FLORI'IDA MUSEUM BULLETIN OF NATURAL HISTORY ®
A MIDDLE EOCENE FOSSIL PLANT ASSEMBLAGE (POWERS CLAY PIT) FROM WESTERN TENNESSEE
DavidL. Dilcher and Terry A. Lott
Vol. 45, No. 1, pp. 1-43 2005
UNIVERSITY OF FLORIDA GAINESVILLE -
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Publication Date: October 31,2005
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Managing Editor of the BULLETIN Florida MuseumofNatural-History University offlorida PO Box 117800 Gainesville, FL 32611 -7800 U.S.A. Phone: 352-392-1721 Fax: 352-846-0287 e-mail: [email protected] A MIDDLE EOCENE FOSSIL PLANT ASSEMBLAGE (POWERS CLAY PIT) FROM WESTERN TENNESSEE
David L. Dilcher and Terry A. Lottl
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 Claibome 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
Dedication 2 Introduction 3 Materials and Methods X Systematic Paleontology 10 I mirnlpq 10 Magnoliales...... „.,~,-,..,...... „,„~...... ,..~-...... ,10 Imdeq 11 Proteales 11 Saxifragales Myrtales 17 Fabales 1713 Fagalp.R Malpighiales 14 Rosales 1 5 Sapindales ...... ,.„„.,...... „,~,...... „,...... ,16
Cornales 1617 F.ricales Gentianales 17 Apiales 18 I Amialeq Incertae sedis 18 Insecta 19 Discussion 10 Acknowledgments 73 Literature Cited 73
'Florida Museum of Natural History, University of Florida, Gainesville, FL 32611-7800, email:
IN DEDICATION TO: BRUCE ROGER MOORE
1r
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 ofthe Florida Museum ofNatural History. His field-work, as part ofa team effort, resulted in collections ofseveral ofthe 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 ofother fossil localities. DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 3
INTRODUCTION are many localities in this region (Fig. 2) and each local- There are numerous clay deposits in western Tennes- ity yielded hundreds to thousands, even tens of thou- see, many which contain abundant plant fossils. These sands, of plant fossil remains. This report is the first clay deposits are of middle Eocene age and occur as publication in what we expect to become a series of isolated clay lenses of the Claiborne Group deposited papers detailing the floras o f each locality. Reasons for during the Mississippi Embayment of southeastern North breaking this very large flora of middle Eocene plant- America. These numerous clay lenses are distributed rich deposits into discreet units are elaborated upon in in the ancient coastal plains sediments o f the northeast- the discussion of this paper. ern part of the Mississippi Embayment (Fig. 1). There The fossil plants in the present report were col
i ~ ILLINOIS-J KANSAS MISSOURI ' - ENTUCKY~
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Ter Jackson group Claiborne group Wilcox group Tj Jackson Formation Tc Cockfield Formation Th Hatchetigbee Formation Eocene Ty Yazoo Clay ~ Tco Cook Mountain Formation Tw Wilcox Formation May include Paleocene locally Trn Moodys Branch Formation Ts Spana Sand Tpr Tz Zilpha Clay Tt Tallahatta Formation Paleocene Trm Reklaw Formation Kr F1 Cretaceous
Figure 1. Geologic map of Mississippi Embayment. Stratigraphic units as recognized by U.S. Geological Survey. Modified from Tschudy (1973). 4 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
KENTUCKY 17··1 •2 GRA\'ES
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Powers I Pit #14 N SR 54 Como A 190 M CR m Fr nc CR 140 -~ La e
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-HEV-coMS_ B cKe e Carroll County Weakley (0199
Figure 2. A, Localities of clay pits in Kentucky, Tennessee, and Mississippi, Eastern United States. 1, Lamkin (UF15815). 2, South-40 (UF 15824). 3, Bell City (UF15803). 4, New Puryear (UF15819). 5, Puryear (UF 15820). 6, Martin (UF15809. 7, Foundry Hill(UF15810). 8, Buchanan (UF15806). 9, Warman (UF15826). 10, Gleason (UF15811). 11, Miller(UF15817). 12, Lawrence (UF15816). 13,Willbank 1&2(UF18884, UF18927). 14, Powers OF I 8810). 15, Bovay (UF 1 5737) 16, Bolden (UF15738). 17, Old Hickory (UF15742). 18, Richies Black (UF15828). 19, New Lawrence (UF15818). 20,New Haynes (Rancho, UF15921). B, Locality ofPowers Pit (#14), Weakley County, Tennessee. DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 5
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Figure 3. Powers Pit. A, Northeast comer. B, East side of pit. Photographed by Hongshan Wang. 6 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1) S3IhIEIS WHITLATCH SCHNEIDER STEARNS HARDEMAN PARKS PARKS AND AND ETAL. AND BLANKENSHIP ARMSTRONG CARMICHAEL (surface) (subsurface) (subsurface) (surface) (surface)
1940 1950 1955 1966 1975 1990 JACKSON JACKSON JACKSON(?) JACKSON (?) FM. FORMATION FORMATION FORMATION o
JACKSON COCKFIELC
GRENADA AND FM.
FM. 3N3003 CLAIBORNE ~ COOK CLAIBORNE GROUPS ...... D- ..18.. .1 ~ MOUNTAIN GROUP CLAIBORNE r HOLLY AND CLAIBORNE ~ FM.
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WILCOX MEMPHIS WILCOX ACKERMAN WILCOX WILCOX SAND (?) FM. FM. GROUP GROUP
PORTERS PORTERS PORTERS PORTERS PORTERS WILCOX 3N30031Vd CREEK CREEK CREEK CREEK CREEK CLAY 5 CLAY 0- CLAY 0- CLAY a. CLAY GROUP ODD 0 0 0 0 GROUP 1 0 0M r H ? 24 / > >s 07 ~ CLAYTON 5 CLAYTON ~ CLAYTON ~ CLAYTON MIDWAY o FM. 8 FM. a FM. g FM MIDWAY 5 E 5 GROUP 0
UPPER CRETACEOUS
Figure 4. Stratigraphic chart ofwestern Tennessee. Modified from Parks (1975) and Parks and Carmichael (1990). lected at Powers Clay Pit in Weakley County, Tennes- site. The well preserved fossil leaves, flowers, fruits, see (Figs. 2,3). This clay pit was selected because we seeds, and insect larval cases occur as compressions/ have accumulated, during the past 5 years, a collection impressions, casts, and carbonized organic material. ofoyer400 specimens consisting mainly ofleaves, fruits, Many of the fossils collected in Powers Clay Pit can be and seeds through the efforts of B. Roger Moore, the related to the extant families Lauraceae, Fabaceae, authors, and numerous colleagues who have visited the Fagaceae, Theaceae, and Araliaceae, with less abun- DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 7
GROUP FORMATION Jackson; upper Eocene Moodys Branch
Cockfield Fm. Cook Mt. Fm. Claiborne Group; Sparta sand- Zilpha Fm. middle Eocene Neshoba Mbn REGIONAL Tallahatta Fm.
MARINE Reklaw DISCONFO
Wilcox; lower Eocene
TRANSGRESSION REGRESSION
Figure 5. Transgression-regression schematic. From Potter and Dilcher 1980.
dant fossils related to Annonaceae, Moraceae, 2000a). The entire concept of presenting huge fossil Rhamnaceae, Smilacaceae, and Myrtaceae. A few rare floras that demand only brief comparisons with living specimens are related to Platanaceae, Altingiaceae, plant genera are now understood to be filled with identi- Salicaceae, Sapindaceae, Nyssaceae, Apocynaceae, fication errors. As Dilcher (1974) mentioned, Berry's Rubiaceae, and Oleaceae. There are also some caddisfly identifications for the Eocene floras ofsoutheastern North cases associated with the plant fossils. America were about 60% incorrect or held some mar- E.W. Berry (1916,1930,1941) originally consid- gin of systematic errors and thus Berry's conclusions ered the plant fossils from numerous localities extending are often not supported. With this in mind, the flora from Texas, Arkansas, western Tennessee, western Ken- presented here is done so with tentative assignments to tucky, Mississippi and Alabama to be of lower Eocene, extant genera and more certain assignments to extant belonging to the Wilcox Formation. Later, the age of families as in Dilcher (2001). This is logical because this flora was clearly shown based on palynological cor- many extant families can be recognized and validated relations to belong to the middle Eocene Claiborne Group by Eocene times, but fewer extant genera can be vali- (Elsik & Dilcher 1974; Frederiksen 1988). As well as dated by that time. The use of leafrnorphotypes (Johnson the age, the depositional settings ofthese plant-rich clay 1989; Ellis et al. 2003; Barclay et al. 2003) is a practical deposits has been revised over the last 60 years. These way to proceed in paleobotanical studies of fossil leaves revisions will be discussed in the material and methods for which a modern generic affinity is uncertain. Sev- section of this report. eral angiosperm leaf morphotypes are presented in this Since the original reports ofthe Eocene floras from study. southeastern North America (Berry 1916,1930,1941), This report represents the first attempt in over 75 methods for the study of fossil plant remains have been years of presenting an entire known fossil flora based revised, and the philosophy ofreporting on specific fos- upon megafossil evidence, from the Eocene sediments sil plants and floras has undergone extensive modifica- of southeastern North America. It is limited to a single tion (Hickey 1973; Dilcher 1974; LAWG 1999; Dilcher locality, represents a change in age, presents new ideas 8 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1) in the deposition of the flora, and a new philosophy in known fossil floras. The use of such comparisons in the our aRproach to this floristic study from previous floras understanding of the paleoenvironment, paleoclimate, ofthis region by Berry (1916,1930). The more recent stasis and change in floristic composition through time research of these fossil plants, limited to studies of spe- will be presented in the discussion. As floras from other cific taxa or specific families, is indicated below. localities are published, we expect that the plant species composition ofthis locality will be used to compare and PREVIOUS PALEOBOTANICAL INVESTIGATIONS contrast with species lists from other middle Eocene fossil There is a long and rich history of investigations of plant localities. A leaf morphology key is provided for plant megafossils collected in western Tennessee and preliminary field identification for this one locality (Table Kentucky, starting in the mid 1800's (Lesquereux 1859, 3). This will allow the leafmorphotypes presented here 1869; Owen 1860; Safford 1869; Berry 1914,1916,1922, to be used in field studies ofplant fossil diversity, overall 1924,1930,1937,1941). Brown (1939,1940,1944,1946, species richness and abundance. 1960) provided limited taxonomic revisions ofsome of the fossil material previously published by Berry. Sub- MATERIALS AND METHODS sequently, systematic revisions and additions to the The plant megafossils and insect larval cases presented Claiborne megafossil flora can be found in Dilcher (1963, in this report were collected mostly in July 2000, May 1969,1971, 200Ob), Dilcher & McQuade (1967), Dilcher 2001 and April 2002 from the Powers Clay Pit, Weakley & Mehrotra (1969), Dilcher & Dolph (1970), Dilcher et County, Tennessee, 36° 16.94 N, 88° 31.78 W, in sedi- al. (1974,2001), Crepet et aL (1974,1975,1980), Dolph ments belonging to the middle Eocene, Claiborne Group. (1975), Crepet & Dilcher (1977), Dilcher & Daghlian Specimens were also collected from a commercial shed (1977), Crepet (1978), Roth & Dilcher (1979), Crepet with piles of clay mined from Powers Clay Pit, at the & Daghlian (1980,1982), Jones & Dilcher (1980,1988), headquarters ofH. C. Spinks Clay Company in July 2002. Potter & Dilcher (1980), Weiss (1980), Roth (1981), Terminology for leafdescription follows Hickey (1973), Zavada & Crepet (1981), Jones (1984), Dilcher & Radford et al. (1974), Dilcher (1974) and LAWG (1999). Manchester (1986, 1988), Jones et al. (1988), Taylor The systematic classification used follows APG II (2003). (1988), Grote & Dilcher (1989, 1992), Sun (1989), The fossil material is numbered with the University of Herendeen & Dilcher (19908,b,c, 1991), Herendeen et Florida acronym (UF) followed by the locality number al. (1990), Call & Dilcher (1992), Herendeen (1992), (18810 for Powers Clay Pit), then the individual speci- Moore (2001), & Moore et al. (2003). Palynological men number (e.g. UF 18810-34381). Modern leaves cited reports from western Tennessee can be found in for comparison are numbered as the University of Florida McLaughlin (1957), Tschudy (1973), Elsik & Dilcher Modern Reference Leaf Collection followed by the (1974), Potter (1976), & Taylor (1987,1989) while dis- specimen number (e.g. UF5437). Herbarium specimens persed cuticle of one locality was presented by Kovach of modern leaves were examined at the University of & Dilcher (1984). These publications since 1963 repre- Florida Herbarium (FLAS). Some specimens are fig- sent work on individual taxa or limited families common ured but not described such as Entire Margin in the Eocene floras of western Tennessee. The reason Morphotype 3 (UF 18810-34450) and Entire Margin for this systematic rather than floristic emphasis is given Morphotype 4 (UF 18810-34449). in the discussion ofthis paper. Changing concepts of stratigraphic nomenclature for the Claiborne Group in western Tennessee since the ~ OBJECTIVES 1940's are illustrated in Figure 4. The clay lenses in the This study is an attempt to document the megafossil upper Claiborne Group were considered to represent flora (with minor faunal elements included) of one spe- near-marine paleoenvironments because ofthe possible cific clay pit in western Tennessee, by presenting each coastal elements, such as sabal palms in the fossil floras type of megafossil plant found that is now in the Paleo- (Berry 1916, 1930), sedimentary environment types botany and Palynology Collection ofthe Florida Museum (Stearns 1957), and the presence of hystrichospheres of Natural History. Leaf morphotypes are presented and dinoflagellates (Parks 1975). Dilcher (1974) sug- and morphological descriptions ofeach taxon are corre- gested that the clay lens of some of the plant-rich clay lated with modern genera when some similarities are pits represented oxbow lake deposits. Dolph (1975) also noticed. As taxa can be related to a specific extant envisioned back-swamp paleoenvironments on ancient family, extant genus, and extinct fossil plant morphotypes, floodplains while Potter (1976) also suggested abandoned the overall floral composition of this fossil locality can river channels. Potter & Dilcher (1980) presented nu- be compared with floras of modern ecosystems and merous detailed outlines of several clay pits with some ]DILCHER,a®(1261*,Af6§~il 65»inEl@66*Rowey*Clay. R.it . 9
Ta816 1.: S*tEmafit# liat 6f thefflofa· midfmiiia:,With ,6xemplafyl*edimeirs.. -* Indicatess,an extincf fossil genus or unla~vn- imidern Btfietaisimilh-r'fla this, fess#
.
