THE FOSSIL BIRDS OF THE LATE MIOCENE AND EARLY PLIOCENE
JONATHAN J. BECKER
A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN
PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
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
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
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
TABLE OF CONTENTS
ACKNOWLEDGMENTS . . . . . . . . . . ...
ABSTRACT . . . . . . . . . . . . . .
I. INTRODUCTION AND PREVIOUS WORK . . . . . . .
Introduction . . . . . . . . . ...
Limitations of Study . . . . . . . . ...
Previous Work . . . . . . . . . ....
II. METHODS . . . . ........ . . .
Measurements . . . . . ... .
Computer Software . .. . . . . . ....
Nomenclature . .. . . . . .
Systematics . ...... . . . .
Paleoecology . . . .. . .............
Biochronology and Faunal Dynamics . . . . . .
Specimens Examined . . . . . . . . . .
Abbreviations . . . . . . . . . . .
III. GEOLOGY . . . . . . . . . . . .
Biochronology . . . . . . . . . ...
Local Faunas . . . . . . . . .....
Eustatic Sea-level Changes . . . . . . . .
IV. SYSTEMATIC PALEONTOLOGY . . . . . . . . .
Order Podicipediformes .
Order Pelecaniformes .
Order Ciconiiformes .
Order Anseriformes . .
Order Galliformes . .
Order Ralliformes . .
Order Strigiformes . .
Order Passeriformes .
.......... . . 49
. . . . . . . . .. 65
. . . . . . . . 90
S. . . . . . . 116
. . . . . . . . 136
. . . . . . . . . 155
* . . . . . . . . 158
. . . . . . 175
S. . . . . . . . 191
. . . . . .. . .. 195
V. PALEOECOLOGY . . .
Introduction . . .
Local Faunas . . .
VI. BIOCHRONOLOGY AND FAUNAL
Introduction . . .
Faunal Dynamics ...
Biochronology . . .
VII. SUMMARY . . ..
Systematics . . .
Paleoecology . . .
Biochronology . . .
LITERATURE CITED . . .
BIOGRAPHICAL SKETCH . . .
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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
THE FOSSIL BIRDS FROM THE LATE MIOCENE AND EARLY PLIOCENE
Jonathan J. Becker
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,
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.
INTRODUCTION AND PREVIOUS WORK
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.
The earliest published report of fossil birds from Florida is
Sellard's (1916) description of a supposed jabiru (Jabiru? weillsi
=Ciconia maltha) from Vero, closely followed by Shufeldt's (1917a, 1917b)
study of this local fauna. Wetmore (1931) reviewed the Pleistocene
avifauna of Florida and was also the first to report (19h3) on the
Tertiary birds from Thomas Farm and the Bone Valley Mining District.
Numerous studies have appeared since then, primarily by Brodkorb or his
students. A few other faunal and systematic studies have also included
avian material from Florida.
Table 1.1 lists many of the fossil localities in Florida which have
a notable record of fossil birds. Reference to Brodkorb's Catalogue of
Fossil Birds (1963-1978), where the avifaunas from many Florida
Pleistocene localities were first reported, is omitted to conserve space.
Table 1.1. Avian Fossil Localities of Florida.
Locality Reference (s)
MIOCENE TO EARLIEST PLIOCENE (Pre-Blancan)
Bone Valley, Polk Co. Becker, 1985a; Brodkorb, 1953b, 1953c,
1953d, 1953e, 1955a, 1970; Olson, 1981b;
Steadman, 1980; Wetmore, 1943; this study
Haile VI, Alachua Co.
Haile XIXA, Alachua Co.
Love Bone Bed, Alachua Co.
Manatee Co.Dam Site
McGehee Farm, Alachua Co.
Mixson Bone Bed, Levy Co.
Seaboard Airline Railroad,
SR-64, Manatee Co.
Thomas Farm, Gilchrist Co.
Withlacoochee River 4A,
LATE PLIOCENE (Blancan)
Haile XVA, Alachua Co.
Santa Fe IB, Gilchrist Co.
