wetmore, 1931


Figure 4.8. Anas sp. A, B. Left coracoid, UF 25805. A. Dorsal
view. B. Ventral view. C, D. Left carpometacarpus, UF 25756.
C. Ventral view. D. Dorsal view. E, F. Left humerus, UF 25738.
E. Cranial view. F. Caudal view. Scale (top) A-D = 2 cm.;
(bottom) E, F = 3 cm.


16
7. D-PROX.Depth of proximal end from most caudal edge of medial
articular face (Facies articularis medialis) to the most cranial
point of the cranial cnemial crest.
8. W-PROX-L.Transverse width of proximal end from medial border
of cranial cnemial crest to lateral border of lateral cnemial
crest (Crista cnemialis lateralis).
9. W-DIST-CR.Transverse width of distal end, measured across
cranial portion of condyles.
10. W-DIST-CD.Transverse width of distal end, measured across
caudal portion of condyles.
11. D-MCON.Greatest depth of medial condyle.
12. D-LCON.Greatest depth of lateral condyle.
13. D-ICON.Depth of area intercondylaris.
Tarsometatarsus
1. LENGTH.Greatest length from intercondylar eminence (Eminentia
intercondylaris) through trochlea for digit III (Trochlea
metatarsi III).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. FLEXOR.Intercondylar eminence to middle of tubercle for
tibialis anterior (Tuberositas m. tibialis cranialis).
5 W-PROX.Greatest transverse width proximal articular surface,
measured across dorsal surface.
6. D-MCOT.Greatest depth medial cotyla.
7. D-LCOT.Greatest depth lateral cotyla.


56
a few specimens, none of which are directly comparable. Additional
material will eventually determine the validity of these assignments.
The occurrence of Tachybaptus in Florida is not surprising
considering the present range of Tachybaptus dominicus throughout the
Caribbean. Storer (1976a:124) suggests that T. dominicus has long been
separated from its Old World relatives (T. ruficollis subgroup).
Supporting this view is the lack of an extra canal in the hypotarsus of
T. dominicus (present in T. ruficollis subgroup). Storer (1976a)
considers this a derived character of T. dominicus.
The absence of small grebes the size of Tachybaptus from the Bone
Valley is probably due to a sampling bias toward large specimens or
possibly a general rarity of grebes due to the ecology of the area during
the deposition of the Bone Valley Formation.
The presence of Rollandia in the late Miocene of Florida suggests
that this genus, like Tachybaptus, had a far greater range in the past.
None of the fossil material of this family now known gives any
indication of the higher level systematic relationships of this group.
Postulated relationships include the Gaviidae, Hesperornithiformes,
Sphenisciformes (Cracraft, 1982), based on primarily foot-propelled
swimming adaptations; and the Rhinochetidae and Eurypygidae, on an
apparently unique configuration of the M. longus colli (Zusi and Storer,
1969).
Considering the distribution and diversity of the living species of
grebes (cf. Storer, 1963) it is probable that this group originated in
either North or South America. Supporting this view is the presence of
two genera unique to the Americas (Aechomorphus and Podilymbus) and the
diversity of the fossil record (ten out of eleven fossil species occur in


6
Table 1.1continued.
Locality Reference (s)
EARLY PLEISTOCENE (irvingtonian)
Coleman IIA, Sumter Co.
Ritchie 1980; Steadman, 1980
Haile XVIA, Alachua Co.
Steadman, 1980
Inglis IA, Citrus Co.
Carr, 1981; Ritchie, 1980; Steadman, 1980
Santa Fe River IIA,
Gilchrist Co.
Steadman, 1980
Williston, Levy Co.
Holman, 1959, 1961; Steadman, 1980
LATE PLEISTOCENE (Rancholabrean)
Arredondo, Alachua Co.
Brodkorb, 1959; Holman, 196l; Olson, 1974b,
1977b; Steadman, 1976, 1980; Storer, 1976b
Aucilla River IA,
Jefferson Co.
Steadman, 1980
Bowman IA, Putnam Co.
Steadman, 1980
Bradenton, Manatee Co.
Becker, 1984; Steadman, 1980; Wetmore,
1931
Catalina Lake,
Pinellas Co.
Storer, 1976b
Coleman III, Sumter Co.
Ritchie, 1980
Crystal Spring Run,
Pasco Co.
Brodkorb, 1956b
Davis Quarry, Citrus Co.
Steadman, 1980
Econfina River, Taylor Co.
Steadman, 1980
Eichelberger Cave,
Marion Co.
Brodkorb, 1955b; Holman, 1961
Florida Lime Company,
Marion Co.
Steadman, 1980
Haile IA, Alachua Co.
Brodkorb, 1953a, 1954b; Olson, 1974b, 1977b
Haile IIA, Alachua Co.
Holman, 1961; Steadman, 1980
Haile VIIA, Alachua Co.
Steadman, 1980


131
Family Pandionidae (Sclater and Salvin, 1893)
Remarks. The following account briefly establishes the presence and
distribution of the late Miocene and early Pliocene ospreys in Florida
for the paleoecological and biochronological aspects of this study.
Detailed descriptions and systematic remarks may be found in Becker
(1985b).
Genus Pandion Savigny, 1809
Pandion lovensis Becker, 1985
Material. Love Bone Bed local fauna; UF 25950, nearly complete left
tarsometatarsus (holotype); UF 25766, distal half right femur; UF 2588U,
distal end of right tibiotarsus; UF 25928, complete left tibiotarsus; UF
25863, right tarsometatarsus lacking proximal end; UF 26055, UF 26056, UF
29660, ungual phalanges (paratypes).
Remarks. This species is more generalized, with longer and more
slender pelvic limb elements, than the living Pandion haliaetus. See
additional comments in Becker (1985b).
Pandion sp.
Material. Bone Valley Mining District, Palmetto Mine; UF 123^6,
ungual phalanx (claw).
Remarks. This specimen is not identifiable to species, but is
sufficently distinct to document a species of Pandion being present in
the Bone Valley Avifauna.
Remarks on the Family Pandionidae.
Warter (1976) has reviewed the fossil history of this family and
described the first fossil species of Pandion from the mid-Barstovian
Sharkstooth Hill local fauna, California. Recently, another fossil


231
Brodkorb, P. 1953e. A Pliocene grebe from Florida. Annals and Magazine
of Natural History, Series 12:953-954.
Brodkorb, P. 1954a. A chachalaca from the Miocene of Florida. Wilson
Bulletin, 66:l80-l83.
Brodkorb, P. 1954b. Another new rail from the Pleistocene of Florida.
Condor, 56:103-104.
Brodkorb, P. 1955a. The avifauna of the Bone Valley Formation. Florida
Geological Survey, Report of Investigations, 14:1-57.
Brodkorb, P. 1955b. Pleistocene birds from Eichelberger Cave, Florida.
Auk, 1:136-137.
Brodkorb, P. 1956a. Two new birds from the Miocene of Florida. Condor,
58:367-370.
Brodkorb, P. 1956b. Pleistocene birds from Crystal Springs, Florida.
Wilson Bulletin, 68:158.
Brodkorb, P. 1957- New passerine birds from the Pleistocene of Reddick,
Florida. Journal of Paleontology, 31:129-138.
Brodkorb, P. 1959* The Pleistocene avifauna of Arredondo, Florida.
Bulletin of the Florida State Museum, Biological Sciences, 4:269-
291.
Brodkorb, P. i960. Great auk and common murre from a Florida midden.
Auk, 77:342-343.
Brodkorb, P. 1963a. Fossil birds from the Alachua clays of Florida.
Florida Geological Survey, Special Publication No. 2, Paper No. 4:1-
17.
Brodkorb, P. 1963b. Miocene birds from the Hawthorne formation.
Quarterly Journal Florida Academy Science, 26:159-167.
Brodkorb, P. 1963c. Catalogue of Fossil Birds. Part 1.
(Archaeopterygiformes through Ardeiformes). Bulletin Florida State
Museum, Biological Science, 7:180-293.
Brodkorb, P. 1963d. A giant flightless birds from the Pleistocene of
Florida. Auk, 80:111-115.
Brodkorb, P. 1963e. An extinct grebe from the Pleistocene of Florida.
Quarterly Journal of the Florida Academy of Sciences, 26:53-55*
Brodkorb, P. 1964. Catalogue of Fossil Birds. Part 2. (Anseriformes
through Galliformes). Bulletin Florida State Museum, Biological
Sciences, 8:195-335*


55
California, and the Curtis Ranch, San Pedro Valley, Arizona, to this
species. Aechmophorus elasson, Murray, from the Blancan Hagerman local
fauna, was described on the distal end of an humerus and an associated
left ulna. It is similar to the living A. occidentalis. Podilymbus
ma,i us cuius Murray, also from the Hagerman local fauna, is based on a
nearly complete tarsometatarsus. It is larger than Podilymbus podiceps.
He also tentatively refers material from the Rexroad and Saw Rock Canyon
local faunas to this species.
Pleistocene species include Podiceps parvus (Shufeldt), based on a
lectotype right tarsometatarsus selected by Wetmore (1937) from the
Fossil Lake local fauna, Oregon. It is similar to the living P.
grisegena but is appreciably smaller (Howard, 1946). It is also know
from a well-core in the Tulane Formation of Kern County, California
(Wetmore, 1937). Podiceps dixi Brodkorb is known only from the proximal
end of a right carpometacarpus from Reddick, Florida. It was named after
the Dixie Lime Products Company which owned the quarry in which it was
found (Brodkorb, pers. coram., 1984; etymology omitted in Brodkorb,
1963e). It resembles the living Podiceps auritus, but is somewhat
larger (Brodkorb, 1963e). Podilymbus wetmorei Storer is based on a type
left tarsometatarsus, also from Reddick, Florida, and from a referred
tarsometatarsus and two femora from the Itchtucknee River, Florida. This
species is diagnosed as being the size of Podilymbus podiceps but more
robust. It is only known from these four elements.
The distribution of fossil grebes from the late Miocene through the
early Pliocene of Florida is shown in Table 4.1. Either Podilymbus cf.
P. podiceps or Podiceps sp. from Bone Valley could possibly be
conspecific with Pliodytes lanquisti. These species are only known from


Table 6.3 Faunal dynamics of the marine Neogene birds from North America. Abbreviations as in Table 6.1
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Duration (MA)
3
3.5
5
2.5
4.5
2.7
Localities (Published)
3 (3)
1 (1)
3 (3)
8 (8)
11 do)
1(1)
Sampling Index
1.00
0.29
0.60
3.20
2.22
0.37
Number of genera (Si)
3
3
12
21
26
26
Originations (No.)
3
2
9
11
10
2
Extinctions (No.)
2
0
2
5
2
2
Running mean (Rm)
0.50
2.00
6.50
13.00
20.00
24.00
Origination Rate
1.00
0.86
1.80
4.40
2.22
0.74
Extinction Rate
0.67
0.00
0.40
2.00
0.44
0.74
Turnover Rate (T)
0.84
0.43
1.10
3.20
1.33
0.74
T/Rm
1.68
0.21
0.17
0.25
0.07
0.03
T/Si
0.28
0.l4
0.09
0.15
0.05
0.03
i
217


21k
HEMPHILLIAN. First Appearance: Podiceps, Podilymbus, Pelecanus,
Eudocimus, Nycticorax, Cygnini, Bucephala, Oxyura, Haematopus, Calidris,
Larus, Pinguinus, Manca 11a. Last Appearance: Pelagornithidae,
Balearicinae, Premancalla, Megapaloelodus.
BLANCAN. First Appearance: Aechmophorus, Botaurus, Colinus, Meleagris,
Ptychoramphus, Titanis, Sterna. Last Appearance: Miosula, Pliogyps,
Mancalla.


178
may be distinguished from that of P. floridanus by (l) larger size, (2)
proportionally wider distal end, (3) more robust shaft, (4) papilla just
medial to the proximal opening of the tendinal canal more expanded.
Comparisons of the distal end of a right tibiotarsus (USNM 242202)
from the Lee Creek local fauna with the large subset of tarsometatarsi
from the Love Bone Bed show it to be similar in all characters except for
the notch on the distal surface of the external condyle being slightly
more pronounced in distal end view. Distal opening of the tendinal canal
more transversely elongated and nutrient canal lateral to papilla is
larger and more pronounced in the Love Bone Bed specimens than in the Lee
Creek specimen in cranial view. Groove usually deeper and more
pronounced in the Love Bone Bed specimens than in the one from Lee Creek
in lateral view. In caudal view, the most cranial portion of the lateral
edge of the articular surface is expanded cranially and laterally in Lee
Creek specimens and not in specimens from the Love Bone Bed.
Distal ends of tarsometatarsi from the Love Bone Bed and McGehee may
be distinguished from the distal ends of tarsometatarsi of P. floridanus
by (l) averging larger and more robust (P. floridanus smaller and more
gracile) (2) caudal portion of the articular surface of trochlea III is
raised above shaft (P. floridanus blends relatively smoothly with shaft)
(3) Distal foramen elliptical and larger (P. floridanus smller, more
nearly circular) (4) Prominent nutrient foramen pierces the caudal
surface of the shaft cranial to the distal foramen (P. floridanus
nutrient foramina very small or absent).
Remarks. The flamingos from the Love Bone Bed show a distinct
bimodal distribution in the size classes of the tibiotarsi, with the
smaller group of tibiotarsi being very similar, if not identical with,


Figure 4.9 A C. Rallid, undescribed genus. A, B. Right
coracoid, UF 29717. A. Ventral view. B. Dorsal view. C.
Proximal end right humerus, UF 9494, caudal view. D. Philomachus
sp., proximal end left humerus, UF 60062, caudal view. E.
Tytonid, undescribed genus, distal end right tibiotarsus, UF 25926,
cranial view, stereopair. Scale A D (top) = 1.5 cm; E (bottom) =
1.5 cm.


34
Mixson Bone Bed
The Mixson Bone Bed is located approximately 2 miles northeast of
Williston, Levy County, in the NE 1/4, SW 1/4, Sec. 29, T. 12 S., R. 19
E., Williston Quadrangle, U. S. Geological Survey 7*5 minute series
topographical map, 1969 This is the type locality of the Alachua
Formation (Dali and Harris, 1892). There have been many studies on this
site, including papers by Leidy and Lucas (1896), Sellards (1916), Hay
(1923), Simpson (1930), Webb (1964, 1969, in press), and Harrison and
Manning (1983). Additional references are listed in Ray (1957).
Genera in common with McGehee include Calippus, Pediomeryx
(Yumaceras), Aepycamelus, Osteoborus, and Aelurodon (Webb, 1969) The
early Hemphillian rqylodontid sloth Thinobadistes is present. Two other
early Hemphillian index genera are absent from this local fauna, although
they are present in other Florida sitesPliometanastes, and Indarctos.
The first appearance of Indarctos occurs late in the early Hemphillian
and its absence in this local fauna probably has temporal significance.
The absence of Pliometanastes is usually considered an ecological
sampling bias.
McGehee Farm
This locality is almost exactly three miles north of Newberry,
Alachua County, along State Highway 45, Sl/2, NW1/4, Sec. 22, T. 9 S., R.
17 E., Newberry Quadrangle, U. S. Geologic Survey 7.5 series
topographical map, 1968, in northcentral Florida. This locality was
first discovered in 1958 and was extensively collected by the University
of Florida, with support of the Frick. Corporation. This local fauna
originates from the Alachua Formation, and the geology of this locality
is briefly discussed by Webb (1964) and Hirschfeld and Webb (1968). The


228
Biochronology and Faunal Dynamics
The final aspect of this dissertation focused on the biochronology
and faunal dynamics of the Neogene North American fossil record of birds.
From the examination of 133 Neogene localities which have produced fossil
birds, I show that the fossil localities which have produced fossil birds
are not uniformly distributed in time, with nearly 75% of the localities
occurring in the last 4l% of the Neogene. A majority of the living
families of North American birds which have a fossil record appear by the
Barstovian. Generic diversity increases throughout the Neogene. The
marine avifauna is essentially established by the Clarendonian at a level
of 20 25 genera, while the non-marine avifauna increases continually.
Generic origination rates peak for marine birds in the Clarendonian, but
extinction rates remain low throughout the Neogene. Generic origination
rates for non-marine birds show a cyclic nature every alternate Land
Mammal Age.
A preliminary list of biochronologically important genera of birds
is presented. As our knowledge of the North American Neogene avifauna
expands and becomes more complete, this list can be revised and updated.


Table 4.l4. Measureemnts of the tibiotarsi of the fossil storks from Florida; Ciconia sp. A, from the Love
Bone Bed Local fauna, Ciconia sp. B. from the Bone Valley Mining District and Mixson local fauna, Ciconia
sp. C. from theBone Valley Mining District, and Ciconia maltha from various Pleistocene localities in
Florida. Data are mean +_ standard deviation and range. Abbreviations are defined in the methods section.
Compare measurements with those in Table 4.13.
Measurements
Ciconia sp. A
Ciconia sp. B
Ciconia sp. C
Ciconia maltha
Tibiotarsus
W-SHAFT
9.04 + 0.66 (5)
8.1 9*6
9.8; 10.2; 11.8*
12.3
12.60 + 1.09 (6)
11.7 12.7
D-SHAFT
7.80 + 0.53 (5)
7.1 8.5
8.4; 8.3; 9-1
11.3
10.01 + 0.75 (7)
9.0 11.1
W-DIST-CR
14.23 + 0.84 (3)
13.7 15.2
17.5; l6.6; 18.0
19.1
19.47 + 0.90 (6)
18.5 20.6
W-DIST-CD
*11.5
14.2; 13.7; 13.2
15.0
15.98 + 0.68 (5)
15.3 17.0
D-MCON
18.5; 18.9
22.8: 21.7; 22.5
26.4
24.80 + 1.57 (5)
22.9 26.9
D-LCON
l8.8; 18.9
22.6; 20.8; 22.4
25.5
24.62 + 1.67 (5)
23.0 26.7
D-ICON
11.48 + 0.28 (5)
11.2 11.9
13.9; 13.0; 13.8
16.5
15.51 + 0.99 (7)
14.5 17.1
i
105


l6l
.laponensis, and G. antigone. These last three species have very similar
distal tibiotarsi and I have not been able to find consistent qualitative
characters on this element to separate these species from each other and
from the fossil specimens above. The proximal and distal ends of the
tarsometatarsus are also indistinguishable from those of G. americana, G.
Japonensis, and G. antigone. Two specimens (distal tibiotarsi; UF 25908,
UF 25911) are larger than all other fossil specimens and may represent
either sexual variation or a specific difference. The lack of an
adequate series of skeletons of Recent species prevents determination of
this question.
A large species of Grus is known from the Lee Creek local fauna
(Olson, ms). Grus sp. B. probably is closely related, or identical with
this species.
Subfamily Balearicinae (W. L. Sclater, 1924)
Balearicinae, Genus et. species indet. _
Material. Bone Valley Mining District, Nichols Mine; UF 24586,
distal end of right tibiotarsus.
Description. Tibiotarsus slightly smaller than that of a female
Anthropoides virgo, but having characters typical of the subfamily
Balearicinae. Comparisons with the tibiotarsus of an undescribed
balearicinae crane from the Hemphillian Long Island local fauna, Kansas,
(YPM 4662) shows UF 24586 to share a similar shape and position of the
condyles, although UF 24586 is much larger and is more anterior-
posteriorly compressed.
Remarks. Comparisons of this tibiotarsus with unpublished fossil
material in the Frick and USNM collections shows that this specimen is
distinct from that of both Probalearica and Aramornis. Additional


V. PALEOECOLOGY 198
Introduction 198
Local Faunas 199
VI. BIOCHRONOLOGY AND FAUNAL DYNAMICS 205
Introduction 205
Faunal Dynamics 205
Biochronology . 213
VII. SUMMARY 224
Systematics 224
Paleoecology 227
Biochronology 228
LITERATURE CITED 229
BIOGRAPHICAL SKETCH 245
vii


Rancholabrean Little Box Elder local fauna (Eraslie, 1985)* This species
is significantly smaller than the living A. crecca. Anas schneideri is
near the size of A. pullulans described from the mid to late Clarendonian
Black Butte local fauna, Oregon, but differs from this species by having
metacarpal I relatively higher (Emslie, 1985)* Campbell (1979) described
three new species of Anas (A. talarae, A. amotape, and A. sanctahelenae)
from the late Pleistocene of Bolivia and Ecuador. Differences between
these South American species of Anas and other living species of Anas
seem slight. Campbell (1979) also described, from the same localities, a
new genus and species (Nannonetta invisitata) of Tadorine.
Anatids are rare as fossils until the Neogene. Olson (ms) notes
that the major adaptive radiation of anatids took place in the Miocene;
for it is difficult to place fossils before this time in modern tribes
and genera.


186
proximal end right tarsometatarsus; Palmetto Mine; UF 21092, distal end
right tibiotarsus.
Remarks. Assignment of additional specimens to Limosa ossivallis
very tentative as most of the skeletal elements are abraded. These
skeletal elements represent a scolopacid of the correct size for this
species (see Brodkorb, 1955a for measurements). The type material, and
the material referred above, should be compared with Limosa vanrossemi L.
Miller, which was described from the Lompoc local fauna of Mohinian age
(=late Miocene) of California. Miller (1925) states that it is closest
to the living Limosa fedoa and shows but slight divergence.
Genus Erolia Vieillot, l8l6
Erolia penepusilla Brodkorb, 1955
Material. Bone Valley Mining District, near Brewster; PB 6ll,
distal end left humerus (holotype).
Remarks. Brodkorb (1955a) found this species to be larger than Erolia
temninckii, E. ruficollis, E. minutilla, Ereunetes pusillus, and
Ereunetes mauri; smaller than Erolia bairdii and E. fusicollis. E.
penepusilla appears to be closest to E. minutilla.
Genus Ereunetes Illiger, l8ll
Ereunetes rayi Brodkorb, 1963
Material. McGehee Farm local fauna; UF 3978, humeral end left
coracoid (holotype).
Remarks. Brodkorb (1963a) found this species to be larger than that
of Ereunetes pusillius, E. mauri, and Erolia minutilla; but smaller than
Erolia bairdii and E. fuscicollis. Ereunetes rayi falls within the same


192
Remarks on the Family Tytonidae
There have been many fossil species of Tyto described (listed in
Brodkorb, 1971; Olson, in press); based mainly on size differences. The
morphology of the distal end of the tibiotarsus of these fossil species
(T. sanctialbani, T. ostologa, and T. pollens examined) is remarkably
uniform within this genus and is also very similar to that of Pholidus
badius.
There is only one fossil tytonid genus now known. Prosybris
Brodkorb is based on the type species P. antigua (Milne-Edwards, 1863)
from the Aquitanian of St.-Gerand-le-Puy, France. It was described on an
tarsometatarsus and is not directly comparable with the above specimen
from the Love Bone Bed local fauna. The tibiotarsus of Prosybris is
unknown.
A number of undescribed Oligocene owls are known. The above
specimen should be compared with these before allocating the Love Bone
Bed owl to a genus.
J


172
from the late Miocene of Oregon, should he compared further with this
undescribed genus and species.
Remarks on the Family Rallidae.
The fossil record of rails has been recently reviewed (Feduccia,
1968; Cracraft, 1973; Olson, 197^+b, 1977b; Kurochkin, I98O; and references
therein). The generic or higher status of living rails has been reviewed
by Olson (1973b) and accounts have appeared for all living species
(Ripley, 1977).
The obvious lack of rails from many of the fossil localities in this
study is probably size related, whether due to inadequate sampling while
collecting, or bone destruction during deposition (or both).
Several fossil species already described should be re-examined as
additional material becomes available from Florida. The fossil species
Rallus phillipsi Wetmore, 1957, from the Wickieup 1. f. of Arizona was
described as intermediate in size between Rallus limicola and Rallus
longirostris. Olson (1977b) notes that when a larger series of recent
comparative skeletons are examined, Rallus phillipsi falls well within
the lower size range of the living Rallus longirostris. Rallus sp. A and
perhaps Rallus sp. B possibly has affinites with Ib_ phillipsi, but
without more material, this cannot be determined.
Rallus prenticei Wetmore 19^ from the Blancan of Kansas and Idaho,
was described as being somewhat larger and heavier than the living Rallus
limicola. Rallus sp. C may have affinities with this species. Better
material is again needed to identify this species with confidence.


Table 4.1continued.
Family Rallidae
Rallus sp. A (LOV)
Rallus sp. B (BV)
Rallus (cf.) sp. C (LOV)
Undescribed genus (LOV, MCG)
Order Charadriiformes
Family Phoenicopteridae
Phoenicopterus floridanus (BV)
Phoenicopterus sp. A (LOV, MCG)
Family Jacanidae
Jacana farrandi (LOV, MCG)
Family Scolopacidae
Limosa ossivallis (BV)
Erolia penepusilla (BV)
Ereunetes rayi (MCG)
Calidris pacis (BV)
"Calidris" sp. indet. 1 (LOV, MCG, BV)
"Calidris" sp. indet. 2 (LOV)
"Calidris" sp. indet. 3 (MCG)
"Calidris" sp. indet. 4 (LOV)
??Actitis sp. indet. 5 (LOV)
??Arenaria sp. indet. 6 (LOV)
Genus indet. sp. indet. 7 (LOV)
Genus indet. sp. indet. 8 (LOV)
?Philomachus sp. (BV)
Order Strigiformes
Family Tytonidae
Undescribed genus (LOV)
Family Strigidae
Bubo sp. (BV)
Order Passeriformes
Suborder indet. sp. A (LOV)
Suborder indet. sp. B (LOV)
Family Fringillidae
Palaeostruthus eurius (H 6)


Table 4.10continued
Measurements
P. a. auritus
P. a. floridanus
W-DIST-CR
11.64 + 0.36
11.06 + 0.50
11.1 12.3
10.2 11.9
W-DIST-CD
11.16 + 0.40
10.47 + 0.50
10.3 11.6
9.8 11.6
D-MCON
10.98 + 0.42
10.41 + 0.60
10.5 11.6
9.2 12.0
D-LCON
10.15 + 0.28
9.76 + 0.45
9.8 10.T
9.0 10.7
D-ICON
6.31 + 0.44
5.70 + 0.34
5.4 6.7
5.1 6.4
i
P. wetmorei
Phalacrocorax sp.
11.57 + 0.49 (25)
10.6- 12.4
11.1; 10.6
11.04 + 0.60 (21)
9.8 12.0
10.7
11.59 + o.4o (38)
10.5 12.3
10.6
10.14 + 0.35 (31)
9.6 10.8
9.4
6.43 + 0.26 (40)
6.1 7.0
6.1; 6.0
00
Ln


CHAPTER VII
SUMMARY
This study has examined three aspects of avian paleontology
systematics, paleoecology, and biochronology and faunal dynamics.
Systematics
It has first focused on the systematics of the non-marine fossil
birds from the late Miocene and early Pliocene of Florida. 78 taxa have
been identified in this study. Grebes (Family Podicipedidae) are
represented by 6 taxa. A species of Tachybaptus is abundant in the Love
Bone Bed local fauna and is also present from McGehee Farm. Rollandia is
known from a few specimens from the Love Bone Bed, Mixson, and McGehee
Farm. The skeleton of this species is slightly more robust than that of
the living Rollandia rolland chilensis. A small species of Podilymbus is
known from Mixson. The Bone Valley Mining District has produced
specimens of 3 grebesPodilymbus cf. JP. podiceps, Podiceps sp. and
Pliodytes lanquisti. All are rare members of the Bone Valley avifauna.
Pelecaniformes were represented by 2 or possibly 3 species of
cormorants (Family Phalacrocoracidae) and 1 or possibly 2 species of
anhinga (Family Anhingidae). The cormorant Phalacrocorax wetmorei is one
of the best represented Neogene fossil species, with well over 500
specimens known. Almost every skeletal element is known.
Phalacrocorax sp. A. is known from the Love Bone Bed, McGehee and
probably Haile XIXA. The anhinga, Anhinga grandis, is known from the
22 k


92
probable that these fossil herons were members of the same diurnal fish
eating guild as are their modern counterparts. It is not suprising that
three herons occur together. As many as 12 species of herons occur
sympatrically in Florida today. They coexist by partitioning resources
such as size and kind of prey, use of habitat (e.g. water depth), and
foraging behavior (Recher and Recher, 1980).
The two herons from Bone Valley could possibly be viewed as marine
specialistsone medium-sized specalizing on small prey and the other
specializing on larger prey, as the living Egretta rufescens and Ardea
herodias occidentalis do today.
Little can be said of the paleoecology of Egretta sp. from the
Withlacoochee River bA but perhaps it was similar to E. ibis in its
ecology, as it is similar to this species in its morphology.
Family Ciconiidae (Gray, l8U0)
Remarks. There are only a few characters on the skeletal elements
preserved here that can distinguish between the genera Mycteria, Ciconia,
and Jabir (sensu Kahl, 1972). Size is of no generic value as evidenced
by species of Ciconia overlapping with species of all other ciconiid
genera.
Characters on the distal end of the tibiotarsus include the internal
ligamental prominence (= medial epicondyle) well-developed in Jabir
(less so in Mycteria and Ciconia); distal end laterally compressed in
Mycteria (less so in Ciconia, except the atypical C. abdimii and C.
episcopus; somewhat compressed in Jabir); distal opening of the tendinal
canal more medially placed in Jabir than in Ciconia or Mycteria;
tubercle slightly elevated above the surface of a strong ridge connecting


8
Table 1.1continued
Locality
Reference (s)
Steinhatchee River,
Taylor/Dixie Co.
Steadman, 1980
Venice Rocks, Manatee Co.
Wetmore, 1931
Vero (Stratum 2),
Indian River Co.
Holman, 1961; Sellards, 1916; Shufeldt,
1917; Steadman, 1980; Storer, 1976b; Weigel,
1962; Wetmore, 1931
Warren's Cave, Alachua Co.
Holman, 1961
Wekiva Run III, Levy Co.
Steadman, 1980
West Palm Beach,
Palm Beach Co.
Becker, 1985c
Withlacoochee River,
Citrus Co.
Steadman, 1980
Zuber, Marion Co.
Holman, 1961
HOLOCENE
Cotton Midden, Volusia Co.
Hay, 1902; Neill et al., 1956
Castle Windy Midden,
Volusia Co.
Weigel, 1958
Good's Shellpit,
Volusia Co.
Steadman, 1980
Green Mound Midden,
Volusia Co.
Hamon, 1959
Nichol's Hammock, Dade Co.
Hirschfeld, 1968; Steadman, I98O
Silver Glenn Springs,
Lake Co.
Neill et al., 1956; Steadman, 1980
Summer Haven Midden,
St. Johns Co.
Brodkorb, i960
Vero (Stratum 3),
Indian River Co.
see Vero, Stratum 2, above
Wacissa River
Jefferson Co.
Steadman, 1980


-T5S
174


evolved more than once, as evidenced by its being open in some genera of
mergansers (always open in Mergus, almost always open in Lophodytes) and
closed in others (Mergellus) (Woolfenden, 1961).
Before the exact systematic position of this species can be
determined, additional comparisons are needed with small living and
fossil species of Anas. Of special interest is Anas luederitzensis
Lambrecht 1929, from the mid-Tertiary of southwest Africa. This species
is said to be distinguished from Anas querquedula and Anas cyanoptera by
having a pneumatic fossa not markedly perforated as in most anatids
(Howard, 1964).
?Anas, size near A. acuta
Material. Love Bone Bed local fauna; UF 26001, right coracoid; UF
25798, UF 25799, UF25800, UF 25801, left coracoids.
McGehee Farm local fauna; UF 9481, UF 31784; right coracoids; UF
9486, UF 9489, left coracoids.
Remarks. Typical Anas morphology. Skeletal elements slightly
larger than A. acuta; definately smaller than modern specimens of Anas
platyrhynchos.
Anatini, Genus indet., species A.
Material. Love Bone Bed local fuana; UF 29767, complete left
coracoid; UF 25808, UF 25854, humeral ends coracoids, tentatively
referred.
Remarks. Coracoids proportionally long and slender; slightly
smaller than females of A. crecca carolinensis. See remarks below under
"Genus indet., sp. B.".


35
latter publication includes a preliminary list of the fossil vertebrates
from this local fauna. No faunal revision of this locality has been
undertaken, but several papers treating specific groups have appeared
(sloths: Hirschfeld and Webb, 1968; mylagaulids: Webb, 1966; canids:
Webb, 1969; and protoceratids: Patton and Taylor, 1973).
The faunal composition of this local fauna indicates an early
Hemphillian age (Hirschfeld and Webb, 1968; Webb, 1969; Marshall et al.,
1979), based primarily on the presence of Pliometanastes, the early
Hemphillian megalonychid ground sloth. Typical early Hemphillian
mammalian genera present include Calippus, Pedimeryx (Yumaceras),
Aepycamelus, Osteoborus, and Aelurodon (Webb, 1969).
The birds of this locality were first studied by Brodkorb (1963a)
who reported Phalacrocorax wetmorei, and described two new species
Nycticorax fidens and Ereunetus rayi. Later, Olson (1976) described
Jacana farrandi from here.
Withlacoochee River 4a
The Withlacoochee River 4a local fauna lies approximately 8 km.
southeast of Dunnellon (center of Nl/2, NW1/4, Sec. 30, T. 17 S., R. 20
E., Stokes Ferry Quadrangle, U. S. Geologic Survey 75 minute series
topographical map, 195*+, Marion County), in northcentral Florida. The
fossil vertebrates originate from a massive green clay filling of a
sinkhole in the late Eocene Inglis Formation of the Ocala Group (Webb,
1969, 1973, 1976). This deposit of green clay is being eroded by the
Withlachoochee River; the fossils were collected in approximately 20 feet
of water using scuba equipment. Deposition almost certainly occurred in
a pond environment near sea-level as shown by the fine-grained sediments


242
Tedford, R. 1970. Principles and practices of mammalian geochronology in
North America. Proceedings North American Paleological Convention,
Part F:666-703.
Tedford, R. H., T. Galusha, M. F. Skinner, B. E. Taylor, R. W. Fields, J.
R. Macdonald, J. Rensberger, S. D. Webb, and D. P. Whistler, in
press. Faunal succession and biochronology of the Arikareean
through Hemphillian interval (late Oligocene through late Miocene
Epochs), North America. University of California Press, Berkeley.
Terres, J. K. 1980. The Audubon Society Encyclopedia of North American
Birds. Alfred A. Knopf, Inc. New York. 1110 pp.
Tonni, E. P. 1980. The present state of knowledge of the Cenozoic birds
of Argentina. Contributions in Science, Natural History Museum of
Los Angeles County, 330:105-114.
Tordoff, H. B. 1959* A condor from the Upper Pliocene of Kansas.
Condor, 61:338-343.
Van den Berge, J. C. 1975* Aves myology, pp. l802-l848. I_n R. Getty
(ed.) Sisson's and Grossman's The Anatomy of the Domestic Animals.
W. B. Saunders, Philadelphia.
van den Driesch, A. 1976. A guide to the measurements of animal bones
from archaeological sites. Peabody Museum of Archeology and
Ethnology. Harvard University Bulletin l:ix + 137 pp. + 62 figs.
Voorhies, M. 1984. 'Citellus kimballensis' Kent and 'Propliophenacomys
uptegrovensis' Martin, supposed Miocene rodents are Recent
intrusives. Journal of Paleontology, 53:254-258.
Vuilleumier, F. 1984. Faunal turnover and development of fossil
avifaunas in South America. Evolution, 38:1384-1396.
Vuilleumier, F. in press. What birds can tell us about continental
connections. I_n F. G. Stehli and S. D. Webb (eds.). The Great
American Interchange. Plenum Press, Geobiology Series, New York.
Warter, S. L. 1976. A new osprey from the Miocene of California
(Falconiformes: Pandionidae). Smithsonian Contributions to
Paleobiology, 27:133-139.
Webb, S. D. 1964. The Alachua Formation. 1964 Field trip. Society of
Vertebrate Paleontologists Guidebook. Gainesville, Florida,
pp. 22-29.
Webb, S. D. 1966. A relict species of the burrowing rodent, Mylagaulus,
from the Pliocene of Florida. Journal of Mammalogy, 47:401-412.
Webb, S. D. 1969. The Pliocene Canidae of Florida. Bulletin of the
Florida State Museum, Biological Sciences, 14:273-308.


