FLORIDA STATE MUSEUM
THE PLEISTOCENE AVIFAUNA
OF ARREDONDO, FLORIDA
UNIVERSITY OF FLORIDA
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Published 22 May 1959
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HE PLEISTOCENE AVIFAUNA OF ARREDONDO, FLORIDA
PIERCE BRODKORB 1
SYNOPSIS: An extensive fauna of Pleistocene vertebrates occurs at Arredondo,
Florida. The bone bed lies in a fresh water clay which is here named the
Arredondo clay member of the Wicomico formation. The geographic extent of
this member includes 10 localities in 3 counties of northern Florida. The sedi-
ments were deposited below the present 100-foot contour under cooler climatic
conditions during the Illinoian glacial stage. The environment included a fresh
water marsh community with nearby scrub. The avifauna of 43 species includes
forms still living, interglacial relicts, and glacial indicators. Five living species
of birds are added to the fossil record, and six others to the Pleistocene fauna of
Florida. Four species are described as new: Falco readei, Colinus suilium, Dory-
paltus prosphatus (n. gen., Charadriidae), and Cremaster tytthus (n. gen., Icteridae).
The name Podilymbus magnus Shufeldt is revived. Limicolavis pluvianella Shu-
feldt, from the Oligocene of Oregon, is removed from the Charadriidae; it must
be left in incertae sedis until the type is restudied.
Arredondo is a settlement about 6 miles southwest of the center
of Gainesville, Alachua County, Florida. An extensive fauna of
Pleistocene vertebrates occurs here in overburden of two mines of
the Levy County Lime Rock Corporation.
Certain of the amphibians and reptiles from the deposit have been
referred to in papers by Goin and Auffenberg (1955) and Auffenberg
(1956, 1958). Much of the mammalian fauna has been reported by
Bader (1957) and Olsen (1958). Fish remains are fairly numerous,
but these have not yet been studied. Fresh water and terrestrial
snails of the genera Polygyra and Physa occur locally.
In the 1955 paper cited the name Kanapaha was used for the
locality, but later papers have all employed the term Arredondo,
Which is the closer of the two settlements.
The two fossil localities lie just northeast of Arredondo (see map),
Sin the NW V4 sec. 22, T 10 S, R 19 E, Alachua County, Florida. Pit
1 is on property owned by W. H. Damron, and is just north of the
, right-of-way of the Seaboard Air Line Railway, 0.2 mile southwest
-\of railroad milepost 709. Pit 2 lies directly across the railroad from
S The author is Professor of Biological Sciences at the University of Florida.
\ Manuscript received 8 April 1959-ED.
BULLETIN FLORIDA STATE MUSEUM
the first locality. Both quarries have surface elevations of about
90 feet (Gunter, 1948, p. 22) and are on the northeast edge of a terrace
separating Hogtown Sink and Lake Kanapaha to the north from Levy
Prairie and Paynes Prairie to the south.
MAP. Arredondo, Florida, and vicinity. Mines with Arredondo clay ex-
posed: A, Arredondo; F, Fort Clark; H, Haile; J, Kanapaha; K, Kendrick; O,
Orange Lake; R, Reddick; W, Williston; Z, Zuber. 100 foot contour indicates
approximate shore of Wicomico sea.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 271
The surficial layer (see table 1) is composed of reddish brown
sand. This is the "Newberry sand" of Matson and Sanford (1913, p.
32), which has been correlated with the Wicomico terrace formed
during the standstill of the sea at 100 feet elevation during the Sanga-
mon interglacial stage of the Pleistocene (Cooke, 1945, p. 281; Mac-
Neil, 1950, p. 99). Matson and Sanford's use of the term "Newberry
sand" antedates that of Dennison (1923, p. 628) for a deposit of Per-
mian age, but it is not in current use and is not necessarily endorsed
here as distinct from the Wicomico formation.
STRATIGRAPHIC SECTION AT ARREDONDO, FLORIDA
Bed Description Thickness
1 Pleistocene series, Wicomico formation
Reddish brown, massive, unfossiliferous marine terrace sand 9 feet
2 Wicomico formation, Arredondo clay member. Blue clay,
streaked with yellow; weathers to brown. Grades locally into 14 feet
sandy clay, white sand with alternating bands of yellowish gray,
argillaceous sand, and at bottom of bed into blue and blackish
brown clay. Surface irregular. Contains shells of terrestrial
and fresh water gastropods and vertebrate fossils.
3 Eocene series, Ocala group, Crystal River formation.
White, soft and friable, fossiliferous limestone. Surface irreg- 14 feet
ular with many steep-walled solution pipes. Bottom of bed not
exposed beyond water table.
The bedrock consists of limestone of the Crystal River formation
of the Ocala group (Puri, 1953, p. 130). Its surface contains many
steep-walled solution pipes and sinks with depths up to 20 feet.
Where the surface of the limestone is relatively low, and particu-
larly in solution pits, a bone-bearing clay layer intervenes between
the Wicomico sand and the Crystal River limestone. This clay is a
fresh water deposit, as it contains remains of amphibians (Siren, Pseu-
dobranchus, Bufo, and Rana) as well as land and fresh water gastro-
pods (Polygyra and Physa). Evidence will be presented below which
indicates that it was deposited under cooler climatic conditions,
namely during the Illinoian glacial stage.
Although the clay at Arredondo agrees with the Alachua clay (Dall
and Harris, 1892, p. 127) in being of fresh water or terrestrial origin,
the two clays differ in lithology, in cycles of sedimentation, and in
BULLETIN FLORIDA STATE MUSEUM
faunas. The Alachua formation, at least in its bone-bearing facies, is
more yellowish and often contains a high concentration of phosphate.
