• TABLE OF CONTENTS
HIDE
 Main
 Table of Contents
 Dedication
 Introduction
 Materials and methods
 Systematic paleontology
 Liliales
 Proteales
 Fagales
 Malpighiales
 Rosales
 Sapindales
 Ericales
 Apiales
 Insecta
 Discussion
 Acknowledgments
 Back Cover






Group Title: Bulletin of the Florida Museum of Natural History
Title: A Middle Eocene fossil plant assemblage (Powers clay pit) from western Tennessee
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00087417/00001
 Material Information
Title: A Middle Eocene fossil plant assemblage (Powers clay pit) from western Tennessee
Series Title: Bulletin of the Florida Museum of Natural History ; vol. 45, no. 1
Physical Description: 43 p. : ill., maps ; 28 cm.
Language: English
Creator: Dilcher, David L.
Lott, T. A. ( Terry A. )
Publisher: Florida Museum of Natural History
Place of Publication: Gainesville, FL
Publication Date: 2005
Copyright Date: 2005
 Subjects
Subject: Angiosperms, Fossil -- Tennessee -- Weakley County   ( lcsh )
Plants, Fossil -- Tennessee -- Weakley County   ( lcsh )
Paleobotany -- Eocene   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Includes bibliographical references (p. 23-28).
General Note: Caption title.
General Note: "Publication date October 31, 2005"- - P. 2 of cover.
Statement of Responsibility: David L. Dilcher and Terry A. Lott.
 Record Information
Bibliographic ID: UF00087417
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 - 62458627
lccn - 2006475618
issn - 0071-6154 ;

Table of Contents
    Main
        Page i
        Page ii
    Table of Contents
        Page 1
    Dedication
        Page 2
    Introduction
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
    Materials and methods
        Page 8
        Page 9
    Systematic paleontology
        Page 10
    Liliales
        Page 11
    Proteales
        Page 12
    Fagales
        Page 13
    Malpighiales
        Page 14
    Rosales
        Page 15
    Sapindales
        Page 16
    Ericales
        Page 17
    Apiales
        Page 18
    Insecta
        Page 19
    Discussion
        Page 20
        Page 21
        Page 22
    Acknowledgments
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
        Page 41
        Page 43
    Back Cover
        Page 44
Full Text





FLORIDA
_MUSEUM
OF NATURAL HISTORY


BULLETIN


A MIDDLE EOCENE FOSSIL PLANT ASSEMBLAGE
(POWERS CLAY PIT) FROM WESTERN TENNESSEE



David L. Dilcher and Terry A. Lott


Vol. 45, No. 1, pp. 1-43


2005


UNIVERSITY OF FLORIDA GAINESVILLE


UNIVERSITY OF FLORIDA


GAIN ESVI LLE







The FLORIDA MUSEUM OF NATURAL HISTORY is Florida's state museum of natural history, dedicated to
understanding, preserving, and interpreting biological diversity and cultural heritage.

The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY is a peer-reviewed publication
that publishes the results of original research in zoology, botany, paleontology, and archaeology. Address all inquiries
to the Managing Editor of the Bulletin. Numbers of the Bulletin are published at irregular intervals. Specific volumes
are not necessarily completed in any one year. The end of a volume will be noted at the foot of the first page of the
last issue in that volume.




Richard Franz, Managing Editor
Erika H. Simons, Production



Bulletin Committee
Richard Franz, Chairperson
Ann Cordell
Sarah Fazenbaker
Richard Hulbert
William Marquardt
Susan Milbrath
Irvy R. Quitmyer
Scott Robinson, Ex officio Member




ISSN: 0071-6154

Publication Date: October 31, 2005






Send communications concerning purchase or exchange
of the publication and manuscript queries to:

Managing Editor of the BULLETIN
Florida Museum of Natural History
University of Florida
PO Box 117800
Gainesville, FL 32611-7800 U.S.A.
Phone: 352-392-1721
Fax: 352-846-0287
e-mail: dfranz@flmnh.ufl.edu







A MIDDLE EOCENE FOSSIL PLANT ASSEMBLAGE (POWERS CLAY PIT)

FROM WESTERN TENNESSEE





David L. Dilcher and Terry A. Lott'





ABSTRACT

Plant megafossils are described, illustrated and discussed from Powers Clay Pit, occurring in the middle Eocene, Claiborne Group
of the Mississippi Embayment in western Tennessee. Twenty six species and eight types of plants, and two species of insect
larval cases are represented in this study. They include Lauraceae, Annonaceae, Smilacaceae, Platanaceae, Altingiaceae, Myrtaceae,
Fabaceae, Fagaceae, Salicaceae, Moraceae, Rhamnaceae, Sapindaceae, Nyssaceae, Theaceae, Apocynaceae, Rubiaceae, Araliaceae,
Oleaceae, entire margin morphotype 1-5, tooth margin morphotype 1, reproductive structure morphotype 1, Folindusia, and
Terrindusia. Specimens collected from Powers Pit are compared to those from previous studies from western Tennessee, the
Claiborne Group in general, and assessed in terms of extant relationships. The extant relationships of plant megafossils described
in this study provide clues to the paleoenvironment of western Tennessee during the middle Eocene. The paleoenvironment may
have been subtropical accommodating warm tropical to cool temperate plant species.


Key words: Eocene, Claiborne Group, Powers Clay Pit, Tennessee, Angiosperms





TABLE OF CONTENTS

D education ........................................................................ ........................ 2
Introduction ......................................................................... .................... . 3
M materials and M ethods ....................................................... ....................... 8
Systematic Paleontology .................... ...................................10
L aurales ........................................................................... .................. . 10
M agnoliales ................................................................... ....................... 10
L ilia le s ........................................................ ................................................
P ro teales ........................................................... .......................................... 12
Saxifragales ..................................................................... ...................... 12
M y rtales ...................................................... ..............................................12
Fabales ........................................................................... .................... . 12
F ag ales .............................................................. .......................................... 13
M alpighiales ................................................................... ...................... 14
R o sales .......................................................... ............................................. 15
Sapindales ...................................................................... ................... . 16
C o r ales ........................................................ ............................................. 16
E rica les ............................................................ ........................................... 17
G entianales ...................................................................... ..................... 17
A p iales ........................................................... ............................................ 18
L am iales ......................................................................... .................... . 18
Incertae sedis ................................................................... ..................... 18
Insecta .................. ....................................................... 19
D iscu ssio n .......................................................................... .............................2 0
A know ledgm ents .............................................................. .......................23
L literature C ited ................................................. ........................................ 23




'Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800, email: , .
Dilcher, D.L. & T.A. Lott. 2005. A middle Eocene fossil plant assemblage (Powers Clay Pit) from western Tennessee. Bull. Florida Museum Nat.
Hist. 45(1):1-43.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


IN DEDICATION TO: BRUCE ROGER MOORE


This paper is dedicated to Mr. Bruce Roger Moore, a student of Paleobotany whose interest in the fossil plants
of western Tennessee and collections from the Powers Clay Pit helped contribute many fossils to the Paleobotany
and Palynology Collection of the Florida Museum of Natural History. His field-work, as part of a team effort, resulted
in collections of several of the fossils presented in this paper. His intense passion for collecting and inquiries about the
fossil plants and their depositional history encouraged continued study of fossil plants from the middle Eocene of
western Tennessee. Roger Moore is a native of western Tennessee and completed a Masters of Science degree
from the University of Tennessee in Agronomy. Subsequently, he went into business in western Tennessee. His
interest in plants continued, and when Professor Michael Gibson introduced him to the fossils of western Tennessee
and he saw how beautifully preserved the plants were, it was like striking a spark under dry kindling. The spark took
and flamed into an intense interest in these fossil plants. We are happy to dedicate this paper to him in recognition of
his significant contributions of collecting so many fossil plants and making them available for this study and future
reports of other fossil localities.





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

INTRODUCTION
There are numerous clay deposits in western Tennes-
see, many which contain abundant plant fossils. These
clay deposits are of middle Eocene age and occur as
isolated clay lenses of the Claiborne Group deposited
during the Mississippi Embayment of southeastern North
America. These numerous clay lenses are distributed
in the ancient coastal plains sediments of the northeast-
ern part of the Mississippi Embayment (Fig. 1). There


are many localities in this region (Fig. 2) and each local-
ity yielded hundreds to thousands, even tens of thou-
sands, of plant fossil remains. This report is the first
publication in what we expect to become a series of
papers detailing the floras of each locality. Reasons for
breaking this very large flora of middle Eocene plant-
rich deposits into discreet units are elaborated upon in
the discussion of this paper.
The fossil plants in the present report were col


Ter Jackson group
erl Tj Jackson Formation
Eocene Ty Yazoo Clay
May include Paleocene locally Tm Moodys Branch Formation
Tpr l
Paleocene

Kretac
Cretaceous


Claiborne group Wilcox group
Tc Cockfield Formation Th Hatchetigbee Formation
Tco Cook Mountain Formation Tw Wilcox Formation
Ts Sparta Sand
Tz Zilpha Clay
Tt Tallahatta Formation
Trm Reklaw Formation


Figure 1. Geologic map of Mississippi Embayment. Stratigraphic units as recognized by U.S. Geological
Survey. Modified from Tschudy (1973).





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


Powers I
Powers I


Henry County B
cKe e Carroll County
Weakley County





Figure 2. A, Localities of clay pits in Kentucky, Tennessee, and Mississippi, Eastern United States. 1, Lamkin (UF 15815). 2,
South-40 (UF 15824). 3, Bell City (UF15803). 4, New Puryear (UF15819). 5, Puryear (UF15820). 6, Martin (UF15809. 7, Foundry
Hill (UF15810). 8, Buchanan(UF15806). 9, Warman(UF15826). 10, Gleason (UF15811). 11, Miller(UF15817). 12, Lawrence
(UF15816). 13,Willbank I &2(UF18884,UF18927). 14, Powers (UF18810). 15,Bovay(UF15737). 16,Bolden(UF15738). 17,
Old Hickory (UF15742). 18, Richies Black (UF15828). 19, New Lawrence (UF15818). 20, New Haynes (Rancho, UF 15921). B,
Locality of Powers Pit (#14), Weakley County, Tennessee.





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


Figure 3. Powers Pit. A, Northeast corer. B, East side of pit. Photographed by Hongshan Wang.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


WHITLATCH SCHNEIDER STEARNS HARDEMAN PARKS PARKS C)
AND AND ET AL. AND W
BLANKENSHIP ARMSTRONG CARMICHAEL
(surface) (subsurface) (subsurface) (surface) (surface) W

1940 1950 1955 1966 1975 1990 1U


JACKSON

FORMATION


GRENADA
FM.


HOLLY
SPRINGS
FM.


ACKERMAN
(?) FM.


PORTERS
CREEK
CLAY



iiZ


uj-II


JACKSON (?)

FORMATION


CLAIBORNE
GROUP


JACKSON
AND
CLAIBORNE

GROUPS


V 9


WILCOX

GROUP


PORTERS
CREEK
CLAY


CLAYTON
FM.


WILCOX

GROUP


UPPER


PORTERS
CREEK
CLAY


CLAYTON
FM.


JACKSON (?)

FORMATION


CLAIBORNE
AND
WILCOX
FMS.


PORTERS
CREEK
CLAY


CLAYTON
FM.


..A..


CLAIBORNE
FM.


WILCOX

FM.


PORTERS
CREEK
CLAY


CLAYTON
FM.


JACKSON

FM.


COCKFIELE
FM.

COOK

MOUNTAIN
FM.



