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Group Title: Two mid-Pleistocene avifaunas from Coleman, Florida (FLMNH Bulletin v.26, no.1)
Title: Two mid-Pleistocene avifaunas from Coleman, Florida
Full Citation
Permanent Link: http://ufdc.ufl.edu/UF00095812/00001
 Material Information
Title: Two mid-Pleistocene avifaunas from Coleman, Florida
Physical Description: 36 p. : ill. ; 23 cm.
Language: English
Creator: Ritchie, Thomas L.
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1980
Copyright Date: 1980
Subject: Birds, Fossil   ( lcsh )
Paleontology -- Pleistocene   ( lcsh )
Paleontology -- Florida -- Coleman   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
Bibliography: Bibliography: p. 35-36.
General Note: Bulletin of the Florida State Museum, Volume 26, Number 1
Statement of Responsibility: Thomas L. Ritchie.
 Record Information
Bibliographic ID: UF00095812
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 08218957
lccn - 81623585

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Full Text

of the

Biological Sciences


Number 1




Volume 26


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Copyright 1980 by the Florida State Museum of the University of Florida

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SYNOPSIS: Avifaunas from two sites near Coleman, Florida, are Irvingtonian in age and con-
tain a total of 38 species. The two sites, Coleman IIA and Coleman IIIC and D, are probably
related temporally, but they are analyzed separately. Ten species (Anhinga cf. A. grandis,
Buteo lagopus, Aquila chrysaetos, Falco columbarius, Bubo virginianus, Protocitta cf. P.
ajax, Dendroica sp., Pheucticus ludovicianus, Pooecetes gramineus, and Aimophila aestivalis)
are new to the Pleistocene of Florida. Pheucticus ludovicianus and Aimophila aestivalis are
new to the fossil record. New material of two populations of Colinus suilium, one early to
middle Pleistocene in age (from the Inglis IA site in Citrus County, Florida) and the other
middle Pleistocene in age (from Coleman IIA), is analyzed, and indicates C. suilium under-
went a gradual size increase throughout the Pleistocene. This is the opposite of what was
previously thought and suggests that C. suilium was not derived from Colinus hibbardi as has
been hypothesized.


INTRODUCTION .............. . ................................................ 2
ACKNOWLEDGMENTS .... ........ ................................................. 3
SYSTEM ATIC L IST .............. .. .............................................. 4
Coleman IIA ............................................... ... 24
Coleman III ................... ..... ........................ 28
PALEOECOLOGY ...... ............................................................. 32
EVOLUTIONARY TRENDS ...... ........ .............................................. 33
M ODE OF FOSSIL DEPOSITION ............. . ................................. .... 34
L ITERATURE C ITED ............... .............................................. 35

'The author is currently a Naturalist/Guide on board the M.S. Lindblad EXPLORER, as well as a pro-
fessional wildlife artist based in Palm Beach, Florida. This study was performed while he was a Graduate
Assistant in the Department of Zoology, University of Florida, Gainesville 32611. It represents, in part, a
thesis prepared in partial fulfillment of the requirements for the degree Master of Science.

RITCHIE, THOMAS L. 1980. Two Mid-Pleistocene Avifaunas from Coleman, Florida.
Bull. Florida State Mus., Biol. Sci. 26(1): 1-36.


Two fossil deposits collected near Coleman, Sumter County, Florida,
have proved important in the study of middle Pleistocene birds and mam-
mals of Florida. The first, Coleman IIA, was discovered in 1966 during
limestone mining operations at the Coleman II mine of the Dixie Lime
and Stone Corporation. The site was collected by Norm Tessman, S.
David Webb, Robert Allen, Charles Collins, and Sue Hirschfeld (Webb
1974) and was destroyed that same year by the ensuing mining opera-
The deposit represents a sinkhole filling that extended into late Eo-
cene Ocala limestone; it was deposited during the late Illinoian Glacial
Stage (Webb 1974). Allen (in Martin 1974) gave the following description
of the site:
The opening of the paleosink was approximately 30
yards long and 25 yards wide. Only the upper 8 to 10 feet
of sediments filling the sink were exposed. Thin alter-
nating beds of two different types appeared: a wet
orange-brown clay and a coarse gray-white sand.
Although the clay facies appeared homogeneous, the
sand facies were filled with pebble-to-boulder-sized
limestone rubble. Bones found in the clay were
characteristically colored white or tan, with all grada-
tions in between; the bone found in gray-white sand was
characteristically black, but in a few instances, white.
Bones often transgressed these lithologic units and are
characteristically colored half tan and half black, testify-
ing to relatively rapid accumulation of successive
lithologic units.
A study by Tessman (Martin 1974) indicated a large number of the
bigger mammalian bones have "green breaks," meaning breaks that oc-
curred before fossilization. No evidence indicates carnivores caused these
breaks, so it is assumed that fossil remains from some of the larger mam-
mals were accumulated by their falling a considerable distance into the
sinkhole. To verify this hypothesis he made comparisons of the Coleman
IIA fossils with freshly broken bones and bones recently fallen into a
sinkhole 65 feet deep.
Martin (1974) reported on the mammalian fauna of Coleman IIA, and
noted that bat remains, especially Myotis cf. M. austroriparius, indicated
the sink was "extensive and not filled with water." Martin identified 37
mammalian species, including 5 species (Dasypus bellus, Pampatherium
septentrionalis, Palaeolama mirifica, Platygonus cumberlandensis, and
Arctodus pristinus) indicative of a late Irvingtonian age.

Vol. 26, No. 1


In 1968 Robert Martin and H. Kelly Brooks discovered and collected
another site near Coleman, Coleman III. According to Brooks (pers.
comm.) this site was approximately 300 meters due north of Coleman IIA,
in the Coleman III mine of the Dixie Lime and Stone Corporation.
Unlike the karst deposit of IIA, Coleman III was a small spring deposit
that has been dry since the middle Pleistocene (Brooks, pers. comm.). The
site was divided into four horizontal sections, A, B, C, and D. A sample
from IIIA (the spring mouth) yielded a light gray, fine-grained marl with
freshwater snail shell fragments. IIIB (Viviparus zone) consisted of a
brownish-gray, fine grained sandstone with slightly darker silt stringers
throughout, and many snail shell fragments, mostly of Viviparus
georgianus. IIIC is identical to IIIA. Unfortunately no substrate samples
were collected from IID. This site, too, was later destroyed by mining
The bird fossils of Coleman III were found only in IIIC and IID.
Fourteen non-avian vertebrates (Amia calva, Micropterus sp., Rana sp.,
Chelydra sp., Trionyx ferox, Terrapene carolina, Chrysemys concinna,
Alligator mississipiensis, Crotalus adamanteus, Pampatherium septen-
trionalis, Proboscidea, Hemiauchenia cf. H. macrocephala, Tapirus sp.,
and Equus sp.) are identified from IIIC. The non-avian vertebrates iden-
tified from IID are Chrysemys scripta, Chrysemys concinna, Chrysemys
cf. C. floridanus, and Sigmodon sp.
Both Coleman IIA and Inglis IA, a more recently discovered
Pleistocene site of central Florida, have yielded large samples of the ex-
tinct quail Colinus suilium. Holman (1961) made a thorough study of the
osteology of the New World quails, and the samples of these two popula-
tions of C. suilium have added to the knowledge of evolution of Colinus in
North America.
Inglis IA has been dated to both the late Blancan age (Klein 1971) and
the early Irvingtonian age (Webb 1974). No matter which assessment of
its age is correct, Inglis IA is considerably older than Coleman IIA. Inglis
IA is located along the Gulf Coast near Inglis, Citrus County, Florida, at
about the present sea level. Jean Klein and Robert Martin discovered this
extremely fossiliferous sandy fissure-fill deposit (Webb 1974), and Klein
(1971) made a thorough study of its geology, paleoecology, and
Material from the three localities (Coleman IIA, Coleman IIIC and
D, and Inglis IA) is presented below. Osteological terminology follows
that of Howard (1929). All specimens are in the Florida State Museum.
Table 1 lists the bird species from Coleman, Florida.
I wish to express my deepest thanks to Pierce Brodkorb for his invaluable advice and
supervision during this study. H. Kelly Brooks answered numerous questions concerning the



two Coleman sites and was especially helpful in describing Coleman III. Jon Baskin assisted
in the description of the geology of Coleman IIIA, B, and C, and S. David Webb, H. Gregory
McDonald, and Michael Frazier provided much helpful aid and technical advice.
Financial support was provided through Teaching Assistantships with the Department of
Zoology, University of Florida. Essential comparative skeletal material was studied on loan
from Pierce Brodkorb of the Department of Zoology, University of Florida, and Storrs L.
Olson of the Smithsonian Institution.

Table 1.-Pleistocene birds from Coleman, Florida.