CHAPTER IV
SYSTEMATIC PALEONTOLOGY
Table 4.1 lists the non-marine avian taxa now known to occur in
Florida during the late Miocene and early Pliocene. In the following
systematic section, I have tried to present osteological characters which
define the taxonomic groups in a hierarchical fashion. However,
considering the relatively small number of fossil and recent specimens
available for some species, and the restricted geographical area from
which many of the Recent species were collected, I would not be surprised
if some "diagnostic" characters do not hold when a larger number of
specimens are examined.
Order Podicipediformes (Ftirbringer, 1888)
Family Podicipedidae (Bonaparte, 1831)
Tribe Podilymbini Storer, 1963
Characters. Separate canal through hypotarsus for the tendon of
insertion of M. flexor perforatus digiti II (Storer, 1963). Murray
(1967) gives additional osteological characters for the separation of the
different taxa of this family.
Genus Rollandia Bonaparte, 1856
Rollandia sp.
Material. Love Bone Bed local fauna, questionably referred; UF
29670, UF 29673, right coracoids; UF 25815, left coracoid.
Mixson Bone Bed local fauna; F:AM FLA-120-2183, complete right
tarsometatarsus.
49


212
relative time (NALMAs) to one keyed to absolute time. This database
could then be clustered into "natural" groups of geological ranges of
species to discriminate between the lack of localities and real turnover.
Vuilleumier (198*0 examined the faunal turnover of the South
American avifauna and has recently (in press) expanded his investigations
to include the late Neogene North American fossil record of birds. It is
difficult to make comparisons between his study and the results presented
above because of the time scales used. The faunal parameters which
Vuilleumier (1984, in press) presents, are based on a division of
geologic time into Epochs, which have vastly uneven durations (Miocene,
IT MA; Pliocene, 2.5 MA). Many of the finer scale events presented above
would not be apparent in Vuilleumier's parameters.
The faunal dynamics of mammals have been extensively investigated
(Webb, 1976, Marshall, et al., 1982). Table 6.5 compares the most recent
information on faunal dynamics of mammals (Marshall, et al., 1982) with
those for birds. Generic extinction rates, generic origination rates,
and turnover rates are substantially higher for mammals than for birds.
One possible explanation is that mammals are evolving at a much
faster rate than are birds. However, considering the great diversity of
the Recent avifauna and the comparable amount of morphological difference
between equivalent taxonomic ranks of birds and mammals (Wyles et al.,
1983), this explanation seems at best, only partial.
A more likely explanation is that the differences in faunal dynamic
parameters between birds and mammals are due to the use of different
types of taxonomic characters. In systematics of fossil avian species,
the post-cranial skeleton is used almost exclusively. Conversely, in
fossil mammals, dental morphology is usually used. As the post-cranial


59
Table 4.1continued.
Family Pandionidae
Pandion lovensis (LOV)
Pandion sp. (BV)
Family Accipitridae
Haliaeetus (?) sp. (BV)
Buteo near B. jamaciensis (WITH 4a)
Aguila sp. TbV)
Accipitrid, genus indet. sp. A (LOV)
Accipitrid, genus indet. sp. B (BV)
Accipitrid, genus indet. sp. C (LOV)
Accipitrid, genus indet. (WITH 4a, BV)
Order Anseriformes
Family Anatidae
Dendrocygna sp. (LOV)
Branta sp. A (LOV)
Anserinae, genus indet. sp. B (LOV)
Anserinae, genus indet. sp. C (LOV, BV)
Anserinae, genus indet. sp. D (BV)
Tadorini, genus indet. sp. A (BV)
Anas undescribed sp. A (LOV, MCG)
Anas size near A. acuta (LOV, MCG)
Anatini, genus indet. sp. A (LOV)
Anatini, genus indet. sp. B (LOV)
Aythya sp. A (BV)
Qxyura cf. 0_. dominica (BV)
Order Galliformes
Family Phasianidae
Meleagridinae, genus indet. (LOV)
Meleagris sp. (BV)
Order Gruiformes (auct.)
Family Gruidae
Grus sp. A (LOV)
Grus sp. B (LOV)
Balearicinae, genus indet. (BV)
Aramornis (cf.) (LOV)


120
family. They include: Eocathartes robustus Lambrecht, (too crushed);
Teracus littoralis Aymard, (incertae sedis; Olson 1978); and Phasmagyps
patritus Wetmore.
This' leaves Diatropornis ellioti (Milne-Edwards) and Pleiocathartes
europaeus Gail lard from the late Eocene and early Oligocene of the
phosphorites du Quercy as the oldest certain records of vultures in
Europe. Also known is Plesiocathartes? gaillardi from the early Miocene
of Spain and an unreported specimen of a large member of this family from
the early Oligocene of Mongolia (Kurochkin, in litt., to Olson, ms.).
In South America, the oldest current record of this family is
Dryornis pampeanus Moreno and Mercerat, from the Monte Hermoso Formation
in Argentina. This species was originally described as a species of
phorusrachid, but was later moved to the family Vulturidae (Brodkorb,
1967). Tonni (1980) states that it is close to the living Vultur. I
also note that Campbell (1979) synonymzed the fossil Vultur patruus
L&inberg, from the Pliocene of Tarija, Bolivia, with the living Vultur
gryphus.
In North America, the oldest records of vultures are Sarcoramphus
kernensis from the late Miocene (mid-HemphiIlian) of Kern River,
California, the species of Pliogyps from the Love Bone Bed local fauna,
discussed above, and an unreported species of vulturid from the mid-
Barstovian Sharkstooth Hill local fauna (Howard, in litt., 1984).
Sarcoramphus kernensis was originally described, and is still only known,
from a crushed distal end of a humerus. It was compared only with S.
papa (then Vultur papa). It should be re-examined and differentially
diagnosed to determine its correct generic position. There are several
described species of vultures from the Pliocene and Pleistocene of North


20h
the primary formation of phosphate grains from the upwelling of nutrient-
rich waters. I suggest that the fossil birds in the Bone Valley Deposits
are present as a result of the abundant food supplies in these nutrient-
rich waters, rather than being the direct cause of it.
Manatee County Dam Site. The only avian taxon known from this local
fauna is one specimen representing Phalacrocorax cf. P. wetmorei. Based
on the available information pertaining to the geology, vertebrate fauna,
and geographic location, this locality is in all aspects essentially an
outlier of the Bone Valley local fauna discussed above. All comments
pertaining to the paleoecology of Bone Valley also apply to this local
fauna.
SR-6U. Fossil birds from this locality include the cormorant
Phalacrocorax wetmorei, loons, the flamingo, Phoenicopterus cf. P.
floridanus, and alcids. As with the Manatee County Dam Site above, all
available information about the geology, vertebrate fauna and geographic
location indicates this locality is an outlier of the Bone Valley local
fauna. See paleoecological comments under Bone Valley Mining District
above


63
Table 4.4. Measurements of the femora of the grebes Rollandia rolland
chilensis (N = 6, 2 males, 1 female, 3 unsexed), Tachybaptus dominicus
(N = 7, 4 males, 3 females), and Tachybaptus sp. from the Love Bone Bed.
Data are mean _+ standard deviation and range. (*) Specimen
damaged. Abbreviations defined in the methods section.
Measurements
R. r. chilensis
T. dominicus
Tachybaptus sp
Femora
M-LENGTH
30.73 + 1.55
27.9 -32.0
25.03 + 1.53
23.5 27-8
26.1
L-LENGTH
32.73 + 1.64
29-7 33.5
26.81 + 1.6l
25.1 29.7
27.5
W-SHAFT
2.80 + 0.15
2.6- 3.0
2.53 + 0.22
2.3 2.9
2.6
D-SHAFT
3.18 + 0.24
2.8 3.4
2.67 + 0.30
2.3 3.0
2.6
W-PROX
7.65 + 0.39
7.1 8.3
6.53 + 0.30
6.2 6.9
6.8
D-HEAD
3.30 + 0.24
3.0 3.7
2.74 + 0.20
2.4 3.0
2.8
W-DIST
8.08 + 0.44
7.5 8.4
6.84 + 0.53
5.9 7.4
*6.0
W-M&LCON
5.97 + 0.4l
5.4"- 6.4
4.96 + 0.40
4.4 5-5
*5.l~
D-LCON
6.03 + 0.24
5.7 6.4
4.97 + 0.37
4.3 5.5

D-MCON
4.40 + 0.24
4.1 4.7
3.44 + 0.26
3.1 3.8
3.5


197
the only reports of nine-primaried oscines in the Tertiary (Steadman and
McKittrick, 1982).
No fossil passeriformes are recorded before the Miocene (Brodkorb,
1978; Olson and Feduccia, 1979) This could be explained simply as a
sampling bias, but considering the amount of detailed paleontological
field work done both in North America and in Europe, this absence seems
real, at least on these two continents. Prior the Miocene, a great
diversity of small non-passeriforms (mainly Coraciiformes) are known and
they evidently occupied many of the niches which are now filled by
passerines (Mourer-Chauvire^ 1982; Olson and Feduccia, 1979; Feduccia and
Olson, 1982). Feduccia and Olson (1982) noting the great radiation of
suboscines in South America, speculate that they were present in South
America for most of the Tertiary. They also argue that, in addition to
the suboscines, the entire order Passeriformes is of South American
origin.


68
Bone Valley Mining District, Chicora Mine.UF 29733, humeral end
left coracoid.
Bone Valley Mining District, Fort Green Mine.UF 61958, associated
(?) partial skeleton; UF 58062, right quadrate; UF 60047, caudal portion
left mandible; UF 53912, caudal portion right mandible; UF 57248, sternal
fragment with coracoidal sulci; UF 52415, UF 53938, proximal ends right
scapulae; UF 53873, proximal end left scapula; UF 62025, complete left
coracoid; UF 52413, UF 57332, UF 55838, UF 57246, UF 58058, UF 61960, UF
65711, humeral ends right coracoids; UF 53913, UF 55872, UF 57331, UF
58059, UF 58339, UF 61961, UF 65655, humeral ends left coracoids; UF
524l4, UF 61962, UF 65656, sternal ends right coracoids; UF 53934, UF
55831, UF 55875, UF 61963, sternal ends left coracoids; UF 55810, UF
558II, UF 58378, UF 61964, proximal end right humeri; UF 58304, UF 60048,
proximal ends left humeri; UF 53937, shaft left humerus; UF 52410, UF
53914, UF 60050, UF 65657, distal ends right humeri; UF 55865, UF 58060,
UF 58338, UF 58419, UF 58420, UF 60049, distal ends left humeri; UF
55867, UF 57242, UF 58061, UF 60051, UF 65712, proximal ends right ulnae;
UF 55866, UF 61965, proximal ends left ulnae; UF 57243, UF 57247, UF
57249, UF 60052, UF 61966, distal ends right ulnae; UF 52411, UF 55812,
UF 55871, UF 57334, UF 57335, UF 58340, distal ends left ulnae; UF 55813,
UF 55832, UF 55869, UF 60053, UF 65661, proximal ends right
carpometacarpi; UF 52412, UF 55833, UF 55834, UF 57333, UF 584l8, UF
61967, UF 61968, UF 61969, UF 58407, distal ends right carpometacarpi; UF
81959, partial synsacrum; UF 55814, UF 60054, complete right femora; UF
57336, complete left femur; UF 57244, proximal end right femur; UF 53872,
UF 53872, UF 55873, UF 55874, UF 57337, UF 60055, UF 61970, UF 55835,
distal ends right femora; UF 57391, UF 61971, distal ends left femora; UF


equipment. I gratefully acknowledge these departments and institutions
for their support.
Last, I thank my parents, Elwood W. and Nita E. Becker, for their
encourgement and support over the years. They have provided not only the
opportunity, but much of the impetus, that has allowed me to finish my
formal education.
v


32
In Florida, common late Clarendonian taxa include Barbourofelis,
Nimravides, Mylagaulus, Pseudhipparion, Amebelodon, advanced species of
Eucastor, Pediomeryx, and Aelurodon.
The early Hemphillian is defined (Tedford et al., in press) by the
earliest appearance of Arvicolinae (limited occurrence of
Microtoscoptes and Paramicrotoscoptes), Pliotomodon, and
Megalonychidae (Pliometanastes), limited occurrence late in
interval of Simocyon, Indarctos, Plionarctos, Lutravus, and
Eomellivora, and earliest appearance, late in interval, of
Mylodontidae (Thinobadistes), Machairodus and the Bovidae
(NeotragocerusT. Characterization earliest appearance of
Dipoides, Pliosaccomys, Pliotaxidea, Vulpes, 'Cams',
Osteoborus, and Cranioceras (YumacerasT!latest occurrence of
Amphicyonidae, Leptarctus, Sthenictis, Nimravides,
Barbourofelis, Epicyon, Pliohippus, Protohippus,
Cormohipparion, Prosthenops, Aepycamelus, Pseudoceras and
Plioceras.
In Florida, common early Hemphillan taxa include Calippus,
Pediomeryx (Yumaceras), Aepycamelus, primitive species of Osteoborus,
Cormohipparion, and Nannippus, and advanced species of Epicyon. The
early sloths Pliometanastes and Thinobadistes are present. The
Eurasiatic immigrants Indarctos and Machairodus appear in the later part
of the early Hemphillian.
The late Hemphillian is defined (Tedford et al., in press) by the
limited occurrence of Promimomys, 'Propliophenacomys',
Plesiogulo, Agriotherium, and Plionarctos and the earliest
appearance of Megalonyx, Ochotona, Megantereon, Felis,
Enhydriodon, and Cervidae. Characterization earliest
appearance of Taxidea, Borophagus, Rhynchotherium, Platygonus,
and Mylohyus, limited occurrence of Pediomeryx, latest
occurrence of Mylagaulidae, Osteoborus, Astrohippus,
Neohipparion, Dinohippus and Rhinocerotidae.
In Florida, common late Hemphillian taxa include advanced species of
Osteoborus, Neohipparion, Nannippus, Pseudhipparion, Teleoceras, and
Hipparion. Also present are Plesiogulo, Megalonyx, antilocaprids,
Agriotherium, Enhydriodon s.l., Megantereon, Machairodus, Rhynchotherium,
Pliornastodon, Dinohippus, Plionarctos, Felis, and cervids.


188
"Calidris" sp. 2
Material. Love Bone Bed local fauna, UF 26018, right coracoid.
Remarks. Coracoid similar in size to that of Calidris melanotos
and morphologically indistinguishable.
"Calidris" sp. 3
Material. McGehee Farm local fauna; UF 31776, distal end left
tarsometatarsus.
Remarks. Distal tarsometatarsus larger than that of Calidris
minutilla and C. ruficollis; smaller than that of £. fusicollis.
Specimen very tentatively referred to genus.
Genus indet. sp. k
Material. Love Bone Bed local fauna, UF 25760, right
carpometacarpus missing minor metacarpus.
Remarks. Carpometacarpus similar to that of Calidris minutilla.
Genus Actitis Illiger, l8ll
??Actitis sp. indet. sp 5
Material. Love Bone Bed local fauna, UF 29701, distal end right
tibiotarsus.
Remarks. Tibiotarsus is about the size of, and very similar in
morphology to Actitis macularia. Species 3 above may belong here.
Assignment to Actitis very tentative.
Genus Arenaria Brisson, 1760
??Arenaria sp. indet. sp. 6
Material. Love Bone Bed local fauna, UF 258l4, UF 29693, humeral
ends left coracoids.


156
Material. Bone Valley Mining District, Palmetto Mine; UF 21033,
distal end tibiotarsus.
Remarks. I have not been able to locate this specimen, but have
followed Steadman (I980:l4l) for this identification. There is additional
material of Meleagris in the UF collections from the Bone Valley Mining
District, but it is likely that it originated from Pleistocene deposits.
Remarks on the Family Phasianidae (Subfamily Meleagridinae).
See Steadman (1980) for an in-depth discussion of the osteology,
paleontology, systematics, and evolution of the subfamily Meleagridinae.


30
Table 2.1. List of acronyms of institutions and abbreviations of terms
used in the text.
Acronym
Institutions/Collections
AMNH
American Museum of Natural History
F:AM
Frick Collections, American Museum of Natural History
LACM
Los Angeles County Museum of Natural History
MCZ
Museum of Comparative Zoology, Harvard University
PB
Collection of Pierce Brodkorb
ROM
Royal Ontario Museum
UF
Florida State Museum, University of Florida
UM
University of Michigan
UMCP
University of California, Museum of Paleontology,
Berkeley
UNSM
University of Nebraska State Museum
USNM
United States National Museum
YPM
Yale Peabody Museum
Abbreviation
Terms
BMDP
Biomedical Statistical Program, P-series
1. f.
local fauna
M.
muscuius
MA
megannum (or million years)
max.
maximum
min.
minimum
mm.
millimeter
MYBP
million years before present
N
number (of specimens)
NALMA
North American Land Mammal Age


1+2
feet over 30 miles (the distance between the Manatee County Dam Site and
the "classic" Bone Valley exposures). This would also account for the
mixture of land and marine vertebrates in these sites, presuming the
tilting is post-depositional.
There is no question that there were eustatic sea-level changes
during this time period, such as the Messinian. But the elevations of
the Florida fossil vertebrate localities are of very doubtful value as
evidence. It should be noted that the rejection of the above hypothesis
does not prevent using the faunal composition of these localities to
determine their relative proximity to the ocean at the time of
deposition. They simply cannot be used as an absolute scale to measure
vertical sea-level changes.


Figure 4.4. Plot of greatest length (LENGTH) versus width of proximal end (W-PROX) of the tarsometatarsi
of the following species of vultures: (l) Vultur gryphus, (2) Sarcoramphus papa, (3) Gymnogyps
californianus, (4) Coragyps atratus atratus, (5) Cathartes aura, (A) Pliogyps fisheri, (B) Pliogyps
sp. from the Love Bone Bed 1^ fTj [c] Breagyps clarki, (D) Geranogyps reliquus~ (e) Gymnogyps howardae,
and (F) Gymnogyps amplus.
i


182
Family Jacanidae (Stejneger, 1885)
Characters. The Jacanidae may he distinguished from other families
in this order by having a tarsometatarsus with an extremely large distal
foramen with a deep tendinal groove leading into it and a deep pit
present on the medial surface of the inner trochlea (Olson, 1976).
Coracoid with procoracoid not perforated by a coracoidal fenestra
(perforated in all Charadriform families except the Rostratulidae,
Scolopacidae, Thinocoracidae, and Pedionomidae). Elongated tuberosity
(for attachment of part of the membrana sternocoracoclavicularis) present
on the procoracid and well developed in Jacanidae (less developed in the
Rostratulidae, small but distinct in Pedionomus; lacking in other
charadriforms).
Genus Jacana Brisson, 1760
Jacana farrandi Olson, 1976
Material. McGehee Farm local fauna (referred by Olson, 1976); UF
21219 (holotype), distal end of left tarsometatarsus, missing trochlea
IV; UF 11108 (paratype), left coracoid.
Love Bone Bed local fauna (referred by Becker); UF 25824, UF 29694,
UF 29696, UF 29698, humeral ends left coracoids; UF 260l6, UF 26026, UF
29690, UF 29691, humeral ends right coracoids; UF 29700, extreme proximal
end right humerus (tentatively referred); UF 25728, proximal end of left
humerus; UF 67806, distal end left tibiotarsus.
Description/ Remarks. Proximal end and shaft of humerus (UF 25728)
differs from Jacana spinosa by having shaft more gracile, pneumatic fossa
less deep, head of humerus not undercut by capital groove, and the small
papilla-like process not present. Deltoid crest broken. Coracoid
similar to the paratype coracoid described by Olson (1976). The


89
4.5 million years later than did A. grandis. Unfortunately, this species
is only known from ulnae and not from more diagnostic elements.
Remarks on the Family Anhingidae.
The earliest record of the Anhingidae is Protoplotus beauforti
Lambrecht based on a skeletal impression, from the late Eocene of
Sumatra. It is presently being restudied and will probably be referred
to a new family (P. V. Richin litt., cited in Olson, ms).
Anhinga pannonica Lambrecht was described from a cervical vertebra
and carpometacarpus from the late Miocene of Tataros, Hungary. Rich
(1972) also assigned another cervical vertebra and a partial humerus from
the late Miocene of Tunisia to this species. The only other anhinga
known from the late Miocene, Anhinga grandis, originally described from
the Hemphillian Cambridge (Ft.- 40) locality, Nebraska, is discussed
above.
The validity of A. laticeps Devis from the late Pleistocene of
Australia is somewhat questionable (Brodkorb and Mourer-Chauvir',, 1982;
Olson, ms.). Anhinga hadarensis Brodkorb and Mourer-Chauvire'", 1982 from
the Upper Pliocene Kadar Hadar member of the Hadar Formation, Ethiopia is
also known from the Omo Basin, Ethiopia, and Olduvai Gorge, Tanzania. It
appears to be the immediate ancestor to A. rufa (Brodkorb and Mourer-
Chauvire, 1982). Two Pleistocene fossil species of anhingas have been
shown to be cormorants. Anhinga parva Devis from Australia was shown by
Miller (1966) to be the cormorant Phalacrocorax melanoleucos and A. nana
Newton and Gadow, from Mauritius was shown by Olson (1975a) to be another
cormorant, Phalacrocorax africanus.


Family Strigidae Visors, 1825
Remarks. Femur referrable to the Strigidae by having a smooth
antero-dorsal condyle slope and having the postero-dorsal portion of the
external condyle joining the shaft abruptly (Ford, 1967).
Subfamily Buboninae Vigors, l8l5
Genus Bubo Burneril,l806
(cf.) Bubo sp.
Material. Bone Valley Mining District, Tiger Bay Mine. UF 29782,
distal end left femur.
Remarks. Distal end of the femur intermediate in size between that
of males and females of Bubo virginianus. Agrees with Bubo (and differs
from Strix and Asio) in placement of muscle scars, especially those above
the lateral condyle. Differs from Bubo virginianus and Nyctea scandica
by having the posterio-medial portion of the medial condyle merging
smoothly with the posterior intercondylar sulcus. In other Bubonini
examined, the posterior-medial portion of the medial condyle sharply
overhangs the popliteal fossa.
Remarks on the Family Strigidae.
There is several undescribed strigids awaiting a comprehensive
review. Olson (in press) briefly discusses much of this material.


2
Biochronology and Faunal Dynamics
1. What is the temporal distribution of the fossil localities
producing birds in the Neogene of North America?
2. What is the temporal distribution of the North American Neogene
avifauna?
3. When, and at what rate, do the North American Neogene avian
families and genera appear and become extinct?
4. How do the marine and non-marine localities and avifaunas differ
in questions 1 to 3 above?
5. What avian species are biostratigraphically useful in the
Neogene of North America?
6. What degree of temporal resolution does avian biochronology
offer?
Limitations of Study.
There have been several limitations imposed on this study by the
current knowledge of avian systematics and paleontology, and to a lesser
degree by the lack of previous studies dealing with the paleoecology and
biochronology of birds. Birds are often considered ". . the best known
and most completely described class of animals . (Welty 1975:14).
In reality, this statement applies only to the simple designation of
living forms and to the obvious aspects of their behavior and ecology,
but definitely not to their evolution, their superspecific relationships,
and to many aspects of internal morphology. Many living groups of birds
still lack modern systematic revisions based on internal morphology, or
in many, even a description of their internal morphology. Most modern
orders have never been shown to be monophyletic, although many are
doubtlessly so. Modern classifications of birds above the specific level


232
Brodkorb, P. 1967 Catalogue of Fossil Birds. Part 3*
(Ralliformes, Icthyornithiformes, Charadriiformes). Bulletin
Florida State Museum, Biological Sciences, 11:99-220.
Brodkorb, P. 1970. (Abstract). New discoveries of Pliocene birds in
Florida, p. 74. In. E. J. Brill (ed). Proceedings of the XVth
International Ornithological Congress, Leiden.
Brodkorb, P. 1971a. Catalogue of Fossil Birds. Part 4. (Columbiformes
through Piciformes). Bulletin Florida State Museum, Biological
Sciences, 15:163-266.
Brodkorb, P. 1971b. Origin and Evolution of Birds, pp. 19-55* In.
Avian Biology, Volume 1. Academic Press, New York and London.
Brodkorb, P. 1978. Catalogue of Fossil Birds. Part 5.
(Passeriformes). Bulletin Florida State Museum, Biological Sciences,
23:139-228.
Brodkorb, P. 1980. A new fossil heron (Aves: Ardeidae) from the Omo
Basin of Ethiopia, with remarks on the position of some other
species assigned to the Ardeidae. Contributions in Science, Natural
History Museum of Los Angeles County, 330:87-92.
Brodkorb, P. and C. Mourer-Chauvire". 1982. Fossil anhingas (Aves:
Anhingidae) from Early Man sites of Hadar and Omo, Ethiopia, and
Olduvai Gorge, Tanzania. Geobios, 15:505-515
Brodkorb, P. and C. Mourer-Chauvirel 1984. A new species of cormorant
(Aves: Phalacrocoracidae) from the Pleistocene of Olduvai^Gorge,
Tanzania. Geobios, 17:331-337.
Brown, L. H. and D. Amadon. 1969. Eagles, Hawks, and Falcons of the
World. Country Life, London.
Campbell, K. E., Jr. 1976. An early Pleistocene avifauna from Haile
XVA, Florida. Wilson Bulletin, 88:345-347.
Campbell, K. E., Jr. 1979- The non-passerine Pleistocene avifauna of
the Talara Tar Seeps, northwestern Peru. Publications in Life
Sciences, Royal Ontario Museum, 118:1-203.
Campbell, K. E., Jr. 1980. A review of the Rancholabrean avifauna of
the Itchtucknee River, Florida. Contributions in Science, Natural
History Museum of Los Angeles County, 330:119-129.
Campbell, K. E., Jr. and E. P. Tonni. 1983. Size and locomotion in
teratorns (Aves: Teratornithidae). Auk, 100:390-403.
Carr, G. S. 1981. An early Pleistocene avifauna from Inglis, Florida.
Ph. D. Dissertation. University of Florida, Gainesville.


Figure 2.3. Schematic diagrams illustrating measurements of the
femur and tibiotarsus. A. Tibiotarsus, caudal view. B.
Tibiotarsus, proximal end view. C. Tibiotarsus, distal end view.
D. Femur, cranial view. E. Femur, proximal end view. F. Femur,
distal end view. G. Femur, caudal view of distal end. Figures
are not drawn to scale. Measurements are defined in the text.


53
Tribe Podicipedini Storer, 19^3
Characters. Absence of a separate canal in the hypotarsus for the
tendon of insertion of the M. flexor perforatus digiti II (Storer, 1963).
See Murray (1967) for additional osteological characters.
Genus Podiceps Latham, 1787
Podiceps sp. indet.
Material. Bone Valley Mining District, Payne Creek Mine; UF 21205,
proximal end of left tarsometatarsus.
Remarks. Agrees with Podiceps in hypotarsal configuration (no
extra canal). Waterworn and abraded, and is not identifiable to species.
Similar in size to Podiceps nigricollis or P. o. occipitalis. (£. auritus
much larger).
Tribe indet.
Genus Pliodytes Brodkorb, 1953
Pliodytes lanquisti Brodkorb, 1953
Material. Bone Valley Mining District, Palmetto Mine; PB 299,
complete right coracoid (holotype).
Remarks. This species is known only from the holotype. Brodkorb
(l953e) states that it possesses characters in common with both
Podilymbus and Podiceps but also has its own unique characters. As the
tribes of grebes are defined on tarsometatarsal characters (Murray, 1967;
Storer, 1963), it is not possible to assign this genus to a tribe. As
more fossil material of grebes from the Bone Valley becomes available,
this species should be restudied to determine its generic validity and
relationships to other species.


birds then known, primarily the more numerous marine birds. Table 3.2
gives a partial list of birds now known from the Bone Valley Mining
District and notes the taxa (marine) not included here. The majority of
birds here studied have been collected in the last 15 years and mainly
include the rarer, non-marine members of this avifauna.
Manatee County Dam Site
This local fauna originates from a borrow pit south of the Manatee
River in Sec. 30, T. 3k S., R. 20 E., Verna Quadrangle, U. S. Geologic
Survey 7*5 minute series topographical map, 19kk, Manatee County. Like
the nearby SR-6k local fauna discussed below, the Manatee County Dam Site
is essentially an outlier of the classic Bone Valley local fauna. All
three share a similar fauna, geology, and paleoecology. Webb and Tessman
(1968) describe this local fauna and report the presence of one bird,
Phalacrocorax wetmorei.
SR-6k
This locality is located 6 miles east of 1-75 along State Road 6k in
Sec. 35, T. 3k S., R. 19 E., Manatee County, Lorraine Quadrangle, U. S.
Geological Survey 7.5 minute series topographical map, 1973, Florida. It
was discovered by Philip Whisler, of Venice, Florida, in 1983- The
majority of fossil vertebrates are in his private collection, except for
a few speciemens in the Florida State Museum collections. Numerous
fossil vertebrates are recorded from here, including several species of
bony fish, sharks, rays, and several different turtles. Mammals include
odobenids, phocids, Schizodelphis, Megalodelphis, tremarctine ursids, cf.
Agriotherium, felids, Nannippus minor, Neohipparion eurystyle,
Rhinocerotids, and Hexameryx. Based on the presence of Nannippus minor,


Copyright 1985
8y
Jonathan J
Becker


135
Owing to the long time interval between the Bone Valley local faunas and
the Love Bone Bed, it is unlikely that this specimen represents one of
the Bone Valley eagles.
Accipitrid, Genus indet.
Material. Withlacoochee River 4a local fauna; UF 67809, pedal
phalanx I, digit I; Bone Valley Mining District, District Grade (Agrico)
Mine; UF 57303, ungual phalanx (claw); Fort Green Mine, UF 61957, pedal
phalanx I, digit I (?); Palmetto Mine, UF 21123, UF 21125, ungual
phalanges (claws).
Remarks. All material is representive of a large accipitrid, but is
undiagnostic at the generic level.
Remarks on the Family Accipitridae.
The family Accipitridae had approximately 205 Recent species and 62
fossil species when Brodkorb published his Catalogue of Fossil Birds
(1964). Other species have been described since then. The systematics
of this family is based primarily on external characters (Brown and
Amadon, 1968), and probably does not accurately reflect the evolution of
this diverse group. Several recent studies (Jollie, 1976-1977; Rich,
1980) have described, in exhaustive detail, the morphology of this
family. But at the present, there is not a phylogeny of the family as a
whole based on internal morphology (but see comments in Olson, in press;
and Jollie, 1976-1977:309ff).
In Florida, there is also unreported accipitrid material from the
Hemingfordian Thomas Farm 1. f. and from several late Pleistocene
localities (Campbell, 1980; Carr, 1981).


