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Fruits, seeds and flowers from the Bovay and Bolden clay pits (early Eocene Tallahatta Formation, Claiborne Group), northern Mississippi, USA

Jane Blanchard, Hongshan Wang, and David L. Dilcher

Plain Language Abstract

This report is the second of a series of comprehensive reviews of the flowers, fruits, and seeds from localities of the Eocene Claiborne Group of the southeastern United States. We examine 193 specimens from two localities, the Bovay and Bolden clay pits, in northern Mississippi, USA. The investigation has resulted in the recognition of 52 different types. Based upon comparisons of characters of these specimens with available modern plant material and the literature, we relate 17 types to 11 modern plant families. In addition, 35 types are not assigned to any family due to the limited number of defining characters. Of the 52 types recognized, 38 are unique to the Bovay locality and nine are unique to Bolden. Only five types are common to both Bovay and Bolden. Compared with the 62 types recognized from the Warman clay pit, Tennessee, only eight are shared with the Bovay and/or Bolden localities. This study has expanded our knowledge of the Eocene plant fossils of the southeastern United States and has shown that there is a remarkable lack of species overlap among the three localities examined to date.

Resumen en Español

Frutos, semillas y flores de los pozos de arcilla de Bovay y Bolden (Formación Tallahatta del Eoceno temprano, Grupo Claiborne), norte de Mississippi, EE.UU.

El presente informe es el segundo de una serie de revisiones exhaustivas del material reproductivo de las localidades del Grupo Claiborne del Eoceno del sureste de los Estados Unidos. Examinamos restos fósiles reproductivos de plantas de dos localidades, los pozos de arcilla de Bovay y Bolden, en el norte de Mississippi, EE.UU. El estudio de 193 especímenes ha dado como resultado el reconocimiento de 52 taxones/morfotipos. De acuerdo con las comparaciones de la morfología macroscópica de estos especímenes con el material disponible de plantas actuales y la bibliografía, podemos relacionar 17 taxones/morfotipos con las siguientes 11 familias actuales: Annonaceae (1), Araliaceae (1), Caprifoliaceae (1), Eucommiaceae (1), Fabaceae (6), Fagaceae (2), Icacinaceae (1), Sapindaceae (1), Salicaceae (1), Rhamnaceae (1) y Theaceae (1). Además, 35 morfotipos no se asignan a ninguna familia debido al número limitado de caracteres de valor diagnóstico. De los 52 taxones/morfotipos reconocidos, 38 (73,1%) son únicos para la localidad de Bovay y nueve (17,3%) son únicos para la de Bolden. Sólo dos (9,6%) taxones/morfotipos son compartidos por ambas localidades. De los 62 taxones/morfotipos reconocidos en el pozo de arcilla de Warman, Tennessee, sólo ocho taxones/morfotipos se comparten con las localidades de Bovay y/o Bolden, Mississippi. Este estudio ha ampliado el conocimiento de la diversidad de plantas eocenas del sureste de los Estados Unidos y ha demostrado que hay una notable falta de superposición de especies entre las tres localidades examinadas hasta la fecha.

PALABRAS CLAVE: Nuevas especies; material reproductivo; Formación Tallahatta; Eoceno temprano; Mississippi

Traducción: Enrique Peñalver (Sociedad Española de Paleontología)

Résumé en Français

Fruits, graines, et fleurs des carrières d'argile de Bovay et Bolden (Éocène ancien de la formation Tallahatta, groupe Claiborne), partie nord de l'État du Mississippi, États-Unis d'Amérique

Ce rapport est le deuxième d'une série de synthèses exhaustives de l'appareil reproducteur des plantes des localités éocènes du groupe Claiborne, sud-est des États-Unis d'Amérique. Nous examinons l'appareil reproducteur des plantes fossiles de deux localités, les carrières d'argile de Bovay et Bolden, au nord de l'État du Mississippi, États-Unis d'Amérique. L'étude de 193 spécimens a conduit à reconnaître 52 taxons/morphotypes. D'après des comparaisons de la morphologie macroscopique de ces spécimens avec le matériel disponible pour les plantes actuelles et avec la littérature, nous pouvons associer 17 taxons/morphotypes aux 11 familles actuelles suivantes : Annonaceae (1), Araliaceae (1), Caprifoliaceae (1), Eucommiaceae (1), Fabaceae (6), Fagaceae (2), Icacinaceae (1), Sapindaceae (1), Salicaceae (1), Rhamnaceae (1), et Theaceae (1). De plus, 35 morphotypes ne sont attribués à aucune famille à cause d'un nombre limité de caractères diagnostiques. Parmi les 52 taxons/morphotypes reconnus, 38 (73,1 %) ne sont trouvés que dans la localité de Bovay et neuf (17,3 %) ne sont trouvés que dans la localité de Bolden. Seuls cinq (9,6 %) taxons/morphotypes sont communs aux deux localités. Parmi les 62 taxons reconnus dans la carrière d'argile de Warman, Tennessee, seuls huit taxons/morphotypes sont communs avec les localités de Bovay et/ou Bolden, Mississippi. Cette étude a permis d'améliorer notre connaissance de la diversité des plantes éocènes du sud-est des États-Unis d'Amérique et a montré qu'il y avait un très faible recoupement d'espèces entre les trois localités examinées jusqu'à présent.

Mots-clés : nouvelle espèce ; appareil reproducteur ; formation Tallahatta ; Éocène ancien ; Mississippi

Translator: Antoine Souron

Deutsche Zusammenfassung

Früchte, Samen und Blüten aus den Tongruben Bovay und Bolden (frühes Eozän Tallahatta Formation, Claiborne Gruppe), nördliches Mississippi, USA