Podilymbus podiceps
Botaurus lentiginosus
Ixobrychus sp.
Ciconia maltha weillsi
Anas platyrhynchos
Branta canadensis cf. B. c. hutchinsii
Coragyps occidentalis
Buteo platypterus
Buteo jamaicensis
Buteo lagopus
Aquila chrysaetos
Falco columbarius
Colinus suilium
Agriocharis anza
Rallus elegans
Coturnicops noveboracensis
Philohela minor
Zenaida macroura
Bubo virginianus
Otus asio
Colaptes auratus
Progne subis

Protocitta cf. P. ajax
Dendroica sp.
Agelaius phoeniceus
Pandanaris floridana
Pheucticus ludovcianus
Pipilo erythrophthalmus
Passerculus sandwichensis
Pooecetes gramineus
Aimophila aestivalis
Spizella pusilla
Spizella sp.
Emberizinae, genus and species indeterminate

Anhinga cf. A. grandis
Ardea herodias
Aythya affinis
Aythya sp.
Gallinula chloropus
Agriocharis anza





Podilymbus podiceps (Linnaeus), Pied-billed Grebe
MATERIAL -Tarsometatarsus (UF 11816).
REMARKS -Podilymbus is distinguished from Podiceps by the nar-
rower and deeper proximal end of the tarsometatarsus; ridges surround-
ing both cotylae less pronounced; shaft wider and stouter, with the ridge

Vol. 26, No. 1


on the internal side of anterior metatarsal groove not as well developed;
calcaneal ridges shorter, with inner and outer proximal foramina aligned
approximately side by side (in the tarsometatarsus of Podiceps the outer
proximal foramen is much lower than the inner one).

Table 2.-Tarsometatarsal measurements (in mm), with mean and range, of Recent
Podilymbus podiceps podiceps (Brodkorb Collection) and fossil Podilymbus
podiceps from Coleman IIA (UF 11816).

Proximal Width Proximal Depth
female (5) 7.1 (6.9-7.5) 6.2 (5.8-6.8)
male (3) 7.9 (7.8-8.1) 6.8 (6.3-7.3)
overall 7.4 (6.9-8.1) 6.4 (5.8-7.3)
UF 11816 9.1 7.5

The single tarsometatarsus was not completely ossified before
fossilization took place, which shows it was from an immature bird. In
spite of this, the specimen is considerably larger than the tarsometatarsi of
all recent P. podiceps specimens it was compared with (Table 2). Storer
(1976) demonstrated that P. p. magnus, a supposedly large Pleistocene
subspecies, is totally untenable, for its size falls within the normal size
variation for living P. p. podiceps. The poor state of the fossil does not
permit comparing its overall size with the measurements Storer (1976)


Botaurus lentiginosus (Rackett), American Bittern
MATERIAL.-Tibiotarsus, distal end (UF 11821).
REMARKS.-The tibiotarsus of Botaurus differs from that of Nycticorax
by having shaft stouter and less round in cross section; flange overlap on
both internal and external condyles better developed; internal condyle,
when viewed medially, more round; anterior intercondylar fossa nar-
rower; internal condyle projecting approximately even with external edge
of tendinal groove (the condyle is located more externally in that of Nyc-
ticorax); depression just proximal to anterior articulating surface of exter-
nal condyle less excavated; depression on internal condylar half of
anterior intercondylar fossa more excavated; internal ligamental prom-
inence more proximally located, generally.



Ixobrychus sp.
MATERIAL.-Tibiotarsus, distal end (UF 22101).
REMARKS.-The tibiotarsus of Ixobrychus is separated from that of
Butorides by having flange overlap of condyle poorly developed along
outer edges of both condyles; tuberosity on ridge forming internal bound-
ary of tendinal groove located closer to supratendinal bridge; flange form-
ing posterior ridge of internal condyle less extended, resulting in a nar-
rower posterior intercondylar sulcus; ridge separating anterior intercon-
dylar fossa from posterior intercondylar sulcus depressed along external
half (this ridge is straighter and more defined in Butorides).
The fossil was compared with Recent skeletal material from six species
of Ixobrychus: exilis, involucris, minutus, sinensis, sturmii, and cin-
namomeus. It differs from the tibiotarsi of all six species by having the
anterior intercondylar fossa more excavated. The fossil is most similar to
the tibiotarsi of the first four Ixobrychus species listed above, but is fur-
ther separated as follows: In I. exilis the tibiotarsus has less distance be-
tween the condyles; condyles relatively thinner; smaller size (Table 3). In

Table 3.-Measurements (in mm) of the tibiotarsus of fossil Ixobrychus sp. from Coleman
IIA, Recent I. exilis and I. minutus (Brodkorb Collection), and Recent I. in-
volucris, I. sinensis, I. sturmii, and I. cinnamomeus (Smithsonian Institution).

Distal Greatest Depth Greatest Depth
Species Width External Condyle Internal Condyle
I. sp. 5.6 4.7 5.4
I. exilis
mean 5.0 4.5 4.8
range (6) 4.9-5.2 4.2-4.6 4.5-5.0
I. minutus 5.9 5.1 5.3
I. involucris 5.5 4.8 5.1
I. sinensis 5.3 4.5 4.4
I. sturmii 5.9 5.3 5.8
I. cinnamomeus 5.8 5.5 5.8

I. involucris and I. minutus the tibiotarsi have both condyles (anterior
view) wider; internal condyles extended in antero-posterior plane (the in-
ternal condyle of the fossil extends more medially). In I. sinensis the
tibiotarsus has more excavation on external face of external condyle; both
condyles (anterior view) wider; relatively greater distance between con-
This fossil tibiotarsus possibly represents a new species, but more
material is necessary for an accurate diagnosis. The fossil is most similar to
the tibiotarsus of I. exilis in overall characters, and may be simply an
aberrant form referable to exilis (Fig. 1).

Vol. 26, No. 1


i 5mm I
Figure 1. (A) Anterior and (B) distal views of the tibiotarsus of Ixobrychus sp. from Coleman
IIA (UF 22101).

Ciconia maltha weillsi (Sellards)
MATERIAL.-Carpometacarpus, proximal end (UF 11619).
REMARKS.--Ciconia is separated from Mycteria by having metacarpal
I larger and more robust with shorter process; carpal trochlea wider
through lateral extension of external face of trochlea; region just posterior
to external ligamental attachment more depressed; anterior carpal fossa
located more anteriorly and more deeply excavated; ligamental attach-
ment of pisiform process less prominent; flexor attachment longer and
more raised from metacarpal II shaft.


Anas platyrhynchos Linnaeus, Mallard
MATERIAL.-Tibiotarsus, lacking the proximal end (UF 11642).
REMARKS.-Anas rubripes, Anas diazi, and Anas fulvigula are here
considered subspecies of A. platyrhynchos, following Johnsgard's (1961)
suggested classification. Woolfenden (1961) made a thorough study that is
useful in separating the various genera within this family, and Ligon
(1965) separated A. platyrhynchos from other species of Anas by its larger
The slimness of the shaft shows it is that of a female. The fossil record



of this species is well documented from Pleistocene and prehistoric sites
around the world, including about a dozen sites in Florida alone
(Brodkorb 1964).

Branta canadensis cf. B. c. hutchinsii (Richardson), Canada Goose
MATERIAL.-Carpometacarpus, proximal end (UF 22103).
REMARKs.-The specimen was compared to carpometacarpi of Branta,
Anser, Anabernicula, and Brantadorna species, and found to be most
similar to those of Branta and Anser. The fossil agrees with Branta in hav-
ing a large pollical facet; internal edge of pollical facet extends more
media; external edge of pollical facet extends more distad (the car-
pometacarpus of Anser has a shorter and narrower pollical facet, with the
external edge of the facet blunter).
The size of the fossil is identical to that of Hutchins' Canada goose,
which is among the smallest subspecies of B. canadensis. This subspecies
has been recorded from three other Pleistocene sites in Florida: Seminole
Field, Melbourne, and Itchtucknee River (Wetmore 1931).


Coragyps occidentalis (L. Miller)
MATERIAL.-COracOid (UF 11799), 2 humeri (UF 11591T, 22102),
radius (UF 11593), ulna (UF 22096), 2 tarsometatarsi (UF 11800A,
11800B), and hallux (UF 22097).
REMARKS.-Most of the fossils are fragmentary, so only a few
measurements (Tables 4 and 5) could be compared with those published

Table 4.-Tarsometatarsal measurements (in mm) of Coragyps occidentalis from seven
Pleistocene sites. indicates mean measurements of Howard (1968).

Trochlea Trochlea Trochlea
Distal for Digit for Digit for Digit
Locality Width II Width III Width IV Width
Coleman IIA 16.8 4.8 7.2 4.1
mean 16.9 4.8 6.7 4.1
range (8) (16.7-17.3) (4.5-5.2) (6.5-6.9) (3.9-4.3)
Rancho La
Brea 17.5 -- -
McKittrick 18.5* -
Carpinteria 17.65* -- -
Smith Creek 17.8* -
San Josecito 16.8* --

Vol. 26, No. 1


by Howard (1968). Because of the broken nature of the specimens,
measurements other than those used by Howard were also made and com-
pared with measurements of specimens from the Brodkorb Collection of
both C. occidentalis from Reddick, Florida, and Recent Coragyps atratus
(Tables 5 and 6).
There appear to be no diagnostic characteristics, other than size,
(elements of C. occidentalis are larger and/or stouter than those of C.
atratus) and intermembral proportions (the wing bones are longer in C.
occidentalis, but the leg bones are about the same length in both species)
that can be used to separate C. occidentalis from C. atratus. C. occiden-
talis was obviously a larger and heavier bird with longer wings and com-
paratively shorter legs than C. atratus (Howard 1968).
The two species have not been found together in any excavation sites,
and it has been suggested that C. occidentalis gave rise to C. atratus
(Fisher 1944). Most authorities, including Howard (1968) and Brodkorb
(pers. comm.), agree with this hypothesis. It is interesting to note that the
two species overlap temporally, but not spatially, in the late Pleistocene
and Recent. In western North America, C. occidentalis was displaced, at
least in an ecological sense, not by C. atratus, but rather by Cathartes
aura (Howard 1968). This apparently did not occur in Florida, because
C. atratus and C. aura have been recovered together from several sites of
Pleistocene and Recent ages (Brodkorb 1964).