The clay at Arredondo is mostly bluish when freshly exposed and is
without important phosphate content. The Alachua clay has no defi-
nite relationship to Pleistocene marine terraces, but the Arredondo
clay invariably lies below the Wicomico terrace as a member of the
Wicomico formation. The vertebrates of the Alachua clay have been
assigned to the lower Pliocene (G. Simpson, 1929, p. 259), whereas
the Arredondo clay member contains Pleistocene indicators (table 2).
PLEISTOCENE INDICATORS AT ARREDONDO, FLORIDA
Pseudobranchus robustus Goin and Auffenberg. Ex-
Testudo sellardsi Hay. Extinct giant tortoise
Terrapene putnami Hay. Extinct box turtle
Podilymbus magnus Shufeldt. Extinct grebe
Falco readei, n. sp. Extinct falcon
Colinus suilium, n. sp. Extinct quail
Porzana auffenbergi Brodkorb. Extinct rail
Fulica minor Shufeldt. Extinct coot
Dorypaltus prosphatus, n. g. and sp. Extinct lapwing
Tachycineta speleodytes Brodkorb. Extinct swallow
Cremaster tytthus, n. g. and sp. Extinct hangnest
Desmodus, n. sp., Gut (in press). Extinct vampire bat
Dasypus bellus (Simpson). Extinct armadillo
Synaptomys australis Simpson. Extinct bog lemming
Aenocyon ayersi (Sellards). Extinct dire wolf
Mammut americanum (Kerr). Extinct mastodon
Tapirus veroensis Sellards. Extinct tapir
Equus sp. Extinct one-toed horse
Mylohyus cf. gidleyi Simpson. Extinct peccary
Tanupolama mirifica Simpson. Extinct camel
Tanupolama cf. americana Wortman. Extinct camel
The type section is exposed at Pit 1, NW 1/4 sec. 22, T 10 S, R 19 E,
Arredondo, Alachua County, Florida. The term Arredondo clay mem-
ber is proposed for the 14 feet represented by Bed 2.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 273
The Arredondo clay member has been studied along a northwest-
southeast transect 40 miles in length in Marion County (mines near
Kendrick, Zuber, Reddick, and Orange Lake) and Alachua County
(mines near Arredondo, Kanapaha, Fort Clark, and Haile), and it
also occurs at Williston in Levy County (see map). All exposures
seen lie in localities whose surface elevation is below the 100-foot con-
The bottom of the Arredondo clay member, in the areas studied,
lies between 55-68 feet above sea level, with its top at 74-89 feet.
The upper surface of the Wicomico terrace at these localities is be-
tween 90-95 feet above present sea level.
The Ocala uplift began in early Miocene times, and subaerial
erosion has since removed most of the younger sediments, to ex-
pose the Crystal River formation and expedite sink formation (Vernon,
1951, pp. 184-185). This process, although in a sense continuous
since Miocene time and still taking place today, was interrupted by
periodic transgressions of the sea. During the subsequent regressions
of the sea, fresh water sediments were deposited under a terrestrial
environment on the eroded surface of the Crystal River formation
during the Miocene, Pliocene, and Pleistocene (Vernon, 1951, p. 193).
None of the deposits contain mixed faunas, however, and it is thought
that previous reports of mixed vertebrate faunas in these areas re-
sulted from lack of stratigraphic control.
Podilymbus magnus Shufeldt.
Pit. 1: right humerus, left ulna. Pit 2: two right coracoids,
right and left humeri, two right and one left carpometacarpi, two
right and one left tarsometatarsi.
Shufeldt (1913, p. 136) separated the Pleistocene Pied-billed Grebe
on the basis of large size, but Wetmore (1937, p. 198) synonymized
it with the living Podilymbus podiceps because of the sexual dimorph-
ism existing in this genus. Compared with 8 modern specimens, 4
of each sex, the Arredondo material is also large. Measurements are
as follows: width of humerus through epicondylar prominences, 7.5-
8.0 (P. podiceps, 6.4-7.6); height of carpometacarpus through first
metacarpal, 6.9 (P. podiceps, 5.6-6.3); length of first metacarpal, 5.0
(P. podiceps, 3.8-4.6); least width of shaft of tarsometatarsus, 3.0-3.5
BULLETIN FLORIDA STATE MUSEUM
(P. podiceps, 2.7-3.0 mm.). It therefore seems advisable to revive
Querquedula discors (Linnaeus).
Pit 2: left coracoid, left carpometacarpus.
Nettion carolinense (Gmelin).
Pit 2: left coracoid.
Spatula clypeata (Linnaeus).
Pit 2: left tibiotarsus. The first record of the Shoveller from the
Aythya collaris (Donovan).
Pit. 1: left humerus. Pit. 2: left humerus, right tarsometatarsus.
Accipiter cooperii (Bonaparte).
Pit 1: right carpometacarpus.
Buteo jamaicensis (Gmelin).
Pit 1: left tibiotarsus.
Genus Falco Linnaeus
Tibiotarsus with (1) two distal openings under bridge; (2) condyles
parallel with each other and with axis of shaft; (3) posterior intercon-
dylar fossa shallow; (4) a pit in anterior intercondylar fossa at medial
side of external condyle.
Subgenus Hierofalco Cuvier
Tibiotarsus with (1) intercondylar fossa relatively deep distally;
(2) intercondylar pit very deep, excavating medial side of external
Falco readei n. sp.
HOLOTYPE.-Distal end of left tibiotarsus, Brodkorb collection
no. 1692. From Illinoian stage of Pleistocene (Arredondo clay mem-
ber), at Pit 2, Arredondo, Alachua County, Florida. Collected by
Ernest H. Reade, Jr., 21 September 1956.