Dieser Report ist der zweite einer Reihe umfassender Bewertungen des reproduktiven Materials aus den Fundstellen der eozänen Claiborne Gruppe in den südöstlichen Vereinigten Staaten. Wir untersuchen fossiles reproduktives Pflanzenmaterial aus zwei Fundstellen, den Tongruben Bovay und Bolden im nördlichen Mississippi, USA. Die Untersuchung der 193 Stücke resultierte in der Identifizierung von 52 Taxa/Morphotypen. Durch Vergleiche der groben Morphologie dieser Stücke mit verfügbarem heutigen Pflanzenmaterial und der Literatur sind wir in der Lage 17 Taxa/Morphotypen folgenden 11 heutigen Familien zuzuordnen: Annonaceae (1), Araliaceae (1), Caprifoliaceae (1), Eucommiaceae (1), Fabaceae (6), Fagaceae (2), Icacinaceae (1), Sapindaceae (1), Salicaceae (1), Rhamnaceae (1) und Theaceae (1). 35 Morphotypen sind aufgrund der limitierten Anzahl diagnostischer Merkmale keiner Familie zugeordnet. Von den 52 erkannten Taxa/Morphotypen sind 38 (73.1%) einzigartig in Bovay und neun (17.3%) einzigartig in Bolden. Nur fünf (9.6%) Taxa/Morphotypen kommen in beiden Fundstellen zugleich vor. Von den 62 aus der Warman Tongrube (Tennessee) identifizierten Taxa/Morphotypen kommen nur acht Taxa/Morphotypen in Bovay und/oder Bolden (Mississippi) vor. Diese Untersuchung hat unser Wissen über die eozäne Pflanzenvielfalt der südöstlichen Vereinigten Staaten erweitert und zeigte, dass sich die Arten in den drei Fundstellen bemerkenswert wenig überlappen.

Schlüsselwörter: neue Art; reproduktives Material; Tallahatta Formation; frühes Eozän; Mississippi

Translator: Eva Gebauer

Arabic

Translator: Ashraf M.T. Elewa

 

 

TABLE 1. All taxa/morphotypes reported from the Bovay and Bolden localities in northern Mississippi, and the Warman locality in western Tennessee, USA (Wang et al., 2013). The table also lists each taxon/morphotype’s organ type, proposed modern affinity, and its figure numbers in this paper and in Wang et al. (2013). “+” indicates a taxon/morphotype is present at a locality.

Proposed modern affinity Taxa/morphotypes Figure numbers in this paper Figure numbers in Wang et al. (2013) Organ type Warman Bovay Bolden
Lauraceae Lauroflorum warmanensis Wang, Blanchard, and Dilcher, 2013   figure 3 flower +    
Annonaceae Anonaspermum robertsi (Berry) Wang, Blanchard, and Dilcher, 2013 Figure 4 figure 4.1-3, 4.5-8 seed +   +
Anonaspermum warmanensis Wang, Blanchard, and Dilcher, 2013   figure 4.4 seed +    
Magnoliaceae cf. Magnolia sp.   figure 5 seed +    
Araceae cf. Acorites heeri Crepet, 1978   figure 6 inflorescence +    
Altingiaceae Claiborne Altingiaceae Infructescence Type 1   figure 7 infructescence +    
Hamamelidaceae Corylopsis grotei Wang, Blanchard, and Dilcher, 2013   figure 8 seed +    
Hamamelis sp.   figure 9 seed +    
Euphorbiaceae Hippomaneoidea warmanensis Crepet and Daghlian, 1982   figure 20 inflorescence +    
Malpighiaceae Eoglandulosa warmanensis Taylor and Crepet, 1987   figure 21 flower +    
Salicaceae cf. Populus sp. Figure 13   fruit     +
Fabaceae Caesalpinia claibornensis Herendeen and Dilcher, 1991   figure 13.4-5 fruit +    
Crudia grahamiana Herendeen and Dilcher, 1990a   figure 13.6 fruit +    
Diplotropis claibornensis Herendeen and Dilcher, 1990b   figure 14.1 fruit +    
Eliasofructus claibornensis Herendeen and Dilcher, 1990c   figure 14.2 fruit +    
Eomimosoidea plumosa Crepet and Dilcher, 1977   figures 10-11 inflorescence +    
Ormosia sp.   figure 14.5-6 fruit +    
Protomimosoidea buchananensis Crepet and Taylor, 1986 Figures 5-7 figure 12 flower + +  
Swartzia sp.   figure 13.1 fruit +    
cf. Acrocarpus sp. Figure 8   fruit   +  
cf. Erythrophleum sp.   figure 13.2 fruit +    
cf. Gleditsia? mississippiensis (Berry) Berry, 1930 Figure 9 figure 14.3 fruit +   +
cf. Leguminosites phyllocarpoides Berry, 1930   figure 13.3 fruit +    
Claiborne Fabaceae Floral Type 1 Figure 11   flower   +  
Claiborne Fabaceae Floral Type 2 Figure 12   flower   + +
"Short, broad winged fruit" Figure 10   fruit   + +
"Thick walled fruit"   figure 14.4 fruit +    
Fagaceae Quercus oligocenensis Daghlian and Crepet, 1983   figure 15 staminate inflorescences +    
cf. Trigonobalanoidea americana Crepet and Nixon, 1989 Figure 14   infructescence   +  
Claiborne Fagaceae Fruit Type 1   figure 16.1-2 fruits +    
Claiborne Fagaceae Fruit Type 2   figure 16.3 fruit +    
Claiborne Fagaceae Fruit Type 3 Figure 15.1-3 figure 17.4 fruit +    
Claiborne Fagaceae Fruit Type 4 Figure 16   fruit   +  
Claiborne Fagaceae Fruit Type 5 Figure 15.5 figure 17.1-3 fruit +    
Juglandaceae Eokachyra aeolius Crepet, Dilcher and Potter, 1975   figure 18 inflorescence +    