Buteo platypterus (Vieillot), Broad-winged Hawk
MATERIAL--Ulna (UF 11823).
REMARKS.-The ulna of B. platypterus is separated from that of Buteo
lineatus by having internal cotyla rounder; external cotyla smaller; prom-
inence for anterior articular ligament larger and more pronounced;
olecranon shorter and rounder; impression of brachialis anticus deeper
and better developed distally along shaft; external condyle rounder (ex-
ternal view) with its edge forming obtuse angle with shaft (the ulna of B.
lineatus has the external condyle extending more proximally, forming an
acute or right angle with the shaft); ligamental attachment at proximal
end of external condyle much more excavated; carpal tuberosity rising
much less abruptly from shaft and not projecting as far.
Buteo jamaicensis (Gmelin), Red-tailed Hawk
MATERIAL.-Tibiotarsus, lacking proximal end (UF 11621), and tar-
sometatarsus (UF 11622).
REMARKS.-Sexual dimorphism is considerable within this species, so
the elements were compared with bones from both B. j. umbrinus and B.
j. borealis to try to determine sex. The two fossils are definitely not from


Table 5.-Measurements of humerus (in mm) of Coragyps occidentalis from seven Pleistocene sites and Recent C. atratus (Brodkorb Collection).
indicates mean measurements of Howard (1968).

Width at Width of Shaft Height of Brachial
Distal Upper End of above Ectepi- Scar through Midshaft
Species Width Brachial Scar Condylar Process Internal Condyle Width

C. atratus
mean 24.6 13.4 16.9 27.3 10.8
range 23.1-26.3 12.4-15.0 16.3-19.0 25.7-30.3 10.4-11.1
C. occidentalis
UF 11591, Coleman IIA 26.6 14.4 18.7 29.6
UF 22102, Coleman IIA -- -- 11.8
933-1601, Tex. Mem. Mus.,
Friesenhahn Cave, Texas 28.0 14.4 19.0 30.8
PB 9003 (juv.), Reddick, Florida. 25.1 14.3 18.2 28.8
Rancho La Brea, California. 27.4* -- -11.5'
McKittrick, California. -- -- -- 11.0
Conkling Cavern, California. 27.5* -- -- 11.4*
San Josecito Cave, California. 26.4 -- -- 11.5*


Table 6.-Measurements of the ulna, radius, and tarsometatarsus (in mm) comparing
Coragyps atratus (Brodkorb Collection) to C. occidentalis from Coleman IIA.
These unconventional measurements of the tarsometatarsus were made to
facilitate fragmental material.

Ulna Midshaft Radius Midshaft
Species Width Width Depth

C. atratus
mean (11) 8.2 5.1 4.9
range 7.2-8.8 4.4-5.9 4.4-5.4
C occidentalis
(UF 11593) -5.5 5.9
(UF 22096) 8.2

Tarsometatarsus Distal End of Ridge Ex- Proximal End of
Distal tending from Hypotarsus Metatarsal Facet
Species Width Width Depth Width Depth

C. atratus
mean (11) 16.3 7.8 5.5 8.1 5.1
range (14.9-17.7) (7.1-8.3) (5.6-6.0) (7.5-8.8) (4.6-5.5)
C. occidentalis
(UF 11800A) 16.8 -8.8 5.8
(UF 11800B) 8.6 5.8 --

the same individual, because the tibiotarsus corresponds in size to a small
male and the tarsometatarsus is from a much larger bird, most likely a

Buteo lagopus (Pontoppidan), Rough-legged Hawk
MATERIAL.-Humerus, distal end (UF 11823).
REMARKS.-The fossil is badly worn, but agrees with the humerus of B.
lagopus, and differs from that of B. jamaicensis, by having distal portion
of shaft flatter and straighter (ventral view); olecranal fossa more ex-
cavated; internal tricipital groove better excavated; external tricipital
groove wider. This is the first fossil record of B. lagopus from Florida.

Aquila chrysaetos (Linnaeus), Golden Eagle
MATERIAL.-Femur, distal end (UF 22095).
REMARKS.-The femur of A. chrysaetos is distinguished from that of
Haliaeetus leucocephalus by having fibular groove more deeply ex-
cavated; fibular condyle extends more laterally and distad, causing distal
portion of fibular condyle to be more extensive and pointed; external con-
dyle narrower (distal view); depression just anterior to distal-most point
of external condyle tends to be more deeply excavated.



The size of the specimen suggests it is a male. More importantly, this is
the earliest record of the genus from the New World, and the first fossil
record of A. chrysaetos from Florida.


Falco columbarius Linnaeus, American Merlin
MATERIAL.-Humerus, distal end (UF 16704), and ulna, distal half
(UF 22099).
REMARKS.-Wing elements of F. columbarius are very similar to those
of Falco sparverius, but are larger (Tables 7 and 8). Although both species
exhibit sexual dimorphism, Recent specimens show no size overlap in
wing elements. The fossils are slightly smaller than Recent male F. colum-
barius, but are considerably larger than Recent female F. sparverius.
The humerus of F. columbarius differs from that of F. sparverius by
having olecranal fossa less excavated near external condyle; attachment
site on ectepicondylar prominence larger, more dorsal (external) and
proximal in its location on shaft; attachment of pronator brevis larger and
more ventral; internal ridge of external tricipital groove less pronounced.
The fossil humerus has a narrow shaft like that of F. sparverius, but this
character is probably variable in F. columbarius. The ulna agrees with F.
columbarius in having trochlea wider; small groove just proximal and
anterior to internal condyle more apparent.
Table 7.-Measurements of the humerus (in mm) of Recent Falco sparverius sparverius
(Brodkorb Collection), fossil F. columbarius from Coleman IIA, and Recent
F. columbarius (Brodkorb Collection).

Shaft Width just Depth of Depth of
Distal Proximal to Ect- External Internal
Species Width epicondylar Pro. Condyle Condyle

F. sparverius
mean 6.7 4.5 4.2 2.0
range (5) 6.4-7.1 4.2-4.7 4.0-4.5 1.9-2.1
mean 7.5 4.9 4.3 2.2
range (5) 6.9-7.3 4.5-5.3 4.2-4.3 2.1-2.2
mean 6.9 4.7 4.2 2.1
range (10) 6.4-7.3 4.2-5.3 4.0-4.3 1.9-2.2
F. columbarius
Fossil 8.1 5.3 4.8 2.5
Male 8.4 5.4 5.3 2.6
Male 8.6 5.6 5.3 2.7

Vol. 26, No. 1


It is of interest to note that Ligon (1965) demonstrated that F.
sparverius from the Haile XIB Pleistocene was as large or larger than Re-
cent F. sparverius, while the Coleman IIA F. columbarius specimens are,
for the most part, smaller than those of the modern F. columbarius. This
is the first fossil record of F. columbarius from Florida. The species was
previously known from only two Pleistocene sites in North America: Ran-
cho La Brea (L. Miller 1921) and McKittrick (L. Miller 1925), both in



Colinus suilium Brodkorb
MATERIAL.-Cranium (UF 11604), 2 mandibles (UF 11810, 11811), 3
cervical vertebrae (UF 11604), 2 furculae (UF 11605), 7 scapulae (UF
11610), 13 coracoids (UF 11609), 2 sterna (UF 11606), 18 humeri (UF
11611), 16 ulnae (UF 11814), 4 radii (UF 11613), 9 carpometacarpi (UF
11614, 11813), 5 fused thoracic vertebrae (UF 11615, 11815), 2 synsacra
(UF 11608, 11818), 16 femora (UF 11616), 17 tibiotarsi (UF 11617), 9 tar-
sometatarsi (UF 11618), 5 ribs, 2 ulnares, 3 proximal phalanges, hallux,
and 2 pes terminal digits.
REMARKs.-Holman (1961) made a thorough study of the New World
quails, including both fossil and living species. He compared 94

Table 8.-Measurements of the ulna (in mm) of Recent Falco sparverius sparverius
(Brodkorb Collection), fossil F. columbarius from Coleman IIA, and Recent
F. columbarius (Brodkorb Collection).