DrAGNOSIs.-Similar to living Falco (Hierofalco) mexicanus Schle-
gel of western North America, but (1) external ligamental prominence
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 275
a small knob; (2) no shelf connecting
external ligamental prominence and
groove for peroneus profundus; (3) in-
tercondylar pit elongated in an oblique-
ly transverse direction toward anterior
edge of external condyle; (4) size about
half. Distal width, 8.0; depth of ex-
ternal condyle, 6.5; anterior height of
external condyle, 4.7; depth of internal
condyle, about 6.5 mm.
Differs from all subgenera men-
tioned below in having distal end of
intercondylar fossa deeper.
Resembles living Falco (Rhyncho-
falco) fuscocaerulescens Vieillot of Figure 1
tropical America in size and in very
deep intercondylar pit, but differs further in having pit not round, but
obliquely transverse, extending toward anterior edge of condyle; ex-
ternal ligamental prominence much smaller and not forming a shelf.
Resembles the living forms Falco (Falco) albigularis Daudin of
tropical America, Falco (Falco) subbuteo Linnaeus of the Palearctic
region, Falco (Tinnunculus) columbarius Linnaeus of the Holarctic
region, Falco (Erythropus) amurensis Radde of eastern Asia, and
Falco (Cerchneis) tinnunculus Linnaeus of the Palearctic region in
having external ligamental prominence knob-like, but differs in hav-
ing intercondylar pit deep and size larger.
Differs from Falco (Cerchneis) sparverius Linnaeus of America in
lacking external ligamental shelf; intercondylar pit deeper; size much
larger. Differs from the nearly cosmopolitan Falco (Rhynchodon)
peregrinus Tunstall in the same characters, except size much smaller.
The new species is much smaller than the Pleistocene Falco swarthi
L. Miller (1927, p. 152) of California and Falco oregonus Howard
(1946, p. 178) of Oregon. It is much larger than Falco ramenta Wet-
more (1936, p. 75) from the Miocene of Nebraska.
The records of Falco sp. from the Pleistocene of California (L.
Miller, 1912, p. 78, 95, 114; 1925, p. 99) were referred by Lambrecht
(1933, p. 751) to Falco fuscocaerulescens, although later Miller and
DeMay (1942, p. 67, 105) denied this determination. It seems possible
that they may belong to the present species.
Figure 1.-Falco readei, n. sp. Holotype tibiotarsus, no. 1692. X 5.
BULLETIN FLORIDA STATE MUSEUM
Falco (Rhynchodon) peregrinus Tunstall.
Pit 2: unguis.
Falco (Cerchneis) sparverius Linnaeus.
Pit 2: right carpometacarpus, right tibiotarsus, left tarsometa-
Bonasa umbellus (Linnaeus).
Pit 1: sternum. The Ruffed Grouse is new to the fauna of Florida.
Genus Colinus Goldfuss.
Humerus (1) longer than tarsometatarsus, with ratio of metatarsus
to humerus 86.94-98.42 per cent; (2) ratio of ulna to humerus 87.79-
91.01 per cent; (3) ratio of distal width of humerus to proximal width
69.39-76.14 per cent; (4) ratio of depth of caput humeri to proximal
width of humerus 34.69-39.13 per cent; (5) ratio of distance from proxi-
mal end of origin of brachialis to distal end of internal condyle and
distal width of humerus 79.10-89-71 per cent; (6) medial bar at angle
of about 105 degrees to shaft.
Colinus suilium n. sp.
HoLoTYPE.-Complete left humerus, Brodkorb collection, no. 1291.
From Illinoian stage of Pleistocene (Arredondo clay
member), at Pit 1, Arredondo, Alachua County,
Florida. Collected by Ernest H. Reade, Jr., 21 Sep-
ETYMOLOGY.-Latin, genitive plural of suile, pig-
sty, in reference to the former course of Hogtown
Creek through the type locality. Gainesville was
called Hogtown in 1830, after the Seminole chief
Hogmaster (C. Simpson, 1956).
DIAGNOSIs.-Humerus similar to that of living
Central American Colinus leucopogon (Lesson) and
North American C. virginianus (Linnaeus), but (1)
larger; (2) region around scar of infraspinatus pro-
duced; (3) scar of brachialis larger; (4) entepicondy-
lar prominence large and rising abruptly from shaft;
(5) ectepicondyle more rotated; (6) ectepicondylar
Figure 2 prominence larger and more deeply notched.
Figure 2.-Colinus suilium, n. sp. Holotype humerus, no. 1291. X 1.6.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 277
Differs from Colinus hibbardi Wetmore (1944, p. 96) from the
upper Pliocene of Kansas in being smaller and in having scar of
brachialis smaller. Other critical areas are missing in the material
of C. hibbardi.
PARATYPE.-Complete right humerus (no. 757), Pit 1.
REFERRED HUMER.-Pit 1: left proximal (nos. 758, 1290, 1292, 1624).
Pit 2: right distal (nos. 1672, 1703), left proximal (nos. 1671, 1673).
Measurements of comparative material are based on 2 modern
specimens of C. leucopogon, 19 modern specimens of C. virginianus
(represented by the subspecies C. v. virginianus, C. v. floridanus, C. v.
coyolcos, and C. v. insignis), and Tordoff's (1951, pp. 25-28) measure-
ments of C. hibbardi.