Order Family, ISpefiss, Magnoliids Laurales Lauraceae Ocotea sp Magfialiales Atinonaceae Duide-lib :ip: M0*0-eets I1iaja# Smi_lheaceae Sm#*sp.1&2 Uhdetermined Monocot* sp. Eudicots Proteales Platanaceae Platanus* sp. Core Eudicots Saxifragales Altingiaceae Liquidambar sp: Rosids Myrtales M~rtaceae A*rcia sp, Euro#da I Fabales Fabaceae Cladrastis-like sp. Ormosiaelike sp. 1&2 Swartzia sp. Fagales Fagaceae Berryophyllum* tenugolia (Berry) Jones et Dilcher Berryophyllum* sp. Knightiophyllum* wilcoxianum Berry Malpighiales Salicaceae Populus sp. Rosales Moraceae Pseudolmedia sp. Ficus.like sp. Rhamnaceae Berhamniphyllum* claibornense Jones et Dilcher Berhamniphyllum* sp. Eurosids II Sapindales Sapindaceae Cupanites sp. Asterids Cornales Nyssaceae Nyssa eolignitica* Berry . Ericaies Theaceae Ternstroemites* sp. €f Gordonia sp. Euasterids I Gentianale5 Apocynaceae Apocynophyllum* sp. Rubiaceae ef Paleorubiaceophyllum* sp. Euasterids H Apiales Araliaceae Dendropanax eocenensis Dilcher et Dolph Lamiales Oleaceae Oleaceae.leafmorphotype Inceriae sedis 'Entire margin morphotype 1 * Entire marginmorphotype 2* Entire margin morphotype(3* Entife margin morphotyp'e 4* Entire margin
Specimen description.-Leaf incomplete, palmately intramarginal vein. Some intersecondary veins present, lobed, appears to be 3-lobed; apex missing; base angle a few admedially branched. Leaf margins with paired obtuse, with rounded flanks. Leaf margin occasionally intramarginal veins, one running along the margin, the serrate, teeth straight to curved, sinuses rounded. Peti- second 1 -2 mm inside the leaf margin and·slightly seal- ole missing. Venation palinactinodromous; secondary loped. Tertiary veins sometimes joining secondary veins venation camptodromous, angle ofdivergence 35-40°. at acute angles. Discussion.-The lobed nature of the leaf is simi- Discussion.-The spacing and angle of the sec- lar to Lauraceae (Sassafras), Altingiaceae (Liquidam- ondary veins, along with the intramarginal venation is bar), and Platanaceae (Platanus ), although in Sassa- similar to Qualea in Vochysiaceae, and Calyptranthes, fras and Liquidambar the primary venation is Syzygium and A*rcia in Myrtaceae (Klucking 1988). suprabasally actinodromous, and in Liquidambar the The difference is in the intersecondary veins , in margins are continuously serrate, and with basal lobing Vochysiaceae they branch toward the margin, while in (Schwarzwalder 1986). General characters such as Myrtaceae some are admedial, branching towards the lobes, venation, and occasional teeth are similar to midvein. Also, in Vochysiaceae, the secondary veins Platanus sp., although Platanaceae venation is some- curve toward the margin (Sajo & Rudall 2002). Vena- times superbasally actinodromous. The secondary veins tion is similar to Myrcia and Syzygium, butin Syzygium are not sufficiently preserved near the margins to deter- the secondary veins are recurved to sigmoidal (Klucking mine i f they extend into the teeth ("Platanoid") or not, 1988). Leafsize, along with secondary and intramarginal but the slope does suggest that they curve away from venation is similar to Calyptranthes eocenica Berry the teeth as in Platamis kerrii Gagnep. (Schwarzwalder (Berry 1916), but in Calyptranthes, both intramarginal 1986). This is the first occurrence ofa lobed Platanus vein pairs run 1 -2 mm inside the leaf margin (Klucking leaf in the Claiborne flora. Pollen of Platanus is re- 1988). Pollen of Myrtaceae (Myrtaceidites)is recorded corded from the upper Claiborne of the Mississippi from the upper Claiborne of Mississippi and Alabama Embayment (Frederiksen 1980a, 1988). (Gray 1960; Engelhardt 1964; Frederiksen 1980a, 1988). Material examined.-UF 1 88 10-34439. Material examined.-UF 18810-34386.
Saxifragales Fabales Altingiaceae Fabaceae Liquidambar sp. Cladrastis-\ike sp. (Fig. 18, D, H) (Fig. 10, A-C) Specimen description.-Fruit laterally compressed, Description.-Lamina of leaflet lanceolate, sym- subglobose, 2 cm diameter. Convoluted carbonaceous metrical to slightly asymmetrical, 6.5-9.5 cm long by 2- material with scattered circular impressions ofstyle bases 2.3 cm wide, length/width ratio 3.3:1 -4.1:1; apex angle with, or without acute impressions of style tips. acute; base angle obtuse, with rounded flanks; margin Discussion.-Impressions in the fruit appear simi- entire. Petiolule 3 mm long, with cross striations. Vena- lar to Liquidambar slyraciflua L before dehiscence tion pinnate; secondary venation brochidodromous, 9- 10 of the basal portion of the persistent styles (Godfrey pairs of secondary veins, angle of divergence 30-60°, 1988). Liquidambar fruit have been recorded from the secondary veins straight or arching from midvein to near Claiborne flora but were not described or figured (Berry margin then arching up to superadjacent vein. One to 1930). Pollen of Liquidambar is recorded from the two intersecondary veins per intercosta. Midvein deeply upper Claiborne of Alabama (Gray 1960). impressed. Tertiary veins orthogonal. Excostal vena- Material examined.-UF 18810-34432. tion in 1-2 series. Discussion.-The size, shape, apex, and venation Myrtales are similar to Sapindus (Sapindaceae) and Cladrastis Myrtaceae (Fabaceae) but in Sapindus the base is cuneate, curved, Myrcia sp. and asymmetrical (UF335, 340, 343, 345, 352; Herendeen (Fig. 10, F-H) 1992; Yu & Chen 1991). The venation and size of the Specimen description.-Leaf incomplete, lamina smaller leaflets are similar to Gleditsiophyllum possibly elliptic, - 7 cm long by 3.2 cm wide; apex and eocenicum Berry (Berry 1916; Herendeen 1992). Ex- base missing; margin entire. Petiole missing. Midvein tant species of Cladrastis have a temperate distribution stout; venation pinnate; secondary veins narrowly spaced, (Herendeen 1992). angle of divergence 70-80°, straight from midvein to Material examined.-UF 18810-34402, 34413a, DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 13
34421,34428. Discussion.-Leaflet petiolules with prominent striations are found in Fabaceae and Connaraceae, but Ormosia-like sp. 1 the shape of this leaflet is similar to extant Ormosia (Fig. 11 , A-C) ( Fabaceae) such as Ormosia nitida Vog. and 0. Description.-Lamina of leaflet elliptic, symmetri- monosperma (Sw.) Urb. (Rudd 1965). The shape, base, cal, 11 cm long by 4.7-5.6 cm wide; apex angle acute, margin, and venation is similar to fossil Ormosia sp. 2 with acuminate flanks; base angle acute, with convex (Herendeen 1992). Extant species of Ormosia can be flanks; margin entire. Petiolule missing. Venation pin- found in the tropics ofAsia, Australia, Madagascar, and nate; secondary venation weak brochidodromous, 6-8 America (Herendeen 1992). pairs of secondary veins, angle of divergence 40-60°, Material examined.-UF 18810-34395. secondary veins alternate, broadly arching from the midvein to near the margin then arching sharply and di- Swartzia sp. rected medially toward midvein joining superadjacent (Fig. 7, D-H, J, L; Fig. 13, A, B) vein. Weak basal secondary veins, and 1 -3 weak to Description.-Lamina of leaflet narrowly elliptic strong intersecondary veins per intercosta. Tertiary veins to elliptic, symmetrical, 3.7-8 cm long by 1.3-4 cm wide, percurrent, straight to slightly sinuous, unbranched to length/width ratio 2:1 - 2.8:1; apex angle acute to ob- branched, tertiary angle obtuse. tuse, retuse; base angle acute to obtuse, with convex to Discussion.-The absence of a petiolule makes concave flanks; margin entire. Petiolule missing. Ve- the determination to family difficult because the leaflets nation pinnate; secondary venation strongly of legumes frequently have short, wrinkled appearing brochidodromous, angle of divergence 50-60°, second- petiolules. The general shape, size, apex, margin, and ary veins alternate, with 5-6 pairs, straight from midvein venation ofthese specimens suggest leaflets in the fami- to near margin, then arching upwards. One pair of weak lies Fabaceae and Connaraceae. These two families acute basal secondary veins, and 1-2 strong seem very close in the morphological characters oftheir intersecondary veins per intercosta. Tertiary veins or- leaflets, although in Connaraceae the tertiary venation thogonal reticulate. Associated pod incomplete (UF is generally more prominent as in Connarus 34419a, not figured), apex missing, base curving with fasciculatus (de Candolle) Planchon (Forero 1983) and basal portion missing. Parallel oblique striations near Rourea minor (Gaertn.) Alston (Jongking 1989). Within base, marginal suture well developed. Fabaceae, this fossil leafiet morphology and venation Discussion.-Although the morphology and vena- are similar to Swartzia racemosa Benth. (FLAS 165336) tion are similar to numerous species of Swartzia and Ormosia grandiflora (Tul.) Rudd although in S. (Fabaceae) such as Swartzia parvifolia Schery, S racemosa the margins have a few teeth. Further study apetala Raddi, and S. corrugata Bentham (Cowan of the anatomy would help to clarify the generic and 1968), future anatomical work is needed to confirm this species determination of these fossils. At present the determination. The shape, size, margin, apex, base, and leaflets appear most similar to Ormosia sp. 1 (Herendeen venation ofthe leaflets, and pod shape and size are simi- 1992; Fig. 119) and so are placed in that genus. lar to the fossil species Swartzia sp. 1 (Herendeen 1992), Material examined.-UF 18810-34404a, 34406. although the leaftet determination was based mainly on anatomical characters. Fabaceous pollen is recorded in Ormosia-like sp. 2 the upper Claiborne of the Mississippi Embayment (Fig. 7, I, K) (Frederiksen 198Oa). Extant species of Swartzia are Specimen description.-Leallet incomplete, lamina distributed in the Neotropics and Africa (Herendeen narrowly elliptic, symmetrical, - 6 cm long by 1.4 cm 1992). wide, length/width ratio-4.3:1; apex missing; base angle Material examined.-UF18810-34388a, 34391, acute, with slightly convex flanks; margin entire. Peti- 34393,34394,34419a, 34420. olule prominently cross-striated, 0.8 mm long by 0.15 mm wide. Venation pinnate; secondary venation Fagales brochidodromous, angle of divergence 30-40°, second- Fagaceae ary veins alternate, extending straight from midvein to Berryophyllum tenuifolia (Berry) Jones et margin or curving abruptly upward near the margin. One Dilcher pair of weak acute basal secondary veins, and 1 -2 (Fig. 17, H; Fig. 18, B) intersecondary veins per intercosta. Tertiary veins Specimen description.-Leaf incomplete, lamina percurrent. Quaternary veins random and reticulate. linear, symmetrical, - 9 cm long by 0.4 cm wide at 14 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1) middle; apex missing; base angle acute, with cuneate Description.-Lamina narrowly elliptic, symmetri- ftanks; margin serrate for most of leaf length. Petiole cal,- 11-13 cm long by 1.8-2.5 cmwide; apex and base missing. Venation pinnate; secondary venation angle acute, with cuneate flanks. Margin serrate and craspedodromous, angle of divergence 40°, secondary confined to the middle and distal half of the leaf; teeth veins alternate, straight to leafmargin, one entering the short, less than 1 mm, basal flank convex, apical flank tooth, the adjacent vein running alongside leaf margin. concave. Petiole missing. Venation pinnate; secondary Discussion.-The shape, size, margin, and vena- venation camptodromous to semicraspedodromous, angle tion are similar to leaves of Fagaceae, and the fossil of divergence 40-60°, secondary veins alternate, arch- species Banksia tenuifolia proposed by Berry which ing gradually from midvein to near margin then joining was renamed Berryophyllum tenuifolia (Berry) Jones the superadjacent vein. One to two intersecondary veins et Dilcher (Berry 1916; Jones & Dilcher 1988). per intercosta. Tertiary veins orthogonal, tertiary veins Fagaceous pollen (Quercoidities, Cupuliferoi- arising from a secondary marginal loop enter the teeth. pollenites) is recorded in the upper Claiborne of the Ultimate veins freely ending and one branched. Mississippi Embayment (Frederiksen 1980a, 1988) and Discussion.-The shape, size, apex, base, margin, from the Lawrence ( Quercus ) and Warman and venation are similar to Knightiophyllum (Quercoidities) clay pits (Elsik & Dilcher 1974; Potter wilcoxianum Berry (Berry 1916; Dilcher & Mehrotra 1976). 1969) except in this material the teeth are only in the Material examined.