The avifauna of the Bone Valley formation ( FGS: Report of investigations 14 )

STATE OF
STATE BOARD OF
ERNEST MITTS

FLORIDA
CONSERVATION
i, Director

FLORIDA GEOLOGICAL SURVEY
HERMAN GUNTER, Director

REPORT OF INVESTIGATIONS
No. 14

THE AVIFAUNA OF
THE
BONE VALLEY FORMATION

By
Pierce Brodkorb
Department of Biology
University of Florida
Gainesville, Florida

Published for the
Florida Geological Survey

TALLAHASSEE, FLORIDA
1955

:I
i-
'i I.-
.
I~ `

AGRI,

CULTURAL
FA S E RAR
FLOI II)A STATE IIOA ID

OF

CO,{NSElIVATION

LeROY COLLINS
Governor

R. A. GRAY
Secretary of State

J. EDWIN LARSON
Treasurer

NATHAN MAYO
Commissioner of Agriculture

THOMAS D. BAILEY
Superintendent Public Instruction

RAY E. GREEN
Comptroller

ERNEST MITTS
Director

RICHARD ERVIN
Attorney General

LETTER OF TRANSMITTAL

August 20, 1955

Mr. Ernest Mitts, Director
Florida State Board of Conservation
Tallahassee, Florida
Dear Mr. Mitts:
The Bone Valley formation is the source of most of the com-
mercial phosphate in Florida. The stratigraphic relationship of the
formation to older and younger formations is being determined
through studies being conducted by the Atomic Energy Commission
in cooperation with the United States Geological Survey. The de-
termination of the age of the Bone Valley formation is important
economically to the phosphate industry in that the time of formation
of the phosphate can thus be determined and a possible lead to
future prospecting and an expansion of the phosphate reserve may
be obtained.
This paper, "The Avifauna of the Bone Valley Fo mation," by
Dr. Pierce Brodkorb of The Department of Biology, University of
Florida, Gainesville, contributes additional data on the age of the
Bone Valley formation. The data will be welcomed by geologists,
stratigraphers, and ornithologists of the State. We are pleased to
publish this contribution to Florida stratigraphy as Report of In-
vestigations No. 14.
Very truly yours,

Herman Gunter, Director

Printed by Rosr PRINTING COMPANY, TALLAHASSEE, FLORIDA

CONTENTS

Letter of Transmittal .....
Introduction ..............
Acknowledgments ..
Location ........
Stratigraphy .......

. . .. . . . . .. .. . . iii
. . . . . . . . . ..

. .o .. . .. . . ..1.. .. .. 2
. .. .. .. .. 1. .

List of Species ................................................ 4
Gavia palaeodytes Wetmore .............................. 5
Gavia concinna Wetmore ................. ..... ...... 5
Pliodytes lanquisti Brodkorb ............................. 6
Diomedea anglica Lydekker ............................. 7
Morus peninsularis new species ................. ........ 8
Sula guano new species .............. .................. 9
Sula phosphata new species ............................... 11
Phalacrocorax wetmorei new species ........................ 12
Phalacrocorax idahensis (Marsh) ..........................14
Ardea polkensis new species .............................. 17
Phoenicopterus floridanus Brodkorb ....................... 18
Bucephala ossivallis new species .........................18
Palostralegus sulcatus new genus and species ................ 19
Calidris pacis new species ..................................22
Erolia perpusilla new species ............................ 23
Lim osa sp. ...............................................24
Larus elmorei Brodkorb ....................................25
Australca grandis new genus and species ................... 25

Age of the deposit ...............................

................29

Paleoecology ..................................................33
Census .................................................. 33
Dominance of species ...................................33
Habitat requirements ....................................33
Effect of bird life on the production of phosphate......... 36

Sum m ary ................................... ............ ......38
Literature cited ...................................... .. .. ....55

ILLUSTRATIONS

Tables Page
1. Section at Locality 1 ........................................ 3
2. Section at Locality 2 ...................................... 4
3. Measurements of humerus in Gavia ....................... 6
4. Measurements of coracoid of Morus and Sula ................ 13
5. Measurements of Phalacrocorax wet-morei ............... 16-17
6. Ratios of wing elements to length of Coracoid ............ 29
7. Proportion of extinct species in Quarternary
and late Tertiary Avifaunas ..............................31
8. Census of birds of the Bone Valley formation ................ 34
Plates 1- 11 ............................................. 41 -53

THE AVIFAUNA OF THE BONE VALLEY FORMATION
Pierce Brodkorb
INTRODUCTION

Lying unconformably upon the lower Miocene Hawthorn forma-
tion and covered by Pleistocene terrace sands is the Bone Valley
gravel, named by Matson and Clapp (1909: 138), to which the term
Bone Valley formation was later applied by Cooke (1945: 203). In
Polk and Hillsborough counties, in southwestern Florida, this forma-
tion is being exploited through extensive phosphate mining operations.
Remains of fossil vertebrates were first reported from the Bone
Valley phosphate by Leidy (1889). Sellards (1915:73) reported a
new species of gavial, a large land tortoise, several large land mam-
mals, cetaceans, and teeth of elasmobranchs. The pelagic mammals
have been worked up by Allen (1921), Hay (1922), Kellogg (1924),
Simpson (1932), and Case (1934). The land mammals, comprising
a rather extensive fauna, were studied by Simpson (1930) and by
White (1941, 1942).
The birds heretofore known from this deposit consist of four
fragmentary bones of three species, preserved in the Museum of
Comparative Zoology (Wetmore, 1943). Until the present study was
undertaken they represented all that was known of the avifauna
attributed to the Pliocene east of the Mississippi River.

In November 1951 George C. Elmore began to collect bird ma-
terial from two localities near Brewster, Florida. Thanks to his
interest and diligence, about 200 bird bones were assembled, so that
the Bone Valley formation now has the largest known avifauna of
Tertiary age in this country, both as to species and specimens. New
species of grebe, flamingo, and gull in this collection have been
described in previous papers (Brodkorb, 1953A, 1953B, 1953D).
These forms, as well as Wetmore's records, are included in the pres-
ent report in order to present a complete survey of the avifauna of
the Bone Valley formation. About five specimens not identified gen-
erically are omitted. All holotypes are retained in the Brodkorb col-
lection at the University of Florida.

Acknowledgments. Thanks are due George C. Elmore, who
assembled the collection. Valuable advice was given by Dr. Robert
0. Vernon and Herbert Winters, Florida Geological Survey; by E. C.
Pirkle, Jr., Department of Geology, University of Florida; and by

FLORIDA GEOLOGICAL SURVEY

John L. Rich, Department of Geology, University of Cincinnati.
James B. Cathcart, United States Geological Survey, generously sup-
plied stratigraphic sections of the two collecting localities. Drs. Loye
Miller and Alden H. Miller kindly made a comparison of specimens
with the type of Limosa vanrossemi at the University of California
Museum of Paleontology.
For the loan of recent or fossil comparative material I am in-
debted to Drs. Herbert Friedmann and Alexander Wetmore, United
States National Museum; Dr. Hildegarde Howard, Los Angeles Coun-
ty Museum; S. J. Olsen, Museum of Comparative Zoology; Dr. Frank
A. Pitelka, Museum of Vertebrate Zoology, University of California;
and Dr. Harrison B. Tordoff, Museum of Natural History, University
of Kansas.
The drawings are the work of Miss Esther Coogle, of the University
of Florida staff. The photographs of a bird rookery were supplied by
Dr. Ernest H. Lund, Department of Geology, Florida State University.

Location. Bird fossils were collected at two localities on holdings
of the American Agricultural Chemical Company, with headquarters
at Pierce, Florida. Both localities lie somewhat south of the post
office of Brewster, in southwestern Polk County, Florida.

Locality 1 is an area of about five acres in extent in the center
of sec. 32, T. 31 S., R. 24 E., or about two and one-half miles east-
southeast of Brewster; surface elevation about 145 feet above sea
level.

Locality 2 is in the N 1/2 NE 4 NW /4, sec. 5, T. 32 S., R. 24 E.,
or about three miles southeast of Brewster; elevation about 145 feet.

I have not been able to ascertain the exact locality at which Dr.
Theodore E. White collected the four bird specimens reported by
Wetmore. They were said to have been collected near Pierce from
screenings of the company, and may or may not have come from
one of Elmore's localities. It is my understanding that some of El-
more's material was also obtained from screenings.

Stratigraphy. Stratigraphic sections taken by James B. Cathcart
at the two Elmore localities are given in Tables 1 and 2. Bed 6 at
Locality 1 is the equivalent of Beds 7 and 8 at Locality 2. It is in this
lowest part of the Bone Valley formation that vertebrate fossils occur.
They are white in color and are heavily mineralized.

REPORT OF INVESTIGATIONS No. 14

In these sections the bottom of the Bone Valley lies at an alti-
tude of about 102-104 feet above sea level, and the top of the forma-
tion lies at about 128-134 feet above sea level. If no actual movement
of the land took place, sea level at the beginning of Bone Valley
time must have been at least 104 feet higher and at the close of
Bone Valley time at least 134 feet higher than at present.

Table 1.-SECTION AT LOCALITY 1: CENTER OF SEC. 32, T. 31 S., R. 24 E.; SURFACE
ELEVATION ABOUT 145 FEET ABOVE SEA LEVEL.

Depths
Bed in
Feet

3) 17-24

7) 41-44 2

Sand, brown
loose. ... .

and black, slightly carbonaceous,

Sand, brown and white, some iron-cemented
lumps, slightly clayey ..................... ..
SLithologic break -- -----
Sand, gray-green and white, slightly clayey, but
less clay than bed 2; minor aluminum phosphate
as cement.. ...............................

Sand, gray-green, slightly clayey, with 10% cal-
cium phosphate nodules, coarse sand to gran-
ule size.....................................
-- Base of the "Overburden" of the Company--
Sand, gray, with thin interbeds of greenish clayey
sand. Contains thin lens-like beds of highly
phosphatic sand. Phosphate is about 15% of
matrix ...................................

Sand, gray, loose, cross-bedded, with 45-50% of
black and brown phosphate, sand to gravel size
Unconformity (Base of "Matrix") ---
Sand, slightly clayey, blue-gray; contains abund-
ant borings filled with material from bed 6.
Some black and brown phosphate nodules. Phos-
phate is 15 % of matrix..................

Surface soil,
Recent or
Pleistocene

Pleistocene?
terrace sands

Upper Bone
Valley
formation

Lower Bone
Valley
formation

Miocene:
Hawthorn
formation

_--- ------_ --- Base of Exposure -------------

0-5

5-17

4)

5)

6)

24-31

31-34

34-41

I

'I --- -

FLORIDA GEOLOGICAL SURVEY

Table 2.---SECTION AT LOCALITY 2: N NE NW4 SEC. 5, T. 32 S., R. 24 E.; SUR-
FACE ELEVATION ABOUT 145 FEET ABOVE SEA LEVEL.

Depths
l'd in
Feet

1) 0-8

2) 8-11

3) 11-13,2

4) 13 --18 '/

5) I8' 30 14

6) 30: 32

7) 32 38

S) :38-43

9) J 43 ?

Sand, loose, white, quartz .................. .. Surface sand,
possibly
Recent or
Pleistocene
wind-blown
sand

Sand, brown and white, loose, with non-cemented Pleistocene?
lumps ................ .................... terrace sands
-.---...-- -. Litholoyic Break ----- --- --
Clay, sandy, to sand, clayey, blue-green, with
minor tan and white phosphate nodules.......

Sand, slightly clayey, light green, with trace of Upper Bone
phosphate nodules ............. ............ Valley
formation
Sand, gray, mottled with light green, trace clay,
trace black phosphate nodules-unit massive..
Base of the "Overburden" of the Company ---
C(ravel, )phosphatic, gray, contains abundant
quartz sand but little or no clay.............
Lower Bone
Sand, white, strongly cross-bedded, very abund- Valley
ant black and brown phosphate nodules, from formation
coarse sand to granule size.............. ......

As above, except for abundant fossil bones.......
nconformity (Base of "Matrix") ----- -
(lay, greenish-gray, very sandy, with fine black Miocene:
phosphate nodules. Upper surface irregular and Hawthorn
filled with borings. Exposed in base of pit. ..... formation

LIST OF SPECIES

Order GAVIIFORMES

Family GAVIIDAE

The loons have a Holarctic distribution and a time record from
the upper Eocene to the Recent. There are four living and seven
fossil species.

