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Neotropical Rainforest Origins

Permanent Link: http://ufdc.ufl.edu/UFE0022146/00001

Material Information

Title: Neotropical Rainforest Origins Paleoflora and Paleoclimate
Physical Description: 1 online resource (155 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: colombia, fossil, leaves, paleocene, paleoclimate, rainforest
Geological Sciences -- Dissertations, Academic -- UF
Genre: Geology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Little is known about the early history of Neotropical rainforests, although this vegetation types are extremely diverse, they have a distinctive spectrum of leaf size and shape, a characteristic floral composition at the family level, no fossil floras with these unique features have been reported from low latitudes yet. Here I compile a monograph on a middle to late Paleocene (58-60 Ma) flora from the Cerrej?n Formation of northeastern Colombia that is the oldest record of Neotropical rainforest. Fifty-seven fossil morphospecies were described; they include 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. The Cerrej?n assemblage has a high proportion of large, entire-margined, thin-textured leaves indicating a moist tropical climate with mean annual temperature ~24 ?C and rainfall ? 2.3 m/y. Family-level taxonomic composition is similar to that of extant Neotropical rainforests. Three monocot morphospecies were indentified at the genus level. The fossil leaves belong to the family Araceae, and they are the oldest report of this family from the Paleogene in northern South America. The three leaf taxa are related to two extant Neotropical genera, two of them closely related to Anthurium and the other fossil taxa was placed within the extant genus Montrichardia. The fossil leaves provide new information about patterns of angiosperm diversification in the Neotropics.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Dilcher, David L.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0022146:00001

Permanent Link: http://ufdc.ufl.edu/UFE0022146/00001

Material Information

Title: Neotropical Rainforest Origins Paleoflora and Paleoclimate
Physical Description: 1 online resource (155 p.)
Language: english
Publisher: University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 2008

Subjects

Subjects / Keywords: colombia, fossil, leaves, paleocene, paleoclimate, rainforest
Geological Sciences -- Dissertations, Academic -- UF
Genre: Geology thesis, M.S.
bibliography   ( marcgt )
theses   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
born-digital   ( sobekcm )
Electronic Thesis or Dissertation

Notes

Abstract: Little is known about the early history of Neotropical rainforests, although this vegetation types are extremely diverse, they have a distinctive spectrum of leaf size and shape, a characteristic floral composition at the family level, no fossil floras with these unique features have been reported from low latitudes yet. Here I compile a monograph on a middle to late Paleocene (58-60 Ma) flora from the Cerrej?n Formation of northeastern Colombia that is the oldest record of Neotropical rainforest. Fifty-seven fossil morphospecies were described; they include 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. The Cerrej?n assemblage has a high proportion of large, entire-margined, thin-textured leaves indicating a moist tropical climate with mean annual temperature ~24 ?C and rainfall ? 2.3 m/y. Family-level taxonomic composition is similar to that of extant Neotropical rainforests. Three monocot morphospecies were indentified at the genus level. The fossil leaves belong to the family Araceae, and they are the oldest report of this family from the Paleogene in northern South America. The three leaf taxa are related to two extant Neotropical genera, two of them closely related to Anthurium and the other fossil taxa was placed within the extant genus Montrichardia. The fossil leaves provide new information about patterns of angiosperm diversification in the Neotropics.
General Note: In the series University of Florida Digital Collections.
General Note: Includes vita.
Bibliography: Includes bibliographical references.
Source of Description: Description based on online resource; title from PDF title page.
Source of Description: This bibliographic record is available under the Creative Commons CC0 public domain dedication. The University of Florida Libraries, as creator of this bibliographic record, has waived all rights to it worldwide under copyright law, including all related and neighboring rights, to the extent allowed by law.
Thesis: Thesis (M.S.)--University of Florida, 2008.
Local: Adviser: Dilcher, David L.

Record Information

Source Institution: UFRGP
Rights Management: Applicable rights reserved.
Classification: lcc - LD1780 2008
System ID: UFE0022146:00001


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NEOTROPICAL RAINFOREST ORIGINS: PALEOFLORA AND PALEOCLIMATE


By

FABIANY ALBERTO HERRERA-TOLOSA




















A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2008


































2008 Fabiany Alberto Herrera-Tolosa




























To my family, my mom, my dad, my nephew Joseph and my niece Sofia for their happiness. To
my friends









ACKNOWLEDGMENTS

This research was supported by grants from Carbones del Cerrej6n, a graduate research

fellowship of the University of Florida, NSF Deep Time grant DEB-0090283 to DLD, NSF grant

DEB-0733725, Smithsonian Paleobiology Endowment Fund, the Unrestricted Endowments SI

Grants, the Fondo para la Investigaci6n de Ciencia y Tecnologia del Banco de la Republica de

Colombia, The Explorers Club, Colombian Petroleum Institute, and Corporaci6n Geol6gica

ARES. I deeply thank David Dilcher, Carlos Jaramillo, Scott Wing and Carolina Gomez for their

helpful discussions and contributions. Steven Manchester, Ellen Martin and Mark Brenner for

comments improving the manuscript. Fernando Chavez, Camilo Montes, and the geology team at

the Cerrej6n mine for their help and support during field trips. I thank Dan Nicolson for helping

at the National Herbarium and Stanley Yankowski for his assistance clearing the extant

Montrichardia leaf. I thank Josef Bogner, Marcela Mora, and Thomas Croat for helpful

discussions about the systematics of the fossils, and Giovanni Bedoya for his suggestions about

the nomenclature. I thank everybody in the Paleobotany lab for their support, finally special

thanks go to Edwin Cadena for his friendship and Benjamin Himschoot for reading part of this

thesis and such great happiness.










TABLE OF CONTENTS

page

A C K N O W L E D G M E N T S ..............................................................................................................4

LIST OF TABLES ............................ ............ ........ ..................7

LIST O F FIG U RE S ................................................................. 8

ABSTRAC T ........................................................................ 9

CHAPTER

1 MONOGRAPH OF THE CERREJON FLORA ........................................... ............... 10

A b stra c t ................... ...................1...................0..........
In tro d u ctio n ................... ...................1...................0..........
M materials and M methods ..................................... .............................. .. ... ......... 11
F o ssil L eaf D description s .......................................................................... .......................... 15
Ferns .......................................15
Cycads .........................................17
C o n ife r s ...................................................................................................................... 1 7
D ic o ts ................... ...................1...................8..........
M o n o c o ts .................................................................................................................... 4 8
Paleoclimate........................ ............................... ........ 55
Floristic Com position of the Cerrej6n Flora .......................................................... ............ 56
C onclu sions..... ..........................................................57

2 FOSSIL ARACEAE FROM A PALEOCENE NEOTROPICAL RAINFOREST IN
C O L O M B IA .............. .. .................................................................5 9

A b stra ct .......................................... ... .....59..........
In tro d u c tio n ............................................ ........................................................................... 5 9
M materials an d M eth o d s ...................................................................................................... 6 1
Systematics .........................................62
D iscu ssion ................................. .............................2
Sedim entary D position and Paleoclim ate ................................................. ...............72
P a le o e c o lo g y ............. .. ....... .. .............. .. ...................................................7 3
Paleobiogeography and Palynology ........................................ ................. 74
Comments on Aroid Evolution...................................... ......... 76

APPENDIX

A PHOTOGRAPHIC PLATES ............. .......... ..... ............... 80

LIST OF REFERENCES ...................................................... 147









B IO G R A PH IC A L SK E T C H ............................................................................... ............... ..... 155










LIST OF TABLES


Table


2-1 Morphological features of fossil leaf taxa and extant Araceae ......................................78


page









LIST OF FIGURES


Figure page

1-1 Map and location of Cerrej6n mine and Cerrej6n flora localities (stars). .........................13

1-2 Stratigraphic column of the Cerrej6n Formation showing lithology and stratigraphic
positions of the plant megafossil localities. Modified from Bayona et al (2004).............14









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

NEOTROPICAL RAINFOREST ORIGINS: PALEOFLORA AND PALEOCLIMATE

By

Fabiany Alberto Herrera-Tolosa

May 2008

Chair: David Dilcher
Major: Geology

Little is known about the early history of Neotropical rainforests, although this vegetation

types are extremely diverse, they have a distinctive spectrum of leaf size and shape, a

characteristic floral composition at the family level, no fossil floras with these unique features

have been reported from low latitudes yet. Here I compile a monograph on a middle to late

Paleocene (58-60 Ma) flora from the Cerrej6n Formation of northeastern Colombia that is the

oldest record of Neotropical rainforest. Fifty seven fossil morphospecies were described; they

include 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. The Cerrejon assemblage has a

high proportion of large, entire-margined, thin-textured leaves indicating a moist tropical climate

with mean annual temperature -24 C and rainfall > 2.3 m/y. Family-level taxonomic

composition is similar to that of extant Neotropical rainforests.

Three monocot morphospecies were identified at the genus level. The fossil leaves belong

to the family Araceae, and they are the oldest report of this family from the Paleogene in

northern South America. The three leaf taxa are related to two extant Neotropical genera, two of

them closely related to Anthurium and the other fossil taxa was placed within the extant genus

Montrichardia. The fossil leaves provide new information about patterns of angiosperm

diversification in the Neotropics.









CHAPTER 1
MONOGRAPH OF THE CERREJON FLORA

Abstract

Little is known about the early history of Neotropical rainforests, although this vegetation

types are extremely diverse, they have a distinctive spectrum of leaf size and shape, a

characteristic floral composition at the family level, no fossil floras with these unique features

have been reported from low latitudes yet. Here I compile a monograph on a middle to late

Paleocene (58-60 Ma) flora from the Cerrej6n Formation of northeastern Colombia that is the

oldest record of Neotropical rainforest. Fifty seven fossil morphospecies were described; they

include 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. The Cerrej6n assemblage has a

high proportion of large, entire-margined, thin-textured leaves indicating a moist tropical climate

with mean annual temperature -24 C and rainfall > 2.3 m/y. Family-level taxonomic

composition is similar to that of extant Neotropical rainforests.

Introduction

Rainforests in tropical America have the highest plant diversity of any region in the world

(Morley, 2000), and much of this plant diversity occurs in the flowering plants. However, the

paleobotanical fossil record from the tropics have remained scarce and nearly unexplored,

leaving the origin of the Neotropical rainforest as one the biggest mysteries in plant evolution

and paleoecology.

An exhaustive review of the fossil record in northern South America (Burnham and

Graham, 1999; Dilcher, 2000; Burnham and Johnson, 2004) suggests that the Neotropical

rainforests could have appeared during the early Cenozoic (-50-65 million years ago). This

evidence comes mainly from the fossil pollen record (Jaramillo et al., 2006). But this

fragmentary fossil record still has not addressed many questions: when did the Neotropical









rainforest appear in South America? Under what climatic conditions did those ancient rainforests

grow? Did ancient rainforests have the leaf morphologic characters seen in modem rainforests?