MEMPHIS

SAND


WILCOX
GROUP


MIDWAY
GROUP


S. ..........


CRETACEOUS


Figure 4. Stratigraphic chart of western Tennessee. Modified from Parks (1975) and Parks and Carmichael (1990).


elected at Powers Clay Pit in Weakley County, Tennes-
see (Figs. 2, 3). This clay pit was selected because we
have accumulated, during the past 5 years, a collection
of over 400 specimens consisting mainly of leaves, fruits,
and seeds through the efforts of B. Roger Moore, the
authors, and numerous colleagues who have visited the


site. The well preserved fossil leaves, flowers, fruits,
seeds, and insect larval cases occur as compressions/
impressions, casts, and carbonized organic material.
Many of the fossils collected in Powers Clay Pit can be
related to the extant families Lauraceae, Fabaceae,
Fagaceae, Theaceae, and Araliaceae, with less abun-


.........





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


Figure 5. Transgression-regression schematic. From Potter and Dilcher 1980.


dant fossils related to Annonaceae, Moraceae,
Rhamnaceae, Smilacaceae, and Myrtaceae. A few rare
specimens are related to Platanaceae, Altingiaceae,
Salicaceae, Sapindaceae, Nyssaceae, Apocynaceae,
Rubiaceae, and Oleaceae. There are also some caddisfly
cases associated with the plant fossils.
E.W. Berry (1916, 1930, 1941) originally consid-
ered the plant fossils from numerous localities extending
from Texas, Arkansas, western Tennessee, western Ken-
tucky, Mississippi and Alabama to be of lower Eocene,
belonging to the Wilcox Formation. Later, the age of
this flora was clearly shown based on palynological cor-
relations to belong to the middle Eocene Claibome Group
(Elsik & Dilcher 1974; Frederiksen 1988). As well as
the age, the depositional settings of these plant-rich clay
deposits has been revised over the last 60 years. These
revisions will be discussed in the material and methods
section of this report.
Since the original reports of the Eocene floras from
southeastern North America (Berry 1916, 1930, 1941),
methods for the study of fossil plant remains have been
revised, and the philosophy of reporting on specific fos-
sil plants and floras has undergone extensive modifica-
tion (Hickey 1973; Dilcher 1974; LAWG 1999; Dilcher


2000a). The entire concept of presenting huge fossil
floras that demand only brief comparisons with living
plant genera are now understood to be filled with identi-
fication errors. As Dilcher (1974) mentioned, Berry's
identifications for the Eocene floras of southeastern North
America were about 60% incorrect or held some mar-
gin of systematic errors and thus Berry's conclusions
are often not supported. With this in mind, the flora
presented here is done so with tentative assignments to
extant genera and more certain assignments to extant
families as in Dilcher (2001). This is logical because
many extant families can be recognized and validated
by Eocene times, but fewer extant genera can be vali-
dated by that time. The use ofleafmorphotypes (Johnson
1989; Ellis et al. 2003; Barclay et al. 2003) is a practical
way to proceed in paleobotanical studies of fossil leaves
for which a modern generic affinity is uncertain. Sev-
eral angiosperm leaf morphotypes are presented in this
study.
This report represents the first attempt in over 75
years of presenting an entire known fossil flora based
upon megafossil evidence, from the Eocene sediments
of southeastern North America. It is limited to a single
locality, represents a change in age, presents new ideas







in the deposition of the flora, and a new philosophy in
our approach to this floristic study from previous floras
of this region by Berry (1916, 1930). The more recent
research of these fossil plants, limited to studies of spe-
cific taxa or specific families, is indicated below.

PREVIOUS PALEOBOTANICAL INVESTIGATIONS
There is a long and rich history of investigations of
plant megafossils collected in western Tennessee and
Kentucky, starting in the mid 1800's (Lesquereux 1859,
1869; Owen 1860; Safford 1869; Berry 1914,1916,1922,
1924, 1930, 1937, 1941). Brown(1939, 1940, 1944, 1946,
1960) provided limited taxonomic revisions of some of
the fossil material previously published by Berry. Sub-
sequently, systematic revisions and additions to the
Claibome megafossil flora can be found in Dilcher (1963,
1969, 1971,2000b), Dilcher & McQuade (1967), Dilcher
& Mehrotra (1969), Dilcher & Dolph (1970), Dilcher et
al. (1974, 2001), Crepet et al. (1974, 1975, 1980), Dolph
(1975), Crepet & Dilcher (1977), Dilcher & Daghlian
(1977), Crepet (1978), Roth & Dilcher (1979), Crepet
& Daghlian (1980, 1982), Jones & Dilcher (1980, 1988),
Potter & Dilcher (1980), Weiss (1980), Roth (1981),
Zavada & Crepet (1981), Jones (1984), Dilcher &
Manchester (1986, 1988), Jones et al. (1988), Taylor
(1988), Grote & Dilcher (1989, 1992), Sun (1989),
Herendeen & Dilcher (1990a,b,c, 1991), Herendeen et
al. (1990), Call & Dilcher (1992), Herendeen (1992),
Moore (2001), & Moore et al. (2003). Palynological
reports from western Tennessee can be found in
McLaughlin (1957), Tschudy (1973), Elsik & Dilcher
(1974), Potter (1976), & Taylor (1987, 1989) while dis-
persed cuticle of one locality was presented by Kovach
& Dilcher (1984). These publications since 1963 repre-
sent work on individual taxa or limited families common
in the Eocene floras of western Tennessee. The reason
for this systematic rather than floristic emphasis is given
in the discussion of this paper.

OBJECTIVES
This study is an attempt to document the megafossil
flora (with minor faunal elements included) of one spe-
cific clay pit in western Tennessee, by presenting each
type of megafossil plant found that is now in the Paleo-
botany and Palynology Collection of the Florida Museum
of Natural History. Leaf morphotypes are presented
and morphological descriptions of each taxon are corre-
lated with modern genera when some similarities are
noticed. As taxa can be related to a specific extant
family, extant genus, and extinct fossil plant morphotypes,
the overall floral composition of this fossil locality can
be compared with floras of modem ecosystems and


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

known fossil floras. The use of such comparisons in the
understanding of the paleoenvironment, paleoclimate,
stasis and change in floristic composition through time
will be presented in the discussion. As floras from other
localities are published, we expect that the plant species
composition of this locality will be used to compare and
contrast with species lists from other middle Eocene fossil
plant localities. A leaf morphology key is provided for
preliminary field identification for this one locality (Table
3). This will allow the leafmorphotypes presented here
to be used in field studies of plant fossil diversity, overall
species richness and abundance.

MATERIALS AND METHODS
The plant megafossils and insect larval cases presented
in this report were collected mostly in July 2000, May
2001 and April 2002 from the Powers Clay Pit, Weakley
County, Tennessee, 36' 16.94 N, 880 31.78 W, in sedi-
ments belonging to the middle Eocene, Claibome Group.
Specimens were also collected from a commercial shed
with piles of clay mined from Powers Clay Pit, at the
headquarters ofH. C. Spinks Clay Company in July 2002.
Terminology for leaf description follows Hickey (1973),
Radford et al. (1974), Dilcher (1974) and LAWG (1999).
The systematic classification used follows APG II (2003).
The fossil material is numbered with the University of
Florida acronym (UF) followed by the locality number
(18810 for Powers Clay Pit), then the individual speci-
men number(e.g. UF 18810-34381). Modem leaves cited
for comparison are numbered as the University of Florida
Modern Reference Leaf Collection followed by the
specimen number (e.g. UF5437). Herbarium specimens
of modem leaves were examined at the University of
Florida Herbarium (FLAS). Some specimens are fig-
ured but not described such as Entire Margin
Morphotype 3 (UF18810-34450) and Entire Margin
Morphotype 4 (UF 18810-34449).
Changing concepts of stratigraphic nomenclature
for the Claibome Group in western Tennessee since the
1940's are illustrated in Figure 4. The clay lenses in the
upper Claiborne Group were considered to represent
near-marine paleoenvironments because of the possible
coastal elements, such as sabal palms in the fossil floras
(Berry 1916, 1930), sedimentary environment types
(Steams 1957), and the presence of hystrichospheres
and dinoflagellates (Parks 1975). Dilcher (1974) sug-
gested that the clay lens of some of the plant-rich clay
pits represented oxbow lake deposits. Dolph (1975) also
envisioned back-swamp paleoenvironments on ancient
floodplains while Potter (1976) also suggested abandoned
river channels. Potter & Dilcher (1980) presented nu-
merous detailed outlines of several clay pits with some





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


Table 1. Systematic list of the flora and fauna with exemplary specimens. Indicates an extinct fossil genus or
unknown modem genera similar to this fossil.


Magnoliids


Monocots


Eudicots
Core Eudicots
Rosids
Eurosids I

















Eurosids II
Asterids


Order
Laurales
Magnoliales
Liliales


Proteales
Saxifragales
Myrtales
Fabales




Fagales




Malpighiales
Rosales





Sapindales
Comales
Ericales


Euasterids I Gentianales


Euasterids II Apiales
Lamiales
Incertae sedis










Insect cases


Family Species
Lauraceae Ocotea sp.
Annonaceae Duguetia sp.
Smilacaceae Smilar sp. 1 & 2
Undetermined Monocot* sp.
Platanaceae Platanus* sp.
Altingiaceae Liquidambar sp.
Myrtaceae Myrcia sp.
Fabaceae Cladrastis-like sp.
Ormosia-like sp. 1 & 2
Swartzia sp.
Fagaceae Berryophyllum* tenuifolia (Berry) Jones et Dilcher
Berrvophyllum* sp.
Knightiophyllum* wilcoxianum Berry
Salicaceae Populus sp.
Moraceae Pseudolmedia sp.
Ficus-like sp.
Rhamnaceae Berhamniphyllum* claibornense Jones et Dilcher
Berhamniphyllum* sp.
Sapindaceae Cupanites sp.
Nyssaceae Nyssa eolignitica* Berry
Theaceae Ternstroemites* sp.
cf Gordonia sp.
Apocynaceae Apocynophyllum* sp.
Rubiaceae cf Paleorubiaceophyllum* sp.
Araliaceae Dendropanax eocenensis Dilcher et Dolph
Oleaceae Oleaceae leafmorphotype
Entire margin morphotype 1 *
Entire margin morphotype 2*
Entire margin morphotype 3*
Entire margin morphotype 4*
Entire margin morphotype 5*
Tooth margin morphotype *
Reproductive structure morphotype 1*
Folindusia sp.
Terrindusia sp.







cross sections of their shapes, and the lithology of the
clay lenses. They concluded that they conformed to
clay plugs typical of oxbow lakes in an ancient middle
Eocene river systems, or braided stream distributary
migrating across their flood plains. Taking into account
multiple overlapping clay lenses surrounded by cross-
bedded sands with frequent clay rip-up, noted in two
clay pits, Moore et al. (2003) suggested clay plugs of
oxbow lakes in a braided river system in the ancient
flood plain. Regional and local pollen data (Engelhardt
1964; Tschudy 1973; Elsik 1974; Elsik & Dilcher 1974;
Potter 1976), and repeated marine transgression/regres-
sions in southwestern Tennessee (Stears 1957; Murray
1961) (Fig. 5) indicate that the clay lenses are corre-
lated with the mainly nonmarine (Fisher 1964;
Frederiksen 1988) Cockfield Formation of the Claibome
Group (Potter & Dilcher 1980).

SYSTEMATIC COMPOSITION
The taxa we have recognized are listed in Table I
and organized according to the APG system of classifi-
cation.