REPORT OF INVESTIGATIONS NO. 14

Genus GAVIA Forster
Gavia palaeodytes Wetmore
Gavia palaeodytes Wetmore, 1943: 64, figs. 1-2 (orig. descr.; Middle
Pliocene, Pierce, Florida; type coracoid, M.C.Z. 2329) .-Wetmore, 1951:65
(Middle Pliocene, Pierce, Florida) .-Brodkorb, 1953C: 212, fig. 1C (near
Brewster, Florida; descr. coracoid, humerus, femur).
Material. Six specimens, three individuals. Represented in White's
locality by M.C.Z. 2329, in Elmore's Locality 1 by No. 88, and in
Locality 2 by the remaining specimens.
Coracoid: left distal M.C.Z. 2329 (cast of type); right complete
No. 132.

Humerus: right proximal No. 306; left distal Nos. 88, 524.
Femur: right complete No. 133.
All of the material listed except No. 524 has been described in
my paper cited above. The measurements of the latter specimen are
included in Table 3.
Gavia palaeodytes was a small species, about the size of the living
red-throated loon, G. stellata. The latter has a Holarctic distribution,
breeding in the far north and wintering south to the Gulf of Mexico
and the Mediterranean. Thus far G. palaeodytes is known only from
the Bone Valley.
Gavia concinna Wetmore
Gavia concinna Wetmore, 1940: 25, figs. 1-4 (orig. descr.; lower Pliocene,
Sweetwater Canyon, east of King City, California; type proximal portion
of ulna, U.S.N.M.).-Brodkorb, 1953C: 211, fig. 1A (Bone Valley formation,
near Brewster, Florida; descr. humerus, ulna, femur).
Material. Five specimens, two individuals. Represented in Locali-
ty 1 by Nos. 89, 90, and 593; in Locality 2 by Nos. 297, 298.
Humerus: left distal 90, 297; right proximal 593.
Ulna: right distal 89.
Femur: left complete 298.
The proximal humerus (593) was received since completion of
my paper cited above. Its measurements are included in Table 3.
This new material confirms the assignment of the Bone Valley speci-
mens to G. concinna.

The present species was somewhat larger than G. palaeodytes. It
had a continent-wide distribution. The type locality is in the Etchegoin
formation, which Wood et al. (1941: 19) and Woodring, Stewart, and
Richards (1940: 112, insert) consider middle Pliocene.

FLORIDA GEOLOGICAL SURVEY

Table 3. -MEASUREMENTS (IN MILLIMETERS) OF HUMERUS IN Gavia.

Breadth through epicondyles........
Breadth through condyles.........
Depth through internal condyle.....
Depth through external condyle.....
Depth through brachial depression...
Depth above brachial depression....
Width above brachial depression....
Length of attachment for anterior
ligament ........................
Length of internal condyle..........
Diagonal length of external condyle..
External tuberosity to capital groove
External tuberosity to internal
tuberosity ...................
Maximum depth of head ...........
l)epth through internal tuberosity...
Length of capital groove............

G. howardxs

(3 distals)

12.0-14.4
9.2-10.2
8.8- 9.3
7.8- 8.4
4.2- 4.7
4.7- 5.2
6.2- 6.8

9.5-1.0.2
4.3- 4.8
6.8- 7.4

G. palxodytes
(2 distals,
1proximal)

14.3-14.7
11.7-12.4
10.1
9.1- 9.2
5.1- 5.5
5.8- 6.1
6.9- 7.1

8.6-
5.2-
8.4-
16.6

8.7
5.4
8.8

19.3
9.5
6.4
8.6

I

Order COLYMBIFORMES

Family COLYMBIDAE

The record of this cosmopolitan family extends from the Oligocene
to the Recent. Eighteen living and five extinct species of grebes are
recognized.

Genus PLIODYTES Brodkorb

Pliodytes lanquisti Brodkorb

Pliodytes lanquisti Brodkorb, 1953D (orig. descr.; Bone Valley formation,
near Brewster, Florida; type coracoid).
Material. One specimen, one individual, Locality 2.
Corocoid: right complete, No. 299 (type).

This new genus combines some of the characters found in the
genera Colymbus and Podilymbus, besides having some unique char-
acters. It was about the size of the living pied-billed grebe, Podilymbus
podiceps. Colymbus pisanus (Portis), of the upper Pliocene of Italy, is
larger than the Bone Valley grebe (see Lambrecht, 1933: 262).

G. concinna
(2 distals,
2 proximals)

15.5-16.7
12.5-13.1
10.5-11.6
9.7-10.4
6.0- 6.1
6.5- 7.0
8.4- 8.8

10.2-10.7
5.4- 5.5
9.2-10.5
18.5-18.8

21.5-22.0
10.1-10.2
7.4- 7.6
9.4- 9.5

REPORT OF INVESTIGATIONS No. 14

Order PROCELLARIIFORMES
Family DIOMEDEIDAE
The albatrosses, represented by 14 living species, frequent all
oceans except the North Atlantic, where they are only of accidental
occurrence. It is therefore of considerable interest that the only
named fossil species, Diomedea anglica, has been recorded on both
sides of the North Atlantic.

Genus DIOMEDEA Linnaeus
Diomedea anglica Lydekker
Diomedea anglica Lydekker, 1891: 189, fig. 42 (orig. descr.; Red Crag,
Foxhall, Suffolk, England; type tarsometatarsus and associated toe phalanx,
Ipswich Mus.).-Lambrecht, 1933: 273 (type material assigned to middle
Pliocene; ulna from Coralline Crag, Lower Pliocene).- Wetmore, 1943: 66,
pl. 12, figs. 10-15 (middle Pliocene, near Pierce, Florida; descr. tibiotarsus).
Material. One specimen, one individual, White's locality.
Tibiotarsus. right distal, M.C.Z. 2328 (not examined by me).
It is unfortunate that the albatross was not represented among
the material collected by Elmore. The reference of the Florida speci-
men to Lydekker's species was made simply on the basis of size,
since the tibiotarsus is unknown in European collections. The two
English localities are now considered to be of late Pliocene (Astian)
and Middle Pliocene (Plaisancian) age.

Order PELECANIFORMES
Family SULIDAE

The family Sulidae, represented by nine living species, now has
a cosmomarine distribution. The present material includes specimens
of a gannet (Morus) and two boobies (Sula). The 15 fossil species
are all from the Holarctic Region. The fossil record of the family
is as follows:
Pleistocene: Morus bassanus (Linnaeus). Recent species
recorded as fossil from Norway.
Morus reyanus Howard. California.
Middle Pliocene: Miosula recentior Howard. California.
Bone Valley: Morus peninsularis Brodkorb. Florida.
Sula guano Brodkorb. Florida.
Sula phosphata Brodkorb. Florida.

FLORIDA GEOLOGICAL SURVEY

Upper Miocene: Miosula media L. Miller. California.
Morus lompocanus (L. Miller). California.
Morus stocktoni (L. Miller). California.
Morus vagabundus Wetmore. California.
Sula willetti L. Miller. California.
Middle Miocene: Sula pygmaea Milne-Edwards. France.
Lower Miocene: Morus loxostyla (Cope). Maryland, New
Jersey.
Sula avita Wetmore. Maryland.
Upper Oligocene: Sula arvernensis Milne-Edwards. France.
Lower Oligocene: Sula ronzoni (Gervais). France.
Genus MORUS Vieillot

Morus peninsularis new species
Figs. 1, 4, 7
Type. No. 148, collection of Pierce Brodkorb; nearly complete
left coracoid. Bone Valley formation, from Locality 2 near Brewster,
Polk County, Florida. Collected in February 1952 by George C.
Elmore.
Diagnosis. Agrees with Morus in having the lower anterior face
of the coracoid broad and plane toward inner side, instead of being
narrower and more rounded as in Sula (see Wetmore, 1926B).
Differs from Morus reyanus Howard (1936), from the Pleistocene
of California, in having the head of the coracoid narrower and more
pointed; length of bone somewhat shorter, but distance from head
to procoracoid somewhat greater.
Differs from Morus lompocanus (L. Miller, 1925) and Morus stock-
toni (L. Miller, 1935), both from the Miocene of California, in being
considerably smaller. The coracoids of these species have not been
described in detail nor well figured, so a further comparison is im-
practical. My assignment of Sula stocktoni to the genus Morus is
made on the basis of the humerus exceeding the ulna in length, the
reverse being the case in Sula (sensu stricto.

Differs from Morus loxostyla (Cope, 1870), from the Miocene of
Maryland and New Jersey, in greater size and wider shaft. A further
comparison with this species is likewise not possible at this time.

REPORT OF INVESTIGATIONS No. 14

The coracoid of Morus vagabundus Wetmore (1930), from the
Miocene-of California, is unknown. This species is described as being
of about the size of Sula sula, and thus the Florida gannet is a much
larger bird.
The new species is smaller than the three living species of gan-
nets, Morus bassanus (Linnaeus), M. capensis (Lichtenstein), and
M. serrator (Gray). It has the anterior intermuscular line located
relatively more posteriorly, and the head of the bone is more pointed.
In the Miocene and Pliocene genus Miosula the coracoid is larger
than in Morus (see L. Miller, 1925).
Referred material. No. 613, nearly complete left coracoid (para-
type), and No. 614, cervical vertebra, both from the type locality.
The cervical vertebra is about the size of that of Morus serrator.
but the length through the zygapophyses (25.4 mm.) and the narrow-
est posterior width of the centrum (6.0) are less than in that species,
whereas the length of the body of the vertebra (25.4) and the width
through the prezygapophyses (16.6) are greater.
Genus SULA Brisson
Sula guano new species
Figs. 2, 5, 8
Type. No. 301, collection of Pierce Brodkorb; nearly complete left
coracoid. Bone Valley formation, from Locality 2 near Brewster,
Polk County, Florida. Collected in September 1952 by George C.
Elmore.
Diagnosis. Agrees with Sula in having the lower anterior face of
the coracoid narrower and more rounded than in Morus.
Compared with living species, the fossil is similar in size to Sula
sula (Linnaeus), but the breadth of the head, breadth at level of
scapular facet, and breadth of shaft are greater in the fossil, and
the distance from head to procoracoid is somewhat greater. The
new species is smaller than S. nebouxii, but the head of the bone
and the internal sternal facet are both broader than in nebouxii.
The fossil is likewise smaller than S. dactylatra.
The present species agrees with Sula sula in having the internal
sternal facet shallow, with its medial margin passing gently toward
the shaft in sternal aspect. It differs in having the external sternal

FLORIDA GEOLOGICAL SURVEY

facet shorter and the internal sternal facet longer. The external facet
makes a more pronounced shelf media, and the shaft above the
shelf is more excavated. The anterior intermuscular line is situated
more laterad than in S. sula, but swings farther media at its lower
end as it meets the sternal facet, being parallel with the facet be-
fore joining it. A condition similar to that in the fossil occurs in
S. nebouxii and S. dactylatra, whereas S. leucogaster resembles S.
sula in that the line joins the facet without running parallel to it.
From the Oligocene forms of France, Sula ronzoni (Gervais, 1851)
and Sula arvernensis Milne-Edwards (1868), it differs in being much
smaller, since those species exceed Morus bassanus in size. Their
retention here in Sula merely follows custom and is not necessarily
a reflection of their true systematic position.
From the Miocene species, Sula pygmaea Milne-Edwards (1874)
from France, Sula willetti L. Miller (1925) from California, and Sula
avita Wetmore (1938) of Maryland, it differs in being much larger,
since the Miocene forms were all smaller than any living booby.
Referred material. In addition to the coracoid there are two
fragmentary right ulnae (Nos. 123, 529) and the distal end of a
tibiotarsus (No. 309), all from the type locality.
The ulna of both fossil and living species of Sula differs from that
of Morus in being more pneumatic in both the humeral and radial
depressions. Compared with the living species of Sula, the proximal
end of the ulna in the present species has the humeral depression
more excavated, making the olecranon more pointed and hooked
toward the inner side. There is a large pneumatic foramen in the
humeral depression. This depression is non-pneumatic in Sula sula
and S. leucogaster and shows only a slight pneumaticity in S. nebouxii
and S. dactylatra. The two ulnar fragments indicate a species be-
tween S. sula and S. nebouxii in size.
The tibiotarsus of Sula differs from that of Morus in having the
intercondylar fossa less excavated (i.e., shallower and wider), with
the condyles less protruding. In Sula the external condyle is short,
whereas in Morus it extends distad almost as far as the internal
condyle. Further, in Sula the inner margin of the shaft, in anterior
aspect, merges gradually into the internal condyle, so that the latter
is located directly below the inner margin of the lower part of the
shaft. In Morus the inner margin of the shaft swings abruptly media,
so that the internal condyle is located more media than the inner

REPORT OF INVESTIGATIONS No. 14

margin of the lower part of the shaft. The fossil tibiotarsus comes
from a bird near nebouxii in size. The intercondylar fossa is wider;
the internal condyle narrower; and the tibial bridge narrower and
more elevated on the shaft than in S. nebouxii.
Because of the resemblance of these elements to those of nebouxii
and because of the apparent affinity of nebouxii to S. guano, the
ulnar and tibial fragments are referred to this species.
Sula phosphata new species
Figs. 3, 6, 9
Type. No. 302, collection of Pierce Brodkorb; right coracoid, lack-
ing head and part of sternal end. Bone Valley formation, from
Locality 2 near Brewster, Polk County, Florida. Collected in Septem-
ber 1952 by George C. Elmore.
Diagnosis. Differs from the Oligocene and Miocene species of
Sula as described for Sula guano. From the latter species it differs
in having the procoracoid process situated higher on the shaft; shaft
slightly deeper; breadth at level of scapular facet somewhat less.
In particular it differs from S. guano in having the external sternal
facet longer and the internal facet shorter; the internal facet with
its margin more arched in sternal aspect, and meeting the shaft at
a more pronounced angle, as in S. leucogaster. The lower portion of
the anterior intermuscular line swings farther forward in the pres-
ent species than in S. guano.
Among living species it most closely resembles S. leucogaster,
but differs in having the shaft deeper; breadth at level of scapular
facet greater; and in having the internal sternal facet shorter. The
lower portion of the anterior intermuscular line lies parallel to the
sternal facet before joining it, which is not the case in S. leucogaster.
In both this species and the one just described, the coracoid is
about the same size as in Morus loxostyla, although of a different
form, as described above.
Referred material. No. 138, lower portion of left coracoid (para-
type); Nos. 120 and 597, upper portions of left coracoidd, all from
the type locality.
Specimen No. 138 resembles the type of S. phosphata. The two
specimens of the upper end of the bone agree among themselves and
differ from the type of S. guano; they are therefore likewise referred to
S. phosphata.