What were the first families that inhabited the Neotropical rainforest? Did the families that are

dominant in the Neotropical rainforest today originate in South America or did they immigrate

from other latitudes?

Here, I report on a middle-late Paleocene flora from Northern Colombia: Cerrejon

paleoflora that is the oldest record of a Neotropical rainforest. A complete description of the leaf

fossils, a paleoclimatic reconstruction based on the fossil leaf morphology and their implications

on the Paleocene climate, and finally a review of the floristic composition are presented in this

monograph.

Materials and Methods

Fossil plants from the Cerrej6n mine were first described by Doubinger and Pons in 1970

which consisted of a report on the remains of fossil leaf cuticle. After this report there was no

additional research on the Cerrej6n fossil plants until they were collected during 2003-2007. The

Cerrej6n Formation is exposed in the Cerrej6n mine, located in Northern Colombia (Guajira

Peninsula, Fig. 1-1). The Paleogene age of the Cerrej6n flora is based upon the palynological

assemblages, which demonstrates that the pollen flora belong to the Foveotricolpitesperforatus

zone and to zone Cu-02 (Bayona et al., 2004; Jaramillo et al., 2007) based on the presence of

Foveotricolpites perforatus and Bombacacidites annae throughout the interval. These zones are

considered to be middle to late Paleocene, based on their co-occurrence with marine microfossils

in shallow marine rocks and stable carbon isotope stratigraphy.

The -800-m-thick stratigraphic sequence of Cerrej6n Formation is composed of thick

coals, fluvially deposited sandstones, and lacustrine siltstones that were deposited in a rapidly

subsiding basin (Bayona et al., 2004). Most of the Cerrej6n Formation consists of a depositional









cycle of three parts (Bayona et al., 2004). The lowest part is fossiliferous black shale and

laminated black mudstone with thin lenticular laminae of sandstone (interpreted as anoxic

lagoonal, or flooded coastal-plain environments). The middle part of the type section includes

slightly bioturbated mudstones and sandstones with flaser and heterolithic lamination and

dispersed plant remains, interpreted as deposits of subtidal and tidal flats. The upper part of the

type section is dominated by fine-grained, massive to lenticularly-laminated, bioturbated

mudstones and siltstones with abundant plant remains, which are cut by thick to very thick

massive to cross-bedded sandstones (interpreted as coastal plains crossed by channels). The

thickest and extensive coal beds in the Cerrej6n Formation occur in the middle part of the

section, these coal deposits have low-ash and low-sulfur contents.

The megafloral localities occur from 100-750 meters above the base of the Cerrejon

Formation (Fig. 1-2). In total, I recovered 1347 megafossil specimens. The collections of fossil

plants were made from six lens-shaped, siltstone/sandstone bodies with inclined heterolithic

strata that suggest deposition in low-energy channels (Fig. 1-2; localities 0410, 0317, 0315,

0322, 0319, and 0323). Two other localities preserve leaves in thin but tabular beds of flat-

laminated siltstone that indicate deposition in small lakes (Fig. 1-2; localities 0318 and 0324).

The classification and description of fossil leaf morphotypes is based upon leaf

architectural features such as the characters presented in the Leaf Architecture Working Group

Manual (LAWG, 1999). The botanical affinities suggested for some of the leaf morphotypes

were determined after extensive comparisons with cleared leaves of living plants in the National

Cleared Leaf Collection held in the Department of Paleobiology, National Museum of Natural

History, Smithsonian Institution, the modern reference Leaf Collection of Florida Museum of

Natural History, and the Herbarium of the Smithsonian Tropical Research Institute.

































311 ,
! !!






All .-- --


Figure 1-1. Map and location of Cerrejon mine and Cerrej6n flora localities (stars).





















































13
















coal 175


coal 155


coal 150







coal 115


0319 coal 110
0322
400
0315 i coal 100
0317


500




600
coal 45


0410 coal 40
700





cooal
= oalsandstone
sandy mudstone-siltstone
claystone-shales
r limestones


Figure 1-2. Stratigraphic column of the Cerrej6n Formation showing lithology and stratigraphic
positions of the plant megafossil localities. Modified from Bayona et al (2004).









Fossil Leaf Descriptions


Ferns

CJ42 (Plate 1)

Diagnosis. Blade elliptic and symmetrical, emarginated apex. Tertiaries forming

quadrangular areolation. Entire margin.

Description. The only specimen found reaches -2 cm of length and width of 1.7 cm; the

specimen is missing part of the leaf base, a midvein is visible with a curved course that ends in

an emarginated apex. Tertiary veins forming a very tiny quadrangular areolation. The margin is

entire.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal

beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, two specimens.

Systematic affinity suggested. The leaf closely resembles the pantropical, floating-aquatic

fern Salvinia (Salviniaceae)

CJ57 (Plate 2)

Diagnosis. Pinnate leaf, secondary veins are craspedodromous, decurrent on midvein, and

closely spaced. Irregular polygonal reticulate tertiaries. Serrate margin, cc/st and cc/cv teeth with

long basal sides.

Description. The specimen is not complete but reaches -16 cm in length and a maximum

width -6 cm. Leaf texture coriaceous. Laminar shape oblong. Secondary veins are numerous,

very closely spaced (0.5 mm), decurrent on midvein, and craspedodromous. The margin is

serrate, teeth have long basal sides and the morphology varies between apical sides concave and

basal sides straight and convex. Tertiary veins are the highest order veins present on the









specimen, these veins are irregular polygonal reticulate aerolations with elongate and rounded

shapes.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0322, Pit La Puente, locality placed between coal beds

103 and 105, middle to late Paleocene, coordinates 11.9 N, 72.3 W, one specimen.

Systematic affinity suggested. The specimen closely resembles members of the family

Stenochlaenaceae, common climbing ferns in Old World tropical swamps.

CJ61 (Plate 3)

Diagnosis. Lobate leaf with many veins of equal gauge branching toward apex.

Description. The only specimen found has a maximum length of 2 cm and a maximum

width of 3 cm (measured apically); leaf shape resembles a palmate leaf with four asymmetrical

lobes preserved, their basal sinus departs either very to close to the base of the blade or from

more apical sides; the lobes miss prominent midveins. Many thin veins depart from the base and

branch in random patterns toward apex.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0322, Pit La Puente, locality placed between coal beds

100 and 115, middle to late Paleocene, one specimen.

Systematic affinity suggested. The specimen closely resembles members of the

cosmopolitan genus Lygodium.

CJ66 (Plate 4)

Diagnosis. Pinnate leaf. Secondaries anastomose near the midvein to form relatively short

cells, and then these cells become more elongated toward the margin of the leaf. Secondaries are

weekly decurrent on mid-rib.









Description. The specimen is about 8 cm in length and the maximum width is about 4 cm;

laminar shape elliptic; margin entire; primary vein thick; secondary veins thin, weekly decurrent

on midvein and anastomose near midvein to form relatively short cells, then the cells become

more elongated toward the margin of the leaf.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0322, Pit La Puente, locality placed between coal beds

103 and 105, middle to late Paleocene, coordinates 11.9 N, 72.3 W, two specimens.

Systematic affinity suggested. The leaf closely looks like Acrostichum, a pantropical fern

common in freshwater and brackish mangrove swamps today.

Cycads

CJ46 (Plate 5)

Diagnosis. Rachis stout; leaflets non-articulate, decurrent at the base, opposite; a midrib is

absent from the leaflets. Secondary veins are thin and parallel each other. Entire margined.

Description. Fragment of a leaves that reach -15 in length and -18 in width. Rachis is

about 3 cm width. The maximum number of leaflets preserved on the same specimen are sixteen;

leaflets more than 22 mm wide, opposite, non-articulate, decurrent at the base and lack a

midvein. Secondary veins are numerous, closely spaced, and parallel each other. Entire

margined.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0319, Pit Tabaco High Dip, locality placed below coal

bed 110, middle to late Paleocene, coordinates 11.66 N, 73.31W, three specimens.

Systematic affinity suggested. The cycad is similar to extant Dioon.

Conifers

CJ60 (Plate 6)









Diagnosis. Shoot of conifer with scale leaves.

Description. Length -11 cm and maximum width reaches -0.8 cm; scales leaves are

crowded and overlapped through the axes; scales arranged in a spiraled course. Maximum length

of the scales reach -5-7 mm and maximum width measured -4-5 mm.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, three specimens.

Systematic affinity suggested. The shoot closely resembles members of living

Taxodiaceae and Araucariaceae.

Dicots

CJ1 (Plate 7)

Diagnosis. Petiole transversely striated, short and pulvinate. Secondary angle to primary

increases basally, and secondary veins fork at low angles near margin. Tertiaries closely spaced

and mostly opposite percurrent.

Description. Laminar size range from notophyll to macrophyll; leaves elliptic, oblong and

obovate; Petiole transversely striated, short and pulvinate; base shapes from convex, concave and

weekly asymmetrical; apex rounded and acuminate; margin entire; unlobed; leaves pinnate;

secondary veins brochidodromous, between 19-22 pair of 2's; Secondary vein angle to primary

increases basally, and secondary veins fork at low angles near margin; tertiary veins mostly

opposite percurrent with courses straight and convex and their angles respect to the primary

range from obtuse to perpendicular; fourth order veins mostly alternate percurrent, higher order

veins regular polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0315; 0317, 0318, 0319, 0322, 0323, and 0410; Pits









Tabaco 1, Tabaco High Dip, La Puente, Expanded West, localities placed between coal beds 40

and 161, middle to late Paleocene, 173 specimens.

Systematic affinity suggested. Based on the leaf morphology and the presence of a petiole

transversely striated, short and pulvinate the morphotype closely resembles the family

Leguminosae.

CJ2 (Plate 8)

Diagnosis. Strongly asymmetrical blade. Secondary veins are weak brochidodromous at

the base and brochidodromous apically. Secondary vein angle on primary increases apically to

almost 900. Falcate apex. Thick margin.

Description. Laminar size range from notophyll to mesophyll, laminar shape elliptic;

strongly asymmetrical blades; base shape complex; apex falcate; margin entire; leaves pinnate;

secondary veins are weak brochidodromous at the base and brochidodromous apically, between

23 and 18 pairs; the angle of the 2's on primary increase apically to almost 900; inter-secondary

veins are week; tertiary veins are alternate percurrent; the angle of the 3's on primary veins range

from perpendicular to acute; a thick margin is visible easily.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0317 and 0318; Pits Tabaco 1 and Tabaco High Dip;

localities placed between coal beds 100 and 161, middle to late Paleocene, coordinates 11.13N,

72.57W and 11.12 N, 73.55 W, 8 specimens.

Systematic affinity suggested. The fossil leaves seem similar to living species within the

family Meliaceae.