SYSTEMATIC PALEONTOLOGY
Laurales
Lauraceae
Ocotea sp.
(Fig. 12, C-E, G)
Description.-Majority of leaves incomplete,
lamina linear to narrowly elliptic, symmetrical, 8.5-
14.7 cm long by 0.9-3.1 cm wide, length/width ratio 4.7:1
- 9.4:1; apex angle acute, with cuneate flanks; base angle
acute, with cuneate to slightly concave flanks. Margin
entire and thickened or with a marginal vein. Petiole
stout, 1.2 cm long, widening gradually towards lamina.
Venation pinnate; secondary venation weak
brochidodromous with 6-9 pairs, angle of divergence 20-
40. One pair of acrodromous basal to suprabasal sec-
ondary veins extending 1/3 1/2 distance of leaf length.
Tertiary veins reticulate, with a continuous row of ter-
tiary looping along the marginal side of the upturned sec-
ondary veins.
Discussion.-The general venation patterns are
similar to those found in the families Lauraceae and
Monimiaceae, although in Monimiaceae the orientation
of tertiary veins in the upper 1/3 of the leaf is toward the
midvein (Klucking 1987). Within Lauraceae, the num-
ber of secondary vein pairs, acrodromous venation, and
tertiary looping are similar to Oreodaphne
pseudoquianensis Berry (Ocotea pseudoquianensis
[Berry] LaMotte), 0. obtusifolia Berry (Ocotea
obtusifolia [Berry] LaMotte; Berry 1916; Dilcher 1963),


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

and Ocotea species of Klucking (1987). In specimen
34410a, the shape, and suprabasal acrodromous vena-
tion is similar to 0. quianensis Aublet (Klucking 1987).
The venation matches the course of the acrodromous
venation, such as one vein extends 1.5x further than the
other. In specimen 34409, the venation pattern is close
to 0. fasciculata (Nees.) Mez. (Klucking 1987).
Ocotea pollen is also recorded from the upper Claiborne
of Alabama (Gray 1960). The isolated leaves of
Lauraceae are well placed within the family based upon
venation and form, however they are notoriously diffi-
cult to place within a genus and one can do little more
than picture match unless some unique features of ve-
nation and cuticle are present. In this leaf the strong
paired basal veins provide a distinct character that can
be identified from pit to pit and also holds close similari-
ties to a leaf from the middle Eocene Messel pit in Ger-
many (Sturm 1971). The leaves from Messel are gen-
erally similar to those illustrated by Dilcher (1963) from
other clay pits, including the nature of the cuticle but
with the exception of the abundance and placement of
the trichome bases along the anticlinal cell walls. In the
future, cuticle preparations of the fossils discussed here
from Powers Pit may also have similarities in the nature
of the cuticle. The leaves illustrated in Fig. 12 C and G
may represent one species while the specimen illustrated
in Fig. 12 D, E is certainly a different species. Cuticular
features will be important to distinguish them.
Material examined.-UF18810-34381, 34385a,
34409,34410a,34411.

Magnoliales
Annonaceae
Duguetia sp.
(Fig. 10, D, E; Fig. 14, C; Fig. 16, A)
Description.-Leaf incomplete, lamina elliptic,
symmetrical, -10-17 cm long by 4.2-9 cm wide, length/
width ratio 1.9:1 2.4:1; apex missing; base obtuse;
margin entire. Venation pinnate; secondary venation
brochidodromous, -14 pairs of secondary veins, angle
of divergence 50-80o, secondary veins mostly alternate,
with basal venation sigmoidal, and mid to upper venation
arching slightly or straight, from the midvein to near the
margin then arching sharply upward joining
superadjacent vein. Up to two strong intersecondary
veins in each intercosta, sometimes flanked by weaker
and shorter intersecondary veins. Tertiary veins
percurrent to lineate, sometimes forked and at right
angles to secondary veins. Excostal veins uniseriate to
biseriate, close to margin. Ultimate venation open.
Discussion.-The characters of shape, margin, and
narrowly spaced secondary veins are characteristic in




DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

the families Annonaceae and Magnoliaceae. The dif-
ference is in the angle of the secondary veins, and ulti-
mate venation. In Magnoliaceae, the secondary veins
have moderate angles (<550) and the ultimate venation
is closed, while in Annonaceae and the Powers Pit speci-
mens, the secondary veins are high angled (>550) and
the ultimate venation is open (Klucking 1986). Speci-
men UF34418 is similar to Magnolia leei Knowlton
(Berry 1916, 1922, 1930, 1941) and Duguetia leei
(Knowlton) Roth et Dilcher (Roth 1981) in the nature of
the secondary venation and the percurrent tertiary veins.
Specimen UF34407 is similar to D. argentea Fries in
secondary and excostal venation, and in that the tertiary
veins are lineate. Although the genus was identified
from other Claiborne clay pits using characters of epi-
dermal anatomy (Roth 1981), the specimens from Pow-
ers Pit were not investigated anatomically. The extant
species Duguetia is distributed in the Neotropics (Roth
1981; Dilcher 2000b).
Material examined.-UF 18810-34407, 34418.

Liliales
Smilacaceae
Smilax sp. 1
(Fig. 13, C, D)
Specimen description.-Lamina ovate, symmetri-
cal, 6.5 cm long by 3.6 cm wide; apex missing; base
angle obtuse, with concave flanks; margin entire. Peti-
ole missing. Venation basal acrodromous, of three pri-
mary veins, two secondary veins basal acrodromous.
Tertiary veins mixed opposite, alternate percurrent, ter-
tiary vein angle acute.
Discussion.-The general shape, size, margin, and
prominent basal acrodromous venation is similar to
Smilacaceae, Rhamnaceae, and Dioscoreaceae. In
Rhamnaceae, such as Ziziphus cinnamomum Triana &
Planch. (Heald 2002), there are three prominent basal
acrodromous veins, but they lack the outer less promi-
nent basal acrodromous veins present in Smilacaceae
(Conran 1998). In Dioscoreaceae, such as Dioscorea
decipiens J. D. Hooker (Zhizun & Gilbert 2000) and D.
bulbifera L. (UF6173), the tertiary venation is distinctly
opposite percurrent. The presence of five prominent basal
acrodromous veins, and tertiary veins of mixed opposite
and alternate percurrent is characteristic of Smilacaceae,
and is similar to Berry's Smilax wilcoxensis (1930) of
the Wilcox Flora, but the basal characters are different
from our specimen. Berry's type material is incomplete,
but diagramed to show that the secondary veins arise
from the midvein independently. This is not the case
with Smilax sp.1, as a pair of secondary veins arise from
the very base of the midvein and then each bifurcates


into the lateral acrodromous veins. If this character is
demonstrated to be one of species importance, and if
the details of Berry's S. wilcoxensis can be determined
from other material, then we can ascertain whether these
are conspecific. Smilax cuticle is known to differ be-
tween similar leaves from different localities so that spe-
cies differences sometimes can be determined only by
cuticular anatomy (Sun & Dilcher 1988).
Material examined.-UF 18810-34415.

Smilax sp. 2
(Fig. 13, E; Fig. 14,A)
Specimen description.-Lamina ovate-cordate,
symmetrical, 8.5 cm long by 6.2 cm basal width; apex
missing; base angle obtuse, with cordate flanks; margin
entire. Petiole missing. Venation campylodromous, with
three primary veins, two secondary veins basal
acrodromous. Tertiary venation barely discernable.
Discussion.-The conspicuous characters of Smi-
lax sp. 2, such as basal acrodromous venation with a
cordate leaf base and entire margin, can be found in
Smilacaceae and Dioscoreaceae, although in
Dioscoreaceae they can have up to 11 primaries and the
tertiary venation is distinctly opposite percurrent (Zhizun
& Gilbert 2000; Raz 2002). Although the leaf shape of
extant Smilax bona-nox L. is highly variable (Godfrey
1988), some forms with a cordate base are similar to
our specimen. There are about 300 extant species of
Smilax worldwide. Many species are common in south-
eastern North America extending into Mexico and Cen-
tral America.
Material examined.-UF 18810-34435

Undetermined Monocot sp.
(Fig. 19, B)
Specimen description.-Leaf blade fragment, 1.3
cm wide. One strong midvein and one strong parallel
marginal vein, otherwise other parallel veins barely
discernable. No cross venation evident.
Discussion.-Lack of definitive characters makes
further determination of this specimen impossible at this
time. If cross veins were preserved, this character might
be attributed to the palms. The lack of characters is not
due to poor preservation alone, as a palm leaf fragment
would normally show parallel secondary veins and the
typical cross veins.
Material examined.-UF 18810-34400

Proteales
Platanaceae
Platanus sp.
(Fig. 7, A-C)







Specimen description.-Leaf incomplete, palmately
lobed, appears to be 3-lobed; apex missing; base angle
obtuse, with rounded flanks. Leaf margin occasionally
serrate, teeth straight to curved, sinuses rounded. Peti-
ole missing. Venation palinactinodromous; secondary
venation camptodromous, angle of divergence 35-40.
Discussion.-The lobed nature of the leaf is simi-
lar to Lauraceae (Sassafras), Altingiaceae (Liquidam-
bar), and Platanaceae (Platanus), although in Sassa-
fras and Liquidambar the primary venation is
suprabasally actinodromous, and in Liquidambar the
margins are continuously serrate, and with basal lobing
(Schwarzwalder 1986). General characters such as
lobes, venation, and occasional teeth are similar to
Platanus sp., although Platanaceae venation is some-
times superbasally actinodromous. The secondary veins
are not sufficiently preserved near the margins to deter-
mine if they extend into the teeth ("Platanoid") or not,
but the slope does suggest that they curve away from
the teeth as in Platanus kerrii Gagnep. (Schwarzwalder
1986). This is the first occurrence of a lobed Platanus
leaf in the Claiborne flora. Pollen of Platanus is re-
corded from the upper Claiborne of the Mississippi
Embayment (Frederiksen 1980a, 1988).
Material examined.-UF18810-34439.

Saxifragales
Altingiaceae
Liquidambar sp.
(Fig. 18, D, H)
Specimen description.-Fruit laterally compressed,
subglobose, 2 cm diameter. Convoluted carbonaceous
material with scattered circular impressions of style bases
with, or without acute impressions of style tips.
Discussion.-Impressions in the fruit appear simi-
lar to Liquidambar styraciflua L. before dehiscence
of the basal portion of the persistent styles (Godfrey
1988). Liquidambar fruit have been recorded from the
Claiborne flora but were not described or figured (Berry
1930). Pollen of Liquidambar is recorded from the
upper Claiborne of Alabama (Gray 1960).
Material examined.-UF 18810-34432.

Myrtales
Myrtaceae
Myrcia sp.
(Fig. 10, F-H)
Specimen description.-Leaf incomplete, lamina
possibly elliptic, 7 cm long by 3.2 cm wide; apex and
base missing; margin entire. Petiole missing. Midvein
stout; venation pinnate; secondary veins narrowly spaced,
angle of divergence 70-80o, straight from midvein to


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

intramarginal vein. Some intersecondary veins present,
a few admedially branched. Leaf margins with paired
intramarginal veins, one running along the margin, the
second 1-2 mm inside the leaf margin and slightly scal-
loped. Tertiary veins sometimes joining secondary veins
at acute angles.
Discussion.-The spacing and angle of the sec-
ondary veins, along with the intramarginal venation is
similar to Qualea in Vochysiaceae, and Calyptranthes,
Syzygium and Myrcia in Myrtaceae (Klucking 1988).
The difference is in the intersecondary veins, in
Vochysiaceae they branch toward the margin, while in
Myrtaceae some are admedial, branching towards the
midvein. Also, in Vochysiaceae, the secondary veins
curve toward the margin (Sajo & Rudall 2002). Vena-
tion is similar to Myrcia and Syzygium, but in Syzygium
the secondary veins are recurved to sigmoidal (Klucking
1988). Leaf size, along with secondary and intramarginal
venation is similar to Calyptranthes eocenica Berry
(Berry 1916), but in Calyptranthes, both intramarginal
vein pairs run 1-2 mm inside the leaf margin (Klucking
1988). Pollen of Myrtaceae (Myrtaceidites) is recorded
from the upper Claiborne of Mississippi and Alabama
(Gray 1960; Engelhardt 1964; Frederiksen 1980a, 1988).
Material examined.-UF 18810-34386.