FLORIDA GEOLOGICAL SURVEY

In S. guano the overhang of the lip of the brachial tuberosity at
its upper end makes a rather clean sweep upward. In the two speci-
mens of S. phosphata in which this region is preserved the lip is
distinctly bifid at its upper end. Furthermore, in S. phosphata there
is a more pronounced depression, in posterior view, between the
medial margin of the head and the furcular facet. This is only
weakly indicated in S. guano,

Family PHALACROCORACIDAE
The abundant cormorant remains enable me to state that two
species are represented in the Bone Valley, and further that the
small form is not identical with the living double-crested species,
as was thought by Wetmore, but rather represents a distinct species,
closely allied and probably ancestral to the living bird.
The record of this cosmopolitan family, of which there are 30
Recent and 17 fossil species, extends back to the Oligocene.

Genus PHALACROCORAX Brisson
Phalacrocorax wetmorei new species
Figs. 10, 11
Phalacrocorax auritus, Wetmore, 1943: 68 (Middle Pliocene, near Pierce,
Florida; two distal metatarsi).
Type. No. 530, collection of Pierce Brodkorb; nearly complete
right coracoid. Bone Valley formation, from Locality 2 near Brewster,
Polk County, Florida. Collected in December 1952 by George C.
Elmore.

Diagnosis. Very similar to modern Phalacrocorax auritus flor-
idanus (Audubon) but bones in general less robust.

Coracoid with anterior intermuscular line situated farther laterad.
Humerus with head shallower, ligamental furrow relatively longer,
pneumatic fossa narrower and deeper, and condyles averaging less
deep. Ulna decidedly less robust. Carpometacarpus somewhat more
delicate, but with first metacarpal more produced. Femur averaging
longer and narrower. Tibiotarsus with both proximal and distal ends
more slender, but with internal condyle relatively deeper. Tarsome-
tatarsus averaging more slender, but shaft slightly deeper. The frag-
mentary specimens of synsacrum, scapula, and radius do not show
differences from P. auritus, nor do the cervical vertebrae and digits.

i,

Table 4.---MEASUREMENTS (MM.) OF CORACOID OF Morus AND Sula.

M. peninsularis..
M.loostyla.....
M. reyanus......
M. lompocanus...
M. bassanus.....
M. capensis.....
M. serrator......
S. guano .......
S. phosphata.....
S. willetti......
S. leucogaster ...
S. eula.........
S. nebouxii.... .
S. dactylatra......

Length
along axial
border

54.0-55.6
48.2-51.3
56.6
62.0
58.6-61.3
55.8
54.7
50.0

45.0
49.1
48.7-50.5
55.2
58.2

Length
of external
sternal facet

14.2-14.5
14.2

17.4-20.8
17.2
18.2
10.7
11.1-11.5

11.2
11.5-12.1
12.2
14.7

Length
of internal
sternal facet

8.6-10.8

.. . .. .o .
13.8-14.3
14.0
12.0
10.3
7.5- 7.7
. . .. .
8.4
9.4- 9.8
10.0
12.0

Least dep
of shaft

7.0- 7.
5.0- 5.

7.4- 8.
7.4
7.8
5.6
5.7- 5.

5.3
5.2- 5.
5.8
5.8

th Head to
procoracoid

,4 25.0-25.1
.8 ........... .
24.7

2 .27.9-29.6
27.4
27.6
21.0
8 21.0
. . . . .
20.0
5 20.5-20.7
23.5
25.0

Breadth
of head

14.2

15.5
............
14.4-15.4
15.4
15.5
13.4
12.7-13.5

11.4
11.0-11.6
12.5
13.8

Breadth at
leiel of
scapular facet

17.0-17.7

17.1

17.5-20.4
19.0
18.7
14.7
13.8-14.5

12.6
13.5-13.8
14.8
15.6

0
I
0

M

0
'.
14
0r
Co
z~

FLORIDA GEOLOGICAL SURVEY

Material. 135 specimens, at least 15 individuals. Represented in
White's locality by two specimens, two individuals; Locality 1 by
48 specimens, 5 individuals; Locality 2, 85 specimens, 8 individuals.
Coracoid: right complete 87 (paratype), 530 (type), right proxi-
mal 95, 312, 314, 315, 531; right distal 94, 179, 180, 181, 316, 532,
602; left complete 168 (paratype), left proximal 602; left distal 119,
313, 533.
Scapula: right proximal 97, 326; left proximal 96, 185, 607, 657.
Humerus: right proximal 99, 144; right shaft 319; right distal 102,
103, 183; left proximal 98, 182, 317; left shaft 100, 101; left distal 104,
121, 122, 184, 303, 608, 658.
Radius: right proximal 659; left distal 325, 534, 660.
Ulna: right proximal 105, 189, 190, 320, 321, 535, 536, 537, 603,
619, 620; right distal 186, 187, 621; left proximal 106, 124, 169, 188, 322,
538, 604; left distal 323, 324, 539, 540, 605.
Carpometacarpus; right proximal 541, 618; left proximal 125, 327,
617; left distal 328, 542, 606.
Alar digit: 117, 126.
Femur: right complete 662; right proximal 127, 193, 331; left
complete 145, 543, 544; left proximal 191, 192, 661.
Tibiotarsus: right distal 108, 109; left proximal 107; left distal
110, 128, 170, 194, 195, 545, 609, 616.
Tarsometatarsus: right proximal 113, 114, 129, 198; right distal
131, 199, 334, 335, M.C.Z. 2326, M.C.Z. 2327; left proximal 111, 112,
115, 130; left distal 116, 196, 197, 332, 333.
Synsacrum: 118, 329, 615, 656.
Cervical vertebrae: 200, 201, 330, 546, 547, 599, 600.

Phalacrocorax idahensis (Marsh)
Fig. 12
Graculus idahensis Marsh, 1870: 216 (orig. descr.; Pliocene: Castle Creek,
Idaho; type fragmentary carpometacarpus, Yale Univ.)
Referred material. No. 311, proximal portion of left ulna, Locality
2.
This tremendous ulna came from a cormorant much larger than

REPORT OF INVESTIGATIONS No. 14

any living species and differing from the mean of P. wetmorei by
about six standard deviations. Its measurements are as follows: proxi-
mal width 13.7, depth through internal cotyla 12.8, maximum width
of shaft 11.8 mm.
Two large fossil cormorants have been described from North
America.
Phalacrocorax macropus (Cope, 1878) was based on material from
the Pleistocene of Fossil Lake, Oregon. A number of the elements
of the skeleton have been described (Howard, 1946) but these un-
fortunately do not include the ulna. However, the proximal portion
of an ulna has been figured by Shufeldt (1913, pl. 21, fig. 269).
Compared with the figure, the Bone Valley specimen is a trifle
smaller, with the olecranon more pointed and deflected.
Phalacrocorax idahensis (Marsh, 1870) was described from a
fragmentary carpometacarpus from the Pliocene of Castle Creek,
Idaho. Later Wetmore (1933: 5) referred the distal portion of an
ulna from the upper Pliocene Hagerman Lake beds to the same
species. He stated that the ulna is not quite as heavy as in the extinct
Phalacrocorax perspicillatus of Bering Island but gave no measure-
ments. Comparison of the figures of the carpometacarpi of P. macropus
(Shufeldt, 1913, pi. 21, fig. 262, 263) and P. idahensis (Shufeldt, 1915,
pl. 6, fig. 44) shows the latter to be somewhat the smaller. As the
Bone Valley specimen is also somewhat smaller than P. macropus, I
refer it to P. idahensis, following the precedent of Wetmore in refer-
ring the Hagerman Lake specimen.
Order CICONIIFORMES
Family ARDEIDAE
The 66 living species of herons give this family a cosmopolitan
distribution. The record of the family extends back to the Eocene,
but it is unrecorded from the Pliocene epoch, except for two occur-
rences of Ardea, species uncertain, from the upper Pliocene of Europe
(Lambrecht, 1933: 734). The heron described below, the seventeenth
fossil species, thus helps fill in the chronology of the family.

FLORIDA GEOLOGICAL SURVEY

Table 5.-MEASUREMENTS (MM.) OF Phalacrocorax wetmorei

Coracoid:
Length along medial side.......
Head to procoracoid ...........
Least width of shaft ...........
Least width of blade........
Anterior intermuscular line to
medial end of sternal facet. .
Width of head ...............
Humerus:
Proximal width ...............
Depth of head ................
Length of ligamental furrow ....
W idth of shaft ................
I istal w idth ..................
Height of internal condyle......
Height of external condyle ..

Ulna:
Proximal width ...............
Depth through internal cotyla. .
Maximum width of shaft.......
Maximum distal diameter......
Depth of exte'rnal condyle......
Distal depth of shaft ..........
Least distal width of shaft......
Carpometacarpus:
Length of metacarpal two......
Width of shaft, metacarpal two.
Height through metacarpal one.
Width through trochlel........
W idth of distal end ............
Femur:
Length .......................
Width through condyles........
Narrowest width of shaft.......
Depth of external condyle......
Width through head...........
Tibiotarsus:
Length of outer cnemial crest...
Width of proximal end........
greatestt width through fibular
crest........................
Breadth through condyles. ...
Narrowest breadth of shaft.....
Narrowest depth of shaft.......
Depth of internal condyle......
Height of internal condyle.......