CJ4 (Plate 9)









Diagnosis. Decurrent base. Leaf shape narrower than CJ32. Semicraspedodromous

secondaries, they become acute basally more than in CJ32. Tertiary veins opposite percurrent

and about 90 degrees to primary, epimedial tertiaries forming intersecondaries. Fourth order

veins mostly irregular polygonal and alternate percurrent. Teeth closely spaced and have long

basal sides.

Description. Long petioles; laminar shapes from microphyll to mesophyll; laminar shape

elliptic to oblong; base shape strongly decurrent; margin entire; leaves pinnate; secondary veins

are semicraspedodromous, between 23 to 24 pairs of 20s; the secondaries become acute basally,

secondary vein spacing decreases toward base; intersecondary veins are weak on the specimens;

tertiary veins are opposite percurrent with sinuous to straight courses, the angle of the 3's on

primary is mainly perpendicular to it; higher order veins range from irregular to regular

polygonal reticulate; areolation is well developed; margin type crenate, teeth are closely spaced

and have short convex apical sides and long convex basal sides; leaf texture is chartaceous.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0317 and 0318; Pits Tabaco 1 and Tabaco High Dip;

localities placed between coal beds 100 and 161, middle to late Paleocene, coordinates 11.13N,

72.57W and 11.12 N, 73.55 W, 24 specimens.

Systematic affinity suggested. No extant affinity identified.

CJ5 (Plate 10)

Diagnosis. Suprabasal or basal acrodromous, two lateral primaries running in convergent

arches toward the leaf apex. Eucamptodromous 20s. One secondary vein running just inside the

margin. Two specimens covered with small laminar resin dots.









Description. Laminar size range from microphyll to macrophyll; laminar shape elliptic;

base shape from concavo-convex to convex; apex acuminate; margin entire; primary veins are

either suprabasal or basal acrodromous (three veins only); costal secondary veins are

eucamptodromous, at about 10 pairs of 2's, one secondary vein runs from the base parallel and

close to the margin as an intramarginal veins which collects the costal 2's, the spacing of the

costal secondaries decreases toward base; tertiary veins are mixed opposite/alternate percurrent,

the course of the opposite ones range from convex to straight and their angles on primaries are

obtuse; the variability of the angle of 3's increase basally (undercostals) and increase exmedially

(epimedial); fourth order veins are alternate percurrent; a fimbrial vein is present along the

margin, only two specimens are covered with small laminar resin dots.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0317, 0319, and 0410; Pits Tabaco 1, Tabaco High

Dip, and Expanded West, localities placed between coal beds 40 and 107, middle to late

Paleocene, 22 specimens.

Systematic affinity suggested. Leaves with suprabasal or basal acrodromous venation and

covered with resin glands are very common in the family Lauraceae. The fossil leaves closely

resemble this family.

CJ6 (Plate 11)

Diagnosis. Deeply cordate or truncate base. Brochidodromous 20s that loop just inside the

margin both costal and minor secondaries. Opposite percurrent tertiaries concentric. Fifth and

sixth order veins are regular polygonal reticulate. Tiny and squarish areoles. Thick fimbrial vein.

Description. Petiole with wiry thickening; leaves size range from microphyll to

macrophyll; laminar shape ovate; base shape are deeply cordate, cordate and truncate; apex









acuminate, margin entire; primary veins are basal actinodromous with 5 to 7 veins; agrophic

veins compound; intersecondary veins are absent; tertiary veins mostly opposite percurrent with

straight to strongly convex courses, the angle of the 3's on primaries is mostly perpendicular to

it, the tertiary veins angle variability decreases exmedially and basally; fourth order veins are

strongly alternate percurrent; higher order veins (5th and 6th orders) regular polygonal reticulate;

areolation is well developed, tiny, and squarish (3-4 sided); a thick fimbrial veins is present

along the margin.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315; 0317, 0318, 0319, 0322, and 0323; Pits Tabaco

1, Tabaco High Dip, and La Puente, localities placed between coal beds 100 and 161, middle to

late Paleocene, 68 specimens.

Systematic affinity suggested. Based on the actinodromous leaf venation, blade shape,

higher order veins and the presence of a thick fimbrial vein the morphotype closely resembles

the family Menispermaceae.

CJ8 (Plate 12)

Diagnosis. Primary vein and petiole very thick. Secondaries become more obtuse basally

(almost 90), and 3-4 pairs crowded, secondary vein spacing decreases toward base but apically

increases. Tertiaries thin in mid-course.

Description. Petiole very thick, sometimes this reaches at about 2 cm in width; laminar

size range from notophyll to macrophyll; laminar shape between elliptic but mostly obovate

leaves; base shape concave; apex acuminate to rounded; margin entire; leaves pinnate, the

midvein is stout; secondary veins are eucamptodromous; secondary vein spacing crowded

basally or decreases toward base but apically increases; secondary veins also become more









obtuse basally (almost 900); tertiary veins mostly opposite percurrent with straight to sinuous

courses, their angles on primaries are perpendicular to acute; the tertiary vein angle variability on

the rest of the blade increases exmedially and basally; tertiary veins are as well thin compared to

secondaries in mid-course; higher order veins are regular polygonal reticulate and not strongly

visible.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315; 0317, 0318, 0319, and 0323; Pits Tabaco 1,

Tabaco High Dip, and La Puente, localities placed between coal beds 100 and 161, middle to late

Paleocene, 36 specimens..

Systematic affinity suggested. Leaves with these characters occur in several families such

as Sapotaceae, Bombacaceae, and Annonaceae.

CJ10 (Plate 13)

Diagnosis. Primary vein is very thick at the base and thins distally and develops a zig-zag

course with the primary being deflected at each secondary juncture. Brochidodromous 2s

forming prominent arches almost festooned brochidodromous. Poorly organized tertiaries.

Fimbrial vein.

Description. Petiole is very long, reaches at about 5 cm in lenght; laminar size mesophyll;

laminar shape elliptic; base shape varies between concave-convex and convex; apex shape either

retuse or convex; margin entire; leaves pinnate, midvein very thick at the base and thins distally

and develops a zig-zag course with the primary being deflected at each secondary juncture;

secondary veins are brochidodromous, these veins form prominent arches almost like festooned

brochidodromous venations, 6 to 7 pairs of 2's, sometimes two or three of these veins depart

from the base of the leaf; secondary vein spacing decreases toward base and the angle of these









veins smoothly increases toward base; tertiary veins are mixed opposite/alternate percurrent but

poorly organized; tertiary vein angle variability decreases apically and more acute relative to the

midvein; higher order veins are irregular and regular polygonal reticulate; areolation well

developed (3-4 sided); a fimbrial vein runs parallel to margin.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0317, 0318, and 0323; Pits Tabaco 1 and Tabaco High

Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 13 specimens.

Systematic affinity suggested. No extant affinity identified.

CJ11 (Plate 14)

Diagnosis. Distinctly lobate leaf, thick primary veins. Brochidodromous 20s closely

spaced, and forming loops at the margin.

Description. Laminar size occurs from mesophyll to macrophyll; laminar shape elliptic;

apex angle odd lobed acute; apex acuminate; margin entire; primary veins are basal

actinodromous, at least 7 veins are recognized in one of the specimens; at least three lobes are

preserved; agrophic veins are simple; costal secondary vein spacing decreases toward base;

tertiary veins are mixed opposite/alternate percurrent.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, seven specimens.

Systematic affinity suggested. Three-lobate leaves with entire margin are common in the

order Malvales.

CJ12 (Plate 15)









Diagnosis. Tertiaries are alternate percurrent and form composite intersecondaries,

Tertiaries admedially ramified on the midvein. Specimens covered with irregular black dots.

Description. Petiole with base swollen; laminar size mesophyll; laminar shape elliptic;

base shape convex; margin entire; leaves pinnate; secondary veins are eucamptodromous;

secondary vein spacing is weekly irregular through the blade, 2's vein angle on primary

smoothly increases towards apex; tertiary veins are strongly alternate percurrent and form

composite intersecondaries, then the tertiaries ramify admedially on the midvein, the angle of the

3's respect to the midvein varies between perpendicular and acute; higher order veins are regular

polygonal reticulate; the specimens are covered with irregular black dots.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315 and 0323; Pit Tabaco 1 and Tabaco High Dip,

localities placed between coal beds 100 and 120, middle to late Paleocene, seven specimens.

Systematic affinity suggested. Some characters such as type of tertiary veins and its

courses are similar to those seen in the families Moraceae and Ulmaceae, but more characters are

necessary to prove this affinity.

CJ13 (Plate 16)

Diagnosis. Petiole with base swollen and short. 13-14 pairs of weakly brochidodromous

20s, secondaries diverge from one another exmedially. Tertiaries are opposite percurrent, mostly

convex or sinuous and wider spaced than in CJ1.

Description. Petiole short with a swollen base; laminar size varies between mesophyll and

macrophyll; laminar and base shape constantly elliptic and convex respectively; apex shape from

acuminate to convex; margin entire; leaves pinnate; secondary veins are weak brochidodromous

looping very close to margin, 10 to 13 pairs of secondaries that diverge from one another









exmedially with a course almost sinuous; secondary vein spacing is uniform, and secondary vein

angle smoothly increases towards base; tertiary veins are opposite percurrent with strong convex-

sinuous course, the angles of the 3's on primaries go from obtuse to perpendicular, tertiary vein

angle variability increases exmedially; higher order veins regular polygonal reticulate; areolation

moderately developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0315, 0317, 0323 and 0410; Pit Tabaco 1, Tabaco

High Dip and Expanded West, localities placed between coal beds 40 and 120, middle to late

Paleocene, 16 specimens.

Systematic affinity suggested. No extant affinity identified.

CJ15 (Plate 17)

Diagnosis. Similar to CJ8 but secondaries are not perpendicular near base of primary,

secondary veins spacing increases toward (2/3) of blade and then decreases. Fourth order veins

are regular polygonal reticulate.

Description. Petiole thick and short; laminar size from mesophyll to megaphyll; laminar

shape obovate to elliptic; base shape cuneate; apex shape is either rounded or acuminate; margin

entire; leaves pinnate; secondary veins are eucamptodromous, between 20 to 25 pairs of 2's;

secondary vein spacing increases toward (2/3) of blade and then decreases, and secondary vein

angle smoothly decreases towards base; tertiaries mostly opposite percurrent with straight to

sinuous courses and their angles on primary range from perpendicular to obtuse; tertiary vein

angle variability decreases exmedially; higher order veins regular polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0315; 0317, 0318, 0323, and 0410; Pits Tabaco 1,









Tabaco High Dip, Expanded West, localities placed between coal beds 40 and 161, middle to late

Paleocene, 28 specimens.

Systematic affinity suggested. Leaves with these characters occur in several families such

as Anacardiaceae, Apocynaceae, and Sapotaceae.

CJ16 (Plate 18)

Diagnosis. Pinnate with cordate base. Secondaries (3-4 pairs) crowded at the base,

secondary angle at the base is more obtuse than CJ28. More closely spaced opposite percurrent

tertiaries than CJ28.