-UF18810-34389. distal half of the lamina. The systematic placement of this genus appears to be with Fagaceae but no modern Berryophyllum sp. genera have been found that are similar to K. (Fig. 17, E-G) wilcoxianum (Dilcher & Mehrotra 1969). Morphologi- Description.-Leaves incomplete, lamina narrowly cal features such as leaf margin with a few teeth, gen- elliptic, symmetrical, - 20 cm long by 2 cm wide at middle; eral looping of the secondary veins, tertiary vein from apex and base missing; upper 25% of leaf attenuate, secondary looping entering teeth, and intersecondary lower 25% of leaf cuneate. Margin serrate in upper veins common, are characteristic of Fagaceae, notably 75% of leaf; teeth sharp, prominent, 1 mm long, basal Quercus section Erythrobalanus (Jones 1984), near side straight to convex, apical side concave. Petiole Quercus viminea but the teeth are not as prominent as missing. Venation pinnate; basal secondary venation in Q. viminea (Villarreal 1986). camptodromous, secondary venation craspedodromous Material examined.-UF 18810-34424, 34425a in mid and upper portions of leaf, angle of divergence 50-60°, secondary veins alternate, arching gradually from Malpighiales midrib to margin, one vein entering each tooth, adjacent Salicaceae 1 -2 veins running parallel to margin. Secondary veins Populus sp forked just below tooth, of equal strength, one entering (Fig. 16, D; Fig. 17, A) tooth medially and the other running parallel to margin. Specimen description.-Leaf incomplete, lamina One to two intersecondary veins per intercosta. Ter- elliptic, symmetrical, - 6 cm long by - 3 cm wide; apex tiary veins percurrent, simple to branched. Quaternary and base missing. Margin serrate/dentate; teeth apex veins random. obtuse (they appear to be glandular) to acute, sinus Discussion.-The leaf characters seem to be in- rounded. Petiole missing. Venation pinnate; secondary termediate between Berryophyllum warmanense Jones venation semicraspedodromous, with one et Dilcher and B. saffordil (Lesq.) Jones et Dilcher. subactinodromous vein, angle ofdivergence 60° basally, The tooth shape and the way the secondary veins enter 40-50° apically. Intersecondary veins per intercosta one the teeth are characters that match B. warmanense, 0r absent. Tertiary veins not well preserved. while the leafsize, margin, and secondary venation match Discussion.-Although the leafis incomplete, and B. saffordii (Jones & Dilcher 1988). It has been sug- tertiary veins are not well preserved, characters such as gested that Trigonobalanus and Colombobalanus may shape, subactinodromous and semicraspedodromous be related extant genera that are found in tropical Asia, venation, and toothed margins are similar to extant leaves and the mountains ofColombia (Dilcher 200Ob). of Populus ( Salicaceae ; Eckenwalder 1980) and Material examined.-UF 18810-34396, 34408a. "Tiliaceous" species Dicraspidia donnell-smithii Standley (Robyns 1964) and young seedlings of Tilia Knightiophyllum wilcoxianum Berry americana L. (UF6174). In D. donnell-smithii and T. (Fig. 16, B, C, G) americana, the secondary venation is mostly DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 15 craspedodromous, the teeth lack glands, and the sinus Ficus-like sp. of the teeth are angular. The shape of some forms of (Fig. 8, C, D) Cercidiphyllum japonicum Siebold & Zucc. with Specimen description.-Lamina elliptic, symmetri- toothed margins, and teeth with a rounded sinus is simi- cal, 9.5 cm long by 3.7 cm wide; apex angle obtuse, with lar to the fossil, but in C japonicumthe secondary ve- convex flanks; base angle acute, with concave ftanks; nation is brochidodromous (UF 1965). The presence of margin entire. Petiole missing. Venation pinnate; sec- glandular teeth is characteristic of Populus. The vena- ondary venation brochidodromous, 14 pairs of second- tion and teeth are similar to middle/upper portions of ary veins, angle of divergence basally 60-70°, apically Grewiopsis tennesseensis Berry (1916) which is con- 40-50°, secondary veins subopposite basally to alternate sidered similar to Populus cinnamomoides (Lesq.) apically, straight from midvein to near margin, then arch- MacGinitie (Eckenwalder 1977, 1980). The fossil de- ing upward parallel to margin, then arching medially join- scribed here may be an intermediate form between a ing superadjacent vein, also a prominent secondary loop linear juvenile and deltoid adult leaf of Populus above this lower loop. One pair of low angled basal (Eckenwalder 1980). secondary veins present, 1-3 strong to weak Material examined.-UF 18810-34397. intersecondary veins per intercosta, extending to less than !/2 distance to margin. Excostal veins in 2-3 series. Rosales Tertiary veins reticulate. Moraceae Discussion.-The general shape and size, apex, Pseudolmedia sp. base, margin, brochidodromous venation with high angled (Fig. 14, B, D) basal secondary veins, 1-3 strong and weak Description.-Lamina obovate, slightly intersecondary veins, and reticulate tertiary veins are inequilateral,-14 cm long by 6.3 cm wide; apex angle characteristic to the families Moraceae and acute (not illustrated, UF 34425b); base angle acute, with Lecythidaceae, although in Lecythidaceae the asymmetrical flanks; margin entire. Petiole missing. intersecondary veins can extend to more than M leaf Venation pinnate; secondary venation brochidodromous, distance from midrib to margin, and they lack a pair of - 14 pairs of secondary veins present, angle of diver- basal low angle secondary veins. In Moraceae, the low- gence 40-50°, secondary veins alternate, straight from ermost basal secondary vein angle is different frpm other midvein to near margin then arch sharply upward joining basal secondary veins, and a prominent secondary loop superadjacent veins. Basal pair of secondary veins, over the junction of the secondary vein with the occasional 1-2 intersecondary veins per intercosta usu- superadjacent vein is evident in many species. Leaf ally extending less than ~/2 the distance from midrib to shape, size, base shape, and venation ofthis specimen is margin. Tertiary veins alternate percurrent and forked, similar to the fossil Ficus puryearensis Berry (1916) at right angles to secondary veins. Excostal veins and extant Ficus (Burger 1977), although in Berry's uniseriate, close to leaf margin. description the secondary venation angle is nearly 90° Discussion.-The large size, margin, and the basal while the figures (Plate XXVIII) are closer to our mea- pair of secondary veins are typical characters of surements. The secondary vein angle is in the low range Moraceae. The combination ofcharacters such as shape for extant Ficus. Ficus pollen has been recorded in the of the apex and base, margin, number and angle of sec- upper Claibome ofAlabama (Gray 1960). Berry's (1916, ondary vein pairs, intersecondary veins, and percurrent 1930) floras included an excessive number of species in tertiary veins are common in Pseudolmedia. These the genus. A detailed re-evaluation ofthe genus is needed characters also can be found in Naucleopsis but in in order to evaluate the real nature of all the so-called Naucleopsis uie percurrent tertiary veins are in a dis- Ficus leaves from the Claiborne of southeastern North tinct double layer in the intercostal area (Berg 1972; America. FLAS 132816; UF1483). Berry (1916) recognized Material examined.-UF18810-34446. Pseudolmedia inthe Wilcox flora (now Claibome Flora) of Tennessee but the leaves he described are narrower Rhamnaceae and the secondary veins arise from the midvein at a 90° Berhamniphyllum claibornense Jones et angle. The extant genus Pseudolmedia is distributed in Dilcher the Neotropics (Berg 1972; Burger 1977). (Fig. 12, A, B) Material examined.-UF 18810-34417, 34425b. Specimen description.-Leaf incomplete, lamina narrowly elliptic, symmetrical, - 11 cm long by 4 cm wide; apex angle acute, with cuneate flanks; base miss- 16 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1) ing; margin entire. Petiole missing. Midvein straight; Sapindales venation pinnate; secondary venation eucamptodromous, Sapindaceae angle of divergence 20-30°, secondary veins opposite Cupanites sp. basally to alternate apically, straight to slightly curved (Fig. 17, B, D) from midvein to near margin then arching abruptly up- Specimen description.-Leaftet incomplete, lamina ward to the margin. Secondary veins rarely divided near narrowly oblong, asymmetrical, - 7 cm long by 2.1 cm the margin. Tertiary venation barely perceptible, wide; apex missing; base angle acute, with convex and percurrent, straight, perpendicular to the midrib. asymmetrical flanks. Margin with a few serrate teeth Discussion.-Leaf venation characters such as from middle to apical portion of leaf. Petiole - l mm secondary venation eucamptodromous and percurrent long and expanded at base. Midvein stout; venation pin- tertiary veins perpendicular to the midrib, are character- nate; secondary venation semicraspedodromous, angle ~ istic ofRhamnaceae. The general shape, apex, margin, of divergence 50-60°, secondary veins alternate to sub- and venation are similar to the fossil leaves opposite, straight from midvein to near margin, then di- Berhamniphyllum claibornense Jones et Dilcher (Jones vided, one continuing toward margin and teeth, one arch- & Dilcher 1980). Berhamniphyllum claibornense is ing upward joining superadjacent vein. One to two closely related to the tribe Rhamneae (Richardson et al. intersecondary veins per intercosta. Tertiary veins at 2000), and possibly the extant genera Berchemia and right angle to secondary veins. Excostal veins in one Rhamnidium (Jones & Dilcher 1980) that have prima- series. rily tropical distributions (Record 1939; Brizicky 1964). Discussion.-The general shape and size, margin The fossil leaves have frequently been reported as Rham- with serrate teeth, asymmetrical base, and venation is nus however identifications to extant genera can not be similar to leaflets of Sapindaceae and Juglandaceae, al- validated without fruiting material, even ifperfectly pre- though in Juglandaceae the secondary vein divides just served leaves complete with cuticular material (even before entry into the tooth, one entering the tooth, the extant leaves) are preserved. A new genus was pro- other going towards the sinus. The leaflet is similar to posed by Jones and Dilcher (1980) to indicate that only the fossil species Cupanites (Berry 1916, 1930) and the family Rhamnaceae and tribe Rhamneae can be extant Cupania glabra (Standley & Steyermark 1949; determined based upon leaf material. UF310), although in Berry's Cupanites, the venation is Material examined.-UF18810-34384. craspedodromous and the teeth extend along the full length of the leaf. Cupania-like pollen (Cupanieidites) Berhamniphyllum sp. and megafossils are recorded in the upper Claiborne of (Fig. 9, D, (1 H) Mississippi andAlabama (Gray 1960; Engelhardt 1964; Specimen description.-Lamina elliptic, symmetri- Frederiksen 1980a, 1988). Distribution of extant cal, 6.3 cm long by 3 cm wide; apex angle acute, retuse; Cupania is Neotropical (Standley & Steyermark 1949). base angle acute, with convex flanks; margin entire. Material examined.-UF18810-34401. Petiole missing. Venation pinnate; secondary venation eucamptodromous, angle of divergence 40°, basal sec- Cornales ondary veins opposite, becoming alternate towards the Nyssaceae apex, secondary veins straight to slightly curved as they Nyssa eolignitica Berry extend from the midvein then arch abruptly upward to (Fig. 18, G, K) near the leaf margin. Specimen description.-Endocarp ovate, 2 cm long Discussion.-Although the lack of tertiary vena- by 0.8 cm wide, apex acute, base rounded. Fruit trilocu- tion creates difficulty in the identification of this fossil, lar with numerous longitudinal ridges on the surface. the general shape and size, base, margin, and secondary Discussion .-This fruit is similar to Nyssa venation is very similar to Berhamniphyllum eolignitica Berry (Berry 1916; Dilcher & McQuade claibornense Jones et Dilcher (Jones & Dilcher 1980). 1967) and N. wilcoxiana Berry ( 1916, 1930). Nyssa- The apex is retuse, which is not in the species descrip- like pollen (Nyssapollenites) is recorded in the upper tion of B. claibornense (Jones & Dilcher 1980), but is Claiborne of Mississippi and Alabama (Gray 1960; found in species of Rhamnidium such as R. eliipticum Engelhardt 1964; Frederiksen 1980a). Distribution of Britton et R Wilson and R. shaferi Britton et R Wilson the extant genus Nyssa is temperate, cool/warm tem- (Britton 1915). perate of the United States and China to the tropics of Material examined.-UF 18810-34427. Mexico/Central America, East Asia and Malaysia/Indo- nesia (Dilcher 200Ob). DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 17