Shaft Width just Depth of
Distal Proximal to External
Species Width External Condyle Condyle

F. sparverius
mean 4.4 2.9 3.8
range (5) 4.1-4.7 2.8-3.2 3.6-3.8
mean 4.6 3.1 3.9
range (5) 4.4-4.7 2.9-3.3 3.8-4.1
mean 4.5 3.0 3.9
range (10) 4.1-4.7 2.8-3.3 3.6-4.1
F. columbarius
Fossil 5.1 3.4 4.3
Male 5.3 3.4 4.5



osteological characters and 15 intermembral proportions for each of the
living genera and included as many as possible for the fossil genera. He
also listed qualitative characters useful in separating both fossil and living
species of Colinus (C. hibbardi, C. suilium, C. virginianus, C. leucopo-
gon, C. cristatus, and C. nigrogularis) and included measurements of all
species of Colinus except C. cristatus.
Unfortunately all of Holman's specimens of C. suilium were late
Pleistocene in age and revealed no physical changes or trends with respect
to time. An early Pleistocene site, Inglis IA, contains a very large sample
of C. suilium, and along with the specimen from Coleman IIA, has pro-
vided important data helpful in tracing a more complete study of the
species development in Florida throughout the Pleistocene.
In the two populations of C. suilium presented here, the major
variables (other than the ages of the deposits) are minimized. Both
populations are from late in glacial stages, which would tend to decrease
any differences that are due to effects of "Bergmann's principle." The two
localities where the fossils were collected are near each other, separated
by an east-west distance of about 60 kilometers, eliminating any possible
north-south size gradient, such as that found today in C. virginianus (see
Holman 1961). The removal of these two factors would make any dif-
ferences between the two populations attributable to age, i.e. different
stages of evolution.
Species of Colinus cannot be separated by intermembral proportions
(Holman 1961:152, 206). Intermembral proportions calculated for the
two populations of C. suilium showed no substantial differences, except
for a difference of 7% in the tarsometatarsus/humerus ratios (Table 9).
However other leg/wing ratios and wing/leg ratios showed no substantial
proportional differences. Therefore the 7% difference in the tar-
sometatarsus/humerus ratios is considered to be insignificant. No reliable
osteological characters can be used to separate the two populations,
although some slight average differences in intramembral proportions are
As mentioned earlier, a gradual size gradient occurs in C. virginianus
from north to south. In the eastern United States, the largest specimens
(C. v. virginianus) occur in Michigan and the smallest (C. v. floridanus)
occur in southern Florida. Specimens of C. suilium from Arredondo,
Williston, Orange Lake, Reddick, and Haile average as large or larger
than Recent C. v. virginianus from Michigan in most measurements
(Table 10, and Holman 1961:222-223). Specimens of C. suilium from
Coleman IIA and Inglis IA are of interest because they are smaller than
those of the later populations of C. suilium. The Inglis quail is about
equal in size to C. virginianus of northern Florida, while the Coleman
quail is about midway in size between the quail of northern Florida and
C. v. virginianus of Illinois.

Vol. 26, No. 1


Table 9.-Intermembral proportions (in percent) of Colinus suilium from Coleman IIA and
Inglis IA, and overall range for genus Colinus (from Holman 1961:152).

Coleman IIA Inglis IA Overall

wing/leg 65 67 63-69
ulna/humerus 91 88 86-91
tibiotarsus 58 55 57-61
femur 79 77 73-83
humerus/femur 84 88 82-88
tibiotarsus 62 63 62-66
ulna/femur 76 78 72-79
tibiotarsus 56 56 53-59
tarsometatarsus 97 101 92-104
femur 45 46 43-47
tibiotarsus 33 33 33-35
tarsometatarsus 57 60 55-62
humerus 94 87 86-98

Table 10.-A chronological list comparing mean measurements (in mm) of Colinus suilium
from eight Pleistocene sites of central Florida with the number of specimens
(in parenthesis) for each. (Arredondo I and II, Orange Lake, and Reddick
measurements from Holman 1961; Haile XIB measurements from Ligon 1965;
Williston measurements from Holman 1969.)


Locality Length Proximal Width Distal Width

Orange Lake 36.40(1) 9.85(4) 7.58(3)a
Reddick 35.91(31) 9.75(66) 6.98(84)a
Arredondo I 37.75(2) 10.02(6) 7.45(2)a
Arredondo II 10.00(1) 7.10(3)a
Haile XIB -- 9.31(15) 6.52(14)
Williston 35.84(4) 9.83(4) 7.45(4)b
Coleman IIA 34.71(8) 9.35(15) 7.09(14)
Inglis IA 34.93(40) 9.07(70) 6.99(68)

aHolman (1961) published the distal width as "width below epicondyles". His measurements have been altered to con-
form to the greatest distal width published for the other sites, Coleman IIA, and Inglis IA. The alteration involved the addi-
tion of an average value of 0. 35 mm.
bHolman (1959) did not include the mean value for this measurement, but did give the overall range. From this, a me-
dian value was figured and substituted in place of a mean value.



Table 10 continued

Locality Length Proximal width

Orange Lake -6.00(1)
Reddick 32.33(11) 6.11(23)
Arredondo I 33.05(2) 6.30(2)
Arredondo II 33.00(1) 5.90(1)
Haile XIB 32.42(4) --
Williston 34.00(1) 6.60(1)
Coleman IIA 31.56(9) 5.99(11)
Inglis IA 30.89(12) 5.72(24)

Locality Length Proximal height

Orange Lake 19.60(1) 5.90(1)
Reddick C 19.05(10) 5.63(10)
Reddick 18.93(16) 5.75(26)
Arredondo I 19.30(1) 5.50(1)
Arredondo II
Haile XIB 18.99(10) 5.87(15)
Williston 20.00(1) 5.90(1)
Coleman IIA 18.67(5) 5.65(9)
Inglis IA 18.22(20) 5.54(22)

Locality Proximal width

Orange Lake 7.13(3)
Reddick 6.94(7)
Arredondo I
Arredondo II 7.10(1)
Haile XIB --
Coleman IIA 6.87(7)
Inglis IA 6.73(44)

Locality Length Distal width

Orange Lake 29.53(3) 9.00(2)
Reddick 28.99(16) 8.79(7)
Arredondo I 30.82(2) 9.00(2)
Arredondo II 29.80(1) 8.70(1)
Haile XIB
Williston -- --
Coleman IIA 28.08(5) 8.75(6)
Inglis IA 27.21(49) 8.60(18)

Vol. 26, No. 1


Table 10 continued

Locality Length Proximal width Distal width

Orange Lakec 42.40(4) -7.07(7)
Orange Laked -- -6.60(1)
Reddick 42.41(6) -7.00(30)
Arredondo I -- --
Arredondo II 42.50(1) -- 6.90(1)
Haile XIB -- 7.39(8) 7.01(9)
Williston 43.60(1) -7.05(2)
Coleman IIA 41.27(7) 6.98(13) 6.84(11)
Inglis IA 39.61(12) 7.04(21) 6.68(21)

Locality Length Proximal width Distal width

Orange Lake 58.50(3) 9.93(3) 5.65(4)
Reddick A 57.20(1) 9.23(12) 5.58(36)
Arredondo I 60.00(1) 9.65(2) 5.45(2)
Arredondo II --- 5.57(3)
Haile XIB -9.23(6) 5.33(12)
Williston 56.00(1) 8.90(1) 5.50(3)
Coleman IIA 56.29(1) 9.27(8) 5.35(7)
Inglis IA 55.42(2) 8.83(19) 5.21(21)

Locality Length Proximal Width Distal Width

Orange Lake -- 6.15(2) 6.00(3)
Reddick 32.87 5.88(24) 6.03(27)
Arredondo I 34.60(1) 6.20(2) 6.50(2)
Arredondo II -- 6.00(1) -
Haile XIB 33.65(2) 5.80(7) 6.22(12)
Williston -- --
Coleman IIA 32.48(4) 5.73(9) 6.02(7)
Inglis IA 30.46(10) 5.64(9) 5.84(19)
dOrange Lake, Arredondo clay.
Orange Lake, Arredondo-Wicomico contact.

As Holman pointed out, C. hibbardi, from the upper Pliocene of Kan-
sas, is the most distinctive species of the genus because of its large size and
qualitative osteological characters (see Wetmore 1944). C. suilium is less
distinct from C. virginianus than from C. hibbardi and, as Brodkorb
(1960) noted, is probably a temporal equivalent to C. virginianus. The
evolutionary trend of species in the genus Colinus has been thought to be
simply a gradual size decrease through time, but the new material from


Coleman IIA and Inglis IA demonstrates this is not correct. It has been
hypothesized that C. suilium was derived from C. hibbardi, but consider-
ing these new data, it is possible to hypothesize that a different species of
Colinus, much smaller than C. hibbardi, gave rise to C. suilium. The im-
portant point to be made here is that C. suilium apparently underwent a
gradual size increase, rather than a decrease as previously thought,
throughout the Pleistocene of Florida.