Description. Petiole long, reaches at about 6 cm; laminar size mesophyll; laminar shape

constantly ovate; base shape cordate; apex shape acuminate; margin entire; leaves pinnate;

secondary veins are eucamptodromous (13-15 pairs of 2's) having five to six of those veins

crowded and departing from the base; secondary vein spacing decreases toward base, and their

angles abruptly increase toward base; tertiary veins are opposite percurrent, closely spaced each

other and having straight to sinuous courses, their angles on primary are obtuse; the tertiary vein

angle variability decreases exmedially and apically; higher order veins probably regular

polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0318 and 0319, Pit Tabaco High Dip, localities placed

between coal beds 105 and 161, middle to late Paleocene, five specimens.

Systematic affinity suggested. No extant affinity identified.

CJ17 (Plate 19)

Diagnosis. Pinnate with slightly cordate base with 5 basal veins. Compound agrophic

veins. Tertiaries mixed opp/alt percurrent. Areolation very small and squarish.









Description. Long petiole, reaches at about 5.5 cm; laminar size and shape mesophyll and

elliptic respectively; base shape slightly cordate; apex shape acuminate; margin entire; leaves

pinnate; secondary veins weak brochidodromous and looping very close to margin, five

secondaries are crowded at the base; secondary vein spacing apparently decreases toward base,

and the angle seems to increase toward base; agrophic veins are compound; tertiary veins are

mixed opposite/alternate percurrent with courses sinuous to straight, and their angles on primary

vein are perpendicular to obtuse; fourth order veins are alternate percurrent; higher order veins

regular polygonal reticulate; areolation well developed, small, and squarish.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315 and 0318; Pit Tabaco 1 and Tabaco High Dip,

localities placed between coal beds 100 and 61, middle to late Paleocene, three specimens.

Systematic affinity suggested. Pinnate leaves with cordate bases common in the order of

Malvales.

CJ18 (Plate 20)

Diagnosis. Like CJ8 or CJ15 but about half as many secondaries ( 8-10 pairs), widely

spaced secondaries, and the angle of 2s smoothly increasing toward base, one pair acute basal

secondaries. Percurrent tertiaries.

Description. Petiole long (5.3 cm) and thin (0.5 cm); laminar size and shape mesophyll

and elliptic respectively; base shape concave-convex and decurrent; margin entire; leaves

pinnate; secondary veins are eucamptodromous, at least eleven pair of 2's are preserved;

secondary vein spacing is widely spaced, and the secondary vein angle smoothly increases

toward base; tertiary veins are opposite percurrent with straight to sinuous courses; 4th and 5th

order veins are alternate percurrent and opposite percurrent respectively.









Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, seven specimens.

Systematic affinity suggested. No extant affinity identified.

CJ19 (Plate 21)

Diagnosis. Pulvinate petiole. Very large number of closely spaced secondaries that are

decurrent on the primary and arise from it at about 90 degrees. Tertiaries are random reticulate

and also decurrent on primary.

Description. Petiole pulvinate; laminar size from microphyll to mesophyll; laminar shape

elliptic; base shape convex and rounded; apex acuminate; margin entire; leaves pinnate;

secondary veins are eucamptodromous, numerous, closely spaced, and decurrent on the primary

and they arise from it at about 90 degrees; secondary vein angle increases toward base, and their

angles smoothly increase toward base; intersecondary veins are common, usually one to two

pairs per secondaries; tertiary veins are random reticulate and as well decurrent on primary; their

angles on primary range from perpendicular to acute; tertiary vein angle variability is

inconsistent' 4th and 5th order veins are irregular and regular polygonal reticulate

correspondingly; areolation is well to moderately developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0315; 0317, 0318, 0319, 0323 and 0324; Pits Tabaco

1, Tabaco High Dip, La Puente, localities placed between coal beds 100 and 161, middle to late

Paleocene, 22 specimens.

Systematic affinity suggested. Based on the secondary and tertiary venation and the

presence of a pulvinate petiole the morphotype closely resembles the family Leguminosae.









CJ20 (Plate 22)

Diagnosis. Like CJ22 in major venation, but fourth and fifth order veins less distinct than

in CJ22. Secondary spacing mostly irregular. Tertiaries with minutely irregular course, leaf rank

2r.

Description. Laminar size from mesophyll to macrophyll; laminar shape elliptic; base and

apex shape convex; margin entire; leaves pinnate; secondary veins are eucamptodromous, 10-15

pairs of 2's; secondary veins spacing mostly irregular, and their angles smoothly increase toward

base; intersecondary veins occur but they are weak; tertiary veins mostly mixed

opposite/alternate percurrent with minutely irregular course, and their angles respect of the

primary oscillate between perpendicular to acute; tertiary vein angle variability increases

exmedially and apically; 4th and 5th order veins are irregular and regular polygonal reticulate;

areolation well developed, leaf rank 2r.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315; 0318, 0319; Pits Tabaco land Tabaco High Dip,

localities placed between coal beds 100 and 161, middle to late Paleocene, nine specimens.

Systematic affinity suggested. No extant affinity identified.

CJ21 (Plate 23)

Diagnosis. Decurrent base. Very closely spaced (-1 mm) opposite percurrent tertiaries,

course mostly sinuous, angle to primary strongly perpendicular and increases exmedially.

Description. Laminar size and shape mesophyll and elliptic correspondingly; base angle at

almost 90 degrees and base shape very decurrent; apex acuminate; margin entire; leaves pinnate;

secondary veins are eucamptodromous, at least 12 pairs of 2's are preserved on the specimens;

secondary vein angle increases from base until 2/3 length of leaf and their angles are uniform;









tertiary veins are constantly opposite percurrent and very closely spaced (-lmm), with course

sinuous and the angle of the 3's on primary vein is strongly perpendicular; tertiary vein angle

variability increases exmedially, 4th vein order are irregular polygonal reticulate; areolation is

moderately developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315; 0318, 0323; Pits Tabaco land Tabaco High Dip,

localities placed between coal beds 100 and 161, middle to late Paleocene, 10 specimens.

Systematic affinity suggested. Some species in Chrysobalanaceae and Annonaceae have

similar characters as the fossil leaf.

CJ22 (Plate 24)

Diagnosis. Between 7 and 14 pairs of secondaries, crowded basal secondaries are

decurrent and more acute than apical secondaries. Fourth and fifth order veins are alternate

percurrent and regular polygonal reticulate unlike CJ20. Most specimens covered with laminar

resin glands (<0. mm). Leaf rank 3r.

Description. Laminar size vary from notophyll to macrophyll leaves; laminar shape

principally elliptic; base shape from convex, concave to weakly decurrent; apex shape

acuminate; margin entire; leaves pinnate; secondary veins are eucamptodromous, between 7 to

14 pairs of 2's were counted on the specimens, generally a few pair of 2's are crowded at the

base and they are decurrent on primary veins and more acute than apical secondaries; one pair

acute basal secondaries is common within the morphotype; intersecondaries are strong and

common; tertiary veins are mixed alternate/opposite percurrent, the opposite percurrents having

courses straight to sinuous and their angles are mostly perpendicular on primary; tertiary vein

angle variability is perpendicular for both on midvein and ending; 4th order veins are alternate









percurrent; 5th order veins are regular polygonal reticulate; most specimens covered with

laminar resin glands (<0. mm); leaf rank 3r.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0315; 0317, 0318, 0319, and 0323; Pits Tabaco 1,

Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 72

specimens.

Systematic affinity suggested. The fossil leaf closely resembles the typical characters

seen in Lauraceae, pinnate leaves, eucamptodromous venation and resin glands.

CJ24 (Plate 25)

Diagnosis. Long petiole. One pair of strictly agrophic lateral primaries, sometimes it may

be difficult to distinguish from costal secondaries. Brochidodromous looping arches close to

margin. Teeth are low closely spaced crenations mostly fed by agrophic veins.

Description. Maximum length of the petiole measured reached at about 9 cm; laminar size

from notophyll to macrophyll; laminar shape elliptic; base shape oscillates from truncate to

cordate; apex shape acuminate; primary veins basal actinodromous; secondary veins are

brochidodromous, 12-14 pairs were counted on the specimens; secondary vein spacing increases

toward base and their angles remain uniform; agrophic veins are simple; intersecondary veins are

weak; tertiary veins are opposite percurrent with courses straight to sinuous; tertiary vein angle

variability decreases exmedially; 4th order veins are alternate percurrent; margin toothed, teeth

are closely spaced with crenations mostly fed by agrophic veins; the spacing is irregular and the

sinus of the teeth are rounded.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0317, 0318, and 0323; Pits Tabaco 1, and Tabaco









High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 30

specimens.

Systematic affinity suggested. The fossil leaf closely resembles the typical characters

seen in many species of Euphorbiaceae, leaves with primary veins basal actinodromous, long

petioles, and margin toothed.

CJ25 (Plate 26)

Diagnosis. Mucronate or rounded apex, base shape highly variable (concavo-convex,

cordate, truncate), sometimes almost are lobate, 3rd and 4th order veins are mostly opposite

percurrent. Teeth distantly spaced, and formed by simple extension of the vein beyond the

margin of the lamina, teeth sometimes retuse.

Description. Petiole has a base swollen, and it is long and thick; laminar size from

microphyll to macrophyll; laminar shape varies from elliptic to ovate; base shape highly variable

(concavo-convex, cordate, truncate); apex shape highly variable acuminate, mucronate and

rounded; primary veins are basal actinodromous, between 5 to 7 veins from the base; agrophic

veins compound; secondary veins are craspedodromous; and the costal secondaries have a vein

spacing and angle uniform; tertiary veins mainly opposite percurrent, their courses vary from

straight to sinuous, angles of tertiary veins on primaries are perpendicular to obtuse; tertiary vein

angle variability increases exmedially and basally; 4th order veins mostly opposite percurrent,

5th order veins are regular polygonal reticulate; a fimbrial vein runs parallel to margin.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315; 0317, 0318, 0319, 0323 and 0324; Pits Tabaco

1, Tabaco High Dip, and la Puente; localities placed between coal beds 100 and 161, middle to

late Paleocene, 67 specimens.









Systematic affinity suggested. The fossil leaf closely resembles the typical characters

seen in the group of Malvales. Leaves actinodromous, with long petioles, and margin toothed.

CJ26 (Plate 27)

Diagnosis. Almost lobate, primaries branching near margin. Fourth and fifth order veins

orthogonal reticulate. Teeth closely spaced and convex/convex and almost symmetrical.