Material examined.-UF18810-34392 Discussion.- The flower is tentatively allied with the leaves, Ternstroemites. However, a total re- Ericales study ofthe fossil record ofthis complex is in progress. Theaceae Grote and Dilcher (1989) described fruits that are also Ternstroemites sp. related to these leaves and flowers . One species, (Fig. 16, E, F; Fig. 17, C, I; Fig. 18, A, E) Andrewsiocarpon henryense Grote & Dilcher was Description.-Lamina mostly elliptic to narrowly recognized as totally extinct today but closely related to elliptic, sometimes obovate, symmetrical, 5 .8- 14.5 cm Franklinia alatamaha Marsh. The leaves and flow- long by 3.2-3.3 cm wide, length/width ratio 2.6: 1 - 4.4: 1 ; ers mentioned here form part of this complex and are apex angle acute to obtuse, with emarginate to acumi- typical of G lasianthus found in the swamp forests of nate flanks; base angle acute, with cuneate to decurrent north Florida today. The flower width of G lasianthus flanks. Margin crenulate to serrate to near the base; is 8 cm including petals, absent in the fossil (Godfrey some teeth with glands (UF34405). Petiole 2 cm long 1988), 3-4 cm in Ternstroemia sp. (Kobuski 1942a,b, and expanded at base. Midvein stout, - 1 mm wide at 1943), and 2 cm in Adinandra parvifolia Ridley middle section ofleaf. Venation pinnate; secondary ve- (Kobuski 1947). A similar flower (Antholithes nation camptodromous to semicraspedodromous, 9-10 ternstroemioides Berry), slightly larger than our speci- pairs of secondary veins, angle of divergence 40-60°, men, was reported by Berry (1930) as similar to G secondary veins alternate to subopposite, straight to lasianthus. slightly curved from midvein dividing 1 -2 times before Material examined.-UF18810-34438. joining superadjacent vein. Excostal veins in two series, some terminating in crenulate to serrate teeth. One to Gentianales two intersecondary veins per intercosta. Tertiary veins Apocynaceae percurrent and forked. Apocynophyllum sp. Discussion.-The shape and size, venation, and (Fig. 11, D, E) presence of glandular teeth are characteristic of Specimen description.-Leaf incomplete, lamina Theaceae and Ternstroemiaceae, but whether the glands elliptic, symmetrical, - 12 cm long by - 4.5 cm wide; are deciduous (Ternstroemiaceae) or permanent apex missing; base angle acute, with slightly decurrent (Theaceae) is not evident in the fossil specimens flanks; margin entire. Petiole width at blade base 1 mm, (Weitzman et al. 2004; Stevens et al. 2004). Characters width at petiole base 2 mm. Venation pinnate; second- such as venation, crenate to serrate margin, and glandu- ary venation brochidodromous, angle of divergence 40- lar teeth are similar to extant Gordonia lasianthus (L.) 50°, secondary veins widely spaced, straight from Ellis (Theaceae, UF5191). Overall, these characters midvein to near margin, then arching upward and medi- can be found in Eurya, Adinandra and Ternstroemia ally to superadjacent vein. Tertiary veins reticulate. (Ternstroemiaceae, Kobuski 1939, 1942a,b, 1943,1947), Discussion.-The incomplete and poorly preserved although not in any one particular species. For exact nature ofthis specimen makes further determination dif- systematic placement of the fossil Ternstroemites sp., a ficult. The leaf is similar to the acute basal form of leaf venation comparison study is needed for Theaceae Apocynophyllum mississippiensis Berry, except for the and Ternstroemiaceae. A detailed study of the leaves widely spaced secondary veins and reticulate tertiary of Gordonia and related genera is in progress by Dilcher veins (Berry 1916, 1930; Dolph 1975). Dolph ( 1975) and Wang. Distribution of extant species of G lasianthus undertook an extensive study of Apocynophyllum leaf and Ternstroemia is warm temperate to predominately types from the Claiborne sediments of several clay pits Neotropical (Kobuski 1942a,b, 1943; Dilcher 200Ob). in western Tennessee and Kentucky. He recognized Material examined.-UF 18810-34390, 34399, two distinct subspecies of this extinct genus and after 34405,34426,34429. extensive research has been unable to ally it systemati- cally with any extant family. d. Gordonia sp. Material examined.-UF18810-34441. (Fig. 18, C, I) Specimen description.-Flower, - 1.8 cm by - Rubiaceae 5 mm basal width, 5 sepals, the lobes with 3-5 longitudi- 4 Paleorubiaceophyllum sp. nal striations per 1 mm. Petals and carpels missing. (Fig. 9, A, B) About 46 stamen impressions in 2-3 series, ovary 3.5 Specimen description.-Leaf incomplete, lamina mm basal width. elliptic, symmetrical, - 7.5 cm long by 3.5 cm wide; apex 1 8 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1) missing; base angle obtuse, with short cuneate flanks; ondary veins opposite to alternate, straight from midvein margin entire. Petiole incomplete, - 5 mm long. Midvein to near margin, then arching upward to margin. One to stout; venation pinnate; secondary venation two intersecondary veins per intercosta, traversing up brochidodromous, angle ofdivergence 50-60°, more than to 1/2 distance to leaf.margin. Tertiary veins at right 10 pairs of secondary veins, sub-opposite to alternate, angle to secondary veins, straight and divide to adjacent straight from midvein to near margin then arching abruptly secondary. upward to superadjacent vein. One to two Discussion.-An acute and asymmetrical leaf intersecondary veins per intercosta. Excostal veins in base, plus a slender petiole are traits of Oleaceae and one series and looped. Tertiary veins inconspicuous. many other families. Further samples are needed for a Discussion.-The incomplete nature of this leaf clear indication ofthe distal portion ofthe leaf for char- makes our determination uncertain but the general shape acterization and possible generic determination. Cuticular and size, base, margin, and secondary venation are simi- characters might also be helpful. Berry (1916) assigned lar to Paleorubiaceophyllum eocenicum (Berry) Roth leaves to Fraxinus and Osmanthus, while Berry ( 1916) et Dilcher (Roth & Dilcher 1979). Additional speci- and Call and Dilcher (1992) described samaras of mens are needed to determine i f the petiole does have a Fraxinus. Oleaceaous pollen (Salixipollenites) is re- stipule attached to its base. This feature is common to corded in the upper Claiborne of the Mississippi all species of Paleorubiaceophyllum. Embayment (Frederiksen 1980a, 1988). Materialexamined.-UF18810-34431 Materialexamined.-UF18810-34430
Apiales Incertae sedis Araliaceae Entire Margin Morphotype 1 Dendropanax eocenensis Dilcher et Dolph (Fig. 9, C, E, F) (Fig. 6, A-D) Specimen description.-Leaf incomplete, lamina Description.-Lamina obovate, palmately 3-4 narrowly elliptic, symmetrical, - 5 cm long by 2.1 cm lobed, 5.4-6.7 cm long by 5.8-8.5 cm wide, lobes entire wide; apex and base missing; margin entire. Petiole and sometimes with secondary lobes; apex angle acute, missing. Midvein stout; venation pinnate; secondary with cuneate flanks; base angle acute, with concave venation brochidodromous, angle ofdivergence 40-50°, flanks. Petiole missing. Venation actinodromous, single - 10 pairs of secondary veins mostly alternate, arching vein entering each lobe; secondary venation interior, basal from midvein to near margin, then arching upward and sub-primary vein running along margin of basal lobe. parallel with the margin, then arching medially to join the Tertiary venation reticulate. superadjacent vein. One to three weak to strong Discussion.-The shape and size, lobing, and ve- intersecondary veins per intercosta, the strong vein ex- nation are similar to Dendropanax eocenensis Dilcher tending /2 to % width ofleafbefore dividing andjoining et Dolph (Dilcher & Dolph 1970) except for the lobing secondary veins. Excostal veins in one series. Tertiary of one specimen that has four lobes, while the speci- veins at right angle to secondary veins, joining mens described as D. eocenens is have three or five intersecondary and secondary veins. lobes. Rare Araliaceous pollen is recorded in the Discussion.-The lack ofan apex and base makes Cockfield Formation of Mississippi (Engelhardt 1964). determination to any known leaf type difficult. The The distribution of extant Dendropanax is tropical lamina of the fossil leaf appears to have a slight V-con- America and eastern Asia (Mabberley 1989). figuration in which the midvein is depressed. In the speci- Material examined.-UF18810-34380, 34382a, men figured in Figure 9, C, E, and F, we see the abaxial 34383. side of the leaf so the midvein appears to stand out and the lamina is an inverse V-configuration. Overall shape, Lamiales and venation are similar to those of extant Maranthes Oleaceae leaf morphotype (Chrysobalanaceae) (Prance 1972 ), Allophylus, (Fig. 11 , F; Fig. 12, F) Deinbollia, Meliococca, Scyphonychium, and Talisia Specimen description.-Leaf incomplete, lamina (Sapindaceae) (Standley & Steyermark 1949), and the possibly elliptic, symmetrical, - 7.5 cm long by 3.5 cm fossil Chrysobalanus inaequalis (Lesq.) Berry (Berry wide; apex missing; base angle acute, with asymmetri- 1916). cal and slightly decurrent flanks; margin entire. Petiole Material examined.-UF 18810-34433. slender, 1 mm wide. Venation pinnate; secondary vena- tion eucamptodromous, angle of divergence 60°, sec- Entire Margin Morphotype 2 DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 19
(Fig. 8, A, B) Specimen description.-Leaf incomplete, - 2 cm Description.-Lamina ofleallet elliptic, symmetri- wide; apex angle acute, with attenuate flanks; base miss- cal, - 6.5-9.5 cm long by 2.5-3.2 cm wide; apex angle ing. Margin variable, midsection of leaf serrate, apical acute, with acuminate flanks; base angle acute, with section entire. Venation pinnate; secondary venation convex flanks; margin entire and undulate. Petiole miss- variable, midsection venation semicraspedodromous, ing. Venation pinnate; secondary venation apical section brochidodromous, angle ofdivergence 40- eucamptodromous, angle ofdivergence 40-50°, second- 50°, secondary veins alternate, arching slightly from ary veins mostly alternate, arching from midvein to very midvein to near margin, secondary vein divides at base near margin, extending to but not merging with the mar- of tooth, one branch entering tooth, other branch joins ginal vein or one areole loop away from the margin. superadjacent vein as a tertiary vein. One to two Secondary veins connected to superadjacent veins by a intersecondary veins per intercosta. Tertiary veins series oftertiary cross veins. One or two intersecondary percurrent and forked. veins per intercosta, intersecondary veins traversing up Discussion.-The fragmentary nature of this to % distances to the leaf margin. Tertiary veins reticu- specimen limits the characters available for systematic late. placement ofthis leafmorphotype. However, the shape, Discussion.-Our specimens lack a petiole which margin, apex, and secondary venation are similar to Ilex results in fewer characters useful in systematic place- species living in the swamps near Gainesville, Florida. ment. The general shape and size, apex, base, margin, Material examined.-UF18810-34445. and venation are similar to Aegiphila (Lopez-Palacio 1977) in the Verbenaceae but petiole characters are Reproductive Structure Morphotype 1 important. The Anacardiaceae may have leaves or leaf- (Fig. 18, F, J) lets with revolute or undulate margins that also have Specimen description.-Reproductive structure secondary veins that arch very close to the leaf of leaf- elliptic, 11 mm long by 5 mm medial width. Apex short let margins. acuminate, base obtuse. Numerous closely spaced lon- Material examined.-UF18810-34444a, b, 34412. gitudinal striations or veins, eight near the base dividing to 20 at the midpoint and then rejoining adjacent veins or Entire Margin Morphotype 5 striations distally. (Fig. 15, B, D) Discussion.-Berry (1916) placed this type of Specimen description.