Paleooreomunnea stoneana Dilcher, Potter and Crepet, 1976   figure 19 fruit +    
Moraceae Cornerocarpon copiosum Grote, 1989   figures 22, 23 fruit +    
Rhamnaceae “Solanites” pusillus Berry, 1930 Figures 17-18 figure 35 flower + + +
Sapindaceae Bridgesia bovayensis Manchester and O’Leary, 2010 Figure 19   fruit   +  
Theaceae Andrewsiocarpon henryense Grote and Dilcher, 1989   figure 27 fruit +    
Gordonia warmanensis Grote and Dilcher, 1989 Figure 20 figure 24 fruit + +  
cf. Gordonia sp. 1   figure 25 flower +    
cf. Gordonia sp. 2   figure 26 flower +    
Icacinaceae cf. Natsiatum wilcoxiana (Berry) Stull, Moore and Manchester, 2011 Figure 21   endocarp   +  
Eucommiaceae Eucommia eocenica (Berry) Brown, 1940 Figure 22   fruit   + +
Bignoniaceae Grotea warmanensis Wang, Blanchard and Dilcher, 2013   figure 28.1 fruit +    
Oleaceae Fraxinus wilcoxiana (Berry) Call and Dilcher, 1992   figure 29 fruit +    
Caprifoliaceae Dipelta bovayensis sp. nov. Figure 23   fruit   +  
Araliaceae Araliasemina Berry, 1930   figure 30 seed +    
Paleopanax boldensis sp. nov. Figure 24   fruit   +  
Angiosperm incertae sedis - flowers Antholithes wilcoxensis Berry, 1930   figure 34 flower +    
Solanitessarachaformis Berry, 1930 Figure 26   flower   +  
cf. “Solanitessaportanus (Berry) Martínez-Millán, 2010 Figure 25   flower   +  
Claiborne Floral Type 1   figure 36.1-2 flower +    
Claiborne Floral Type 2   figure 36.3-4 flower +    
Claiborne Floral Type 3   figure 37 flower +    
Claiborne Floral Type 4   figure 38 flower +    
Claiborne Floral Type 5 Figure 27 figure 39 flower +   +
Claiborne Floral Type 6   figure 40 flower +    
Claiborne Floral Type 7 Figure 28   flower   +  
Claiborne Floral Type 8 Figure 29   flower   +  
Claiborne Floral Type 9 Figure 30   flower   +  
Claiborne Floral Type 10 Figure 31   flower   +  
Claiborne Floral Type 11 Figure 32   flower   +  
Claiborne Floral Type 12 Figure 33   flower     +
Claiborne Floral Type 13 Figure 34   flower   +  
Claiborne Floral Type 14 Figure 35   flower     +
Claiborne Floral Type 15 Figure 36   flower   +  
Claiborne Floral Type 16 Figure 37   flower   +  
Claiborne Floral Type 17 Figure 38   flower     +
Claiborne Floral Type 18 Figure 39   flower   +  
Claiborne Floral Type 19 Figure 40   petal?   +  
Claiborne Floral Type 20 Figure 41   stamens   +  
Angiosperm incertae sedis - infructescences " Sparganium " sp. Figure 42 figure 49 infructescence + +  
Claiborne Infructescence Type 1   figure 31 infructescence +    
Claiborne Infructescence Type 2   figure 32 infructescence +    
Claiborne Infructescence Type 3   figure 33 infructescence +    
Claiborne Infructescence Type 4 Figure 43   infructescence   +  
Claiborne Infructescence Type 5 Figure 44   infructescence   +  
Claiborne Infructescence Type 6 Figure 45   infructescence   +  
Claiborne Infructescence Type 7 Figure 46   infructescence     +
Angiosperm incertae sedis - fruits and seeds Carpolithus boldensis sp. nov. Figure 47   seed     +
Carpolithus collinsi (Berry) Wang, Blanchard, and Dilcher, 2013   figure 41 fruit +    
Carpolithus complanata (Lesquereux) Wang, Blanchard, and Dilcher, 2013   figure 42 seed +    
Carpolithus prangosoides Berry, 1916b   figure 43 fruit +    
Carpolithus sophorites Berry, 1916b   figure 44 seed +    
Carpolithus warmanensis Wang, Blanchard, and Dilcher, 2013   figure 45 fruit +    
Palmocarpon wilcoxiana (Berry) Wang, Blanchard, Dilcher, 2013 Figure 55 figure 47, 48 fruit + +  
Acerknowltoni (Berry) Brown, 1940 Figure 49   fruit   + +
Copaiferayeguana Berry, 1915   figure 46 fruit +    
cf. Saportaspermum sp. Figure 48   seed   +  
Claiborne Fruit/Seed Type 1   figure 50 fruit/seed +    
Claiborne Fruit/Seed Type 2   figure 51 fruit/seed +    
Claiborne Fruit/Seed Type 3   figure 52 fruit/seed +    
Claiborne Fruit/Seed Type 4   figure 53 fruit/seed +    
Claiborne Fruit/Seed Type 5   figure 54 fruit/seed +    
Claiborne Fruit/Seed Type 6 Figure 50   fruit/seed   +  
Claiborne Fruit/Seed Type 7 Figure 51   fruit/seed   +  
Claiborne Fruit/Seed Type 8 Figure 52   fruit/seed   +  
Claiborne Fruit/Seed Type 9 Figure 53   fruit/seed   +  
Claiborne Fruit/Seed Type 10 Figure 54   fruit/seed   +  
Plant incertae sedis Equisetum?   figure 55 unknown +    
Undetermined Claiborne Undetermined Structure 1 Figure 56   unknown   +  
Claiborne Undetermined Structure 2 Figure 57   unknown   +  
Claiborne Undetermined Structure 3 Figure 58   unknown   +  
Claiborne Undetermined Structure 4 Figure 59   unknown   +  
Number of taxa/morphotypes from each locality         62 43 14
Number of taxa/morphotypes from all three localities         106    