Description. Petiole long and having a swollen base; laminar size from notophyll to

macrophyll; laminar shape constantly ovate; base shape cordate; apex shape convex; primary

veins are basal actinodromous, between 8 to 9 veins depart from the base; agrophic veins

compound; secondary veins are craspedodromous and their angles are irregular; tertiary veins

mainly opposite percurrent with straight to sinuous courses, their angles on primaries vary from

perpendicular to obtuse; tertiary vein angle variability increases exmedially and basally;

distinctly 4th and 5th order veins are orthogonal reticulate; fimbrial vein parallel to margin;

margin type crenate; the teeth are closely spaced and convex/convex and almost symmetrical, 6

to 7 teeth per cm, teeth sinuous rounded with simple apex.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, six specimens.

Systematic affinity suggested. The fossil leaf closely resembles the typical characters

seen in the group of Malvales. Leaves actinodromous, with long petioles, and margin toothed.

CJ27 (Plate 28)

Diagnosis. Naked basal lateral pimaries, 7-8 pairs of semicraspedodromous secondaries,

teeth closely spaced and convex/convex with elongate basal sides.









Description. Laminar size oscillates from notophyll to macrophyll; laminar shape ovate;

base shape cordate; apex shape acuminate; primary veins are basal actinodromous with 5 basal

veins; agrophic veins are compound; secondary veins are semicraspedodromous (7-8 pairs of

2's); the costal secondary veins increases toward base; tertiary veins are opposite percurrent with

a constant straight course and an angle acute; tertiary vein angle variability increases exmedially

and basally; 4th vein order are orthogonal reticulate; 5th order vein regular polygonal reticulate,

areolation is well developed, margin toothed, teeth closely spaced and convex/convex with

elongate basal sides.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0317 and 0319 Pit Tabaco 1 and Tabaco High Dip;

locality placed between coal beds 100 and 115, middle to late Paleocene, six specimens.

Systematic affinity suggested. The fossil leaf closely resembles the typical characters

seen in the group of Malvales. Leaves actinodromous, with long petioles, and margin toothed.

CJ28 (Plate 29)

Diagnosis. Pinnate with cordate base. Secondary vein angle at the base is more acute than

in CJ16. Tertiary veins are opposite percurent and are less closely spaced than in CJ16. Fourth

order veins are mostly mixed opp/alter and very thin.

Description. Laminar size mesophyll; laminar shape elliptic; base shape varies from

rounded to slightly cordate; margin entire; leaves pinnate with a very thick midvein; secondary

veins are eucamptodromous and five of these veins depart from the base; secondary vein spacing

decreases toward base and their angles smoothly increases toward base; tertiary veins are

opposite percurrent with straight courses and their angles on primary mainly perpendicular;

tertiary vein angle variability are perpendicular for both veins on the midvein and margin; 4th









vein category mostly opposite/alternate percurrent, veins very thin; 5th order type from regular

polygonal reticulate to dichotomizing.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, seven specimens.

Systematic affinity suggested. No extant affinity identified.

CJ30 (Plate 30)

Diagnosis. Asymmetrical blade. Secondary vein spacing and angles are irregular; however

the angle abruptly increases toward base. Tertiaries opposite percurrent, very sinuous and widely

spaced, epimedial tertiaries form intersecondaries.

Description. Laminar size mesophyll; blade asymmetrical; laminar shape obovate; base is

incomplete; margin entire; leaf pinnate; secondary veins are eucamptodromous; secondary vein

spacing irregular; secondary vein angle abruptly increases toward base; intersecondary veins are

strong and common; tertiary veins are opposite percurrent with very sinuous course and widely

spaced, their angles on primary are mostly perpendicular; tertiary vein angle variability increases

exmedially; 4th order vein are mainly opposite percurrent; 5th order veins are irregular

polygonal reticulate, areolation moderately developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen.

Systematic affinity suggested. No extant affinity identified.

CJ32 (Plate 31)









Diagnosis. Shape wider than CJ4. Semicraspedodromous 20s decurrent on primary.

Tertiaries opposite percurrent but do not form intersecondaries, tertiaries more closely spaced

and obtuse than CJ4. Fourth order veins mostly opposite percurrent. Fimbrial vein. Teeth similar

to CJ4.

Description. Long petioles, reaches up to 10 cm; laminar size from microphyll to

macrophyll; laminar shape elliptic; base shape strongly decurrent; apex shape acuminate; laves

pinnate; secondary veins are semicraspedodromous, at least 15 pairs of 2's; secondary vein

spacing uniform and secondary vein angle smoothly increases toward base; tertiary veins are

opposite percurrent with courses straight to sinuous, their angles on primary are mostly obtuse;

tertiary veins angle variability decreases exmedially; distinctly opposite percurrent; 5th order

veins are regular polygonal reticulate; areolation well developed; fimbrial veins parallel to

margin; margin toothed, teeth similar to CJ4, but spacing is irregular or closely spaced.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315, 0317, 0318, and 0319; Pits Tabaco 1, and

Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 34

specimens.

Systematic affinity suggested. Some species in Violaceae has the characters seen on the

morphotype.

CJ33 (Plate 32)

Diagnosis. Two pairs of long acrodromous secondaries in basal half of leaf, no apical

secondaries (hemieucamptodromous).









Description. Laminar size microphyll; laminar shape elliptic; apex shape rounded; margin

entire; leaf pinnate; secondary veins are suprabasal acrodromous with no apical secondaries

(hemieucamptodromous); secondary vein angle is low relative to primary.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen.

Systematic affinity suggested. No extant affinity identified.

CJ34 (Plate 33)

Diagnosis. Craspedodromous secondaries at almost 90 degrees to primary and spacing

decreasing toward base. Perpendicular and thin tertiaries at the midvein. Teeth have long basal

sides.

Description. Laminar size from notophyll to mesophyll; laminar shape from elliptic to

ovate; base shape concavo-convex to truncate; apex shape convex; leaves pinnate; secondary

veins are craspedodromous, with no or two basal veins; secondary vein spacing decreases toward

base; secondary vein angle smoothly increases toward base; intersecondaries are weak; tertiary

veins mainly opposite percurrent with sinuous to straight courses, their angles on primary are

perpendicular; 4th order vein are alternate percurrent; 5th order vein are regular polygonal

reticulate, areolation well developed; margin toothed, teeth have long basal sides, 1-2 teeth per

cm and spacing regular.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal

beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, two specimens.









Systematic affinity suggested. Leaves pinnate, craspedodromous secondaries and

tertiaries perpendicular and thin at the midvein are very common in the family Anacardiaceae.

CJ36 (Plate 34)

Diagnosis. Actinodromous, three primary veins, lateral primaries forming loops. Weak

brochidodromous 20s that loop just inside the margin both, costal and minor secondaries. Fourth

order veins alternate percurrent and leaf texture different from CJ25.

Description. Long petiole; laminar size notophyll to mesophyll; laminar shape elliptic;

base shape varies from truncate to convex; margin entire; primary veins are basal actinodromous

with three veins at the base; agrophic veins are simple; secondary veins are weak

brochidodromous, the spacing of costal 2's increases toward base and their angles are acute;

tertiary veins are opposite percurrent having sinuous to convex courses, their angles are

perpendicular on primaries; tertiary vein angle variability uniform; 4th order veins are alternate

percurrent; 5th order veins are regular polygonal reticulate; areolation well developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315, 0318, and 0319; Pits Tabaco 1, and Tabaco

High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, nine

specimens.

Systematic affinity suggested. The fossil leaf closely resembles the typical characters

seen in the group of Malvales, leaves actinodromous with long petioles.

CJ37 (Plate 35)

Diagnosis. Few secondary veins (5 or 6 pairs), secondary spacing decreasing toward base

but not crowded basally like CJ22. Secondary angles more acute than CJ20 and CJ22. Lacking

laminar resin dots.









Description. Petiole short; laminar size mesophyll; laminar shape elliptic; base shape

decurrent; margin entire; leaf pinnate; secondary veins are eucamptodromous, their spacing

decreases toward base; secondary vein angle uniform; tertiary veins are mixed opposite/alternate

percurrent, their angles on primary are perpendicular; tertiary vein angle variability mostly

uniform; 4th order vein alternate percurrent; 5th order vein are regular polygonal reticulate;

areolation well developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal

beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, one specimen.

Systematic affinity suggested. No extant affinity identified.

CJ38 (Plate 36)

Diagnosis. Pulvinate and striated petiole. Secondaries are perpendicular basally on

midvein, intersecondary veins are common. Tertiaries alternate percurrent and wider spaced than

CJ8. Tertiary angle increases basally on midvein, tertiaries decurrent on midvein.

Description. Petiole pulvinate and striated; laminar size from notophyll to mesophyll;

laminar shape elliptic; base shape rounded; margin entire; leaves pinnate; secondary veins

eucamptodromous, sometimes having three basal veins; secondary vein spacing decreases

toward base, and their angles smoothly increases toward base; intersecondary veins are weak to

strong; tertiary veins are alternate percurrent with their angles perpendicular to base; tertiary

veins angle variability increases basally on midvein; 4th order vein regular polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0317 and 0318; Pits Tabaco 1 and Tabaco High Dip,

localities placed between coal beds 100 and 161, middle to late Paleocene, 20 specimens.









Systematic affinity suggested. Based on the leaf morphology and the presence of a petiole

transversely striated, short and pulvinate the morphotype closely resembles the family

Leguminosae.

CJ40 (Plate 37)

Diagnosis. Pinnate with cordate to rounded base. Costals 20s veins appear near base, one

pair of agrophic lateral secondaries. Higher order veins are regular polygonal reticulate with

areoles of rounded shape and tiny size.

Description. Long petiole (-6 cm); laminar size from microphyll to macrophyll; laminar

shape is elliptic to ovate; base shape cordate to rounded; apex shape acuminate; margin entire;

leaves pinnate; agrophic veins compound ;secondary veins are brochidodromous, 3 to 5 veins

depart at the base; costal secondary vein spacing decreases toward base; tertiary veins are

opposite percurrent with sinuous to convex courses, their angles on primary are perpendicular to

obtuse; tertiary vein angle variability increases basally; 4th and 5th vein category regular

polygonal reticulate; areolation well developed, tiny and rounded.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315; 0317, 0318; 0319 and 0323; Pits Tabaco 1 and

Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 37

specimens.

Systematic affinity suggested. No extant affinity identified.

CJ41 (Plate 38)

Diagnosis. Six or seven pairs of costals 20s, crowded basal secondaries with relative long

space between basal 2-3 and more apical secondaries, secondaries curved up smoothly.









Secondaries and tertiaries are decurrent on midvein, opposite percurrent tertiaries perpendicular

to primary and the angle increases basally.

Description. Long petiole (-6 cm); laminar size from microphyll to macrophyll; laminar

size from elliptic to ovate; base shape either cordate or rounded; apex shape acuminate; margin

entire; leaves pinnate; agrophic veins compound; secondary veins are brochidodromous with 3 to

5 veins at the base; costal secondary vein spacing decreases toward base; tertiary veins distinctly

opposite percurrent, their courses vary from sinuous to convex; the angles of 3's on primary are

perpendicular to obtuse; tertiary vein angle variability increases basally; 4th and 5th regular

polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0318 and 0324; Pit Tabaco High Dip; localities placed

between coal beds 125 and 161, middle to late Paleocene, seven specimens.