Brodkorb, 1963a; this study
Becker, 1985a, 1985b; Webb et al., 1981;
Webb and Tessman, 1968; this study
Brodkorb, 1963a; Hirschfeld and Webb, 1968;
Olson 1976; 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
Locality Reference (s)
EARLY PLEISTOCENE (Irvingtonian)
Coleman IIA, Sumter Co. Ritchie 1980; Steadman, 1980
Haile XVIA, Alachua Co. Steadman, 1980
Inglis IA, Citrus Co. Carr, 1981; Ritchie, 1980; Steadman, 1980
Santa Fe River IIA, Steadman, 1980
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
Bowman IA, Putnam Co.
Bradenton, Manatee Co.
Coleman III, Sumter Co.
Crystal Spring Run,
Davis Quarry, Citrus Co.
Econfina River, Taylor Co.
Florida Lime Company,
Haile IA, Alachua Co.
Haile IIA, Alachua Co.
Haile VIIA, Alachua Co.
Becker, 1984; Steadman, 1980; Wetmore,
Brodkorb, 1955b; Holman, 1961
Brodkorb, 1953a, 1954b; Olson, 1974b, 1977b
Holman, 1961; Steadman, 1980
Haile XIB, Alachua Co.
Hog Cave, Sarasota Co.
Hog Creek, Manatee 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,
Santa Fe River IA,
Santa Fe River IVA,
Ligon, 1965; Olson, 1974b
Steadman, 1980; Wetmore, 1931
Campbell, 1980; McCoy, 1963; Olson, 1974a,
1974b, 1977b; Storer, 1976b; Wetmore, 1931
Holman, 1961; Storer, 1976b
Holman, 1961; Steadman, 1980; Wetmore, 1931
Holman, 1961; Steadman, 1980
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
Holman, 1961; Olson, 1974b; Steadman, 1980;
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.
Zuber, Marion Co.
Cotton Midden, Volusia Co.
Castle Windy Midden,
Green Mound Midden,
Nichol's Hammock, Dade Co.
Silver Glenn Springs,
Summer Haven Midden,
St. Johns Co.
Vero (Stratum 3),
Indian River Co.
Holman, 1961; Sellards, 1916; Shufeldt,
1917; Steadman, 1980; Storer, 1976b; Weigel,
1962; Wetmore, 1931
Hay, 1902; Neill et al., 1956
Hirschfeld, 1968; Steadman, 1980
Neill et al., 1956; Steadman, 1980
see Vero, Stratum 2, above
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
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
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
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
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
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
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
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.
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
20 67 :8~
S22 21 17 18 MED
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.
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.
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
Biomedical Statistical Program, P-series
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.
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
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.
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.
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.
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
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
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,
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.
MANATEE CO. ISR-64
DAM L.F. L.F.
RIVER 4A L.F.
Figure 3.2. Location of Included Local Faunas.
ALACHUA 1 MCGEHEE
2 HAILE SITES
1 3 LOVE BONE BED
02 e 4 MIXSON BONE BED
4* 5 WITHLACOOCHEE
LEVY 6 BONE VALLEY
8 MANATEE CO. DAM
0 CITRUS ----
^ MANATEE \
Table 3.1. A partial list of Bone Valley mines, their mine codes,
approximate location, and the stratigraphic codes commonly used.
Payne Creek 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
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
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
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
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
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
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
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).
Rollandia sp. (LOV, MIX, MCG)
Tachybaptus sp. (LOV, MCG)
Podilymbus cf P. podiceps (BV)
Podilymbus sp. A (MIX)
Podiceps sp. (BV)
Pliodytes lanquisti (BV)
Phalacrocorax sp. A (LOV, MCG, H19A)
Phalacrocorax wetmorei (BV, MD, SR-64)
Phalacrocorax cf. idahensis (BV)
Anhinga grandis (LOV, MCG, H19A)
Anhinga sp. (BV)
Ardea polkensis (BV)
Areda sp. indet.(LOV)
Egretta sp. indet. (LOV, BV)
Egretta subfluvia (WITH hA)
Ardeola sp. (LOV)
Nycticorax fidens (MCG)
Mycteria sp. (LOV,MCG)
Ciconia sp. A (LOV)
Ciconia sp. B (MIX,BV)
cf. Ciconia sp. C (BV)
Eudocimns sp. A. (BV)
Plegadis cf. P. pharangites (LOV)
Threskiornithinae, genus et species indet. (LOV)
Order Falconiformes (auct.)