 

TABLE 2. Comparison of major characters of Protomimosoidea buchananensis, Claiborne Fabaceae Floral Types 1 and 2, Claiborne Floral Types 17 and 18. All dimensions are in millimeters. “?” indicates that a measurement is not available or a character is not observed.

  Ovary size (length x width) Number of stamens Filament length Anther size
(length x width)
Anther connective Perianth
(length x width)
Pedicel
(length x width)
Protomimosoidea buchananensis 2 x 1 10 5 0.75-1 x 0.5 strong 2 x 1 up to 6 x 0.3
Claiborne Fabaceae Floral Type 1 expanded ? 9 1.5 x? ? ? ?
Claiborne Fabaceae Floral Type 2 2 x 2.5 > 10 1 0.5 x 0.5 strong 1 x 1 5-5.5 x 1
Claiborne Floral Type 17 2 x 2.5 > 10 4-5 ? ? 4 x 2 3.5 x 1
Claiborne Floral Type 18 2 x 2 4-6 2, fused 0.5-1 x 0.5 very strong ? 1 x 0.5


TABLE 3.
Major characters of Claiborne Fagaceae Fruit Types 1 to 5 and Catahoulea grahamii. All dimensions are in millimeters. “?” indicates that a measurement is not available or a character is not observed.

  Fruit size (length x width) Number of nutlets Nutlet size (length x width) Spines Cupule valve Web-like pattern of cupule Peduncle (length x width) Specimen number and references
Claiborne Fagaceae Fruit Type 1 15 x 35 ? ? simple or branched ? ? 10 x 1 UF15826-51476 (figure 16.1-2 in Wang et al., 2013) and six other specimens from Warman clay pit, Tennessee
Claiborne Fagaceae Fruit Type 2 20 x 20 ? ? simple ? no ? UF15826-51231 (figure 16.3 in Wang et al., 2013)
Claiborne Fagaceae Fruit Type 3 10 x 10 at least 2 2 x 1 simple or branched 3 yes 0.5-1 x? UF15815-005347 (Figure 15.1-2); UF15826-029215 (figure 2.F in DeVore et al., 2014); UF15826-51307 (Figure 15.3; figure 17.4 in Wang et al., 2013; figure 2.C DeVore et al., 2014).
Claiborne Fagaceae Fruit Type 4 12 x 6.5 at least 2 7 x 5 branched ? spines intertwined 2 x 0.5 UF15737-59211 (Figure 16)
Claiborne Fagaceae Fruit Type 5 25 in diameter ? ? branched 3 yes 10 x 1 UF15826-29216 (Figure 15.5; figure 17.1-3 in Wang et al., 2013)
Catahoulea grahamii 5.5-10 x 13 3 one large, 3 x 2; two small 3 x 1 simple or branched 3 yes 9.5 x 0.3 UF 15841-062191 (Figure 15.4) and other 23 specimens from Huntsville, Texas (DeVore et al., 2014)


TABLE 4.
Comparison of characters of Claiborne Floral Type 5, Claiborne Floral Types 8 to 11, " Diospyros " mirafloriana, Eoglandulosa warmanensis, “Solanitespusillus, “Solanitessaportanus, and “Solanitessarachaformis. All dimensions are in millimeters. “?” indicates that a measurement is not available or a character is not observed.

 

Taxa/morphotypes Flower -merous Size in diameter when vertically compressed Number of sepals Sepal base Sepal shape Sepal size (length x width) Sepal venation Number of petals Petal shape Petal size (length x width)
Claiborne Floral Type 5 5-merous 22 ? ? ? ? ? ? ? ?
Claiborne Floral Type 8 ?4-merous 8 ? ? ? ? ? ? ? ?
Claiborne Floral Type 9 5-merous 6 ? ? ? ? ? ? ? ?
Claiborne Floral Type 10 6-merous 15 ? ? ? ? ? 6 ovate 5 x 2-2.5
Claiborne Floral Type 11 ?4-merous 11 ? ? ? ? ? ? ? ?
"Diospyros" mirafloriana 5-merous 35-40 5 separate ? 12.5 x 9.5 ? ? ? ?
Eoglandulosa warmanensis (Wang et al. 2013) 5-merous 10 ?5 connate oblong to triangular 4 x 2 ? ? ? ?
“Solanites”  pusillus 5-merous 8-10 5 separate isosceles-triangular   2.5-3 x 2.5-3 mid-vein present 5 ? 3 x 1-2
“Solanites” saportanus 6-merous 9 6 separate triangular 3 x 2 mid-vein present ? ? ?
“Solanites” sarachaformis 4-merous 10 4 separate ovate 4.5 x 3.5 5 veins ? ? ?

 TABLE 4 (continued).

Taxa/morphotypes Petal venation Arrangement Number of periath parts when sepals and petals can not be distinguished Perianth part shape Perianth part size (length x width) Perianth part venation Hypanthium Hypanthium rim Floral cup diameter Ovary Pedicel (length x width)
Claiborne Floral Type 5 ? ? 5 ovate 7 x 5 faint veins present ? ? superior ?
Claiborne Floral Type 8 ? alternate, 2 whorls 8 triangular 1.5 -2.5 x 1.5-2 5 veins present present 3 superior ? x 0.5
Claiborne Floral Type 9 ? ? 5 triangular 2 x 1 not observed present present ? 2-3 ? x 0.2
Claiborne Floral Type 10 thin ? ? ? ? ? ? ? ? inferior ?
Claiborne Floral Type 11 ? ? ? ovate 5 x 3 fine veins ? ? ?  ? long; 10 x 0.5
"Diospyros" mirafloriana ? ? ? ? ? ? present present 12.5 ? ?
Eoglandulosa warmanensis (Wang et al. 2013) ? ? ? ? ? ? ? ? ? ? ?
“Solanites”  pusillus ? alternate, two whorls ? ? ? ? present present 3.5 – 4.5  inferior 1 x 0.5
“Solanites” saportanus ? ? ? ? ? ? present absent 4 ? ? x 0.5
“Solanites” sarachaformis ? ? ? ? ? ? present absent ? ? ?


TABLE 5.
Total numbers of specimens, leaves, and reproductive material from the Bovay and Bolden clay pit, Mississippi, and the Warman clay pit, Tennessee.

  Bovay Bolden Bovay and Bolden Warman
Total number of specimens 505 187 692 3166
Leaves 359 140 499 2560
Reproductive material Total number of reproductive material 146 47 193 606
Identified 89 18 107 338
Unidentified 57 29 86 268
 

FIGURE 1. Locations of the Bovay clay pit (UF15737) in Marshall County, and Bolden clay pit (UF15738) in Benton County, northern Mississippi, USA.

 figure1

FIGURE 2. Sketch map showing collection sites in the Bovay clay pit and the Bolden clay pit, northern Mississippi: 1) Bovay clay pit. Numbers indicate collecting sites within the clay pit; 2) Bolden clay pit. * indicates the place where the section is measured in Johnston (1993, p. 73, text-figure 2).

figure2 

FIGURE 3. Simplified stratigraphic column of the middle Eocene Claiborne Group in northern Mississippi (after Dockery, 1996). Specimens examined in this paper were collected from the Tallahatta Formation at Bovay and Bolden localities, Mississippi. Specimens from other localities used for comparison to the Tallahatta Formation, including the Warman clay pit, were collected from the Cockfield Formation in western Tennessee and southwest Kentucky.

figure3 

FIGURE 4. Anonaspermum robertsi (Berry) Wang, Blanchard, and Dilcher: 1-2) UF15738-060635. Part and counterpart of a specimen showing reniform seed, transverse endosperm ruminations, and thick testa. Scale bar equals 3 mm.

figure4 

FIGURE 5. Protomimosoidea buchananensis Crepet and Taylor, 1986, UF15737-026430. Flower showing at least three valvate petals, 10 stamens, and dorsifixed anthers. Note two stamens with missing anthers (indicated by arrows). Scale bar equals 0.5 mm.

 figure5

FIGURE 6. Protomimosoidea buchananensis Crepet and Taylor, 1986, UF15737-060709. Specimen showing three petals and five stamens. Scale bar equals 0.5 mm.