Length, 37.6-37.9 (C. leucopogon, 33.7-34.4; C. virginianus, 31.7-
35.8); proximal width, 9.6-10.4 (C. leucopogon, 9.2-9.3; C. virginiaus,
9.3-9.8); width of shaft, 3.4-3.8 (C. leucopogon, 3.1-3.2; C. virginianus,
2.6-3.3); distal width, 7.5-7.9 (C. leucopogon, 6.8-7.0; C. virginianus, 6.3-
6.8; C. hibbardi, 8.0); depth of head, 3.7-4.1 (C. leucopogon, 3.5-3.6;
C. virginianus, 3.1-3.5); proximal end of brachialis to distal end of
internal condyle, 6.3 (C. leucopogon, 5.6-5.8; C. virginianus, 5.3-6.1;
C. hibbardi, 6.9).
STERNUM.-Pit 1 (no. 1622). Estimated width of ventral lip of
Figure 3 Figure 4 Figure 5
Figure 3.-Colinus suilium, n. sp. Referred sternum, no. 1622. X 2.
Figure 4.-Colinus suilium, n. sp. Referred coracoid, no. 1623. X 2.
Figure 5.-Colinus suilium, n. sp. Referred ulna, no. 1324. X 1.5.
BULLETIN FLORIDA STATE MUSEUM
coracoidal sulcus, 11.5 (C. leucopogon, 9.6-10.8; C. virginianus, 9.2-
11.2; C. hibbardi, 13.4).
SCAPULA.-Pit 2: left (no. 1648). Acromion narrower and recurved
toward medial side, compared with living species. Scapula of C.
CORACOiD.-Pit 1: right (no. 1623) and left (no. 1294). Pit 2: right
(no. 1629) and left (no. 1701). Head depressed as in C. hibbardi.
Length along medial edge, 29.7-30.3 (C. leucopogon, 26.4-26.7; C. vir-
ginianus, 24.5-28.1); sternal width, 9.2 (C. leucopogon, 8.0; C. virgini-
anus, 7.3-9.1; C. hibbardi, 11.3); head to glenoid facet, 8.7-8.8 (C. leu-
copogon, 7.7-8.2; C. virginianus, 7.6-8.7; C. hibbardi, 9.1-10.0); depth
posterior to furcular facet, 3.4-3.5 (C. leucopogon, 2.6-3.0; C. virgini-
anus, 2.6-3.3; C. hibbardi, 3.4-3.8).
ULNA.-Pit 1: right (no. 1625) and left (no. 1324). Pit 2: left (no.
1702). Brachial impression deep, as in C. leucopogon and C. hibbardi.
Figure 6.-Colinus suilium, n. sp. Referred synsacrum, no. 756. X 1.5
Figure 7.-Colinus suilium, n. sp. Referred tibiotarsus, no. 1545. X 1.5.
Figure 8.-Colinus suilium, n. sp. Referred tarsometatarsus, no. 1626. X 1.4.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 279
Length, 32.7-33.1 (C. leucopogon, 30.0-30.2; C. virginianus, 28.5-32.2;
C. hibbardi, 37.4-38.7); proximal width, 4.1-4.6 (C. leucopogon, 3.8-
4.1; C. virginianus, 3.4-4.3; C. hibbardi, 4.8-4.9); distal width, 3.1-3.3;
(C. leucopogon, 3.4-3.5; C. virginianus, 2.6-3.2; C. hibbardi, 4.2-4.8);
distal depth, 3.9-4.0 (C. leucopogon, 3.7; C. virginianus, 3.3-3.6).
SYNSACRUM.-Pit 1 (no. 756). Fourth and fifth synsacral vertebrae
with transverse processes extending to innominates. Length from
first synsacral centrum through fourth transverse process, 12.6 (C.
leucopogon, 11.7-12.1; C. virginianus, 11.2-12.5).
TIBIOTARSUS.-Pit 1: right (no. 1545) and left (no. 1634). Pit 2:
right (no. 1649) and left (no. 1704). In proximal view, posterior notch
between internal and external articular surfaces deeper than in living
species, cutting farther into internal articular surface; in medial view,
inner cnemial crest with anterior margin strongly curved, not angled.
Length, 58.1 (C. leucopogon, 50.3-52.4; C. virginianus, 49.8-53.5); width
through condyles, 5.4 (C. leucopogon, 4.8-5.1; C. virginianus, 4.6-5.2);
depth of external condyle, 5.5 (C. leucopogon, 5.0; C. virginianus, 4.6-
5.1); depth of internal condyle, 5.2-5.7 (C. leucopogon, 5.2-5.3; C. vir-
TARSOMETATARSUs.-Pit 1: right (no. 1626). Pit 2: left (no. 1647).
Second trochlea lower on shaft than in C. virginianus, thus resembling
C. leucopogon. Length, 34.5 (C. leucopogon, 29.3-30.3; C. virginianus
29.2-32.2); proximal width, 5.8-6.0 (C. leucopogon, 5.5; C. virginianus,
5.0-5.7; C. hibbardi, 6.5); least width of shaft, 2.6-2.7 (C. leucopogon,
2.5-2.6; C. virginianus, 2.2-2.5; C. hibbardi, 3.0-3.1); distal width, 6.5
(C. leucopogon, 5.2-5.8; C. virginianus, 5.3-5.6; C. hibbardi, 6.6-7.0);
proximal depth, 6.0-6.1 (C. leucopogon, 5.6; C. virginianus, 5.2-5.8);
width of middle trochlea, 2.5 (C. leucopogon, 2.0-2.1; C. virginianus,
1.9-2.3; C. hibbardi, 2.4-2.9); depth of middle trochlea, 3.4 (C. leuco-
pogon, 2.8; C. virginianus, 2.6-3.2; C. hibbardi, 3.1-3.4).
Meleagris gallopavo Linnaeus.
Pit 1: left humerus. Pit 2: left humerus.
Rallus elegans Audubon.
Pit 2: left scapula.
Rallus limicola Vieillot.
Pit 2: left carpometacarpus, right tibiotarsus.