Agriocharis anza Howard
MATERIAL.-3 skulls and premaxilla (UF 11588), 58 vertebrae (UF
11597), 18 coracoids (UF 11590), 12 sterna (UF 11589), 37 humeri (UF
11591), 6 scapulae (UF 11595), 23 ulnae (UF 11592), 9 radii (UF 11593),
12 carpometacarpi (UF 11594), 7 synsacra (UF 11598), 27 femora (UF
11599), 33 tibiotarsi and 2 fibulae (UF 11600), 35 tarsometatarsi (UF
11601), 4 manus phalanges, and 16 pes digits.
REMARKS.-A study by Steadman (1975) consisted of detailed
osteological comparisons of both fossil and living turkeys, and included
the specimens of A. anza from Coleman IIA. He made comparisons be-
tween Proagricocharis kimballensis, Agriocharis progenes, A. ocellata, A.
anza. Parapavo californicus, and Meleagris gallopavo. Steadman con-
cluded, on the basis of osteological similarities and because most of the
qualitative osteological differences are only average ones, that the genera
Proagriocharis, Agriocharis, and Parapavo should be synonymized with
the genus Meleagris. Paynter (1955) found Agriocharis not deserving of a
separate genus from Meleagris, and Steadman (1975) stated that there is
little doubt M. gallopavo evolved from A. anza (or as he referred to it,
Meleagris anza). This would certainly indicate that Agriocharis, at least,
should be synonymized with Meleagris, but for the purposes of this paper
the Coleman turkeys are still referred to Agriocharis.

Rallus eleganus Audubon, King Rail
MATERIAL.-Humerus, distal end (UF 16703), ulna, proximal end (UF
16702), femur, distal end (UF 11820).
REMARKs.-Ligon (1965) listed characters by which elements of the
larger rails (Rallus, Fulica, and Gallinula) can be segregated, but he did
not include elements of Porphyrula, which also overlaps these rails in size.
The humerus of Rallus is separated from that of Porphyrula by hav-
ing the distal end narrower; entepicondyle more pointed and located
more distally; intercondylar furrow and furrow between entepicondyle

Vol. 26, No. 1


and internal condyle both wider; internal condyle less constricted ven-
trally; ectepicondyle less pointed.
The ulna of Rallus differs from that of Porphyrula by having depres-
sion just proximal to prominence for anterior articular ligament much
more excavated; olecranon more pointed and extending more proximally.
The femur of Rallus is separated fom that of Porphyrula by having
ridge connecting posterior articular surface of internal condyle to shaft
better developed and connecting to shaft laterally; a deep depression on
medial side of shaft just posterior to anterior ridge extending from inter-
nal condyle (this depression is absent in Porphyrula); rotular groove
wider; ligamental attachment just proximal to fibular groove larger.
The tibiotarsus of Rallus differs from that of Porphyrula by having
ridge connecting anterior face of external condyle to shaft better
developed, forming a more gradual anterior extension of condyle; shaft
thicker where it meets with internal condyle; shaft more rounded in cross
section; external condyle more pointed anteriorly.

Coturnicops noveboracensis (Gmelin), Yellow Rail
MATERIAL.-Tibiotarsus, distal end (UF 16709).
REMARKS.--Feduccia (1968) compared Coturnicops noveboracensis
with C. avita from the Upper Pliocene of Idaho and concluded that C.
avita is probably the ancestor of C. noveboracensis. Laterallus exilis oc-
curred in Florida during the Pleistocene (Olson 1974), and its elements
are similar in both size and characters to those of C. noveboracensis.
The tibiotarsus of C. noveboracensis is separated from that of
Laterallus by having intercondylar fossa more narrow; ridges extending
posteriorly from both condyles protruding farther from shaft, making
posterior intercondylar sulcus more excavated; external and internal
ligamental prominences both better developed; bridge over peroneus pro-
fundus groove located more proximally; external condyle more rounded
(lateral view); a distinct furrow between medial connection of supraten-
dinal bridge and medial ridge of tendinal groove (in that of Laterallus the
supratendinal bridge connects directly to the medial ridge of the tendinal
groove and forms no furrow).
C. noveboracensis previously was known as a fossil from the Reddick
(Brodkorb 1957) and Haile (Ligon 1965) sites of Florida.

Philohela minor (Gmelin), Woodcock
MATERIAL.-Mandible (UF 11796), humerus (UF 11798), 2 coracoids
(UF 11797), and tibiotarsus, distal end (UF 16701).


REMARKS.-Ligon (1965) listed osteological characters that separate
four genera, all of similar size, of the family Scolopacidae: Philohela,
Capella, Limnodromus, and Totanus. This species was previously known
from only three Pleistocene sites, including Haile and Eichelberger Cave
(Brodkorb 1957) of Florida.

Zenaida macroura (Linnaeus), Mourning Dove
MATERIAL.-Humeri, proximal half and distal end, probably from a
single bone (UF 16706), and tibiotarsus, proximal end (UF 22104).
REMARKS.-A fossil dove from the Upper Pliocene of Kansas, originally
identified as Z. macroura by Wetmore (1944), was later described as a
new species, Zenaida prior, by Brodkorb (1969). Brodkorb (1969)
separated the two species and stated that Z. prior is allied and apparently
ancestral to Z. macroura.

Bubo virginianus (Gmelin), Great Horned Owl
MATERAL.-Tibiotarsus, external condyle (UF 22098).
REMARKS.-The fossil is separated from the tibiotarsus of Strix by hav-
ing a well-defined depression just proximal to external condyle. The
tibiotarsi of both Bubo and Nyctea have a ridge between the condyles
separating the intercondylar fossa into two depressions, with one of them
located just proximal to the external condyle (the fossil shows evidence of
this). The fossil is separated from the tibiotarsus of Nyctea by having
posterior ridge of external condyle wider; a convex medial side and a con-
cave lateral side on external condyle (the external condyle of Nyctea has
two flat sides converging at the ridge); external condyle less elongated in
antero-posterior plane; more flange overlap on lateral face of external
The fossil is more robust than the tibiotarsus of Recent B. virginianus
(Table 11). Husband (1924) demonstrated that the supposed large extinct
Bubo sinclairi of late Pleistocene actually falls well within the size range
of B. virginianus of Rancho La Brea, and Howard (1947) suggested B.
sinclairi may simply be a geographical variant of B. virginianus from
Rancho La Brea. B. sinclairi is probably not a viable taxon, so the Cole-
man specimen is referred to B. virginianus.
This fossil is particularly important because it represents (1) the first
fossil record for B. virginianus in Florida, and (2) the earliest fossil record

Vol. 26, No. 1


for the genus in the New World. The earliest known species of Bubo is
Bubo incertus from the Upper Eocene or Lower Oligocene of France
(Brodkorb 1971).

Otus asio (Linnaeus), Screech Owl
MATERIAL.-Synsacrum (UF 11608), and tarsometatarsus (UF 11623).
REMARKS.-O. asio is similar to Aegolius funereus in all elements except
for the tarsometatarsus. The tarsometatarsus of Otus differs from that of
Aegolius by having a relative length 1/3 longer; shaft much narrower.
The neural spines of the fused sacral vertebrae are longer in Otus, giving
the anterior dorsal surface a higher, arched appearance. The anterior
neural spines are broken away in the fossil, but remnants of the iliac
fossae indicate it had the same condition as in Otus.
0. asio has been recorded extensively from the North American
Pleistocene (Brodkorb 1971).



Colaptes auratus (Linnaeus), Common Flicker
MATERIAL--Humerus (UF 16708), and carpometacarpus (UF 16707).
REMARKS.-All North American flickers are now considered conspecific
(Short 1965, Brodkorb 1971). The humerus of Colaptes is separated from
that of Melanerpes by having pneumatic fossa deeper, with fewer and

Table 11.-Measurements (in mm) of the tibiotarsus of Recent Bubo virginianus (Brodkorb
Collection, fossil Bubo virginianus from Coleman IIA (UF 22098), and Recent
Nyctea scandiaca (Brodkorb Collection).

Depth of Depth of Shaft
Distal Width of External just above
Species External Condyle Condyle External Condyle

B. virginianus
mean 5.4 11.8 5.0
range (7) 5.0-5.7 11.2-12.3 4.5-5.3
mean 6.0 13.1 5.5
range (6) 5.9-6.2 12.6-13.9 5.4-5.7
(UF 22098) 6.6 13.8 6.0
N. scandiaca
mean 5.65 12.3 4.85
range (2) 5.6,5.7 12.0,12.6 4.7,5.0
Female (1) 5.7 13.6 5.5



smaller foramina; region about attachment of infraspinatus thicker;
deltoid crest larger; impression of brachial anticus larger; entepicondyle
more extensive. The carpometacarpus of Colaptes differs from that of
Melanerpes by having ligamental attachment of pisiform process larger
and more extensive; intermetacarpal tuberosity larger and more distad;
process of metacarpal I longer.

Hamon (1964) made a detailed study of osteological characters for
those genera of each passerine family recovered from the Pleistocene
deposits of eastern North America. The elements he studied are the
humerus, ulna, carpometacarpus, coracoid, femur, tibiotarsus, and tar-
sometatarsus. Hamon's work proved of great value during the study of
passerine birds from Coleman IIA.