figure6 

FIGURE 7. Protomimosoidea buchananensis Crepet and Taylor, 1986: 1) UF15737-027593. Two flowers with attached anthers. Note long pedicel. Scale bar equals 2 mm; 2) Enlargement of Figure 7.1 (indicated by upper arrow) to show anther. Scale bar equals 1 mm; 3) Enlargement of Figure 7.1 (indicated by left arrow) to show anthers and filaments. Scale bar equals 1 mm.

 figure7

FIGURE 8. cf. Acrocarpus sp.: 1-2) UF15737-008276, 008276'. Part and counterpart of a specimen showing a single valve of a fruit with alternating outlines of three seeds (s) and four seed chambers (sc). Scale bar equals 5 mm.

figure8 

FIGURE 9. cf. Gleditsia? mississippiensis Berry, 1916b: 1-2) UF15738-008791, 008791'. Part and counterpart of a specimen showing stipe (s), calyx (c), calyx lobes (cl), and pod with rounded apex. Scale bar equals 2 mm.

 figure9

FIGURE 10. "Short, broad winged fruit" (1-3) UF15737-008279: 1) Showing stipe (s), calyx (c), and pod with attenuate apex (a). Scale bar equals 5 mm; 2) Counterpart of UF15737-008279. Scale bar equals 5 mm; 3) Enlargement of Figure 10.2 to show valve venation. Scale bar equals 2 mm; 4) UF15738-008790. Showing stipe (s), calyx (c), and pod with attenuate apex (a). Scale bar equals 5 mm.

 figure10

FIGURE 11. Claiborne Fabaceae Floral Type 1, UF15737-059328. Laterally preserved flower showing a short pedicel (pd), calyx (c), 2 petals (pt), standard petal (sp), expanding ovary (o) with an apical style (st), and a stamen with a filament (f) and an anther (a). Scale bar equals 2 mm.

figure11 

FIGURE 12. Claiborne Fabaceae Floral Type 2: 1-2) UF15738-008215, 008215'. Part and counterpart of a laterally preserved flower showing style (s) and a pedicel (p). Scale bars equal 2 mm; 3) Enlargement of Figure 12.2 to show anthers (a), filaments (f), and calyx (c). Scale bar equals 1 mm; 4-5) UF15737-059312, 059312'. Part and counterpart of a laterally preserved flower showing a petal (indicated by arrows) and stamens above. Scale bar equals 2 mm; 6) UF15737-059252. A laterally compressed, immature flower showing a style (s). Scale bar equals 2 mm.

figure12 

FIGURE 13. cf. Populus sp., UF15738-008213. Specimen showing a four-valved capsule. Scale bar equals 1 mm.

 figure13

FIGURE 14. cf. Trigonobalanoidea americana Crepet and Nixon, 1989: 1) UF15737-059224. An immature infructescence with at least 32 sessile fruits. Scale bar equals 2 mm; 2) Enlargement of a fruit indicated by the upper arrow in Figure 14.1 to show two styles (s, indicated by arrow). Scale bar equals 0.5 mm; 3) Enlargement of a fruit indicated by the lower arrow in Figure 14.1 to show three styles (s, indicated by arrow). Scale bar equals 0.5 mm; 4) Counterpart of Figure 14.1 showing an immature infructescence. Scale bar equals 2 mm.

 figure14

FIGURE 15. Claiborne Fagaceae Fruit Type 3 (1-3): 1) UF15815-005347', Lamkin clay pit, Kentucky. Fruit showing two nutlets (n) and spiny cupule valve (c) with a web-like pattern. Scale bar equals 2 mm; 2) UF15815-005347, Lamkin clay pit, Kentucky. Counterpart of Figure 15.1 showing a short peduncle (p). Note the overlap of spiny cupule valves (c). Scale bar equals 2 mm. 3) UF15826-051307, Warman clay pit, Tennessee. Fruit showing spiny cupule valve (c) and short peduncle (p). Scale bar equals 2 mm. Catahoulea grahamii DeVore, Pigg, Dilcher and Freile, 2014: 4) UF15841-062191, Huntsville, Texas. Fruit showing spiny cupule valve (c) and long peduncle (p). Scale bar equals 2 mm. Claiborne Fagaceae Fruit Type 5 (5) UF15826-29216, Warman clay pit, Tennessee. Specimen showing peduncle (p) and two overlapping spiny cupule valves (c). Scale bar equals 2 mm.

figure15 

FIGURE 16. Claiborne Fagaceae Fruit Type 4: 1) UF15737-059211. Showing interwined spines, fruit body, a short peduncle (p). Scale bar equals 1 mm; 2) UF15737-059211'. Counterpart of Figure 16.1 showing three ridges converging at both ends of the fruit body and a short peduncle (p). Scale bar equals 1 mm.

figure16 

FIGURE 17.Solanitespusillus Berry, 1930: 1) UF15737-049568. Vertically compressed flower showing five triangular sepals (s) alternating with petals (p, only three are visible), and a floral cup at the center. Scale bar equals 1 mm; 2) UF15737-049568'. Counterpart of Figure 17.1 showing sepals and membranaceous petals with conspicuous venation. Scale bar equals 1 mm; 3) UF15737-008226. A laterally compressed flower showing an obtriangular hypanthium with an inferior ovary (o), a short pedicel (indicated by arrow), and three sepals (s), and three petals (p). Scale bar equals 1 mm; 4) UF15737-049566. Vertically compressed flower showing a circular floral cup (dark concentric rings) and five triangular sepals alternating with five petals. Scale bar equals 1 mm; 5) UF15737-049566'. Counterpart of Figure 17.4 showing sepals alternating with petals, and a floral cup with radiating veins. Note the center hole resulting from vertical compression of the pedicel. Scale bar equals 1 mm; 6) Enlargement of Figure 17.2 to show venation of sepal (s) and petal (p). Scale bar equals 0.5 mm.

 figure17

FIGURE 18.Solanitespusillus Berry, 1930: 1) UF15738-008214. Vertically compressed flower showing four sepals. Note two filaments (f) between two adjacent sepals (s) indicated by arrows. Scale bar equals 1 mm; 2) UF15738-008214'. Counterpart of Figure 18.1 showing five sepals and the radiating veins on the hypanthium extending to the sepals (indicated by arrow). Scale bar equals 1 mm.

figure18 

FIGURE 19. Bridgesia bovayensis Manchester and O’Leary, 2010: 1) UF15737-30660. Fruit showing pedicel, hypogynous perianth scar (ps), stylar protrusion (sp), and two mericarps each with a large darkened locular area. Scale bar equals 3 mm; 2) UF15737-30661. Mericarp showing reticulate veins on the wing and locular area. Scale bar equals 3 mm.