202
Buteo jamaicensis and the holotype of Egretta subfluvia, am egret about
the size of Egretta ibis. This locality probably represents a pond
environment with some marine influence (Becker, 1985a).
Haile VB local fauna. The meager avifauna from this local fauna
consists of several specimens of an indeterminate species of anatid. The
paleoenvironment of this site was highly aquatic as shown by the abundant
material of the crocodylian, Gavialosuchus.
Haile VI local fauna. Birds known from here include the holotype of
Palaeostruthus eurius and several specimens of an indeterminate species
of duck. Based on the birds, little can be said of the paleoecology of
this locality.
Haile XIXA local fauna. Avian taxa from here include a cormorant
and a few specimens of anatids. Further studies on the paleoecology of
this site await additional systematic studies on the vertebrates present.
Bone Valley Mining District. While large numbers of fossil birds
have been collected from here, it represents the most limited for
paleoecological analysis because of collection techniques. This mining
district, which is well over 100 square miles in extent and is being
systematically strip-mined, resulting in the moving of hundreds of
thousands of cubic yards of sediment. Many specimens of fossil birds
were collected from spoil areas in conjunction with the commerical
collection of the more numerous shark teeth and mammalian cranial
fragments. Because of these collection techniques, usually it is not
possible to quantify the relative abundance of fossil specimens of birds,
the volume of sediments from which they originated, the sedimentary


211
the Clarendonian. In the Hemphillian and the Blancan it decreases
again to less than 0.1. In the non-marine avifauna the turnover
rate is 0.6 and 0.4 in the late Arikareean and Hemingfordian
respectively, and a rate of 0.2 in the Clarendonian. Other mammal
ages have a rate of 0.1 or less. The extremely high per-genus
turnover rate in the Arikareean may be an artifact of the low
sample size in this NALMA.
Discussion
Several factors could produce the results presented above, including
real changes in fossil avifaunas, but also including invalid assumptions
or errors in sampling. Certainly an obvious cause for some of the
changes in faunal dynamics is the unequal distribution of localities
within each mammal age. The absence of extinctions in the Hemingfordian
and Barstovian is probably an artifact resulting from an impoverished
prior record of taxa. The Clarendonian is the first NALMA in ihe Neogene
with a large number of both marine and non-marine localities. This
accounts at least for part of the peak in Or (marine) and the resultant
peak in turnover.
The mammalian fossil record in the Neogene has long been known to
consist of sequences of relatively long-lived "chronofaunas" whose
boundaries do not coincide with the boundaries of the NALMAs. It is
equally possible that birds exhibit a similar pattern of long periods of
faunal stasis with relatively rapid periods of turnover at the ends of
these periods. There is no a priori reason that these "avian" boundaries
would correspond with the "mammalian" boundary of the NALMA, or even with
the boundary of the mammalian chronofauna. Further work is needed to
reconstruct the database of Neogene localities from one keyed into


?Actitis, ?Arenaria, ?Philomachus, Tytonid, undescribed genus, Bubo,
Passeriformes (2).
The largest avifaunas are from the Love Bone Bed local fauna (44
taxa present) and the Bone Valley local fauna (4l taxa present, 31 here
included). These two localities are the most diverse non-marine and
marine avifaunas, respectively, known in North America prior to the
Pleistocene.
An analysis of the faunal dynamics of the Neogene fossil birds from
North America shows the following results, (l) Localities which have
produced fossil birds are not uniformly distributed through time74.4%
of the localities are from the last 4l% of the Neogene. (2) By the
Barstovian, a majority of the living families which have a fossil record,
have appeared. (3) Generic diversity increases from 10 to 98 during the
Neogene. (4) The marine avifauna is essentially established at a
diversity of 20 to 25 genera by the Clarendonian. (5) The non-marine
avifauna increases continually throughout the Neogene. (6) Origination
rates for marine birds peak in the Clarendonian with 4.4 genera appearing
per million years. (7) Extinction rates for marine birds are
consistently low throughout the Neogene. (8) Origination rates for non
marine birds show a 2- to 4-fold increase in alternate Land Mammal Ages.
(9) Turnover rates parallel the origination rates described above.
ix


Table 6.5 Comparisons of avian and mammalian faunal dynamics. Avian parameters include only non-marine
faunas, from Table 6.4. Mammalian parameters are from Marshall et al. (1982). The mammalian parameters for
the Hemphillian and Blancan are recalculated to account for the use of a different duration of these NALMAs.
AVIAN
CLAR.
HEMP.
Running mean (Rm)
26.50
39.00
Origination rate (Or)
8.00
4.22
Extinction rate (Er)
2.80
I.56
Turnover rate (T)
5.40
2.89
Per-genus Turnover (T/Rm)
0.20
0.07
T/Si
0.14
0.06
BLANC.
CLAR.
MAMMALIAN
HEMP.
BLANC.
56.00
52.00
50.00
52.00
10.00
17.20
16.67
20.00
1.85
14.80
18.00
l4.8l
5-93
16.00
17.34
17.41
0.11
0.31
0.35
0.33
0.08
0.17
0.14
0.18
1
219


last three years, he has donated hundreds of specimens of fossil birds
from the Bone Valley Mining District to the Florida State Museum.
Without his generous contributions, the Bone Valley portion of this study
would have been impossible to complete. John Waldrop is also gratefully
acknowledged for providing information about the avian localities in the
Bone Valley.
I also thank the following individuals and institutions for making
fossil and/or Recent specimens available for study: A. Andors, G.
Barrowclough, C. Houlton, R. H. Tedford, F. Vuilleumier, American Museum
of Natural History; P. Brodkorb, Department of Zoology, University of
Florida; J. Chenval, C. Mourer, Universite Claude Bernard; J. Hardy, B.
J. MacFadden, G. S. Morgan, S. D. Webb, T. Webber, Florida State Museum;
R. Mengel, University of Kansas; R. Payne, University of Michigan; M.
Voorhies, University of Nebraska; H. James, S. L. Olson, D. Steadman,
United States National Museum. Helen James, Storrs L. Olson, and David
Steadman generously provided accommodations during a lengthy stay in
Washington, D.C.
Financial support received while at the University of Florida,
includes teaching and research assistantships from the Department of
Zoology, College of Liberal Arts and Sciences; teaching assistantships
from the Department of Physiological Sciences, College of Veterinary
Medicine; a curatorial assistantship from the Department of Vertebrate
Paleontology, Florida State Museum; and grants from the Frank M. Chapman
Memorial Fund, American Museum of Natural History, and from Sigma Xi
Grants-In-Aid of research. The Department of Zoology, College of Liberal
Arts and Sciences, and the Florida State Museum supplied all expendable
iv


241
Shufeldt, R. W. 1917a. Report on fossil birds from Vero, Florida. In
Sellars, E. H. On the association of human remains and extinct
vertebrates at Vero, Florida. Journal of Geology, 25:18-19
Shufeldt, R. W. 1917b. Fossil birds found at Vero, Florida. Florida
Geological Survey 9th Annual Report, Tallahassee, pp. 35-42.
Shufeldt, R. W. 1918. Notes on some bird fossils from Florida. Auk,
35:357-358.
Simpson, G. 1930. Tertiary land mammals of Florida. Bulletin American
Museum of Natural History, 59:149-211.
Simpson, G. G., A. Roe, and R. C. Lewontin. i960. Quantitative Zoology.
Harcourt, Brace, and Co., Inc. New York.
Steadman, D. W. 1976. An addition to two Florida Pleistocene avifaunas.
Auk, 93:645-646.
Steadman, D. W. I98O. A review of the osteology and paleontology of
Turkeys (Aves: Meleagridinae). Contributions in Science, Natural
History Museum of Los Angeles County, 330:131-207
Steadman, D. W. 1981. A re-examination of Palaeostruthus hatcheri
(Shufeldt), a late Miocene sparrow from Kansas. Journal of
Vertebrate Paleontolgy, 1:171-173.
Steadman, D. W. 1984. A middle Pleistocene (Late Irvingtonian) avifauna
from Payne Creek, central Florida. Carnegie Museum of Natural
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Chihuahua, Mexico. Condor, 84:240-241.
Storer, R. W. 1963. Courtship and mating behavior and the phylogeny of
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Storer, R. W. 1967. Observations on Rol land's Grebe. El Hornero,
10:339-350.
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153.
Strauch, J. G., Jr. 1978. The Cladistic Relationships of the
Charadriiformes. Ph.D. dissertation. University of Michigan.


113
Table 4.15continued
Measurements Eudocimus ruber Eudocimus albus Eudocimus sp.
Coracoid
HEAD-CS
14.46 + 0.51
13.7 15.1
15.52 + 1.02
13.4 17.0
W-SHAFT
5.51 +0.31
5.2 6.1
5.93 + 0.4i
5.0 6.5

D-SHAFT
4.84 + 0.32
4.4 5.4
5.04 + 0.33
4.4 5.4

L-GLEN
9.50 + 0.35
8.9 9.9
10.50 + 0.44
9-8 11.1

Tarsometatarsus
W-PROX
10.76 + 0.83
10.1 12.6
11.47 + 0.98
10.2 12.8
10.3
D-MCOT
5.03 + 0.27
4.7 5.4
5.47 + 0.37
4.7 5.9
5.0
W-HYPOTS
5.24 + 0.37
4.8 5.8
5.65 + 0.54
4.9 6.5
5.5
D-PROX-L
9-33 + 0.49
10.33 + 0.73
9-5
8.8 10.4
9.1 11.3
_


93
the lateral condyle and tubercle in Mycteria (much more elevated in
Jabir and Ciconia, excluding C. abdimii and episcopus). Proximal
tarsometatarsus with sulcus ligamentosus sloping gently to the hypotarsus
in Mycteria (sharply sloping and usually deeply excavated in Ciconia and
Jabir). Tibiotarsi and tarsometatarsi of all species of ciconiids (l4
species total) were examined except those of Ciconia nigra,
Ephippiorhynchus senegalensis, and Leptoptilos crumeniferus.
Subfamily ftycteriinae American Ornithologists' Union, 1908
Genus Mycteria Linnaeus, 1758
Mycteria sp. A
Material. Love Bone Bed local fauna; UF 25990, proximal end right
tarsometatarsus (questionally referred).
McGehee Farm local fauna: UF 29^75, proximal end right
tarsometatarsus.
Description. UF 25990 with hypotarsus broken. Tentatively referred
to Mycteria by having the intercotylar prominence sharp and elevated, as
in modern species of Mycteria (prominence more rounded in Ciconia and
Jabir). UF 297^5 agrees with Mycteria by having the intercotylar
prominence highly raised and the sulcus ligamentosus sloping gradually
toward the hypotarsus (sharply notched in Ciconia). There are no
qualitative characters outside the range of variation of the modern
species Mycteria americana, except possibly having the hypotarsus
slightly lower on the shaft.
Remarks. As Table 4.12 shows, this fossil material is within the
range of a modern population of Mycteria americana. However, the
proportions are slightly different, with the proximal tarsometatarsus


MCOT (mm)
1 1-
10-
l
Q
9-
13
j p
14 15
W-PROX (mm)
8
16
nr
17
100


121
America which are listed in Brodkorb (1964b). Exactly how many of these
species are valid remains to be determined.
The earliest certain records for this family are therefore of late
Eocene and early Oligocene age in Europe and Asia; of mid to late Miocene
age in North America; and of Pliocene age in South America. The fossil
record would therefore suggest that this family had an Eurasian origin,
then invaded North America, and more recently South America.
An examination of the characters and proportions of the
tarsometatarsus and their variation in living and Neogene fossil species
suggests that the genera of vultures are oversplit. I agree with Mayr
and Short (1970) that Vultur Linnaeus and Gymnogyps Lesson (including G.
ampulus, G. howardae, and G. californianus) are congeneric. As Figure
4,4 and 4.5 show, the proportions of the tarsometatarsus of these species
differ little from each other. I would also tentatively include Breagyps
L. Miller and Howard and Geranogyps Campbell in the genus Vultur
Linnaeus, as they too are large vultures with similar tarsometatarsal
proportions.
However, I strongly disagree with Mayr and Short's (1970) opinion
that Pliogyps Tordoff should be included within the genus Vultur. The
tarsometatarsi of both species of Pliogyps differ greatly from Vultur in
proportions (Figure 4.4 and 4.5).
Therefore, on tarsometatarsal proportions, I would recognize the
following genera: Cathartes Illiger, Coragyps Geoffroy, Sarcoramphus
Dumeril, Pliogyps Tordoff, and Vultur Linnaeus, (including Gymnogyps,
Breagyps, and Geranogyps). If other skeletal elements show a similar
trend, I would urge adoption of these genera and their use as described
above.


3
(e.g., American Ornithologists' Union, 1983) has changed very little in
the 90 years since Gadov's (1893) classification. Olson (l98la:193),
addressing this problem, states
Many ornithologists appear to believe that the higher-
level systematics of birds is a closed book, the sequence of
orders and families in their field guides being an immutable
constant that was determined long ago according to some
infallible principle. In reality, the present classification
of birds amounts to little more than superstition and bears
about as much relationship to a true phylogeny of the Class
Aves as Greek mythology does to the theory of relativity. A
glance at the Gadov-Wetmore classification now in use shows
that there is still no concept in ornithology of what
constitutes a primitive bird.
Certainly correct phytogenies are impossible to develop without an
accurate knowledge of primitive character states, the distribution of
primitive and derived characters within the group being studied, and
meaningful outgroup comparisons.
Many fossil species are described from single, non-diagnostic
elements, making useful comparisons between, or among, species
impossible. But even diagnostic elements, when singly preserved, add
little to our understanding of the phylogeny and evolution of that taxon.
Other species are arbitrarily allied with the wrong family, the wrong
order, and even in some cases, with the wrong class of vertebrates (cited
in Brodkorb, 1978:211-228), many because of lack of proper comparisons.
Only recently have many of these errors been recognized, due in part to
an increased number of workers in the field, and from greater
availability and use of comparative skeletal material. Adequate skeletal
collections are still lacking for many common species (Zusi et al.,
1982).
It is outside the scope of this study to revise the many recent and
fossil genera and families which need such treatment. In such cases, I


236
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World. University of Nebraska Press, Lincoln. 493 pp.
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horse from the Miocene Phosphate Mining District of Central Florida.
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local fauna (Hemphillian, Mexico) and implications for the phylogeny
of one-toed horses. Journal of Vertebrate Paleontology, 4:273-283.
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Felidae) from Bone Valley Formation of Central Florida. Journal of
Paleontology, 55:218-226.


148


46


237
MacFadden, B. J. and J. S. Waldrop. 1980. Nannipus phlegon (Mammalia,
Equidae) from the Pliocene (Blancan) of Florida. Bulletin of the
Florida State Museum, Biological Sciences, 25:1-37.
MacFadden, B. J. and S. D. Webb. 1982. The succession of Miocene
(Arikareean through Hemphillian) terrestral mammalian localities and
faunas in Florida, pp. 186-199. In. Scott, T. M. and S. B.
Upchurch. Miocene of the southeastern United States. Tallahassee,
Florida Department of Natural Resources, Bureau of Geology, Special
Publication 25, 319 PP
Marshall, L. G., R. F. Butler, R. E. Drake, C. H. Curtis, and R. H.
Tedford. 1979 Calibration of the Great American Interchange.
Science, 204:272-279
Marshall, L. G., S. D. Webb, J. J. Sepkoski, Jr., and D. M. Raup. 1982.
Mammalian evolution and the Great American Interchange. Science,
215:1351-1357
Martin, L. and R. M. Mengel. 1975 A new species of anhinga
(Anhingidae) from the Upper Pliocene of Nebraska. Auk, 92:137-140.
Martin, L. and R. M. Mengel. 1980. A new goose from the Late Pliocene
of Nebraska with notes of variability and proportions in some Recent
geese. Contributions in Science, Natural History Museum of Los
Angeles County, 330:75-85
Mayr, E. 1969 Principles of Systematic Zoology. McGraw-Hill, New
York. 428 pp.
Mayr, E. 1981. Bioligical classification: toward a synthesis of
opposing methodologies. Science, 214:510-516.
Mayr, E. and G. W. Cottrell. 1979 Check-list of Birds of the World.
Volume 1. 2nd Edition. Harvard University Press, Cambridge,
Massachusetts.
Mayr, E. and L. L. Short. 1970. Species taxa of North American birds.
A contribution to comparative systematics. Publication of the
Nuttall Ornithological Club, 9:1-127.
McCoy, J. J. 1963. The fossil avifauna of Itchtucknee River, Florida.
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Miller, A. H. 1944. An avifauna from the lower Miocene of South Dakota.
University California Publication, Bulletin Department of Geological
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Miller, A. H. 1966. An evaluation of the fossil Anhingas of Australia.
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Miller, L. 1925. Avian remains from the Miocene of Lompoc. Carnegie
Institution of Washington, Publication, 349:107-117


183
coracoids range in size from similar to the paratype of Jacana farrandi
to decidely larger. This supports Olson's assertion (1976:261-262) that
the paratype of J. farrandi (UF 11108) is from a male. Tibiotarsus
similar to Jacana spinosa except for a reduced lateral epicondyle. For
type description and additional remarks see Olson (1976).
Remarks on the Family Jacanidae.
Rhegminornis calobates was originally described by Wetmore (l9*+3b)
as a jacana from the early Hemingfordian Thomas Farm local fauna. Olson
and Farrand (197*0 and later Steadman (1980) showed that this species has
affinities with the Phasianidae (Meleagridinae). Olson (1976) later
described the only fossil species of this family now known, Jacana
farrandi (discussed above).
The Jacanidae have long been allied with the Rostratulidae of South
America. In these two families, the females are larger than the males
and the young are similar (downy, strongly marked dorsally with black-
edged stripes from the forehead to tail). The similar osteology of these
two families support their association. Strauch (1978) presented a
subordinal classification of the Charadriiformes and associated the
Jacanidae, Rostratulidae, Scolopacidae, Phalaropodidae, and Thinocoridae
in his suborder Scolopaci. Characters used to support this
classification included the absence of the maxillo-palatine strut A,
absence of the coracoidal fenestra, and presence of a ridge in the
capital groove of the humerus. Olson and Steadman (1981) showed that
Pedionomus (Pedionomidae) was not only a Charadriiform, but had
affinities with at least one of these families (Thinocoracidae). The
relationships of these families need additional study.


6l
Table 4.2. Measurements of humeri and coracoids of the grebes Rollandia
rolland chiliensis (N =6, 2 males, 1 female, 3 unsexed), Tachybaptus
dominicus (N = 7, 4 males, 3 females), and Rollandia sp. from McGehee
Farm local fauna. Data are mean +_ standard deviation and range. (*)
specimen damaged. Abbreviations defined in methods section.
Measurement
R. r. chilensis
T. dominicus
Rollandia sp
Humerus
W-SHAFT
2.67 + 0.15
2.5 2.8
2.39 + 0.15
2.2 2.6
2.8
D-SHAFT
2.U7 + 0.12
2.3 2.6
2.11 + 0.15
1.9 2.3
2.6
W-DIST
5.35 + 0.23
5.1 5.7
5.00 + 0.31
1+.6-- 5.5
5.9
Coracoid
HEAD-FAC
25.1+5 + 0.71
24.3 26.2
21.80 + 1.57
20.1 24.2
*24.6
HEAD-IDA
21+.50 + 0.68
23.3 25.3
21.17 + 1.26
19-4 23.2
*23.3
HEAD-CS
7.03 + 0.23
6.7 7.1+
6.10 + 0.34
5.6"- 6.6
7.6
D-HEAD
2.17 + 0.10
2.17 + 0.17
__
2.0 2.3
2.0 2.4

W-SHAFT
2.30 + 0.18
2.1 2.6
1.94 + 0.21
1.7 2.3
2.1
D-SHAFT
1.65 + 0.10
1.5 1.8
1.29 + 0.09
1.1 1.4
1.7
FAC-IDA
8.53 + 0.38
8.2 9.1
7.44 + 0.4l
7.1 8.3

L-GLEN
1+.55 + 0.16
1+.1+ 5-8
4.31 + 0.25
3.8 4.5
5.3


44


26


96
Remarks. The presence of this species, species A. above, and
species C. below, shows that the genus Ciconia was much more diverse in
North America than was previously known.
cf. Ciconia sp. C
Material. Bone Valley Mining District, Swift Mine (=Estech); UF
52958, distal end left tibiotarsus. Palmetto Mine; UF 12470, distal end
left tarsometatarsus missing trochlea IV (tentatively referred).
Description. Distal end of tibiotarsus (UF 52958) extremely large,
in the size range of Jabir mycteria. Distal end slightly laterally
compressed (as in Jabir). As in Ciconia, the external ligamental
attachment is ridge-like and distal opening of tendinal bridge is toward
the edge of the bone (contra Jabir). See Figure 4.2 for comparisons
with other species of Ciconia.
Tarsometatarsus assigned here on the basis of size.
Remarks. Although referrable to the genus Ciconia on the basis of
the above characters, this specimen bears a striking resemblance to that
of Jabir mycteria. Were it not for Howard's (1942) study of Ciconia
maltha and Jabir nycteria, I would be tempted to suggest that C. maltha
and J. mycteria are congeneric. Ciconia sp. C. is probably closely
related to the clade which gave rise to Ciconia maltha.
Remarks on the Family Ciconiidae.
Living species of storks have been revised by Kahl (1971, 1972) who
synomized a number of raonotypic genera in Peters (1931). Wood (1983,
1984) has analyzed the phenetic relationships within the Ciconiidae.
Apart from the substantial criticisms which have be made on the use of
phenetics as a basis for classifications (Mayr, 1969; Hull, 1970;


163
relationships of the cranes above the species level. Wood's paper (1979)
unfortunately adds little new information; rather it addresses the
concordance between previously proposed classifications and the clusters
produced from multivariate analysis of different suites of characters.
The family Aramidae is here considered to be closely related to the
subfamily Balearicinae (Olson, ms). Fossil species of cranes (Gruidae +
Aramidae) are numerous throughout much of the Tertiary of North America
and Europe. They are listed in Brodkorb (1967), reviewed formally in
Cracraft (1973), and reviewed less formally by Olson (ms). Systematic
changes to Brodkorb (1967) may be found in the latter two publications.
My survey of mainly unpublished collections in the Frick
Collections, American Museum of Natural History, indicates there was a
great radiation of balearicinae cranes in North America during most of
the Tertiary (Becker, in prep; Olson, in press). Many fossil species
originally assigned to the Gruinae should be re-assigned to the
Balearicinae; and probably would have been long before now, were this
subfamily still extant in North America. The latest known occurrence of
this subfamily in North America is from the latest Hemphillian Bone
Valley local fauna.


164
Table 4.22. Measurements of the tibiotarsi and tarsometatarsi of Grus
canadensis tabida (N = 7, 4 males, 3 females), Grus canadensis canadensis
(N = 6, 3 males, 3 females), and Grus species A. from the Love Bone Bed
local fauna. Data are mean +_ standard deviation and range. Abbreviation
are defined in methods section.
Measurements
G. c. tabida
G. c. canadensis
Grus
Tibiotarsus
W-SHAFT
9.46 + 0.16
9.2 9.6
8.27 + 0.88
7.2 8.8
9.5
D-SHAFT
8.24 + 0.38
7.7 8.8
6.97 + 0.95
5.9 8.4
7.8
W-DIST-CR
18.73 + 0.84
17.7 20.1
17.33 + 2.09
15.2 20.3
18.8
W-DIST-CD
14.30 + 0.81
12.7 15.0
13.05 + 1.48
11.5 15.1
l4.i
D-MCON
18.73 + 0.76
17.5 19.7
17.52 + 2.14
15.1 20.6
17.9
D-LCON
18.39 + 0.60
17.3 19.1
16.80 + 1.86
14.7 19.6
17.0
D-ICON
10.29 + 0.38
9-7 10.9
9.32 + 0.86
8.3 10.4
9.7
Tarsometatarsus
W-PROX
20.83 + 0.99
19.8 22.2
19.23 + 1.93
17.3 21.6
22.5
D-PROX
13.37 + 0.33
12.9 13.8
11.70 + 1.01
10.4 13.1
12.7
D-PROX-L
19.57 + 0.80
18.5 20.5
17.82 + 1.59
l6.4 20.2
19.0


24


208
Families. The easiest statistic to calculate is the number of
families present in a given NALMA. In the late Arikareean 8 familes are
present of which 5 (63%) survive to the present. The number of families
rapidly increase to 18 (IT surviving; 94^) in the Hemingfordian, 31 (30
surviving; 97%) in the Barstovian, 37 (36 surviving; 97%) in the
Clarendonian, 40 (39 surviving; 98%) in the Hemphillian, and 4l (40
surviving; 98%) in the Blancan. By the Barstovian a majority of the
living families with a fossil record have appeared (Table 6.1).
Genera. The diversity (Si) of the genera in the Neogene of North
America begins with 10 (2 surviving; 20%) in the late Arikareean, 28 (ll
surviving; 39%) in the Hemingfordian, 38 (22 surviving; 56%) in the
Barstovian, 6l (43 surviving; 70%) in the Clarendonian, 78 (65 surviving;
83%) in Hemphillian, and increases to 98 (88 surviving; 90%) in the
Blancan.
Originations (Oi) and Extinctions (Ei) are given in Table 6.2.
These parameters are the simple counts of the first and last appearance,
respectively. Nine genera first appear in the late Arikareean. This
increases to about 20 in both the Hemingfordian and Barstovian and then
increases again to about 30 in the Clarendonian, Hemphillian, and
Blancan. Five extinctions occur in the late Arikareean, then 8
extinctions occur in each of the succesive NALMAs, with the exception of
the Clarendonian, which records 12 extinctions.
The raw counts of origination and extinctions are adjusted for the
unequal time interval in each of the NALMAs by dividing each count by the
duration of the given NALMA to produce Origination rates (Or) and
Extinction rates (Er) in Table 6.2.


200
shores of rivers and often in ponds with floating vegetation. Species of
Dendrocygna may also be found in shallow ponds with floating vegetation,
but they have a greater tolerance to brackish water. Scolopacids may be
found in grassy marshes, mudflats, estuaries, and edges of ponds.
Flamingos are found in shallow water to mudflats. It would seem likely
therefore, that the environments around the Love Bone Bed during the time
of deposition would include freshwater ponds and streams, but probably
also with wet marshes, streams, estuaries, and mudflats nearby.
Although the avifauna from the Love Bone Bed is substantial, the
usefulness of this collection for qualitative paleoecological analysis is
limited by collection techniques. As in most fossil vertebrate
localities, this site was excavated to maximize the number of mammalian
specimens recovered per unit time spent digging. While sediment samples
were screenwashed for small specimens, it was not done in a systematic
fashion. By using these methods, many of the smaller specimens
apparently were never collected. In the original study, only two one-
quarter cubic meter samples were collected for paleoecological analysis
(Webb et al. 1981: 553ff). Each of these small samples was from a
different stratigraphic unit; vertebrate remains were reported by weight
only with no indication of number of specimens given, or of the weight of
the sediment.
It should also be noted that the MNI (minimum number of individuals)
of birds as given by Webb et al. 1981: 538) based on preliminary
identifications is not MNI but rather the estimated number of species of
birds. Neither MNI nor number of specimens are valid indicators of
abundance when samples are strongly biased toward larger specimens. It


187
size class as Erolia penepusilla (above) and should be further compared
with this species.
Genus Calidris Merrem, 180^
Calidris pacis Brodkorb, 1963
Material. Bone Valley Mining District, near Brewster; PB 59*+
proximal end left humerus (Holotype).
Remarks. Brodkorb (1955a) states this species is almost identical
in size to Calidris canutus (see Brodkorb, 1955a:22 for measurements),
but differs significantly in the characters of the proximal end of the
humerus. Brodkorb (l955a:22) further suggests that this species may
require generic separation from Calidris when additional material is
known.
"Calidris" sp. 1
Material. Love Bone Bed local fauna; UF 25807, UF 29697, humeral
ends left coracoids; UF 26008, right coracoid; UF 29692, humeral end
right coracoid.
McGehee Farm local fauna; UF 3978, humeral end left coracoid; UF
9J+87, right coracoid.
Bone Valley Mining District, Ft. Green Mine (# 13 dragline); UF
539*+*+, right humerus lacking distal end.
Remarks. Skeletal elements similar in size to those of Calidris
minutilla, C. minuta, C. pusilla, and C. mauri. Slightly smaller than
those of C. subminuta females, C. ruficollis, £. bairdi, and C.
fusicollis; slightly larger than that of C. temnrinckii females.


ACKNOWLEDGMENTS
I would first like to thank my advisor, Pierce Brodkorb, for his
support, guidance, and friendship during the course of this project. He
has provided much encouragement and councel throughout ny researches on
fossil birds, as have the members of ny committeeDrs. Richard A.
Kiltie, S. David Webb, Elizabeth S. Wing, and Ronald G. Wolff. I thank
them for the many hours spent reading this manuscript and for their many
helpful comments pertaining to my research.
A number of friends and colleagues contributed to this study with
their comments, suggestions, and discussions. They include S. Emslie, R.
Hulbert, and A. Pratt. Gary S. Morgan has taken much time to discuss the
geology of Florida and systematics, evolution, and biochronology of
fossil and Recent mammals, in addition to many other aspects of
vertebrate paleontology. I especially thank him for his comments on
Chapter III. Storrs Olson has freely shared his considerable knowledge
and insights of avian systematics, evolution and anatomy. David Steadman
also provided many helpful comments. I also thank Cynthia West for her
support and aid in completing this dissertation.
I thank the many amateur fossil collectors who have generously
donated fossil birds from the included localites to the Florida State
Museum. They include Danny Bryant, George Hess lop, and Ron Love. Phil
Whisler, of Venice, Florida, originally discovered the SR-6U locality and
brought it to the attention of the staff of the Florida State Museum.
Rick Carter, of Lakeland, Florida, deserves special mention, for over the
iii


119
Tordoff's (l959:3^1ff) conclusions that Pliogyps is relatively a very
heavy-bodied, short-legged bird.
The intergeneric relationships of living and fossil vultures in
general, and Pliogyps in particular, are very difficult to determine,
owing to a paucity of pre-Pleistocene fossil specimens required to
determine believable character states. Pliogyps shares tarsometatarsal
characters with Sarcoramphus (anterior fossa continuing down shaft to the
distal foramen, a similar size of the distal foramen and a similar shape
of the hypotarsal ridge). Characters in which Pliogyps differs from
Sarcoramphus include the degree of elevation of trochlea III (proximal
border merging smoothly with shaft (plantar surface) in all modern
skeleton of Sarcoramphus examined), and the amount of excavation of the
lateral parahypotarsal sulcus. Pliogyps also shares characters with
Vultur and Gymnogyps including the lateral side of the area proximal to
trochlea IV being inclined, and the excavation of the anterior fossa
extending to the distal foramen (although to a lesser degree than in
Sarcoramphus).
Remarks on the Family Vulturidae.
Lign (1967) and Rea (1983) discuss the relationships of the family
Vulturidae with other avian families. There have been many fossil
species described as vultures; these are listed in Brodkorb (1961+b).
Those which have been subsequently moved to other families and orders
include: Lithornis vulturinus Owen, volant paleognath (Olson, ms.);
Palaeogyps prodromus Wetmore and Meocathartes grallator Wetmore, Family
Bathornithidae (Olson, ms.). Olson (ms.) also considers several other
"vultures", not to be sufficently diagnostic to be maintained in this


90
Order Ciconiiformes Garrod, 1874 (Auct.)
Family Ardeidae Visors, 1825.
Remarks. The following account briefly establishes the presence and
distribution of the late Miocene and early Pliocene herons in Florida for
the paleoecological and biochronological aspects of this study. More
detailed descriptions and systematic remarks may be found in Becker
(1985a). Systematic nomemclature follows Payne and Risley (1976).
Genus Ardea Linnaeus, 1758
Ardea polkensis Brodkorb, 1955
Material. Bone Valley Phosphate Mining District, Palmetto Mine; PB
380, proximal end of right tarsometatarsus (type), UF 21138, distal end
right tarsometatarsus; Payne Creek Mine; PB 7924, humeral end of right
coracoid.
Remarks. This heron is about the size of A. cinerea and is a rare
member of the Bone Valley avifauna. On the material now known for this
species, it is not possible to determine its relationship to other
members of the genus Ardea.
Ardea sp. indet.
Material. Love Bone Bed local fauna; UF 25939, distal end left
tarsometatarsus, missing trochlea IV.
Remarks. This specimen represents a species of Ardea about the size
of A. herodias occidentalis.
Genus Egretta T. Forster, 1817
Egretta subfluvia Becker, 1985
Material. Withlacoochee River 4a local fauna; UF 19001, right
tarsometatarsus lacking only trochlea IV and hypotarsus.
Remarks. This species is a small heron about the size of Egretta


166
Table 4.23continued
Measurements
G. americana
G. japonensis
Grus sp. B.
Tarsometatarsus
W-PROX
24.83 + 0.93
23.4 26.0
26.33 + 2.49
21.9 28.6
26.36 + 0.81 (5)
25.3 27.5
D-PROX
16.02 + 1.36
14.5 17.5
19.8
17.86 + 1.25 (5)
15.7 18.7
D-PROX-L
23.14 + 1.33
22.0 24.4
25.7
23.70 + 0.62 (5)
23.0 24.5
TRIII-TRIV
20.47 + 1.40
18.1 22.5
22.93 + 1.23
22.1 25.4
19.8; 21.8
TRII-TRIV
20.58 + 1.90
17.9 22.9
23.62 + 0.87
22.3 24.9
19.9
W-TRII
6.93 + O.58
6.5 8.0
7.95 + 0.29
7.6 8.4
7.48 + 0.35 (4)
7.3 8.0
D-TRII
13.60 + 0.45
12.8 14.0
15.15 + O.56
l4.4 15.9
*12.2; 13.3; l4.l
W-TRII
9.95 + 0.36
9.6 10.0
11.33 + 0.49
10.9 12.2
10.25 + 0.44 (4)
9.8 10.8
D-TRIII
12.33 + 0.56
11.5 13.0
13.98 + 0.59
13.0 14.5
13.3; *12.T; 13.4
W-TRIV
6.57 + 0.49
6.1 7.3
7.52 + 0.32
7.1 7-9
*6.0; 6.4; 6.6
D-TRIV
14.43 + 0.52
13.6 15.O
15.65 + 0.42
15.1 16.2
13.2; *13.4; 14.7


13
8. CPTB.Length from carpal tuberosity (Tuberculum carpale)
through lateral face of distal end.
9. ECON-CPTB.Length from external condyle through carpal
tuberosity.
10. EC0N-IC0N.Length from external condyle through internal
condyle.
Radius
1. LENGTH.Greatest length from the radial head (Caput radii)
through distal end of the radius (Extremitas distale radii).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. W-PROX.Greatest transverse width of proximal end.
5. D-PROX.Greatest depth of proximal end.
6. W-DIST.Greatest transverse width of distal end.
7. D-DIST.Greatest depth of distal end.
Carpometacarpus
1. LENGTH.Greatest length from most proximal portion of the
carpal trochlea (Facies articularis radiocarpalis of trochlea
carpalis) through facet for digit III (Facies articularis
digitalis minor).
2. W-PROX.Transverse width proximal end from ligamental
attachment of pisiform process (Processus pisiformes) to dorsal
surface (Facies dorsalis), measured at right angles to the long
axis of the shaft.
2a. W-CARPAL.Transverse width carpal trochlea measured at the
proximal edge of the articular facet.


Table 4.11. Measurements of the tarsoraetatarsi of the cormorants Phalacrocorax auritus auritus (N = l4, 7
males, 7 females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), and Phalacrocorax
wetmorei, from the Bone Valley Mining District, and Phalacrocorax species from the Love Bone Bed and McGehee Farm
local faunas. Data are mean +_ standard deviation, (N), and range. Abbreviations are defined in. the methods
section.
Measurements
P. a. auritus
P. a. floridanus
P. wetmorei
McGehee
Farm
Love
Tarsometatarsus
LENGTH
62.01 + 2.42
55.2 65.0
59.50 + 2.41
54.7 63.6
64.62 + 3.68 (5)
60.3 69.2
57.7

W-SHAFT
5.94 + 0.37
5.3 6.6
5.83 + 0.44
5.1 6.9
6.17 + 0.38 (13)
5.8 7.0
5.5;
6.2

D-SHAFT
5.24 + 0.24
4.9 5.6
5.27 + 0.52
4.6 6.2
5.53 + 0.44 (13)
4.9 -6.2
5.1;
5.1

W-PROX
12.84 + 0.44
12.0 13.6
12.20 + 0.57
11.0 13.4
12.96 + 0.47 (22)
12.1 14.1
11.5;
, 11.7

D-MCOT
8.70 + 0.38
8.0 9.2
8.44 + 0.29
7.9 9.0
9.01 + 0.58 (21)
8.1 10.0
7.9;
8.1

D-LCOT
7.42 + 0.44
6.8 7.9
7.25 + 0.49
6.5 8.3
7.22 + 0.42 (20)
6.iT 8.3
6.7;
7.2

W-HYPOTS
5.61 + 0.21
5.3 6.0
5.52 + 0.27
5.0 5.9
|
6.38 + 0.65 (9)
5.6" 7.8
5.5;
6.1

L-HYPOTS
9.67 + 0.67
9.1 10.7
9.74 + 0.62
9.0 11.0
10.62 + 1.02 (11)
8.7 12.6
9.6;
9.7



171
Humerus with shallow pneumatic foramen; wide and deep capital
groove. Crest-like margo caudalis extending from distal end of capital
groove (Fig. 4.9). Deltoid crest greatly expanded. Tarsometatarsi worn
and badly abraded and are assigned here mainly on the basis of size.
Distinguished from all genera examined (listed below) by having the
following combination of characters: coracoid with a large procoracoid,
flared medially, causing the triosseal canal to be open; dorsal surface
of coracoid not deeply excavated. Humerus with large deltoid crest and
well-developed capital shaft ridge.
Remarks. In trying to place this fossil material in a genus, I have
examined the skeletons of a large number of genera (number of species in
parentheses). For convience, nomenclature follows Peters (1934), but see
Olson (1973b) for phylogenetic relationships. Genera examined include:
Rallus (6), Alantisia (l), Ortygonax (2), Amaurolimnas (l), Ral lina (l),
Aramides (3), Gymnocrex (l), Gallirallus (2), Habroptila (l), Himantornis
(l), Canirallus (l), Crex (l), Limnocorax (l), Porzana (7), Laterallus
(4), Micropygia (l), Coturnicops (l), Neocrex (l), Sarothrura (l),
Poliolimnas (l), Porphyriops (l), Tribonyx (l), Amaurornis (l), Gallicrex
(l), Gall inula (2), Porphyrula (l), Porphyrio (3), and Flica (6).
None of the genera above match the characters of the fossil
material. There is a superficial resemblance of the coracoid (especially
the enlarged procoracoid process) to Canirallus, Gymnocrex, Aramides, and
Amaurolimnas. This appears to be due to shared primitive characters
between the fossil rail and these primitive living rails (Olson, 1973b).
Additional comparisons with other living and fossil rails are needed
to determine the systematic position of this fossil rail. Possibly
Flica infelix Brodkorb, known only from the distal end of a tibiotarsus,


52
Genus Podilymbus Lesson, 1831
Podilymbus cf. P. podiceps
Material. Bone Valley Mining District, Gardinier Mine; UF 65678,
distal half of left tibiotarsus; Palmetto Mine; UF 21147, distal half of
left tibiotarsus.
Description. Tibiotarsus similar in size and general morphology to
that of Podilymbus podiceps. Distinguished from the tibiotarsus of
Podiceps by having a smooth medial border of the medial condyle (notched
in Podiceps) and a less distinct depressio epicondylaris medialis (very
deep in Podiceps). Differs from that of Podilymbus podiceps as follows:
Shaft of UF 21147 slightly more robust than in males; tubercle slightly
proximal to medial attachment of supratendial bridge better developed;
deeper depressio epicondylaris lateralis than in most specimens. Other
characters within range of variation of modern populations of Podilymbus
podiceps. Measurements are given in Table 4.5.
Podilymbus sp. A
Material. Mixson Bone Bed local fauna; F:AM FLA 66-1115, proximal
end of right tarsometatarsus; F:AM FLA 66-1116, proximal end of left
tarsometatarsus.
Remarks. Both specimens are juvenile, at a similar stage of
ossification, and have identical morphology; they may represent the same
individual. Tarsometatarsal morphology similar to that of Podilymbus
podiceps, but differs by being smaller, having a sharper intercondylar
knob, and a smooth cranio-dorsal border of the medial cotyla (notched in
P. podiceps). Both specimens are too small to correspond to that
expected of Podilymbus cf. P. podiceps from the Bone Valley.
Measurements given in Table 4.3.