BULLETIN FLORIDA STATE MUSEUM
Porzana carolina (Linnaeus).
Pit 2: left coracoid.
Porzana auffenbergi Brodkorb.
Pit 2: left tibiotarsus (no. 1717).
This species was described from Haile, Florida, on the basis of a
humerus (Brodkorb, 1954, p. 103). The presently referred tibiotarsus
agrees with that of P. carolina in having a relatively stout shaft com-
pared with Rallus limicola, but it is larger. Width through condyles,
4.7; depth of external condyle, 4.7; depth of internal condyle, 4.8;
least width of shaft, 2.2.
Porphyrula martinica (Linnaeus).
Pit 2: right coracoid.
Gallinula chloropus (Linnaeus).
Pit 2: left ulna, right tibiotarsus.
Fulica minor Shufeldt.
Pit 2: right humerus, left femur, left tibiotarsus, right tarsometa-
Most of these specimens are too fragmentary for specific deter-
mination, but the tibiotarsus has the internal condyle relatively deeper
than in F. americana. Depth of external condyle, 8.7 mm.; depth
of internal condyle, 9.0 mm.; ratio, 96.7 per cent.
Humerus with (1) ectepicondylar process long, with its anconal
face somewhat concave; (2) external condyle rotated 45 degrees or
less along its internal margin; (3) facet for anterior articular ligament
merging gently at its lower end with distal extension of brachial de-
pression, without intervening shelf.
Humerus with (1) internal condyle a flattened oval, little produced
distally; (2) entepicondyle oblique, not produced distally and internal-
ly; (3) facet for anterior articular ligament abruptly included lat-
erally; (4) scar of pronator brevis large, expanded laterally, equal to
one-third to half width of facet for anterior articular ligament; (5)
scar of brachialis deeply excavated, located relatively high with half
to one-third its length above level of ectepicondylar process, its distal
end above level of articular ligament; (6) foramen in lower portion
of brachial depression located directly above internal margin of ex-
ternal condyle; (7) external condyle rotated less than 45 degrees.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 281
Genus Dorypaltus n. gen.
TYPE OF GENUS.-Dorypaltus prosphatus n. sp.
ETYMOLOGY.-Greek, dorypaltos, masculine, a spear-wielder. Greek,
prosphatos, not decomposed, of a corpse miraculously preserved.
DIAGNosIS.-Resembles living South American Belonopterus Reich-
enbach and differs from living Palearctic Vanellus Brisson in having (1)
entepicondyle only moderately produced medially in area of pit for
pronator longus; (2) side of entepicondyle strongly concave opposite
scar of anterior articular ligament; (3) area of origin of pronator brevis
forming a lengthened oval shelf; (4) ectepicondyle rounded at proximal
end, not forming a spur; (5) anconal face of ectepicondyle more con-
Differs from Belonopterus in having (1) scar of brachialis still more
deeply excavated and wider, extending laterally to meet scar for pal-
mar branch of extensor metacarpi radialis; (2) the latter muscle scar
longer and wider, extending distally nearly to level of brachial fora-
men, and laterally from scar of brachialis nearly to end of ectepicon-
dyle; (3) entepicondyle even less produced laterally, in area of origin
of pronator longus.
Dorypaltus prosphatus n. sp.
HOLOTYPE.-Distal portion of left humerus, Brodkorb collection,
no. 1712. From Illinoian stage of Pleistocene (Arredondo clay mem-
ber), at Pit 2, Arredondo, Alachua County,
Florida. Collected by Ernest H. Reade, Jr.,
12 September 1956.
DIAGNOSIS.-Humerus slightly smaller .
than in Belonopterus chilensis (Molina), with
ectepicondyle relatively longer. Distal width,
9.5; width through proximal end of ectepi- -
condylar process, 7.4; width of shaft above
ectepicondylar process, 6.3; length through
external condyle and ectepicondylar process,
REFERRED SPECIMEN.-Proximal end of
right humerus (no. 1713), collected with the
holotype; lacks external tuberosity. Slightly Figure 9
smaller than Belonopterus chilensis; smaller
Figure 9.-Dorypaltus prosphatus, n. g., n. sp. Holotype humerus, no. 1712.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 283
Melanerpes carolinus (Linnaeus).
Pit 2: right ulna. The first fossil record of the Red-bellied Wood-
Tachycineta speleodytes Brodkorb.
Pit 2: right coracoid, four right and two left humeri, one right
and five left ulnas, left carpometacarpus, right femur. Two of the
humeri are from juvenile individuals of this recently described swal-
low (Brodkorb, 1957, p. 131).
Corvus brachyrhynchos Brehm.
Pit 2: right tarsometatarsus.
Corvus ossifragus Wilson.
Pit 1: right humerus. Pit 2: left ulna.
Cyanocitta cristata (Linnaeus).
Pit 2: left tarsometatarsus.
Aphelocoma coerulescens coerulescens (Bosc).
Pit 1: right tibiotarsus.
Although Aphelocoma coerulescens californica has been reported
from the Pleistocene of California (A. H. Miller, 1932B, p. 173), this
is the first fossil record of the Florida Scrub Jay. These two races,
formerly considered distinct species, are separable osteologically by
Lanius ludovicianus Linnaeus.
Pit 1: left tarsometatarsus. Previously unrecorded from the Ple-
istocene of Florida.
Agelaius phoeniceus (Linnaeus).
Pit 2: right humerus, right ulna, right femur, two right and one
Sturnella magna (Linnaeus).
Pit 2: two left ulnas.
Tibiotarsus with (1) distal margin of intercondylar space oblique,
with internal sulcus the deeper one; (2) anterior wall of intercondylar
BULLETIN FLORIDA STATE MUSEUM
fossa separated from distal opening of tibial bridge by a sharply raised
ridge connecting condyles.