figure19 

FIGURE 20. Gordonia warmanensis Grote and Dilcher, 1992: 1-2) UF15737-059235, 059235'. Part and counterpart of a specimen showing a winged seed. Note slightly concave ventral margin and convex dorsal margin. Scale bar equals 1 mm; 3-4) UF15737-059239, 059239 '. Part and counterpart of a specimen showing slightly concave ventral margin and convex dorsal margin. Scale bar equals 1 mm.

figure20 

FIGURE 21. cf. Natsiatum wilcoxiana (Berry) Stull et al., 2011: 1-2) UF15737-059325, 059325'. Part and counterpart of a specimen showing irregular, blindly ending ridges. Scale bar equals 1 mm.

 figure21

FIGURE 22. Eucommia eocenica (Berry) Brown, 1940: 1) UF15737-008218. Fruit showing apical stigmatic cleft (sc), fertile carpel (fc) bearing a conspicuous reticulum (r) composed of fascicles of vascular strands, infertile carpel (ic), and carpel suture (cs). Scale bar equals 2 mm; 2, 3) UF15737-011033, 011033’. Part and counterpart of a specimen showing stout stipe (s). Scale bar equals 3 mm; 4) UF15737-011032 - Incomplete fruit showing longer stipe. Scale bar equals 2 mm; 5) UF15737-011063. Well developed fruit showing vascular strands of the reticulum and stout stipe. Scale bar equals 2 mm; 6) UF15737-011035. Distal half of fruit showing compression of latex strands and strong carpel suture. Scale bar equals 2 mm; 7) UF15738-011027. Fruit showing stout stipe. Scale bar equals 3 mm; 8-9) UF15737-011022, 011022’. Fruit showing impression of reticulum, and compression of vascular and latex strands. Scale bar equals 3 mm; 10) UF15737-008219. Fruit showing well developed fertile carpel on the right and infertile carpel on the left. Scale bar equals 3 mm; 11) UF15737-011065. Fruit showing a complete stipe. Scale bar equals 3 mm. 12-13) UF15738-011030a, 011030'a. Part and counterpart of a specimen showing two fruits. Note stigmatic cleft (sc). Scale bar equals 3 mm; 14-15) UF15738-0052526, 0252526'. Part and counterpart of a specimen showing stigmatic cleft, reticulum, and stipe. Scale bar equals 3 mm; 16-17) UF15738-011028, 011028'. Fruit showing short stipe. Scale bar equals 3 mm.

figure22 

FIGURE 23. Dipelta bovayensis sp. nov.: 1-2) UF15737-049026, 049026'. Part and counterpart of a specimen showing a fruit with two overlapping bracts (indicated by arrow), venation, and a slender pedicel. Note obtuse apex of the top bract. Scale bar equals 3 mm; 3) Enlargement of Figure 23.1 to show reticulate venation. Scale bar equals 3 mm.

 figure23

FIGURE 24. Paleopanax boldensis sp. nov.: 1) UF15738-059493. Showing one central acrocidal capsule flanked by two lateral, younger capsules and a long peduncle. Note dehiscence scar of the capsule (indicated by white arrow). Scale bar equals 2 mm; 2) Enlargment of Figure 24.1 (indicated by black arrow) to show persistent apical style. Scale bar equals 1 mm; 3) UF15738-059493'. Counterpart of Figure 24.1 showing persistent style and dehiscence gap (indicated by arrow. Scale bar equals 2 mm; 4) Enlargement of Figure 24.3 to show the three branches of the style. Scale bar equals 1 mm.

 figure24

FIGURE 25.Solanitessaportanus (Berry) Martínez-Millán, 2010: 1) UF15737-008222. Vertically compressed flower showing six perianth parts and pedicel (indicated by arrow). Note that each perianth part has a midvein and two or three faint lateral veins. Scale bar equals 1 mm; 2) UF15737-008222’. Counterpart of Figure 25.1 showing bases of six perianth parts. Scale bar equals 1 mm.

 figure25

FIGURE 26.Solanites sarachaformis Berry, 1930: 1) UF15737-049569. Vertically preserved flower showing four floral parts each with four or five converging veins and pedicel (indicated by arrow). Scale bar equals 1 mm; 2) UF15737-049569’. Counterpart of Figure 26.1 showing a shallow floral cup formed by fusion of basal portions of the four floral parts. Note relatively strong veins extend to the center of the cup. Scale bar equals 1 mm.

 figure26

FIGURE 27. Claiborne Floral Type 5: 1-2) UF15738-008216, 008216'. Part and counterpart of a specimen showing five perianth parts and an ovary. Scale bar equals 2 mm.

 figure27

FIGURE 28. Claiborne Floral Type 7. UF15737-059208a. A flower showing three sepals (s), three petals (p), and an ovary (indicated by arrow). Scale bar equals 1 mm.

 figure28

FIGURE 29. Claiborne Floral Type 8: 1) UF15737-049570. Specimen showing seven floral parts (indicated clockwise by numbers) and a pedicel (indicated by arrow). The eighth floral part is proably positioned between 6 and 7. Scale bar equals 2 mm; 2) UF15737-049570'. Counterpart of Figure 29.1 showing two filaments (f) and the hypanthium rim (indicated by arrow). Scale bar equals 2 mm; 3) UF15737-049567a. Flower showing three floral parts of different sizes and five filaments attached to the hypanthium rim (indicated by arrow). Scale bar equals 2 mm; 4) Enlargement of Figure 29.3 to show attachment of filaments. Scale bar equals 0.5 mm.

 figure29

FIGURE 30. Claiborne Floral Type 9: 1) UF15737-059240. Flower showing five floral parts, floral rim (r), and pedicel (p). Scale bar equals 1 mm; 2) UF15737-059240'. Counterpart of Figure 30.1 showing two filaments (indicated by arrow) and impression of an ovary rim defined by two dark concentric rings with the pedicel impression at the center. Scale bar equals 1 mm; 3) UF15737-059247. Flower showing several filaments attached to the floral rim (arrow). Scale bar equals 1 mm.

 figure30

FIGURE 31. Claiborne Floral Type 10. UF15737-059243b. Vertically preserved flower showing six perianth parts, ovary (dark area), and pedicel (indicated by arrow on the right). The sixth perianth part is indicated by the arrow on the left. Scale bar equals 1 mm.