Mean

63.50
21.44
5.07
3.12

11.24
12.14

22.22
7.10
14.07
7.83
15.77
6.17
10.52

11.28
10.42
9.08
10.75
8.16
5.51
5.26

- .42
- .13

.18
.20
.06
.10

.10
.08
.07
.13
.12
.07
.09

54.0
4.60 =: .04
13.80
6.20
6.90

58.18
15.25
6.14 A .03
9.92
13.46 .18

16.2
10.2

10.2
11.67
6.64
4.45
10.92
9.07

.13
.06
.04
.12
.10

Range

60.7-65.3
20.5-22.4
4.4- 5.4
2.7- 3.4

9.4-13.6
11.2-12.7

20.7-23.8
7.0- 7.3
13.6-14.6
7.0- 8.5
14.7-17.3
5.8- 6.5
9.9-11.1

10.1-11.8
9.9-10.8
8.4- 9.5
10.0-11.2
7.5-- 8.4
5.0- 5.7
4.6- 5.5

4.4- 4.8
13.6-14.2
6.0- 6.3
6.6- 7.1

57.0-61.0
14.6-15.8
6.0- 6.3
9.1-10.3
12.6-14.4

11.0-12.3
6.3- 7.0
4.2- 4.7
10.4-11.6
8.3- 9.7

Standard
DeViation

.55
.27
.19

1.39
.40

.51
.67
.22
.34

.42
.34
.29
.36
.35
.19
.26

.12

.10
. . .
.56

.40
.16
.14
.36
.31

Num-
ber

3
9
9
13

11
9

3
4
3
8
11
11
11

18
17
18
8
8
8
8

1
8
5
5
3

4
4
9
4
10

1
1

1
9
8
10
9
9

___~_

REPORT OF INVESTIGATIONS No. 14

Standard Num-
Mean Range Deviation ber

Tarsometatarsus:
Depth through hypotarsus..... 17.63 17.2-18.0 ............ 4
Breadth of proximal end. ...... 12.43 11.6-13.3 ............. 7
Narrowest width of shaft....... 5.97 = .10 5.4- 6.5 .32 11
Breadth through trochler....... 14.65 = .17 13.4-15.5 .58 11
Narrowest depth of shaft....... 4.41 .08 3.8- 4.8 .27 11

Genus ARDEA Linnaeus
Ardea polkensis new species
Figs. 13, 14, 15
Type. No. 308, collection of Pierce Brodkorb; proximal portion
of right tarsometatarsus. Bone Valley formation, from Locality 2,
near Brewster, Polk County, Florida. Collected in September 1952
by George C. Elmore.
Diagnosis. Similar to living Ardea herodias Linnaeus of North
America and Ardea cocoi Linnaeus of South America, but differs
in smaller size; intercotylar knob more pointed and its outer edge
more abruptly ascending; groove on inner side of intercotylar knob
extending upward (as in Ardea cinerea) instead of being obliquely
transverse; inner hypotarsal ridge relatively longer and continuing
more proximad.
Differs from living Ardea cinerea Linnaeus of the Palearctic Region
in larger size; more pointed intercotylar knob; longer inner hypotarsal
ridge; relatively lower inner articular surface and rim.
Measurements of the type (compared in parentheses with those
of Ardea herodias, cocoi, and cinerea, respectively) are as follows:
proximal width 14.7 (16.4-16.6, 16.8, 14.2); proximal depth 15.0 (17.0-
17.2, 16.8, 14.2); depth through middle of hypotarsus 13.7 (14.4-15.0,
16.2, 12.5), length of first hypotarsal ridge 13.0 (12.0-13.0, 11.4, 9.3),
width of shaft below hypotarsus 7.6 (7.5-8.2, 9.6, 6.8 mm.).
The only other American fossil herons are Botauroides parvus
Shufeldt (1915: 33) and Eoceornis ardetta Shufeldt (1915: 39), both
from the Eocene of Wyoming and both considerably smaller forms.
Ardea paloccidentalis Shufeldt (1892: 820), from the Pleistocene of

FLORIDA GEOLOGICAL SURVEY

Oregon, has been synonymized by Howard (1946: 157) with the living
American bittern, Botaurus lentiginosus, likewise a smaller bird.
Family PHOENICOPTERIDAE
The six living species of flamingoes reach their northern limits
in the West Indies and the Mediterranean area. The 14 fossil forms
are entirely Holarctic in distribution, with a record going back to
the upper Eocene.
Genus PHOENICOPTERUS Linnaeus
Phoenicopterus floridanus Brodkorb
Phoenicopterus floridanus Brodkorb, 1953A; 1, figs. 1-2 (orig. descr.; Bone
Valley formation, near Brewster, Florida; type tibiotarsus, tarsometatarsus).
Material. Four bones, two individuals, Locality 2.
Tibiotarsus: right distal 147 (type); right shaft 202.
Tarsometatarsus: right distal 146, 300.
This is the first fossil flamingo material from eastern North Amer-
ica. Four species, referred to two genera, have been described from
the west (South Dakota, Oregon, California, and Chihuahua) in the
Miocene, Pliocene, and Pleistocene.

Order ANSERIFORMES
Family ANATIDAE
This large family, of 228 Recent species, has a cosmopolitan dis-
tribution. The fossil record goes back to the Cretaceous, and with
the one described below there are now 84 species known.

Genus BUCEPHALA Baird
Bucephala ossivallis new species
Figs. 16, 17
Type. No. 172, collection of Pierce Brodkorb; proximal half of left
coracoid. Bone Valley formation, from Locality 2 near Brewster,
Polk County, Florida. Collected in April 1952 by George C. Elmore.
Diagnosis. Referable to the Subfamily Aythyinae on the basis of
the head of the coracoid rising forward and upward from the anterior
plane of the shaft.
Closely agrees with the living Bucephala clangula (Linnaeus)

REPORT OF INVESTIGATIONS No. 14

in general appearance, particularly in the shape of the brachial tuber-
osity and the truncated upper margin of the head. Differs from B.
clangula in much smaller size; more curved coraco-humeral groove;
better developed procoracoid process; and more excavated triosseal
canal.

Differs from living Bucephala albeola (Linnaeus) in larger size;
less pronounced medial overhang of brachial tuberosity; relatively
shallower head, which is more inclined from plane of shaft; stouter
shaft; more curved coraco-humeral groove; better developed procora-
coid process; more excavated triosseal canal.

Also resembles Melanitta in truncate head and decidedly curved
coraco-humeral groove. Differs in much smaller size and in lacking
the massiveness of the bone of that genus; more excavated triosseal
canal; less overhanging medial end of lip of brachial tuberosity.

Differs more widely from Aythya, although resembling the smaller
forms of that genus in size. Aythya has the head more rounded and
less inclined forward from the plane of the shaft; less curved coraco-
humeral groove; and shallower triosseal canal.

Measurements. Width of head 6.0; maximum depth of head 3.2;
head to lower end of scapular facet 13.2; least width of shaft 4.2 mm.

Order CHARADRIIFORMES
Family HAEMATOPODIDAE
The oystercatchers are represented in the living fauna by a single
genus with four species. The rather spotty distribution along the sea
coasts of the world suggests some antiquity for the group. The only
fossil heretofore described is Paractiornis perpusillus Wetmore (1930),
from the Lower Miocene of Nebraska. The discovery of this family
in the Florida phosphate is therefore a welcome addition to our scanty
knowledge of the group.

Genus PALOSTRALEGUS new genus
Diagnosis. Distal portion of tibiotarsus agrees with Haematopus
in having the external ligamental prominence only moderately angular
and situated well above condyle; groove for peroneus profundus mod-
erately developed; internal condyle in distal aspect nearly perpendi-
cular, that is slanting only slightly away from shaft; internal liga-

FLORIDA GEOLOGICAL SURVEY

mental prominence situated high with relation to the condyle, that
is at the anterior edge of the lower end of the shaft.
Differs from Haematopus in having the intercondylar sulcus more
excavated; tibial bridge ossified; ligamental groove above bridge nar-
rower; internal ligamental prominence slightly higher and slightly
better developed.
Differs from the Scolopacidae, as exemplified by Numenius, in
having the intercondylar sulcus more excavated and without raised
medial portion; external ligamental prominence less angular and situ-
ated higher on shaft; groove for peroneus profundus less distinct;
tibial bridge and its openings more oblique; ligamental groove above
tibial bridge narrower; internal ligamental prominence better devel-
oped and located higher and more anteriorly; internal condyle in
distal aspect nearly perpendicular, not slanting so abruptly away from
shaft.
Differs from the Burninidae in having the intercondylar sulcus
wider and without raised medial area; internal condyle with upper
end more inclined toward shaft; internal ligamental prominence more
angular and situated above condyle; groove for peroneus profundus
less distinct; internal condyle in distal aspect much deeper than ex-
ternal condyle (only slightly so in Burhinus).
From the Recurvirostridae it differs more markedly. Its long in-
ternal condyle, stout shaft, and raised internal ligamental prominence
remove it from that family immediately.
Type. Palostralegus sulcatus new species.

Palostralegus sulcatus new species
Fig. 18
Type. No. 177, collection of Pierce Brodkorb; distal third of right
tibiotarsus. Bone Valley formation, at Locality 1 near Brewster, Polk
County, Florida. Collected in May 1952 by George C. Elmore.
Description. Anterior face with surface of shaft flat with a ridge
rising along distal portion of internal edge, its distal portion with a
tendinal groove and with an anterior prominence; supratendinal bridge
with its upper margin sloping medially, its lower margin more nearly
straight; a pronounced external ligamental prominence above sup-
ratendinal bridge; a well-marked flattened knob for muscle attach-

REPORT OF INVESTIGATIONS No. 14

ment .between external ligamental prominence and supratendinal
bridge; a groove for peroneus profundus between this and external
ligamental prominence; distal opening under supratendinal bridge
obliquely rounded, with lower end sloping toward medial edge; in-
ternal ligamental prominence angular, with its apex (slightly im-
perfect) below projected line from distal opening of supratendinal
bridge, its anterior face concave; external condyle broad and up-
right; internal condyle narrower, sloping proximally toward sup-
ratendinal bridge, and extending somewhat farther distad than ex-
ternal condyle; intercondylar sulcus broadly rounded, with little divi-
sion into lateral and median portions.
Internal face with internal condyle lengthened anteriorly, its sur-
face concave and its edges distinctly raised; internal ligamental
prominence located on anterior edge of bone, immediately above
condyle; distal border of condyle slightly indented forward of mid-
line of shaft projected.
Posterior face with shaft sloping toward medial edge; intercondy-
lar sulcus broad, shallow, and practically undivided.
External face with shaft gently rounded toward rear; groove for
peroneus profundus distinct; external condyle rounded, its surface
concave, with prominent raised anterior margin, a papilla in center,
and another above groove for peroneus profundus.
Measurements. Width through condyles 8.0; width through in-
ternal ligamental prominence 8.2; width of shaft 4.4; depth of ex-
ternal condyle 7.5; depth of internal condyle 9.4 approximately; depth
of shaft 3.6; width of distal end of posterior intercondylar sulcus
6.0 mm.
Characters. Resembles modern Haematopus palliatus and H.
bachmani, but width through condyles less; shaft more robust (wider
and deeper); external ligamental prominence situated higher on shaft
and more pronounced (rising more abruptly from shaft, both distally
and proximally); internal ligamental prominence more angular and
situated slightly higher on shaft; tendinal groove rather more pro-
nounced; internal condyle deeper; intercondylar sulcus narrower and
deeper; groove for peroneus profundus deeper; tibial bridge com-
pletely ossified.
Paractiornis perpusillus is known only from the tarsometatarsus.
It is a tiny bird, about the size of a sanderling.

22 FLORIDA GEOLOGICAL SURVEY

Family SCOLOPACIDAE
Although 21 fossil species of sandpipers have been previously
described, only one has heretofore been found in the Pliocene de-
posits of North America. This is Micropalama hesternus Wetmore
(1924), from the Upper Pliocene of Arizona. The discovery of three
species of scolopacids in the Bone Valley gravel is therefore of con-
siderable interest.
Genus CALIDRIS Merrem
Calidris pacis new species
Figs. 19, 20
Type. No. 594, collection of Pierce Brodkorb; proximal half of
left humerus, with internal tuberosity broken. Bone Valley forma-
tion, from Locality 2 near Brewster, Polk County, Florida. Collected
January 18, 1953, by George C. Elmore.
Characters. Similar in general size and conformity to modern
Calidris canutus (Linnaeus), but head of humerus more rounded, less
elongate; capital groove shallower; coraco-humeral groove broader
and deeper, the outer part of its proximal margin straight, and with
more pronounced overhang proximally; medial bar lying more oblique
to mid-line; capital-shaft ridge straighter, not deflected inwardly;
surface of deltoid ridge slanted toward external edge of bone, instead
of toward inner edge, the scar about the same width throughout,
instead of tapering distally; bicipital groove shorter; bicipital furrow
deeper.
Measurements. Proximal width 10.6; least width of shaft 3.2; least
depth of shaft 2.8; depth of head 2.8; length from outer end of bici-
pital groove to end of caput humeri 8.8 mm.
Comparisons. Although its measurements are almost identical
with those of Calidris canutus, this species may require generic sep-
aration when more specimens are collected.
Micropalama hesternus Wetmore is a smaller species with the
proximal width of the humerus 7.7 mm., and the length of the bici-
pital groove 7.1 mm.
The humerus of Calidris gracilis (Milne-Edwards, 1868), from the
upper Oligocene of France, is described as being much smaller than
that of C. canutus. It may belong in the genus Erolia rather than
Calidris.

REPORT OF INVESTIGATIONS No. 14

Totanus grivensis Ennouchi (1930), from the upper Miocene of
France, is larger than C. pacis, and Tringa numenoides (Serebrovsky,
1941), from the Pliocene of the Ukraine, is very much larger.