Order Anseriformes (Wagler, 1831)
136
Family Anatidae Vigors, 1825
Remarks. There is a large amount of fossil material representing
anatids from nearly all local faunas included in this study, especially
from the Love Bone Bed local fauna. Unfortunately, much of it consists
of fragmentary, waterworn specimens or specimens of slight diagnostic
value (radii, ulnae, vertebrae, etc.). The correct taxonomic assignment
of these specimens is further complicated by the presence of both goose
like ducks (Anatinae: Tadornini) and duck-like geese (Anserinae:
Dendrocygnini) which precludes assigning specimens to subfamily on the
basis of size. Therefore, I do not assign material to a taxonomic rank
and list it under referred material unless it is clearly diagnostic. The
classification proposed by Woolfenden (1961) is followed.
Subfamily Anserinae Vigors, 1825
Tribe Dendrocygnini Reichenbach, "1850"
Genus Dendrocygna Swainson, 1831 _
Generic characters. The coracoids of Dendrocygna may be
distinguished from those of all other anseriform genera by the presence
of a pit-like depression on the ventro-lateral surface of the sternal
end. Other characters of the coracoid are listed in Woolfenden (1961).
The carpometacarpus of Dendrocygna may also be distinguished from all
other anseriform genera by its narrow and elongate proportions, the
metacarpal II incurved in dorsal view, the external rim of the carpal
trochlea only slightly notched, and a prominent neck present between the
carpal trochlea and metacarpal III (Woolfenden, 1961).


18o
Table 4.26. Measurements of the tibiotarsi and tarsometatarsi of the
fossil species Phoenicopterus floridanus from the Bone Valley Mining
District, Phoenicopterus sp. A~ (large), and Phoenicopterus sp. B.
(small) from the Love Bone Bed local fauna. Data are mean +_ standard
deviation (number) and range. Abbreviations defined in the methods
section.
Measurement P. floridanus P. sp. A (lg.) P. sp. A (sm.)
Tibiotarsus
W-DIST-CR
15.09
+ 0.4i
(4)
17.53
+ 0.70
(8)
15.0;
15.8
14.5
- 15.4
16.3
- 18.2
W-DIST-CD
10.39
+ 0.55
(4)
12.23
+ 0.33
(8)
9.6;
11.1
9.9
- 11.0
11.8
- 12.8
D-MCON
17.49
+ 0.81
(4)
20.33
+ 0.58
(6)
16.8;
19.5
16.4
- 18.3
19-3
- 20.8
D-LCON
17.51
+ 0.74
(5)
20.32
+ 0.63
(7)
16.8;
19.1
16.8
- 18.5
19.5
- 21.5
Tarsometatarsus
W-TRIII
TRIII-TRIV
7.83 + 0.64 (3)
7.1- 8.3
8.3; 9.4
8.08 + 0.60 (5)
7.1 8.6
14.25 + 0.24 (4)
14.1 14.6
Combined
in column
to the
left
13.4; 15.0
15.60 + 0.35 (3)
15.4 16.O
TRII-TRIV


29
Er)/2; per-genus turnover rate is the turnover rate adjusted for average
diversity, calculated by dividing the total turnover rate (T) by the
total running mean (P.m). A sampling index was calculated as the number
of localities per Land Mammal Age divided by the duration of that
interval. Marshall et al. (1982) should be referenced for additional
qualifications of each statistic.
Specimens Examined
Fossil specimens included in this study are housed in the
collections of the Florida State Museum (UF), the collection of Pierce
Brodkorb (PB), and the Frick collections of the American Museum of
Natural History (F:AM). I have tried to include all known avian material
from the late Miocene through early Pliocene from Florida within these
collections. Non-diagnostic skeletal elements (vertebrae, phalanges,
etc.) were not considered. Fossil material accessioned into the UF
collections after 01 June 1984 (primarily from Bone Valley) was not
included in this study.
I relied on Recent comparative material in the collections of P.
Brodkorb, Florida State Museum, United States National Museum, American
Museum of Natural History, University of Michigan, and Royal Ontario
Museum.
Abbreviations
Table 2.1 lists the common abbreviations and acronyms used in this
dissertation. Anatomical abbreviations were given earlier.


74
genus Phalacrocorax (presently with 26 paleospecies), many without
extensive comparisons with other recent and fossil species to determine
the variability of the of the characters used. It would be desirable to
revise the paleospecies of Phalacrocorax and integrate this extensive
fossil record with the recent species to produce a phylogeny of the
family. There are large amounts of fossil material available, permitting
analysis of variation of several fossil species (e.g. P. wetmorei, P.
oweri, etc.), a recent descriptive osteological study (Ono, 1980), and
papers identifying osteological characteristics and proportions of the
various subgenera of cormorants (Howard, 1932a, 1965; Brodkorb and
Mourer-Chauvirfe, 1984). However, such a revision is beyond the scope of
this dissertation.
Species listed by Brodkorb (1963c) that are not cormorants include
all species of the genus Graculavus (moved to Charadriiformes, Olson and
Parris, ms), Actiornis anglicus (not a cormorant, nor ibis, Olson,
198lb), and Phalacrocorax mediterraneus (Gruiformes, Family
Bathornithidae = Paracrax antigua, Cracraft, 1971)
The earliest cormorants appear in the early Miocene. Phalacrocorax
subvolans Brodkorb from the mid-Hemingfordian Thomas Farm local fauna,
Florida, is known only from a proximal humerus. It is currently under
study (Becker, in prep.). Phalacrocorax marinavis Shufeldt from the
Arikareean John Day Formation, Oregon, is known from a humerus, ulna,
tarsometatarsus, and part of a femur. It is somewhat smaller than P.
carbo but is reported to be allied with this species. Phalacrocorax
miocaenus (Milne-Edwards) from the Aquitanian of Langy, Vaumas, St.-
Grand-le-Puy, and Montaigu, France, is known from most skeletal
elements. It was moved to a new genus Nectornis and is said to share


20.
18
D-TRIII.Greatest depth of trochlea III.
21. W-TRIV.Greatest transverse width of trochlea IV.
22. D-TRIV.Greatest depth of trochlea IV.
Computer Software
Biomedical Statistical Software, P-Series (Dixon, 198l) was used to
analyze many of the measurements. Programs used included BMDP1D (simple
descriptive statistics), BMDP6D (bivariate plots), and BMDP2M (cluster
analysis). Computations were made at the Northeast Regional Data Center
(NERDC) at the University of Florida, Gainesville.
Nomenclature
Common names have not been used for species. Few nomenclatural
systems have a more unstable, inaccurate, or confusing set of terms than
the American Ornithologists' Union's (1983) list of common names for
birds. I agree with J. L. Peters (l93^:ii)
. . inventing common English names for birds that do not have
them is a waste of time. After all, the primary reason for a
scientific name is to have a name intelligible to scientists
the world over.
Systematic nomenclature generally follows the Checklist of Birds of
the World (Mayr and Cottrell, 1979; Peters, 1931-1951) for Recent species
and Brodkorb's Catalogue of Fossil Birds (1963-1978) for fossil species.
Departures are accompanied by full citation.


TABLE OF CONTENTS
PAGE
ACKNOWLEDGMENTS iii
ABSTRACT viii
CHAPTER
I. INTRODUCTION AND PREVIOUS WORK 1
Introduction 1
Limitations of Study 2
Previous Work U
II. METHODS 9
Measurements 9
Computer Software 18
Nomenclature 18
Systematics 27
Paleoecology 27
Biochronology and Faunal Dynamics 27
Specimens Examined 29
Abbreviations 29
III. GEOLOGY 31
Biochronology 31
Local Faunas 33
Eustatic Sea-level Changes 1+1
IV. SYSTEMATIC PALEONTOLOGY 1+9
Order Podicipediformes 1+9
Order Pelecaniformes 65
Order Ciconiiformes 90
Order Accipitriformes Il6
Order Anseriformes 136
Order Galliformes 155
Order Ralliformes 158
Order Charadriiformes 175
Order Strigiformes 191
Order Passeriformes 195
vi


128


38
from one mine, or a single dragline operating within one mine, are
considered as coming from one broad "locality." The exact geographic
position of these localities range from precisely known (1/4, 1/4, 1/4
section) to generally known (from one minei.e. somewhere within 1 to
10 sections). For a few amateur collections where even the mine is
unknown, the only designation can be the Bone Valley Mining District
(i.e. somewhere within 100 square miles), but the majority of specimens
are identified as coming from one mine. Table 3.1 lists the mines, mine
codes, describes their approximate location, and lists the stratigraphic
codes used.
The age of the Bone Valley fossil vertebrates was much debated from
the 1920s through the 1950s (discussed in Brodkorb 1955a), with the
proposed age ranging from the Miocene through the Pleistocene. Brodkorb
(1955a), using a Lyellian method of percent extinct species in the fauna
and the temporal ranges of three species, suggested that the age of the
Bone Valley avifauna was between the late Miocene and middle Pliocene,
probably early or middle Pliocene (i.e. Clarendonian or Hemphillian; =
late Miocene of current usage).
The majority of fossil land mammals are late Hemphillian in age and
compare well with others of similar age in North America. There is no
evidence to suggest that the fossil birds, which are found in association
with these land mammals, are of a different age. Recently several older
local faunas (Barstovian, Clarendonian, and early Hemphillian) have been
found (MacFadden, 1982; MacFadden and Webb, 1982; Webb and Crissinger,
1984; Berta and Galiano, 1984; and Tedford et al., in press). These
older occurrences are from the Phosphoria, Nichols, Silver City, and
Kingsford mines and to ny knowledge have produced no fossil birds. There


Figure 4.2. Plot of transverse width of the distal end, across the
cranial surface (W-DIST-CR) versus the depth of the area
intercondylaris (D-ICON) of the tibiotarsi of the following
ciconiid species: (l) Ciconia abdimii, (2) C. episcopcus, (3) C.
nigra, (4) C. ciconia, (5) C. maguari, (6) C. maltha from the
Pleistocene of Florida, (7?~Jabir mycteria, (triangles) Ciconia
sp. A from the Love Bone Bedj (squares) Ciconia sp. B from the
Mixson and Bone Valley local faunas, and (open circle) Ciconia sp.
C from the Bone Valley local fauna.


153
Table 4.20. Measurements of coracoids, humeri, carpometacarpi, and
tarsometatarsi of males of Anas hottentata and Anas sp. A from the Love
Bone Bed and McGehee local faunas. Data are mean _+ standard deviation
and range. Abbreviations defined in methods section.
Measurement
Anas hottentata
Anas sp. A
Coracoid
HEAD-IDA
29.7; 30.6; 29.7
25.85 + 0.72 (4)
24.8 26.4
HEAD-CS
9.7; 9.5; 9.7
8.93 + 0.43 (4)
8.3 9-2
W-SHAFT
3.2; 3.1; 3.0
3.06 + 0.23 (7)
2.8 3.4
D-SHAFT
2.2; 2.5; 2.6
2.56 + 0.50 (7)
2.1 3.6
IDA-PP
22.8; 24.0; 22.9
20.37 + 0.34 (6)
19.8 20.7
L-GLEN
5-8; 6.0; 6.1
6.10 + 0.23 (7)
5.B- 6.5
Humerus
LENGTH
54.6; 56.2; 54.2
50.8
W-SHAFT
4.1; 4.0; 4.3
3.63 + 0.31 (4)
3.2 3.9
D-SHAFT
3.5; 3.4; 3.7
3.03 + 0.49 (4)
2.Â¥ 3.5
W-PROX
11.5; 11.6; 11.6
12.2; 11.6
D-PROX
6.2; 6.5; 6.3
6.4; 5.3
D-HEAD
4.1; 4.4; 4.4
3.80 + 0.13 (4)
3.7 4.0
W-DIST
8.2; 8.3; 8.6
7.8
D-DIST
4.8; 4.8; 5.2
4.8
D-ENTEP
3.5; 3.5; 4.0
4.0


184
Table 4.28. Measurements of the humeri and tibiotarsi of Jacana spinosa
(N = 12, 6 males, 6 females) and Jacana farrandi. For measurements of
the type (tarsometatarsus) and paratype (coracoid) see Olson (1976).
Data are mean +_ standard deviation and range. Abbreviations are defined
in the methods section.
Measurements J. spinosa J. farrandi
males females
Humerus
W-SHAFT
2.67
+ 0.05
3.10
+ 0.13
3.4
2.6
- 2.7
3.0
- 3.3
D-SHAFT
2.25
+ 0.08
2.70
+ 0.09
2.9
2.1
- 2.3
2.6
- 2.8
W-PROX
8.25
+ 0.23
9.70
+ 0.52
9-6
7.9
- 8.4
9-0
- 10.6
D-PROX
4.68
+ 0.12
5.48
+ 0.18
5.9
4.5
- 4.8
5.2
- 5.7
D-HEAD
2.45
+ 0.08
2.82
+ 0.08
3.0
2.4
- 2.6
2.7
- 2.9
Tibiotarsus
W-SHAFT
2.33
+ 0.10
2.67
+ 0.14
2.8
2.2
- 2.5
2.5
- 2.8
D-SHAFT
1.93
+ 0.08
2.25
+ 0.14
2.6
1.8
- 2.0
2.1
- 2.4
W-DIST-CR
4.70
+ 0.11
5.35
+ 0.08
5.6
4.5
- 4.8
5.3
- 5.5
W-DIST-CD
3.52
+ 0.12
3.92
+ 0.04
3.9
3.4
- 3.7
3.9 4.0
D-MCON
5.28
+ 0.17
5.70
+ 0.20
5.8
5.1
- 5.5
5.4
- 5.9
D-LCON
4.90
+ 0.13
5.37
+ 0.10
5.6
4.7
- 5.0
5.2
- 5-5
D-ICON
3.32
+ 0.18
3.50
+ 0.l4
3.7
3.1
- 3.6
3.3
- 3.7


122
Anti 1 lovultur Arredondo was described from a late Pleistocene cave
deposit in Cuba. The type is a 42.5 mm tarsometatarsal fragment which
lacks both the proximal end and the distal one-half of the bone. It is
also known from a referred distal portion of a humerus, a trochlea IV of
a tarsometatarsus, and a single cervical vertebra. The description
(Arredondo, 1976) does not clearly differentiate this material from that
of the genus Vultur.


190
visitor to North America today and has never been noted from Florida, I
hesitate to report it as a fossil until I make additional comparisons
with other recent and fossil species; and can distinguish the proximal
humerus of all genera of scolopacids with certainty by a differential
diagnosis.
Remarks on the Family Scolopacidae.
There are some 20 fossil species of scolopacids, many of which are
in need of revision. Reference to these may be found in Olson (ms) and
Brodkorb (1967). Living species are treated in Johnsgard (1981).


Table 6.4 Faunal dynamics of non-marine Neogene birds of North America. Abbreviations as in Table 6.1.
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Duration (MA)
3
3.5
5
2.5
4.5
2.7
Localities (Published)
2 (2)
10 (5)
15 (9)
22 (14)
29 (15)
28 (15)
Sampling Index
0.67
1.43
1.80
5.60
3.33
5.56
Number of genera (Si)
7
25
26
4o
52
72
Originations (No.)
6
21
9
20
19
27
Extinctions (No.)
3
8
6
7
7
5
Running mean (Rm)
2.50
10.50
18.50
26.50
39-00
56.00
Origination Rate
2.00
6.00
1.80
8.00
4.22
10.00
Extinction Rate
1.00
2.29
1.20
2.80
1.56
1.85
Turnover Rate (T)
1.50
U.15
1.50
5.40
2.89
5-93
T/Rm
0.60
0.40
0.08
0.20
0.07
0.11
T/Si
0.21
0.17
0.06
0.14
0.06
0.08
i
218


Figure 4.3. Plot of transverse width of distal end across the
caudal surface (W-DIST-CD) versus depth of medial condyle (D-MCON)
of the tibiotarsi of the following species of ibis: (l) Eudocimus
albus, (2) E. ruber, (3) Plegadis falcinellis, (4) F. chihi, (3)
P. ridgwayi, () Eudocimus sp. from the Lee Creek 1. f., (b)
Eudocimus sp. A from the Bone Valley 1. f., and (C) Plegadis cf. P.
pharangites from the Love Bone Bed local fauna.


Table 4.8. Measurements of carpometacarpi of the cormorants Phalacrocorax auratus auritus (N = 14, 7
males,? females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), Phalacrocorax vetmorei,
from the Bone Valley, and Phalacrocorax species from McGehee Farm and Haile XIXA. Data are mean +_ standard
deviation, (N), and range. Abbreviations defined in methods section.
Measurements
P. a. auritus
P. a. floridanus
P. vetmorei
Haile XIXA
McGehee
Carpometacarpus
LENGTH
69.OI + 2.49
63.8 73.3
66.51 + 2.74
62.6- 70.1
69.6 (1)
'
W-PKOX
7.79 + 0.25
7.4 8.1
7.31 + 0.36
6.7 8.0
6.93 + 0.36 (26)
5.9 7.6
6.8
8.3
W-CARPAL
6.29 + 0.30
5.9 6.9
6.00 + 0.27
5.5 6.4
6.o4 + 0.26 (28)
5.F 6.4
6.1
5.6; 7.2
D-PROX
13.30 + 0.40
12.7 14.1
12.54 + 0.51
11.5 13.3
13.68 + 0.39 (24)
13.1 14.2
12.6
12.9; 16.4
L-MCI
10.64 + 0.47
9.4 11.3
10.06 + 0.68
8.9 11.4
9.87 + 0.38 (23)
9.3 10.5
9.1
9.7; 11.5
D-SHAFT
3.54 + 0.19
3.3 3.9
3.18 + 0.17
2.9 3.4
3.69 + 0.34 (8)
3.3 4.3

4.6
W-SHAFT
4.69 + 0.23
4.3 5.2
4.34 + 0.21
4.0 4.8
4.68 + 0.12 (8)
4.^ 4.9

5.4
D-DIST
5.01 + 0.23
4.6 5.3
4.71 + 0.29
4.0 5.2
4.91 + 0.33 (8)
4.F 5.4

4.8
W-DIST
7.44 + 0.22
7.1 7-7
7.08 + 0.37
6.3 7.7
7.42 + 0.21 (9)
7.2 7.8

7.1


132
species of Pandion was described from the late Clarendonian Love Bone Bed
local fauna and a phylogeny was proposed for this family (Becker, 1985b).
Family Accipitridae (Vieillot, l8l6)
Genus Haliaeetus Savigny, l809
? Haliaeetus sp.
Material. Bone Valley Mining District, Palmetto Mine; UF 21136,
distal end left humerus. Fort Green Mine; UF 55819, distal end left
tibiotarsus; UF 01956, distal end left tibiotarsus.
Description. Humerus similar in size and morphology to a small
Haliaeetus _1. leucocephalus. Provisionally assigned to the genus
Haliaeetus. Distinguished from Necrosyrtes and Neophron by having a
broader attachment for the anterior articular ligament. Distinguished
from Aguila, Necrosyrtes, and Neophron by having a narrow intercondylar
furrow (=incisura intercondylaris).
Tibiotarsi also the size of the modern Haliaeetus _1. leucocephalus.
There is little variation in size between the two tibiotarsi. UF 55819
has a slightly more slender shaft and a more horizontally placed tendinal
bridge.
Remarks. The distal tibiotarsus is rather undiagnostic in
accipitrids (Jollie, 1976-1977: 226-227), although it has been used in
the past to define species. This material is tentively assigned to
Haliaeetus on size, overall similarity, and the differences noted above.
The variablity of the characters used above has not been determined.
Additional analysis of the characters used by Rich (1980) to separate the
various subfamilies and genera of accipitrids is needed before this
material can be correctly placed within this family.


179
that of P. floridanus. Along with these small specimens are a group of
larger specimens which represent a much larger flamingo. If this
variation is considered to be sexual in origin, then there is more size
variation than has ever been observed in any single living or fossil
species of flamingo. This suggests that there are two species of
flamingos present at the Love Bone Bed, which overlap in size. The
similarity of the smaller flamingo from the Love Bone Bed to that of P.
floridanus from the Bone Valley (4 MA later in time) suggest that
flamingos have had a slow evolutionary rate. Pending further studies on
other Miocene flamingos, particularly P. stocki, I have left these
Florida specimens unassigned to species.
Remarks on the Family Phoenicopteridae.
Fossil species of Phoenicopterus are now known from the Aquitanian
in Europe and the late Miocene to the late Pleistocene of North America.
Phoenicopterus croizeti Gervais, was described from the Aquitanian of
France. It is known from abundant material and has recently been
restudied (Cheneval, 1984). Phoenicopterus stocki Miller, is based on a
distal tibiotarsus from the Hemphillian Yepomera local fauna. It is said
to have the morphological characters of the genus, but is of pigny size
(Miller, 1944: 77). Phoenicopterus floridanus Brodkorb, is discussed
above. Other Neogene species of Phoenicopterus include only P.
novaehollandiae A. Miller, from the late Oligocene or early Miocene of
Australia. Pleistocene species of Phoenicopterus include P. copei
Shufeldt, from Fossil Lake in Oregon and Manix Lake in California and £.
minutus from Manix Lake in California. All North American fossil species
should be revised and analysed with a larger database of skeletons to
account adequately for the large size variation present.


103
Table 4.12. Measurements of the tarsometatarsi of the storks Mycteria
americana (N = 10, 5 males, 5 females) and Mycteria species A. from the
Love Bone Bed and the McGehee Farm local fauna. Data are mean +_ standard
deviation and range. Abbreviations are defined in the methods section.
(*) Specimen broken.
Measurements Mycteria americana Mycteria sp. A
Tarsometatarsus
W-PROX
15.19 + 0.90
13.9 16.9
17.0;
l6.1
D-MCOT
9.30 + 0.56
8.4 10.3
9-4;
8.6
D-LCOT
8.88 + 0.60
8.1 9.8
9.6;
9.5*
W-HYPOTS
9.66 + 0.62
8.5 10.5
9.5
D-PROX-L
16.57 + 0.94
15.3 17.8
17.2


Table 4.13. Measurements of the tibiotarsi of the Recent storks Jabir
mycteria (N= 13, 2 males, 5 females, 6 unsexed), Ciconia maguari (N = 5,
1 male, 1 female, 3 unsexed), and Ciconia ciconia (N= 6, 2 males, 4
unsexed). Data are mean +_ standard deviation and range. Abbreviations
are defined in the methods section.
Measurement
Ciconia ciconia
Ciconia
maguari
Jabir nycteria
Tibiotarsus
W-SHAFT
8.38 + 0.56
9.50+1
3.48
11.13
+ 0.91
7.6 -
9.0
8.8 :
10.0
9.6 -
- 12.7
D-SHAFT
7.22 + C
5.48
8.26 + (
D.30
10.37
+ 0.64
6.6 -
7.7
7.8 i
3.6
9-3
- 11.1
W-DIST-CR
14.98 +
0.47
17.72 +
0.66
19.21
+ 1.58
14.4 -
15.8
17.0 -
18.5
16.3
- 22.3
W-DIST-CD
11.87 +
0.95
14.56 +
0.35
16.35
+ 1.31
10.7 -
13.0
14.0 -
14.9
13.6
- 18.9
D-MCON
18.97 +
1.13
21.46 +
0.95
25.78
+ 1.76
17.9 -
20.4
20.6 -
22.8
23.4
- 28.7
D-LCON
18.65 +
0.55
21.14 +
0.69
24.37
+ 1.68
18.4 -
19.3
20.4 -
22.1
22.2
- 27.4
D-ICON
11.60 +
0.56
13.52 +
0.6l
16.31
+ 1.07
11.1 -
12.3
12.8 -
14.5
15.1
- 18.3


130
Table 4.18. Ratios of intertarsal flexion and tarsometatarsal robustness
of species of vultures. Flexion ratio is calculated by (FLEXOR + LENGTH)
X 100; Robustness ratio calculated by (W-SHAFT + LENGTH) X 100.
Increasing values are correlated with increasing force of flexion and
increasing robustness, respectively. (*) approximate.
RATIO
Species
LENGTH
FLEXOR
W-SHAFT
FLEXION
ROBUSTNESS
P. undesc. sp
86.6
14.9
10.8
17.2
12.4
P. fisheri
94
14.5
11.4
15.4
12.1
S. papa
*91
*16
*10.5
*16.5
*10.8
G. californica
*116
*22
*13.5
*19.0
*11.6
V. gryphus
*126
*22
*14
*17.4
*11.1
C. atratus
88.4
12.8
7.2
14.5
8.1
C. aura
65.9
11.3
7.5
17.1
11.4


Table 6.1. Number of families and genera present in each North American Land Mammal Age and the percentage
of these which are still living. See text for discussion of the division of families and genera into marine
and non-marine groups. Abbreviations: L. ARIK. late Arikareean (23.0 20.0 nybp), HEMING.
Hemingfordian (20.0 16.5 mybp), BARST. Barstovian (16.5 11.5 mybp), CLAR. Clarendonian (11.5 9*0
nybp), HEMP. Hemphillian
(9*0 4.5 mybp)
, BLAN. Blancan (4.5
- 1.8 mybp).
North American
genera
which lack a fossil record ¡
are omitted from
this table.
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Number families present
Marine
3
3
9
10
12
11
Non-marine
5
15
22
27
28
30
Total
8
18
31
37
40
4i
Number (%) living families
Marine
1 (33)
2 (66)
8 (89)
9 (90)
11 (92)
11 (100)
Non-marine
4 (80)
15 (100)
22 (100)
27 (100)
28 (100)
29 (97)
Total
5 (63)
17 (94)
30 (97)
36 (97)
39 (98)
40 (98)
Number genera present
Marine
3
3
12
21
26
26
Non-marine
7
25
26
40
52
72
Total
10
28
38
6l
78
98
Number {%) living genera
Marine
1 (33)
1 3 (100)
9 (75)
14 (67)
22 (85)
24 (92)
Non-marine
1 (14)
8 (32)
13 (50)
29 (73)
43 (83)
64 (89)
Total
2 (20)
11 (39)
22 (58)
43 (70)
65 (83)
88 (90)
215


32
30
28
E 26
E
X
24
22-
oc
Q.
I 20-
£
18-
16
14
12
104-
60
\
5
70
80
i 1 1 1 1
90 100 110 120 130
LENGTH (mm)
140
124


234
Ford, N. L. 1967 A Systematic Study of the Owls based on Comparative
Osteology. Ph. D. Dissertation. University of Michigan, Ann Arbor.
Friedmann, H. 19^7 Geographic variations of the Black-bellied,
Fulvous, and White-faced Tree Ducks. Condor, 49:189-195
Frbringer, M. 1888. Untersuchengen zur Morphologie und Systematik der
Vttgel, zugleich ein Beitrag zur Anatomie der Sttttz und
Bewegungsorgane. 2 vol. Van Holkema, Amsterdam. 1751 pp. 30 pis.
Gadow, H. 1893. V8gel. II. Systematischer Theil. Iri Dr. H. G.
Bronn's Klassen und Ordnungen der Thier-Reichs, Vol. 6(4). C. F.
Winter, Leipzig. 303 pp.
Gilbert, B. M., L. D. Martin, and H. G. Savage. 1981* Avian Ostelogy.
Author, Laramie, Wyoming. 252 pp.
Hamon, J. H. 1959* Northern birds from a Florida Indian Midden. Auk
76:533-534
Hamon, J. H. 1964. Osteology and paleontolgy of the passerine birds of
the Reddick, Florida, Pleistocene. Florida Geological Survey,
Geology Bulletin, 44:1-210.
Harrison, C. J. 0. 1981. A re-assignment of Amphipelargus [sic] majori
from Ciconiidae (Ciconiiformes) to Ergilornithidae (Gruiformes).
Tertiary Research, 3:111-112.
Harrison, J. A. 1981. A review of the extinct wolverine, Pleisogulo
(Carnivora: Mustelidae), from North America. Smithsonian
Contributions to Paleobiology, 46:1-27.
Harrison, J. A. and E. M. Manning. 1983* Extreme carpal variability in
Teleoceras (Rhinocerotidae, Mammalia). Journal of Vertebrate
Paleontology, 3:58-64.
Hay, 0. P. 1902. On the finding of the bones of the Great Auk (Plactus
impennis) in Florida. Auk, 19:255-258.
Hay, 0. P. 1923. The Pleistocene of North America and its vertebrate
animals from the states east of the Mississippi River and from the
Canadian provinces east of longitude 95 degrees. Carnegie
Institution of Washington, Publication, 322:i-viii,1-499.
Hirschfeld, S. E. 1968. Vertebrate fauna of Nichol's Hammock, a natural
trap. Quarterly Journal of the Florida Academy of Sciences, 31:177-
189.
Hirschfeld, S. E. and S. D. Webb. 1968. Plio-Pleistocene megalonychid
sloths of North America. Bulletin of the Florida State Museum,
Biological Sciences, 5:213-296.