Genus Cremaster n. gen.
TYPE OF GENus.-Cremaster tytthus n. sp.
ETYMOLOGY.-Greek, cremaster, masculine, something hanging
down like a basket or a bunch of grapes, in allusion to the supposed
habits. Greek, tytthos, little.
DIAGNOSIS.-An arboreal icterid agreeing with the living Neotropi-
cal oropendolas and caciques of the genera Ostinops, Gymnostinops,
Zarhynchus, and Cacicus in having tibiotarsus with (1) distal portion
of shaft straight, stout, and nearly as wide as condyles; (2) internal
ligamental ridge proximal to tibial bridge only moderately developed.
Differs in having (1) external condyle in anterior view parallel
with and scarcely protruding from shaft; (2) external ligamental prom-
inence reduced, not forming a shelf; (3) tendinal canal under tibial
bridge confined to lateral portion of bone; (4) tibial bridge transverse,
with its lateral margin produced proximally to form a bluntly rounded
projection; (5) surface of tibial bridge slanting in a smooth plane to
meet anterior portion of shaft, which hence continues as a flat surface
toward medial edge of bone; (6) upper opening leading under bridge
rounded; (7) lower opening with proximal margin forming an obtuse
angle, with the short leg lateral; (8) internal condyle but slightly pro-
truding from medial line of shaft; (9) anterior intercondylar fossa deep,
forming a pit with sharply defined distal as well as
Less closely allied to the living caciques of the Neo-
tropical genera Archiplanus, Amblycercus, and Cassicu-
lus, but shaft stouter, with condyles scarcely protruding;
internal ligamental ridge more weakly developed prox-
imal to bridge.
t. Bears a superficial resemblance to the tibiotarsus of
the larger species of orioles of the genus Icterus, particu-
larly I. icterus and I. gularis, but immediately separable
Sby its stouter shaft with the condyles less protruding
from it in anterior view, transverse instead of oblique
bridge, and reduced external ligamental prominence.
V Cremaster tytthus n. sp.
Figure 10 HOLOTYPE.-Distal portion of left tibiotarsus, Brod-
Figure 10.-Cremaster tytthus, n. g., n. sp. Holotype tibiotarsus, no. 1663. X 3.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 285
korb collection, no. 1663. From Illinoian stage
of Pleistocene (Arredondo clay member), at Pit
2, Arredondo, Alachua County, Florida. Col-
lected by Pierce Brodkorb, 6 May 1956.
DIAGNOSIS.-Structurally closest to Ostinops
decumanus (Pallas) but separable on generic
characters; size much less. Distal width, 3.7;
least width of shaft, 2.0; depth of internal con-
dyle, 3.4; depth of external condyle, 3.4 mm.
REFERRED MATERIAL.-Pit 2: right humerus
(no. 1664) and right femur (no. 1694).
Humerus agrees with that of Ostinops, Caci-
cus, and Icterus icterus in having proximal por-
tion contracted, with attachment for infraspi-
natus only slightly produced. Differs in hav-
ing (1) subtrochanteric and tricipital fossae (fos-
sae I and II) without partially roofing shelf
from medial bar; (2) floor of fossae at a single
level; (3) medial bar extending to floor of fossae
rather than to external side, and directed at its
base toward distal end of bone at angle of 120
degrees to shaft; (4) distal portion of bone Figure 11
strongly rotated and only slightly expanded;
(5) brachial depression forming a deep, com-
pressed, steep-walled pit; (6) ectepicondylar prom-
inence extending above level of ectepicondylar
process as a shelf which is bent at an angle before
reaching shaft. Length, 27.1; proximal width, 7.9;
width of shaft, 2.5; distal width, 6.2 mm. The posi-
tion of the medial bar and the excavation of the
pneumatic fossae indicate (Ashley, 1941, pp. 192-
194) that the humerus of Cremaster is more primi-
tive than that of Ostinops or Cacicus.
The referred femur agrees more closely with
that of Icterus icterus than with Ostinops in having
the condyles lengthened and compressed from front
to rear, and the notch for the femoral head of the
tibialis anticus more anterior in position. It differs
from Icterus in having (1) fibular condyle produced Figure 12
Figure 11.-Cremaster tytthus, n. g., n. sp. Referred humerus, no. 1664. X 3.
Figure 12.-Cremaster tytthus, n. g., n. sp. Referred femur, no. 1694. X 3.3.
BULLETIN FLORIDA STATE MUSEUM
proximally toward pit of origin of external part of gastrocnemius,
leaving only a narrow channel between, devoid of pneumatic foramina;
(2) wing of internal condyle somewhat produced internally; (3) inter-
condylar fossa with excavation confined to its external portion. Width
through condyles, 5.6; width of shaft, 2.6 mm.
Richmondena cardinalis (Linnaeus).
Pit 1: right tibiotarsus. The first fossil record of the Cardinal.
Pipilo erythrophthalmus (Linnaeus).
Pit 1: mandible, left humerus, left ulna, left tibiotarsus.
Passerherbulus henslowii (Audubon).
Pit 2: left ulna.
Passerculus sandwichensis (Gmelin).
Pit 2: right carpometacarpus. The first fossil record of the Sa-
Spizella passerina (Bechstein).
Pit 1: right carpometacarpus. The Chipping Sparrow was known
previously in the fossil record only on the basis of a tentative deter-
mination from California (Sibley, 1939, p. 127).
Spizella pusilla (Wilson).
Pit 1: left humerus. The first fossil record of the Field Sparrow.