Genus EROLIA Vieillot
Erolia penepusilla new species
Fig. 21
Type. No. 611, collection of Pierce Brodkorb; left humerus, lack-
ing the head. Bone Valley formation, from Locality 2 near Brewster,
Polk County, Florida. Collected by George C. Elmore, March 10, 1953.
Characters. Agrees with the smaller living species of Erolia in
having the external condyle relatively small and the internal condyle
relatively large. Larger than the Asiatic Erolia temminckii (Leisler),
the Palearctic E. ruficollis (Pallas), and the American E. minutilla
(Vieillot). Smaller than the American species Erolia bairdii (Coues)
and E. fuscicollis (Vieillot). Appears to be closest to Erolia minutilla,
from which it differs in larger size and in having a higher internal
condyle/external condyle ratio.
Larger also than the American genus Ereunetes, represented by
the two living species, E. pusillus (Linnaeus) and E. mauri Cabanis.
The humeri of Erolia and Ereunetes are very close, but differ in pro-
portions of the condyles. In various species of Erolia the ratio internal
condyle/external condyle is 36.00-50.00 per cent. In Ereunetes the
ratio is 33.33-41.67 percent, reflecting the relatively large external
condyle and the small internal condyle. The new species, with a
ratio of 44.00 percent, falls within the range of Erolia and beyond
that of Ereunetes.
The only fossil species of comparable size is Totanus minor En-
nounchi (1930), described from the Miocene of France. Inspection
of Ennounchi's plate shows that penepusilla differs in having the
distal end of the bone more on a plane, with the internal condyle
less deep as seen from below, and its shaft is narrower. Ennouchi's
species appears to have little in common with the genus Totanus, now
known as Tringa. Furthermore, the name minor is preoccupied in
Tringa by Tringa cinclus minor Schlegel (1844).
Ennouchi's bird is very similar to the smaller species of Erolia,
and probably should be referred to that genus. However, the name
minor is also preoccupied in Erolia by Tringa cinclus minor Schlegel,

FLORIDA GEOLOGICAL SURVEY

since the latter is a synonym of Erolia alpina (Linnaeus). In view of
these circumstances, it is greatly to be desired that Dr. Ennouchi
supply a new name for his Totanus minor.
Measurements. Length (estimated), about 26.1 mm.; length from
bicipital crest 20.7; narrowest width of shaft 1.7; distal width 4.0;
diagonal length of external condyle 2.5; height of internal condyle
1.1; upper base of spur to distal end 3.7; narrowest depth of shaft
1.5 mm.
Genus LIMOSA Brisson
Limosa sp.
Figs. 22, 23
Material. Two specimens, one individual.
Nos. 526, 527, distal and proximal portions of right tibiotarsus
(Locality 2). There is a segment of about an inch of the shaft miss-
ing between the two pieces.
Characters. Differs from the living Limosa lapponica and L. fedoa
in having smaller condyles and narrower posterior intercondylar
sulcus, but with the shaft of about the same breadth and depth as
in those species.
Because of the inadequate description of Limosa vanrossemi Miller
(1925) from the Miocene of California, I am unable to differentiate
my bird from that form, the only measurement of the tibiotarsus
given by Miller being the length. The type is an impression of a
skeleton, and it is not feasible to take accurate measurements. Dr.
Miller kindly compared my material with his type but was unable
to come to a conclusion. This is unfortunate, because the difference
in age of the Florida bird makes it practically certain that it repre-
sents a new species.
Measurements. Width through condyles 6.3; narrowest width of
shaft 3.2; depth of shaft 2.5; depth of external condyle 5.7; depth
of internal condyle 6.5; width of posterior intercondylar sulcus 4.7;
width of shaft above fibular crest 3.7; width of head 5.7 mm.

Family LARIDAE
Gulls are relatively rare as fossils, there being 84 living and
14 fossil ones. The record goes back to the Oligocene.

REPORT OF INVESTIGATIONS NO. 14

Genus LARUS Linnaeus
Larus elmorei Brodkorb
Larus elmorei Brodkorb, 1953B: 94, fig. 1 (orig. descr.; Bone Valley
formation, near Brewster, Florida; type distal portion of humerus; descr.
coracoid, carpometacarpus).
Material. Six specimens, two individuals. Nos. 176 and 178 are
from Locality 1, the others from Locality 2.
Coracoid: right proximal 134.
Humerus: right distal 140 (type); left distal 176.
Ulna: left proximal 307.
Carpometacarpus: left distal 178; left proximal 528.
Measurements. Two of the above specimens (Nos. 307 and 528)
were received after the description of this species was published.
Their measurements are given below.
Ulna: width through cotylae 11.0, depth of shaft 5.2 mm.
Carpometacarpus: width of metacarpal two 3.8, height of proximal
end 11.7, width through trochleae 5.0.
Remarks. The phosphate species is closely related to the living
ring-billed gull, Larus delawarensis, of which it was probably the
direct ancestor.
Family ALCIDAE
At the present time the Alcidae occur in Florida only as rare
or accidental stragglers along the east coast. The dovekie (Plautus
alle), a species subject to sporadic migrations, has been collected a
few times in Florida, and two bones of the great auk (Pinguinus
impennis) have been discovered in an Indian shell heap at Ormond
(Hay, 1902). Considerable interest; therefore, is attached to alcid
material in the phosphate deposits of the Gulf region. The Bone
Valley specimens do not necessarily indicate a cooler climate, how-
ever, since several species breed at present in the Pacific as far
south as Latitude 27 along the Mexican coast.
The family is entirely Holarctic. There are 23 Recent species and
the Bone Valley auk is the tenth fossil species.
Genus AUSTRALCA new genus
Diagnosis. Coracoid agrees with that of the Alcidae and differs
from the Mancallidae in having the sternal facet with a broad mesial

25

FLORIDA GEOLOGICAL SURVEY

flare. The brachial tuberosity and the sternal facet are elongated,
as in Pinguinus, Alca, Synthliboramphus, Uria, and Cepphus, in con-
trast with the group typified by Plautus, Fratercula, Lunda, and
Cerorhinca, in which these parts are reduced.
The proximal end of the bone is wide, with the ratio of the dis-
tance from head to brachial tuberosity greater than in any living
form examined. The shaft is also relatively wide.
Differs from Pinguinus in narrower dorsal end, but relatively more
produced brachial tuberosity; relatively greater depth of head; less
elongate and straighter neck of the coracoid.
Differs from Alca in somewhat shorter relative distance from head
to scapular facet; more produced brachial tuberosity; greater depth
of head; wider sternal end.
Differs from Synthliboramphus in more produced and less de-
flected brachial tuberosity, and less rotated procoracoid.
Differs from Uria in more produced and less deflected brachial
tuberosity, wider shaft, and deeper head.
Differs from Cepphus in more produced and less deflected brachial
tuberosity, deeper shaft, wider sternal end, and in having a fenest-
rate, not notched, procoracoid.
Type. Australca grandis, new species.
Relationships. In general appearance the coracoid of this genus
falls between that of Pinguinus on the one hand, and Uria and Alca
on the other.
An index to the coracoid was obtained by summing four intramem-
bral ratios. These were the percentage of the length of the bone in-
volved in the distances of the head to scapular facet, glenoid facet
to brachial tuberosity, depth of shaft, and sternal width, respectively.
The indices thus computed for various alcids are as follows: Pinguinus
139, Alca 127, Synthliboramphus 122, Australca 121, Uria 117, Cep-
phus 116, Plautus 98, Fratercula 95, Lunda 92, and Cerorhinca 91 per-
cent. The index of Australca confirms its intermediate position be-
tween Pinguinus and the Uria-Alca group.

REPORT OF INVESTIGATIONS No. 14

Australca grandis new species
Figs. 24, 29
Type. No. 141, collection of Pierce Brodkorb; right coracoid, lack-
ing the hyosternal facet. Bone Valley formation, from Locality 2, near
Brewster, Polk County, Florida. Collected in February 1952 by George
C. Elmore.
Diagnosis. General size of coracoid near that of Lunda, shorter
than in Pinguinus, and larger than that of any other living alcid.
Length of bone to medial side 44.4, brachial tuberosity to sternal
facet 44.2, head to procoracoid 17.0, glenoid facet to brachial tuber-
osity 14.1, head to brachial tuberosity 8.5, depth of shaft below
glenoid facet 5.4, greatest depth of head 7.0, width of sternal facet
17.5 mm.
Referred material. The total alcid material consists of three speci-
mens and one individual from Locality 1, and 15 specimens and four
individuals from Locality 2.
Humerus: right proximal 137, 310; right distal 91, 304, 305, 595;
left proximal 135, 136, 525; left distal 173.
Radius: left distal 93.
Ulna: right proximal 142; left distal 92, 596.
Carpometacarpus: right distal 143.
Tibiotarsus: left distal 612.

Humerus with capital lip but little overhanging capital groove
surface, as in Alca and Pinguinus, less so than in Uria, much less
so than in Cepphus. However, the medial bar is only faintly indicated,
and in this respect the fossil more nearly resembles Cepphus and
Uria. None of the humeri is complete, but two (Nos. 91 and 304) lack
only the proximal end. These indicate that the humerus was similar
in length to that of Uria, being decidedly longer than in Alca or Cep-
phus. The distal portion of the humerus has a very well developed
ectepicondylar process, whose outer margin is parallel to the shaft,
as in Cepphus, instead of being inclined toward the shaft proximally
as Uria and Alca. The proximal margin of the ectepicondylar process
is therefore wider and is also more truncate than in living genera.
Possibly No. 525 may represent another species, since its shaft nar-
rows abruptly distal to the middle of the bone. This character, how-
ever, shows some variation in living alcids.

FLORIDA GEOLOGICAL SURVEY

The ulna is represented by two distal and one proximal portions.
The entire bone seems to have been about the size of the ulna of
Uria, and resembled that genus in its conformity, without the pro-
nounced shortening which occurs in Pinguinus. The proximal end is
a little smaller than in Uria, but wider than in Alca. The two distal
ends differ considerably in size, No. 596 equaling the great auk in
most measurements. The shaft in both distal specimens tapers gently
distad, without any pronounced decrease near the condyles.
The fragmentary radius resembles that of Alca and Uria, without
the great deepening and compression of Pinguinus. There is a pro-
nounced neck just before the distal end, more so than in any other
alcid examined.
The fragmentary carpometacarpus is relatively short compared
with the other elements, and therefore shows a tendency toward the
condition in Pinguinus. In size it comes closest to Cepphus columba,
being smaller than the carpometacarpus of Uria, and also being smaller
than Alca except in the length of the shaft of the second metacarpal,
which is the same. It therefore may be said that the carpometacarpus
resembles that of Alca in length, but its distal portion is narrower
and less deep.
The distal portion of the tibiotarsus resembles that of Uria. The
internal condyle extends far inward, as in Uria, more so than in
Alca and Cepphus. The posterior intercondylar sulcus is broad as in
Uria and Pinguinus; in Alca it is somewhat, and in Cepphus it is
much narrower. The tibial bridge is incompletely ossified, as in Pin-
guinus, Alca, and some specimens of Uria; the bridge is ossified in
Cepphus.
Measurements. Humerus: length to pectoral attachment (1), 66.0;
proximal width (1), 18.3; depth of head (4), 6.1-6.7; least width of
shaft (6), 5.5-6.6; depth of shaft (8), 3.3-3.8; distal width (5), 8.0-8.5.
Ulna: proximal width (1), 8.4; depth though external cotyla (1),
10.3; distal depth (2), 8.7-9.8.
Radius: distal width (1), 5.4.
Carpometacarpus: length of second metacarpal (1), 28.2; distal
depth (1), 6.4.
Tibiotarsus: width through condyles (1), 7.1.
Conclusions. Considerable study was required before it was de-

28

REPORT OF INVESTIGATIONS NO. 14

termined that all the alcid material represented a single species. It
gradually became apparent that the Bone Valley auk was a large
bird with the wings reduced in size compared with the living mem-
bers of the family. The reduction is especially evident in the distal
elements of the wing.
All alcids use the wings in swimming under water (Bent, 1919:
206). Cepphus, Uria, and Alca are strong fliers. The recently extinct
great auk, Pinguinus impennis, had lost the power of flight and had
carried the reduction of the distal wing elements to a remarkable
degree (see Table 6).
In Australca the reduction of the wings was about half-way be-
tween the condition in the flying alcids and Pinguinus. It was thus
already well on the road to flightlessness, and because of other similari-
ties it may even have been the ancestor of the great auk.

