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


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The fossil birds of the late Miocene and early Pliocene of Florida
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ix, 245 leaves : ill. ; 28 cm.
Becker, Jonathan J., 1955-
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Subjects / Keywords:
Birds, Fossil   ( lcsh )
Paleontology -- Florida -- Miocene   ( lcsh )
Paleontology -- Florida -- Pliocene   ( lcsh )
Zoology thesis Ph. D
Dissertations, Academic -- Zoology -- UF
bibliography   ( marcgt )
theses   ( marcgt )
non-fiction   ( marcgt )


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

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University of Florida
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All applicable rights reserved by the source institution and holding location.
Resource Identifier:
aleph - 021166423
oclc - 12891016
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Full Text







Copyright 1985


Jonathan J. Becker


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


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.


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

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



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 . . . . . . . .


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





Introduction . . .
Local Faunas . . .


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


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



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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



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+),


?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


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.



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:


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?


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


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


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.,


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)


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.


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)


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,

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


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,

Holman, 1961; Wetmore, 1931

Storer, 1976b

Steadman, 1980

Steadman, 1980

Steadman, 1980

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

Table 1.1--continued


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.


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



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).


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.


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


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


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).


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

humeri) through the midpoint of the lateral condyle (Condylus


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


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


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.


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


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


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


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



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.


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


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.


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

scapular tuberosity.

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


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


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

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

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


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


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.


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.


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.





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.




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.


1 13 12


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



A N \~10



2- 1


13 12


Is-- 7
F/19- E

20 67 :8~

S22 21 17 18 MED
k- 16


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.


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



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.





University of Nebraska State Museum

United States National Museum

Yale Peabody Museum



1. f.










Biomedical Statistical Program, P-series

local fauna


megannum (or million years)




million years before present

number (of specimens)

North American Land Mammal Age



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

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


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.


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.



2 >w





Figure 3.2. Location of Included Local Faunas.



7 SR-64

0 CITRUS ----




| ~7"


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



District Grade


Ft. Green

Ft. Meade


Hooker's Prairie


New Palmetto



Payne Creek

Peace River


Tiger Bay

Stratigraphic Codes






































Range Sections

Payne Creek Below

Swift Below










Ft. Meade Above















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


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


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


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


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


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,


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.













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.









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.










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.










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.












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.










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












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


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


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.


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.


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


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.


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.-
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.


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


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















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
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8.40 + 0.50
7.5 9.2

9.78 + 0.57
8.7 10.5

8.59 + 0.48
7.6 9.4

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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
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6.76 + 0.28
6.3 7.2


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10.8 12.3

2.99 + 0.20
2.7 3.3

6.85 + 0.32
6.4 7.5

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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


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


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