Progne subis (Linnaeus), Purple Martin
MATERIAL--Humerus (UF 11625).
REMARKS.-The large size of the fossil (length: 2.2 cm) separates it
from the humeri of all other North American swallows. P. subis was
previously reported from the Pleistocene only from Reddick (Brodkorb

Protocitta cf. P. ajax Brodkorb
MATEIAL.-Humerus, distal end (UF 16705).
REMARKS.-The specimen is most similar to the humeri of two extinct
species of jays, Protocitta dixi and Henocitta brodkorbi, both known only
from Pleistocene deposits of Florida. The Coleman IIA humerus differs
from that of H. brodkorbi by having the internal condyle more spherical
and the entepicondyle located less aconally. The Coleman IIA humerus
differs from those of both P. dixi and H. brodkorbi by having ectepicon-
dylar prominence more extensive and nearly perpendicular to shaft; ex-
ternal condyle wider; internal condyle smaller; external tricipital groove
narrower; attachment of anterior articular ligament small; medial spur of
ectepicondylar process longer; entepicondyle smaller and thinner;
olecranal fossa less excavated into entepicondyle.
The projection of the ectepicondylar process and the well developed
medial spur are similar to those found in Pica. The Coleman IIA humerus
differs from that of Pica by having olecranal fossa less excavated; both
tricipital grooves better developed; entepicondyle (ventral view) shorter
and wider; attachment of anterior articular ligament smaller. Although

Vol. 26, No. 1


very little of the shaft remains on the fossil specimen, it was apparently
straighter (as in other jays) than that of the humerus of Pica.
Brodkorb (1972) described Protocitta ajax from a single tarsometatar-
sus and stated that it is stouter than that of P. dixi. Unfortunately no
humeri were recovered. It is quite possible that the Coleman IIA Proto-
citta is referable to P. ajax. At this time it is inappropriate to describe a
new species on the basis of this single element (Fig. 2).
Measurements of the specimen are: distal width 10.8 mm; depth exter-
nal condyle 4.6 mm; length external condyle 4.1 mm; and length internal
condyle 3.0 mm.
P. ajax has been reported from the late Pliocene of both Texas (Palo
Duro Falls) and Kansas (Rexroad Ranch) (Brodkorb 1972).


Figure 2. (A) Anconal and (B) palmar views of the humerus of Protocitta ef. P. ajax from
Coleman IIA (UF 16705).

Dendroica sp., Wood Warbler
MATERIAL.-Tibiotarsus, distal end (UF 22107).
REMARKS.-The tibiotarsus of Dendroica is very similar to that of
Geothlypis, but differs by having supratendinal bridge wider; external
condyle (external view) less round; protuberance located at external foot
of bridge less developed; external ligamental prominence less developed.
Similarity among the members of this genus prevents specific identifica-
tion. This is the first fossil record for Dendroica in Florida.
Agelaius phoeniceus (Linnaeus), Red-winged Blackbird
MATERIAL.-Humerus (UF 11808), and tarsometatarsus (UF 11812).
REMARKS.-Elements of Agelaius phoeniceus are similar to those of
Euphagus carolinus and E. cyanocephalus. The humerus of A.


phoeniceus is separated from that of E. carolinus by having internal
tricipital groove less excavated; brachial depression less excavated; no
distinct ridge connecting internal condyle with attachment of anterior ar-
ticular ligament: attachment of infraspinatus smaller; region of external
condyle and ectepicondylar prominence (distal view) wider; depression
on connection between attachment of infraspinatus and internal tubero-
sity more developed. The humerus of A. phoeniceus differs from that of
E. cyanocephalus by having brachial depression less excavated and
shorter; external ridge of brachial depression (ventral view) not straight,
but bending where it meets with shaft; depression extending from exter-
nal tuberosity onto head less developed.
The tarsometatarsus of A. phoeniceus is separated from those of both
E. carolinus and E. cyanocephalus by having shaft more robust; metatar-
sal facet better excavated; trochlear region (internal view) bending more
A. phoeniceus in known extensively from the late Pleistocene. It has
been reported from Ontario, California, possibly from New Mexico and
Virginia, as well as Seminole Field, Itchtucknee River, Reddick, and Ar-
redondo in Florida (Brodkorb 1978).

Pandanaris floridana Brodkorb
MATERIAL.--Femur, proximal end (UF 22108).
REMARKs.-The femur of P. floridana is separated from that of
Quiscalus mexicanus by having a depression in distal surface of neck
underneath head; depression on anterior face just medial to trochanteric
ridge less excavated (this depression extends proximally underneath the il-
iac facet of the femur of Q. mexicanus); neck generally shorter.
The fossil was compared with a large number of specimens of P.
floridana from Reddick, now in the Brodkorb Collection. Apparently, P.
floridana had a considerable sexual size dimorphism, like that found in Q.
mexicanus. The size of the fossil proximall depth 4.1 mm and proximal
width of shaft 3.1 mm) is rather small compared to the Reddick
specimens, so it probably represents a female. Brodkorb (1957), in his
description of the species, stated that its closest living relatives are prob-
ably the two cowbird genera, Molothrus and Tanganavius, but neither of
these genera exhibit the size variation found in both P. floridana and Q.
mexicanus. P. floridana was previously known only from Reddick and
Haile XIB (Ligon 1965).

Pheucticus ludovicianus (Linnaeus), Rose-breasted Grosbeak
MATERIAL.-Ulna (UF 11806).
REMARKS.-The ulna of P. ludovicianus is separable from that of

Vol. 26, No. 1


Pheucticus melanocephalus by having depression between tricipital at-
tachment and olecranon more excavated, with excavation extending into
base of olecranon at region of humeroulnar depression narrower and
ridged; ridge located ventrally to impression of brachialis anticus gener-
ally straight, extending from prominence for anterior articular ligament
along shaft (this ridge is more curved in the ulna of P. melanocephalus);
distal radial depression more smoothly excavated (it is pit-like in the ulna
of P. melanocephalus, with a sharp edge along the distal border).
The ulna of P. ludovicianus differs from the ulna of Cardinalis car-
dinalis by having entire distal end, including distal end of shaft, com-
paratively wider; depression between tricipital attachment and olecranon
much less excavated, giving base of olecranon a thicker appearance.
The ulna of P. ludovicianus is separated from that of Cardinalis
sinuata by having external cotyla much larger; carpal tuberosity smaller.
This is the first fossil record for P. ludovicianus.
Pipilo erythrophthalmus (Linnaeus), Rufous-sided Towhee
MATERIAL.-2 ulnae, distal ends (UF 22100A and B), and tibiotarsus,
distal end (UF 22100C).
REMARKs.-Hamon (1964) compared elements from Pipilo with those
of Passerculus, Ammodramus, Passerherbulus, and Spizella. For each
genus he described 13 given characters for the ulnae and 18 given
characters for the tibiotarsi. P. erythrophthalmus has been previously
reported from late Pleistocene sites in California and New Mexico, as well
as Reddick, Arredondo, and Haile XIB (Brodkorb 1978).

Passerculus sandwichensis (Gmelin), Savannah Sparrow
MATERIAL-Ulna, proximal end (UF 11809A), and femur (UF
REMARKS.-Hamon (1964) compared 13 given characters for the ulnae
and 15 given characters for the femora of Passerculus, Ammodramus,
Passerherbulus, and Spizella. The ulna of P. sandwichensis is separated
from that of Melospiza melodia by having tricipital attachment closer to
proximal edge of external cotyla; prominence for anterior articular liga-
ment more extensive; external cotyla more extensive at region of proximal
radial depression, giving external cotylar edge a more concave shape.
The femur of P. sandwichensis differs from that of M. melodia by hav-
ing attachment of adductor profundus less developed; attachment of
gastrocnemius (external head) smaller and less extensive; internal condyle
narrower (less extensive from shaft), making distal end relatively nar-
rower; shaft slenderer; attachment of ischio-femoralis smaller; iliac facet
narrower anteroposteriorly; trochanter less ridged, giving the proximal
end region more rounded appearance; depression located on shaft at base
of neck smaller.



P. sandwichensis has been previously reported as a Pleistocene species
only from Reddick (Brodkorb 1978).

Pooecetes gramineus (Gmelin), Vesper Sparrow
MATERIAL.-Humerus (UF 11804A).
REMARKS.-The humerus of P. gramineus is separated from the
humerus of Chondestes grammacus by having no well-developed pit be-
tween ectepicondylar prominence and shaft; longer and narrower
brachial depression; olecranal fossa more excavated, extending entire
length of internal condyle (the olecranal fossa only extends from intercon-
dylar region to midway along internal condyle in the humerus of C.
grammacus); proximal edge of pneumatic fossa I circular; deltoid crest
closer to shaft; bone connection between head and internal tuberosity
thicker, making capital groove longer.
The humerus of P. gramineus differs from that of Passerella iliaca by
having distal end relatively narrower; ligamental attachment site just
proximad to external condyle less developed; external condyle (external
view) less round. The humeri of P. iliaca and C. grammacus are similar in
that they are both separated from the humerus of P. gramineus by having
medial spurs better developed; pneumatic fossae I and II larger; distal
edges of articulating surfaces of heads better developed.
Dawson (1948) identified this species from Rancho La Brea; this is its
first fossil record for Florida.