Fabales
Fabaceae
Cladrastis-like sp.
(Fig. 10, A-C)
Description.-Lamina of leaflet lanceolate, sym-
metrical to slightly asymmetrical, 6.5-9.5 cm long by 2-
2.3 cm wide, length/width ratio 3.3:1 -4.1:1; apex angle
acute; base angle obtuse, with rounded flanks; margin
entire. Petiolule 3 mm long, with cross striations. Vena-
tion pinnate; secondary venation brochidodromous, 9-10
pairs of secondary veins, angle of divergence 30-60,
secondary veins straight or arching from midvein to near
margin then arching up to superadjacent vein. One to
two intersecondary veins per intercosta. Midvein deeply
impressed. Tertiary veins orthogonal. Excostal vena-
tion in 1-2 series.
Discussion.-The size, shape, apex, and venation
are similar to Sapindus (Sapindaceae) and Cladrastis
(Fabaceae) but in Sapindus the base is cuneate, curved,
and asymmetrical (UF335, 340, 343,345,352; Herendeen
1992; Yu & Chen 1991). The venation and size of the
smaller leaflets are similar to Gleditsiophyllum
eocenicum Berry (Berry 1916; Herendeen 1992). Ex-
tant species of Cladrastis have a temperate distribution
(Herendeen 1992).
Material examined.-UF 18810-34402, 34413a,





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

34421,34428.

Ormosia-like sp. 1
(Fig. 11,A-C)
Description.-Lamina of leaflet elliptic, symmetri-
cal, 11 cm long by 4.7-5.6 cm wide; apex angle acute,
with acuminate flanks; base angle acute, with convex
flanks; margin entire. Petiolule missing. Venation pin-
nate; secondary venation weak brochidodromous, 6-8
pairs of secondary veins, angle of divergence 40-60,
secondary veins alternate, broadly arching from the
midvein to near the margin then arching sharply and di-
rected medially toward midvein joining superadjacent
vein. Weak basal secondary veins, and 1-3 weak to
strong intersecondary veins per intercosta. Tertiary veins
percurrent, straight to slightly sinuous, unbranched to
branched, tertiary angle obtuse.
Discussion.-The absence of a petiolule makes
the determination to family difficult because the leaflets
of legumes frequently have short, wrinkled appearing
petiolules. The general shape, size, apex, margin, and
venation of these specimens suggest leaflets in the fami-
lies Fabaceae and Connaraceae. These two families
seem very close in the morphological characters of their
leaflets, although in Connaraceae the tertiary venation
is generally more prominent as in Connarus
fasciculatus (de Candolle) Planchon (Forero 1983) and
Rourea minor (Gaertn.) Alston (Jongking 1989). Within
Fabaceae, this fossil leaflet morphology and venation
are similar to Swartzia racemosa Benth. (FLAS 165336)
and Ormosia grandiflora (Tul.) Rudd although in S.
racemosa the margins have a few teeth. Further study
of the anatomy would help to clarify the generic and
species determination of these fossils. At present the
leaflets appear most similar to Ormosia sp. 1 (Herendeen
1992; Fig. 119) and so are placed in that genus.
Material examined.-UF 18810-34404a, 34406.

Ormosia-like sp. 2
(Fig. 7, I, K)
Specimen description.-Leaflet incomplete, lamina
narrowly elliptic, symmetrical, 6 cm long by 1.4 cm
wide, length/width ratio 4.3:1; apex missing; base angle
acute, with slightly convex flanks; margin entire. Peti-
olule prominently cross-striated, 0.8 mm long by 0.15
mm wide. Venation pinnate; secondary venation
brochidodromous, angle of divergence 30-40o, second-
ary veins alternate, extending straight from midvein to
margin or curving abruptly upward near the margin. One
pair of weak acute basal secondary veins, and 1-2
intersecondary veins per intercosta. Tertiary veins
percurrent. Quaternary veins random and reticulate.


Discussion.-Leaflet petiolules with prominent
striations are found in Fabaceae and Connaraceae, but
the shape of this leaflet is similar to extant Ormosia
(Fabaceae) such as Ormosia nitida Vog. and 0.
monosperma (Sw.) Urb. (Rudd 1965). The shape, base,
margin, and venation is similar to fossil Ormosia sp. 2
(Herendeen 1992). Extant species of Ormosia can be
found in the tropics of Asia, Australia, Madagascar, and
America (Herendeen 1992).
Material examined.-UF 18810-34395.

Swartzia sp.
(Fig. 7, D-H, J, L; Fig. 13, A, B)
Description.-Lamina of leaflet narrowly elliptic
to elliptic, symmetrical, 3.7-8 cm long by 1.3-4 cm wide,
length/width ratio 2:1 2.8:1; apex angle acute to ob-
tuse, retuse; base angle acute to obtuse, with convex to
concave flanks; margin entire. Petiolule missing. Ve-
nation pinnate; secondary venation strongly
brochidodromous, angle of divergence 50-60, second-
ary veins alternate, with 5-6 pairs, straight from midvein
to near margin, then arching upwards. One pair of weak
acute basal secondary veins, and 1-2 strong
intersecondary veins per intercosta. Tertiary veins or-
thogonal reticulate. Associated pod incomplete (UF
34419a, not figured), apex missing, base curving with
basal portion missing. Parallel oblique striations near
base, marginal suture well developed.
Discussion.-Although the morphology and vena-
tion are similar to numerous species of Swartzia
(Fabaceae) such as Swartzia parvifolia Schery, S.
apetala Raddi, and S. corrugata Bentham (Cowan
1968), future anatomical work is needed to confirm this
determination. The shape, size, margin, apex, base, and
venation of the leaflets, and pod shape and size are simi-
lar to the fossil species Swartzia sp. 1 (Herendeen 1992),
although the leaflet determination was based mainly on
anatomical characters. Fabaceous pollen is recorded in
the upper Claiborne of the Mississippi Embayment
(Frederiksen 1980a). Extant species of Swartzia are
distributed in the Neotropics and Africa (Herendeen
1992).
Material examined.-UF 18810-34388a, 34391,
34393, 34394,34419a, 34420.

Fagales
Fagaceae
Berryophyllum tenuifolia (Berry) Jones et
Dilcher
(Fig. 17, H; Fig. 18, B)
Specimen description.-Leaf incomplete, lamina
linear, symmetrical, 9 cm long by 0.4 cm wide at







middle; apex missing; base angle acute, with cuneate
flanks; margin serrate for most of leaf length. Petiole
missing. Venation pinnate; secondary venation
craspedodromous, angle of divergence 400, secondary
veins alternate, straight to leaf margin, one entering the
tooth, the adjacent vein running alongside leaf margin.
Discussion.-The shape, size, margin, and vena-
tion are similar to leaves of Fagaceae, and the fossil
species Banksia tenuifolia proposed by Berry which
was renamed Berryophyllum tenuifolia (Berry) Jones
et Dilcher (Berry 1916; Jones & Dilcher 1988).
Fagaceous pollen (Quercoidities, Cupuliferoi-
pollenites) is recorded in the upper Claiborne of the
Mississippi Embayment (Frederiksen 1980a, 1988) and
from the Lawrence (Quercus) and Warman
(Quercoidities) clay pits (Elsik & Dilcher 1974; Potter
1976).
Material examined.-UF18810-34389.

Berryophyllum sp.
(Fig. 17, E-G)
Description.-Leaves incomplete, lamina narrowly
elliptic, symmetrical, 20 cm long by 2 cm wide at middle;
apex and base missing; upper 25% of leaf attenuate,
lower 25% of leaf cuneate. Margin serrate in upper
75% of leaf; teeth sharp, prominent, 1 mm long, basal
side straight to convex, apical side concave. Petiole
missing. Venation pinnate; basal secondary venation
camptodromous, secondary venation craspedodromous
in mid and upper portions of leaf, angle of divergence
50-60, secondary veins alternate, arching gradually from
midrib to margin, one vein entering each tooth, adjacent
1-2 veins running parallel to margin. Secondary veins
forked just below tooth, of equal strength, one entering
tooth medially and the other running parallel to margin.
One to two intersecondary veins per intercosta. Ter-
tiary veins percurrent, simple to branched. Quaternary
veins random.
Discussion.-The leaf characters seem to be in-
termediate between Berryophyllum warmanense Jones
et Dilcher and B. sqffordii (Lesq.) Jones et Dilcher.
The tooth shape and the way the secondary veins enter
the teeth are characters that match B. warmanense,
while the leaf size, margin, and secondary venation match
B. saffordii (Jones & Dilcher 1988). It has been sug-
gested that Trigonobalanus and Colombobalanus may
be related extant genera that are found in tropical Asia,
and the mountains of Colombia (Dilcher 2000b).
Material examined.-UF 18810-34396, 34408a.

Knightiophyllum wilcoxianum Berry
(Fig. 16, B, C, G)


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

Description.-Lamina narrowly elliptic, symmetri-
cal, -11-13 cm long by 1.8-2.5 cm wide; apex and base
angle acute, with cuneate flanks. Margin serrate and
confined to the middle and distal half of the leaf; teeth
short, less than 1 mm, basal flank convex, apical flank
concave. Petiole missing. Venation pinnate; secondary
venation camptodromous to semicraspedodromous, angle
of divergence 40-60o, secondary veins alternate, arch-
ing gradually from midvein to near margin then joining
the superadjacent vein. One to two intersecondary veins
per intercosta. Tertiary veins orthogonal, tertiary veins
arising from a secondary marginal loop enter the teeth.
Ultimate veins freely ending and one branched.
Discussion.-The shape, size, apex, base, margin,
and venation are similar to Knightiophyllum
wilcoxianum Berry (Berry 1916; Dilcher & Mehrotra
1969) except in this material the teeth are only in the
distal half of the lamina. The systematic placement of
this genus appears to be with Fagaceae but no modem
genera have been found that are similar to K.
wilcoxianum (Dilcher & Mehrotra 1969). Morphologi-
cal features such as leaf margin with a few teeth, gen-
eral looping of the secondary veins, tertiary vein from
secondary looping entering teeth, and intersecondary
veins common, are characteristic of Fagaceae, notably
Quercus section Erythrobalanus (Jones 1984), near
Quercus viminea but the teeth are not as prominent as
in Q. viminea (Villarreal 1986).
Material examined.-UF 18810-34424, 34425a

Malpighiales
Salicaceae
Populus sp.
(Fig. 16, D; Fig. 17, A)
Specimen description.-Leaf incomplete, lamina
elliptic, symmetrical, 6 cm long by 3 cm wide; apex
and base missing. Margin serrate/dentate; teeth apex
obtuse (they appear to be glandular) to acute, sinus
rounded. Petiole missing. Venation pinnate; secondary
venation semicraspedodromous, with one
subactinodromous vein, angle of divergence 60 basally,
40-500 apically. Intersecondary veins per intercosta one
or absent. Tertiary veins not well preserved.
Discussion.-Although the leaf is incomplete, and
tertiary veins are not well preserved, characters such as
shape, subactinodromous and semicraspedodromous
venation, and toothed margins are similar to extant leaves
of Populus (Salicaceae; Eckenwalder 1980) and
"Tiliaceous" species Dicraspidia donnell-smithii
Standley (Robyns 1964) and young seedlings of Tilia
americana L. (UF6174). In D. donnell-smithii and T
americana, the secondary venation is mostly





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

craspedodromous, the teeth lack glands, and the sinus
of the teeth are angular. The shape of some forms of
Cercidiphyllum japonicum Siebold & Zucc. with
toothed margins, and teeth with a rounded sinus is simi-
lar to the fossil, but in C. japonicum the secondary ve-
nation is brochidodromous (UF1965). The presence of
glandular teeth is characteristic of Populus. The vena-
tion and teeth are similar to middle/upper portions of
Grewiopsis tennesseensis Berry (1916) which is con-
sidered similar to Populus cinnamomoides (Lesq.)
MacGinitie (Eckenwalder 1977, 1980). The fossil de-
scribed here may be an intermediate form between a
linear juvenile and deltoid adult leaf of Populus
(Eckenwalder 1980).
Material examined.-UF 18810-34397.