189
Remarks. The fossil coracoids are very similar in size and
morphology to the males of Arenaria melanocephala, but are slightly more
robust. Assignment to genus very tentative.
Genus indet. sp. 7
Material. Love Bone Bed local fauna, UF 25871, distal end left
tibiotarsus; UF 25858, distal end right tarsometatarsus.
Remarks. Fossil skeletal elements similar to those of females of
Limosa fedoa. Brief comparisons with Limosa ossivallis shows the
tibiotarsus from the Love Bone Bed local fauna to have a smaller
transverse width of both the shaft and the anterior portion of the distal
end and is of a slightly smaller size. Additional comparisons are needed
with Limosa vanrossemi from the Miocene of California.
Genus indet. sp. 8
Material. Love Bone Bed local fauna, UF 25810, UF 25822, UF 29695,
UF 29699, left coracoids, broken and abraded; UF 26009, UF 26013, UF
26028, UF 29867, UF 29688, UF 29689, right coracoids, broken and abraded.
Remarks. Coracoids similar in size to that of females of Tringa
flavipes, but slightly more slender. As all coracoids are broken or
abraded to varying degrees, assignment to genus is unwarranted.
Genus Philomachus Merrem, 1804
?Philomachus sp.
Material. Bone Valley Mining District, Ft. Green Mine; UF 60062,
proximal end left humerus.
Remarks. After an extended survey of scolpacid genera, I have found
this humerus (Figure 4.9) from Ft. Green to be most similar to that of
males of Philomachus pugnax. Because Philomachus is only a casual


107
Olson erroneously reported PB 7749 as a distal end rather than a proximal
end.
Nothing prevents the two specimens listed above from representing
the same species. Despite the apparent difference in size, the Bone
Valley specimens (see Table 4.15) and USNM 181027 from Lee Creek could
easily represent the same species of Eudocimus. The geologic ages are
similar, both fall within the size variation of a single species (see
Figure 4.3, where both specimens fall within the size range of E. albus),
and both USNM 181027 and UF 60040 have the proximal border of the
posterior articular surface extending farther proximally on the external
side. The distribution of this last character in some specimens of
Plegadis species cannot be evaluated with the skeletal material presently
available.
Genus Plegadis Kaup, 1829
Plegadis cf. £. pharangites (A. H. Miller and Bowman, 1956)
Material. Love Bone Bed local fauna; UF 25870, distal end right
tibiotarsus.
Description. Tibiotarsus similar in size to a very small Plegadis
chihi or P. ridgwayi. Distinguished from these species by having a more
distinct tuberculum on the distal end, a more gracile shaft, lateral
condyle merging with the shaft less abruptly, and the surface cranial to
the tendinal bridge more excavated.
Remarks. The material from the Love Bone Bed is not directly
comparable to that of Plegadis pharangites. Skeletal elements of both P,
pharangites and P. cf. P. pharangites are approximately 10 to 12 percent
smaller than P. mexicana (=P. chihi). While I doubt that this material
from the late Clarendonian of Florida is conspecific with that from the


50
McGehee Farm local fauna; UF 9488, shaft and distal end of right
humerus; UF 12468, right coracoid.
Description. Humerus (UF 9488) poorly preserved; larger and much
more robust than both species of Tachybaptus (dominicus and ruficollis).
Morphology of distal end similar to that of Rollandia rolland, but shaft
more robust. Measurements given in Table 4.2.
Coracoids from the Love Bone Bed are extremely worn and are here
assigned strictly on the basis of their size being near Rollandia (larger
than all species of Tachybaptus and smaller than the smallest males of
Podilymbus podiceps). Shaft similar to, or slightly more robust than,
that of Rollandia rolland. Proximal edge of ventral sternal articulation
flared (as in Rollandia rolland, less so in Tachybaptus). Coracoid from
McGehee Farm local fauna (UF 12488) similar to those from the Love Bone
Bed, except by having a proportionally longer shaft. Measurements are
given in Table 4.2.
Overall length of tarsometatarsus similar to Rollandia rolland
chilensis. Distinguished from Podilymbus by lacking the cranial
expansion of the proximal end of the tarsometatarsus and by having
trochlea II placed lower on the shaft. Distinguished from Podiceps
species by having the shaft less laterally compressed. Tarsometatarsus
differs from that of Rollandia rolland chilensis by having a more deeply
excavated lateral parahypotarsal fossa, a hypotarsus with a smaller
transverse width, a more distinct ridge extending distally from the
hypotarsus, and trochlea II slightly more narrow. Measurements given in
Table 4.3.
Remarks. This species appears to be slightly more robust than the
modern Rollandia rolland chilensis. The lack of a series of modern sexed


139
Remarks. Skeletal elements about the size of those of the living
males of Anser albifrons. Material badly waterworn, leaving no
diagnostic characters to permit assignment to genus.
Anserinae, Genus indet. sp. C. (or B.?)
Material. Love Bone Bed local fauna; UF 25750, proximal end right
carpometacarpus; UF 25763, distal end right carpometacarpus.
Bone Valley Mining District, near Brewster; PB 17^, proximal end
left carpometacarpus.
Remarks. The above skeletal elements are approximately the size of
those of Anser rossii. These elements may represent Anserinae, genus and
species indet. sp B. (above), depending upon how much size variation is
allowed within a single fossil species.
Anserinae, Genus indet. sp. D
Material. Bone Valley Mining District, specific locality unknown;
UF 61598, left coracoid.
Remarks. Coracoid much smaller than that of Anser rossii (or
species B or C above). Furcular facet deeply undercut as is typical of
the tribe Anserini (Woolfenden, 196l).
Subfamily Anatinae (Vigors, 1825)
Remarks. A large amount of material of anatines exists from nearly
all the localities included in this dissertation, but it is not of
sufficent diagnostic character to allow further identification. Rather
them arbitrarily assign specimens to size categories which would not
reflect the true species composition, I prefer to leave this material
undesignated pending further investigations of the anatid material from


244
Williams, K., D. Nicol, and A. Randazzo. 1977 The geology of the
western part of Alachua County, Florida. Bureau of Geology,
Division of Natural Resources, Florida Department of Natural
Resources, Report of Investigations No. 85:1-98
Wolff, R. G. 1973. Hydrodynamic sorting and ecology of a Pleistocene
mammalian assemblage from California (U. S.A.). Paleogeography,
Paleoclimatology, and Paleoecology, 13:91-101.
Wolff, R. G. 1975 Sampling and sample size in ecological analyses of
fossil mammals. Paleobiology, 1:195-204.
Wolff, R. G. 1978. Function and phylogenetic significance of cranial
anatomy of an early bear (indarctos), from Pliocene sediments of
Florida. Carnivore, 1:1-12.
Wood, D. S. 1979 Phenetic relationships within the family Gruidae.
Wilson Bulletin, 91:384-399
Wood, D. S. 1983. Phenetic relationships within the Ciconiidae (Aves).
Annals of the Carnegie Museum, 52:79-112.
Wood, D. S. 1984. Concordance between classifications of the Ciconiidae
based on behavioral and morphological data. Journal Ornithologie,
125:25-37
Wood, H., R. Chaney, J. Clark, E. Colbert, G. Jepson, J. Reeside, and C.
Stock. 1941. Nomenclature and correlation of the North American
continental Tertiary. Bulletin of the Geological Society of
America, 52:1-48.
Woolfenden, G. E. 1959 A Pleistocene avifauna from Rock Spring,
Florida. Wilson Bulletin, 71:183-187
Woolfenden, G. E. 1961. Postcranial osteology of the waterfowl.
Bulletin of the Florida State Museum, Biological Sciences, 6:1-129.
Wright, D. B. and S. D. Webb. 1984. Primitive Mylohyus (Artiodactyla:
Tayassuidae) from the late Hemphillian Bone Valley of Florida.
Journal of Vertebrate Paleontology, 3:152-159
Wyles, J. S., J. G. Kunkel, A. C. Wilson. 1983. Birds, behavior, and
anatomical evolution. Proceedings of the National Academy of
Sciences, U.S.A. 80:4394-4397
Zusi, R. L. and R. W. Storer. 1969 Osteology and myology of the head
and neck of the Pied-Billed Grebe (Podilymbus). Miscellaneous
Publications, Museum of Zoology, University of Michigan, 139:1-49
Zusi, R. L., D. S. Wood, and M. A. Jenkinson. 1982. Remarks on a world
wide inventory of avian anatomical specimens. Auk, 99:740-757.


65
Order Pelecaniformes Sharpe,1891
Family Phalacrocoracidae (Bonaparte, 1853)
Genus Phalacrocorax Brisson, 1760
Remarks. The following morphological descriptions are based on the
comparisons of a sample of 5 males and 5 females each of Phalacrocorax
auritus auritus and P. auritus floridanus, and' all available fossil
material.
Phalacrocorax sp. A.
Material. Love Bone Bed local fauna; UF 25735, UF 29661, distal
ends left humeri; UF 29662, distal end right ulna; UF 25877, distal end
left tibiotarsus; UF 25861, distal end right tarsometatarsus (badly
worn, tentatively referred); UF 25933, distal end left tarsometatarsus.
McGehee Farm local fauna; UF II569, complete left coracoid; UF
31779, sternal end left coracoid; UF 12351, distal end right humerus; UF
Ul07, proximal end right ulna; UF 9^92, proximal end right ulna
(questionably referred); UF 31778, proximal end right carpometacarpus; UF
11105, distal end left carpometacarpus; UF 297^6, complete left
tarsometatarsus; UF 31777, proximal end right tarsometatarsus. PB 796U,
proximal end left carpometacarpus.
Haile XIXA; UF 2977^, proximal end left humerus; UF 473^0, proximal
end right carpometacarpus.
Description. Coracoids from McGehee Farm differ from those of both
subspecies of Phalacrocorax auritus (auritus and floridanus) examined,
and from P. wetmorei by having a more elliptical facies articularis
clavicularis, a more robust shaft in relation to the length of coracoid,
the brachial tuberosity more undercut, the impression for the attachment


116
Order Accipitriformes (Vieillott, l8l6) (Auct.)
Family Vulturidae (illiger, l8ll)
Remarks. Vulture tarsometatarsi and tibiotarsi, the only elements
considered here, are characterized by Cracraft and Rich (1972).
Genus Pliogyps Tordoff, 1959
Emended Generic Diagnosis. The tarsometatarsus of Pliogyps differs
from that of other living and fossil genera of vultures in having a
proportionally large trochlea for digit III, the proximal articular
surface wide and deep in comparison to the length of the bone, a
generally columnar form, with symmetrical lateral and medial flaring both
proximally and distally; shaft wide in comparison to length of bone;
hypotarsus merging distally with shaft by means of a broad, rounded ridge
(as in Vultur, Breagyps, Gymnogyps, and Geranogyps; narrow in Coragyps,
Cathartes, and Sarcoramphus). This last character may be strictly size
dependent and if so, not of value as a generic character.
Remarks. Two other proposed generic characters (Tordoff,- 1959)
appear to be variable within a species, and are of no generic value.
They are the shaft less deeply and extensively excavated anteriorly and
groove of trochlea for digit III ending anterioproximally in a shallow,
but distinct pit.
Pliogyps species
Referred Material. Love Bone Bed local fauna; UF 25719, fragment of
shaft of left humerus; UF 25886, distal end left tibiotarsus; UF 25952,
complete right tarsometatarsus missing a small portion of hypotarsus.
Diagnosis. Tarsometatarsus relatively more robust than any living
or fossil genus of vulturid except Pliogyps. Tarsometatarsus
distinguished from Pliogyps fisheri Tordoff 1959 by smaller size, by
having a narrow ridge extending from the hypotarsus farther down shaft


206
supposed relationships of fossil birds suggested by previous authors and
to correct erroneous locality information. However, the value of such
investigations on fossil birds include:
(1) Demonstration of general trends in avian faunal dynamics in the
Neogene of North America. Major trends should still be evident,
even though they are based on incomplete or partially correct data.
(2) Taxa which do not parallel the general trend may be selected for
further investigations.
(3) Specific geographic areas and/or time intervals can be featured in
future work.
(U) Specific groups can be easily isolated, such as those of interest
from a zoogeographical standpoint.
Formulae used to calculate the parameters appearing in these table
are presented and defined in the methods section, and abbreviations are
given in Table 6.1. The separation of taxa into marine and non-marine
groups is based on the habitats used by the majority of living congeners.
The Local Faunas
The Neogene localities that have produced avian specimens are not
uniformly distributed in geologic time. Of 133 local faunas surveyed,
3.87 (5) are late Arikareean, 8.3 1 (ll) are Heraingfordian, 13.5 % (18)
are Barstovian, 22.6 % (30) are Clarendonian, 30.1 % (4o) are
HemphiIlian, and 21.8% (29) are Blancan in age. Or stated slightly
differently, 'J'h.hlo (99/130) of the localities examined are from the last
4l.O % (Clarendonian to Blancan; 8.7 MA / 21.2 MA) of the Neogene. If
only the published localities are used, instead of the total number of
localities, the percentages are roughly comparable, but the absolute
values are much lower (see Table 6.2).


33
Local Faunas
Love Bone Bed
The Love Bone Bed is located near the town of Archer, Alachua
County, along State Road 24l, in the NW 1/4, SW 1/4, NW 1/4, Sec. 9, T.
11 S., R. 18 E., Archer Quadrangle, U. S. Geologic Survey 7.5 minute
series topographical map, 1969. Excavation and collection of fossil
vertebrates by the Florida State Museum took place from its discovery in
1974 until the quarry was closed during the summer of 1981. This local
fauna originated from the Alachua Formation (Williams et al., 1977).
The Love Bone Bed is considered latest Clarendonian in age (Webb et
al., 198l) Studies published to date on the Love Bone Bed and its
vertebrate fauna include a general overview of the geology and
paleontology of the locality, including a preliminary faunal list (Webb
et al., 1981); studies on the turtles Pseudemys caelata, and Deirochelys
carri (Jackson, 1976, 1978); a description of the rodent Mylagaulus
elassos Baskin (1980); description of the carnivores Barbourofelis lovei
and Nimravides galiani Baskin (1981); descriptions of the procyonids
Arctonasua floridana and Paranasua biradica Baskin (1982); the
description of the ruminant Pedimeryx hamiltoni Webb (1983); and a
population study on the three-toed horse Neohipparion cf. N. leptode
(Hulbert, 1982). Studies on fossil birds include papers on the fossil
herons (Becker, 1985a), a description of a new species of osprey (Becker,
1985b), and one on the fossil anhinga (Becker, ms.). Additional studies
are in progress.


118
Humerus fragment tentatively referred.
Remarks. Table L.l8 lists indices for flexion of the intertarsal
joint (power-arm ratio of Jollie, 1976-1977; [(FLEXOR * LENGTH) X 100)
and for robustness of shaft [W-SHAFT - LENGTH) X 100]. These indices
show that Pliogyps sp. from the Love Bone Bed has an average flexion
ratio but a very broad tarsometatarsus. If this is considered with the
distinct, excavated muscle attachments discussed above, it is suggestive
of a powerful pelvic limb, more so than in most vulturids, reminiscent of
some accipitrids. Certainly the interpretation of this is very tentative
(see Fisher, 19^5; Becker, 1985b), but possibly Pliogyps sp. from the
Love Bone Bed was more rapacious than other living or fossil vultures, as
rapacious birds tend to have a higher flexor ratio than do non-rapacious
birds of equal size. As additional fossil material of this species
becomes available, this suggestion should be examined further.
Campbell and Tonni (1983) develop further ideas of Prange et al.
(1979) on the correlation between the cross-sectional area of the
tibiotarsus of a given species and its live weight. They empirically
determined the following regression
log Y = 2.51* log X 0.19906
where Y is the live body weight (gms) and X is the least shaft
circumference of the tibiotarsus (mm). The correlation coefficient for
this relationship is O.986, showing that the predictions of the live
weight should be very accurate. The least shaft circumference of
Pliogyps species (UF 25886) from the Love Bone Bed is 32 mm yielding a
predicted weight of 5*2 kg. Sarcoramphus papa, which has a
tarsometatarsus approximately as long as this species, weights between
3.0 and 3.75 kg (5 individuals, Brown and Amadon, 1968). This supports


Figure 4.5. Ratio diagram (after Simpson et al. i960) of measurements of the tarsometatarsi of the
following species of vultures: (A) Cathartes aura, (B) Coragyps atratus atratus, (C) Pliogyps sp. from
the Love Bone Bed 1. f., (D) Pliogyps fisheri, (E) Sarcoramphus papa, (f1 Gymnogyps californianus, (G)
Geranogyps reliquus, (H) Gymnogyps howardae, (i) Vultur gryphus, and (J) Breagyps clarki. Measurements,
defined in the methods section, are abbreviated as follows: (l) LENGTH, (2; W-PROX, (3) D-PROX-L, (4)
FLEXOR, (5) W-SHAFT, (6) D-SHAFT, (7) W-TRIII, (8) D-TRIII, (9) W-DIST.
1


240
Peters, J. L. 1951 Check-list of Birds of the World. Volume VII.
Harvard University Press, Cambridge, Massachusetts. 318 pp.
Prange, H. D., J. F. Anderson, and H. Rahn. 1979* Scaling of skeletal
mass to body mass in birds and mammals. American Naturalist,
113:103-122.
Ray, C. E. 1957. A list, bibliography, and index of the fossil
vertebrates of Florida. Florida Geological Survey Special
Publication No. 3. 175 pp.
Rea, A. M. 1983 Cathartid affinities: a brief overview, pp. 26-53.
In S. R. Wilbur and J. A. Jackson (eds.). Vultur Biology and
Management. University of California Press, Berkeley.
Recher, H. F. and J. A. Recher. 1980. Why are there different kinds of
herons? Transactions of the Linnaean Society of New York, 9:135-
158.
Rich, P. V. 1972. A fossil avifauna from the Upper Miocene Beglia
Formation of Tunisia- Notes Serv. Geol. Tunisie 35 Trav. Geol.
Tunisienne No. 5 Formation Beglia-Fasc. 1. p. 29-66. [not seen].
Rich, P. V. 1976. The history of birds on the island continent
Australia. Proceedings l6th International Ornithological Congress,
Canberra, Australia, pp. 53-65.
Rich, P. V. 1980. 'New World Vultures' with Old World affinities?
Contributions to Vertebrate Evolution, 5:viii + ll6.
Rich, P. V. and C. A. Walker. 1983. A new genus of Miocene flamingo
from East Africa. Ostrich, 54:95-104.
Riggs, S. R. 1984. Paleoceanographic model of the Neogene phosphorite
deposition, U. S. Alantic continental margin. Science, 223:123-131.
Ripley, S. D. 1977. Rails of the World. A Monograph of the Family
Rallidae. Godine, Boston, Massachusetts. 406 pp.
Ritchie, T. 1980. Two mid-Pleistocene avifaunas from Coleman, Florida.
Bulletin of the Florida State Museum, Biological Sciences, 26:1-36.
Schultz, C., M. Schultz, and L. Martin. 1970. A new tribe of saber-
toothed cats (Barbourofelini) from the Pliocene of North America.
Bulletin University of the Nebraska State Museum, 9:1-31.
Sel lards, E. H. 1916. Fossil vertebrates from Florida: A new Miocene
fauna; new Pliocene species; the Pleistocene fauna. Florida
Geological Survey 8th Annual Report, Tallahassee, pp. 77-102, pis.
10-14.
Shipman, P. 1981. Life history of a fossil. Harvard University Press.
Cambridge, Massachusetts. 222 pp.


91
ibis, and is only known from the holotype. The tarsometatarsus is
, proportionally narrower than in other members of this genus.
Egretta sp. indet.
Material. Love Bone Bed local fauna; UF 25759, proximal end of left
carpometacarpus, UF 26082, distal end right ulna. Bone Valley Phosphate
Mining District, Payne Creek Mine; PB 7925, coracoid.
Remarks. These specimens fall within the size range of the living
E. rufescens. Because of the large time interval (4.5 MA) between the
Love Bone Bed and the Bone Valley, it is unlikely that these elements
represent the same species.
Genus Ardeola Boie, 1822
Ardeola sp. indet.
Material. Love Bone Bed local fauna; UF 25940, distal one-third
left tarsometatarsus.
Remarks. Small, similar in size to Ardeola striata. Taxonomic
assignment based entirely on size.
Subfamily Nycticoracinae Payne and Risley, 1976
Genus Nycticorax T. Forster, l8l7
Nycticorax fidens Brodkorb, 1963
Material. McGehee Farm local fauna; UF 3285, complete left femur.
Remarks. See Brodkorb (1963a) for description and remarks.
Remarks on the Family Ardeidae.
Table 4.1 summarizes the distribution of the fossil herons from
Florida. The Love Bone Bed local fauna has produced three heronsa
small Ardeola, a large Egretta, and a very large Ardea. It is most


Table 4.10. Measurements of the tibiotarsi of the cormorants Phalacrocorax auritus aurltus (N = 14, 7
males, 7 females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), Phalacrocorax vetmorei,
from the Bone Valley Mining District, and Phalacrocorax species from the Love Bone Bed. Data are mean
standard deviation
, (N), and range.
Abbreviations are
defined in the methods
section.
Measurements
P. a. auritus
P. a. floridanus
P. vetmorei
Phalacrocorax sp.
Tibiotarsus
FIBULAR
43.18 + 1.99
38.5 45.8
41.26 + 1.94
36.7 43.9
37.7 (1)

W-SHAFT
6.92 + 0.22
6.6 7.4
6.63 + 0.28
6.0 7.1
7.02 + 0.25 (5)
6.6- 7.2

D-SHAFT
5.41 + 0.46
4.8 6.5
4.88 + 0.34
4.3 5.4
5.14 + 0.09 (5)
5.0 5.2

W-PROX-M
11.70 + 0.67
10.7 12.7
10.84 + 0.8I
8.9 12.2
11.70 + 0.80 (5)
10.5 12.7

D-PROX
17.09 + 0.75
16.2 18.4
15.87 + 0.79
i4.1 17.6
17.22 + 0.78 (5)
16.5 18.3

W-PROX-L
11.80 + 0.58
10.6 12.7
11.09 + 0.56
10.1 12.2
10.14 + 0.51 (5)
9.3 10.7

i
00
-o


4
try to note the systematic problems in each group, briefly review its
fossil record, and describe the fossil material from Florida. This
material should be reexamined as the state of systematics of these groups
improves and as additional Recent and fossil material becomes available.
Previous Work
The earliest published report of fossil birds from Florida is
Sellard's (1916) description of a supposed jabir (Jabir? weillsi
=Ciconia maltha) from Vero, closely followed by Shufeldt's (1917a, 1917b)
study of this local fauna. Wetmore (1931) reviewed the Pleistocene
avifauna of Florida and was also the first to report (19^3) on the
Tertiary birds from Thomas Farm and the Bone Valley Mining District.
Numerous studies have appeared since then, primarily by Brodkorb or his
students. A few other faunal and systematic studies have also included
avian material from Florida.
Table 1.1 lists many of the fossil localities in Florida which have
a notable record of fossil birds. Reference to Brodkorb's Catalogue of
Fossil Birds (1963-1978), where the avifaunas from many Florida
Pleistocene localities were first reported, is omitted to conserve space.


CHAPTER VI
BIOCHRONOLOGY AND FAUNAL DYNAMICS
Introduction
The following analysis of the biochronology and faunal dynamics of
the Neogene (23 nybp to 1.8 mybp) avifauna of North America takes much of
its information from an accompanying project (Becker, ms.), which updates
and reviews the Neogene records of the North American avifauna. This
fossil record is still very incomplete in many areas, and its
intrepretation will doubtlessly change as new material becomes available
and previously described material is restudied. I should also note that
the current record of fossil birds suffers from decades of a typological
systematic approach, with many species, and even genera, being based more
on geography and on geologic age than on differences in morphology.
Faunal Dynamics
Remarks. Among the assumptions necessary to calculate faunal
dynamic parameters are the following:
(1) All taxa are correctly identified, are taxonomically valid, and
are correctly placed systematically.
(2) The stratigraphic context of the fossil specimens is known and the
specimen is correctly placed in a local fauna.
(3) The local fauna is correctly placed in geologic time.
(U) Taxa are correctly divided into marine and non-marine groups.
In one or another instance, each of these assumptions is surely
violated. Much work still remains to be done to verify some of the
205


Remarks. Brodkorb (1955a) states that this coracoid agrees with
that of Bucephala clangula in general appearance and details of the
brachial tuberosity, but differs by being smaller and in details of the
procoracoid and triosseal canal.
Anatids are fairly rare members of the Bone Valley avifauna. As
additional material becomes available, this poorly known species should
be re-examined.
Tribe Oxyurini J. C. Phillips, 1926
Genus Oxyura Bonaparte, 1828
Oxyura cf. £. dominicus
Material. Bone Valley Mining District, Ft. Green Mine; UF 61950,
proximal ends left humerus.
Remarks. Size of a small Oxyura dominicus. Humeral head slightly
more undercut by the external head of the m. triceps in the fossi
specimen than in the series of recent skeletons of 0. dominicus. All
other characters within the range of variation of 0. dominicus.
Remarks on the Family Anatidae.
Howard (1964, 1973) has reviewed the extensive fossil record of the
Anatidae, so there is little need once again to review all taxa coverd by
Howard. New Miocene fossil species described since 1973 include
Cygnopterus alphonsi from the Aquitanian of St.-Gferand-le-Puy, France by
Cheneval (1984); and a Blancan goose from the Broadwater local fauna,
Anser thompsoni by Martin and Mengel (1980). Both of these species are
known from several elements.
Four new species have also been described from the late Pleistocene.
Anas schneideri was described from a single carpometacarpus from the


67
than in P. auritus. Posterior opening of the lateral proximal vascular
foramen is located lateral to the ridge extending down from the lateral
calcaneal ridge. This ridge not extending as far down the shaft as in P.
auritus.
Remarks. If the carpometacarpus above is correctly assigned to the
same species as is represented by the other skeletal elements, then this
cormorant is quite different in proportions than Phalacrocorax auritus
and related species such as Phalacrocorax wetmorei.
Phalacrocorax wetmorei Brodkorb, 1955
Material. This material is very well represented in the Bone Valley
Fauna. Only Florida State Museum and Florida State Geological specimens
are included in the following referred material section. Material
accessioned into the Florida State Museum collections after 26 April 1984
has not been included in the list of referred specimens; material
accessioned after 5 March 1984 has not been included in the tables of
measurements. Additional material from Bone Valley (type material) is
listed in Brodkorb (1955a).
Manatee County Dam Site.UF 11916, distal end right humerus.
SR-64 local fauna.UF 67805, complete left coracoid; UF 64l43,
humeral end left coracoid; UF 64l44, humeral end right coracoid; UF
64l46, humeral end right scapula; UF 64l45, partial sternum; UF 64l47,
proximal end right tibiotarsus; UF 64l48, UF 64l49, distal ends left
tarsometatarsi.
Bone Valley Mining District, Brewster Mine.UF 61987, humeral end
right coracoid; UF 61988, distal end right tarsometatarsus; UF 65691,
proximal end left ulna.


66
of the coraco-brachialis more distinct, and in medial view, the shaft
more rotated ventrally.
Distal end of the humeri of the fossil species is generally smaller
than that of females of P. a. floridanus, and the shaft more slender
(much smaller than P. a. auritus). Other characters are within the range
of variation of £. auritus. Differs from the humeri of P. wetmorei by
being smaller, having a more shallow fossa brachialis of a different
angle and a much wider attachment of the anterior articular ligament
(=tuberculum supracondylare ventrale).
The two ulnae that are sufficiently perserved to make comparisons
(UF 29662, UF 4107) appear small, about the size of small females of P.
a. floridanus. Characters are within the range of variation of this
species.
Carpometacarpus larger than that of the largest male P. a. auritus.
Process of metacarpal I more nearly square than in that of P. auritus.
Shaft of metacarpal II more robust and angular; anterior carpal facet not
extending up the carpal trochlea as in P. auritus. Pollical facet with a
small papilla .
Tibiotarsus indistinguishable from that of small females of P. a.
floridanus.
Tarsometatarsus description based on UF 29746 (UF 25933 broken and
badly worn; UF 31777 missing distal half, but both agree with UF 29746 in
all discernable characters). Tarsometatarsus short, about equal to that
of females of P. a. floridanus. Transverse width of shaft very narrow.
Lateral face of shaft much more flattened than P. auritus, causing the
posterior intermuscular line to be on the lateral edge of the shaft.
Proximal end narrow, lateral calcaneal ridge more narrow and elongate


185
Family Scolopacidae Vigors, 1825
Remarks. Many of the specimens discussed below have been
arbitrarily assigned to the genus Calidris because of their similarity in
size and overall morphology to this genus. I have been unable to find
generic characters (i. e. non-size related) on the elements here
preserved which will confidently separate Calidris from other genera of
scolopacids. Possibly with study of additional series of all scolopacid
genera, characters could be isolated. But for the present, the following
generic assignments should be regarded as tentative.
Brodkorb (1955a, 1963a, 1967) described four species of scolopacids
from the late Miocene and early Pliocene of FloridaCalidris pacis,
Erolia penepusilla, and Limosa ossivallis from the Bone Valley Mining
District and Ereunetes rayi from the McGehee Farm local fauna. They are
listed below for the sake of completeness, but additional specimens
usually have not been assigned to them, pending further comparisons with
living and fossil species. All specimens are assigned at the species
level on the basis of size. This approach has the effect of
overestimating the diversity of scolopacids now known from in the late
Miocene and early Pliocene of Florida.
Genus Limosa Brisson, 1760
Limosa ossivallis Brodkorb, 1967
Material. Bone Valley Mining District, near Brewster (referred by
Brodkorb, 1955a); PB 526, distal end right tibiotarsus (holotype); PB 527
proximal end right tibiotarsus ("paratype"). Specimens referred by
Becker: Bone Valley Mining District, specific locality unknown; UF
60817, distal end left humerus; UF 61597, left coracoid; UF 6l600,


162
material of balearicine cranes is now under study by Feduccia. As much
of this material consists of complete, articulated skeletons, it seems
unneccessary to further describe this single, partial element. From the
material which I have seen, there are at least two species of
balearicinae cranes in North America at the end of the Hemphillian.
Based on the above specimen, the last occurrence now known of this
subfamily of cranes in North America is from the late Hemphillian Bone
Valley local fauna.
Genus Aramornis Wetmore, 1926
cf. Aramornis, sp. A.
Material. Love Bone Bed local fauna; UF 25949, distal end left
tarsometatarsus, missing trochlea IV.
Description/ Remarks. Tarsometatarsus compares well with the type
of Aramornis longurio (F:AM 6269) in relative proportions and positions
of the trochlae II and III, but differs by having a larger distal foramen
and a greater overall size. See Table 4.25 for measurements. Additional
material is needed to verify this generic assignment.
Remarks on the Family Gruidae.
Johnsgard (1983) considers the following species to be closely
related: Grus .japonensis and G. americana, Grus rubicundus and G.
antigone; Bugeranus carunculatus and B. leucogeranus; Antropoides virgo
and A. paradisea; and Balerica pavonina and B. regulorum. Additionally,
he considers Grus grus, G. monacha, G. canadensis, and G. vipio to form a
loose species cluster. Outside of these obvious groupings, he makes no
attempt to place them within a phylogeny. With the exception of Wood's
(19T9) phenetic study, little work has been done concerning the


Table 4.l6. Measurements of tibiotarsi, coracoids, and tarsoraetatarsi of the ibises Plegadis ridgwayi
(N = 2, unsexed), Plegadis chihi (N = 7, 2 males, 2 females, 3 unsexed; number may be less due to incomplete
specimens), Plegadis falcinellis (N =10, 7 males, 3 females), Plegadis species (tibiotarsus) and
Threskiornithinae, genus and species indet. (coracoid) from the Love Bone Bed local fauna. Data are
mean _+ standard deviation and range. Abbreviations are defined in the methods section. (*) specimen
damaged.
Measurements
P. ridgwayi
P. chihi
P. falcinellis
Plegadis sp.
Tibiotarsus
W-SHAFT
4.0;
4.3
4.33
+
0.47
4.52
+
0.37
3.7
3.8
-
4.9
3.9
-
5.0
D-SHAFT
3. 4;
3.6
3.80
+
0.35
4.05
+
0.34
3.2
3.5
-
4.3
3.6
-
4.7
W-DIST-CR
7.5;
7.9
8.45
+
0.66
8.63
+
O.56
7.2
7.6
-
9.1
7.8
-
9.7
W-DIST-CD
5.7;
5.5
5.93
+
0.32
6.28
+
0.43
*5.1
5-7
-
6.4
5.5
-
7.0
D-MCON
8.4;
8.4
9.20
+
0.69
9.93
+
O.58
*7-9
8.7
-
10.2
8.9
-
10.8
D-LCON
8.1;
8.3
9.18
+
0.75
9.56
+
0.71
*7.1
8.3
-
10.1
8.5
-
10.5
D-ICON
5.6;
5-5
6.34
+
0.54
6.39
+
0.43
4.9
5.8
-
'7.0
5.8
-
7.0
114


213
skeleton is likely to be much more conservative as a systematic character
than tooth morphology, a slower turnover rate in birds would not be
unexpected.
Biochronology
The following preliminary list of taxa represents the first attempt
at using fossil birds as biochronologically useful taxa. This list is
based on the current record of avian genera from the Neogene of North
America, with taxa of questionable validity being omitted. The following
list will doubtlessly change and increase as our knowledge of avian
evolution in the Neogene becomes more complete.
LATE ARIKAREEAN. First Appearance: Morus.
Last Appearance: Plotopteridae.
HEMINGFORDIAN. First Appearance: Puffinus, Anhinga, Sula,
Dendrocygnini, Anatini, Palaeoborus, Neophrontops, Falco, Rallidae,
Megapaloelodus, Burhinus, Strigidae. Last Appearance: None identified.
BARSTOVIAN. First Appearance: Gavia, Podicipedidae, Diomedea,
Fulmarus, Phalacrocorax, Microsula, Qsteodontornis, Ardea, Anserini,
Mergini, Vulturidae, Pandion, Rallus, Laridae, Stercorariidae, Alcinae,
Corvidae. Last Appearance: None identified.
CLARENDONIAN. First Appearance: Rollandia, Tachybaptus, Oceanodroma,
Miosula, Plegadis, Egretta, Ardeola, Mycteria, Ciconia, Pliogyps, Flica,
Grus, Limosa, Phoenicopterus, Jacana, Alca, Cepphus, Cerorhinca, Uria,
Aethia, Praemancalla, Tytonidae. Last Appearance: Qsteodontornis,
Microsula


11
10. L-GLEN.Length of glenoid facet from the most cranial portion
of glenoid through the most caudal point of scapular facet.
11. IDA-FNS.Length from internal distal angle through the most
sternal' edge of coracoidal fenestra (Foramen nervus
12.ANG-HEAD.Angle formed between axis of the head, as seen in
proximal view, and the plane parallel to the dorsal surface
(Facies dorsalis).
Humerus
1. LENGTH.Greatest length from the head of the humerus (Caput
humeri) through the midpoint of the lateral condyle (Condylus
ventralis).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. W-PROX.Transverse width of proximal end from the external
tuberosity (Tuberculum dorsale) to the most ventral face of the
bicipital crest (Crista bicipitalis).
5. D-PROX.Depth of proximal end, from the bicipital surface
(Facies bicipitalis) to the internal tuberosity (Tuberculum
ventrale), measured at right angles to the long axis of the
shaft.
5a. D-HEAD.Depth of head, measured parallel to the axis of the
head.
6. L-DELTOID.Length of deltoid crest (Crista pectoralis),
measured from the external tuberosity to the most distal
extension of the deltoid crest


CHAPTER II
METHODS
Measurements
Measurements made in this study are listed below and are illustrated
in Figures 2.1 2.k. They have been selected from the literature
dealing with the osteology and identification of both Recent and fossil
species of birds. Previous authors have dealt with only one specific
group (Steadman, 1980; Howard, 1932b; Ono, 1980) or with Recent birds
commonly found in archeological sites (von den Driesch, 1979; Gilbert, et
al., 1981). The most applicable anatomical studies on fossil birds are
by Ballmann (1969a, 1969b). I have modified measurements from the above
studies to make them applicable to the range of morphologies encountered
in the fossil birds of this study. Measurements for the less diagnostic
elements and for the cranium were not included. Measurements presented
in the systematic section of this dissertation depend on the fossil
material available and the morphology of the species considered.
Anatomical terminology follows Baumel et al. (1979) and Howard
(1929, 1980). Many Latin terms have been anglicized for ease of
communication, but the original Latin is given parenthetically when the
term is first used (below). Terms for soft tissue anatony come from
Feduccia (1975) and Van den Berge (1975 )
9


169
Family Rallidae Vigors, 1825
Family Characters. See references in family remarks section; also
Gilbert, et al. (1981).
Material. Love Bone Bed local fauna; UF 26015, UF 29714, UF 29709,
UF 25787, coracoid fragments.
Remarks. The above material appears to be rallid, but is not
identifiable to the generic level.
Genus Rallus Linnaeus, 1758
Rallus sp. A
Material. Love Bone Bed local fauna; UF 25936, distal end left
tarsometatarsus.
Remarks. Near Crex crex in size. Decidely larger than Rallus
limicola, but smaller than Rallus longirostris. I have not been able to
find qualitative characters in the distal ends of the tarsometatarsus to
distinguish between Rallus and Crex and have therefore arbitrally
assigned these specimens to Rallus.
Rallus sp. B
Material. Bone Valley Mining District, Palmetto Mine; UF 21060,
humeral end right coracoid. Payne Creek Mine; UF 21204, proximal end
left tarsometatarsus.
Remarks. Coracoid slightly smaller than that of females of Rallus
longirostris. Agrees with Rallus by having a small procoracoid process.
Tarsometatarsus near Crex crex in size, or intermediate between Rallus
limicola and Rallus longirostris. These two specimens could possibly
represent two different species, but owing to the slight difference in size
and the lack of additional specimens, I have listed them together as


199
Love Bone Bed and Bone Valley. Although these two localities have large
samples of fossil birds, the techniques used to collect the fossil
vertebrates prevent a quantitative paleoecological analysis. Hence I am
limited to qualitative statements about the paleoecology of each of the
local faunas considered in this investigation. These statements are
based primarily on the birds present, the habitats used by their Recent
congeners (Blake, 1977; Palmer, 1962, 1975, 1976; Terres, 1980), and on a
general knowledge of the geology of the locality and its vertebrate
fauna. In general, the paleoecological reconstructions based on fossil
birds are similar to those based on other fossil groups.
Local Faunas
Love Bone Bed local fauna. The avifauna of the Love Bone Bed is
richly aquatic, with but a minor influence of more terrestrial species.
Taxa presently identified include 2 species of grebe, a species of
cormorant and one of anhinga, 3 species of heron, 2 species of~stork, 2
ibses, one vulture, one osprey, 2 accipitrids, 4 species of geese, 4
species of duck, a turkey, 3 cranes, 3 rails, 2 flamingos, a jacana, 7
species of shorebirds, a barn owl, and 2 species of passerine.
The more abundant species of those listed above include the grebe,
Tachybaptus sp.; a tree-duck, Dendrocygna sp.; a tiny species of teal,
Anas sp. A.; a larger duck, Anas near A. acuta; a crane, Grus sp. B.; two
species of rail, one a species of Rallus cf. sp. C., and the other an
undescribed genus; 2 flamingos in the genus Phoenicopterus; a jacana,
Jacana farrandi; and a indeterminate genus and species of scolopacid.
These abundant taxa suggest that more than one type of habitat has
been sampled. Modern species of grebe typically occur in freshwater,
from lakes to shallow ponds. Jacanas occur in freshwater marshes to