Aimophila aestivalis (Lichtenstein), Bachman's Sparrow
MATERIAL.-Humeri, proximal 3/4 and distal end (probably parts of
the same bone) (UF 11803), and 2 carpometacarpi (UF 11801).
REMARKS.-The humerus of A. aestivalis differs from that of Melo
spiza melodia by having less distance between shaft and ectepicondylar
prominence; entepicondyle larger; ectepicondylar prominence smaller;
brachial depression shallower; articulating surface of head larger, causing
opening of fossa I to be more constricted; internal tuberosity less exten-
sive. The humerus of A. aestivalis is separable from that of Junco by hav-
ing entire element less robust; proximal and distal ends narrower; shaft
lighter; ectepicondylar prominence smaller; brachial depression
shallower. The humerus of A. aestivalis differs from that of Aimophila
cassinii by having both pneumatic fossae smaller; brachial depression less
excavated; external condyle smaller; capital groove longer; entepicondyle
shorter; less distance between medial spur and shaft. The humerus of A.
aestivalis is separable from that of Aimophila ruficeps by having entire
proximal end less robust; attachment of infraspinatus less developed;
brachial depression less excavated; external condyle smaller; entepicon-
dyle shorter. The humerus of A. aestivalis differs from that of Ammospiza

Vol. 26, No. 1


maritima by having pneumatic fossae I and II larger and deeper; head
larger; entepicondyle smaller; brachial depression larger and deeper.
The carpometacarpus of A. aestivalis is separable from that of M.
melodia by having intermetacarpal tuberosity much larger; depression at
proximal end of shaft of metacarpal II (internal view) more excavated;
depression at proximal end of metacarpal III connection with internal
condyle more excavated. The carpometacarpus of A. aestivalis differs
from that of Junco by having intermetacarpal tuberosity larger; entire
element more robust; facet for digit III shorter; flange midway along ten-
dinal groove better developed. The carpometacarpus of A. aestivalis is
separable from that of A. cassinii by having proximal end less robust; ex-
ternal carpal trochlea less extensive; external carpal trochlea (external
view) rounder. The carpometacarpus of A. aestivalis differs from that of
A. ruficens by having entire element comparatively more robust;
metacarpal III more parallel with metacarpal II; facet for digit III
shorter. The carpometacarpus of A. aestivalis is separable from that of A.
maritima by having intermetacarpal tuberosity larger; facet for digit III
more robust; depression between carpal trochlea and metacarpal II (ex-
ternal view) more excavated.
This is the first fossil record for A. aestivalis.

Spizella pusilla (Wilson), Field Sparrow
MATERIAL-Humerus (UF 22109).
REMARKs.-Hamon (1964) compared the humeri of Spizella, Passer-
culus, Ammodramus, and Passerherbulus. The humerus of S. pusilla dif-
fers from that of Spizella passerina by having region between attachment
of infraspinatus and internal tuberosity thicker, making pneumatic fossa
II smaller; head comparatively smaller; brachial depression shorter. The
humerus of S. pusilla is separable from that of Spizella arborea by having
entire element less robust; brachial depression smaller and shallower;
pneumatic fossae I and II smaller; ectepicondylar prominence less
developed. The humerus of S. pusilla differs from that of Spizella pallida
by having region between attachment of infraspinatus and internal
tuberosity thicker; ectepicondyle smaller; brachial depression shallower.
S. pusilla has been recorded as a fossil only from the Pleistocene of Ar-
redondo (Brodkorb 1978).

Spizella sp.
MATERIAL.-Carpometacarpus, lacking metacarpal III (UF 11805).
REMARKS.-The broken nature of the specimen and the great amount
of similarity within the various species of Spizella make specific iden-
tification impossible.



Genus and species indeterminate
MATERIAL-Ulna, proximal half (UF 11804B), and ulna, proximal and
distal halves to same bone (UF 11807).
REMARKS.-The specimens are from two different species of sparrows,
but because they are both broken more accurate identification is impossi-

Anhinga cf. A. grandis Martin and Mengel
MATERIAL-Pit C; ulna, distal half (UF 16664).
REMARKS.-The ulna of Anhinga is separable from that of
Phalacrocorax by having trochlea comparatively longer and narrower;
distal articulating surfaces less raised from shaft; radial depression smaller
and shallower; internal condylar process and carpal tuberosity less exten-
sive; external condylar ridge less round; carpal tuberosity extending
nearly parallel to the shaft (in the ulna of Phalacrocorax the carpal
tuberosity extends almost perpendicular to the shaft); tuberosity at prox-
imal end of internal condyle (internal face) less developed; shaft
straighter. In Phalacrocorax, the anconal and inner papilae at each
secondary feather region of the ulna are separated by a distance nearly
twice that found in Anhinga, which causes the inner papillae to be
located on the flat internal surface of the shaft.
The specimen differs from the ulna of A. anhinga by size (Table 12)
and by having tuberosity on external face of external condyle extending
more proximad, making most distal papilla closer to tendinal pit; ridge of
external condyle attachment to shaft in shape of smooth curve, giving ex-
ternal condyle a more elongated shape (in the ulna of A. anhinga the ex-
ternal condylar ridge attaches to the shaft at an angle approaching 900);
articulation surface of external condyle extends more proximad, to most
proximal point of ligamental attachment; projection of external condylar
ridge (internal face) more anterior and less curved; internal condylar
process less developed; carpal tuberosity longer with more anterior pro-
jection, forming an acute angle between internal condylar process and
carpal tuberosity (this angle is obtuse in the ulna of A. anhinga); ligamen-
tal attachment (ventral surface) better developed and more excavated;
tuberosity at proximal end (internal face) of internal condyle less
developed; distal radial depression better excavated, and its extension
more external (Fig. 3).
Anhinga grandis, from the late Pliocene of Nebraska, is about the
same size as the Anhinga from Coleman, but is known only by the distal

Vol. 26, No. 1


Table 12.-Measurements of the ulna (in mm) of Recent Anhinga anhinga (Brodkorb Collec-
tion) and fossil Anhinga cf. A. grandis (UF 16664) from Coleman IIA.

Height of Height of Width
Greatest External Internal through
Species Distal Width Condyle Condyle Trochlea

A. anhinga
mean 10.6 7.8 5.5 6.3
range (11) 10.0-11.3 7.6-8.1 5.2-5.8 5.7-6.6
UF 16664 11.8 9.1 6.2 7.6

Length of Carpal Central
Length of Tuberosity Ridge from Height of
Species Tendinal Pit Intermuscular Line Shaft

A. anhinga
mean 4.8 6.3 5.1
range (11) 4.3-5.1 5.8-6.5 4.7-5.6
UF 16664 5.9 8.5 5.4

Length from Distal Point
of Internal Condyle to Central
Distal Part of Carpal Width of
Species Tuberosity Shaft

A. anhinga
mean 3.9 6.2
range (11) 3.5-4.2 5.7-6.6
UF 16664 3.6 6.7

Length from Proximal Edge
of Ligamental Attachment
Species to Center of Trochlea

A. anhinga
mean 9.7
range (11) 8.6-10.5
UF 16664 11.1

portion of a single humerus (Martin and Mengel 1975). The distal width
of the humerus of A. grandis is 1.2 times as large as that of A. anhinga.
This compares favorably to some of the proportions between the Anhinga
from Coleman and Recent A. anhinga (Table 13). Rather than describe
this fossil as being that of a new species, it is here referred to A. grandis.



Figure 3. Palmar view of the ulna of Anhinga cf. A. grandis from Coleman IIIC (UF 16664).

Table 13.-A comparison of the ratio Coleman IIA Anhinga cf. A. grandis/A. anhinga for
the ulnar measurements (from Table 11), showing the inconsistency of the
various ratios.



Distal Width 1.07 1.04-1.18
Height of External Condyle 1.04 1.11-1.20
Height of Internal Condyle 1.13 1.07-1.19
Width of Trochlea 1.21 1.13-1.29
Length of Tendinal Pit 1.23 1.16-1.37
Length of Carpal Tuberosity 1.36 1.31-1.46
Length from Internal Condyle
to Carpal Tuberosity 0.92 0.86-1.03
Length from Ligamental
Attachment to Trochlea 1.14 1.06-1.29
Width of Shaft 1.08 1.02-1.17
Height of Shaft 1.06 0.96-1.15



Ardea herodias Linnaeus, Great Blue Heron
MATERIAL-Pit C; tibiotarsus, distal end (UF 15160).
REMARKS.-Ardea herodias (including what was formerly Ardea oc-
cidentalis) is separable from all other North American herons by the much
larger size of its elements. A. herodias has previously been reported from
Pleistocene sites in California and Oregon,and from Seminole Field, Itch-
tucknee River, Melbourne, Bradenton, and Rock Spring in Florida, as
well as numerous prehistoric sites (Brodkorb 1963).


Vol. 26, No. 1


Aythya affinis (Eyton), Lesser Scaup
MATERIAL.-Pit C; humerus (UF 15160).
REMARKS.-The humerus of A. affinis differs from that of Aythya col-
laris by having pectoral attachment larger with more distal extension; in-
ternal tuberosity more robust; bicipital crest less curved; angle at which
bicipital crest connects to shaft generally obtuse (in A. collaris the angle of
this connection is approximately 900); attachment of anterior articular
ligament larger.
A. affinis is usually smaller than A. collaris (Johnsgard 1975), but it
tends to have longer wing elements. A. affinis has been reported exten-
sively from Pleistocene sites all over North America (Brodkorb 1964).