Rosales
Moraceae
Pseudolmedia sp.
(Fig. 14, B, D)
Description.-Lamina obovate, slightly
inequilateral, -14 cm long by 6.3 cm wide; apex angle
acute (not illustrated, UF 34425b); base angle acute, with
asymmetrical flanks; margin entire. Petiole missing.
Venation pinnate; secondary venation brochidodromous,
- 14 pairs of secondary veins present, angle of diver-
gence 40-50o, secondary veins alternate, straight from
midvein to near margin then arch sharply upward joining
superadjacent veins. Basal pair of secondary veins,
occasional 1-2 intersecondary veins per intercosta usu-
ally extending less than 2 the distance from midrib to
margin. Tertiary veins alternate percurrent and forked,
at right angles to secondary veins. Excostal veins
uniseriate, close to leaf margin.
Discussion.-The large size, margin, and the basal
pair of secondary veins are typical characters of
Moraceae. The combination of characters such as shape
of the apex and base, margin, number and angle of sec-
ondary vein pairs, intersecondary veins, and percurrent
tertiary veins are common in Pseudolmedia. These
characters also can be found in Naucleopsis but in
Naucleopsis the percurrent tertiary veins are in a dis-
tinct double layer in the intercostal area (Berg 1972;
FLAS 132816; UF1483). Berry (1916) recognized
Pseudolmedia in the Wilcox flora (now Claibome Flora)
of Tennessee but the leaves he described are narrower
and the secondary veins arise from the midvein at a 900
angle. The extant genus Pseudolmedia is distributed in
the Neotropics (Berg 1972; Burger 1977).
Material examined.-UF 18810-34417, 34425b.


Ficus-like sp.
(Fig. 8, C, D)
Specimen description.-Lamina elliptic, symmetri-
cal, 9.5 cm long by 3.7 cm wide; apex angle obtuse, with
convex flanks; base angle acute, with concave flanks;
margin entire. Petiole missing. Venation pinnate; sec-
ondary venation brochidodromous, 14 pairs of second-
ary veins, angle of divergence basally 60-70, apically
40-50', secondary veins subopposite basally to alternate
apically, straight from midvein to near margin, then arch-
ing upward parallel to margin, then arching mediallyjoin-
ing superadjacent vein, also a prominent secondary loop
above this lower loop. One pair of low angled basal
secondary veins present, 1-3 strong to weak
intersecondary veins per intercosta, extending to less
than distancee to margin. Excostal veins in 2-3 series.
Tertiary veins reticulate.
Discussion.-The general shape and size, apex,
base, margin, brochidodromous venation with high angled
basal secondary veins, 1-3 strong and weak
intersecondary veins, and reticulate tertiary veins are
characteristic to the families Moraceae and
Lecythidaceae, although in Lecythidaceae the
intersecondary veins can extend to more than V leaf
distance from midrib to margin, and they lack a pair of
basal low angle secondary veins. In Moraceae, the low-
ermost basal secondary vein angle is different from other
basal secondary veins, and a prominent secondary loop
over the junction of the secondary vein with the
superadjacent vein is evident in many species. Leaf
shape, size, base shape, and venation of this specimen is
similar to the fossil Ficus puryearensis Berry (1916)
and extant Ficus (Burger 1977), although in Berry's
description the secondary venation angle is nearly 90
while the figures (Plate XXVIII) are closer to our mea-
surements. The secondary vein angle is in the low range
for extant Ficus. Ficus pollen has been recorded in the
upper Claibome of Alabama (Gray 1960). Berry's (1916,
1930) floras included an excessive number of species in
the genus. A detailed re-evaluation of the genus is needed
in order to evaluate the real nature of all the so-called
Ficus leaves from the Claiborne of southeastern North
America.
Material examined.-UF 18810-34446.

Rhamnaceae
Berhamniphyllum claibornense Jones et
Dilcher
(Fig. 12, A, B)
Specimen description.-Leaf incomplete, lamina
narrowly elliptic, symmetrical, 11 cm long by 4 cm
wide; apex angle acute, with cuneate flanks; base miss-







ing; margin entire. Petiole missing. Midvein straight;
venation pinnate; secondary venation eucamptodromous,
angle of divergence 20-30o, secondary veins opposite
basally to alternate apically, straight to slightly curved
from midvein to near margin then arching abruptly up-
ward to the margin. Secondary veins rarely divided near
the margin. Tertiary venation barely perceptible,
percurrent, straight, perpendicular to the midrib.
Discussion.-Leaf venation characters such as
secondary venation eucamptodromous and percurrent
tertiary veins perpendicular to the midrib, are character-
istic of Rhamnaceae. The general shape, apex, margin,
and venation are similar to the fossil leaves
Berhamniphyllum claibornense Jones et Dilcher (Jones
& Dilcher 1980). Berhamniphyllum claibornense is
closely related to the tribe Rhamneae (Richardson et al.
2000), and possibly the extant genera Berchemia and
Rhamnidium (Jones & Dilcher 1980) that have prima-
rily tropical distributions (Record 1939; Brizicky 1964).
The fossil leaves have frequently been reported as Rham-
nus however identifications to extant genera can not be
validated without fruiting material, even if perfectly pre-
served leaves complete with cuticular material (even
extant leaves) are preserved. A new genus was pro-
posed by Jones and Dilcher (1980) to indicate that only
the family Rhamnaceae and tribe Rhamneae can be
determined based upon leaf material.
Material examined.-UF 18810-34384.

Berhamniphyllum sp.
(Fig. 9, D, G, H)
Specimen description.-Lamina elliptic, symmetri-
cal, 6.3 cm long by 3 cm wide; apex angle acute, retuse;
base angle acute, with convex flanks; margin entire.
Petiole missing. Venation pinnate; secondary venation
eucamptodromous, angle of divergence 40, basal sec-
ondary veins opposite, becoming alternate towards the
apex, secondary veins straight to slightly curved as they
extend from the midvein then arch abruptly upward to
near the leaf margin.
Discussion.-Although the lack of tertiary vena-
tion creates difficulty in the identification of this fossil,
the general shape and size, base, margin, and secondary
venation is very similar to Berhamniphyllum
claibornense Jones et Dilcher (Jones & Dilcher 1980).
The apex is retuse, which is not in the species descrip-
tion of B. claibornense (Jones & Dilcher 1980), but is
found in species of Rhamnidium such as R. ellipticum
Britton et P. Wilson and R. shaferi Britton et P. Wilson
(Britton 1915).
Material examined.-UF 18810-34427.


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

Sapindales
Sapindaceae
Cupanites sp.
(Fig. 17, B, D)
Specimen description.-Leaflet incomplete, lamina
narrowly oblong, asymmetrical, 7 cm long by 2.1 cm
wide; apex missing; base angle acute, with convex and
asymmetrical flanks. Margin with a few serrate teeth
from middle to apical portion of leaf. Petiole 1 mm
long and expanded at base. Midvein stout; venation pin-
nate; secondary venation semicraspedodromous, angle
of divergence 50-60, secondary veins alternate to sub-
opposite, straight from midvein to near margin, then di-
vided, one continuing toward margin and teeth, one arch-
ing upward joining superadjacent vein. One to two
intersecondary veins per intercosta. Tertiary veins at
right angle to secondary veins. Excostal veins in one
series.
Discussion.-The general shape and size, margin
with serrate teeth, asymmetrical base, and venation is
similar to leaflets of Sapindaceae and Juglandaceae, al-
though in Juglandaceae the secondary vein divides just
before entry into the tooth, one entering the tooth, the
other going towards the sinus. The leaflet is similar to
the fossil species Cupanites (Berry 1916, 1930) and
extant Cupania glabra (Standley & Steyermark 1949;
UF310), although in Berry's Cupanites, the venation is
craspedodromous and the teeth extend along the full
length of the leaf. Cupania-like pollen (Cupanieidites)
and megafossils are recorded in the upper Claibome of
Mississippi and Alabama (Gray 1960; Engelhardt 1964;
Frederiksen 1980a, 1988). Distribution of extant
Cupania is Neotropical (Standley & Steyermark 1949).
Material examined.-UF 18810-34401.

Cornales
Nyssaceae
Nyssa eolignitica Berry
(Fig. 18, G, K)
Specimen description.-Endocarp ovate, 2 cm long
by 0.8 cm wide, apex acute, base rounded. Fruit trilocu-
lar with numerous longitudinal ridges on the surface.
Discussion.-This fruit is similar to Nyssa
eolignitica Berry (Berry 1916; Dilcher & McQuade
1967) and N. wilcoxiana Berry (1916, 1930). Nyssa-
like pollen (Nyssapollenites) is recorded in the upper
Claiborne of Mississippi and Alabama (Gray 1960;
Engelhardt 1964; Frederiksen 1980a). Distribution of
the extant genus Nyssa is temperate, cool/warm tem-
perate of the United States and China to the tropics of
Mexico/Central America, East Asia and Malaysia/Indo-
nesia (Dilcher 2000b).





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

Material examined.-UF 18810-34392

Ericales
Theaceae
Ternstroemites sp.
(Fig. 16, E, F; Fig. 17, C, I; Fig. 18, A, E)
Description.-Lamina mostly elliptic to narrowly
elliptic, sometimes obovate, symmetrical, 5.8-14.5 cm
long by 3.2-3.3 cm wide, length/width ratio 2.6:1 -4.4:1;
apex angle acute to obtuse, with emarginate to acumi-
nate flanks; base angle acute, with cuneate to decurrent
flanks. Margin crenulate to serrate to near the base;
some teeth with glands (UF34405). Petiole 2 cm long
and expanded at base. Midvein stout, 1 mm wide at
middle section of leaf. Venation pinnate; secondary ve-
nation camptodromous to semicraspedodromous, 9-10
pairs of secondary veins, angle of divergence 40-60,
secondary veins alternate to subopposite, straight to
slightly curved from midvein dividing 1-2 times before
joining superadjacent vein. Excostal veins in two series,
some terminating in crenulate to serrate teeth. One to
two intersecondary veins per intercosta. Tertiary veins
percurrent and forked.
Discussion.-The shape and size, venation, and
presence of glandular teeth are characteristic of
Theaceae and Ternstroemiaceae, but whether the glands
are deciduous (Ternstroemiaceae) or permanent
(Theaceae) is not evident in the fossil specimens
(Weitzman et al. 2004; Stevens et al. 2004). Characters
such as venation, crenate to serrate margin, and glandu-
lar teeth are similar to extant Gordonia lasianthus (L.)
Ellis (Theaceae, UF5191). Overall, these characters
can be found in Eurya, Adinandra and Ternstroemia
(Terstroemiaceae, Kobuski 1939, 1942a,b, 1943, 1947),
although not in any one particular species. For exact
systematic placement of the fossil Ternstroemites sp., a
leaf venation comparison study is needed for Theaceae
and Ternstroemiaceae. A detailed study of the leaves
of Gordonia and related genera is in progress by Dilcher
and Wang. Distribution of extant species of G lasianthus
and Ternstroemia is warm temperate to predominately
Neotropical (Kobuski 1942a,b, 1943; Dilcher 2000b).
Material examined.-UF 18810-34390, 34399,
34405,34426,34429.

cf Gordonia sp.
(Fig. 18, C, I)
Specimen description.-Flower, 1.8 cm by ~
5 mm basal width, 5 sepals, the lobes with 3-5 longitudi-
nal striations per 1 mm. Petals and carpels missing.
About 46 stamen impressions in 2-3 series, ovary 3.5
mm basal width.