Table 6.-RATIOS (PERCENT) OF WING ELEMENTS TO LENGTH OF CORACOID

Ulna
Humerus depth Radius Second
Coracoid length to through distal meta-
length pectoral external width carpal
attachment cotyla length

Cepphus columba......... 100.00 181.61 27.42 16.13 88.71
Uria aalge ............... 100.00 187.18 27.69 14.62 79.23
Uria lomvia............. 100.00 185.75 27.75 15.50 78.00
Alca torda................ 100.00 168.99 29.61 15.36 78.77
Australca grandis.......... 100.00 149.32 23.20 12.22 63.80
Pinguinus impennis....... 100.00 134.62 21.79 9.77 43.22

AGE OF THE DEPOSIT

Geological opinion differs as to the age of the Bone Valley forma-
tion. Cooke (1945: 207) considered it middle Pliocene (Hemphillian
age). Cathcart (1950) allocated the formation to the lower part of the
Pliocene. From a study of the Pleistocene shore lines MacNeil (1950:
106) concluded that there is a possibility that the Bone Valley gravel
might be early Pleistocene (Aftonian), but he too accepted a Pliocene
age. Currently (in litt., January 18, 1954) he favors uppermost Mio-
cene for the upper Bone Valley and middle Miocene (Hawthorn)
for the lower Bone Valley. Vernon (1943: 156) first assigned the

29

FLORIDA GEOLOGICAL SURVEY

Bone Valley to the Pleistocene but now believes it to be late and
middle Miocene (Vernon, 1951: 195, 197).
The cetaceans were studied by Allen (1921), Kellogg (1924), and
Case (1934). These mammals were thought to be of Miocene age, or
more specifically late Miocene. It has been suggested that the ceta-
ceans were redeposited secondarily from reworked upper Miocene
rocks, but this is at variance with the finding of articulated specimens.
Those who have worked with the land vertebrates are in general
agreement as to the Pliocene age of the fauna. Simpson (1930) placed
the land mammals in the lower Pliocene and later stated that the
relationships of the sirenians also supported this view (Simpson, 1932).
White (1941A, 1941B) variously attributed certain land mammals
to the lower or middle Pliocene. Wood et al. (1941: 15) considered
the Bone Valley vertebrates to be of Hemphillian (middle Pliocene)
age.
It is obvious from these accounts that the Miocene-Pliocene
boundary in Florida is in need of further study, and the land verte-
brate chronology may be a portion of an epoch ahead of the chronology
based upon marine sediments. In other words, what is considered
lower Pliocene by vertebrate paleontologists may be the equivalent
of uppermost Miocene in the marine invertebrate chronology.
Age of the Avifauna. In determining the age of the avifauna the
possibility of reworking must be considered. No articulated bird
skeletons were discovered. This in itself cannot be taken as evidence
of reworking, however, since skeletons of birds dying in the rookeries
of the Everglades today are similarly disarticulated and scattered
(see Figs. 30-32). Much of the Bone Valley bird material is in ex-
cellent condition. In nearly every specimen the muscle scars and
processes are perfectly preserved with little or no abrasion. Extensive
reworking is therefore out of the question, and the bird material
must be contemporaneous with the sediments.
The age of the avifauna will be tested by the proportion of ex-
tinct and living species and by the presence of indicators of particular
epochs on both generic and specific levels.

A comparison of the proportion of extinct species to those still
living is given in Table 7 for the Bone Valley and other late Tertiary
and Quaternary deposits in North America. As might be expected,
all the Pleistocene localities have a relatively low proportion of ex-

REPORT OF INVESTIGATIONS No. 14

tinct forms, with a mean of 20.3 percent (range 13.7-24.3). The upper
Pliocene avifaunas of Blancan age are marked by a decided increase
in the proportion of extinct species, with a mean of 72.3 percent
(range 63.6-83.3). All the localities of middle Pliocene (Hemphillian)
age or older are characterized by having their avifaunas composed
entirely of extinct species. Since the Bone Valley avifauna is likewise
composed wholly of extinct species, it must therefore be concluded
to be of middle Pliocene (Hemphillian), age or older.

Table 7.-PROPORTION OF EXTINCT SPECIES IN QUATERNARY AND LATE TERTIARY
AVIFAUNAS.

Percent
Age and Locality A authority Species Extinct

PLEISTOCENE:
Rancho La Brea, California.........
McKittrick, California ....... .....
Carpinteria, California .............
Fossil Lake, Oregon ...............
San Josecito, Nuevo Leon..........
LATE PLIOCENE (Blancan age):
Rexroad fauna, Kansas .............
Hagerman, Idaho..............
Benson, Arizona .................
MIDDLE PLIOCENE (Hemphillian age):
San Diego, California ..............
EARLY PLIOCENE (Clarendonian age):
Snake Creek, Nebraska............
AGE UNCERTAIN:
Bone Valley, Florida.. ............
LATE MIOCENE (Barstovian age):
Snake Creek, Nebraska............
MIDDLE MIOCENE (Hemingfordian age):
Sheep Creek, Nebraska.............
Calvert, Maryland............... ..
Lompoc, California .................
Sharktooth Hill, California. ......
EARLY MIOCENE (Arikareean age):
Flint Hill, South Dakota. ..........
Lower Harrison, Nebraska..........

Miller and DeMay
Miller and DeMay
Miller and DeMay
Howard (1946)....
L. Miller (1943)...

(1942)
(1942)
(1942)

(1944) ....
(1933)....
(1924)....

Howard (1949) ..........

Wetmore (1923).........

Brodkorb (1955).........

Wetmore (1923).........

Wetmore (1923, 1926A)...
Wetmore (1940A)........
Miller and DeMay (1942)
Miller and DeMay (1942)

A. H. Miller (1944)......
Wetmore (1933).........

114
73
58
70
39

11
10
6

8

2

18

3

3
5
6
3

9
9

15.8
13.7
17.2
24.3
30.5

63.6
70.0
83.3

100.0

100.0

100.0

100.0

100.0
100.0
100.0
100.0

100.0
100.0

Although
most of them

fifteen genera of birds are present in
have a long time span. There are three

the collection,
extinct genera,

Pliodytes, Palostralegus, and Australca, which are unknown from

-~--

Wetmorel~
Wetmore
W metmoree

FLORIDA GEOLOGICAL SURVEY

other localities. The other genera are still living in the Recent fauna
of Florida. One of the living genera, Bucephala, was previously re-
ported from the upper Pliocene of Kansas (Wetmore, 1944). The
remaining living genera have records extending back to the Miocene
or Oligocene. In the absence of indicators of a particular epoch, the
analysis of the genera merely limits the age of the deposit as being
Oligocene or later. Since, however, the Bone Valley beds are under-
lain by the Hawthorn formation of middle Miocene (Hemingfordian)
age, the oldest possibility for the Bone Valley must be late Miocene
or younger.
Three species in the Bone Valley collection help to restrict further
the age correlation of the avifauna, since they also occur in other de-
posits. These are Gavia concinna, Diomedea angelica, and Phalacro-
corax idahensis, all reported from localities referred to the Pliocene.
Gavia concinna Wetmore (1940A) was described from the Etche-
goin formation of California, referred by the describer to the lower
Pliocene. According to Woodring, Stewart, and Richards (1940: 112,
insert) and to Wood et al. (1941: 19), this formation is of middle
Pliocene (Hemphillian) age. G. concinna is also reported from the
San Diego formation in San Diego (Howard, 1949; Brodkorb, 1953C).
The San Diego faces of this formation is usually assigned to the
middle Pliocene, but Woodring, Stewart, and Richards (1940: 112)
give it a late early Pliocene age.
Diomedea angelica Lydekker (1891: 189) was described from the
upper Pliocene (Red Crag) of England. Lambrecht (1933: 273) lists
this species from both Plaisancian and Astian ages, which Wood et al.
(1940) correlate with the Hemphillian and Blancan, respectively, and
therefore of middle and late Pliocene age. The Florida record is
based on a tibiotarsus, an element unrepresented in the European
material, and therefore its reference to the present species is some-
what open to question.
Phalacrocorax idahensis (Marsh, 1870) was described from a sup-
posed Pliocene deposit of Idaho and has since been reported from
the upper Pliocene near Hagerman (Wetmore, 1933). As the Bone
Valley and Hagerman ulnas represent different ends of the element,
the reference is likewise not absolutely certain.
On the basis of the above criteria, the age of the Bone Valley
avifauna falls between the late Miocene and middle Pliocene, with
some evidence in favor of an early or middle Pliocene age. While

REPORT OF INVESTIGATIONS No. 14

this conclusion is in agreement with that reached by students of
the land mammals, it does not preclude the possibility that the verte-
brate chronology is not synchronous with the chronology based upon
marine invertebrates and beach lines.

PALEOECOLOGY
Census. A census of specimens and individuals for three Bone
Valley localities is given in Table 8. The minimum number of individ-
uals of each species was computed in the usual way, by counting
the right or left members of the most abundant element of that species
in the locality. In several cases the actual number of individuals was
probably greater.
White's locality is represented by only four specimens and four
individuals of three species. Locality 1 has 58 specimens and ten
individuals of six species. Locality 2 is the most prolific with 133
specimens anA 31 individuals of 16 species. The total material on
which this report is based thus includes 195 specimens from at
least 45 individuals and comprises a total of 18 species.
Dominance of species. The dominant species is the small cormo-
rant Phalacrocorax wetmorei, represented by no less than 135 speci-
mens. In Locality 1 83 percent of the specimens are of this species,
and 64 percent of those in Locality 2.
The next species in point of abundance is the auk Australca
grandis, with 18 specimens. Other relatively common forms are re-
presented by from three to six specimens. These include the two
loons (Cavia palaeodytes and G. concinna), the gannet (Morus penin-
sularis), the two boobies (Sula guano and S. phosphata), the flamingo
(Phoenicopterus floridanus), and the gull (Larus elmorei). These
eight species may all be classed as influent. They comprise 16 per-
cent of the specimens at Locality 1 and 30 percent at Locality 2.
Together the dominant and influent species make up 99 percent
of the collection from Locality 1 and 94 percent at Locality 2.
The remaining nine species may be classed as subinfluent. They
are represented by one or at most two specimens.
Habitat requirements. The entire avifauna of the Bone Valley
presents a fairly homogeneous aspect. All of the species are aquatic.
Further they are all representatives of groups which inhabit salt-
water exclusively or else frequent both salt and fresh water. The
six forms whose allies today are strictly confined to salt-water include

REPORT OF INVESTIGATIONS NO. 14

termined that all the alcid material represented a single species. It
gradually became apparent that the Bone Valley auk was a large
bird with the wings reduced in size compared with the living mem-
bers of the family. The reduction is especially evident in the distal
elements of the wing.
All alcids use the wings in swimming under water (Bent, 1919:
206). Cepphus, Uria, and Alca are strong fliers. The recently extinct
great auk, Pinguinus impennis, had lost the power of flight and had
carried the reduction of the distal wing elements to a remarkable
degree (see Table 6).
In Australca the reduction of the wings was about half-way be-
tween the condition in the flying alcids and Pinguinus. It was thus
already well on the road to flightlessness, and because of other similari-
ties it may even have been the ancestor of the great auk.

Table 6.-RATIOS (PERCENT) OF WING ELEMENTS TO LENGTH OF CORACOID

Ulna
Humerus depth Radius Second
Coracoid length to through distal meta-
length pectoral external width carpal
attachment cotyla length

Cepphus columba......... 100.00 181.61 27.42 16.13 88.71
Uria aalge ............... 100.00 187.18 27.69 14.62 79.23
Uria lomvia............. 100.00 185.75 27.75 15.50 78.00
Alca torda................ 100.00 168.99 29.61 15.36 78.77
Australca grandis.......... 100.00 149.32 23.20 12.22 63.80
Pinguinus impennis....... 100.00 134.62 21.79 9.77 43.22

AGE OF THE DEPOSIT

Geological opinion differs as to the age of the Bone Valley forma-
tion. Cooke (1945: 207) considered it middle Pliocene (Hemphillian
age). Cathcart (1950) allocated the formation to the lower part of the
Pliocene. From a study of the Pleistocene shore lines MacNeil (1950:
106) concluded that there is a possibility that the Bone Valley gravel
might be early Pleistocene (Aftonian), but he too accepted a Pliocene
age. Currently (in litt., January 18, 1954) he favors uppermost Mio-
cene for the upper Bone Valley and middle Miocene (Hawthorn)
for the lower Bone Valley. Vernon (1943: 156) first assigned the

29

Table 8.--('ENSL'S (OF BIRD OF THE BONE VALLEY FORMATION.

Species

Gavia paleodytes..........
Gavia concinna ...........
Pliodytes lanquiti. .......
Diomedea angelica .........
Morus peninsularis.......
Sula guano .............
Sula phosphata...........
Phalacrocoraz wetmorei....
Phalacrocorax idahensis....
Ardea polkensie........
Phrnicopterus .loridanus...
Bucephala ossivallis.......
Palostralegue sulcats ......
Calidris paci............
Erolia penepusilla ........
Limosa sp.................
Larue elmorei............
Australca grandis. ........

LOCALITY 1

Speci- Indi-
nmenr viduals

1 1
3 1
0 0
0 0
0 0
0 0
O 0
48 5
0 0
0 0
0 0
0 0
1 1
0 0
0 0
0 0
2 1
3 1

58 10

LOCALITY 2

Speci- Indi-
mens vidnal.i

4
2
1
0
3
4
4
85
1

1.