Systematic affinity suggested. No extant affinity identified.

CJ43 (Plate 39)

Diagnosis. Asymmetrical blade. Craspedodromous 20s with agrophic branches on mid-leaf

secondaries, 2s become more obtuse basally. Teeth broad convex/convex and almost

symmetrical, apical teeth are simple or foraminate. Fimbrial vein.

Description. Laminar size from mesophyll to macrophyll; laminar shape elliptic to ovate;

laminar symmetry strongly asymmetrical; base shape concave; apex shape concave; leaves

pinnate; agrophic veins compound; secondary veins are craspedodromous; secondary veins

spacing increases toward base and their angles smoothly increases toward base as well; tertiary

veins mostly opposite percurrent with sinuous and straight courses, their angles on primary are

only perpendicular; tertiary vein angle variability increases exmedially; 4th order veins are









mixed opposite/alternate percurrent; 5th order veins are regular polygonal reticulate; areolation

moderately developed; fimbrial vein parallel to margin; margin toothed, two orders of teeth with

irregular spacing, teeth are convex-convex and concave-convex, teeth apex are foraminate or

simple.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0319, 0322; 0323, and 0324; Pits Tabaco High Dip and

La Puente ; localities placed between coal beds 103 and 130, middle to late Paleocene, 62

specimens.

Systematic affinity suggested. No extant affinity identified.

CJ48 (Plate 40)

Diagnosis. Actinodromous with closely spaced nipple-shaped teeth with darkened apices

Description. Laminar size mesophyll; laminar shape ovate; base angle obtuse and base

shape probably cordate; primary veins are basal actinodromous; agrophic veins are simple;

tertiary veins are opposite percurrent with mainly sinuous courses; teeth are closely spaced

nipple-shaped with darkened apices.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0315 and 0319; Pits Tabaco 1 and Tabaco High Dip,

localities placed between coal beds 100 and 107, middle to late Paleocene, three specimens.

Systematic affinity suggested. The leaf and teeth morphology may resemble species in

the order Malvales.

CJ50 (Plate 41)

Diagnosis. Pinnate and cordate base. Percurrent tertiaries, mostly convex and widely

spaced with straight course on the midvein and sinuous on secondaries. Fourth order veins are









alternate percurrent and thick veins which is different from CJ28. Areolation is not squarish as in

CJ17.

Description. Laminar size and shape from mesophyll and elliptic respectively; base shape

is closely cordate to cordate; margin entire; leaves pinnate, sometimes having three basal veins;

agrophic veins are compound; secondary veins are eucamptodromous; secondary vein spacing

decreases toward base and their angles smoothly increases toward base; tertiary veins mainly

opposite percurrent with courses straight to sinuous/convex; the angle of 3's on primary is

mainly perpendicular; tertiary vein angle variability increases exmedially; 4th vein type are

alternate percurrent; 5th order veins are regular polygonal reticulate; areolation is well

developed.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, localities 0315 and 0322; Pits Tabaco 1 and La Puente, localities

placed between coal beds 100 and 105, middle to late Paleocene, three specimens.

Systematic affinity suggested. No extant affinity identified.

CJ53 (Plate 42)

Diagnosis. Actinodromous. Distinct higher order veins with moderately and well

developed areoles, 1-branched F.E.V.s. covered with small laminar resin dots.

Description. Laminar size mesophyll, laminar shape ovate; base shape cordate; margin

entire; primary veins are basal actinodromous, at least three veins are observed; agrophic veins

are compound; tertiary veins are mainly opposite percurrent and their courses are sinuous; the

angles of the 3's on primaries is perpendicular; 4th vein types are mixed opposite/alternate

percurrent; 5th order veins are regular polygonal reticulate; areolation is moderately to well

developed; 1-branched F.E.V.s. covered with small laminar resin dots.









Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal

beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, three specimens.

Systematic affinity suggested. No extant affinity identified.

CJ54 (Plate 43)

Diagnosis. Decurrent base. Acrodromous, three or five primary veins. Secondaries are

mostly opposite percurrent and perpendicular on midvein, marginal secondaries are weak

brochidodromous.

Description. Petiole has a swollen base; laminar size mesophyll; laminar shape distinctly

oblong; base shape decurrent; apex shape rounded; margin entire; primary veins are

parallelodromous, 3 to 5 veins; secondary veins are opposite percurrent and their angles are

perpendicular on primaries, but marginal secondaries are weak brochidodromous; tertiary veins

are regular polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0319 and 0410, Pits Tabaco High Dip and Expanded

West, localities placed between coal beds 40 and 107, middle to late Paleocene, six specimens.

Systematic affinity suggested. No extant affinity identified.

CJ55 (Plate 44)

Diagnosis. Leaflets. Short and pulvinate petiole. Asymmetrical base. Four to six crowded

basal veins, on either side of the midvein, 2 or three secondary veins running with acute angle.

Description. Petiole pulvinate and short; laminar size of the leaflets from microphyll to

mesophyll; laminar shape elliptic; laminar symmetry strongly base asymmetrical; base shape

complex; apex shape acuminate; margin entire; pinnate leaflets; secondary veins are









brochidodromous, four to six of these veins are crowded at the base; secondary vein spacing

decreases toward base and their angles abruptly decrease toward base as well; intersecondaries

are weak; tertiary veins are random reticulate and their angle variability is inconsistent; 4th order

types are regular polygonal reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, localities 0317 and 0410, Pits Tabaco 1 and Expanded West,

localities placed between coal beds 40 and 102, middle to late Paleocene, 21 specimens.

Systematic affinity suggested. Based on the leaf morphology and the presence of a short

and pulvinate petiole the morphotype closely resembles the family Leguminosae.

CJ56 (Plate 45)

Diagnosis. Pinnate with cordate base. Thick primary vein. Four basal veins, compound

agrophic veins. Craspedodromous 20s with straight course and branching at the margin. Teeth

concave/concave. One specimen covered with peltate scales or laminar resin dots.

Description. Laminar size from mesophyll to megaphyll; laminar shape elliptic; base

shape from cordate to convex; leaves pinnate; agrophic veins are compound; secondary veins are

craspedodromous; tertiary veins constantly opposite percurrent with strong sinuous courses, the

angles of 3's on primary are perpendicular; tertiary vein angle variability increases exmedially;

4th order veins are mostly opposite percurrent; teeth concave/concave. One specimen covered

with peltate scales or laminar resin dots.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0324, Pit Tabaco High Dip; locality placed between coal

beds 125 and 130, middle to late Paleocene, coordinates 11.62N, 73.32 W, 11 specimens.









Systematic affinity suggested. Some species in Flacourtiaceae have similar leaf characters

as those seen in the fossil.

CJ58 (Plate 46)

Diagnosis. Thick and slightly curved midvein. Secondaries and intersecondaries strongly

decurrent on midvein. Tertiaries very thin and mostly random reticulate on midvein and mostly

opposite percurrent exmedially. Tertiaries are decurrent on primary and secondaries. 1-2

branched F.E.V.s.

Description. Laminar size mesophyll; laminar shape elliptic; base angle shape acute and

probably decurrent; leaf pinnate; intersecondary veins are thin and weak; tertiary veins are

opposite percurrent and their angles on primary are perpendicular; tertiaries are decurrent on

primary and secondaries; tertiary vein angle variability increases exmedially; 4th and 5th order

types are distinctly irregular polygonal reticulate; areolation is poorly developed; 1-2 branched

F.E.V.s.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen.

Systematic affinity suggested. No extant affinity identified.

CJ59 (Plate 47)

Diagnosis. Primaries are flabellate. Higher order veins mostly random reticulate.

Description. Laminar size notophyll; laminar shape elliptic; base angle obtuse and

probably base shape rounded; margin entire; primary veins are flabellate; tertiary veins are

random reticulate; coriaceous texture.









Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, three specimens.

Systematic affinity suggested. No extant affinity identified.

Monocots

CJ3 (Plate 48)

Diagnosis. Leaf apex rounded, margin entire, cordate or sagittate base, each basal lobe fed

by a secondary vein that branches several times, naked basal veins, midrib and secondary veins

stout and multistranded, secondary veins crowded basally and attached to the midrib at very

obtuse angles basally (-90-100) and decreasing apically to -450, secondaries merging at several

intramarginal veins in a complex brochidodromous pattern, tertiary veins attached to the midrib

and secondaries at angles between 200 and 450 to form a network of intersecondary venation,

higher order veins are strongly irregular polygonal reticulate that have wandering courses and

connections at variable angles.

Description. Maximum length measured is -56 cm, and maximum width measured at the

posterior division -26 cm, margin entire; leaf apex rounded; midrib multistranded and up to 1.5

cm wide; leaf base cordate or sagittate, the basal lobes are fed by the most basal secondary veins,

these secondary veins are very well-developed and form part of the leaf margin at the base as

naked basal veins, the two basal secondary veins also dichotomize within the lobes several times

to form minor secondaries that have perpendicular courses with respect to the midrib and very

obtuse angles basally; secondary veins are multistranded, very thick and decurrent on the midrib,

secondary veins crowded basally and four veins depart from the petiole insertion, secondary

veins branch in a complex brochidodromous pattern, never reaching the margin; the course and

spacing in between the intramarginal veins are very irregular; higher order veins are irregular









polygonal reticulate with wandering courses (sinuous, angular, or straight) and connections at

variable angles.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0322, Pit La Puente, locality placed between coal beds

103 and 105, middle to late Paleocene, six specimens.

Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves are

extremely similar to those found in the family Araceae and the extant genus Montrichardia.

CJ23 (Plate 49)

Diagnosis. Leaves ovate, margin entire, apex acuminate, base cordate or sagittate, naked

basal veins, midrib multistranded, secondary veins eucamptodromous, one or two intramarginal

veins running close to the margin, one or two intersecondary veins running parallel in between

secondaries, tertiary veins mixed opposite/ alternate percurrent and reticulate.

Description. Laminar shape ovate, length 6.2 cm, width 3.8 cm; margin entire; posterior

division of the blade not completely preserved but the presence of laminar tissue proximal to the

petiole attachment shows the base is cordate or sagittate; naked basal veins form part of the leaf

margin near the petiole attachment; anterior division of the blade longer than posterior division;

midrib multistranded; the most basal secondary vein curves in a proximal direction and feeds the

basal lobe. The next most distal secondary is oriented almost perpendicular to the midrib for

much of its course, forms the primary intramarginal vein, and displaces toward the margin a

minor intramarginal vein that arises from the posterior division of the blade. The third secondary

vein forms the secondary intramarginal vein, and the rest of the secondary veins are strongly

eucamptodromous and ascend from the midrib at angles between 290 to 350; one or two

intersecondary veins are present between each pair of secondaries and have courses almost









parallel to the secondaries. Both secondary and intersecondary veins are strongly decurrent on

the midrib and follow a course that curves toward the margin distally. The tertiary veins are

mixed opposite/alternate percurrent, but exmedially most tertiaries are opposite percurrent and

are oriented perpendicularly to the midrib, except for those developed in the basal lobe which

tend to be parallel to the midrib. Higher order veins are irregular polygonal to orthogonal

reticulate.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, two specimens.

Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves are

extremely similar to those found in the family Araceae and the extant genus Anthurium.

CJ47 (Plate 50)

Diagnosis. Multistranded midrib. Submarginal vein. Numerous parallelodromous

secondaries, closely spaced and decurrent on midvein. Thick and darkened fimbrial vein.

Specimen covered with small resin dots.

Description. Fossil fragment; laminar size mesophyll; the fossil is missing the base;

margin entire; primary vein multistranded and it may be pinnate, numerous parallelodromous

secondaries, closely spaced and decurrent on midvein; thick and darkened intramarginal vein that

collects the secondary veins; thick fimbrial vein runs parallel to margin; the specimen is covered

with small resin dots.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds

100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen.









Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves

seem similar to those found in the family Araceae, however a more detailed comparison with

other families is necessary.

CJ49 (Plate 51)

Diagnosis. Blade decurrent on petiole. Numerous parallelodromous secondaries, closely

spaced with perpendiculary cross veins.

Description. Laminar size from mesophyll to macrophyll; margin entire, primary vein

pinnate; secondary veins are numerous and parallelodromous, closely spaced with

perpendiculary cross veins.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal-mine, locality 0322; Pit La Puente, locality placed between coal beds

103 and 105, middle to late Paleocene, seven specimens.

Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves

seem similar to those found in the order Zingeberales and the family Araceae, however a more

detailed comparison with other families is necessary.

CJ62 (Plate 52)

Diagnosis. Primary and/or secondaries decurrent and multistranded. Tertiaries are mostly

opposite percurrent and strongly perpendicular to primary and/or to secondaries. Higher order

veins are regular polygonal reticulate. Specimen covered with resin dots.

Description. Laminar size mesophyll; margin entire; primary and/or secondaries decurrent

and multistranded; tertiaries are mostly opposite percurrent and strongly perpendicular to

primary and/or to secondaries; higher order veins are regular polygonal reticulate; the specimen

is covered with resin dots.









Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0324; High Dip, locality placed between coal beds 125

and 130, middle to late Paleocene, one specimen.

Systematic affinity suggested. No extant affinity identified.

CJ63 (Plate 53)

Diagnosis. Stout midrib. Secondaries multistranded and decurrent on midvein. Tertiaries

very thin, decurrents, irregular paths and with acute angle on secondaries.

Description. Laminar size megaphyll; the fossil misses the base and the margin; midrib

stout and multistranded; secondary veins are multistranded and decurrent on midvein; tertiaries

very are thin, decurrent, and have irregular paths with very acute angle on secondaries.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0322; Pit La Puente, locality placed between coal beds

103 and 105, middle to late Paleocene, one specimen.

Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves

seem similar to those found in the order the family Araceae, however a more complete specimen

is required and a more detailed comparison with other families is necessary as well.

CJ64 (Plate 54)

Diagnosis. Pinnate, multistranded midrib. Secondaries strongly decurrent, secondaries

widely spaced and without fimbrial vein like CJ47.

Description. Laminar size mesophyll; leaf pinnate and multistranded; secondary veins are

strongly decurrent on midrib, and they widely spaced, the specimen misses a fimbrial vein like

seen in CJ47.









Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine; Pit La Puente, locality placed between coal beds 103 and 105,

middle to late Paleocene, one specimen.

Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves

seem similar to those found in the order Zingeberales and the family Araceae, however a more

detailed comparison with other families is necessary.

CJ65 (Plate 55)

Diagnosis. Midrib and secondaries are multistranded. Secondaries and tertiaries are

strongly decurrent on midvein and have parallel paths. Fourth order veins with acute angle on

tertiaries and irregular course.

Description. Laminar size megaphyll; margin entire; leaf pinnate; the midrib and

secondary veins are multistranded; secondaries and tertiaries are strongly decurrent on midvein

and have parallel paths; 4th order veins with acute angle on tertiaries and irregular course.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal-mine, locality 0323; Pit High Dip, locality placed between coal beds

115 and 120, middle to late Paleocene, two specimens.

Systematic affinity suggested. The leaf form and venation patterns of the fossil leaves

resemble to those found in the order Zingeberales.

CJ67 (Plate 56)

Diagnosis. Fragments of costapalmate or palmate leaves. Segments with entire margins, a

midvein, and four orders of parallel venation connected by irregular transverse commisures.

Description. Rachis 23 mm wide in the base and 7 17 mm wide in more apical

fragments, with a longitudinal ridge; leaflets are single fold, with entire margins, regularly









arranged in one plane, opposite to semialternate, separated by 8 42 mm at their insertion;

midrib prominent, 0.5 0.7 mm wide, two orders of parallel veins separated by 1 2.3 mm,

transverse veinlets not evident; the leaflets are joined together in fragments with the thinnest

rachis.

Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrejon coal mine, locality 0319; Pit Tabaco High Dip; locality placed between coal

beds 105 and 107, middle to late Paleocene, three specimens.

Systematic affinity suggested. Arecaceae (palm family).

CJ68 (Plate 57)

Diagnosis. Fragments of pinnate leaves. Leaflets single fold, with entire margins, regularly

arranged in one plane, opposite to semialternate, midrib prominent, two orders of parallel veins,

transverse veinlets not evident.

Description. Fragments of costapalmate or palmate leaves; segments 51.6 112 mm wide,

with entire margins, and a midvein 1.5 mm wide; parallel veins of four orders: first order veins

1.17 1.6 mm wide, separated by 38 38.29 mm; second order veins 0.8 1.1 mm wide; primary

and secondary veins separated by 0.2 mm approximately; third order veins 0.1 0.2 mm wide,

separated by 2.0 3.44 mm; tertiary veins or tertiary and secondary veins separated by 6 8

mm; fourth order veins very thin, almost indiscernible, separated by 0.2 mm and approximately

seven in number between two third order veins; transversal veins 0.10 0.12 mm wide, with an

irregular course, inserted to the middle or primary veins with an acute angle, and separated by 1 -

5 mm; after their insertion, they bend to cross the secondary veins with an almost straight angle;

they can connect two primaries or a primary and a lesser order vein; transversal venation is not

always visible, in some leaf fragments is only visible where a partial decomposition took place.









Source, age, number of specimens, and stratum. Colombia, Rancheria Basin, Cerrej6n

Formation, Cerrej6n coal mine, localities 0317, 0318, and 0323; Pits Tabaco 1 and Tabaco High

Dip; localities placed between coal beds 100 and 107, middle to late Paleocene, 61 specimens.

Systematic affinity suggested. Arecaceae (palm family).

Paleoclimate

I used modern correlations between dicot leaf margin type and temperature (Wilf, 1997)

and leaf size and precipitation (Wilf et al., 1998) to estimate the climatic conditions under which

the Cerrej6n flora grew. Seventy-four percent of the 41 dicot leaf types have entire margins,

yielding a mean annual temperature (MAT) estimate of 23.8 +2.3 C. It is probable that riparian

and wetland floras like those represented by the fossils typically have a higher proportion of

toothed species than adjacent terra fire vegetation and thus can underestimate MAT by up to 5-

7 C (Burnham et al., 2001; Kowalski and Dilcher, 2003).

More than 50% of the dicot leaves in the Cerrej6n flora are mesophyll or larger in size

(>1900 mm2), which is striking given the preservational biases against large leaves (Roth and

Dilcher, 1978). I infer a mean annual precipitation (MAP) of 3240 mm (range 2260-4640 mm),

which is likely an underestimate (Burnham et al., 2005). The abundant, thick beds of low-ash,

low-sulfur coal also indicate a wet climate.

The Cerrej6n fossil leaf physiognomy and the reconstructed paleoclimate prove the

existence of a 58-60 Ma Neotropical Rainforest. Modern rainforests are recognized for having

assemblages with high proportions of large, entire-margined, thin-textured leaves. Modern

rainforests have a MAT that oscillates between 22 and 27 C, and MAP usually is higher than

2200 mm, with a dry season no longer than 2 months (Dolph and Dilcher, 1980; Burnham and

Johnson, 2004).









The Cerrej6n MAT estimate is similar to modern tropical temperatures, and does not

display very high MAT as do some high p-CO2 climate simulations and oxygen isotope estimates

of Paleocene tropical sea surface temperature (Pearson et al., 2001; Huber et al., 2003; Shellito et

al., 2003). The paleoclimatic simulations suggest a low-latitude MAT higher than 28 C with 500

ppm of CO2, and a maximum of 34 C with 2000 ppm of atmospheric CO2 (Shellito et al., 2003).

If leaf margin analysis underestimates MAT at Cerrej6n by 7 C, the equator-to-pole temperature

contrast for the middle to late Paleocene would have been only -20 C, requiring climate

mechanisms that retained heat at high latitudes more effectively than in the present day (Sloan

and Pollard., 1998; Huber et al., 2003; Sewall et al., 2004).

The Cerrej6n MAP estimate is congruent with the predictions by Shellito et al (2003), but

the small scarce differences between the predictions of paleoprecipitation under different levels

of CO2 (from 500 to 2000 ppm) make inferences about the possible CO2 levels reached during

the early Paleogene based solely on precipitation estimates inconclusive.

Floristic Composition of the Cerrej6n Flora

The floristic composition of the Cerrej6n flora compared informally with the most

important tropical families (Burnham and Johnson, 2004) shows that this middle-late Paleocene

flora had a family-level taxonomic composition typical of extant Neotropical rainforests. The

most diverse and abundant families in the Cerrej6n florawere: Fabaceae (4 leaf morphotypes),

Malvales (3), Araceae (3), Arecaceae (2), Lauraceae (2), Zingiberales (2), Anacardiaceae (1),

Cycadaceae (1), and Menispermaceae (1). Also, the four ferns found include: the pantropical,

floating-aquatic Salvinia; Stenochlaena, a common climbing fern in Old World tropical swamps;

the cosmopolitan Lygodium; and Acrostichum, a pantropical fern common in freshwater and

mangrove swamps.









As well as fossil leaves, fruits, seeds and flowers were found in the Cerrej6n Formation;

the most abundant fossil fruits belong to Leguminosae (4 types of pods) and Arecaceae (3

morphotypes) which are related to moderns coconuts and the mangrove palm Nypa. In addition,

the pollen record obtained from the Cerrej6n Formation shows components of modem rainforests

(Jaramillo et al., 2007). Some of the most abundant pollen grains are proxapertites operculatus

(Araceae), Mauritiidites franciscoi var. pachyexinatus (Arecaceae), Psilatriletes types (ferns),

and Momipites africanus (Moraceae).