133
Genus Buteo Lacpde, 1799
Buteo near B. jamaciensis (Gmelin, 1788)
Material Withlacoochee River 4a local fauna; UF 67808, complete
left femur.
Description. Size of Buteo jamaciensis harlani. Contares well with
this subspecies in overall size, position of intermuscular lines.
Differs by having a more narrow patellar sulcus, a slightly more
constricted femoral head, greatest length slightly less, and slightly
more gracile.
Remarks. This specimen has been reported in Becker (1985a).
Genus Aguila Brisson, 1760
Aguila sp. A
Material. Bone Valley Mining District, Hookers Prarie Mine, UF
57299, distal end left tarsometatarsus with posterior wing of trochlea IV
broken off.
Description. Virtually indistinguishable from the range of
variation of that of the living Aguila chrysaetus in size and morphology.
Differs by having a more laterally compressed distal foramen and a
slightly deeper tendinal groove (= outer extensor groove of Howard,
1980).
Accipitrid, Genus indet., species A.
Material. Love Bone Bed local fauna, UF 29676, proximal end right
carpometacarpus.
Description. Similar in size to a large Haliaeetus 1eucocephalus or
a small Aguila chrysaetus. Carpometacarpus agrees with A. chrysaetus by
having a large process on metacarpal I (small in Haliaeetus) and shape of


57
North America). This view is further strengthened by the absence of
grebes in the Early Tertiary European fossil localities which have
otherwise produced a rich aquatic avifauna (St.-Gerand-le-Puy; Cheneval,
1984; Phosphorites du Quercy; Mourer-Chauvire, 1982). The paucity of
knowledge about the evolution of birds in the early Tertiary of South
America makes it premature to decide between a North or South American
origin, although this did not prevent Storer (1967) from suggesting a
South American origin of the family based solely on the diversity of
living species


142
with heavy bone. The fossil humeri particularly agree with the humeri of
the Anatini by having the impression of the brachialis not well defined
(Aythini with impression well-defined, with a distal medial rim sharp)
and by having the entepicondyle approximately equal in anconal height to
the ectepicondyle (Aythini with entepicondyle distinctly higher). In
three out of five fossil specimens, the pneumatic fossa is open and
contains struts (as in Anatini; in Aythini it is usually closed). Distal
end rotated medially as in Anas. Greatest morphological resemblence of
the humerus, coracoid, carpometacarpus, and tarsometatarsus is to the
genus Anas (Fig. 4.J).
Comparisons were made with the smallest living species of Anas, Anas
hottentata, to illustrate the morphological characters of this extremely
small fossil species from the late Miocene of Florida (see Table 4.20).
The humeri from the Love Bone Bed are smaller, less robust, but with
deltoid crest similar. The coracoids from the Love Bone Bed are smaller,
proportionally more stout, with the medial margin (in ventral view)
slightly inflated. Carpometacarpus from the Love Bone Bed is much
smaller, and more gracile. Tarsometatarsus from McGehee Farm is smaller,
more slender, with the lateral parahypotarsal sulcus more excavated.
Sulcus on anterior surface proximal to canal more defined.
Remarks. The above specimens agree with the genus Anas in all
characters except onewhether the humerus has an open or closed
pneumatic fossa. The two extremes of this character are one of the
defining characters which distinguish the tribes Anatini and Aythini
(Woolfenden, 1961: 12). This site is the entrance of the air sac system
into the shaft of the humerus and is highly adaptive as it allows for the
additional regulation of bouyancy in diving birds. It almost certainly


Table 4.19. Measurements of the coracoids and carpometacarpi of Dendrocygna viduata (N = 8, 4 males, 4
females), Dendrocygna arbrea (N = 4, 1 male, 3 females, maximun), Dendrocygna bicolor (N = 9 6 males, 3
females, maximun), Dendrocygna autumnalis (N = 8, 4 males, 4 females), and Dendrocygna species from the Love
Bone Bed local fauna. Data are mean +_ standard deviation (number) and range. Abbreviations are defined in
the methods section.
Measurement
D. viduata
D. arbrea
D. bicolor
D. autumnalis
Dendrocygna sp.
Coracoid
HEAD-IDA
40.79 + 1.74
38.6 44.3
44.98 + 1.97
42.9 46.8
42.14 + 3.97
37.8 52.1
42.46 + 1.6l
4o.l 44.8
44.93 + 1.97 (6)
42.1 47.9
HEAD-CS
14.75 + 0.88
l4.l 16.8
15.18 + 1.30
13.5 16.6
15.01 + 0.49
14.3 15.8
15.26 + 0.65
14.5 16.3
15.78 + 0.87 (9)
13.8 16.8
D-HEAD
3.79 + 0.16
3.5 4.0
4.00 + 0.28
3.8 4.0
3.44 + 0.17
3.2 3.7
3.80 + 0.30
3.4 4.3
4.10 + 0.39 (8)
3.8 5.0
W-SHAFT
3.69 + 0.07
3.6 3.8
4.18 + 0.17
4.0 4.4
4.01 + 0.42
3.3 4.5
4.10 + 0.21
3.8 4.5
4.46 + 0.32 (10)
4.1 5.0
D-SHAFT
3.60 + 0.29
3.2 4.0
3.68 + 0.17
3.5 3.9
3.77 + 0.17
3.6~- 4.4
3.86 + 0.19
3.6- 4.0
4.28 + 0.38 (10)
3.8 5.0
FAC-IDA
14.24 + 0.64
13.4 15.1
16.57 + 0.50
16.1 17.I
14.94 + 0.82
13.6 16.3
15.68 + 0.64
14.6 16.6
16.03 + 0.51 (4)
15.5 16.7
IDA-PP
30.05 + 1.20
28.9 32.7
33.53 + 1.53
32.2 35.0
29.54 + 1.17
27.0 30.7
31.26 + 1.45
29.4 33.4
32.86 + 1.13 (7)
31.8 35.2
i
151


Figure 2.2. Schematic diagrams illustrating measurements of the
radius, ulna, carpometacarpus, and furculum. A. Radius, medial
view. B. Radius, cranial view. C. Ulna, cranial view. D.
Ulna, caudal view. E. Carpometacarpus, ventral view. F.
Carpometacarpus, proximal end view. G. Carpometacarpus, distal
end view. H. Ulna, distal end view. I. Furculum, lateral view.
Figures are not drawn to scale. Measurements are defined in text.


10
Scapula
1. LENGTH.Greatest length from the acromion to the caudal
extremity of the scapula (Extremitas caudalis scapulae).
2. W-NECK.Least width neck of scapula (Collum scapulae).
3. W-PROX.Proximal width from the ventral tip of the glenoid
facet (Facies articularis humeralis) to the dorsal margin of the
scapular head (Caput scapulae).
4. ACR-GLN.Length from tip of acromion through ventral tip of
glenoid facet.
5. D-GLN.Depth of glenoid facet.
Coracoid
1. HEAD-FAC.Length from head (Processus acrocoracoideus) through
external end of sternal facet (Facies articularis sternalis).
2. HEAD-IDA.Length from head through internal distal angle
(Angulus medial is).
3. HEAD-CS.Length from head through scapular facet (Cotyla
scapularis).
4. D-HEAD.Least depth of head.
5. W-SHAFT.Width of midshaft.
6. D-SHAFT.Depth of midshaft.
7. FAC-IDA.Length from external end of sternal facet through
internal distal angle.
8. IDA-PL.Length from internal distal angle to medial most edge
of sternocoracoidal process (Processus lateralis).
9 IDA-PP.Length from internal distal angle to procoracoid
process (Processus procoracoideus).


138
New World tropics. D. autumnalis is widely distributed, highly
discontinuous, with no constant geographical variation (Friedmann, 1947).
Tribe Anserini Vigors, 1825
Genus Branta Scopoli, 1769
?Branta sp. A.
Material. Love Bone Bed local fauna; UF 29751, proximal end right
humerus; UF 29752, distal end right humerus; UF 25797, left coracoid; UF
26005, humeral end right coracoid; UF 25748, UF 25751, proximal ends
right carpometacarpi; UF 25890, UF 25891, distal ends left tibiotarsi; UF
25951, UF 29759, nearly complete left tarsometatarsi.
Remarks. Skeletal elements about the size of those of the Recent
Branta canadensis interior. Assignment to Branta is based on the
following characters: proximal humerus with the attachment of the M.
triceps externus with a distinct border; coracoid with furcular facet not
deeply undercut, with only a few pneumatic foramina present (Woolfenden
1961:49). This assignment is tentative, as similar character states are
approached in specimens of Anser species. Other elements were arbitarily
assigned on the basis of size.
Anserinae, Genus indet. sp. B.
Material. Love Bone Bed local fauna, UF 29753, left carpometacarpus
without metacarpal III; UF 25754, proximal end right carpometacarpus; UF
25758, proximal end left carpometacarpus; UF 29761, right coracoid; UF
25904, distal end right tibiotarsus; UF 25879, distal end left
tibiotarsus; UF 25929, UF 25947, distal ends left tarsometatarsi.


5
Table 1.1. Avian Fossil Localities of Florida.
Locality
Reference (s)
MIOCENE TO EARLIEST PLIOCENE
(Pre-Blancan)
Bone Valley, Polk Co.
Becker, 1985a; Brodkorb, 1953b, 1953c,
1953d, 1953e, 1955a, 1970; Olson, 1981b;
Steadman, 1980; Wetmore, 19^3; this study
Gainesville Creeks,
Alachua Co.
Brodkorb, 1963b
Haile VI, Alachua Co.
Brodkorb, 1963a; this study
Haile XIXA, Alachua Co.
this study
Love Bone Bed, Alachua Co.
Becker, 1985a, 1985b; Webb et al., I98I;
this study
Manatee Co.Dam Site
Manatee Co.
Webb and Tessman, 1968; this study
McGehee Farm, Alachua Co.
Brodkorb, 1963a; Hirschfeld and Webb, 1968;
Olson 1976; this study
Mixson Bone Bed, Levy Co.
this study
Seaboard Airline Railroad,
Leon Co.
Brodkorb, 1963b
SR-64, Manatee Co.
this study
Thomas Farm, Gilchrist Co.
Brodkorb, 1954a, 1956a, 1963b; Cracraft,
1971; Olson and Farrand, 1974; Steadman,
1980; Wetmore, 1943, 1958
Withlacoochee River UA,
Marion Co.
Becker, 1985a; this study
LATE PLIOCENE (Blancan)
Haile XVA, Alachua Co.
Campbell, 1976; Steadman, 1980
Santa Fe IB, Gilchrist Co.
Brodkorb, 1963d


Table 4.20continued
Measurement Anas hottentata Anas sp.
Carpometacarpus
LENGTH
32.7; 32.0
27.9
W-CARPAL
3.3; 3.6
2.9
D-PROX
T.8; 7.8
6.7
L-MCI
4.9; 4.8
4.5
D-SHAFT
2.3; 2.3
2.1
W-SHAFT
2.4; 2.6
2.1
D-DIST
2.8; 3.0
2.2
W-DIST
4.3; 4.2
3.5
Tarsometatarsus
LENGTH
27.5; 27.9; 28.1
25.5
W-SHAFT
3.0; 3.1; 2.9
2.8
D-SHAFT
2.4; 2.5; 2.7
2.3
W-PROX
5.4; 5.8; 5.8
5.1


Remarks on the Family Podicepidadae.
The family Podicipedidae includes 11 fossil and 19 living species.
The earliest certain grebe is Podiceps oligocaenus (Shufeldt), based on a
fragmented left femur (missing the proximal end, distal end badly
abraded) from the Arikareean John Day Formation, Oregon. It is
intermediate in size between the living Podiceps grisegena and P.
nigricollis. Although Wetmore (1937) considers it to be correctly
allocated to genus, next to nothing can be said about its relationships.
Podiceps pisanus (Portis) based on the distal end of a right
humerus, is from the Middle Pliocene (=late Miocene?) of Italy. This
species may also be present at the Hemphillian Lee Creek local fauna
(Olson, ms.). The only other late Miocene species of grebe now known is
Pliodytes lanquisti Brodkorb, discussed above.
There are five species of Pliocene (i.e. Blancan) grebes, all from
North America. Podiceps subparvus (L. Miller and Bowman), from the early
Blancan San Diego Formation, California, is based on a distal end of a
femur. It is approximately the same size as that of the living
Podilymbus podiceps and is now known from additional material. Murray
(1967) in his review of Pliocene grebes, described one new genus and four
new species. Pliolymbus baryosteus Murray, from the Fox Canyon local
fauna, Kansas, of Blancan age, is based on the cranial portion of a
sternum. Murray (1967) states that this is a small grebe with a robust
skeleton but does not suggest any possible relationships between this
species and other living or fossil species of grebes. Podiceps discors
Murray, also from Fox Canyon, is based on a left tarsometatarsus. It is
near the size of Podiceps nigricollis. Murray (1967) also tentatively
refers material from the Hagerman local fauna, the San Diego Formation,


THE FOSSIL BIRDS OF THE' LATE MIOCENE AND EARLY PLIOCENE
OF FLORIDA
BY
JONATHAN J. BECKER
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
1985


76
based on a cervical vertebra to P. brunhuberi. Phalacrocorax intermedius
(Milne-Edwards) from the Oreleanian of Orleanais, France was described
from a proximal end of a humerus. Phalacrocorax brunhuberi may be
synonymous with P. intermedius; only slightly smaller size and slightly
younger age prevented Brodkorb (1980) from placing it in synonymy with
this species.
Phalacrocorax ibericum Villalta, probably from the Lower Pontian of
Spain, is based on the distal end of a humerus. Villalta states that
this cormorant is smaller than the other Aquitanian cormorants of Europe
(P. littoralis, P. miocenaeus, and P. intermedius) and is close to the
living P. carbo.
Phalacrocorax goletensis Howard from the late Hemphillian or early
Blancan La Goletia local fauna, Michocan, Mexico, is known from a
coracoid (type) and a referred distal humerus. It is possibly ancestral
to P. olivaceus.
Five species, all said to be ancestral to P. auritus, have been
described from the late Hemphillian to Mid-Pleistocene of North America.
Phalacrocorax wetmorei Brodkorb, described from the late Hemphillian Bone
Valley District, Florida, is known from all major skeletal elements. See
additional remarks above. Phalacrocorax kennelli Howard from the Blancan
San Diego Formation, was described on a partial coracoid, a humerus, a
furculum, and vertebrae. It agrees in size with P. pelagicus and P.
penicillatus. In morphology, the fossil species resembles P. auritus or
P. pelagicus (Howard, 19^9). Phalacrocorax idahensis (Marsh) from the
Hemphillian Castle Creek, Idaho, is based on a proximal carpometacarpus.
Murray (1970) referred additional material to and redescribed this
species from the (Blancan) Hagerman local fauna. Phalacrocorax macer


196
Remarks on the Order Passeriformes
There have been very few fossil species of passerines described
(Brodkorb, 1978)- With a few exceptions, still fewer are believably
assigned to a genus or family. Most fossil species were compared to very
few other species and were not adequately diagnosed as a member of the
family in which they were placed.
Nine fossil genera of passerines have been described (Brodkorb,
1978). Miocitta Brodkorb is known only from the late Barstovian Kennesaw
local fauna, Colorado. Protocitta Brodkorb is from the Blancan of Texas
and Kansas and from the Pleistocene of Florida and Texas. Henocitta
Holman was described from the Pleistocene of Florida. All of the above
genera were described as medium to large sized jays.
Palaeoscinus Howard, from Mohanian of Tepsquet Creek, California,
was described from a slab and represents an extinct family of passerines.
Howard (1957) states this family has its affinities with the -
Pycnonotidae, Bombycillidae, Corvidae, and Cinclidae.
Necropsar Slater is based on a postcranial skeleton from the
Holocene of Rodriguez Island. It is placed in the family Sturnidae.
Genera of fossil icterids include Cremaster Brodkorb from the
Pleistocene of Florida, Pandanaris A. H. Miller from the Pleistocene of
Florida and California, and Pyelorhamphus A. H. Miller from the
Quaternary of New Mexico.
Palaeostruthus Wetmore was described as a late Clarendonian to early
Hemphillian genus of emberzid finch from Florida and Kansas. Steadman
(1981) synomized this genus with the living genus Ammodramus.
Additionally, Passerina (cf.) is reported from the Hemphillian Yepomera
local fauna (Steadman and McKittrick, 1982). These last two genera are


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.20 .15 .10 .05 0.0 + .05 .10 .15 .20
1 1 1 1 I I I L-
126


137
Dendrocygna sp.
Material. Love Bone Bed local fauna; UF 25992, UF 25997, UF 29765,
UF 29766, right coracoids; UF 29764, UF 29763, UF 25803, left coracoids;
UF 29762, UF 25774, UF 25839, humeral ends left coracoids (tenatively
referred); UF 25845, humeral end right coracoids (tentatively referred);
UF 25755, left carpometacarpus, missing metacarpal III, UF 25757,
prominent end left carpometacarpus.
Remarks. Coracoids and carpometacarpus (Fig. 4.7) typical of
Dendrocygna as described by Woolfenden (1961). I have been unable to
find either qualitative or quantitative characters which will distinguish
the fossil specimens from the Love Bone Bed from the few specimens of the
Recent species of Dendrocygna available, when the variation of modern
populations is taken into account. Measurements are given in Table 4.19.
Fossil species in this tribe include Dendrochen robusta A. H.
Miller, based on humeri from the early Hemingfordian Flint Hill local
fauna, South Dakota; and Dendrocygna eversa Wetmore, based on a proximal
humerus from the Blancan Benson local fauna, Arizona. Dendrocygna
validipinnis DeVis, from the Pleistocene of Australia was shown by Olson
(1977a) to be a junior synonym of Biziura.
Recently, Cheneval (1984) transfered three species previously
referred to Anas from St.-Gerand-le-Puy to the genus Dendrochen as D.
blanchardi, D. consobrina and D. natator. He also noted a possible
relationship between Romainvillia and Dendrochen.
Dendrocygna is predominately tropical, with the greatest diversity
of species occurring in the New World tropics and southeastern Asia. Two
species (D. viduata and D. autumnal is) occur in both the Old World and


22


239
Olson, S. L. in press. The Fossil Record of Birds. Avian Biology.
Vol. 8. Academic Press, New York.
Olson, S. L. and J. Farrand, Jr. 1974. Rhegminornis restudied: a tiny
Miocene turkey. Wilson Bulletin, 86:114-120.
Olson, S. L. and A. Feduccia. 1979 A Old World occurrence of the
Eocene avian family Primobucconidae. Proceedings of the Biological
Society of Washington, 92:494-1+97.
Olson, S. L. and A. Feduccia. 1980. Relationships amd Evolution of
Flamingos (Aves: Phoenicopteridae). Smithsonian Contributions to
Zoology, 31& 73 pp.
Olson S. L. and D. W. Steadman. 1981. The relationships of the
Pedionomidae (Aves: Charadriiformes). Smithsonian Contributions to
Zoology, 337* 25 pp.
Ono, K. 1980. Comparative osteology of three species of Japanese
cormorants of the Genus Phalacrocorax (Aves, Pelecaniformes).
Bulletin National Science Museum, Ser. C (Geology), 6:129-151
Opdyke, N. D., D. R. Spangler, D. L. Smith, D. S. Jones, and R. C.
Lindquist. 1984. Origin of the epeirogenic uplift of Pliocene-
Pleistocene beach ridges in Florida and the development of the
Florida karst. Geology, 12:226-228.
Palmer, R. 1962. Handbook of North American Birds. Volume 1. Loons
through Flamingos. Yale University Press, New Haven. 567 pp.
Palmer, R. 1975* Handbook of North American Birds. Volume 3- Waterfowl
(concluded). Yale University Press, New Haven. 560 pp.
Palmer, R. 1976. Handbook of North American Birds. Volume 2.
Waterfowl (first part). Yale University Press, New Haven. 521 pp.
Patton, T. H. and B. E. Taylor. 1973. The Protoceratinae (Mammalia,
Tylopoda, Protocertidae) and the systematics of the
Protoceratidae. Bulletin of the American Museum of Natural History,
150:347-414.
Payne, R. and C. Risley. 1976. Systematics and evolutionary-
relationships among the herons (Ardeidae). Miscellaneous
Publications, Museum of Zoology, University of Michigan, 150:1-115.
Peters, J. L. 1931. Check-list of Birds of the World. Volume 1.
Harvard University Press, Cambridge, Massachusetts. 345 pp.
Peters, J. L. 1934. Check-list of Birds of the World. Volume II.
Harvard University Press, Cambridge, Massachusetts. 409 pp.
Peters, J. L. 1940. Check-list of Birds of the World. Volume IV.
Harvard University Press, Cambridge, Massachusetts. 291 pp.


36
with articulated fish skeletons, the present elevation, and the marine
taxa present (Becker, 1985a; Berta and Morgan, in press).
Studies on the fossil mammals from here include Webb's (1969) paper
on Qsteoborus ore, Hirschfeld and Webb's (1968) study on Pliometanastes
protistus, Webb's (1973) mention of antilocaprids, and Wolff's (1978)
study of the cranial anatomy of Indarctos. The concurrent range zones of
the first two taxa, and the presence of Indarctos, Machairodus, and
Pseudoceras indicate an age of late early Hemphillian.
Fossil birds from here include a very a small species of Egretta and
an indeterminate species of Buteo (Becker, 1985a). A preliminary faunal
list is included in this paper.
Haile VB
This locality is from the NE 1/4, Sec. 23, T. 9 S., R. 18 E.,
Newberry Quadrangle, U. S. Geologic Survey 7*5 minute series
topographical map, 1968. Auffenberg (1954) discusses the geology of this
locality and describes the abundant material of Gavialosuchus americanus
(Sellards) from this locality. A number of equids are known from this
site including Pliohippus, Cormohipparion, Calippus, and Nannippus.
Haile VI
This locality is in the Nl/2, SW1/4, Sec. 24, T. 9 S., R. 17 E.,
Newberry Quadrangle, U. S. Geologic Survey 7*5 minute series
topographical map, 1968, Alachua County, Florida. It was collected by
the Florida Geologic Survey and Florida State Museum. Auffenberg (1963)
discussed the geology and concluded that this locality represents a
stream deposit. The mammalian fauna includes 'Hipparion', Pseudoceras,
and the type of Mylagaulus kinseyi Webb (1966). A new species of


209
Origination rates show a peak during the Hemingfordian, a very large
increase in the Clarendonian, a decrease in the Hemphillian, although not
to pre-Clarendonian level, then an additional increase in the Blancan.
Extinction rates tend to he much lower than origination rates, hut show a
roughly parallel trend. Again, a high value is noted in the
Clarendonian.
Turnover rates (T), or the numerical average of origination rates
and extinction rates, predictably parallel the trends in origination and
extinction rates described above. Running means (Rm), or the diversity
minus the average of originations and extinctions of a given land mammal
age, increase at a decreasing rate throughout the Neogene.
Turnover rates per genus (T/Rm) show a high value in the late
Arikareean, decrease through the Barstovian, then peak in the
Clarendonian, to decrease and show a low rate through to the end of the
Neogene.
Marine versus Non-marine
A more instructive approach is to divide the North American avifauna
into marine and non-marine subgroups and then compare and contrast these
two broad divisions (Table 6.3; 6.U and Figure 6.1; 6.2). The following
observations result.
(l) The North American marine avifauna is essentially established at a
diversity of 20 to 25 genera by the Clarendonian. The North
American non-marine avifauna is roughly stable at a diversity of 3
genera during the late Arikareean and Hemingfordian. From this
NALMA through the remainer of the Neogene, diversity increases from
13 to 26 genera by the Blancan.


227
A number of shorebirds (Family Scolopacidae) are known. Most are
small and are tentatively referred to the genus Calidris.
Two owls are known. One is an undescribed genus (Family Tytonidae)
which appears to be related to the barn owl and the grass owl of south
east Asia. Another (Family Strigidae), probably in the genus Bubo, is
known from Bone Valley.
Three perching birds (Order Passeriformes) are present. All are
poorly known.
The most diverse localities are the Love Bone Bed local fauna with
approximately 44 taxa and the Bone Valley Mining District with
approximately 4l taxa, 31 of which were studied in this paper. These
localities are the most diverse non-marine and marine avifaunas,
respectively, known in North America prior to the Pleistocene. Other
localities are much less diverse. Approximately l4 taxa are represented
at McGehee Farm, while the other localities included in this study
typically have fewer than 3 or 4 taxa present.
Paleoecology
The second aspect of this study focused on paleoecology. Birds from
two environments dominate the late Miocene and early Pliocene localities
in Florida. Birds from the Bone Valley Mining District, SR-64, and
Manatee County Dam are interpreted as being from near-shore marine
environments. Abundantly represented taxa from other localities, such as
the Love Bone Bed and McGehee Farm, are interpreted as representing
freshwater ponds and streams, with marshes, mudflats, and estuaries not
far removed. More terrestrial localities are either very poorly
represented or lacking for this time period in Florida.


210
(2) The absolute number of genera which become extinct is roughly stable
over most of the Neogene with about 2 generic extinctions per
million years in the marine avifauna and about 7 generic extinctions
per million years in the non-marine avifauna.
(3) In the marine avifauna, the rate of origination peaks during the
Clarendonian with 4.4 new genera appearing per million years.
(4) In the non-marine avifauna, there are peaks in the generic
origination rates during the Hemingfordian, Clarendonian and Blancan
Land Mammal Ages, with 6, 8, and 10 genera appearing per million
years, respectively. These peaks rise above a background rate of 2
to 4 genera appearing per million years in the intervening NALMAs,
or roughly a cyclic 2- to 4-fold change.
(5) Extinction rates are typically low in the marine avifauna throughout
the Neogene. With the exception of the Clarendonian, which shows an
extinction rate of 2 genera per million years, all other NALMAs have
an extinction rate much less than 1.0 extinction per million years.
In the non-marine avifauna, extinction rates average from 1.0 to 2.8
generic extinctions per million years.
(6) Turnover rates are consistently low in the Marine avifauna, with the
highest turnover rate being in the Clarendonian. In non-marine
avifauna, turnover peaks appear in alternating mammal ages (in the
Hemingfordian, Clarendonian, and Blancan). These peaks in turnover
represent a 2- to 3-fold increase over the rates in the intervening
NALMAs. Fluctuations in the turnover rates are governed more by
changes in origination rate than by changes in extinction rates.
(7) The per-genus turnover rate (T/Rm) in marine avifauna is high in the
late Arikareean (1.68) and then drops to approximately 0.20 through


88
Family Anhingidae Ridgway, l88T
Remarks. Skeletal elements of anhingas discussed below may be
distinguished from the Phalacrocoracidae as follows: Humerusby
characters given by Miller (1966) and Martin and Mengel (19T5) Coracoid
head rotated ventrad and mediad, to produce a distinct notch between
head and shaft, when observed from either a ventral or medial view (head
merges smoothly with shaft in the Phalacrocoracidae). Procoracid
expanded and concave cotyla scapularis present.
Genus Anhinga Brisson, 1760
Anhinga grandis Martin and Mengel, 1975
Material. Love Bone Bed local fauna; UF 25739, proximal end right
humerus; UF 25723, UF25725, distal ends right humeri; UF 26000, nearly
complete right coracoid; UF 25873, distal end right tibiotarsus.
McGehee Farm local fauna; UF 11107, distal end right humerus.
Remarks. Elements described in detail in Becker (in prep*). Distal
humeri compare exactly in all features with the type. Coracoid assigned
to this species, as it is of correct size.
Anhinga sp. unknown
Material. Bone Valley Mining District, no specific locality; UF
29781, proximal end left ulna; UF 29780, distal end left ulna.
Remarks. The ulnae are much larger than comparable elements of A.
anhinga and those expected for A. grandis. Parenthetically, the ulna
from Coleman III (UF 1666*0, refered to A. cf. grandis by Ritchie (1980),
is definitely not Anhinga grandis and is not identifiable to a species.
Along with the two specimens from Bone Valley, this Coleman III specimen
represents a much larger species of anhinga which existed approximately


Figure 4.6. Pliogyps sp. A, B. UF 25886, distal end left
tibiotarsus. A. Caudal view. B. Cranial view. C, D. UF
right tarsometatarsus. C. Plantar view. D. Dorsal view.
A, B = 3 cm.; C, D = 5 cm.
25952,
Scale


Love Bone Bed and from McGehee Farm. It is larger than the living New
World anhinga and is fairly completely known.
Six species of heron (family Ardeidae) are present. These include 2
species of Ardea, at least 2 species of Egretta, a species of Ardeola,
and a single species of Nycticorax. Herons are rare members of fossil
avifaunas.
Four storks are present as are 3 species of ibis. Storks (Family
Ciconiidae) include a species of Mycteria from the Love Bone Bed and
McGehee and 3 species of Ciconia, distributed between the Love Bone Bed,
Mixson Bone Bed, and Bone Valley. This shows that storks were more
diverse in the late Miocene and early Pliocene of North America than they
are today. Ibises (Family Plataleidae) include a species of Eudocimus
from the Bone Valley Mining District, and a specimen of Plegadis cf. P.
pharangites and one representing a large species of indeterminate ibis
from the Love Bone Bed.
Nine taxa of accipitriform birds are present, including a species of
New World Vulture (Family Vulturidae), 1 or possibly 2 species of Osprey
(Family Pandionidae), and 7 species of hawk or eagle (Family
Accipitridae). Of note from the Love Bone Bed is an undescribed species
of vulture, Pliogyps, and the most primitive species of osprey, Pandion
lovensis, now known. An indeterminate species of Pandion, ?Haliaeetus
sp., Aguila sp. A., and an indeterminate genus are all known from the
Bone Valley Mining District. A species of Buteo near the size of Buteo
jamaciensis occurs in the Withlacoochee River 4a local fauna.
Waterfowl (Family Anatidae) are common, with 5 species of geese and
8 species of ducks being present. Of note is a well-represented species
of Dendrocygna from the Love Bone Bed which cannot be distinguished from


Table 4.7continued
Measurements
P. a. auritus
P. a. floridanus
Ulna
W-PROX
11.95 + 0.4l
11.4 12.7
11.11 + 0.69
9.8 12.4
D-LENGTH
15.74 + 0.83
14.5 17.2
14.19 + 1.12
12.3 l6.4
D-PROX
12.50 + O.58
11.5 13.5
11.31 + 0.90
9.8 13.0
ECON
8.26 + 0.36
7.8 8.9
7.62 + 0.48
6.9 8.5
CPTB
8.95 + 0.27
8.3 9.3
8.37 + 0.46
7.5 9.0
ECON-CPIB
11.15 + 0.50
10.3 11.9
10.38 + 0.49
9-7 11.3
ECON-ICON
6.29 + 0.27
5.8 6.7
5.83 + 0.23
5.4 6.3
i
P. wetmorei
MCD/HXIXA McGehee LBB
11.52 + 0.42 (27)
10.F 12.8
11.1
14.73 + 0.70 (22)
13.3 l6.0
14.1
11.61+0.43 (27) 11.2
10.5 12.3
8.04 + 0.26 (27) 8.0
7.5 8.6
8.46 + 0.37 (27) 8.4
7-7 9.0
10.70 + 0.35 (28) 10.6
9.7 11.3
6.47 + 0.15 (26) 6.2
6.2 6.7
00


TI
Bone Valley Mining District, Payne Creek Mine.UF 29741, UF 57304,
humeral ends right coracoids; UF 21203, proximal end left
carpometacarpus; UF 29742, distal end right carpometacarpus.
Bone Valley Mining District, Swift Mine.UF 55883, distal end right
tibiotarsus; UF 17687, distal end right tatsometatarsus.
Bone Valley Mining District, specific locality unknown.UF 61549,
caudal portion left mandible; UF 61550, nearly complete right coracoid;
UF 61553, nearly complete left coracoid; UF 61554, UF 61555, humeral ends
left coracoids; UF 61551, humeral end right coracoid; UF 61552, sternal
end right coracoid; UF 61556, sternal end left coracoid; UF 61557, UF
61558, proximal end right humerus; UF 61559, UF 61560, proximal end left
humeri; UF 61561, distal end right humerus; UF 61562, UF 61563, UF 61564,
UF 61565, UF 61566, UF 61567, UF 61568, UF 61569, distal end left humeri;
UF 61570, UF 61571, proximal end left ulnae; UF 61572, UF 61573, UF
61574, distal ends right ulnae; UF 61575, distal end left ulna; UF 61578,
nearly complete left carpometacarpus; UF 61576, UF 61577, proximal ends
right carpometacarpi; UF 61579, proximal end left carpometacarpus; UF
61596, complete left femur; UF 61580, proximal end left tibiotarsus; UF
61581, UF 61582, UF 61583, UF 61584, distal ends right tibiotarsi; UF
61585, UF 61586, UF 61587, distal ends left tibiotarsi; UF 61588, nearly
complete left tarsometatarsus; UF 61589, UF 61590, UF 61591, UF 61592,
proximal end left tarsometatarsus; UF 61593, distal end right
tarsometatarsus; UF 61594, UF 61595, distal ends left tarsometatarsi.
Bone Valley Mining District, specific locality unknown (FGS
collection).V 7311, proximal end left humers; V 7313, distal end left
humerus; V 7309, distal end left ulna; V 7310, proximal end left
carpometacarpus; V 7312, distal end left femur.