Discussion.- The flower is tentatively allied
with the leaves, Ternstroemites. However, a total re-
study of the fossil record of this complex is in progress.
Grote and Dilcher (1989) described fruits that are also
related to these leaves and flowers. One species,
Andrewsiocarpon henryense Grote & Dilcher was
recognized as totally extinct today but closely related to
Franklinia alatamaha Marsh. The leaves and flow-
ers mentioned here form part of this complex and are
typical of G lasianthus found in the swamp forests of
north Florida today. The flower width of G lasianthus
is 8 cm including petals, absent in the fossil (Godfrey
1988), 3-4 cm in Ternstroemia sp. (Kobuski 1942a,b,
1943), and 2 cm in Adinandra parvifolia Ridley
(Kobuski 1947). A similar flower (Antholithes
ternstroemioides Berry), slightly larger than our speci-
men, was reported by Berry (1930) as similar to G
lasianthus.
Material examined.-UF18810-34438.

Gentianales
Apocynaceae
Apocynophyllum sp.
(Fig. 11, D, E)
Specimen description.-Leaf incomplete, lamina
elliptic, symmetrical, 12 cm long by 4.5 cm wide;
apex missing; base angle acute, with slightly decurrent
flanks; margin entire. Petiole width at blade base 1 mm,
width at petiole base 2 mm. Venation pinnate; second-
ary venation brochidodromous, angle of divergence 40-
500, secondary veins widely spaced, straight from
midvein to near margin, then arching upward and medi-
ally to superadjacent vein. Tertiary veins reticulate.
Discussion.-The incomplete and poorly preserved
nature of this specimen makes further determination dif-
ficult. The leaf is similar to the acute basal form of
Apocynophyllum mississippiensis Berry, except for the
widely spaced secondary veins and reticulate tertiary
veins (Berry 1916, 1930; Dolph 1975). Dolph (1975)
undertook an extensive study of Apocynophyllum leaf
types from the Claiborne sediments of several clay pits
in western Tennessee and Kentucky. He recognized
two distinct subspecies of this extinct genus and after
extensive research has been unable to ally it systemati-
cally with any extant family.
Material examined.-UF 18810-34441.

Rubiaceae
cf Paleorubiaceophyllum sp.
(Fig. 9, A, B)
Specimen description.-Leaf incomplete, lamina
elliptic, symmetrical, 7.5 cm long by 3.5 cm wide; apex







missing; base angle obtuse, with short cuneate flanks;
margin entire. Petiole incomplete, 5 mm long. Midvein
stout; venation pinnate; secondary venation
brochidodromous, angle of divergence 50-60o, more than
10 pairs of secondary veins, sub-opposite to alternate,
straight from midvein to near margin then arching abruptly
upward to superadjacent vein. One to two
intersecondary veins per intercosta. Excostal veins in
one series and looped. Tertiary veins inconspicuous.
Discussion.-The incomplete nature of this leaf
makes our determination uncertain but the general shape
and size, base, margin, and secondary venation are simi-
lar to Paleorubiaceophyllum eocenicum (Berry) Roth
et Dilcher (Roth & Dilcher 1979). Additional speci-
mens are needed to determine if the petiole does have a
stipule attached to its base. This feature is common to
all species of Paleorubiaceophyllum.
Material examined.-UF 18810-34431

Apiales
Araliaceae
Dendropanax eocenensis Dilcher et Dolph
(Fig. 6, A-D)
Description.-Lamina obovate, palmately 3-4
lobed, 5.4-6.7 cm long by 5.8-8.5 cm wide, lobes entire
and sometimes with secondary lobes; apex angle acute,
with cuneate flanks; base angle acute, with concave
flanks. Petiole missing. Venation actinodromous, single
vein entering each lobe; secondary venation interior, basal
sub-primary vein running along margin of basal lobe.
Tertiary venation reticulate.
Discussion.-The shape and size, lobing, and ve-
nation are similar to Dendropanax eocenensis Dilcher
et Dolph (Dilcher & Dolph 1970) except for the lobing
of one specimen that has four lobes, while the speci-
mens described as D. eocenensis have three or five
lobes. Rare Araliaceous pollen is recorded in the
Cockfield Formation of Mississippi (Engelhardt 1964).
The distribution of extant Dendropanax is tropical
America and eastern Asia (Mabberley 1989).
Material examined.-UF18810-34380, 34382a,
34383.

Lamiales
Oleaceae leaf morphotype
(Fig. 11, F; Fig. 12, F)
Specimen description.-Leaf incomplete, lamina
possibly elliptic, symmetrical, 7.5 cm long by 3.5 cm
wide; apex missing; base angle acute, with asymmetri-
cal and slightly decurrent flanks; margin entire. Petiole
slender, 1 mm wide. Venation pinnate; secondary vena-
tion eucamptodromous, angle of divergence 600, sec-


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

ondary veins opposite to alternate, straight from midvein
to near margin, then arching upward to margin. One to
two intersecondary veins per intercosta, traversing up
to 1/2 distance to leaf margin. Tertiary veins at right
angle to secondary veins, straight and divide to adjacent
secondary.
Discussion.-An acute and asymmetrical leaf
base, plus a slender petiole are traits of Oleaceae and
many other families. Further samples are needed for a
clear indication of the distal portion of the leaf for char-
acterization and possible generic determination. Cuticular
characters might also be helpful. Berry (1916) assigned
leaves to Fraxinus and Osmanthus, while Berry (1916)
and Call and Dilcher (1992) described samaras of
Fraxinus. Oleaceaous pollen (Salixipollenites) is re-
corded in the upper Claiborne of the Mississippi
Embayment (Frederiksen 1980a, 1988).
Material examined.-UF 18810-34430

Incertae sedis
Entire Margin Morphotype 1
(Fig. 9, C, E, F)
Specimen description.-Leaf incomplete, lamina
narrowly elliptic, symmetrical, 5 cm long by 2.1 cm
wide; apex and base missing; margin entire. Petiole
missing. Midvein stout; venation pinnate; secondary
venation brochidodromous, angle of divergence 40-50,
~ 10 pairs of secondary veins mostly alternate, arching
from midvein to near margin, then arching upward and
parallel with the margin, then arching medially to join the
superadjacent vein. One to three weak to strong
intersecondary veins per intercosta, the strong vein ex-
tending '/2 to 3/4 width of leaf before dividing and joining
secondary veins. Excostal veins in one series. Tertiary
veins at right angle to secondary veins, joining
intersecondary and secondary veins.
Discussion.-The lack of an apex and base makes
determination to any known leaf type difficult. The
lamina of the fossil leaf appears to have a slight V-con-
figuration in which the midvein is depressed. In the speci-
men figured in Figure 9, C, E, and F, we see the abaxial
side of the leaf so the midvein appears to stand out and
the lamina is an inverse V-configuration. Overall shape,
and venation are similar to those of extant Maranthes
(Chrysobalanaceae) (Prance 1972), Allophylus,
Deinbollia, Meliococca, Scyphonychium, and Talisia
(Sapindaceae) (Standley & Steyermark 1949), and the
fossil Chrysobalanus inaequalis (Lesq.) Berry (Berry
1916).
Material examined.-UF 18810-34433.


Entire Margin Morphotype 2





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

(Fig. 8, A, B)
Description.-Lamina of leaflet elliptic, symmetri-
cal, 6.5-9.5 cm long by 2.5-3.2 cm wide; apex angle
acute, with acuminate flanks; base angle acute, with
convex flanks; margin entire and undulate. Petiole miss-
ing. Venation pinnate; secondary venation
eucamptodromous, angle of divergence 40-50, second-
ary veins mostly alternate, arching from midvein to very
near margin, extending to but not merging with the mar-
ginal vein or one areole loop away from the margin.
Secondary veins connected to superadjacent veins by a
series of tertiary cross veins. One or two intersecondary
veins per intercosta, intersecondary veins traversing up
to 4 distances to the leaf margin. Tertiary veins reticu-
late.
Discussion.-Our specimens lack a petiole which
results in fewer characters useful in systematic place-
ment. The general shape and size, apex, base, margin,
and venation are similar to Aegiphila (Lopez-Palacio
1977) in the Verbenaceae but petiole characters are
important. The Anacardiaceae may have leaves or leaf-
lets with revolute or undulate margins that also have
secondary veins that arch very close to the leaf of leaf-
let margins.
Material examined.-UF 18810-34444a, b, 34412.

Entire Margin Morphotype 5
(Fig. 15, B, D)
Specimen description.-Leaf incomplete, lamina
elliptic, symmetrical, -12 cm long by 5.5 cm wide; apex
and base missing; margin entire. Petiole missing. Ve-
nation pinnate; secondary venation brochidodromous, ~
7 pairs of secondary veins, angle of divergence 30-40,
secondary veins alternate, arching slightly to straight from
midvein to near margin, then arch medially joining
superadjacent vein, with further weaker vein looping
above. Excostal veins in one series. Intercosta of basal
portion of leafwith 1-2 mostly weak intersecondary veins
extending less than V2 distance to leaf margin. Intercosta
of middle and apical portion of leaf with 1-3
intersecondary veins extending up to 1/2 distance to leaf
margin. Tertiary veins weak and appear percurrent.
Discussion.-The incomplete set of characters for
this specimen makes it difficult to place systematically.
The shape and size, margin, and venation are similar to
species such as Gonzalagunia dicocca Cham. &
Schltdl. (Rubiaceae), Markea coccinea Rich. (Solan-
aceae) (Mori et al. 2002), or Magnolia (Magnoliaceae).
Material examined.-UF 18810-34448.

Tooth Margin Morphotype 1
(Fig. 15, E; Fig. 19, A)


Specimen description.-Leaf incomplete, 2 cm
wide; apex angle acute, with attenuate flanks; base miss-
ing. Margin variable, midsection of leaf serrate, apical
section entire. Venation pinnate; secondary venation
variable, midsection venation semicraspedodromous,
apical section brochidodromous, angle of divergence 40-
500, secondary veins alternate, arching slightly from
midvein to near margin, secondary vein divides at base
of tooth, one branch entering tooth, other branch joins
superadjacent vein as a tertiary vein. One to two
intersecondary veins per intercosta. Tertiary veins
percurrent and forked.
Discussion.-The fragmentary nature of this
specimen limits the characters available for systematic
placement of this leafmorphotype. However, the shape,
margin, apex, and secondary venation are similar to Ilex
species living in the swamps near Gainesville, Florida.
Material examined.-UF18810-34445.

Reproductive Structure Morphotype 1
(Fig. 18, F, J)
Specimen description.-Reproductive structure
elliptic, 11 mm long by 5 mm medial width. Apex short
acuminate, base obtuse. Numerous closely spaced lon-
gitudinal striations or veins, eight near the base dividing
to 20 at the midpoint and then rejoining adjacent veins or
striations distally.
Discussion.-Berry (1916) placed this type of
structure in Avicennia eocenica Berry but the system-
atic placement was not clearly demonstrated. Avicennia
fruits are much larger, obpyriform or ovate in shape, and
the striations are widely spaced. The characters of this
fossil do not match Avicennia. This structure appears
to be the remains of a fruit that is presently undeter-
mined.
Material examined.-UF 18810-34434.