 figure31

FIGURE 32. Claiborne Floral Type 11 (1-2), UF15737-008227, 008227’. Part and counterpart of a specimen showing a flower with three floral parts and a long pedicel. Arrows point to the gap between two adjacent floral parts, indicating the possible presence of the fourth floral part. Scale bar equals 1 mm.

 figure32

FIGURE 33. Claiborne Floral Type 12: 1-2) UF15738-008217, 008217'. Part and counterpart of a specimen showing a cluster of vertically and laterally compressed flowers subtended by two large bracts or leaves. Scale bar equals 5 mm; 3) Enlargement of Figure 33.2 (indicated by arrow) to show flowers. Scale bar equals 2 mm.

 figure33

FIGURE 34. Claiborne Floral Type 13: 1-2) UF15737-059232, 059232'. Part and counterpart of a specimen showing dispersed, laterally compressed flowers. Scale bar equals 1 mm; 3) Enlargement of Figure 34.2 to show pedicel (pe), ovary (o), perianth (p), and styple (s) of the flower. Scale bar equals 0.5 mm.

figure34 

FIGURE 35. Claiborne Floral Type 14, UF15738-052528. Showing sympetalous, small flower with a cupuliform perianth tube. Note three perianth lobes and remnant of calyx (c). Scale bar equals 1 mm.

figure35 

FIGURE 36. Claiborne Floral Type 15: 1) UF15737-059251. Laterally preserved flower showing three separate perianth parts above and a receptacle (indicated by arrow). Scale bar equals 0.5 mm; 2) UF15737-059251'. Counterpart of Figure 36.1 showing a receptacle (r) and a pedicel (p). Scale bar equals 0.5 mm.

 figure36

FIGURE 37. Claiborne Floral Type 16: 1) UF15737-059226. Showing at least 18 scattered florets (arrows) compressed with an unknown floral structure. Scale bar equals 2mm; 2) Enlargement of Figure 37.1 (indicated by right arrow) to show an individual flower. Scale bar equals 0.5 mm; 3) Enlargement of Figure 37.1 (indicated by bottom arrow) to show a flower with inferior ovary and three perianth parts. Scale bar equals 0.5 mm; 4) Enlargement of Figure 37.1 (indicated by top arrow) to show a stigmatic disc. Scale bar equals 0.5 mm; 5) Enlargement of Figure 37.1 (indicated by left arrow) to show pedicel and three perianth parts. Scale bar equals 5 mm; 6-7) UF15737-059317, 059317'. Part and counterpart of a specimen showing two perianth parts, style and stigmatic disc, ovary, and pedicel. Scale bar equals 0.5 mm.

figure37 

FIGURE 38. Claiborne Floral Type 17: 1) UF15738-060634. Laterally preserved flower showing a sepal with a prominent midvein (indicated by arrow) and numerous filaments. Scale bar equals 2 mm; 2) UF15738-060634’. Counterpart of Figure 38.1 showing a petal (indicated by arrow). Scale bar equals 2 mm.

 figure38

FIGURE 39. Claiborne Floral Type 18. All Scale bar equal 1 mm: 1) UF15737-059241’a. Counterpart of a laterally preserved flower showing a short pedicel (p), fused filaments (f), anthers (a), and a perianth part (pp); 2) UF15737-059241a. Part of Figure 39.1 showing a filament spliting from the column (indicated by arrow); 3) UF15735-059323. Two laterally preserved flowers. Note strong connectives of the anthers (indicated by arrows).

 figure39

FIGURE 40. Claiborne Floral Type 19 (1-2), UF15737-059316, 059316'. Part and counterpart of a specimen showing midvein and lateral veins. Scale bar equals 0.5 mm.

 figure40

FIGURE 41. Claiborne Floral Type 20: 1) UF15737-059209. Showing stamen mass. Note distinct filament zone (indicated by arrow) and anther zone above. Scale bar equals 1 mm; 2) Enlargement of Figure 41.1 to show suborbiculate anthers. Scale bar equals 0.5 mm.

 figure41

FIGURE 42. "Sparganium" sp. : 1-2), UF15737-027592, 027592'. Part and counterpart of a specimen showing an infructescence with a stout stalk. Scale bar equals 2 mm; 3) Enlargement of Figure 42.2 to show small fruits. Scale bar equals 0.5 mm.

 figure42

FIGURE 43. Claiborne Infructescence Type 4 (1-2), UF15737-059313, 059313'. Part and counterpart of specimen showing an infructescence with sessile fruits (f) attached to the axils of bracts (b). Scale bar equals 3 mm.

 figure43

FIGURE 44. Claiborne Infructescence Type 5: 1) UF15737-059250. An infructescence with seven winged fruits (f), a receptacle (r), and a peduncle (p). Scale bar equals 1 mm; 2) UF15737-059234. Showing an infructescence with at least five fruits. Scale bar equals 0.5 mm.

 figure44

FIGURE 45. Claiborne Infructescence Type 6: 1) UF15737-059315. Showing laterally compressed infructescence with an incomplete peduncle. Note the presence of persistent perianth parts (indicated by white arrows) and pedicels on individual fruit (indicated by black arrows). Scale bar equals 1 mm; 2) Enlargement of Figure 45.1 to show trichome bases on peduncle. Scale bar equals 0.5 mm.

figure45 

FIGURE 46. Claiborne Infructescence Type 8: 1) UF15738-060633. Showing a main axis with two lateral branches and a fruit. Scale bar equals 2 mm; 2) Enlargement of Figure 46.1 showing impressions of at least 4 seeds attached to a central point. Scale bar equals 1 mm.

 figure46

FIGURE 47. Carpolithus boldensis sp. nov.: 1) UF15738-059413. Specimen showing lemma (l), awn (a), and seed body (s). Scale bar equals 1 mm; 2) Enlargement of Figure 47.1 to show midvein of lemma (indicated by arrow) and narrow, elliptic seed body. Sclae bar = 0.5 mm.

 figure47

FIGURE 48. cf. Saportaspermum sp. (1-2), UF15737-059314, 059314'. Part and counterpart of specimen showing circular seed body and veinless wing. Note radial striations on the seed body. Scale bar equals 5 mm.

 figure48

FIGURE 49.Acerknowltoni (Berry) Brown, 1940: 1-2) UF15737-059223, 059223'. Part and counterpart showing a samara. Note the protrusion with fibrous tip (indicated by arrow). Scale bar equals 2 mm; 3) A specimen from the Bolden clay pit, stored at the Paleobotany Collection at the Field Museum of Natural History, showing a samara and its venation. Scale bar equals 3 mm; 4) Enlargement of Figure 49.3 to show the smooth curvature of the inner margin of the base of the samara (indicated by arrow) and the seed outline. Scale bar equals 1 mm.