2
4
1
0
1
1
2
4
15

1
1
1
0
2
2
3
8
1
1
2
1
0
1
1
1
1
4

133 31

WHITE'S
LOCALITY

Speci-
mens

1
0
0
1
0
0
0
2
0
0
0
0
0
0
0
0
0
0

4

TOTAL

Speci- Indi-
mens viduals

6 3
5 2
1 1
1 1
3 2
4 2
4 3
135 15-
1 1
1 1
4 2
1 1
1 1
1 1
1 1
2 1
6 2
18 5

195 45

0

0
m
Mn

...................
..................
...................
...................
...................
...................
...................
..................
.............
...................
...................
...................
...........
. . . . . .
...................
...................
...................

_ ~I~ ______

I_________ ___ _~ ~1-1~1_-_-----_

__ __

Indi-
viduals

1
0
0
1
0
0
0
2
0
0
0
0
0
0
0
0
0
0

4

- -----------------'

,I-

REPORT OF INVESTIGATIONS No. 14

the halobionts Diomedea, Morus, Sula guano, S. phosphata, Palos-
tralegus, and Australca. The remaining twelve forms are halocoles, as
their present-day allies are tolerant of both fresh water and salt,
although most of them are perhaps more numerous in the latter
environment.
Wynne-Edwards (1935: 240) has classified the ecological require-
ments of the oceanic birds of the North Atlantic, and Murphy (1936:
326) has made a similar classification for those of South America.
The characteristic birds of the divisions of the ocean, based on the
above-mentioned authors with additional data from Bent (1919 et
seq.), are as follows:
1. Littoral (beaches and rocky foreshores): sandpipers, plovers,
oystercatchers, herons, flamingoes.
2. Inshore (within sight of land): loons, grebes, some cormorants,
sea-ducks, most gulls, skimmers.
3. Offshore (to edge of continental shelf): some diving petrels,
gannets, boobies, some cormorants, pelicans, auks.
4. Pelagic (open ocean): penguins, petrels, shearwaters, albatros-
ses, tropic-birds, skuas, jaegers, phalaropes.
On the basis of this division there are present in the Bone Valley
avifauna six littoral species, Phoenicopterus, Ardea, Palostralegus,
Calidris, Erolia, and Limosa. Only the first of these is an influent
species, the others being subinfluent.
The seven inshore forms are the two species of Gavia, Pliodytes,
two species of Phalacrocorax, Bucephala, and Larus. This group in-
cludes all the dominant Bone Valley species, three influents, and
three subinfluents.
The offshore group contains four birds, all influents. They are
Morus, the two species of Sula, and Australca.
A single pelagic bird, Diomedea, is recorded from the Bone Valley.
It is subinfluent.
The above classification is based on the feeding habits of the birds
during the non-breeding season. It must be remembered that during
the time of nesting they customarily feed farther inshore. There-
fore the presence together in the Bone Valley of species from all four
ecological divisions of the ocean may best be interpreted as a group-

FLORIDA GEOLOGICAL SURVEY

ing of some of them at a coastal or insular breeding site. Further-
more, it is hardly possible to account for such large numbers of
bones being accumulated in any other manner, particularly in the
case of Phalacrocorax wetmorei. I believe that more specimens are
known of this cormorant than of any other Tertiary bird.

Effect of bird life on the production of phosphate. With the ex-
ception of the two loons all of the dominant and influent species, as
well as two of the subinfluents (Diomedea and Phalacrocorax ida-
hensis) are social birds which nest in large colonies.

The presence of large numbers of sea-bird fossils in the phosphate
beds immediately brings to mind the guano islands off the coast of
Peru and other parts of the world. Large rookeries of birds as a
source for the Florida phosphate were suggested long ago by Sellards
(1913: 45), and more recently Vernon (1951: 195-198) has elaborated
on this hypothesis.

The important guano-producing birds today are members of the
order Sphenisciformes, Diomedea among the Procellariiformes, the
order Pelecaniformes, and the families Laridae and Alcidae among
the Charadriiformes (Hutchinson, 1950: 366). According to Murphy
(1936: 293) the Pelecaniformes, in particular species of the genera
Phalacrocorax and Sula, are the most important contributors to the
formation of guano on the Peruvian islands today. Coker (1919)
reports high concentrations of phosphoric acid in cormorant and
pelican guano.

In both their past and present distribution the penguins (Sphenis-
ciformes) are confined to the southern hemisphere and therefore do
not enter into the picture here.

Diomedea is represented in the Bone Valley by one subinfluent
species. The albatrosses today are limited to southern oceans and
the North Pacific and are only of accidental occurence in the North
Atlantic, but are recorded from three British localities of Pliocene
and Pleistocene age (Lambrecht, 1933: 273, 732).

Five of the eighteen Bone Valley species are members of the
Pelecaniformes. This order includes the dominant Phalacrocorax wet-
morei, three of the influent species (Morus peninsularis, Sula guano,
and S. phosphata), and the subinfluent Phalacrocorax idahensis. It is
noteworthy that the two genera which Murphy considers the most

REPORT OF INVESTIGATIONS No. 14

this conclusion is in agreement with that reached by students of
the land mammals, it does not preclude the possibility that the verte-
brate chronology is not synchronous with the chronology based upon
marine invertebrates and beach lines.

PALEOECOLOGY
Census. A census of specimens and individuals for three Bone
Valley localities is given in Table 8. The minimum number of individ-
uals of each species was computed in the usual way, by counting
the right or left members of the most abundant element of that species
in the locality. In several cases the actual number of individuals was
probably greater.
White's locality is represented by only four specimens and four
individuals of three species. Locality 1 has 58 specimens and ten
individuals of six species. Locality 2 is the most prolific with 133
specimens anA 31 individuals of 16 species. The total material on
which this report is based thus includes 195 specimens from at
least 45 individuals and comprises a total of 18 species.
Dominance of species. The dominant species is the small cormo-
rant Phalacrocorax wetmorei, represented by no less than 135 speci-
mens. In Locality 1 83 percent of the specimens are of this species,
and 64 percent of those in Locality 2.
The next species in point of abundance is the auk Australca
grandis, with 18 specimens. Other relatively common forms are re-
presented by from three to six specimens. These include the two
loons (Cavia palaeodytes and G. concinna), the gannet (Morus penin-
sularis), the two boobies (Sula guano and S. phosphata), the flamingo
(Phoenicopterus floridanus), and the gull (Larus elmorei). These
eight species may all be classed as influent. They comprise 16 per-
cent of the specimens at Locality 1 and 30 percent at Locality 2.
Together the dominant and influent species make up 99 percent
of the collection from Locality 1 and 94 percent at Locality 2.
The remaining nine species may be classed as subinfluent. They
are represented by one or at most two specimens.
Habitat requirements. The entire avifauna of the Bone Valley
presents a fairly homogeneous aspect. All of the species are aquatic.
Further they are all representatives of groups which inhabit salt-
water exclusively or else frequent both salt and fresh water. The
six forms whose allies today are strictly confined to salt-water include

REPORT OF INVESTIGATIONS No. 14

important ip the production of guano make up 83 percent of the
collection from Locality 1 and 70 percent from Locality 2.
The Laridae and the Alcidae are each represented in the Bone
Valley by an influent species, Larus elmorei and Australca grandis.
Thus all the dominant and influent species of the Bone Valley,
except the two loons and the flamingo, are guano birds. The eight
guano species comprise 44 percent of the avifauna and 88 percent
of the specimens.
The loons today are not gregarious birds and they occur this far
south only in the non-breeding season. There is no reason to believe
that their habits during the Pliocene were markedly different from
those of the present time.
The formation of guano has on several occasions been attributed
to the flamingoes, but according to Hutchinson (1950: 36, 43, 336)
the cases are not well authenticated. Flamingoes nest on tidal flats,
and it seems probable that their excrement would become dissolved
in the sea-water. Thus while not preserved as guano it would never-
theless help to raise the phosphorus content of the water.
Hutchinson states that marine deposits of phosphate are always
intimately associated with a rich supply of phosphorus from the
land. He continues (p. 373):
"The result of a very large bird colony on a section of coast line
or on an island, whenever climatic conditions and the form of the
substrate permit guano to be returned to the ocean will be to steepen
the nutrient gradient The result will be increased littoral pro-
ductivity and a steady state condition will be set up."
Vernon (1951: 197) suggested that limestone islands with bird
rookeries on them existed in Florida during the Miocene and younger
epochs and formed a ready source of phosphoric acid. Now for the
first time the data are at hand to support this hypothesis.
Whatever the original source of phosphorus in Florida waters may
have been, it would increase the production of phytoplankton, and
this in turn would increase the production of marine invertebrates
and fish on which large bird populations might subsist. Excrement
from the birds would return the phosphorus to the sea and complete
the cycle, attaining an equilibrium.
The general picture derived from a study of the avifauna is of
a guano island near the coast, probably rocky in character. Nesting
on its shores were myriads of cormorants, similar in size to the

38 FLORIDA GEOLOGICAL SURVEY

present-day species. The rookery was also populated by colonies, of
other sea-birds, including a gannet, two species of boobies, and a
large auk, the last already on the road to flightlessness. A second
giant species of cormorant was present in lesser numbers. Flocks of
flamingoes occurred on the tidal flats, where an occasional large'
heron stalked its prey. A medium-sized gull was common and prob-
ably robbed the eggs of the other birds. Along the beach an oyster-
catcher and three species of sandpipers rested and fed. In the waters
off-shore two species of loons were fairly common, and a grebe and
a sea-duck also occurred in lesser numbers off-shore. From still
farther distant an occasional albatross appeared, perhaps attracted
by the teeming invertebrate life, which fed on the plankton fertilized
by the droppings of the nesting birds.

SUMMARY
Eighteen species of birds represented by nearly 200 specimens are
recorded from the Bone Valley formation in Polk County, Florida,
making this the largest North American Tertiary avifauna. The fol-
lowing species are described as new in the present paper: Morus
peninsularis, Sula guano, Sula phosphata, Phalacrocorax wetmorei,
Ardea polkensis, Bucephala ossivallis, Palostralegus sulcatus (new
genus and species), Calidris pacis, Erolia penepusilla, and Australca
grandis (new genus and species). Three other new species were
described in preliminary papers.
Criteria used for the determination of the age of the Bone Valley
avifauna are the proportion of extinct species, the maximum known
age of the various genera, and the presence of index species. A com-
parison of the proportion of extinct versus living species in .major
Quaternary and late Tertiary avifaunas shows the Pleistocene with
about 14-24 percent extinct species, the upper Pliocene with about
64-83 percent extinct, and the Bone Valley and all localities of middle
Pliocene age and older composed entirely of extinct species. Thus
the Bone Valley cannot be younger than middle Pliocene. The con-
clusion derived from study of the age of the genera and from the
stratigraphy indicate that the avifauna cannot be older than late
Miocene. Three species of Bone Valley birds are reported elsewhere
from early, middle, or late Pliocene deposits. The avifauna, therefore,
must be of late Miocene to middle Pliocene age, and the agreement
is closest to other avifaunas recorded from the early or middle parts
of the Pliocene. These conclusions are in agreement with the deduc-

REPORT OF INVESTIGATIONS No. 14

tions of those who have studied the land mammals, but there is a
possibility that the land vertebrate chronology is a portion of an
epoch ahead of the chronology based upon marine invertebrates.
The avifauna is composed of one dominant, eight influent, and
nine subinfluent species. Classified according to feeding habitat, there
are six littoral species, seven inshore forms, four in the offshore group,
and a single pelagic bird. The association of members of these four
groups is explained as representing a breeding colony, and this fur-
ther explains the presence of large numbers of individuals, one species
being represented by 135 specimens.
Groups important today in the production of guano comprise 44
percent of the species and 88 percent of the specimens from the
Bone Valley. Whatever the original source of the phosphorus in
Florida waters may have been, the large colonies of sea-birds added
materially to it and set up stable conditions which may have con-
tinued for a long time.

39

A?

38 FLORIDA GEOLOGICAL SURVEY

present-day species. The rookery was also populated by colonies, of
other sea-birds, including a gannet, two species of boobies, and a
large auk, the last already on the road to flightlessness. A second
giant species of cormorant was present in lesser numbers. Flocks of
flamingoes occurred on the tidal flats, where an occasional large'
heron stalked its prey. A medium-sized gull was common and prob-
ably robbed the eggs of the other birds. Along the beach an oyster-
catcher and three species of sandpipers rested and fed. In the waters
off-shore two species of loons were fairly common, and a grebe and
a sea-duck also occurred in lesser numbers off-shore. From still
farther distant an occasional albatross appeared, perhaps attracted
by the teeming invertebrate life, which fed on the plankton fertilized
by the droppings of the nesting birds.