Minor Faunal Elements
Insecta
Trichoptera
Insect cases
Folindusia sp.
(Fig. 19, D, G)
Specimen description.-Case straight, 3 cm long,
width of posterior end 3 mm, width of anterior end 8
mm. Six transverse leaf pieces cover the case, 2-8 mm
in length.
Discussion.-The use of leaf material to build the
insect case, the number of leaf pieces, and the case size
is similar to extant caddisfly larval cases (Trichoptera)
and fossil cases of Folindusia wilcoxiana Berry (1927)
and Folindusia sp. (Johnston 1998). Such insect cases







are typical of fresh water, shallow lakes.
Material examined.-UF 18810-34436.

Terrindusia sp.
(Fig. 19,E,F)
Specimen description.-Case incomplete, 4 mm
wide. Case covered with minute leaf pieces, woody
material and sand grains.
Discussion.-The use of sand grains to build the
insect case is similar to that illustrated by Johnston (1998)
but in our specimen organic material is mixed with the
sand grains.
Material examined.-UF 18810-34437.


DISCUSSION
The Paleobotany and Palynology Collection of the Florida
Museum contains collections from clay deposits in west-
ern Tennessee and Kentucky, beginning in 1959 with
Dilcher's collections from Puryear. Dilcher continued
to collect fossil plants from this area from the 1960s
onward with the help of assistants, students, colleagues
and collaborators. Thousands of fossils, including leaves,
fruits and seeds, flowers, wood, pollen, and spores have
been collected from over 30 localities in western Ten-
nessee, western Kentucky, Mississippi, Alabama, Ar-
kansas, Texas and Louisiana. Among the most recent
collections are those from the Powers Clay Pit and
nearby localities. This report is a presentation of the
Powers Clay Pit flora that is available in our collection
at this time. It is the first of a series of reports, detailing
the nature of the floras from many of the clay pits with
the best preserved and most complete collections of fossil
plants.
As additional reports of the middle Eocene locali-
ties become available, they will provide the data neces-
sary to ask many interesting and important questions
involving the deposition, evolution, distribution and
paleobiology of plants from the Gulf Coast of North
America. The composition of these floras is expected
to have a direct link to the relationships between fossil
and modem floras of North America, Central America,
northern South America, Africa and tropical Asia (Gra-
ham 1999; Dilcher 2001; Jaramillo & Dilcher 2001).
Many of these fossil localities given in figure 2 repre-
sent a similar depositional environment, but not all of
them fall into the same category. There is a distinct
probability that each locality represents its own specific
time and climate. Therefore, each flora represents a
unique part in the evolution of plants, through change in
the mixture of floristic elements, and a unique climate.
This could occur over a few million years in the upper


BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)

middle Eocene. The Powers Clay Pit flora, presented
here, is just the beginning of assessing the paleofloristics
of this area, the individual plant fossils and their sedi-
ments. It is anticipated that a floristic database for each
locality will be developed at the Florida Museum ofNatu-
ral History, and made available to the public on the
Internet. Also, the plates of this initial report of Powers
Clay Pit will be available at the Paleobotany and Pa-
lynology web site (http://www.flmnh.ufl.edu/paleo-
botany/).
The present discussion is centered upon the Pow-
ers Clay Pit fossils. This includes their relationship to
fossil plants of other clay pits in the area, realizing that
no detailed flora of each locality has yet been done.
Much of the floristics of the Powers Clay Pit is estab-
lished upon leaf morphotypes and comparisons that can
be made with leaf types of extant and fossil taxa. The
authors feel it is better to publish the available data now
rather than to wait until detailed studies have been com-
pleted for each leaf type. Detailed studies of individual
leaf types are labor intensive in the comparisons of fine
venation and cuticle with living and fossil leaves. While
this type of research is continuing and is included here
for those leaves already studied, we feel that the unique
floristic nature of the individual localities of the middle
Eocene is also of primary interest and should be pre-
sented separately.
Taxa described from the Powers Clay Pit flora
allow for character comparisons with other local middle
Eocene localities. Detailed species accounts from Pow-
ers Clay Pit have documented the first occurrence of a
lobed Platanus leaf (Fig. 7, A-C), and a Liquidambar
fruit (Fig. 18, D, H) that is described and illustrated.
Also, new leaf characters presented here enhance our
knowledge of fossil leaf variation. An example is the
intermediate form of Berryophyllum sp. where teeth,
margin, venation and leaf size demonstrates variations
not previously recognized (Jones & Dilcher 1988). Other
character variations include the leaf apex of
Berhamniphyllum sp., and leaf shape in the intermedi-
ate form of Populus sp. Such variations in leaf form
were also found in other localities, along with variations
in cuticular characters (Dilcher & Mehrotra 1969; Roth
& Dilcher 1979). These authors suggested that the rea-
son for these variations involved the possibility that each
fossil locality is of a specific age, with leaf characters
changing through time, demonstrating microfeatures of
evolution. Some variations may be related with sample
size, so an increase in the number of specimens may
widen the range of variation. Also, the differences noted
between localities were consistent over a large number
of specimens from each locality (Roth & Dilcher 1979).




DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

The Powers Pit flora also contains readily identifiable
taxa based upon gross morphological features of the
megafossils. These include Berryophyllum tenuifolia
(Berry) Jones et Dilcher (Berry 1916; Jones & Dilcher
1988), Berhamniphyllum claibornense Jones et Dilcher
(Jones & Dilcher 1980), Nyssa eolignitica Berry (Berry
1916; Dilcher & McQuade 1967), and Dendropanax
eocenensis Dilcher et Dolph (Dilcher & Dolph 1970).
Dilcher and Mehrotra (1969) found that consistent cu-
ticular differences could be observed in
Knightiophyllum leaves between localities but differ-
ences were not apparent in the gross features of the
leaves.
A recurring problem when identifying extant taxa
based upon fossil leaves is the convergence in foliar char-
acters in numerous plant families and genera (Berry
1914). Detailed analyses of venation patterns may help
alleviate this problem, such as differences in
intersecondary vein branching in Vochysiaceae and
Myrtaceae, which in turn helps to identify fossil Myrcia.
Intersecondary venation and basal secondary venation
helps distinguish Moraceae from Lecythidaceae, lead-
ing to the identification of Ficus. Analyses of second-
ary vein angles and ultimate venation helped distinguish
extant Duguetia from Magnolia which lead to the iden-
tification of fossil Duguetia. The determining charac-
ter used to suggest that Berry's Magnolia fossil leaves
were really Duguetia (Annonaceae) was trichome type
(Roth 1981; Dilcher & Roth, unpublished data). Ter-
tiary venation is important, such as in distinguishing similar
leaflet forms of Fabaceae from Connaraceae, where
overall tertiary venation is more prominent in
Connaraceae. Also, Smilax and Dioscorea leaves are
very similar except that in Dioscorea the tertiary veins
are distinctly opposite percurrent. Sometimes, further
morphological studies are needed to distinguish between
two closely related families, such as Theaceae and
Ternstroemiaceae where venation patterns may confirm
placement of Ternstroemites sp. within Theaceae. Other
morphological features are important in distinguishing
between similar leaf forms. Extant Cladrastis and
Sapindus are very similar but the leaf base of Cladrastis
is straight, while Sapindus has a curved, often
inequilateral base, leading to the identification of fossil
Cladrastis. Morphological leaf characters may not be
enough, so future anatomical studies are needed that
incorporate cuticular characters. Such characters are
important in determining Swartzia to the species level
since similar morphological and venation patterns are
present in numerous species of Swartzia. Use of cu-
ticular characters is also important for Ocotea even
though this fossil leaf has been linked with extant O.


quianensis here, and in Berry (1916). Some leaves
from Powers Clay Pit have an uncertain systematic
placement, including Entire Margin Morphotype's 1, 2,
5, Tooth Margin Morphotype 1, and Reproductive Struc-
ture Morphotype 1. This would include Knightiophyllum
wilcoxianum Berry where placement in Fagaceae is
open to question. Knightiophyllum wilcoxianum does
not match any extant genus and represents an extinct
member of Fagaceae. Specimens that are incomplete
and problematic in placement include Apocynophyllum
sp., cf Paleorubiaceophyllum, and Oleaceae, with
Apocynophyllum and Paleorubiaceophyllum repre-
senting extinct taxa (Dilcher 2001).
It has been over 145 years since the first investi-
gation of fossil plants from the Claiborne flora
(Lesquereux 1859) and about 90 years since the first
detailed locality to locality comparisons of these fossil
plants (Berry 1916). Since that time, fossil plants from
new sites have been collected and investigated, increas-
ing the number of localities in western Tennessee, west-
ern Kentucky, Mississippi and Alabama from 10 (Berry
1916) to 30. With the taxa presented here from Powers
Clay Pit, a preliminary comparison can be made with
the limited taxa that have been studied and identified
from other pits in western Tennessee and western Ken-
tucky. Taxa shared with Powers Pit have been found at
the Puryear Clay Pit (15 taxa), Willbanks I Clay Pit (14
taxa), Lamkin Clay Pit (seven taxa), Warman Clay Pit
(seven taxa), and Lawrence Clay Pit (six taxa) (Table
2). As our research continues, it is anticipated that new
morphotypes will be discovered, increasing our under-
standing of the floristics for each locality. Even though
the present data is preliminary, interesting comparisons
have been found.
Of the 35 fossil plant taxa recognized from Pow-
ers Clay Pit, the number of taxa that overlap range from
15 at Puryear Clay Pit to six at Lawrence Clay Pit. In
other words, less than half the known taxa overlap lo-
calities. Although Warman Clay Pit is only 400 meters
north of Powers Clay Pit, the two sites seem to share
very few taxa. Lawrence is 8 KM away, but also shares
few taxa with Powers Pit. Wilbanks Clay Pit shares 14
taxa and is only 1 KM west of the Powers Clay Pit.
Puryear Clay Pit is about 25 KM north north-east, and
Lamkin is about 65 KM north of Powers Clay Pit. It is
interesting to note that distance does not seem to relate
to similarity offloristic elements. This disparity could be
explained by sample size, environmental factors, or that
these clay pits may not represent one common age.
Certainly, more research needs to be done to under-
stand the floristic relationships between the clay pit lo-
calities.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


Table 2. Taxa shared with Powers Clay Pit, and five other clay pits.