 

FIGURE 50. Claiborne Fruit/Seed Type 6: 1-2) UF15737-059237, 059237'. Part and counterpart of a specimen showing a spherical fruit/seed. Note fibers on the surface (indicated by arrows). Scale bar equals 1 mm; 3) UF15737-059242. Showing a spherical fruit/seed. Scale bar equals 1 mm; 4) Enlargement of Figure 50.3 (indicated by arrow) to show fibers associated with the fruit/seed. Scale bar equals 0.5 mm; 5) Enlargement of Figure 50.2 (indicated by arrow) to show fibers associated with the fruit/seed. Scale bar equals 0.5 mm.

 figure50

FIGURE 51. Claiborne Fruit/Seed Type 7 (1-2), UF15837-059319. Showing two possible orientations of a disseminule. It may represent a fruit with a persistent style (indicated by arrow) as in Figure 51.1 or a fruit with a pedicel (indicated by arrow) in Figure 51.2. Scale bar equals 0.5 mm.

figure51 

FIGURE 52. Claiborne Fruit/Seed Type 8 (1-2), UF15737-059245, 059245’. Part and counterpart of a specimen showing compressed ovoid sclerotesta with smooth surface and membranous endotesta (indicated by arrow in Figure 52.2). Scale bar equals 1 mm.

 figure52

FIGURE 53. Claiborne Fruit/Seed Type 9 (1-2), UF15737-059324, 059324'. Part and counterpart of a specimen showing elliptical fruit/seed with longitudinal striations. Scale bar equals 1 mm.

 figure53

FIGURE 54. Claiborne Fruit/Seed Type 10 (1-2), UF15737-059228, 059228’. Part and counterpart of a specimen showing thin layer of tissue (indicated by white arrow) and hilum scar (at the bottom of seed as indicated by black arrows). Scale bar equals 0.5 mm.

figure54 

FIGURE 55. Palmocarpon wilcoxiana (Berry) Wang et al., 2013: 1-2) UF15737-008221, 008221'. Part and counterpart of a specimen showing elliptic endocarp and fiber strands (indicated by arrow). Scale bar equals 2 mm; 3) UF15737-059231. Specimen showing elliptic endocarp. Note fine fiber strands at the distal end (indicated by arrow). Scale bar equals 2 mm; 4) UF15737-059243a. Specimen showing fiber strands/projections at the distal end (indicated by arrow). Scale bar equals 2 mm.

figure55 

FIGURE 56. Claiborne Undetermined Structure 1 (1-2), UF15737-059229, 059229'. Specimen showing a median ridge on part and a median groove on the counterpart. Scale bar equals 1 mm.

 figure56

FIGURE 57. Claiborne Undetermined Structure 2, UF15737-059297. Specimen showing a symmetric structure with a central groove. Scale bar equals 2 mm.

 figure57

FIGURE 58. Claiborne Undetermined Structure 3, UF15737-059244. Specimen showing linear, basally fused units encircling a central area. Note that the apices of the linear units (indicated by arrows) are curved into the matrix. Scale bar equals 1 mm.

 figure58

FIGURE 59. Claiborne Undetermined Structure 4, UF15737-008228. Specimen showing scale-like, irregular hexagonal surface pattern of the structure. Scale bar equals 0.25 mm.

figure59 

FIGURE 60. Salvinia preauriculata Berry, 1930; UF15737-030658. Specimen showing two floating fronds and a finely dissected, rootlike submerged frond. Scale bar equals 1 mm.

figure60 

 

 

author blanchardJane Blanchard. Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA This email address is being protected from spambots. You need JavaScript enabled to view it.

After completing a master's degree in botany at Cornell, I had the great good fortune to work in Harlan Banks' paleobotany lab. Relocating with my husband to Indiana did not result in any botanical opportunities for me so I morphed into a neuroscientist. After nearly 30 years of research work in this field, I retired to Gainesville, FL and found my way back into paleobotany at the Florida Museum of Natural History.

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author wangHongshan Wang. Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611, USA This email address is being protected from spambots. You need JavaScript enabled to view it. (corresponding author)

Hongshan Wang is a paleobotanist at the Florida Museum of Natural History, Gainesville, Florida. He received degrees from Lanzhou University (BA in Geology), China University of Geosciences (MS in Paleontology and Stratigraphy), and the University of Florida (MS in Computer Sciences and Ph.D. in Geology with a minor in Botany). He is interested in plant fossils from the Late Paleozoic to the Cenozoic.

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author dilcherDavid L. Dilcher. Departments of Biology and Geology, Indiana University, Bloomington, Indiana 47405, USA This email address is being protected from spambots. You need JavaScript enabled to view it.

 As a graduate student, I began to collect fossil plants from the Eocene of the Mississippi Embayment in the clay pits of Mississippi, Tennessee and Kentucky in 1959. After completing my Masters degree at the University of Minnesota in 1960, I moved to the University of Illinois to work with Ted Delevoryas. During my two years at Illinois, I collected several hundred fossils from the newly-opened Warman clay pit and other clay pits in the area. When Delevoryas returned to Yale in 1962, I followed him. My bulky collection of angiosperm leaves, fruits, seeds and flowers were too difficult to transport to Yale. However, in the lignitic clay of some clay pits I had found some leaves so well preserved that they could be lifted off the surface of the clay intact. On the surface of these leaves there was an abundance of well-preserved fungi. I collected several hundred of these leaves which fit in two cigar boxes. So I took these to Yale and focused on fossil fungi for my dissertation and after two years at Yale graduated with my Ph.D in 1964. The following year, as an NSF postdoctoral fellow with R. Krause at the Senckenberg Museum of Natural History in Frankfurt am Main, Germany, I again began to work on angiosperm leaves. From 1966 to 1990, while a faculty member at Indiana University, I, my family and my students continued to collect tens of thousands of fossils from the Upper Cretaceous and Eocene clay sediments of the Mississippi Embayment. In 1990 I took this collection with me to the Florida Museum of Natural History.

And for the past twenty years, I have been publishing papers that demonstrate that there is evolution in Cretaceous and Cenozoic fossil angiosperms. The concept that fossil angiosperms often represent extinct life forms at the species, generic and even at the family level was not generally accepted during my graduate student days and my early professional life. The struggle to include evolution as a part of the angiosperm fossil record has been a central theme of my research work.