112
Table 4.15. Measurements of tibiotarsi, coracoids, and tarsometatarsi of
the ibises Eudocimus ruber (N =8, 2 males, 6 females), Eudocimus albus
(N = 12, 6 males, o females), and Eudocimus species from the Bone Valley
Mining District. Data are mean +_ standard deviation and range.
Abbreviations are defined in the methods section.
Measurements
Eudocimus ruber
Tibiotarsus
W-SHAFT
4.90
+
0.20
4.7
-
5.3
D-SHAFT
4.09
+
0.20
3.8
-
4.5
W-DIST-CR
9.09
+
0.64
8.5
-
10.5
W-DIST CD
6.74
+
0.30
6.4 -
- 1
r.3
D-MCON
9.69
+
0.42
9.3
-
10.6
D-LCON
9.14
+
0.50
8.6
-
10.2
D-ICON
6.04
+
0.27
5.8
-
6.6
Eudocimus albus Eudocimus sp.
5.02 + 0.40
4.5 5.6
5-6
4.53 + 0.53
3.9 5.8
4.7
9.74 + 0.74
8.5 10.5
10.7
7.07 + 0.4l
6.3 7-5
7.5
10.76 + 0.78
9.2 11.6
11.2
10.26 + 0.69
9.1 11.1
10.8
6.70 + 0.60
5.7 7.3
6.9


NUMBER OF GENERA
221
GENERA LOCALITIES
L ARIK HEMING BARST 1 CLAR 1 HEMP 1 BLANC
NUMBER OF LOCALITIES


20


51
skeletons, necessary to determine the variability of the characters used
above, prevents the naming of this species as new.
Genus Tachybaptus Reichenbach, 1853
Tachybaptus sp. indet.
Material. Love Bone Bed local fauna; UF 25796, UF 25817, UF 25818,
UF 29671, complete left coracoids; UF 29668, UF 29669, UF 29672, humeral
ends left coracoids; UF 26006, UF 260l4, UF 26017, UF 26019, UF 2966k, UF
29665, UF 29666, UF 29669, complete right coracoids. UF 25773, complete
right femur. UF 29663, proximal end of right tibiotarsus (questionably
referred).
McGehee Farm local fauna; UF 67810, proximal end right tibiotarsus
(questionably referred).
Description. All coracoids are waterworn, abraded, or broken to
varying degrees, making detailed descriptions difficult. All coracoids
near that of Tachybaptus dominicus in size and overall shape. ^Additional
description is not possible.
Femur also near Tachybaptus dominicus in size and general
morphology, but the femoral shaft slightly more gracile. Depression
cranial to the patellar sulcus is absent in fossil. Measurements are
given in Table 4.4.
Both tibiotarsal fragments are questionably referred solely on the
basis of size.
Remarks. The use of generic names follow Storer (1976a). This
material is not diagnostic enough to allow identification to the level of
species.


Figure 6.1. Distribution of avian genera and localities through
geologic time. Abbreviations as in Table 6.1.


235
Holman, J. A. 1959 Birds and mammals from the Pleistocene of
Williston, Florida. Bulletin of the Florida State Museum,
Biological Sciences, 5:1-25
Holman, J. A. 1961. Osteology of living and fossil New World Quails
(Aves, Galliformes). Bulletin of the Florida State Museum,
Biological Sciences, 6:131-233.
Howard, H. 1929* The avifauna of Emeryville shellmound. University
California Publication in Zoology, 32:301-394. 4 pis., 55 text figs.
Howard, H. 1932a. A new species of Cormorant from Pliocene deposits
near Santa Barbara California. Condor, 34:118-120.
Howard, H. 1932b. Eagles and eagle-like vultures of the Pleistocene of
Rancho La Brea, California. Carnegie Instition of Washington,
Publication 429:1-82, 29 pis.
Howard, H. 1942. A review of the American fossil storks. Carnegie
Instition of Washington, Publication, 530:187-203, 1 pi.
Howard, H. 1946. A review of the Pleistocene birds of Fossil Lake,
Oregon. Carnegie Institution of Washington, Publication, 551:l4l-
195.
Howard, H. 1949* New avian records for the Pliocene of California.
Carnegie Institution of Washington, Publication, 584:177-199*
Howard, H. 1957. A new species of passerine bird from the Miocene of
California. Contributions in Science, Natural History Museum of Los
Angeles County, 9:1-16.
Howard, H. 1964. Fossil Anseriformes. pp. 233-326. In J. Delacour
(ed). Waterfowl of the World. Vol. 4. Country Life Ltd., London.
Howard, H. 1965. A new species of cormorant from the Pliocene of
Mexico. Bulletin Southern California Academy Science, 64:50-55.
Howard, H. 1973. Fossil Anseriformes. General Corrections and
Additions, pp. 371-378. In. J. Delacour (ed). Waterfowl of the
World. Second Edition. Vol. 4. Hamlyn Publishing Group Ltd.,
London.
Howard, H. 1980. Illustrations of avian osteology taken from 'The
Avifauna of Emeryville Shellmound'. Contributions in Science,
Natural History Museum of Los Angeles County, 330:xxvii-xxxviii.
Hulbert, R. C. 1982. Population dynamics of the three-toed horse
Neohipparion from the late Miocene of Florida. Paleobiology, 8:159-
167.
Hull, D. L. 1970. Comtempory systematic philosophies. Annual Review of
Ecology and Systematics. 1:19-54.


243
Webb, S. D. 1973. Pliocene pronghorns of Florida. Journal of
Mammalogy, 54:203-221.
Webb, S. D. 1974. Chronology of Florida Pleistocene Mammals, pp. 5-31.
In S. D. Webb (ed.). Pleistocene Mammals of Florida. University
Presses of Florida, Gainesville.
Webb, S. D. 1976. Underwater Paleontology of Florida Rivers, pp. 479-
48l. National Geographic Society Research Reports, 1968 Projects,
National Geographic Society, Washington, D.C.
Webb, S. D. 1981. Kyptoceras amatorum, new genus and species from the
Pliocene of Florida, the last protoceratid artiodactyl. Journal of
Vertebrate Paleontology, 1:357-365.
Webb, S. D. 1983. A new species of Pediomeryx from the late Miocene of
Florida, and its relationships within the subfamily Cranioceratinae
(Ruminantia: Dromomerycidae). Journal of Mammalogy, 64:261-276.
Webb, S. D. 1984. On two kinds of rapid faunal turnover, pp. 417-436.
In W. A. Berggren and J. A. Van Couvering (eds.). Catastrophes and
Earth History: the new uniformitarianism. Princeton University
Press, Princeton.
Webb, S. D. in press. Osteology of Thinobadistes Hay, the oldest
mylodontid sloth from North America. Florida State Museum Bulletin,
Biological Sciences.
Webb, S. D., B. J. MacFadden, and J. A. Baskin. 1981. Geology and
Paleontology of the Love Bone Bed from the Late Miocene of Florida.
American Journal of Science, 281:513-544.
Webb, S. D. and N. Tessman. 1968. A Pliocene vertebrate fauna from low
elevation in Manatee County, Florida. American Journal of Science,
266:777-811.
Weigel, R. D. 1958. Great Auk remains from a Florida shellmidden. Auk,
75:215-216.
Weigel, R. D. 1962. Fossil Vertebrates of Vero, Florida. Florida
Geological Survey Special Publication, 10:1-59.
Welty, J. C. 1975- The Life of Birds. W. B. Saunders, Philadelphia.
623 pp.
Wetmore, A. 1931. The avifauna of the Pleistocene in Florida.
Smithsonian Miscellaneous Collections, 85:1-41.
Wetmore, A. 1943- Fossil birds from the Tertiary Deposits of Florida.
New England Zoological Club, 32:59-68.
Wetmore, A. 1958. Miscellaneous notes on fossil birds. Smithsonian
Miscellaneous Collections, 135:1-11


IT
8. D-PROX.Depth from dorsal edge of proximal articular surface
to closest bypotarsal canal (Canalis hypotarsi), or the closest
hypotarsal groove (Sulcus hypotarsi), if no canals are present as
in the Accipitridae.
9. W-HYPOTS.Greatest transverse width of hypotarsus.
9a- W-HYPOTS-C.Transverse width of tuberosity on medial
hypotarsal crest (Crista medialis hypotarsi), in cormorants only.
10. L-HYPOTS.Length of medial hypotarsal crest.
11. D-PROX-L.Depth of proximal end, measured from dorsal edge of
the proximal articular surface through the lateral hypotarsal
crest (Crista lateralis hypotarsi).
11a- D-PROX-M.Depth of proximal end, measured from dorsal edge
of the proximal articular surface through the medial hypotarsal
crest, if the lateral hypotarsal crest is reduced.
12. L-MTI.Greatest length of metatarsal I facet (Fossa
metatarsi I_).
13. D-D-SHAFT.Depth of shaft at cranial edge of distal canal
(Foramen vasculare distale).
14. W-DIST.Greatest transverse width of distal end (if trochlea
are of equal length).
15. TRIII-TRIV.Greatest transverse width from trochlea III
through trochlea IV (if trochlea II is elevated).
16. TRII-TRIV.Greatest transverse width between plantar portion
of trochlea II and plantar portion of trochlea IV.
IT. W-TRII.Greatest transverse width of trochlea II.
l8. D-TRII.Greatest depth of trochlea II.
19* W-TRIII.Greatest transverse width of trochlea III.


28
major unpublished localities was added to this database. This produced a
total of 133 localities from the Neogene of North America (86 published,
47 unpublished) with a record of fossil birds. (3) The occurrences of
fossil taxa were condensed to tabular form to reflect the actual,
documented range of taxa at the family, subfamily, and generic level. A
given taxonomic range reflects the summation of all lower taxonomic ranks
plus material which is only diagnostic to that given level. (4) From
this table, geological ranges were inferred parsimoniously. For example,
if a genus is known from the early Hemingfordian and the late
Clarendonian, I inferred that it also occurred in North America during
the interval between these endpoints. (5) From this table (#3 above),
biochronologically useful species were identified.
Indices of avian faunal dynamics were calculated following Marshall
et al. (1982): the duration (d) of each land mammal age is given in
millions of years, to the nearest tenth, and is based on all available
data; diversity (Si) represents the total number of genera known for each
land mammal age; originations (Oi) are the number of generic first
appearances in a given land mammal age; extinctions (Ei) are last
appearances of a genus in a given land mammal age; running means (Rm)
compensates for time intervals of unequal duration by subtracting the
average of originations (Oi) and extinctions (Ei) for a given age from
the diversity (Si) of that age, or Rm = Si (Oi + Ei)/2; origination
rates (Or) adjust for unequal time intervals by dividing the total number
of generic originations (Oi) occurring during a given land mammal age by
the duration (d) of that interval; similarly, the extinction rate, Er =
Ei/d; turnover rates (T) are the average number of genera that either
originate or go extinct during a given land mammal age, or T = (Or +


165
Table 4.23. Measurements of the humeri, tibiotarsi, and tarsometatarsi
of Grus americana (N = 9 maximun, 2 males, 2 females, 5 unsexed), Grus
japonensis (N = 6 maximun, all unsexed), and Grus sp. B. from the Love
Bone Bed local fauna. Data are mean +_ standard deviation and range.
Abbreviation are defined in methods section. (*) Specimen abraded or
broken.
Measurements
G. americana
Humerus
W-DIST
35.93
34.0
+ 1.84
- 38.5
D-DIST
20.56
18.8
+ 1.69
- 23.8
Tibiotarsus
W-SHAFT
11.19
10.3
+ 0.8l
- 13.0
D-SHAFT
9-T9
8.6 '
t- 0.83
- 11.3
W-DIST-CR
22.68
21.0
+ 1.00
- 24.1
W-DIST-CD
16.97
15.8
+ 0.71
- 18.1
D-MCON
21.72
21.2
+ 1.16
- 23.6
D-LCON
21.29
19.6
+ 1.33
- 23.1
D-ICON
12.64
11.6
+ 0.62
- 13.6
G. japonensis Grus sp. B.
38.73
+ 1.48
35.7
37.2
- 41.5
21.13
+ 0.94
18.0; *17.9
19.8
- 22.3
12.23
+ 0.34
10.7
11.8
- 12.8
10.33
1 +
0

-P"
V/l
10.7
9.7
- 11.0
24.83
+ 0.91
*23.7; 23.9;
23.9
- 26.4
23.1
18.52
+ 1.11
18.8; *15.2
17.2
- 20.4
24.93
+ 0.88
*24.1; *21.8
23.5
- 25.6
*23.2
23.33
+ 0.96
*22.8; 24.0;
22.2
- 24.5
*21.8; *21.5
13.08
+ 0.48
13.04 + 0.78 (:
12.5
- 13.6
12.3 14.2


48
from the late Miocene and early Pliocene
Asterisks denote marine taxa, which are not
are based on previously published works and
Table 3.2 Checklist of birds
Bone Valley Mining District,
included in this study. Taxa
on original identifications.
^Family Gaviidae
*Gavia palaeodytes
*Gavia concinna
Family Podicipedidae
Podiceps sp.
Pliodytes lanquisti
^Family Diomedidae
*Diomedea anglica
Family Phalacrocoracidae
Phalacrocorax wetmorei
Phalacrocorax idahensis
*Family Sulidae
*Morus peninsularis
*Sula guano
*Sula phosphata
^Family Procellaridae
^Family Pelecanidae
*Pelecanus sp.
Family Plataleidae
Eudocimus sp.
Family Ardeidae
Ardea polkensis
Family Ciconiidae
Ciconia sp.
Family Anatidae
Bucephala ossivallis
Family Pandionidae
Pandion sp.
Family Accipitridae
Haliaeetus sp.
Buteo sp.
Family Scolopacidae
Calidris pacis
Erolia penpusilla
Limosa ossivallis
Family Phoenicopteridae
Phoenicopterus floridanus
*Family Haematopidae
*Haematopus sulcatus
*Family Laridae
*Larus elmorei
^Family Alcidae
*Australea granis


170
representing an undetermined species of Rallus until tetter material is
available.
Rallus (cf.) sp. C.
Material. Love Bone Bed local fauna; UF 25732, distal end left
humerus; UF 26025, UF 26027, UF 26030, UF 29705, UF 29712, humeral ends
right coracoids; UF 29708, left coracoid; UF 26007, UF 26025, UF 29702,
UF 29703, UF 29704, UF 29710, UF 29711, UF 29713, UF 67807, humeral ends
left coracoids; UF 29707, distal ends right tarsometatarsus.
Remarks. All specimens badly water-worn, within the size range of
Rallus limicola or Porzana Carolina or slightly larger. I cannot
distinguish between these two genera on the basis of waterworn elements,
and have therefore arbitrarily assigned these specimens to Rallus on the
basis of size.
Undescribed Genus and Species
Material. Love Bone Bed local fauna; UF 25727, proximal end left
humerus; UF 25836, UF 25849, UF 29715, UF 29716, UF 29717, complete (or
nearly so) right coracoids; UF 29718, humeral end right coracoid; UF
29719, humeral end left coracoid; UF 25865, UF 25866, distal ends right
tarsometatarsi.
McGehee Farm local fauna; UF 9494, proximal end right humerus; UF
29748, nearly complete right coracoid.
Description. Coracoid with a greatly expanded procoracoid process,
extending relatively far down shaft; shaft fairly slender; dorsal surface
not deeply excavated; coracoid fenestra small; lateral process greatly
expanded (Fig. 4.9).


235
Holman, J. A. 1959* Birds and mammals from the Pleistocene of
Williston, Florida. Bulletin of the Florida State Museum,
Biological Sciences, 5:1-25
Holman, J. A. 1961. Osteology of living and fossil New World Quails
(Aves, Galliformes). Bulletin of the Florida State Museum,
Biological Sciences, 6:131-233.
Howard, H. 1929. The avifauna of Emeryville shellmound. University
California Publication in Zoology, 32:301-394. 4 pis., 55 text figs.
Howard, H. 1932a. A new species of Cormorant from Pliocene deposits
near Santa Barbara California. Condor, 34:118-120.
Howard, H. 1932b. Eagles and eagle-like vultures of the Pleistocene of
Rancho La Brea, California. Carnegie Instition of Washington,
Publication 429:1-82, 29 pis.
Howard, H. 1942. A review of the American fossil storks. Carnegie
Instition of Washington, Publication, 530:187-203, 1 pi.
Howard, H. 1946. A review of the Pleistocene birds of Fossil Lake,
Oregon. Carnegie Institution of Washington, Publication, 551:l4l-
195.
Howard, H. 1949 New avian records for the Pliocene of California.
Carnegie Institution of Washington, Publication, 584:177-199*
Howard, H. 1957. A new species of passerine bird from the Miocene of
California. Contributions in Science, Natural History Museum of Los
Angeles County, 9:1-16.
Howard, H. 1964. Fossil Anseriformes. pp. 233-326. _In J. Delacour
(ed). Waterfowl of the World. Vol. 4. Country Life Ltd., London.
Howard, H. 1965. A new species of cormorant from the Pliocene of
Mexico. Bulletin Southern California Academy Science, 64:50-55*
Howard, H. 1973* Fossil Anseriformes. General Corrections and
Additions, pp. 371-378. In J. Delacour (ed). Waterfowl of the
World. Second Edition. Vol. 4. Hamlyn Publishing Group Ltd.,
London.
Howard, H. 1980. Illustrations of avian osteology taken from 'The
Avifauna of Emeryville Shellmound'. Contributions in Science,
Natural History Museum of Los Angeles County, 330:xxvii-xxxviii.
Hulbert, R. C. 1982. Population dynamics of the three-toed horse
Neohipparion from the late Miocene of Florida. Paleobiology, 8:159-
167.
Hull, D. L. 1970. Comtempory systematic philosophies. Annual Review of
Ecology and Systematics. 1:19-54.


Neohipparion eurystyle, and Hexameryx, this locality is late Hemphillian
in age. As for the Manatee County Dam Site, this locality is here
considered an outlier of the classic Bone Valley Formation.
Eustatic Sea-Level Changes
Webb and others (Webb and Tessmann, 1968; MacFadden and Webb, 1982;
Webb, 1984) have proposed a model using the present elevations of fossil
localities with marine or estuarine taxa to reflect the fluctuations of
sea-levels during the late Miocene and early Pliocene in peninsular
Florida. This scheme is predicated on two assumptionsthat the
Florida peninsula had been tectonically stable over the later Cenozoic
and that the sedimentary sequence reflects actual sea-level change. If
these assumptions are valid, then the present elevation of the localities
which contain marine or estuarine taxa should reflect the elevation of
the ocean at the time when these localities were deposited.
The following evidence argues against this model: (l) The relict
Pleistocene shorelines increase in elevation from a low point in southern
Georgia to a maximum elevation in northern peninsular Florida and
gradually lose elevation to the south. This indicates that the northern
part of peninsular Florida has not been stable, and has been
differentially uplifted. Opdyke et al. (1984) argue that this occurred
during the Pleistocene due to the subsurface solution of limestone and
the concomitant isostatic uplift and document an uplift in the magnitude
of 30-50 meters since this time. Most of the high-sea-level localities
are from the northern part of Florida in highly karsted areas.
(2) The elevational changes between most localities could easily be
accounted for by a slight regional dip to the beds. For example, a dip
of 1/20 of 1 degree would account for an elevational difference of 138


117
(caudal view), by having the sulcus extensorius more excavated and
extending farther down shaft (cranial view), and by having the shaft and
trochlea III proportionally less deep. Pliogyps sp. from the Love Bone
Bed has a proportionally greater power-arm ratio (defined below) than P.
fisheri.
Description. Tarsometatarsus proportionally different from all
other vultures except for Pliogyps fisheri (see Figures 4.4 4.6). The
proximal end wide and deep. In cranial view, the proximal vascular
foramina large and approximately equal in size. Papilla for the
attachment of M. tibialis cranialis large, rounded, and in two parts.
Sulcus extensoris extending down the shaft to distal foramen, with a
sharp lateral border. Distinct intermuscular line extending obliquely
through this sulcus (separating attachments for the extensor digitorum
brevis pars hallucis and extensor digitorum brevis pars adductor-extensor
digiti IV; Jollie 1976-1977:243). In caudal view, the tarsometatarsus
has a long ridge extending down the shaft from the hypotarsus,
terminating in an intermuscular line which then extends to the level of
the articular facet of metatarsal I. See Table 4.17 for measurements.
Distal end of tibiotarsus with a broad extensor sulcus. Slight
projection of bone on lateral surface (approximately 4 cm from distal
end) for attachment of the fibula. Intercondylar sulcus broad; with
external condyle merging evenly into it. In distal end view, the
intercondylar sulcus is not symmetrical, with the lateral border sloping
gradually, and the medial border sloping abruptly, up from the base of
the intercondylar sulcus (symmetrical or U-shaped in Coragyps,
Sarcoramphus, Breagyps, Gymnogyps; unsymmetrical or Pliogyps-like in
Cathartes). See Table 4.17 for measurements.


102
i


58
Table 4.1. Checklist of non-marine avian taxa discussed in the text.
Localities where each taxon occurs are given in parentheses Love Bone
Bed (LOV), McGehee Farm (MCG), Mixson Bone Bed (MIX), Bone Valley (BV),
Withlacoochee River 4a (WITH 4a), Manatee County Dam (MD), SR-64, Haile
VB (H5B), Haile VI (h6), and Haile XIXA (H19A).
Class Aves
Order Podicipediformes
Family Podicipedidae
Rollandia sp. (LOV, MIX, MCG)
Tachybaptus sp. (LOV, MCG)
Podilymbus cf. P. podiceps (BV)
Podilymbus sp. A (MIX)
Podiceps sp. (BV)
Pliodytes lanquisti (BV)
Order Pelecaniformes
Family Phalacrocoracidae
Phalacrocorax sp. A (LOV, MCG, H19A)
Phalacrocorax wetmorei (BV, MD, SR-64)
Phalacrocorax cf. idahensis (BV)
Family Anhingidae
Anhinga granis (LOV, MCG, H19A)
Anhinga sp.(BV)
Order Ciconiiformes
Family Ardeidae
Ardea polkensis (BV)
Areda sp. indet.(LOV)
Egretta sp. indet. (LOV, BV)
Egretta subfluvia (WITH 4a)
Ardeola sp. (LOV)
Nycticorax fidens (MCG)
Family Ciconiidae
Mycteria sp. (LOV,MCG)
Ciconia sp. A (LOV)
Ciconia sp. B (MIX,BV)
cf. Ciconia sp. C (BV)
Family Plataleidae
Eudocimus sp. A. (BV)
Plegadis cf. P. pharangites (LOV)
Threskiornithinae, genus et species indet. (LOV)
Order Falconiformes (auct.)
Family Vulturidae
Pliogyps undescribed sp. (LOV)


Table 6.2 Faunal dynamics
of the North American Neogene
avifauna.
Abbreviations as in
Table 6.1
L. ARIK.
HEMING.
BARST.
CLAR.
HEMP.
BLANC.
Duration (MA)
3
3.5
5
2.5
4.5
2.7
Localities (Published)
5 (5)
11 (6)
18 (12)
30 (22)
4o (25)
29 (16)
Sampling Index
1.67
1.71
2.40
8.80
5.56
5.93
Number of genera (Si)
10
28
38
6l
78
98
Originations (No.)
9
23
18
31
29
29
Extinctions (No.)
5
8
8
12
9
7
Running mean (Rm)
3.00
12.50
25.00
39.50
59.00
80.00
Origination Rate
3.00
6.57
3.60
12.40
6.44
10.74
Extinction Rate
1.67
2.29
1.60
4.80
2.00
2.59
Turnover Rate (T)
2.34
4.43
2.60
8.60
4.22
6.67
T/Rm
0.78
0.35
0.10
0.22
0.07
0.08
T/Si
0.23
0.16
0.07
0.14
0.05
0.07
I
216


203
horizon from which they came, or their association with other fossil
vertebrates.
Like the Love Bone Bed, the Bone Valley avifauna is dominated by
aquatic birds. However, in sharp contrast, the most abundant taxa
present are marine. Birds from the Bone Valley Mining District included
in this study are 3 species of grebe, 2 species of cormorant, an anhinga,
2 herons, 2 storks, an ibis, an osprey, 4 species of hawks or eagles, a
goose, 4 species of duck, a turkey, a crane, a rail, a flamingo, 5
species of shorebird, and an owl. There is abundant material of
Phalacrocorax wetmorei, Aythya sp., and Phoenicopterus floridanus. Other
pelagic species, which were not included in this study, but are
abundantly represented, include several (3?) species of alcids, loons,
Larus elmorei, 2 species of Sula, and a species of Morus. Fossil
material still occurs in about the same proportions as was reported by
Brodkorb (1955a), even though the number of taxa in this avifauna have
increased. By far the most common taxon in this deposit is the
cormorant, Phalacrocorax wetmorei. The abundance of cormorants, sulids,
and gulls argues for a near-shore marine environment.
Brodkorb (1955) suggested that the large concentrations of seabirds,
especially cormorants, probably represented a breeding colony. For
future consideration of this hypothesis, I note that there is a very low
frequency of sub-adult specimens in the collections from Bone Valley. I
have been unable to find comparable data on Recent breeding colonies.
It has also been suggested (Brodkorb, 1955; and references therein)
that the large concentrations of fossil birds were responsible for the
formation of the phosphorite deposits. Current hypotheses (Riggs, 1984)
suggest that bacteria at the water-sediment interface are responsible for


95
and UF 25900 are referred here strictly on the basis of size. Both are
too broken and/or worn for further description.
Remarks. See remarks pertaining to Ciconia sp. B., below.
Ciconia sp. B.
Material. Mixson's Bone Bed; F:AM 120-2185, distal end right
tibiotarsus; F:AM 205-3008, distal end left tibiotarsus. Bone Valley
Mining District, Palmetto Mine; UF 21135, distal end right tibiotarsus;
UF 21063, distal end left tarsometatarsus missing trochlea IV
(tentatively referred).
Description. Distal tibiotarsus (F:AM 120-2195) larger than that of
£. mar guar i, similar in size to that of a small Jabir. Agrees with
Ciconia by having the process for the ligamentous attachment above the
distal end of the external condyle ridge-like (papilla-like in Jabir),
and by having the distal opening of the tendinal canal placed more toward
the edge of the bone (more toward the middle of the bone in Jabir).
Agrees with Jabir in having the distal end slightly laterally
compressed.
Distal tibiotarsus (UF 21135) similar to F:AM 120-2185, but with a
robust shaft and having the distal opening of the tendinal canal more
toward the middle of the bone. Very similar to some specimens of Jabir,
but can be distinguished from this genus by having the ridge from the
condyle to tubercle slightly notched, and the ligamentous attachment
above the distal end of the external condyle ridge-like (as in Ciconia).
See Figure 4.2 for comparisons with other species of Ciconia.
Tarsometatarsus fragment assigned on size.


62
Table 4.3. Measurements of the tarsometatarsi of the grebes Rollandia
rolland chilensis (N = 6, 2 males, 1 female, 3 unsexed), Podilymbus
podiceps (N = 14, T males, T females), Rollandia sp., and Podilymbus
sp. A. from the Mixson's Bone Bed. Data are mean + standard deviation
and range. Abbreviations defined
Measurements
R. r. chilensis
Tarsometatarsus
LENGTH
35.58 + 1.30
33.2 36.7
W-PROX
7.00 + 0.36
6.4 7.5
W-HYPOTS
3.65 + 0.20
3.3 3.8
TRIII-TRIV
5.20 + 0.19
5.0 5.4
TRII-TRIV
5.22 + 0.12
5.1 5*4
W-TRII
1.55 + 0.19
1.3 1.8
W-TRIII
2.52 + 0.15
2.3 2.7
D-TRIII
3.78 + 0.21
3.4 3.9
the methods section.
P. podiceps
R. sp.
P.
40.15 + 2.36
36.3 44.9
36.4

8.09 + 0.57
7.1 8.9
6.7
7.2;
7.5
4.6l + 0.28
4.1 5.1
3.5
4.0;
4.0
6.57 + 0.49
5.8 7.2
5.6

6.24 + 0.49
5-3 7.2
5.2

1.96 + 0.24
1.6 2.4
1.7

2.76 + 0.54
1.5 3.3
2.6

5.00 + 0.4i
4.5 5.6
4.1



226
the living species of this genus. A very small species of teal, Anas sp.
A., is also abundant in the Love Bone Bed local fauna. This species may
represent the smallest species of the genus Anas now known. Other
species of ducks and geese are common in these localities. A tadorine
duck is known from one specimen from the Bone Valley Mining District.
Gallinaceous birds are poorly represented. Two species of turkeys
(Family Phasianidae), each known from a single specimen, are known. One
is referable to the genus Meleagris and the other is not referable to a
genus.
Gruiform birds are also common with 4 species of crane (Family
Gruidae) and b species of rail (Family Rallidae) being known. Of note is
an undescribed species of primitive rail, which is known from abundant
material from the Love Bone Bed and from a few specimens from the McGehee
Farm local fauna. A specimen of a balearicine crane from the Bone Valley
Mining District represents the last occurrence of this subfamily of
cranes in North America. Today this subfamily is only known from Africa
south of the Sahara. Cranes are also shown to be much more diverse in
North America in the Miocene than in Pleistocene or Recent avifaunas.
Two species of flamingos (Family Phoenicopteridae; genus
Phoenicopterus) are known. They are present in the Love Bone Bed local
fauna, the McGehee Farm local fauna and from the Bone Valley. At least 2
different species of flamingo survived in North America until the
Pleistocene.
A jacana, or lily-trotter (Family Jacanidae), Jacana farrandi, is
known from the Love Bone Bed and from McGehee Farm. This distinctive
genus is usually not found north of Mexico today.


78
of Mongolia is based on the distal epiphysis of a left femur.
Phalacrocorax reliquus Kurochkin from the middle Pliocene of western
Mongolia, is based on the distal epyphysis of a right humerus. It has
the same dimensions as P. pelagicus. The relationships of these
cormorants have not been determined.
Phalacrocorax rogersi Howard from the Pliocene Veronica Springs
Quarry, California, is known only from the type coracoid. It is a large
species and appears close to P. perspicillatus and P. pelagicus.
Phalacrocorax owrei Brodkorb and Mourer-Chauvire' from the lower
Pleistocene of Olduvai Gorge, Tanzania, is known from nearly all skeletal
elements. It has been assigned to the subgenus Stictocarbo, although its
tarsometatarsus is rather similar to P. fuscicollis (subgenus
Phalacrocorax). Phalacrocorax tanzaniae Harrison and Walker from the
Pleistocene Bed II of Olduvai Gorge, Tanzania, was described from a
tarsometatarsus and appears close to P. carbo. Phalacrocorax pampeanus
Moreno and Mercerat from the upper Pleistocene Lujan local fauna,
Argentina, was described from the proximal end of a humerus. It is very
close to P. olivaceus and may be ancestral to this recent species
(Howard, 1965). Phalacrocorax gregorii and P. vetustus DeVis from the
upper Pleistocene localities near Lake Eyre, South Australia, were
described from many elements.


Figure 3.2. Location of Included Local Faunas


Order Strigiformes (Wagler, 1830)
191
Ordinal Characters. Tibiotarsus without tendinal bridge, with
ligamental tubercle, condyles prominent, nearly parallel, equal in size,
and circular in lateral view (the Psittacidae, the only other New World
family with a tibiotarsus lacking a tendinal bridge, does not have
condyles which are parallel, equal in size and circular in side view).
Family Tytonidae Ridgway, 1914
Characters. In cranial view, tendinal furrow not excavated (deeply
excavated in Strigidae); in caudal view, shaft merges with posterior
intercondylar sulcus evenly (depression present slightly craniad in
Strigidae).
Tytonidae, undescribed genus
Material. Love Bone Bed local fauna. UF 25926, distal end right
tibiotarsus.
Description. Tibiotarsus (Fig. 4.9) distinguished from all species
of Tyto examined (T. alba, glaucops, capensis, sanctialbani, ostologa,
pollens) and Phodilus badius by having a wider anterior intercondylar
sulcus, with the medial condyle sloping gradually into the anterior
intercondylar sulcus, and by having the area intercondylaris much broader
and more shallow (best seen in anterior view of distal end).
Measurements given in Table 4.29
Remarks. Although the above specimen is slightly abraded and is a
solitary specimen, there is no question that it represents a genus
distinct from the two living genera of this family. Additional
comparisons with owls from the early Tertiary are needed to determine the
exact systematic position of this taxon.


39
are also numerous Pleistocene sites in the Payne Creek Mine (Steadman,
1984), Peace River Mine (=Pool Branch; Webb, 197*0, and Nichols Mine.
The Pleistocene fossil birds from these sites can usually be separated
from geologically older specimens by their association with Pleistocene
fossil land mammals. The Pleistocene fossil birds are not considered in
this study.
There have been many recent studies on non-marine mammalian taxa
from Bone Valley (Baskin, 1982; Berta and Galiano, 1983; Berta and
Morgan, in press; Harrison, 1981; MacFadden and Waldrop, 1980; MacFadden
and Galiano, 1981; MacFadden, 1984; Webb, 1969, 1973, 1983; Webb and
Crissinger, 1984; Wright and Webb, 1984), but no single faunal study of
terrestrial mammals from this local fauna. Reference to the earlier
papers published on the Bone Valley Mining District and its vertebrate
fauna are given by Ray (1957)
Bone Valley taxa which are typical of the late Hemphillian age
include the Eurasian immigrant taxa Agriotherium, Plesiogulo,
'Enhydriodon', and Cervidae. Other taxa present are Rhynchotherium,
Hexameryx, and advanced species of Qsteoborus, Gomphotherium,
Pseudhipparion, Nannippus, and Dinohippus (Berta and Morgan, in press).
Other authors (MacFadden and Galiano, 198l; Berta and Galiano, 1983;
Wright and Webb, 1984) suggest that the upper Bone Valley Formation is
very late Hemphillian because of the presence of taxa equally indicative
of an early Blancan age such as Felis rexroadensis, Meganteron hesperus,
Dinohippus mexicanus, Mylohyus elmorei, Antilocapra (Subantilocapra),
Hemiauchenia, Nasua, and Platygonus.
The birds from Bone Valley have been studied by Brodkorb (1953b,
1953c, 1953d, 1953e, 1955a, 1970). His monograph (1955a) dealt with the


12
7. W-DIST.Transverse width of distal end from the entepicondylar
prominence (Epicondylus ventralis) to the ectepicondylar
prominence (Epicondylus dorsalis).
8. D-DIST.Depth of distal end from cranial face of external
condyle (Condylus dorsalis) through ridge slightly mediad from
external tricipital groove (Sulcus scapulotricipitis), measured
at right angles to the long axis of the shaft.
9. D-ENTEP.Depth of entepicondyle (Epicondylus ventralis) from
attachment of the pronator brevis (Tuberculum supracondylare
ventrale) through entepicondyle (Processus flexoris), measured at
right angles to the long axis of the shaft.
Ulna
1. V-LENGTH.Greatest length from olecranon through tip of
internal condyle (Condylus ventralis).
2. W-SHAFT.Transverse width of midshaft.
3. D-SHAFT.Depth of midshaft.
4. W-PROX.Greatest transverse width of proximal articular
surface.
5* D-LENGTH.Length from tip of olecranon to tip of external
cotyla (Cotyla dorsalis).
6. D-PROX.Depth of proximal end from cranial tip of internal
cotyla (Cotyla ventralis) to caudal margin (Margo caudalis) of
shaft of ulna, measured at right angles to the long axis of the
shaft.
7. ECON.Length from external condyle (Condylus dorsalis) through
ventral face of distal end.


16o
specimens. Hovever, in the original description it was not demonstrated
that Grus nannodes actually belongs in the genus Grus. With the
prevalance of balearicinae-like cranes in the Tertiary of North America,
gruid fossil cannot be uncritically assigned to the genus Grus.
Therefore "Grus nannodes" should be re-examined before its current
taxonomic assignment is accepted.
Grus sp. B
Material. Love Bone Bed local fauna; UF 26002, sternal end right
coracoid; UF 29721, humeral end right coracoid; UF 25740, right scapula;
UF 25722, UF 25737, distal ends left humeri; UF 25749, right
carpometacarpus missing part of shaft of metacarpal III; UF 25753,
proximal end right carpometacarpus; UF 29720, proximal end right femur;
UF 25887 (tent, referred), UF 25893 (tent, referred), UF 25908, UF 25911,
distal ends left tibiotarsi; UF 25885, UF 25896 (tent, referred), UF
29722, distal ends right tibiotarsi; UF 25988, UF 25989, UF29725,
proximal ends left tarsometatarsi; UF 26093, UF 29724, proximal ends
right tarsometatarsi; UF 25857, distal end right tarsometatarsi; UF
25931, UF 25944, UF 25945, UF 29726, distal end left tarsometatarsi.
Remarks. Size similar to that of a large Grus americana or Grus
.japonensis. Coracoids, carpometacarpi, humeri, and femur referred here
on the basis of size, as are the several of the above waterworn
tibiotarsi.
Distal tibiotarsus with characters of the genus Grus, but also
similar to Grus (= Bugeranus) lsucogeranus. Larger than Grus canadensis,
G. grus, and G. monacha, slightly larger tha G. vipio and Grus rubicunda
(G. nigricollis not available). Similar in size to G. americana, G.