Puryear

Ocotea

Duguetia


Smilax


Myrcia
(Calyptranthes)

Cladrastis
(Gleditsiophyllum)

Berryophyllum

Knightiophyllum

Populus
(Grewiopsis)

Ficus

Berhamniphyllum
(Rhamnus)


Willbanks


Ocotea


Duguetia

Smilax

Ormosia


Lamkin

Ocotea

Myrcia

Ficus


Berhamniphyllum


Berryophyllum Ternstroemites


Knightiophyllum Apocynophyllum


Populus


Paleorubiaceoyllum


Berhamniphyllum


Nyssa

Ternstroemites?
(Gordonia)


Warman


Smilax? (Fossil type 5)

Berryophyllum

Knightiophyllum


Berhamniphyllum


Ternstroemites


Paleorubiaceophyllum


Lawrence


Berryophyllum

Knightiophyllum

Berhamniphyllum


Nyssa


Paleorubiaceophyllum


Dendropanaxyllum


Cupanites


Nyssa


Ternstroemites

Apocynophyllum


Apocynophyllum

Paleorubiaceophyllum

Dendropanax

Folindusia? (Caddisfly case)


Paleorubiaceophyllum


The relationships of fossil specimens to extant gen-
era can give us a glimpse of past ecosystems. Many of
the fossil specimens from Powers Pit appear to be re-
lated to extant genera that have primarily subtropical or
tropical distributions. These include Ocotea, Swartzia,
Ormosia, Berryophyllum, Ternstroemites, Dendro-
panax, Duguetia, Moraceae, Myrcia, and Berham-
niphyllum. Fossil specimens related to extant genera
that have primarily cool to warm temperate to subtropi-


cal distributions include Liquidambar, Populus, Nyssa,
Cladrastis, and Platanus. Berry (1916, 1924) concluded
that the climate of the "Wilcox" (now Claiborne) flora
was warm tropical. Miranda and Sharp (1950), Sharp
(1951), Gray (1960) and Engelhardt (1964) compared
the Eocene flora to the eastern Mexican escarpment
region and suggested a mixture of temperate and tropi-
cal conditions. Dorf (1960) considered the climate as
tropical, while Axelrod (1966) and Dilcher (1973) con-


Dendropanax





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit

eluded that the climate was warm temperate to cool
subtropical or cooler temperate, and Dilcher (1973) sug-
gested that there were definite seasonal dry periods.
Elsik (1974) suggested a tropical to subtropical climate
while Wolfe (1975, 1978), Wolfe and Poore (1982),
Frederiksen (1980b, 1988) and Graham (1999) consid-
ered a winter dry tropical to marginal humid subtropical
climate. The fossil specimens presented here suggest a
subtropical climate that can accommodate warm tropi-
cal to cool temperate species. While the lowlands pro-
vided tropical plant species, possible uplands with an in-
fluence from the Appalachian area may have provided
cool temperate species (Cain 1943; Dilcher 1973). The
unique mixture of elements living in temperate, subtropi-
cal, and tropical climates today suggests that they may
represent parts of an ancient non-analogue climate. That
is, a climate that is unique in the characteristics of its
temperature and moisture that is found nowhere in the
world today. Plants that may be related to these fossils
continue living today in various places and in slightly dif-
ferent climates as they have shifted their distributions
and evolved various ecological tolerances to accommo-
date present conditions.

ACKNOWLEDGMENTS
We would like to thank the following people for
help in collecting specimens from Powers Pit, Hongshan
Wang, Eizabeth Kowalski and Jonathan Block (Florida
Museum of Natural History), Michael Gibson (Univer-
sity of Western Tennessee), Mihai Popa (University of
Bucharest), and Prakhart Sawangchote (Suranaree
University of Technology). We also like to thank Steven
Manchester, Deborah Matthews, and one anonymous
reviewer for their valuable comments and suggestions,
and Jason McCuiston of Franklin Industries/ H. C. Spinks
Clay Company for access to Powers Pit. This is Uni-
versity of Florida Contribution to Paleobiology number
556.


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DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


Table 3. Leaf Morphology Key


1. Leaves lobed.
2. Lobes entire.
2. Lobes toothed.
1. Leaves simple (leaf and leaflet).
3. Leaf margin entire.
4. Leaves small (leaf length < 8 cm).
5. Leaves elliptic.
6. Venation brochidodromous.
7. Lamina ratio 2-3:1.
8. Secondary veins 5-6 pairs
8. Secondary veins more than 10 pairs
7. Lamina ratio 4:1.
6. Venation eucamptodromous
5. Leaves lanceolate, brochidodromous.
4'. Leaves medium (leaf length 8-15 cm).
9. Leaves elliptic.
10. Intramarginal vein.
10'. Venation brochidodromous
11. Base obtuse
11. Base acute
12. Flanks convex
12'. Flanks decurrent
12". Flanks concave
10". Some venation eucamptodromous.
13. Leaves lack ascending basal veins
14. Angle of divergence 20-30
14. Angle of divergence 600
13. Leaves with ascending basal veins
10"'. Venation acrodromous/campylodromous
9. Leaves obovate.
4". Leaves large, elliptic (>15 cm).
3. Leaf margin toothed.
15. Leaf medium (6-11 cm).
16. Leaf elliptic.
17. Teeth confined to mid-upper portion.
17.Teeth entire length
18. Teeth serrate.
18. Teeth mostly dentate
16'. Leaf oblong.
16". Leaf obovate.
16'". Leaf linear.
15. Leaf large, elliptic (> 11 cm).
19. Teeth confined to mid-upper portion.
20. Teeth prominent.
20. Teeth short
19. Teeth entire length.


Dendropanax eocenensis
Platanus








Swartzia
cf. Paleorubiaeophyllum
Ormosia sp. 2
Berhamniphyllum sp.
Cladrastis



Myrcia

Duguetia

Ormosia sp. 1
Apocynophyllum
Ficus



Berhamniphyllum claibornense
Oleaceae
Ocotea
Smilax
Pseudolmedia
Duguetia




Knightiophyllum wilcoxianum

Ternstroemites sp.
Populus sp.
Cupanites sp.
Ternstroemites sp.
Berryophyllum tenuifolia



Berryophyllum sp.
Knightiophyllum wilcoxianum
Ternstroemites sp.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


Figure 6. Dendropanax eocenensis. A, B, Three lobed leaf, UF18810-34383, Ix, 2x. C, D, Four lobed leaf,
UF18810-34382a, lx, 2x. Bar-10 mm (A, C), 20 mm (B, D).






DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


I.
I`


Figure 7. Lobed and entire leaves. A-C, Platanus sp., UF18810-34439, lx, 2x, 3x. D-H, J, L, Swartzia sp. D, E,
UF18810-34391, lx, 2x. F, J, UF18810-34394, lx, 2x. G H, L, UF18810-34393, 2x, lx, 4x. I, K, Ormosia sp.
2, UF18810-34395, lx, 2x. Bar-5 mm (C-F, I-L), 10 mm (A, B, G, H).





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


Figure 8. Entire leaves. A, B, Entire Margin Morphotype 2, UF18810-34444b, Ix, 2x. C, D, Ficus sp., UF18810-
34446, Ix, 2x. All bars at 5 mm.





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


-4
















Figure 9. Entire leaves. A, B, cf. Paleorubiaceophyllum sp., UF18810-34431, lx, 2x. C, E, F, Entire Margin
Morphotype 1, UF 18810-34433, 2x, lx, 3x. D, G, H, Berhamniphyllum sp., UF18810-34427, lx, 3x, 2x. All
bars at 5 mm.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


Figure 10. Entire leaves. A-C, Cladrastis sp. A, UF18810-34413a, Ix. B, C, UF18810-34402, Ix, 2x. D, E,
Duguetia sp., UF18810-34407, lx, 1.5x. F-H, Myrcia sp., UF18810-34386, lx, 3x, 2x. All bars at 5 mm.





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


Figure 11. Entire leaves. A-C, Ormosia sp. 1. A, UF18810-34406, Ix. B, C, UF18810-34404a, 1.5x, 2x. D, E,
Apocynophyllum sp., UF18810-34441, 1.5x, Ix. F, Oleaceae, UF18810-34430, Ix. All bars at 5 mm.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


A




























F


Figure 12. Entire leaves. A, B, Berhamniphyllum claibornense, UF18810-34384, Ix, 2x. C-E, G, Ocotea sp. C,
UF18810-34409, lx. D, E, UF18810-34410a, lx, 2x. Q, UF18810-34411, 2x. F, Oleaceae, UF18810-34430,
2x. All bars at 5 mm.





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


Figure 13. Entire leaves. A, B, Swartzia sp., UF18810-34388a, Ix, 2x. C, D, Smilax sp. 1, UF18810-34415, 2x,
Ix. E, Smilax sp. 2, UF18810-34435, Ix. All bars at 5 mm.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


B

























Figure 14. Entire leaves. A, Smilax sp. 2, UF18810-34435, 1.5x. B, D, Pseudolmedia sp., UF18810-34417, Ix,
2x. C, Duguetia sp., UF18810-34418, 1.5x. All bars at 5 mm.




DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


D ql

Figure 15. Entire and toothed leaves. A, C, Entire Margin Morphotype 3, UF18810-34450, 2.5x, Ix. B, D, Entire
Margin Morphotype 5, UF 18810-34448, 2x, Ix. E, Toothed Margin Morphotype 1, UF 18810-34445, Ix. Bar-5
mm (A, C), 10 mm (B, D, E).





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Figure 16. Entire and toothed leaves. A, Duguetia sp., UF18810-34418, lx. B, C, G, Knightiophyllum
wilcoxianum, UF18810-34425a, lx, 2x, 2x. D, Populus sp., UF18810-34397, lx. E, F, Ternstroemites sp.,
UF18810-34399, lx, 2x. Bar-5 mm (D-G), 10 mm (A-C).





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


E














Figure 17. Toothed leaves. A, Populus sp., UF18810-34397, 2x. B, D, Cupanites sp., UF18810-34401, lx, 2x.
C, I, Ternstroemites sp. UF18810-34426, lx, 2x. E-G, Berryophyllum sp., UF18810-34396, lx, 3x, 2x. H,
Berryophyllum tenuifolia, UF18810-34389, lx. All bars at 5 mm.





BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)


Figure 18. Toothed leaves and reproductive structures. A, E, Ternstroemites sp. UF18810-34405, Ix, 2x. C, I,
Gordonia sp., UF18810-34438, 4x, Ix. B, Berryophyllum tenuifolia, UF18810-34389, 4x. D, H, Liquidam-
bar sp., UF 18810-34432, 3x, Ix. F, J, Reproductive Structure Morphotype 1, UF18810-34434, 7x, Ix. G, K,
Nyssa eolignitica, UF18810-34392, 3x, Ix. Bar-2 mm (F), 5 mm (B-E, G-K), 10 mm (A).





DILCHER and LOTT: A fossil assemblage of Powers Clay Pit


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Figure 19. Leaves and insect larval cases. A, Tooth Margin Morphotype 1, UF 18810-34445 cpt., 3x. B, Monocot
leaf, UF18810-34400, lx. C, H, Entire Margin Morphotype 4, UF 18810-34449, lx, 2x. D, G, Folindusia sp.,
UF18810-34436, lx, 3x. E, F, Terrindusia sp., UF18810-34437, lx, 6x. Bar-5 mm (A, B, D-H), 10 mm (C).







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RECENT PUBLICATIONS OF THE BULLETIN

Dilcher, D.L. & T.A. Lott. 2005. A middle Eocene fossil plant assemblage (Powers Clay Pit) from western
Tennessee. Bull. Florida Museum Nat. Hist. 45(1): 1-43. Price $7.00

King, F. W. and C. M. Porter, (Editors). 2003. Zooarchaeology: Papers to honor Elizabeth S. Wing. Volume 44, No.1,
pp.1-208. Price $20.00

MacFadden, B. J. and O Carranza-Castaneda. 2002. Cranium of Dinohippus mexicanus (Mammalia: Equidae)
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Kratter, A. W., T. Webber, T. Taylor, and D. W. Steadman. 2002. New specimen-based records of Florida birds.
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MacFadden, B. J. 2001. Three-toed browsing horse Anchitherium clarencei from the early Miocene (Hemingfordian)
Thomas Farm, Florida. Volume 43, No.3, pp. 79-109. Price $5.50

F. G. Thompson and G. P. Brewer. 2000. Land snails of the genus Humboldtiana from northern Mexico (Gastropoda,
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Hayes, F. G. 2000. The Brooksville 2 local fauna (Arikareean, latest Oligocene): Hernando County, Florida. Volume
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