Aythya sp.
MATERIAL.-Pit C; humerus, shaft and partial distal end (UF 22110),
carpometacarpus, distal end (UF 22112), and tibiotarsus, distal half (UF
REMARKS.-Ducks show a tremendous amount of intraspecific varia-
tion, and many characters are not constant enough to separate closely
related species with certainty. The slender shafts of the humerus and
tibiotarsus separate these specimens from other genera of ducks.
Woolfenden (1961) listed characters by which different genera of ducks
are separated.

Agriocharis anza Howard
MATERIAL.-Pit D; femur, proximal half (UF 15151).
REMARKS.-A. anza is the only avian species recovered from both
Coleman II and Coleman III. The fossil is identical to the femora of A.
anza identified from Coleman IIA. A. anza has been reported from the
middle Pleistocene of California and Texas (Brodkorb 1964). It is also
known from Inglis IA, but until now that recovery was unpublished.

Gallinula chloropus (Linnaeus), Common Gallinule
MATERIAL.-Pit C; carpometacarpus, lacking metacarpal I and
metacarpal III (UF 22113).



REMARKS.-Olson (1974) synonymized Gallinula brodkorbi with G.
chloropus, but noted that G. chloropus in the Pleistocene was larger and
heavier than the modern form. This specimen, 35.0 mm in length, falls
within the measurements for both Recent and fossil specimens of G.
chloropus published by Olson (1974).
G. chloropus has been recorded extensively from Pleistocene deposits
in Europe, the Middle East, Brazil, Cuba, Idaho, and possibly Arizona,
as well as six sites in Florida (Brodkorb 1967).


Martin (1974) stated that Coleman IIA was probably more open and
xeric than it is today, and predominantly a savanna. He cited several
species of mammals that would indicate this, such as Peromyscus
floridanus (sandpine scrub/turkey oak sandhill), Cryptotis parva (pine
flatwoods, grass and sedge fields, and Lepus alleni and Platygonus sp.
(open and semi-arid = xeric habitats). Many of the birds, especially the
sparrows Passerculus sandwichensis, Pooecetes gramineus, Aimophila
aestivalis, Spizella pusilla, and Spizella sp., are savanna dwellers, thus
bolstering Martin's conclusion. The two most common bird species, Col-
inus suilium and Agriocharis anza, if one assumes they preferred the
habitats of their living counterparts, give no reason to dispute this. Col-
inus virginianus (C. suilium's modern counterpart) inhabits open
pinelands and fields, and Meleagris gallopavo (A. anza's modern counter-
part) can be found in virtually any type of habitat in Florida (swamps,
open woodlands, particularly oak and hickory forest, and clearings).
Also, most of the birds of prey (Buteo jamaicensis, Buteo lagopus, Aquila
chrysaetos, probably Coragyps occidentalis, and Falco columbarius)
prefer open country for hunting.
The presence of Podilymbus podiceps, Botaurus lentiginosus, Ix-
obrychus sp., Ciconia maltha, Anas platyrhynchos, Branta canadensis,
Rallus elegans, Coturnicops noveboracensis, and Agelaius phoeniceus
suggests a freshwater marsh or pond community must have been nearby.
Martin reported finding remains of Sciurus carolinensis and Ochrotomys
nuttalli, which indicates that a more mesic habitat was also sampled.
Several avian species, most notably Buteo platypterus, Philohela minor,
and Pheucticus ludovicianus, support this conclusion.
Disregarding extinct species, the avifauna from the Pleistocene of
Coleman IIA is very similar to that of central Florida today. Two birds of
prey, Buteo lagopus and Aquila chrysaetos, do not normally range as far
south as central Florida today, but their overall ranges, especially in
eastern North America, are not very different from those of several other

Vol. 26, No. 1


species of birds recovered from Coleman IIA. Their presence is not suffi-
cient to base any significant climatic differences on. One extinct species,
Agriocharis anza, has previously been reported only from the Pleistocene
of western America. However, this species has a modern counterpart that
ranges all across North America, including central Florida, so its presence
does not necessarily indicate any affinity to environmental conditions of
western America.
The paleoecology of Coleman III is more obvious. It was apparently a
small, sluggish spring run. All of the avian fossils are of aquatic species,
with the exception of a single turkey bone from IIID. Most of the non-
avian species (see Introduction for list of non-avian species) are also
aquatic, but the presence of Crotalus adamanteus, Hemiauchenia
macrocephala, and Equus sp. suggest that a drier, more open savanna
habitat was nearby.
Both Coleman IIA and Coleman IIIC and D contained species that in-
dicate dry, open savanna habitats were nearby, and both contained
numerous aquatic species. The two sites were probably contemporaneous,
judging from the extinct species of mammals, Pampatherium septen-
trionalis, Equus sp., and possibly Sigmodon sp., and the one extinct
species of bird, Agriocharis anza, common to both. If so, the spring of
Coleman III was probably the habitat from which many of the aquatic
species of birds of Coleman II came. A relatively complete picture of the
middle Pleistocene habitat of Coleman can be reconstructed. The area
was probably predominantly a savanna or open pineland habitat in
which a freshwater spring gave rise to a small sluggish stream that prob-
ably had a marsh and river swamp habitat along its run. Farther from the
stream was mesic forestland that in turn gave way to the open pineland or
savanna. A short distance from the spring, 300 meters south, a sinkhole
opened into the underlying limestone of the Ocala group. Whether it
formed in the mesic forest or open land is impossible to determine.
If this reconstruction is accurate, the Coleman area during the middle
Pleistocene was very similar to many of the freshwater spring habitats of
central Florida today.


It is well documented that Pleistocene representatives of modern
species are often characterized by larger size. Gallinula chloropus of Cole-
man III is large compared to its modern representative, and Podilymbus
podiceps and Bubo virginianus of Coleman II are larger than their
modern representatives in Florida. Several extinct species, Coragyps oc-
cidentalis and Colinus suilium of Coleman II, Agriocharis anza of Cole-



man II and III, and Anhinga cf. A. grandis of Coleman III, are large
compared to their modern counterparts. Ixobrychus sp. of Coleman II is
larger than a modern species that is closely related or conspecific. Olson
(1974) gave other examples of this phenomenon. The one exception to the
trend is Falco columbarius of Coleman II, which is smaller than its
modern representative of Florida.
Two species, Protocitta cf. P. ajax from Coleman II and Anhinga cf.
A. grandis from Coleman III, show affinities to the late Pliocene, but
neither identification is definite because of a lack of referred material.
Analysis of two large samples of Colinus suilium, from Coleman IIA
and from the earlier deposit of Inglis IA, has demonstrated that the
species underwent an overall increase in size throughout the Pleistocene in
Florida. Until now, it was hypothesized that C. suilium was derived from
Colinus hibbardi, a large species from the late Pliocene of Kansas, and
gradually decreased in size until the late Pleistocene. C. suilium was prob-
ably not derived from C. hibbardi, but from some as yet unknown
species, and then increased in size until the late Pleistocene, at which time
it most likely gave rise to Colinus virginianus and underwent a size


The mode of fossil deposition for Coleman IIA is unclear. It is possible
that roosting or nesting birds of prey were a major source of small animals
in this fauna. A significant portion (24%) of the avifauna in this deposit is
comprised of birds of prey. Buteo platypterus and Falco columbarius prey
almost exclusively on small birds, Bubo preys commonly on birds of all
sizes, and Aquila chrysaetos preys heavily on large birds. Coragyps oc-
cidentalis may have scavenged bird remains.
It is improbable that birds would have fallen into a large sinkhole and
not have been able to escape, unlike many of the large mammals. Food re-
mains from predators seem to be a plausible explanation for the concen-
tration of avian fossils recovered from Coleman IIA. Bubo virginianus
was perhaps most responsible for the extremely varied sample of birds,
especially the larger species, because owls commonly roost in caves in
Florida and can create tremendous stockpiles of animal bones from their
regurgitated pellets. C. occidentalis probably nested on the side of the
sink, as some of the recovered fossils of this species are from immature,
probably nestling birds, and may also have been a source of animal re-
mains. It should be noted that the two most common species from this avi-
fauna, Agriocharis anza and Colinus suilium, are the Pleistocene
representatives of two modern species, Meleagris gallopavo and Colinus

Vol. 26, No. 1


virginianus, which are known to be prey species of Bubo virginianus (Bent
No evidence indicates the water birds of Coleman IIA were deposited
separately during a time when the sinkhole contained water, because
their fossilized bones were recovered in both sand and clay facies along
with the remains of other birds and mammals and vary considerably in
color. Martin (1974:36) stated the sinkhole was "quite extensive and not
filled to the top with water during most of its depositional history" as
evidenced by the presence of a relatively large quantity of bat remains.
The fossils of Coleman III were not in a single concentration, but
spread throughout the area of excavation. Apparently deposition oc-
curred quite randomly along the course of the postulated spring run.


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