Table 4.l6continued
Measurements
P. ridfiwayi
P. chihi
Coracoid
HEAD-CS
12.6; 12.7
13.42 + 0.78
12.3 14.3
W-SHAFT
5-4; 5.+
5.37 + 0.50
4.6 5.9
D-SHAFT
3.7; 4.0
4.45 + 0.38
3.9 4.9
L-GLEN
8.5; 9.0
8.97 + 0.33
8.5 9.4
i
P. falcinellis
Threskiornithinae
Ren. et sp. indet,
13.99 + 0.86
12.2 15.0
16.2
5.52 + 0.54
4.6 6.3
6.1
4.54 + 0.38
4.0 5.1
5.0
9.54 + 0.69
8.1 10.3
11.4
115


15
6. D-HEAD.Greatest depth of femoral head.
7. W-DIST.Greatest transverse width of distal end.
8. W-M&LCON.Transverse width of medial and lateral condyles from
(Crista tibiofibularis) to medial border of medial condyle.
9. W-LCON.Transverse width of lateral condyle.
10. W-L&FCON.Transverse width of lateral condyle and fibular
condyle.
11. D-FCON.Greatest depth of fibular condyle.
12. D-LCON.Greatest depth of lateral condyle.
13. D-MCON.Greatest depth of medial condyle.
Tibiotarsus
1. L-LENGTH.Greatest length from interarticular area (Area
interarticularis) on proximal articular surface through lateral
condyle (Condylus lateralis).
2. M-LENGTH.Greatest length from the most proximal portion of
cnemial crest (Crista cnemialis cranialis) through medial
condyle (Condylus medialis). This includes the patella, if
fused, as in loons and grebes.
3. FIBULAR.Length from interarticular area on proximal articular
surface to the most distal point of fibular crest (Crista
fibularis).
4. W-SHAFT.Transverse width of midshaft.
5. D-SHAFT.Depth of midshaft.
6. W-PROX-M.Transverse width of proximal articular surface from
articular facet for fibular head (Facies articularis fibularis)
to medial border of proximal articular surface.


97
Johnson, 1970) I would also note that Wood used an extremely small
sample size (2-6 individuals per species, many of which were unsexed),
which prevented him from adequately accounting for the considerable
sexual variation which exists in storks.
The fossil species of storks are in critical need of revision. All
fossil species now known were described before Kahl's revisions (1971,
1972) which reduced the number of recent genera from 11 to 6. Because of
the plethora of monotypic genera in this family at the time of the
description of the fossil species, workers have tended to underestimate
the amount of morphological differences within a single genus (sensu
Kahl). Because of this, it is now often impossible to determine
relationships between fossil and recent genera.
A number of species originally described as storks have since been
moved to other families or synomized with other species, or both. These
include Pelargopappus stehlini and P. trouessarti (= Amphiserpentarius
schlosseri; Chauvire, in litt., to Olson, ms), Amphipelargus majori
(Ergilornithidae; Harrison, 1981), Palaeopelargus nobilis,
Xenorhynchopsis tibialis, X* minor, and Xenorhynchus nanus (2 are
flamingos, Rich, 1976; 2 in need of restudy, Olson, ms), and Ibis milne-
edwardsi (= Miophasianus altus; Olson, 197*+b). Lists of the other fossil
storks may be found in Brodkorb (1963c). Most of these are in need of
restudy.
In North America, fossil species of storks include Propelargus
olsoni Brodkorb from the Seaboard Airline Railroad local fauna in
Tallahassee of Barstovian age (see comments on the generic status of
Propelargus in Olson, ms), Ciconia maltha Miller now known from the
Blancan through the Rancholabrean of Idaho, California, and Florida, and


proportionally slightly wider than expected (Figure 4.l). It is very
likely that this material represents a species separate from the living
one. Mycteria wetmorei Howard, 1935, from the Pleistocene of California,
was described on the basis of a lower mandible and is said to be larger
than the living Mycteria americana. As there is unstudied post-cranial
material of this fossil species in many U. S. museum collections, this
material should be examined and Mycteria wetmorei revised before
determining the exact systematic position of the fossil material from
Florida.
Subfamily Ciconiinae Gray, l8h0
Genus Ciconia Brisson, 1760
Ciconia sp. A.
Material. Love Bone Bed local fauna; UF 26102, UF 2967^, distal
ends left tibiotarsi; UF 25906, UF 25909, distal ends right tibiotarsi;
UF 259^6, distal end left tarsometatarsus. UF 29675, distal shaft right
tibiotarsus; UF 25900, distal end left tibiotarsus (tentatively
referred).
Description. Size similar to that of Ciconia ciconia. Distal end
of tibiotarsus agrees with those of Ciconia by having the anterior
intercondylar sulcus broad (narrow in Mycteria) and the distal end not
laterally compressed (laterally compressed in Mycteria). Process for the
ligamental attachment above the distal end of the external condyle is
intermediate between papilla-like (as in Jabir) and crest-like (as in
Ciconia). See Figure U.2 for comparisons with other species of Ciconia.
Distal end of tarsometatarsus agrees with Ciconia in having trochlea
II less rotated ventrally (Mycteria more ventrally rotated). UF 29675


CHAPTER V
PALEOECOLOGY
Introduction
In some situations, the avian specimens from one locality could be
used to reconstruct the fossil avian communities, and the fossil
environments sampled, using methods similar to those of mammalian
paleoecologists (see Shipman, 1981). Each locality could then be
quantitatively compared to other such fossil localities in North America
throughout the latter half of the Cenozoic. Two insurmountable problems,
sample size and collection technique, prevent such an approach being
applied to the localities included in this study.
Localities from which there are relatively few fossils are not
suited, or are severely limited in usefulness, for quantitative
paleoecological analyses because necessary data were never recorded and
because small forms are often poorly represented (Wolff, 1975). He
showed that when collected in a random manner, approximately 12,000 -
25,000 specimens are needed to represent all members of a mammalian
community, and that about 500 identifiable specimens are needed just to
represent the common members of a community. There is no reason to
I
believe that avian communities can be adequately represented by fewer
specimens, certainly when avian communities are as diverse and complex,
if not more so, than mammalian communities. Simply put, avian
communities cannot be reconstructed definitively from a meager handful of
haphazardly collected fossil bird bones. The problem of sample size
applies to all local faunas included in this study except for twothe
198


Table 4.6. Measurements of coracoids and scapulae of the
7 males, 7 females), Phalacrocorax auritus floridanus (N
and Phalacrocorax sp. from the McGehee Farm local fauna,
range. Abbreviations are defined in methods section.
Measurements
P. a. auritus
P. a. floridanus
Coracoid
HEAD-IDA
64.35 + 2.79
59-3 67.9
60.87 + 2.83
56.3 65.6
HEAD-CS
22.74 + 1.08
20.1 -23.8
21.16 + 1.14
19.1 23.7
D-HEAD
8.28 + 0.47
7.6 9.0
7.69 + 0.58
6.5 8.7
W-SHAFT
5.26 + 0.33
4.9 6.0
4.84 + 0.46
4.1 5-5
D-SHAFT
6.21 + 0.47
5.5 7.2
5.36 + 0.66
4.1 6.4
IDA-PP
48.25 + 2.6l
43.4 -51.4
46.04 + 2.07
42.5 49.2
L-GLEN
Scapula
12.38 + 0.46
11.5 13.3
11.68 + 0.57
10.6 12.6
1
W-NECK
6.05 + 0.4l
5.4 6.8
5.34 + 0.37
4.3 -6.0
ACR-GLN
16.46 + 0.98
14.7 18.1
15.26 + 0.94
13.4 17.3
cormorants Phalacrocorax auritus auritus (N = l4,
: 18, 9 males, 9 females), Phalacrocorax wetmorei,
Data are mean +_ standard deviation, (n) and
P. wetmorei
Phalacrocorax sp.
64.38 + 2.60 (4)
60.7 66.8
59.3
22.40 + 0.64 (4l)
21.1 23.8
21.8
8.02 + 0.50 (40)
7.4 9-2
7.0
5.26 + 0.38 (7)
4.8 5.8
5-5
5.16 + 0.30 (7)
4.7 5.6
5.8
48.53 + 2.32 (4)
45.1 50.2
43.6
12.29 + 0.46 (44)
11.3 13.2
12.0
*^J
vO
5.83 + 0.38 (12)
5.1 6.4

15.54 + 1.35 (8)
12.5 16.6



233
Cheneval, J. 1984. Les Oiseaux (Gaviiformes a Anseriformes) du Gisement
Aquitanian de Saint-Ge'rand-le-Puy (Allier, France): Revision
Systematique. Paleovertebrata, l4(2):33-115, 6 fig.
Cracraft, J. 1971a. The humerus of the early Miocene cracid,
Boreortalis laesslei Brodkorb. Wilson Bulletin, 83:200-201.
Cracraft. J. 1971b. Systematics and evolution of the Gruiformes (Class
Aves). 2. Additional comments of the Bathornithidae, with
descriptions of new species. American Museum Novitates, 2449:1-14.
Cracraft, J. 1973. Systematics and evolution of the Gruiformes (Class
Aves). 3. Phylogeny of the suborder Grues. Bulletin of the
American Museum of Natural History, 51:1-127.
Cracraft, J. 1981. Toward a phylogenetic classification of Recent birds
of the world (Class Aves). Auk, 98:681-714.
Cracraft, J. 1982. Phylogenetic relationships and morphology of loons,
grebes, and Hesperornithiform birds, with comments on the early
history of birds. Systematic Zoology, 31:35-56.
Cracraft, J. and P. V. Rich. 1972. The systematics and evolution of the
Cathartidae in the Old World Tertiary. Condor, 74:272-283.
Dali, W. H. and G. D. Harris. 1892. Correlation papers, Neogene. U. S.
Geological Survey. Bulletin 84, 349 pp.
Dixon, W. J. 1981. BMDP Statistical Software. University of California
Press, Berkeley. 725 pp.
Emslie, S. D. 1985. A new species of teal from the Pleistocene
(Rancholabrean) of Wyoming. Auk, 102:201-205.
Feduccia, A. 1968. The Pliocene rails of North America. Auk, 85:441-
453.
Feduccia, A. 1970. A new shorebird from the Upper Pliocene. Journal of
the Graduate Research Center, Southern Methodist University, 38(3
and 4):58-60.
Feduccia, A. 1975. Aves osteology, pp. 1790-1801. In R. Getty (ed.)
Sisson and Grossman's The Anaton^y of the Domestic Animals. W. B.
Saunders, Philidelphia.
Feduccia, A. and S. L. Olson. 1982. Morphological similarities between
the Menurae and the Rhinocryptidae, relict passerine birds of the
Southern Hemisphere. Smithsonian Contribution to Zoology, 366.
22 pp.
Fisher, H. I. 1945. Locomotion in the fossil vulture Teratornis.
American Midland Naturalist, 33:725-742.


3.
. D-PROX.Depth of proximal end from tip of process of
metacarpal I (Processus extensoris) through caudal part of carpal
trochlea (Facies articularis ulnocarpalis), measured at right
angles to the long axis of the shaft.
4. L-MCI.Length metacarpal I (Os metacarpal is alulare) from
process of metacarpal I to pollical facet (Processus alularis).
5. D-SHAFT.Depth of midshaft of metacarpal II ( 0s_ metacarpale
ma.jus).
6. W-SHAFT.Transverse width of midshaft of metacarpal II.
7. D-DIST.Greatest depth of distal end, measured across dorsal
edge of facet for digit II (Facies articularis digitalis major).
8. W-DIST.Transverse width distal end from edge of facet for
digit II through facet for digit III.
Furculum
1. LENGTH.Greatest length, measured from furcular process to
scapular tuberosity.
2. D-PROX.Greatest diameter of coracoidal facet.
Femur
1. M-LENGTH.Greatest length from head of femur (Caput femoris)
through medial condyle (Condylus medialis).
2. L-LENGTH.Greatest length from trochanter (Trochanter femoris)
through lateral condyle (Condylus lateralis).
3. W-SHAFT.Transverse width of raidshaft.
4. D-SHAFT.Depth of midshaft.
5 W-PROX.Transverse width of proximal end, measured from the
head of femur through lateral aspect of trochanter, taken at
right angles to the long axis of the shaft.


Figure 4.1. Plot of the transverse width of the proximal end (W-PROX) versus depth of medial cotyla (D-MCOT)
of the tarsometatarsi of the Recent Mycteria americana and Mycteria sp. A from the Love Bone Bed (L) and
the McGehee Farm (M) local faunas. Sex of Recent individuals is indicated by an arrow (male) or cross
(female) on the symbols.
i


98
Mycteria wetmorei Howard from the Rancholabrean of California and
Florida. These species are also poorly defined on their postcranial
skeleton and are in need of revision.
The fossil species of storks in the late Miocene and early Pliocene
of Florida show Mycteria to be long established (Clarendonian to Recent)
and Ciconia to be more diverse in the past. The relationships between
Ciconia maltha, Ciconia sp. C., and Jabir mycteria should be further
investigated.


D-MCON (mm)
->l 00 (0 o
4^-j i J 1 1 1 L.
i


Table 4.11
continued
Measurements
P. a. auritus
P. a. florida:
D-PROX-M
17.66 + 0.54
16.90 + 0.66
l6.6 18.5
15.5 18.3
D-D-SHAFT
4.59 + 0.34
4.27 + 0.26
4.2 5.3
3.6 4.7
W-DIST
14.64 + 0.36
14.11 + 0.6l
14.0 15.1
12.6 14.9
W-TRII
3.94 + 0.17
3.81 + 0.23
3.7 4.2
3.2 4.0
D-TRII
6.20 + 0.29
5.92 + 0.35
5.7 6.7
5.0 6.3
W-TRIII
4.99 + 0.21
4.78 + 0.38
4.6 5.4
4.0 5.9
D-TRIII
6.89 + 0.25
6.48 + 0.34
6.5 7.3
5.7 6.7
W-TRIV
4.02 + 0.22
3.86 + 0.21
3.6 4.3
3.5 4.2
D-TRIV
8.16 + 0.21
7.84 + 0.35
7.9 8.5
7.0 8.4
P. wetmorei
McGehee Farm
Love
17.54 + 0.72 (15)
l6."iT 19.0
16.1; 16.7

4.54 + 0.21 (30)
3.9 4.9
4.1
4.5
14.42 + 0.49 (25)
13.2 15.8
13.6

4.16 + 0.27 (18)
3.Z 4.5
3.9

6.07 + 0.19 (20)
5.7 6.4
5.6

5.21 + 0.28 (30)
4.3 5.8
4.8
4.9
6.89 + 0.29 (29)
6.1 7.6(7)
6.8
6.7
4.04 + 0.29 (23)
3.17 4.5
3.6
4.3
6.65 + 0.27 (25)
5.9 7.2 (?)
6.2
6.4
00


129
Table 4.17. Measurements of the tibiotarsi and tarsometatarsi of the
vultures Coragyps atratus atratus (N = l6, 8 males, 8 females), Pliogyps
fisheri (holotype) from the Rexroad local fauna, and Pliogyps undescribed
species from the Love Bone Bed. Data are mean _+ standard deviation and
range. Abbreviations are described in the methods section.
Measurements Coragyps a. atratus Pliogyps fisheri Pliogyps sp.
Tibiotarsus
W-DIST-CR
12.79 + 0.4l
12.1 13.6

~
D-MCON
13.67 + 0.4l
13.2 l4.6


Tarsometatarsus
LENGTH
84.43 + 1.54
80.4 87.I
94.0
86.6
W-PROX
15.11 + 0.53
14.1 16.2
21.9
21.1
D-PROX
11.71 + 0.42
11.1 12.4


W-DIST
16.59 + 0.57
15.6 17.6
33.0

W-TRIII
6.43 + 0.21
6.0 6.7
9.6
9.2
D-TRIII
9-99 + 0.32
15.2
13.5


LITERATURE CITED
American Ornithologists' Union. 1983. Check-list of North American
Birds, 6th. edition. 877 pp.
Ammon, L. von 1911. Bayerische Braunkohlen und ihre Verwertung.
Mnich. 82 pp.
Ammon, L. von 1918. TertiSre Vogelreste von Regensburg und die
jungmiBcene Vogelwelt. Abhandl. Naturwiss. Vereines zu Regensburg,
12:1-70.
Arredondo, 0. 1976. The great predatory birds of the Pleistocene of
Cuba. Smithsonian Contributions to Paleobiology, 27:169-187.
Auffenberg, W. 1954. Additional specimens of Gavialosuchus americanus
(Sellards) from a new locality in Florida. Quarterly Journal of the
Florida Acadeiy of Sciences, 17:185-209*
Auffenberg, W. 1963. The fossil snakes of Florida. Tulane Studies in
Zoology, 10:131-216.
Ballmann, P. 1969a. Die VBgel aus der altburdigalen SpaltenfUllung
von Wintershof (West) bei Eichstaett in Bayern. Zitteliana, 1:5-60.
2 taflen.
Ballmann, P. 1969b. Les Oiseaux Mioce'nes de la Grive-Saint-Alban
(isere). Geobios, 2:157-204, 26 fig. pi. 13-15
Baskin, J. A. 1980. Evolutionary reversal in Mylagaulus (Mammalia,
Rodentia) from the late Miocene of Florida. American Midland
Naturalist, 104:155-162.
Baskin, J. A. 1981. Barbourofelis (Nimravidae) and Nimravides
(Felidae), with a description of two new species from the late
Miocene of Florida. Journal of Mammalogy, 62:122-139*
Baskin, J. A. 1982. Tertiary Procyoninae (Mammalia: Carnivora) of North
America. Journal of Vertebrate Paleontology, 2:71-93.
Baumel, J. J., A. S. King, A. M. Lucas, J. E. Breazile, and H. E. Evans.
1979* Nomina Anatmica Avium. Academic Press, London. 637 pp.
Becker, J. J. 1984. Additions to the late Pleistocene avifauna of
Bradenton, Manatee County, Florida. Florida Scientist, 47:201-203.
229


144
Anatini, Genus indet., species B.
Material. Love Bone Bed local fauna; UF 25791, left coracoid; UF
25791, UF 25786, UF 25852, humeral ends coracoids, tentatively referred.
Remarks. Similar morphology as described above for genus and
species indet sp. A, but slightly larger than the coracoids of the males
of A. crecca carolinensis. This material probably represents the same
species as "A" above, but of the opposite sex.
Tribe Aythini (Delacour and Mayr, 1945)
Genus Aythya Boie, 1822
Aythya sp. A
Material. Bone Valley Mining District, Ft. Green Mine (#13
dragline); UF 49695, right carpometacarpus; UF 53945, proximal end right
carpometacarpus; Palmetto Mine; UF 21124, left coracoid, (tentative
referred); Ft. Green Mine (#6 dragline); UF 53866, left coracoid,
(tentatively referred); specific locality unknown, UF 61599, left
coracoid, (tentatively referred).
Remarks. Size similar to that of Aythya collaris. Carpometacarpus
referred to this tribe by lacking the distal swelling on the external rim
of the carpal trochlea (Woolfenden, 1961). Coracoids tenatively referred
here because of general overall similarity with Aythya in characters
(Woolfenden, 1961).
Tribe Mergini (Swainson, 1831)
Genus Bucephala Baird, 1858
Bucephala ossivallis Brodkorb, 1955
Material. Bone Valley Mining District, Palmetto Mine (= locality 2
of Brodkorb, 1955a); PB 172, humeral end left coracoid (holotype).


Table 4.24. Measurements of the tibiotarsi of Balerica pavonica (N = 8,
4 males, 4 females) and Balearicinae, genus and species indet. from the
Bone Valley Mining District. Data are mean + standard deviation and
range. Abbreviation are defined in
or broken.
methods section
Measurement
Balerica pavonica
Balearicinae
Tibiotarsus
W-SHAFT
9.16 + 0.49
8.2 9.8
8.3
D-SHAFT
7.98 + 0.44
7.4 8.6
6.6
W-DIST-CR
19.49 + 0.80
18.1 20.4
15.8
W-DIST-CD
13.10 + 0.79
12.2 l4.4
11.8
D-MCON
19.33 + 0.98
17.4 20.6
*14.0
D-LCON
18.04 + 0.66
17.5 19.0
14.1
D-ICON
10.75 + 0.45
10.1 11.5
8.3


I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
c / 1
Pierce Brodkorb, Chairman
Professor of Zoology
I certify that I have read this study and that in ny opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
udmi
Richard A. Kilti
Assistant Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
S. David Webb
Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as^su-dssBrt^.tion f£>ir~^i^degree of Doctor
of Philosophy.
Ronald G. Wolff
Associate Professor of Zoology
I certify that I have read this study and that in my opinion it
conforms to acceptable standards of scholarly presentation and is fully
adequate in scope and quality, as a dissertation for the degree of Doctor
of Philosophy.
<7
f
Elizabeth
Professor
S. Wing
of Anthropology
This dissertation was submitted to the Graduate Faculty of the Department
of Zoology in the College of Liberal Arts and Sciences and to the
Graduate School, and was accepted for partial fulfillment of the
requirements of the Doctor of Philosophy.
August 1985
Dean for Graduate Studies and Research


Figure 4.7. Dendrocygna sp. A, B. Right coracoid, UF 29765*
A. Dorsal view. B. Ventral view. C, D. Left carpometacarpus, UF
25755 C. Ventral view. D. Dorsal view. Scale = 2.5 cm.


Table 4.7. Measurements of the humeri and ulnae of the cormorants Phalacrocorax auritus auritus (N = 14, 7
males, 7 females), Phalacrocorax auritus floridanus (N = 18, 9 males, 9 females), Phalacrocorax vetmorei
from the Bone Valley Mining District, Manatee Co. Dam (M), and Haile XIXA (H), and Phalacrocorax sp. from
McGehee Farm and the Love Bone Bed. Data are mean _+ standard deviation, (N), and range. Abbreviations are
defined in the methods section.
Measurements
P. a. auritus
P. a. floridanus
P. vetmorei
MCD/HXIXA
McGehee
LBB
Humerus
W-SHAFT
8.04 + 0.50
7.2 8.9
7.16 + 0.47
6.3 7.9
8.09 + 0.36 (18)
7.1 8.6
~
7.4
7.8
D-SHAFT
6.85 + 0.44
6.0 7.7
6.16 + 0.37
5.5 6.8
6.22 + 0.33 (l8)
5.4 7.0

6.2
6.3
W-PROX
22.91 + 1.02
21.0 24.6
21.52 + 1.34
19.7 24.1
23.19 + 0.88 (16)
21.5 24.8
20.4
(H)


D-PROX
7.47 + 0.25
7.1 8.0
6.80 + 0.50
5.9 7.7
6.97 + 0.25 (18)
6.3 7.3
6.4
(H)


D-HEAD
11.10 + 0.54
10.5 11.8
10.21 + 0.75
8.8 11.3
10.85 + 0.18 (10)
10.6~- 11.1
9.5
(H)


L-DELTOID
36.22 + 1.69
33.4 38.8
34.31 + 1.92
31.4 37.6
36.89 + 1.75 (11)
34.1 39-3
34.2
(H)


W-DIST
15.94 + 0.52
15.4 16.8
14.79 + 0.91
12.6 16.4
|
15.63 + 0.43 (37)
14.9 16.8
15.4
(M)
15.2
13.8;
14.9
D-DIST
10.88 + 0.59
10.0 11.9
10.15 + 0.68
8.9 11.8
10.29 + 0.36 (35)
9.4 11.1
10.2
(M)
9.9
9.5
9.5
D-ENTEP
7.25 + 0.25
7.2 7.9
7.10 + 0.45
6.3 8.1
7.18 + 0.25 (39)
6.7 7.7
7.3
(M)
6.8
6.4;
6.9


238
Miller, L. 1929* A new cormorant from the Miocene of California.
Condor, 31:167-172.
Mourer-Chauvire7, C. 1982. Les Oiseaux fossiles des Phosphorites du
Quercy (Eocene superieur a Oligocne superieur): implications
palobiogeographiques. Geobios, Memoire Special 6:413-426.
Murray, B. G., Jr. 1967* Grebes from the late Pliocene of North America.
Condor, 69:277-288.
Murray, B. G., Jr. 1970. A redescription of two Pliocene cormorants.
Condor, 72:293-298.
Neill, W. T., H. J. Gut, and P. Brodkorb. 1956. Animal remains from
four preceramic sites in Florida. American Antiquity, 21:383-395*
Ober, L. D. 1978. The Monkey Jungle, a late Pleistocene fossil site in
southern Florida. Plaster Jacket, 28:1-13.
Olson, S. L. 1973. A classification of the Rallidae. Wilson Bulletin,
85:381-416.
Olson, S. L. 1974a. A reappraisal of the fossil heron Palaeophoyx
Columbiana McCoy. Auk, 91:179-180.
Olson, S. L. 1974b. The Pleistocene Rails of North America. Condor,
76:169-175.
Olson, S. L. 1975a. An evaluation of the supposed Anhinga of Mauritius.
Auk, 92:374-376
Olson, S. L. 1976. A Jacana from the Pliocene of Florida (Aves:
Jacanidae). Proceedings of the Biological Society Washington,
89:259-264.
Olson, S. L. 1977a. The identity of the fossil ducks described from
Australia by C. W. DeVis. Emu, 77:127-131.
Olson, S. L. 1977b. A Synopsis of the Fossil Rallidae. pp. 339-373,
figures 1-26. In_ S. D. Ripley. Rails of the World: A Monograph of
the Family Rallidae. David R. Godine, Boston. 406 pp.
Olson, S. L. 1978. The nomenclatural status of the taxa of fossil birds
attributed to Auguste Aymard. Proceedings of the Biological Society
of Washington, 91:444-449.
Olson, S. L. 198la. The museum tradition in ornithology A response to
Ricklefs. Auk, 98:193-195.
Olson, S. L. 1981b. The generic allocation of Ibis pagana Milne-
Edwards, with a review of fossil Ibises (Aves: Threskiornithidae).
Journal of Vertebrate Paleontology, 1:165-170.


73
more produced. Fovea carpalis caudalis deeper in P. wetmorei than in P.
auritus.
The femora are similar, except that the popliteal fossa is generally
less excavated than in P. auritus. Brodkorb's statement that the femur
is longer and narrower than that of P. auritus is not supported by the
larger sample size now available.
Tibiotarsus with no obvious qualitative morphological differences,
but see Table 4.10 for a few minor quantitative differences.
Tarsometatarsus with a lateral face flat, similar to but not as
extreme as, that found on specimens from the Love Bone Bed local fauna.
Other characters similar. See Table 4.11 for measurements.
Remarks. See comments under Family Remarks (below) pertaining to P.
auritus and related species.
Phalacrocorax cf. P. idahensis
Material. Bone Valley Mining District, Palmetto Mine (locality 2 of
Brodkorb, 1955); PB 311, proximal end left ulna.
Remarks. This proximal ulna (PB 311) is larger than that of other
specimens of P. wetmorei presently known from Bone Valley. Since it was
first reported by Brodkorb (1955), it has been additionally damaged in
transport and is now barely diagnostic at the generic level. Unless
additional material becomes available, the status of this enigmatic
record in Florida will probably never be satisfactorily resolved.
Remarks on the Family Phalacrocoracidae.
The cormorants have an extensive fossil record. Brodkorb (1963c)
lists 23 paleospecies; subsequently eight more have been described, or
are in the process of being described. Most have been referred to the


109
Olson (1981b) notes, doing this would cast considerable doubt on the
validity of Eudocimus peruvianus Campbell. Campbell (1979) states that
E. ruber differs more from E. albus and E. peruvianus than the latter two
do from each other. It is very likely that when a larger series of
recent E. ruber and E. albus are examined and compared with E.
peruvianus, there will be no consistent ostelogical differences between
these three species.
Similarily, Plegadis falcinellus and P. chihi are very similar
osteologically and may possibly be conspecific (see Table 4.l6 for
measurements; Palmer, 1962; Mayr and Short, 1970 for comments).


177
"paratype"), PB 146, PB 300, distal ends right tarsometatarsi (=
"paratype"). New material (referred by Becker); PB 139, proximal end
right carpometacarpus; PB 7980, distal end left tibiotarsus. Nichols
Mine; UF 24623, distal end right tibiotarsus (tentatively referred); Ft.
Green Mine (# 13 dragline), UF 52422, distal end right tibiotarsus; Payne
Creek Mine, UF 67811, distal end right tibiotarsus; No Specific Locality;
UF 29743, distal end right tibiotarsus; Palmetto Mine; UF 21164, distal
end right tarsometatarsus.
Description/ Remarks. For qualitative characters and remarks see
Brodkorb (1953b, 1955a) Measurements given in Table 4.26.
Phoenicopterus sp. A
Material. Love Bone Bed local fauna; UF 25905, UF 25907, UF 29685,
UF 29744, distal ends right tibiotarsi (large); UF 25882, UF 25883, UF
25892, UF 25898, UF 25910, UF 29684, distal ends left tibiotarsi (large);
UF 29686, distal end right tibiotarsus (small); UF 25889, UF 25899, UF
25927; distal ends left tibiotarsi (small); UF 25881, UF 25897, distal
ends right tibiotarsi (abraded); UF 25895, UF 29682, UF 29683, distal
ends left tibiotarsi (abraded); UF 29679, proximal end right
tarsometatarsus; UF 25859, UF 25864, distal ends right tarsometatarsi;
UF 25932, UF 25935, UF 29680, UF 29681, distal ends left tarsometatarsi.
McGehee Farm local fauna; UF 11103, distal end right
tarsometatarsus.
Description. Tibiotarsi with a pronounced bimodal size
distribution, less so in the tarsometatarsi. The group of small
tibiotarsi cannot be distinguished from the tibiotarsi of P. floridanus
when specimens are directly compared. The group of large tarsometatarsi


Table 3.1. A partial list of Bone Valley mines, their mine codes,
approximate location, and the stratigraphic codes commonly used.
Mines
Codes
Township Range
Sections
Chicora
BVC
District Grade
BVPC
see
Payne Creek Below
Estech
BVS
see
Swift Below
Ft. Green
BVFG
32S
23E
2,3,10-14,22-24
Ft. Meade
BVFM
31S
25E
2,13
Gardinier
BVG
32 S
24,24E
unknown
Hooker's Prairie
BVHP
31S
24e
17,18,20,28-30
Kingsford
BVK
31S
23E
3
New Palmetto
BVNP
32S
24E
3
Nichols
BVN
30S
23E
19,28,29
Palmetto
BVP
32S
2UE
9,10,15,16,21,22
Payne Creek
BVPC
32S
24E
13,14,23,24,29-32
Peace River
BVFM
see
Ft. Meade Above
Swift
BVS
31S
24E
14
Tiger Bay
BVTB
31S
24e
12
Stratigraphic Codes
Explaination
0
No
stratigraphic data
1
In place
Hawthorn Fm. dolomitic
2
In place
"lower Bone Valley
Fm."
3
In place
"upper Bone Valley
Fm."
4
In place
Pleistocene sediments
5
Soil zone
: (upper clay)


Table 4.19continued
Measurement
D. viduata
D. arbrea
C arpometacarpus
LENGTH
52.95 + 2.06
51.5 57.7
59.70 + 2.77
56.5 61.3
W-CARPAL
4.79 + 0.20
4.5 5.1
5.57 + 0.06
5.5 5.6
D-PROX
10.60 + 0.26
10.2 11.0
11.70 + 0.20
11.5 11.9
D-SHAFT
3.00 + 0.18
2.8 3.3
3.43 + 0.06
3.4 3.5
W-SHAFT
3.35 + 0.12
3.2 3.5
3.97 + 0.12
3.9 4.1
W-DIST
6.08 + 0.31
5.5 6.5
6.60 + 0.30
6.3 6.9
i
D. bicolor D. autumnalis Dendrocygna sp.
51.93 + 1.51
49.5 54.1
54.63 + 2.64
50.0 58.5
55.0
4.54 + 0.23
4.1 4.8
4.90 + 0.19
4.6 5.1
5.2; 5.4
10.66 + 0.29
10.2 ll.O
11.30 + 0.51
10.7 12.2
11.3; 11.5
3.13 + 0.19
2.9 3.5
3.16 + 0.24
2.9 3.5
3.4; 3.6
3.38 + 0.21
3.1 3.7
3.70 + 0.26
3.3 4.1
4.0; 4.4
5.76 + 0.31
5.4 6.1
6.29 + 0.36
6.0 6.9
6.2
152


Table 4.27. Measurements of the tibiotarsi and tarsometatarsi of the Recent Phoenicopterus jamesi (N = 9, 1
male, 2 females, 6 unsexed), Phoenicopterus chilensis (N = 8, 3 males 2 females, 3 unsexed), Phoenicopterus
ruber (N = 15, 6 males, 7 females, 2 unsexed), and Phoenicopterus minor (N = 7, 3 males, 4 females).Data
are mean _+ standard deviation (number) and range. Abbreviations defined in the methods section.
Measurement
P. ,1amesi
P. chilensis
P. ruber
P. minor
Tibiotarsus
W-DIST-CR
14.49 + 0.67
13.6 15.4
14.96 + 0.65
14.2 16.3
15.70 + 0.87
14.6 17.2
12.79 + 0.65
12.0 13.6
W-DIST-CD
10.21 + 0.39
9.6 10.8
10.26 + O.65
9-5 11.7
10.93 + 0.70
9.8 12.2
8.85 + 0.46
8.0 9.3
D-MCON
15.79 + 0.87
14.3 16.8
16.17 + 0.64
15.4 17.4
18.70 + 1.15
16.3 20.8
14.33 + 0.67
13.2 15.1
D-LCOT
15.29 + 0.69
13.9 16.3
16.29 + 0.53
15.5 17.1
18.80 + 1.19
l6.6 21.2
14.23 + 0.68
13.1 15.0
Tarsometatarsus
W-TRIII
7.09 + 0.44
6.3 7.7
7.55 + 0.38
7.2 8.4
7.85 + 0.50
7.2 8.8
6.31 + 0.40
5.7 6.8
TRIII-TRIV
II.92 + 0.49
11.2 12.5
12.47 + 0.37
11.9 13.1
13.91 + 0.86
12.6 15.9
11.15 + 0.65
10.3 12.0
TRII-TRIV
i4.oi + 0.64
13.2 15.0
14.13 + 0.97
13.0 15.9
14.99 + 0.80
13.7 16.5
12.22 + 0.77
11.6 13.5
i
oo