The birds of the Arredondo local fauna fall naturally into two
groups, a fresh water marsh community with adjacent ecotone of wet
meadows, and a scrub community.
The fresh water marsh community included the grebe (Podilym-
bus), four ducks (Querquedula, Nettion, Spatula, and Aythya), seven
species of rails (Rallus, Porzana, Porphyrula, Gallinula, and Fulica),
and the Red-winged Blackbird (Agelaius).
The ecotone was inhabited by a plover (Dorypaltus), two sand-
pipers (Tringa), meadowlark (Sturnella), and at least two sparrows
(Passerherbulus and Passerculus). The swallow (Tachycineta) prob-
ably nested on rocky outcrops and fed over the wet meadows and
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 287
The scrub community was occupied by the Ruffed Grouse (Bonasa),
Scrub Jay (Aphelocoma), and Loggerhead Shrike (Lanius). The re-
maining species of the Arredondo local fauna, insofar as their habits
are known, are of wider ecological tolerance. They all might be ex-
pected to occur in the ecotone or in the scrub community.
The Scrub Jay is confined to the St. Lucie scrub community. This
community occupies well-drained sandy areas dominated by sand pine
(Pinus clausa) and scrub oaks with evergreen shrubs, including rose-
mary (Ceratiola ericoides), in the understory. The scrub occurs on
low dunes near the coast and on certain fossil dune areas in the inte-
rior of the state.
Much of the original forest at Arredondo has been destroyed, but
the commonest remaining trees include loblolly pine (Pinus taeda),
mockernut hickory (Carya tomentosa), laurel oak (Quercus laurifolia),
and sweet gum (Liquidambar styraciflua). This assemblage of trees
is characteristic of mesophytic hammock, the present climatic climax
community. Since none of the birds of the Arredondo local fauna is
confined to mesophytic hammock, there is no proof of the existence
of this community at Arredondo during Illinoian time.
The Florida scrub represents an early stage in a xerosere (Laessle,
1942, p. 96). It succeeds to xerophytic hammock with live oak (Quer-
cus virginiana), and further succession leads to the present climatic
climax of mesophytic hammock. The presence of the Scrub Jay there-
fore indicates that the environment at Arredondo has passed through
two seral stages and into the present climatic climax since Illinoian
On the basis of time and place of origin the Illinoian fauna of Ar-
redondo may be divided into two categories, interglacial relicts and
The interglacial relicts show a high proportion of endemism. They
include Falco readei, Colinus suilium, Dorypaltus prosphatus, Aphelo-
coma coerulescens coerulescens, and Cremaster tytthus. Four of these
represent species or genera which are now extinct but whose affinities
lie to the southwest. Aphelocoma still survives in Florida today, al-
though its nearest outposts lie 35-40 miles to the south or east of
The species Aphelocoma coerulescens has a discontinuous distri-
bution at present. Seventeen subspecies occupy the area from south-
ern Mexico to the Great Basin, and eastward to the Edwards Plateau.
BULLETIN FLORIDA STATE MUSEUM
There is then a hiatus of a thousand miles to the range of the Florida
subspecies. The survival of this genus in Florida today, as well as
of certain other southwestern animals and plants isolated on the pen-
insula, is explained by the supposedly continuous extension eastward
of the sclerophyll woodland of the southwest during late Tertiary times
(Pitelka, 1951, pp. 383-384).
Several species of the Arredondo local fauna comprise a boreal
element and are therefore indicative of a cooler climate.
The Ruffed Grouse (Bonasa umbellus) is a bird of the northern
coniferous forest and adjacent ecotones. During historical times its
range extended southward through the Appalachians, but in the low-
lands it occurred only as far as Chesapeake Bay (Aldrich and Duvall,
1955, pp. 6-7), and its previous Pleistocene records all lie within its
recent range (Wetmore, 1956, p. 52).
Tachycineta speleodytes was the temporal representative of the
tree swallow (T. bicolor). Today the genus reaches its southern breed-
ing limits on the Atlantic seaboard in Virginia and Maryland, and in
the Mississippi Valley near the line of maximum glaciation (Bent,
1942, p. 398).
A boreal element among the mammals is the lemming (Synap-
tomys australis). At present the genus extends southward on the At-
lantic coastal plain only to Maryland, with a relict colony in the Dis-
mal Swamp (Miller and Kellogg, 1955, pp. 561-565).
As the three genera Bonasa, Tachycineta, and Synaptomys have
their present southern limits 700 miles to the north, their presence
in the Arredondo local fauna is thought to indicate a glacial stage.
Because of the stratigraphic relationship of the Arredondo clay to
deposits of Sangamon age, the fauna is thought to be of Illinoian gla-
The stratigraphy was verified in the field by Robert O. Vernon of
the Florida Geological Survey, E. C. Pirkle, Jr., of the University of
Florida, and William F. Jenks of the University of Cincinnati. Mol-
lusca were determined by Robert M. DeWitt of the University of
Florida. Osteological material was borrowed from the Museum of
Vertebrate Zoology at the University of California through Frank A.
Pitelka, and from the United States National Museum through Her-
bert Friedmann. The photographs were taken by Robert D. Weigel.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 289
Aldrich, John W., and Allen J. Duvall
1955. Distribution of American gallinaceous birds. U. S. Fish and Wildlife
Service, circular 34, pp. 1-30.
Ashley, James F.
1941. A study of the structure of the humerus in the Corvidae. Condor, vol.
43, pp. 184-195, figs. 50-56.
1956. Additional records of Pleistocene lizards from Florida. Quart. Jour.
Florida Acad. Sci., vol. 19, pp. 157-167.
1958. Fossil turtles of the genus Terrapene in Florida. Bull. Florida State
Mus., vol. 3, no. 2, pp. 53-92, 15 figs.