Pliogyps undescribed sp. (LOV)
Pandion lovensis (LOV)
Pandion sp. (BV)
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)
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)
Meleagridinae, genus indet. (LOV)
Meleagris sp. (BV)
Order Gruiformes (auct.)
Grus sp. A (LOV)
Grus sp. B (LOV)
Balearicinae, genus indet. (BV)
Aramornis (cf.) (LOV)
Rallus sp. A (LOV)
Rallus sp. B (BV)
Rallus (cf.) sp. C (LOV)
Undescribed genus (LOV, MCG)
Phoenicopterus floridanus (BV)
Phoenicopterus sp. A (LOV, MCG)
Jacana farrandi (LOV, MCG)
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)
Undescribed genus (LOV)
Bubo sp. (BV)
Suborder indet. sp. A (LOV)
Suborder indet. sp. B (LOV)
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
2.47 + 0.12
5.35 + 0.23
25.45 + 0.71
24.50 + 0.68
7.03 + 0.23
2.17 + 0.10
2.30 + 0.18
1.65 + 0.10
8.53 + 0.38
4.55 + 0.16
2.39 + 0.15
2.11 + 0.15
5.00 + 0.31
21.80 + 1.57
21.17 + 1.26
6.10 + 0.34
2.17 + 0.17
1.94 + 0.21
1.29 + 0.09
7.44 + 0.41
4.31 + 0.25
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
Abbreviations defined in the methods section.
R. r. chilensis
35.58 + 1.30
7.00 + 0.36
3.65 + 0.20
5.20 + 0.19
5.0 -- 5.4
5.22 + 0.12
1.55 + 0.19
2.52 + 0.15
3.78 + 0.21
40.15 + 2.36
8.09 + 0.57
4.61 + 0.28
6.57 + 0.49
6.24 + 0.49
1.96 + 0.24
2.76 + 0.54
5.00 + 0.41
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
32.73 + 1.64
2.80 + 0.15
3.18 + 0.24
7.65 + 0.39
3.30 + 0.24
8.08 + 0.44
5.97 + 0.41
6.03 + 0.24
4.40 + 0.24
25.03 + 1.53
26.81 + 1.61
2.53 + 0.22
2.67 + 0.30
6.53 + 0.30
2.74 + 0.20
6.84 + 0.53
4.96 + 0.40
4.97 + 0.37
3.44 + 0.26
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
68.69 + 4.31
80.17 + 5.27
4.50 + 0.41
3.29 + 0.27
7.00 + 0.44
7.10 + 0.65
6.04 + 0.37
6.86 + 0.44
6.76 + 0.49
4.29 + 0.33
P. cf. podiceps
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
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
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
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
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
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
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
56.58 + 2.63
6.48 + 0.36
8.15 + 0.50
16.13 + 0.60
7.02 + 0.33
15.71 + 0.59
12.21 + 0.65
3.34 + 0.27
7.25 + 0.25
8.81 + 0.47
10.36 + 0.37
9.05 + 0.34
P. a. floridanus
52.12 + 3.15
54.03 + 2.51
5.94 + 0.38
7.38 + 0.46
14.56 + 0.85
6.55 + 0.33
14.93 + 0.84
11.38 + 0.77
3.02 + 0.33
8.40 + 0.50
9.78 + 0.57
8.59 + 0.48
55.46 + 1.58 (9)
57.58 + 1.45 (12)
6.49 + 0.29 (17)
8.04 + 0.41
7.3 -- 9.0
15.43 + 0.59 (21)
6.76 + 0.28
15.11 + 0.47 (17)
11.59 + 0.39 (14)
2.99 + 0.20
6.85 + 0.32
8.68 + 0.40
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I I E E
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
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