-Leaf incomplete, lamina structure in Avicennia eocenica Berry but the system- elliptic, symmetrical, -12 cm long by 5.5 cm wide; apex atic placement was not clearly demonstrated. Avicennia and base missing; margin entire. Petiole missing. Ve- fruits are much larger, obpyriform or ovate in shape, and nation pinnate; secondary venation brochidodromous, - the striations are widely spaced. The characters of this 7 pairs of secondary veins, angle of divergence 30-40°, fossil do not match Avicennia. This structure appears secondary veins alternate, arching slightly to straight from to be the remains of a fruit that is presently undeter- midvein to near margin, then arch medially joining mined. superadjacent vein, with further weaker vein looping Material examined.-UF18810-34434. above. Excostal veins in one series. Intercosta ofbasal portion of leafwith 1 -2 mostly weak intersecondary veins Minor Faunal Elements extending less than '/2 distance to leafmargin. Intercosta Insecta of middle and apical portion of leaf with 1-3 Trichoptera intersecondary veins extending up to /2 distance to leaf Insect cases margin. Tertiary veins weak and appear percurrent. Folindusia sp. Discussion.-The incomplete set ofcharacters for (Fig. 19, D, G) this specimen makes it difficult to place systematically. Specimen description.-Case straight, 3 cm long, The shape and size, margin, and venation are similar to width of posterior end 3 mm, width of anterior end 8 species such as Gonzalagunia dicocca Cham. & mm. Six transverse leaf pieces cover the case, 2-8 mm Schltdl. (Rubiaceae), Markea coccinea Rich. (Solan- in length. aceae) *lori et al. 2002), or Magnolia (Magnoliaceae). Discussion.-The use of leaf material to build the Material examined.-UF 18810-34448. insect case, the number of leafpieces, and the case size is similar to extant caddisfly larval cases (Trichoptera) Tooth Margin Morphotype 1 and fossil cases of Folindusia wilcoxiana Berry ( 1927) (Fig. 15 , E; Fig. 19, A) and Folindusia sp. (Johnston 1998). Such insect cases 2 0 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1) are typical of fresh water, shallow lakes. middle Eocene. The Powers Clay Pit flora, presented Material examined.-UF18810-34436. here, is just the beginning ofassessing the paleofloristics of this area, the individual plant fossils and their sedi- Terrindusia sp. ments. It is anticipated that a floristic database for each (Fig. 19, E, F) locality will be developed at the Florida Museum ofNatu-, Specimen description.--Case incomplete, 4 mm ral History, and made available to the public on the wide. Case covered with minute leaf pieces, woody Internet. Also, the plates of this initial report ofPowers material and sand grains. Clay Pit will be available at the Paleobotany and Pa- Discussion.-The use of sand grains to build the lynology web site (http://www. flmnh.ufl.edu/paleo- insect case is similar to that illustrated by Johnston (1998) botanyO. but in our specimen organic material is mixed with the The present discussion is centered upon the Pow- sand grains. ers Clay Pit fossils. This includes their relationship to Material examined.-UF18810-34437. 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- DISCUSSION lished upon leafmorphotypes and comparisons that can The Paleobotany and Palynology Collection ofthe Florida be made with leaf types of extant and fossil taxa. The Museum contains collections from clay deposits in west- authors feel it is better to publish the available data now ern Tennessee and Kentucky, beginning in 1959 with rather than to wait until detailed studies have been com- Dilcher's collections from Puryear. Dilcher continued pleted for each leaf type. Detailed studies o f individual to collect fossil plants from this area from the 1960s leaf types are labor intensive in the comparisons of fine onward with the help of assistants, students, colleagues venation and cuticle with living and fossil leaves. While and collaborators. Thousands of fossils, including leaves, this type of research is continuing and is included here fruits and seeds, flowers, wood, pollen, and spores have for those leaves already studied, we feel that the unique been collected from over 30 localities in western Ten- floristic nature ofthe individual localities ofthe middle nessee, western Kentucky, Mississippi, Alabama, Ar- Eocene is also of primary interest and should be pre- kansas, Texas and Louisiana. Among the most recent sented separately. collections are those from the Powers Clay Pit and Taxa described from the Powers Clay Pit flora nearby localities. This report is a presentation of the allow for character comparisons with other local middle Powers Clay Pit flora that is available in our collection Eocene localities. Detailed species accounts from Pow- at this time. It is the first of a series of reports, detailing ers Clay Pit have documented the first occurrence of a the nature ofthe floras from many ofthe clay pits with lobed Platanus leaf (Fig. 7, A-C), and a Liquidambar the best preserved and most complete collections offossil fruit (Fig. 18, D, H) that is described and illustrated. plants. Also, new leaf characters presented here enhance our As additional reports ofthe middle Eocene locali- knowledge of fossil leaf variation. An example is the ties become available, they will provide the data neces- intermediate form of Berryophyllum sp. where teeth, sary to ask many interesting and important questions margin, venation and leaf size demonstrates variations involving the deposition, evolution, distribution and not previously recognized (Jones & Dilcher 1988). Other paleobiology of plants from the Gulf Coast of North character variations include the leaf apex of America. The composition of these floras is expected Berhamniphyllum sp., and leaf shape in the intermedi- to have a direct link to the relationships between fossil ate form of Populus sp. Such variations in leaf form and modern floras of North America, Central America, were also found in other localities, along with variations northern South America, Africa and tropical Asia (Gra- in cuticular characters (Dilcher & Mehrotra 1969; Roth ham 1999; Dilcher 2001; Jaramillo & Dilcher 2001). & Dilcher 1979). These authors suggested that the rea- Many of these fossil localities given in figure 2 repre- son for these variations involved the possibility that each sent a similar depositional environment, but not all of fossil locality is of a specific age, with leaf characters them fall into the same category. There is a distinct changing through time, demonstrating microfeatures of probability that each locality represents its own specific evolution. Some variations may be related with sample time and climate. Therefore, each flora represents a size, so an increase in the number of specimens may unique part in the evolution ofplants, through change in widen the range ofvariation. Also, the differences noted the mixture of floristic elements, and a unique climate. between localities were consistent over a large number This could occur over a few million years in the upper ofspecimens from each locality (Roth & Dilcher 1979). DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 21
The Powers Pit flora also contains readily identifiable quianensis here, and in Berry ( 1916). Some leaves taxa based upon gross morphological features of the from Powers Clay Pit have an uncertain systematic megafossils. These include Berryophyllum tenuffolia placement, including Entire Margin Morphotype's 1,2, (Berry) Jones et Dilcher (Berry 1916; Jones & Dilcher 5, Tooth Margin Morphotype 1, and Reproductive Struc- 1988), Berhamaphyllum claibornense Jones et Dilcher ture Morphotype 1. This would include Knightiophyllum (Jones & Dilcher 1980), Nyssa eolignitica Berry (Berry wilcoxianum Berry where placement in Fagaceae is 1916; Dilcher & McQuade 1967), and Dendropanax open to question. Knightiophyllum wilcoxianum does eocenensis Dilcher et Dolph (Dilcher & Dolph 1970). not match any extant genus and represents an extinct Dilcher and Mehrotra (1969) found that consistent cu- member of Fagaceae. Specimens that are incomplete ticular differences could be Observed in and problematic in placement include Apocynophyllum Knightiophyllum leaves between localities but differ- sp., cf. Paleorubiaceophyllum, and Oleaceae, with ences were not apparent in the gross features of the Apocynophyllum and Paleorubiaceophyllum repre- leaves. senting extinct taxa (Dilcher 2001). A recurring problem when identifying extant taxa It has been over 145 years since the first investi- based upon fossil leaves is the convergence in foliar char- gation of fossil plants from the Claiborne flora acters in numerous plant families and genera (Berry (Lesquereux 1859) and about 90 years since the first 1914). Detailed analyses ofvenation patterns may help detailed locality to locality comparisons of these fossil alleviate this problem, such as differences in plants (Berry 1916). Since that time, fossil plants from intersecondary vein branching in Vochysiaceae and new sites have been collected and investigated, increas- Myrtaceae, which in turn helps to identify fossil Myrcia. ing the number of localities in western Tennessee, west- Intersecondary venation and basal secondary venation ern Kentucky, Mississippi and Alabama from 10 (Berry helps distinguish Moraceae from Lecythidaceae, lead- 1916) to 30. With the taxa presented here from Powers ing to the identification of Ficus. Analyses of second- Clay Pit, a preliminary comparison can be made with ary vein angles and ultimate venation helped distinguish the limited taxa that have been studied and identified extant Duguetia from Magnolia which lead to the iden- from other pits in western Tennessee and western Ken- tification of fossil Duguetia. The determining charac- tucky. Taxa shared with Powers Pit have been found at ter used to suggest that Berry's Magnolia fossil leaves the Puryear Clay Pit ( 15 taxa), Willbanks I Clay Pit ( 14 were really Duguetia (Annonaceae) was trichome type taxa), Lamkin Clay Pit (seven taxa), Warman Clay Pit (Roth 1981; Dilcher & Roth, unpublished data). Ter- (seven taxa), and Lawrence Clay Pit (six taxa) (Table tiary venation is important, such as in distinguishing similar 2). As our research continues, it is anticipated that new leaflet forms of Fabaceae from Connaraceae, where morphotypes will be discovered, increasing our under- overall tertiary venation is more prominent in standing of the floristics for each locality. Even though Connaraceae. Also, Smilax and Dioscorea leaves are the present data is preliminary, interesting comparisons very similar except that in Dioscorea the tertiary veins have been found. are distinctly opposite percurrent. Sometimes, further Of the 35 fossil plant taxa recognized from Pow- morphological studies are needed to distinguish between ers Clay Pit, the number of taxa that overlap range from two closely related families, such as Theaceae and 15 at Puryear Clay Pit to six at Lawrence Clay Pit. In Ternstroemiaceae where venation patterns may confirm other words, less than half the known taxa overlap 10- placement of Ternstroemites sp. within Theaceae. Other calities. Although Warman Clay Pit is only 400 meters morphological features are important in distinguishing north of Powers Clay Pit, the two sites seem to share between similar leaf forms. Extant Cladrastis and very few taxa. Lawrence is 8 KM away, but also shares Sapindus are very similar but the leaf base o f Cladrastis few taxa with Powers Pit. Wilbanks Clay Pit shares 14 is straight, while Sapindus has a curved, often taxa and is only 1 KM west of the Powers Clay Pit. inequilateral base, leading to the identification of fossil Puryear Clay Pit is about 25 KM north north-east, and Cladrastis. Morphological leaf characters may not be Lamkin is about 65 KM nofth of Powers Clay Pit. It is enough, ·so future anatomical studies are needed that interesting to note that distance does not seem to relate incorporate cuticular characters. Such characters are to similarity offloristic elements. This disparity could be important in determining Swartzia to the species level explained by sample size, environmental factors, or that since similar morphological and venation patterns are these clay pits may not represent one common age. present in numerous species of Swartzia. Use of cu- Certainly, more research needs to be done to under- ticular characters is also important for Ocotea even stand the floristic relationships between the clay pit 10- though this fossil leaf has been linked with extant 0. calities. 22 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
Table 2. Taxa shared with Powers Clay Pit, and five other clay pits.