SUMMARY
Eighteen species of birds represented by nearly 200 specimens are
recorded from the Bone Valley formation in Polk County, Florida,
making this the largest North American Tertiary avifauna. The fol-
lowing species are described as new in the present paper: Morus
peninsularis, Sula guano, Sula phosphata, Phalacrocorax wetmorei,
Ardea polkensis, Bucephala ossivallis, Palostralegus sulcatus (new
genus and species), Calidris pacis, Erolia penepusilla, and Australca
grandis (new genus and species). Three other new species were
described in preliminary papers.
Criteria used for the determination of the age of the Bone Valley
avifauna are the proportion of extinct species, the maximum known
age of the various genera, and the presence of index species. A com-
parison of the proportion of extinct versus living species in .major
Quaternary and late Tertiary avifaunas shows the Pleistocene with
about 14-24 percent extinct species, the upper Pliocene with about
64-83 percent extinct, and the Bone Valley and all localities of middle
Pliocene age and older composed entirely of extinct species. Thus
the Bone Valley cannot be younger than middle Pliocene. The con-
clusion derived from study of the age of the genera and from the
stratigraphy indicate that the avifauna cannot be older than late
Miocene. Three species of Bone Valley birds are reported elsewhere
from early, middle, or late Pliocene deposits. The avifauna, therefore,
must be of late Miocene to middle Pliocene age, and the agreement
is closest to other avifaunas recorded from the early or middle parts
of the Pliocene. These conclusions are in agreement with the deduc-

PLATES

41

Explanation of Plate I

Figure 1.

Morus peninsularis n. sp. No. 148, type. External view of

coracoid.

Figure 2.

Figure 3.

Sula guano n. sp. No. 301, type. External view of coracoid.

Sula phosphata n. sp. No. 302, type. External view of coro-

coid.

Figure 4. Morus peninsularis n. sp. No. 148, type. Distal view of

coracoid.

Figure 5.

Figure 6.

Sula guano n. sp. No. 301, type. Distal view of coracoid.

Sula phosphata n. sp. No. 302, type. Distal view of cora-

coid.

All figure approximately X 121.i

Plate 1

I__II _I L _II 111

"' ~%'~'
'I
~ ~.I~ NI I~
:.
~.
~;L; ;:
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s:5:: ..

d~ ~ ,. ? '.~
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..

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.
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":
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.: ... ::. ~.

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:

i ':: ''T''~

:

Z'"x".~.: :

Ir.~ t :.~ :
: :

I '
i
:'
i
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i .

i: i: r l'i:

;!
:...~

.;
:
...:~
~ '' ':
'." '
:.
i ~ : ~;...:i.
"'

''''~
i.
:':'~ ': v

4:'.:, .i

j

: ;1

..i

;

1

ut:~ 4
'" .:
:%
r;r ;*
::.':. :

~~ .~i:~ ` :" ~~':: : :~ : ~i :

.:t r: ;:,
~: ~~:~.: 2~":''11
.. ::~ ~^i

c : ~~' .' ~~;~7
: ~~, :
'

tr

i.. ~;

: .. ~ .?

:
'"
'

"

~Bi~
~. ~

.

:: .:

~ :~;

ii;
;, r.~ ,
;I..... '~
'Z~'~~ :

b

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ci;

;:~ .~i::

: ~~I.,.

13r~ :....':. '

~' ~:

'''~ ;

Plate 2

v

Explanation of Plate 2
Figure 7. Morus peninsularis n. sp. No. 148, type. Internal view of
coracoid.
Figure 8. Sula guano n. sp. No. 301, type. Internal view of coracoid.
Figure 9. Sula phosphata n. sp. No. 302, type. Internal view of cora-
coid.
All figures approximately X 1%.

w
^

Plate 3

.. .... 1 0 ..
10

I

12

Explanation of Plate 3
Figure 10. Phalacrocorax wetmorei n. sp. No. 530, type. Coracoid.
Figure 11. Phalacrocorax wetmorei n. sp. No. 124. Ulna.
Figure 12. Phalacrocorax idahensis (Marsh). No.' 311. Ulna.
All figures approximately X 11/2.

. :

Plate 4

14
14

15

Explanation of Plate 4
Figure 13. Ardea polkensis n. sp. No. 308, type. Anterior view of
tarsometatarsus.
Figure 14. Ardea polkensis n. sp. No. 308, type. Proximal view of
tarsometatarsus.
Figure 15. Ardea polkensis n. sp. No. 308, type. Posterior view of
tarsometatarsus.
All figures approximately X 2.

Plate 5

16

Figure 16.

Figure 17.

17

Explanation of Plate 5
Bucephala ossivallis n. sp. No. 172, type. Internal view of
coracoid.
Bucephala ossivallis n. sp. No. 172, type. External view of
coracoid.

All figures approximately X 4.

Plate 6

19

Figure 18.

Figure 19.

Figure 20.

20

Explanation of Plate 6
Palostralegus sulcatzus n. g. et sp. No. 177, type. Tibio-
tarsus.
Calidris pacis n. sp. No. 594, type. Palmar view of hum-
erus.
Calidris pacis n. sp. No. 594, type. Anconal view of hum-
erus.

All figures approximately X 3.

~i s: j., ,
"
'
11
I
II

ff%

.2

Explanation of Plate 7

Figure 21. Erolia penepusilla n. sp. No. 611, type. Humerus.

Figure 22. Limosa sp. No. 526. Distal view of tibiotarsus. .

Figure 23. Limosa sp. No. 526. Anterior view of tibiotarsus.

All figures approximately X 4.

49

N
NY

N

N

i
-;- -- :.'ic:

r:

,~irsl

VT

Plate 8

h'r .~, I ., f
Ir If
I
I
i
J
L
,,I t
1

J

24

25

Explanation of Plate 8
gure 24. Australca grandis n. g. et sp. No. 141, type. Internal view
of coracoid.
gure 25 Australca grandis n. g. et sp. No. 141, type. External view
of coracoid.
All figures approximately X 2.

Fi

Fi

Plate 9

'i
i
.'t
i
e;

r1
i
;i
1
i
6,l;i

26
28

27

29

Explanation of Plate 9
Figure 26. Australca grandis n. g. et sp. No. 310. Anconal view of
humerus.
Figure 27. Australca grandis n. g. et sp. No. 304. Anconal view of
humerus.
Figure 28. Australca grandis n. g. et sp. No. 304. Palmar view of
humerus.
Figure 29. Australca grandis n. g. et sp. No. 612. Tibiotarsus.
All figures approximately X 2.

Plate 10

r 4.
:1 3
rr
;L ~
I- t?:; .z~b. +
IFC1I
,*
II
r"' *- C 'v
:k~t~ P Z ~i~ir~i~r~
~t~~ r
1JI ~r

Explanation of Plate 10

Figures 30-31. Bird bones in rookery key, Cuthbert Lake, Ever-

glades National Park. Figure 30 is at the top.

Plate 11

r; 9 ,
0.r
a''
"'"r~ r
r
4 Cr,'r'.p ?f.k
,~*~:ni I % ua~
'4' I II. it' -E~~ ~; LO'. *
.I Z' '~-
~.a-~-1;I-.
I
~r A c
i r. I'L
t r.'
)i:
CI: .IS ~' I .111
;: i b a
-r
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: :
~s~P i .*i;.: (L'i:L". ~C~~fe*ylq l~i~i~ll!~LII I11 ~1
,,a~,ci a~ w;
\*~ ,,~ ~~~4.f.~ ib'~~~4~2 -'.t r"~
!;r
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ir
rl r .F rii~S1 m :iir~
i.
t: ~lj rl;, ~1 ...I '44
itf~y .r IP
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-I 1~
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ir i i-r---r 'r'
jr p~'tO r t.. rr r r
~I .i~i .. .)' .,
r i~i; r~ 'r .
:~;~1 4 I;
I. ~
..bb~ L.
r -i
'=lf rbrS 9 i~t' )\ ~T,~n ?TC;:
~3~ ,1 .. n~.. :
;ca f"' (I r -C14: ~!~L~a:is~
~ I~r q)': 11:' i
~ arYi~
I i
,. .. r.. .. i ,
81j(,ed ~, ;Ira, rs I; ~ ;is "~~":;.. -.o
'iil-szk.r.::.~,yn;Z1 .
.11~L~Bf:~i~rr (=r ~~ .: ,~r ~~p.'i~;.i`rii k~?n I -;i

Figure 32.

Explanation of Plate 11

Bird bones in rookery key, Cuthbert Lake, Everglades

National Park.

REPORT OF INVESTIGATIONS No. 14

LITERATURE CITED
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1921. Fossil cetaceans from the Florida phosphate beds. Journ. Mamm., 2 (3):
144-159, pl. 9-12.
Bent, Arthur Cleveland
1919. Life histories of North American birds. Order Pygopodes.
U. S. Nat. Mus. Bull. 107. Pp. xiii, 245, 55 pl.
Brodkorb, Pierce
1953A. A Pliocene flamingo from Florida. Nat. Hist. Misc., No. 124: 1-3, fig. 1-2.
1953B. A Pliocene gull from Florida. Wilson Bull., 65 (2): 94-98, fig. 1.
1953C. A review of the Pliocene loons. Condor, 55 (4): 211-214, fig. 1.
1953D. A Pliocene grebe from Florida. Annals and Mag. Nat. Hist., ser. 12, 6:
953-954, 1 fig.
Case, E. C.
1934. A specimen of a long-nosed dolphin from the Bone Valley gravels of
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2 pl.
Cathcart, J. B.
1950. Notes on the land-pebble phosphate deposits of Florida. In Symposium
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Coker, Robert E.
1919. Habits and economic relations of the guano birds of Peru. Proc. U. S.
Nat. Mus., 56: 449-511, pl. 53-69.
Cooke,IC. Wythe
1945. Geology of Florida. Florida Geol. Surv., Geol. Bull., 29. Pp. ix, 339, 1 pl.,
S47 fig.
Cope, D. D.
1870. Synopsis of the extinct Batrachia, Reptilia and Aves of North America.
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1878. Descriptions of new Vertebrata from the Upper Tertiary and Dakota forma-
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Ennouchi, Emile
1930. Contribution a l'6tude de la faune du Tortonien de La Grive-St-Alban
(Isere). Revision Generale.-Etude Ornithologique. Th6ses Faculte des
Sciences de Paris, Ser. A, No. 1266. Pp. 135, 6 pl. Paris, Presses Modernes.
Gervais, P.
1851. Mem. Acad. Montpellier, 1: 220 (cited from Lambrecht).
Hay, 0. P.
1902. On the finding of the bones of the great auk (Plautus impennis) in Flor-
ida. Auk, 19 (3): 255-258.
1922. Description of a new fossil sea cow from Florida, Metaxytherium flori-
danum. Proc. U. S. Nat. Mus., 61, art. 17: 1-4.
Howa::d, Hildegarde
1938. A new fossil bird locality near Playa del Rey, California, with description
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Wash. Publ. 551: 141-195, 2 pl.
1949. New avian records for the Pliocene of California. Carnegie Inst. Wash.
Publ. 584: 177-199, 3 pl.
Hutci'nson, George Evelyn
19650. Survey of contemporary knowledge of biochemistry. 3. The biogeo-
chemistry of vertebrate excretion. Bull. Am. Mus. Nat. Hist., 96. Pp.
554, 16 pl.
Kellogg, A. Remington

FLORIDA GEOLOGICAL SURVEY

1924. Tertiary pelagic mammals of eastern North America. Bull. Geol. Soc.
Amer., 35 (4): 755-766.
Lamnbrecht, Kalman
1933. Handbuch der Palaeornithologie. Pp. xx, 1024, 209 fig. Berlin, Gebriider
Borntraeger.
Leidy, J.
1889. Description of vertebrate remains from Peace Creek, Florida. Trans.
Wagner Free Inst. Sci., 2: 19-31, pl. 3-6.
Lydekker, Richard
1891. Catalogue of the fossil birds in the British Museum. Pp. xxvii, 368, 75
fig. London, Taylor and Francis.
MacNeil, F. Stearns
1950. Pleistocene shore lines in Florida and Georgia. Geol. Surv. Prof. Paper
221-1': 95-107, pl. 19-25.
Marsh, 0. C.
1870. Notice of some fossil birds, from the Cretaceous and Tertiary formations
of the United States. Am. Journ. Sci. Arts, ser. 2, 49 (146): 205-217.
Matson, George Charlton, and Frederick G. Clapp
1909. A preliminary report on the geology of Florida, with special reference
to the stratigraphy. Florida Geol. Surv. Ann. Report, 2: 25-178, map.
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FLRD GEOLOSk ( IC SUfRiW

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	Title Page
	Transmittal letter
	Contents
	Illustrations
	Introduction
	List of species
	Age of the deposit
	Paleoecology
	Summary
	Plates
	Literature cited

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