Bader, Robert S.
1957. Two Pleistocene mammalian faunas from Alachua County, Florida.
Bull. Florida State Mus., vol. 2, no. 5, pp. 53-75, 6 figs.
Bent, Arthur Cleveland
1942. Life histories of North American flycatchers, larks, swallows, and their
allies. Bull. U. S. Nat. Mus., no. 179, pp. i-xi, 1-555, pl. 1-70.
1954. Another new rail from the Pleistocene of Florida. Condor, vol. 56, pp.
103-104, 1 fig.
1957. New passerine birds from the Pleistocene of Reddick, Florida. Jour.
Paleont., vol. 31, pp. 129-138, pl. 20, text-fig. 1.
Cooke, C. Wythe
1945. Geology of Florida. Florida Geol. Surv., Geol. Bull. 29, pp. i-ix, 1-339,
1 pl., 47 text-figs.
Dall, William Healey, and Gilbert D. Harris
1892. Correlation papers; Neocene. U. S. Geol. Surv., Bull. 84, pp. 1-349.
Dennison, A. R.
1923. The Robberson field, Garvin County, Oklahoma. Bull. Amer. Assoc.
Petrol. Geol., vol. 7, pp. 627-644.
Goin, Coleman J., and Walter Auffenberg
1955. The fossil salamanders of the family Sirenidae. Bull. Mus. Comp. Zool.,
vol. 113, no. 7, pp. 497-514, 3 figs.
1949. Elevations in Florida. Florida Geol. Surv., Geol. Bull. 32, pp. i-xxiii,
1946. A review of the Pleistocene birds of Fossil Lake, Oregon. Carnegie Inst.
Wash., Publ. 551, pp. 141-195, pl. 1-2.
BULLETIN FLORIDA STATE MUSEUM
Laessle, Albert Middleton
1942. The plant communities of the Welaka area with special reference to
correlations between soils and vegetational succession. Univ. Florida
Publ., Biol. Sci. Ser., vol. 4, pp. 1-143, 14 pl.
1933. Handbuch der Palaeornithologie. Gebriider Borntraeger, Berlin. Pp.
i-xx, 1-1024, 209 figs.
MacNeil, F. Stearns
1950. Pleistocene shore lines in Florida and Georgia. U. S. Geol. Surv., Prof.
Paper 221-F, pp. 95-107, pl. 19-25.
Matson, George C., and Samuel Sanford
1913. Geology and ground waters of Florida. U. S. Geol. Surv., Water-
Supply Paper 319, pp. 1-445, map.
Miller, Alden H.
1932. The fossil passerine birds from the Pleistocene of Carpenteria, California.
Univ. Calif. Publ., Bull. Dept. Geol. Sci., vol. 21, pp. 169-194, pl. 12-14.
Miller, Gerrit S., Jr., and Remington Kellogg
1955. List of North American Recent mammals. Bull. U. S. Nat. Mus., no.
205, pp. i-xii, 1-954.
1912. Contributions to avian paleontology from the Pacific coast of North
America. Univ. Calif. Publ., Bull. Dept. Geol., vol. 7, pp. 61-115.
1927. The falcons of the McKittrick Pleistocene. Condor, vol. 29, pp. 150-
152, fig. 54.
Miller, Loye, and Ida DeMay
1942. The fossil birds of California, an avifauna and bibliography with anno-
tations. Univ. Calif. Publ. Zool., vol. 47, pp. 47-142.
Olsen, Stanley J.
1958. The bog lemming from the Pleistocene of Florida. Jour. Mammalogy,
vol. 39, pp. 537-540, 2 figs.
Pitelka, Frank A.
1951. Speciation and ecologic distribution in American jayS of the genus
Aphelocoma. Univ. Calif. Publ. Zool., vol. 50, pp. 195-464, pl. 17-30.
Puri, Harbans S.
1953. Zonation of the Ocala group in peninsular Florida. Jour. Sed. Petrol.,
vol. 23, p. 130.
Shufeldt, R. W.
1913. Review of the fossil fauna of the desert region of Oregon, with a de-
scription of additional material collected there. Bull. Amer. Mus. Nat.
Hist., vol. 32, pp. 123-178, pl. 9-43.
1959 BRODKORB: AVIFAUNA OF ARREDONDO, FLORIDA 291
1915. Fossil birds in the Marsh collection of Yale University. Trans. Conn.
Acad. Arts Sci., vol. 19, pp. 1-110, 15 pl.
1939. Fossil fringillids from Rancho La Brea. Condor, vol. 41, pp. 126-127.
Simpson, George Gaylord
1929. The extinct land mammals of Florida. Florida Geol. Surv., 20th Ann.
Report, pp. 229-279, pl. 30-40, text-figs. 1-4.
Simpson, J. Clarence
1956. A provisional gazeteer of Florida place-names of Indian derivation either
obsolescent or retained together with others of recent application.
Florida Geol. Surv., Special Publ. no. 1, pp. i-x, 1-158, frontis., 5 maps.
Tordoff, Harrison B.
1951. Osteology of Colinus hibbardi, a Pliocene quail. Condor, vol. 53, pp.
23-30, figs. 1-2.
Vernon, Robert O.
1951. Geology of Citrus and Levy counties, Florida. Florida Geol. Surv.,
Geol. Bull. 33, pp. i-xi, 1-256, pl. 1-2, figs. 1-40.
1936. Two new species of hawks from the Miocene of Nebraska. Proc. U. S.
Nat. Mus., vol. 84, pp. 73-78, figs. 13-14.
1937. A record of the fossil grebe, Colymbus parvus, from the Pliocene of Cali-
fornia, with remarks on other American fossils of this family. Proc.
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