Puryear Willbanks Lamkin Warman Lawrence
Ocotea Ocotea Ocotea Smilax? (Fossil type 5) Berryophyllum
Duguetia Duguetia Myrcia Berryophyllum Knightiophyllum
Smilax Smilax Ficus Knightiophyllum Berhamniphyllum
Myrcia Ormosia Berhamniphylium Berhamniphyllum Nyssa (Calyptranthes)
Cladrastis Berryophylium Ternstroemites Ternstroemites Paleorubiaceophyllum (Gleditsiophyllum)
Berryophyllum Knightiophyllum Apocynophyllum Paleorubiaceophyllum Dendropanaxyllum
Knightiophyllum Populus Paleorubiaceoyllum Dendropanax
Populus Berhamniphyllum (Grewiopsis)
Ficus Nyssa
Berhamniphyllum Ternstroemitesl (Rhamnus) (Gordonia)
Cupanites Apocynophyllum
Nyssa Paleorubiaceophyllum
Ternstroemites Dendropanax
Apocynophyllum Folindusial (Caddistly case)
Paleorubiaceophyllum
t, The relationships offossil specimens to extant gen- cal distributions include Liquidambar, Populus, Nyssa, era can give us a glimpse ofpast ecosystems. Many of Cladrastis, and Platanus. Berry ( 1916, 1924) concluded the fossil specimens from Powers Pit appear to be re- that the climate of the "Wilcox" (now Claiborne) flora lated to extant genera that have primarily subtropical or was warm tropical. Miranda and Sharp (1950), Sharp tropical distributions. These include Ocotea, Swartzia, ( 1951 ), Gray ( 1960) and Engelhardt ( 1964) compared Ormosia, Berryophyllum, Ternstroemites, Dendro- the Eocene flora to the eastern Mexican escarpment panax, Duguetia, Moraceae, Myrcia, and Berham- region and suggested a mixture of temperate and tropi- niphyllum. Fossil specimens related to extant genera cal conditions . Dorf ( 1960) considered the .climate as that have primarily cool to warm temperate to subtropi- tropical, while Axelrod (1966) and Dilcher (1973) con- DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 23
cluded that the climate was warm temperate to cool sin, Colorado. Rocky Mountain Geology, 38:45- subtropical or cooler temperate, and Dilcher (1973) sug- 71. gested that there were definite seasonal dry periods. Berg, C. C. 1972. Olmedieae, Brosimeae (Moraceae). Elsik (1974) suggested a tropical to subtropical climate FloraNeotropica, 7:1-228. while Wolfe (1975, 1978), Wolfe and Poore (1982), Berry, E. W. 1914. The affinitiesand distribution ofthe Frederiksen (198Ob, 1988) and Graham (1999) consid- Lower Eocene Flora of southeastern North ered a winter dry tropical to marginal humid subtropical America. Proceedings of the American Philo- climate. The fossil specimens presented here suggest a sophical Society, 53:129-250. subtropical climate that can accommodate warm tropi- Berry, E. W. 1916. The Lower Eocene floras of south- cal to cool temperate species. While the lowlands pro- eastern North America. United States Geologi- vided tropical plant species, possible uplands with an in- cal Survey Professional Paper, 91:1-481. fluence from the Appalachian area may have provided Berry, E. W. 1922. Additions to the flora ofthe Wilcox cool temperate species (Cain 1943; Dilcher 1973). The Group. United States Geological Survey Profes- unique mixture ofelements living in temperate, subtropi- sional Paper, 131-A: 1-21. cal, and tropical climates today suggests that they may Berry, E. W. 1924. The Middle and Upper Eocene represent parts of an ancient non-analogue climate. That floras of southeastern North America. United is, a climate that is unique in the characteristics of its States Geological Survey Professional Paper, 92:1- temperature and moisture that is found nowhere in the 206. world today. Plants that may be related to these fossils Berry, E. W. 1927. A new type of caddis case from the continue living today in various places and in slightly dif- Lower Eocene of Tennessee. Proceedings of the ferent climates as they have shifted their distributions United States National Museum, 71:1-4. and evolved various ecological tolerances to accommo- Berry, E. W. 1930. Revision of the Lower Eocene date present conditions. Wilcox flora of the southeastern States with de- scriptions of new species, chiefly from Tennes- ACKNOWLEDGMENTS see and Kentucky. United States Geological Sur- We would like to thank the following people for vey Professional Paper, 156:1-144. help in collecting specimens from Powers Pit, Hongshan Berry, E. W. 1937. Tertiary floras of eastern North Wang, Eizabeth Kowalski and Jonathan Block (Florida America. Botanical Review, 3:30-46. Museum ofNatural History), Michael Gibson (Univer- Berry, E. W. 1941. Additions to the Wilcox flora from sity of Western Tennessee), Mihai Popa (University of Kentucky and Texas. United States Geological Bucharest), and Prakhart Sawangchote (Suranaree Survey Professional Paper, 193-E:83-99. University ofTechnology). We also like to thank Steven Britton, N. L. 1915. Studies ofWest Indian plants-VII. Manchester, Deborah Matthews, and one anonymous Bulletin ofthe Torrey Botanical Club, 42 (9):487- reviewer for their valuable comments and suggestions, 517. and Jason McCuiston ofFranklin Industries/ H. C. Spinks Brizicky, G E. 1964. The genera ofRhamnaceae in the Clay Company for access to Powers Pit. This is Uni- southeastern United States. Journal of the Arnold versity ofFlorida Contribution to Paleobiology number Arboretum, 45:439-463. 556. Brown, R. W. 1939. Fossil leaves, fruits, and seeds of Cercidiphyllum. Journal ofPaleontology, 13 :485- 499. LITERATURE CITED Brown, R. W. 1940. New species and changes ofname APG II (Angiosperm Phylogeny Group II). 2003. An in some American fossil flora. Journal of the update ofthe Angiosperm Phylogeny Group clas- Washington Academy of Sciences, 30:344-356. sification for the orders and families offlowering Brown, R. W. 1944. Temperate species in the Eocene plants: APG II. Botanical Journal ofthe Linnean flora of the southeastern United States. Journal Sodiety, 141:399-436. ofthe Washington Academy of Sciences, 34:349- Axelrod, D. I. 1966. A method for determining the 351. altitudes ofTertiary floras. Palaeobotanist, 14:144- Brown, R. W. 1946. Alterations in some fossil and 171. living floras. Journal of the Washington Academy Barclay, R. S., K. R. Johnson, W J. Betterton & D. L. ofSciences, 36:344-355. Dilcher. 2003. Stratigraphy and megaftora of a Brown, R. W. 1960. Corkwood in the Eocene of the K-T boundary section in the eastern Denver Ba- southeastern United States. Journal ofPaleontol- 24 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
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plants from the Wilbanks Clay Pit (middle Eocene United States Geological Survey Professional Claiborne Formation, Western Tennessee) pale- Paper, 1448:1-68. ontological implications and educational applica- Godfrey, R. K. 1988. Trees, shrubs, and woody vines tions. MS, Department of Educational Studies, of northern Florida and adjacent Georgia andAla- University of Tennessee, Martin, 96 pp. bama. University of Georgia Press, Athens, 734 Dilcher, D. L., F. W. Potter, & W. L. Crepet. 1974. pp. Investigations of angiosperms from the Eocene Graham, A. 1999. Late Cretaceous and Cenozoic His- of North America: Juglandaceous winged fruits. tory of North American Vegetation North of American Journal ofBotany, 63:532-544. Mexico. Oxford University Press, New York, 350 Dolph, G. E. 1975. A statistical analysis of pp. Apocynophyllum mississippiensis. Palaeonto- Gray, J. 1960. Temperate pollen genera in the Eocene grahicaAbt. B, 151:1-51. (Claiborne) flora, Alabama. Science, 132:808-810. Dorf, E. 1960. Climatic changes ofthe past and present. Grote, R J., & D. L. Dilcher. 1989. Investigations of American Scientist, 48: 341-364. angiosperms from the Eocene of North America: Eckenwalder, J. E. 1977. North American cottonwoods a new genus of Theaceae based on fruit and seed (Populus, Salicaceae) of sections Abaso and remains. Botanical Gazette, 150: 190-206. Aigeiros. Journal of the Arnold Arboretum, Grote, R J., & D. L. Dilcher. 1992. Fruits and seeds of 58(3):193-208. Tribe Gordoniae (Theaceae) from the Eocene of Eckenwalder, J. E. 1980. Foliar heteromorphism in North America. American Journal of Botany, Populus (Salicaceae), a source of confusion in 79:744-753. the taxonomy of Tertiary leaf remains. System- Heald, S. V. 2002. Rhamnaceae. Pp. 600-603 in S. A. atic Botany, 5:366-383. Mori, G Cremers, C. A. Gracie, J. J. de Granville, Ellis, B., K. R. Johnson, & R. E. Dunn. 2003. Evidence S. V. Heald, M. Hoff& J. D. Mitchell, eds. Guide for an in situ early Paleocene rainforest from to the Vascular Plants of Central French Guiana. Castle Rock, Colorado. Rocky Mountain Geol- Part 2. Dicotyledons. Memoirs ofthe New York ogy, 38:73-100. Botanical Garden 76, Part 2. The New York Bo- Elsik, W. C. 1974. Characteristic Eocene palynomorphs tanical Garden Press, New York. in the Gulf Coast, U.S.A. Palaeontograhica Abt. Herendeen, P. S. 1992. The fossil history of the B, 149:90-111. Leguminosae from the Eocene of Southeastern Elsik, W. C., & D. L. Dilcher. 1974. Palynology and North America. Pp. 85-160 in R S. Herendeen age ofclays exposed in Lawrence Clay Pit, Henry & D. L. Dilcher, eds. Advances in Legume Sys- County, Tennessee. Palaeontographica Abt. B, tematics: Part 4, The Fossil Record. Royal Botanic 146:65-87. Garden, Kew. Engelhardt, D. W. 1964. Plant microfossils from the Herendeen, R S., & D. L. Dilcher. 1990a. Fossil Eocene Cockfield Formation, Hinds County, Mis- mimosoid legumes from the Eocene and Oligocene sissippi. Mississippi Geologic Research Papers of southeastern North America. Review of Bulletin, 104:65-95. Palaeobotany and Palynology, 62:339-361. Fisher, W L. 1964. Sedimentary patterns in Eocene Herendeen, R S., & D. L. Dilchen 199Ob. Diplotropis cyclic deposits, northern GulfCoast region. Kan- (Leguminosae, Papilionoideae) from the middle sas Geological Survey Bulletin, 169:151-170. Eocene of southeastern North America. System- Forero, E. 1983. Connaraceae. Flora Neotropica, 36: atic Botany, 15:526-533. 1-208. Herendeen, R S., & D. L. Dilcher. 1990c. Reproduc- Frederiksen, N. O. 1980a. Sporomorphs from the Jack- tive and vegetative evidence for the occurrence son Group (upper Eocene) and adjacent strata of of Crudia (Leguminosae, Caesalpinioideae) in the Mississippi and western Alabama. United States Eocene of southeastern North America. Botani- Geological Survey Professional Paper, 1084:1-75. cal Gazette, 151:402-413. Frederiksen, N. O. 1980b. Mid-Tertiary climate of Herendeen, R S., & D. L. Dilcher. 1991. Caesalpmia southeastern United States: The sporomorph evi- subgenus Mezoneuron (Leguminosae, dence. Journal ofPaleontology, 54:728-739. Caesalpinioideae) from the Tertiary of North Frederiksen, N. 0. 1988. Sporomorph biostratigraphy, America. American Journal ofBotany, 78:1-12. floral changes, and paleoclimatology, Eocene and Herendeen, R S., D. H. Less, & D. L. Dilcher. 1990. earliest Oligocene of the eastern Gulf Coast. Fossil CeratophyUum (Ceratophyllaceae) from the
I 26 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
Tertiary of North America. American Journal of Notes on the Mexican and Central American spe- Botany, 77:7-16. cies of Ternstroemia. Journal of the Arnold Ar- Hickey, L. J. 1973. Classification ofthe architecture of boretum, 23:464-478. dicotyledonous leaves. American Journal of Kobuski, C. E. 1943. Studies in the Theaceae, XIV. Botany, 60:17-33. Notes on the West Indian species of Jaramillo, C. A., & D. L. Dilcher. 2001. Middle Paleo- Ternstroemia. Journal of the Arnold Arboretum, gene palynology of Central Colombia, South 24:60-76. America: A study ofpollen and spores from tropi- Kobuski, C. E. 1947. Studies in the Theaceae, XV, a cal latitudes. Palaeontographica Abt. B, 258:87- review of the genus Adinandra. Journal of the 213. ArnoldArboretum, 28:1-98. Johnson, K. R. 1989. A high resolution megafloral bio- Kovach, W. L. & D. L. Dilcher. 1984. Dispersed cu- stratigraphy spanning the Cretaceous-Tertiary ticles from the Eocene of North America. Bo- boundary in the northern Great Plains. Ph.D., Yale tanical Journal ofthe Linnean Society, 88:63-104. University, New Haven. 556 pp. LAWG (Leaf Architecture Working Group). 1999. Johnston, J. E. 1998. Caddisfly cases from the middle Manual of LeafArchitecture - morphological de- Eocene (Lower Lutetian) of Mississippi, USA. scription and categorization ofdicotyledonous and AMBA/AM/PFICM, 98/1.99:61-64. net-veined monocotyledonous angiosperms by Jones, J. H. 1984. Leafarchitectural and cuticular analy- LeafArchitecture Working Group. Smithsonian sis ofextant Fagaceae and' fagaceous' leaves from Institution Press, Washington, D. C., 65 pp. the Paleogene of southeastern North America. Lesquereux, L. 1859. On some fossil plants of recent Ph.D., Indiana University, Bloomington, 328 pp. formations. American Journal of Science 2nd Se- Jones, J. H., & D. L. Dilcher. 1980. Investigations of ries, 27:359-366. angiosperms from the Eocene of North America: Lesquereux, L. 1869. On species of fossil plants from Rhamnus marginams (Rhamnaceae) reexamined. the Tertiary of the state of Mississippi. Transac- American Journal ofBotany, 67:959-967. tions of the American Philosophical Society New Jones, J. H., & D. L. Dilcher. 1988. A study of the Series, 13:411-430. "Dryophyllum" leaf forms from the Paleogene Lopez-Palacio, S. 1977. Verbenaceae, Flora de Ven- of southeastern North America. ezuela. Consejo de Publicaciones, Merida, 654 Palaeontographica Abt. B, 208:53-80. pp. Jones, J. H., S. R. Manchester, & D. L. Dilcher. 1988. Mabberley, D. J. 1989. The Plant Book: A Portable Dryophyllum Debey ex Saporta, Juglandaceous Dictionary ofthe Higher Plants. Cambridge Uni- not Fagaceous. Review of Palaeobotany and versity Press, Cambridge, 706 pp. Palynology, 56:205-211. McLaughlin, R. F. 1957. Plant microfossils from the Jongking, C. C. H. 1989. Rourea Aublet. Pp. 310-368 Bruhn Lignite. Ph.D., University of Tennessee, in F. J. Breteler, ed. The Connaraceae. A taxo- Knoxville, 192 pp. nomic study with emphasis on Africa. Agricul- Miranda, R., & A. J. Sharp. 1950. Characteristics of tural University Wageningen Papers 89-6. the vegetation in certain temperate regions ofeast- Wageningen, The Netherlands. ern Mexico. Ecology, 31:313-333. Klucking, E. R 1986. LeafVenation Patterns. Volume Moore, B. R. 2001. Fossil plants from the Wilbanks 1 Annonaceae. J. Cramer, Berlin, 256 pp. Clay Pit (middle Eocene Claiborne Formation, Klucking, E. R 1987. LeafVenation Patterns. Volume Western Tennessee) paleontological implications 2 Lauraceae. J. Cramer, Berlin, 216 pp. and educational applications. MS, Department of Klucking, E. R 1988. LeafVenation.Patterns. Volume Educational Studies, University of Tennessee, 3 Myrtaceae. J. Cramer, Berlin, 278 pp. Martin, 96 pp. Kobuski, C. E. 1939. Studies in Theaceae. IV, new Moore, B. R., D. L. Dilcher, & M. A. Gibson. 2003. and noteworthy species of Euga. Journal of the Paleoenvironment, depositional setting, and plant ArnoldArboretum, 20:361-374. fossil diversity found in the Claiborne Formation Kobuski, C. E. 1942a. Studies in the Theaceae, XII. (middle Eocene) clay deposits of western Ten- Notes on the South American species of nessee. In R. T. Cox, ed. Field Trip Guidebook, Ternstroemia. Journal of the Arnold Arboretum, Joint Meeting South-Central and Southeastern 23:298-343. Sections Geological Society ofAmerica. State of Kobuski, C. E. 1942b. Studies in the Theaceae, XIII. Tennessee, Department ofEnvironment and Con- DILCHER and LOTT: A fossil assemblage ofPowers Clay Pit 27
servation, Division ofGeology, Report ofInvesti- tion ofRhamnaceae. Kew Bulletin, 55:311-340. gations, 51:187-198. Robyns, A. 1964. Flora of Panama. Part VI. Family Mori, S. A., G Cremers, C. A. Gracie, J.-J. de Granville, 114. Tiliaceae. Annals ofthe Missouri Botanical S. V. Heald, M. Hoff & J. D. Mitchell. 2002. Garden, 51(1-4):1-35. Guide to the vascular plants of central French Roth, J. L. 1981. Epidermal studies in the Annonaceae Guiana. Part 2. Dicotyledons. Memoirs of the and related families. PhD, Department of Biol- New York Botanical Garden, 76 (2):1-776. ogy, Indiana University, Bloomington, 218 pp. Murray, G E. 1961. Geology ofthe Atlantic and Gulf Roth, J. L., & D. L. Dilcher. 1979. Investigations of Coastal province ofNorth America. Harper, New angiosperms from the Eocene of North America: York, 692 pp. stipulate leaves ofthe Rubiaceae including a prob- Owen, D. D. 1860. Second Report of a Geological able polyploid population. American Journal of Reconnoissance ofthe Middle and Southern Coun- Botany, 66:1194-1207. ties of Arkansas. C. Sherman & Son, Philadel- Rudd, V E. 1965 . The American species of Ormosia phia, 433 pp. (Leguminosae). Contributions from the United Parks, W. S. 1975. Stratigraphy of the Paleocene and States National Herbarium 32(5): 1-284. lower part of the Eocene in western Tennessee. Safford, J. M. 1869. Geology of Tennessee. S. C. In E. E. Russel & W. S. Parks, eds. Stratigraphy Mercer, Nashville. 550 pp. ofthe Outcropping Upper Cretaceous, Paleocene, Sajo, M. G., & R J. Rudall. 2002. Leaf and stem and Lower Eocene in western Tennessee (Includ- anatomy ofVochysiaceae in relation to subfamilial ing Descriptions of Younger Fluvial Deposits). and suprafamilial systematics. Botanical Journal Division ofGeology Bulletin, 75:B 1-B39. ofthe Linnean Society, 138:339-364. Parks, W. S., & J. K. Carmichael. 1990. Geology and Schwarzwalder, R. N. 1986. Systematic and early evo- ground-water resources of the Memphis Sand in lution of the Platanaceae. PhI), Department of western Tennessee. United States Geological Biology, Indiana University, Bloomington, 198 pp. Survey Water-Resources Investigation Reports, Sharp, A. J. 1951. The relation of the Eocene Wilcox 88-4182:1-30. flora to some modern floras. Evolution, 5:1-5. Potter, F. W., Jr. 1976. Investigations of angiosperms Standley, R C., & J. A. Steyermark. 1949. Flora of from the Eocene of southeastern North America: Guatemala: Sapindaceae. Fieldiana, Botany, Pollen assemblages from Miller pit, Henry County, 24:234-273. Tennessee. Palaeontographica Abt. B, 157:44- Stearns, R. G 1957. Cretaceous, Paleocene, and lower 96. Eocene geologic history of the northern Missis- Potter, F. W., Jn, & D. L. Dilcher. 1980. Biostratigra- sippi Embayment. Geological Society ofAmerica phy analysis of Eocene clay deposits in Henry Bulletin, 68:1077-1100. County, Tennessee. Pp. 211-225 in D. L. Dilcher Stevens, R F., S. Dressler, & A. L. Weitzman. 2004. & T. N. Taylor, eds. Biostratigraphy of Fossil Theaceae. Pgs. 463-471 in K. Kubitzki, ed. The Plants. Stroudsburg: Dowden, Hutchinson & Families and Genera of Vascular Plants. VI. Ross, Inc. Flowering Plants, Dicotyledons, Celastrales, Prance, G. T. 1972. Chrysobalanaceae. Flora Oxalidales, Rosales, Cornales, Ericales. Berlin, Neotropica, Monograph 9. Hafner Publishing Springer-Verlag. Company, New York, 410 pp. Sturm, M. 1971. Die Eozane flora von Messel bei Radford, A. E., W. C. Dickison, J. R. Massey, & C. R. Darmstadt. I. Lauraceae. Palaeontographica Bell. 1974. Vascular Plant Systematics. Harper Abt. B, 134:1-60. and Row, New York, 891 pp. Sun, Z. 1989. General geology and floristic survey of Raz, L. 2002. Dioscoreaceae. Pg. 479-485 in Flora of the Warman Clay Pit, Weakley County, Tennes- North America Editorial Committee, eds. Flora see. MS, Department of Geology, Indiana Uni- of North America North of Mexico: Volume 26. versity, Bloomington, 77 pp. Magnoliophyta: Liliidae: Liliales and Orchidales. Sun, Z., & D. L. Dilcher. 1988. Fossil Smilax from New York, Oxford University Press. Eocene sediments in western Tennessee. Ameri- Record, S. J. 1939. American woods of the family can Journal ofBotany, 75:118 (abstract). Rhamnaceae. Tropical Woods, 58:6-24. Taylor, D. W. 1987. Survey of biogeographic relations Richardson, J. E., M. F. Fay, Q. C. B. Cronk, & M. W. of North American angiosperms from the Upper Chase. 2000. A revision of the tribal classifica- Cretaceous and Paleogene. Ph.D., University of 2 8 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
Connecticut, Storrs, CT., 338 pp. Oxalidales, Rosales, Cornales, Ericales. Berlin, Taylor, D. W. 1988. Eocene floral evidence of Springer-Verlag. Lauraceae: Corroboration ofthe North American Wolfe, J. A. 1975. Some aspects ofplant geography of megafossil record. American Journal of Botany, the Northern Hemisphere during the Late Creta- 75:948-957. ceous and Tertiary. Annals of the Missouri Bo- Taylor, D. W. 1989. Select palynomorphs from the tanical Gardens, 62:264-279. middle Eocene Claiborne Formation, Tenn., Wolfe, J. A. 1978. A paleobotanical interpretation of (U.S.A.). Review of Palaeobotany and Palynol- Tertiary climates in the Northern Hemisphere. ogy, 58:111-128. American Scientist, 66:694-703. Tschudy, R. H. 1973. Stratigraphic distribution of sig- Wolfe, J. A., & R. Z. Poore. 1982. Tertiary marine and nificant Eocene palynomorphs of the Mississippi nonmarine climatic trends. Pp. 154-158 in Cli- Embayment. United States Geological Survey mate in Earth History. Washington, D. C.: Na- Professional Paper, 743-B: 1-24. tional Academy Press. Villarreal, L. M. G 1986. Contribucion al conocimiento Yu, Cheng-hong, & Ze-lian Chen. 1991. Leafarchitec- del genero Quercus (Fagaceae) en el estado de ture ofthe woody dicotyledons from tropical and Jalisco. Universidad de Guadalajara, Guadalajara. subtropical China. Pergamon Press, Oxford. 414 240 pgs. pp. Weiss, J. 1980. Fossil mimosoid leaflets from the middle Zavada, M. S., & W. L. Crepet. 1981. Investigations Eocene (Claiborne Formation), Graves County, ofAngiosperms from the middle Eocene ofNorth Kentucky. M.S., Department of Geology, Indi- America: flowers ofthe Celtidoideae. American ana University, Bloomington, 76 pp. Journal ofBotany, 68:924-933. Weitzman, A. L., S. Dressler, & R F. Stevens. 2004. Zhizun, D., & M. G Gilbert. 2000. Dioscoreaceae. Ternstroemiaceae. Pgs. 450-462 in K. Kubitzki, Pg. 276-296 in W. Zhengyi & R. H. Raven, eds. ed. The Families and Genera ofVascular Plants. Flora of China, Flagellariaceae through VI. Flowering Plants, Dicotyledons, Celastrales, Marantaceae. Volume 24. Beijing, Science Press, and St. Louis, Missouri Botanical Garden Press. DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 29
Table 3. Leaf Morphology Key
1. Leaves lobed. 2. Lobes entire. Dendropanax eocenensis 2. Lobes toothed. Platanus 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 Swartzia 8 . Secondary veins more than 10 pairs cf. Paleorubiaeophyllum 7. Lamina ratio 4:1. Ormosia sp. 2 6. Venation eucamptodromous Berhamniphyllum sp. 5 . Leaves lanceolate, brochidodromous. Cladrastis 4'. Leaves medium (leaf length 8-15 cm). 9. Leaves elliptic. 10. Intramarginal vein. Myrcia 10'. Venation brochidodromous 11 . Base obtuse Duguetia 11. Base acute 12. Flanks convex Ormosia sp. 1 12 '. Flanks decurrent Apocynophyllum 12". Flanks concave Ficus 10". Some venation eucamptodromous. 13. Leaves lack ascending basal veins 14. Angle of divergence 20-30° Berhamniphyllum claibornense 14. Angle of divergence 60° Oleaceae 13 . Leaves with ascending basal veins Ocotea 10"'. Venation acrodromous/campylodromous Smilax 9. Leaves obovate. Pseudolmedia 4". Leaves large, elliptic (> 15 cm). Duguetia 3. Leaf margin toothed. 15. Leaf medium (6-11 cm). 16. Leafelliptic. 17. Teeth confined to mid-upper portion. Knightiophyllum wilcoxianum 17.Teeth entire length 18. Teeth serrate. Ternstroemites sp. 18. Teeth mostly dentate Populus sp. 16 '. Leaf oblong. Cupanites sp. 16". Leaf obovate. Ternstroemites sp. 16 "'. Leaf linear. Berryophyllum tenuifolia 15. Leaf large, elliptic (> 11 cm). 19. Teeth confined to mid-upper portion. 20. Teeth prominent. Berryophyllum sp. 20. Teeth short Knightiophyllum wilcoxianum 19. Teeth entire length. Ternstroemites sp. 3 0 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
I
B
75 9..
r S
'k>
i
Figure 6. Dendropanax eocenensis. A, B, Three lobed leaf, UF 18810-34383 , lx, 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 31
4 5 B
r G
1
HE t< 1 \ 1 ' 4-fr 1
r
f '.&.:6 K
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. CA H, L, UF18810-34393 , 2x, lx, 4x. I, K, Ormosia sp. 2, UF 18810-34395, l x, 2x. Bar-5 mm (C-F, I-L), 10 mm (A, B, G, H). 32 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45( 1)
I t
f
B
Figure 8. Entire leaves. A, B, Entire Margin Morphotype 2, UF 18810-34444b, l x, 2x. C. D, Ficus sp„ UF 18810- 34446, 1 x, 2x. All bars at 5 mm. DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 33
C
I
H
Figure 9 . Entire leaves. A, B, cf. Paleorubiaceophyllum sp; UF 18810-34431 , l x, 2x . C, E, F, Entire Margin Morphotype 1 , UF18810-34433 , 2x, lx, 3x. D, G, H, Berhamniphyllum sp: UV\ 8810-34427 , lx , Jx, 2x. All bars at 5 rrlrn. 34 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
I =''
D 0-
4 4 4
4 1-4
E H
Figure 10. Entire leaves. A-C, Cladrastis sp. A, Uf \ 8810-34413a, lx . B, C, UF18810-34402 , lx, 2x. D, E, Duguetia sp; UF18810-34407, lx, 1.5x. F-H, Mvrria sp., UF18810-34386, lx, 3x, 2x. All bars at 5 mm. DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 35
,/7 J
I. p I ~.~ b.+ i.'~, ~~ :5- \ ai~ 1HIP
r d
1 -
, @ 8
4
Y..
Y 4 0$44-
D44
Figure 11. Entire leaves. A-C, Ormosia sp. 1. A, UF 1 8810-34406, 1 x. B, C, UF 18810-34404a, 1.5x, 2x. D, E, Apocynophyllum sp., UF18810-34441 , 1 . 5x, lx. F, Oleaceae, UF18810-34430, lx . All bars at 5 mm. 36 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45( 1)
r -i ... n
B
1
A
Figure 12 . Entire leaves. A, B, Berhamniphyllum claibornense, Uf\BB 10-34384, lx, 2x. C-E, G, Ocotea sp. C, UF18810-34409, lx. D, E, UF18810-344108, lx, 2x. 6 UF 18810-34411,2x. F, Oleaceae, UF 18810-34430, 2x. All bars at 5 mm. DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 37
·i
-r A
1
4
A
B
Figure 13 . Entire leaves. A, B, Swartzia sp., UF \ 8810-34388a, lx, 2x. C, D, Smilax sp. 1, UF18810-34415 , 2x, 1 x. E, Smilax sp. 2, UF 18810-34435, l x. All bars at 5 mm. 38 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45(1)
. 'r *'
*9
-e f 4-2
B
D
Figure 14. Entire leaves. A, Smilax sp. 2, UF 188 10-34435, 1.5x. B, D, Pseudolmedia sp., UF18810-34417, lx, 2x. C, Duguetia sp., UF 18810-34418, 1.5x. All bars at 5 mm, DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 39
&1 414, -< ,
F
B
h
A
e
4
t 1%1 D
Figure 15. Entire and toothed leaves. A, C, Entire Margin Morphotype 3, UF18810-34450,2.5x, lx. B, D, Entire Margin Morphotype 5, UF18810-34448,2x, lx. E, Toothed Margin Motphotype 1, UF18810-34445, lx. Bar-5 mm (A, C), 10 mm (B, D, E). 40 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45( 1)
*12
B »4%*~
C
E G
Figure 16. Entire and toothed leaves. A, Ditguetia sp., UF18810-34418 , lx. B, C, G, Knightiophyllum wilcoxianum, UF 1 88 1 0-344258, 1 x, 2x, 2x. D, Populus sp., UF 18810-34397 , lx . E, F, Ternstroemites sp„ UF 18810-34399, l x, 2x. Bar-5 mm (D-G), 10 mm (A-C) DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 41
V A #4. 9
; f'
A
A
/
F D C 13- 36
7
r
G
Figure 17 . Toothed leaves. A, Populus sp., UFl 8810-34397, 2x . B, D, Cupanites sp: Ufl 8810-34401 , lx, 2x. C, 1, Ternstroemites sp. U F 1 8 8 1 0 -3 442 6, 1 x, 2 x . E -G, Ber,yophy/lum s p ., U F 1 8 8 1 0- 3 4 3 9 6. 1 x, 3 x, 2 x . H . Berryophyllum tenuifolia, UF 18810-34389, lx. All bars at 5 mm. 42 BULLETIN FLORIDA MUSEUM NATURAL HISTORY VOL. 45( 1 )
50 .L
C
0 1~ 4 *t:)
**
, 1
Figure 18. Toothed leaves and reproductive structures. A, E, Ternstroemites sp UF 1881 0-34405, lx, 2x. C, I, Gordonia sp., UF 1881 0-34438, 4x, lx. B, Berryophyllum tenuifolia. UF \88 10-34389, 4x. D, H, Liquidam- bar sp., UF 1 8810-34432, 3x, 1 x. F, J, Reproductive Structure Morphotype 1, UF 18810-34434, 7x, 1 x. G, K, Nyssa eolignitica, UF \ 8810-34392, 3x, lx . Bar-2 mm (F), 5 mm ( B- IE, G-K), 10 mm (A). DILCHER and LOTT: A fossil assemblage of Powers Clay Pit 43
2.. f
C
2
4
D
'
H
Figure 19. Leaves and insect larval cases. A, Tooth Margin Morphotype 1, UFI 8810-34445 cpt., 3x. B, Monocot leaf, UF 18810-34400. l x. C, H, Entire Margin Morphotype 4. UF 18810-34449, lx, 2x . D, (3 Folindusia sp., UF18810-34436, lx, 3x. E, F, Terrindlisia sp.. UF 18810-34437, 1 x, 6x. Bar-5 mm (A, B, D-H), 10 mm (C). The BULLETIN OF THE FLORIDA MUSEUM OF NATURAL HISTORY publishes research conducted by our faculty, staff, students, and researcb associates. We also enc6urage appropriate, fully funded manuscripts from external researchers. Manuscripts concerning natural history or systematic problems involving the southeastern United States or the Neotropics are especially welcome, although we will also consider research from other parts of the world. Priority is given to specimen-based research. We consider thirty-five double-spaced pages (excluding figures and tables) as the minimum length for manuscripts, although there can be exceptions as determined by the Editor and Bulletin Committee.
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MacFadden, B. J. and O Carranza-Castaneda. 2002 . Cranium of Dinohippus mexicanus (Mammalia: Equidae) from the early Pliocene (Latest Hemphillian) of central Mexico, and the origin ofEquus. Volume 43, No.5, pp.163- 185. Price $5.00
Kratter, A. W., T. Webber, T. Taylor, and D. W Steadman. 2002. New specimen-based records of Florida birds. Volume 43, No.4, pp. 111-161. Price $5.50
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, Pulmonata, Helicoidea, Humboldtianidae). Volume 43, No. 2, Pp. 49-77. Price $5.00
Hayes, F. G. 2000. The Brooksville 2 local fauna (Arikareean, latest Oligocene): Hernando County, Flofida Volume 43, No.1, pp. 1-47. Price $6.50
*A complete list ofpublications in the Bulletin of the Florida Museum ofNatural History can be found on the Florida Museum web site http://www.flmnh.ufl.edu/bulletin/bulletin_vols.htm. Orderpublications from the Managing Editor. Florida residents are required to add 6.25% sales tax for all purchases. Add $1.50 per publication for·shipping.