When the familial/ordinal taxonomic composition of the Cerrej6n megaflora was compared

statistically with that of extant lowland Neotropical forests, the results showed a remarkable

degree of similarity (Wing et al., in press). The comparison was done with 73 of Gentry's 0.1

hectare transects (Phillips et al., 2002). The modern sites were tabulated for the proportions of

species and individuals belonging to each family/order. The fossils were tabulated for the

proportion of leaf morphotypes in each family/order for the whole flora and the number of leaves

in each family/order for a random sample collected at one site. The results obtained by Wing et

al. (in press) showed that the mean Spearman coefficient between the Cerrej6n and all extant

sites varied from 0.41 to 0.95. Thus, in terms of the rank-order diversity and abundance of plant

families/orders, the Cerrej6n flora is not distinguishable from living Neotropical lowland forests.

Conclusions

I recovered 1347 megafossil specimens and segregated them into 57 morphotypes using

leaf architectural characters. The fossils include foliar organs of 41 dicots, 10 monocots, 4 ferns,

1 cycad, and 1 conifer. In addition, numerous fruits, seeds and flowers have been found from the

Cerrej6n Formation, but they were not described in this monograph.

Cerrej6n plant fossils document the existence 58-60 million years ago of Neotropical

Rainforest with modern family composition. The most important Neotropical families and orders









were present in Cerrej6n such as Fabaceae (the richest family in number of species in the

Neotropics), Malvales, Lauraceae, Menispermaceae, Arecaceae, Araceae, Cycadaceae, ferns, and

Zingiberales.

The Cerrej6n assemblage has a high proportion of large, entire-margined, thin-textured,

leaves, indicating a tropical climate with mean annual temperature > 24 C and rainfall > 2.3

m/y. This paleoclimate and fossil leaf physiognomy is consistent with modern climatic

conditions of tropical rainforests. Also, the abundant, thick beds of low-ash, low-sulfur coal from

the Cerrej6n Formation indicate a wet climate.

The estimated Cerrej6n MAT is consistent with modern tropical temperatures; this result is

much cooler than paleoclimatic simulations for the Paleocene. This paleotemperature inference

should be considered tentatively. There are two factors that may impact the result. First, the

riparian and wetland effect may underestimate MAT by up to 5-7 C in tropical floras. Secondly,

the empirical limitation in the equations available for reconstructing MAT; nowadays there is not

a single forest with a MAT higher than 26-270C, so based on this limitation, it is impossible to

get temperatures as they have been predicted by paleoclimatic simulations during the early

Paleogene. It is necessary to obtain independent proxies of MAT from the same formation. The

reconstructed MAP points to very humid conditions during the low-latitude middle-late

Paleocene.









CHAPTER 2
FOSSIL ARACEAE FROM A PALEOCENE NEOTROPICAL RAINFOREST IN COLOMBIA

Abstract

The family Araceae is very diverse in tropical America. Both the fossil record and

molecular data support a long evolutionary history. Araceous fossils, however, have been

recorded only from mid and high latitudes. Here, we report the oldest fossil leaves of Araceae

from the middle-late Paleocene Cerrej6n flora in northern South America. The three fossil leaf

taxa are related to two extant Neotropical genera, two of them closely related to Anthurium. The

other fossil taxons were placed within the extant genus Montrichardia. The fossil aroids

inhabited a coastal rainforest -60-58 million years ago and provide new information about

patterns of angiosperm diversification in the Neotropics.

Introduction

Araceae is one of the most diverse monocotyledonous families, comprising 9 subfamilies,

106 genera, and about 3300 species (Croat, 1979; Mayo et al., 1997; Govaerts and Frodin, 2002;

Keating, 2003a; 2004). It is most diverse in the tropics, with the Pacific slopes of the Northern

Andes home to the greatest number of species (Croat, 1992b; Vargas et al., 2004; Mora et al.,

2006). Growth forms include herbs, vines, hemiepiphytes, epiphytes, lithophytes, rheophytes,

and helophytes (Croat 1990; 1992a; Mayo et al., 1997; Bown, 2000). Despite high species

diversity, a large range of life forms, and a wide geographic distribution, the origin and evolution

of the Araceae are poorly known. Its fossil record is sparse and controversial, possibly because

the herbaceous life form of most Araceae makes them unlikely to fossilize (Herendeen and

Crane, 1995).

The oldest reported fossils of Araceae are inaperturate, striate pollen grains from the late

Barremian early Aptian (-124-117 Mya) of Portugal (Plate 58-1; Friis et al., 2004, 2006),









although this age has been questioned and remains problematic (Heimhofer et al., 2007). Those

pollen grains were assigned to the fossil genus Mayoa, and may be related to the subfamily

Pothoideae, tribe Monstereae according to the most recent classification of Araceae by Keating

(2003b; 2004). Fossils of Araceae have been reported from a number of Late Cretaceous

localities (Plate 58-2), including several from the Coniacian deposits of Eastern Asia (Krassilov,

1973), the Campanian of Central Europe (Kvacek and Herman, 2004), North America (Bogner et

al., 2005), and from the Maastrichtian of India and South America (Herngreen, 1974; Sarmiento,

1994; Bonde, 2000; Hesse and Zetter, 2007). The number of fossil reports increases during the

Paleocene and Eocene records in both mid and high latitudes (Plate 58-3). Mid latitudes had a

subtropical climate during the early Paleogene, explaining the prevalence of araceous taxa in

present day temperate regions (Dilcher and Daghlian, 1977; Smith and Stockey, 2003; Wilde et

al., 2005).

The minimum age estimate for the origin of the family, using molecular analysis calibrated

with fossils, is between 105 and 128 million years (Bremer, 2000; Janssen and Bremer, 2004).

The extant diversity and subfamilial structure of Araceae is complex and predominantly tropical

(Croat, 1979; Mayo, 1993), suggesting that low latitudes should have been the area of origin and

radiation. The fossil record of tropical aroids has remained nearly unexplored (Plate 58-1-2, and

3). In tropical South America, for example, the only araceous records are dispersed pollen from

the Maastrichtian and Paleocene stages of Spathiphyllum and Proxapertites operculatus types

(Van der Hammen & Garcia, 1966; Sarmiento, 1994; Jaramillo and Dilcher, 2001; Hesse &

Zetter 2007).

Here, I describe three new species of Araceae based upon fossil leaves preserved in

Paleocene sediments in northern South America. Two species are related to the extant genus









Anthurium, and the other is placed within the extant genus Montrichardia. The araceous fossil

leaves are part of the middle-late Paleocene Cerrej6n flora, which provides new data about the

evolution of Neotropical Rainforests (Wing et al., 2004; Herrera et al., 2005).

Materials and Methods

The fossil leaves described in this study were collected from the Cerrej6n Formation,

exposed in the Cerrej6n mine, a large open pit coal mine located in Northeastern Colombia

(1 10N, 720W). The Cerrej6n Formation has been dated by fossil pollen as middle to late

Paleocene, making it about 60-58 million years old (Jaramillo et al., 2007). The 700-m-thick

stratigraphic sequence, composed of thick coals, fluvially-deposited sandstones, and lacustrine

siltstones was deposited in a rapidly subsiding basin (Bayona et al., 2004). The seven specimens

of fossil leaves reported here were found -380 meters above the base of the formation. The fossil

material is stored in the paleontological collections of the Colombian Geological Institute

(INGEOMINAS) in Bogota.

I performed an exhaustive comparison of the fossils with collections of extant Araceae in

the United States National Herbarium, Washington DC, Smithsonian Tropical Research Institute,

Republic of Panama, and University of Florida Herbarium, Gainesville, Fl. We studied 90% (95

of the 106) of the extant genera, as well as all the tribes and subfamilies. Araceae leaf

morphology is extremely variable and diverse; it has not been used commonly as a taxonomic

tool. Nevertheless, recent morphological work by Keating (2003b) considers leaf venation a

potentially useful taxonomic characteristic. Here we follow Keating (2003b, 2004) in using

venation to assign the fossil leaves to genera within subfamilies of Araceae. We used leaf

terminology adapted from LAWG (1999), due to the confusion with the traditional terminology

in aroid descriptions (see Keating, 2003b for a discussion). Consequently, some terms such as

"primary lateral veins" (Croat and Bunting, 1979; Mayo et al., 1997) are called secondary veins,









and submarginal collective veins or collective veins (Croat and Bunting, 1979; Mayo et al.,

1997) are called intramarginal veins. The term midrib was used to describe the primary vein or

midvein; secondary veins are the next narrower class of veins after the midrib (and they originate

from it); and intramarginal veins closely paralleling the leaf margin (LAWG, 1999).

Although two species described in this paper are similar to an extant genus we used form

generic names rather than place them in an extant genus. This is because of the lack of floral and

many leaf structural characters that are used by botanists to identify this extant genus. Although,

the fossil material is good and contains many useful characters, it lacks the total number of

characters to place it closer than to subfamily and suggests a generic relationship at this time.

The standard up until about 35 years ago was to place all fossil angiosperms in extant genera

using a general statement that they "looked most similar" to a particular modem genus. After

Dilcher (1974) pointed out that when examined carefully, many fossil leaves did not belong to

the genera, or even families to which they had been assigned; angiosperm paleobotanists began

to examine and use leaf characters with more care. As a result, we have found that there are

many examples of angiosperm evolution throughout their history. The use of form genus, as

done here, reflects this caution and honesty in establishing relationships as a part of angiosperm

paleobotany (Roth and Dilcher, 1979; Jones and Dilcher 1980).

The leaves of fossil and extant Araceae were photographed for comparison using Nikon

D70 and DXM1200F cameras. Leaves ofMontrichardia arborescens Schott were cleared and

mounted following the techniques of Hickey (1973) and Dilcher (1974). Fossil leaf

reconstructions were made using Adobe Photoshop and Adobe Illustrator CS2.

Systematics

Family. Araceae.

Subfamily. Pothoideae Engler.









Tribe. Pothoeae Bartl.

Morphogenus. Anthurioidea Herrera, Jaramillo, Dilcher, Wing et Gomez-N gen. nov.

Generic diagnosis. Leaves ovate, margin entire, apex acuminate, base cordate or sagittate,

naked basal veins, midrib multistranded, secondary veins eucamptodromous, one or two

intramarginal veins of secondary gauge, one or two intersecondary veins per intercostal region

with courses parallel to those of the secondaries, tertiary veins mixed opposite/alternate

percurrent and reticulate.

Derivation of generic name. From the extant genus Anthurium.

Type species. Anthurioidea wayuuorum Herrera, Jaramillo, Dilcher, Wing et Gomez-N sp.

nov.

Second species. Anthurioidea cerrejonense Herrera, Jaramillo, Dilcher, Wing et Gomez-N

sp. nov.

Species diagnosis. Anthurioidea wayuuorum Herrera, Jaramillo, Dilcher, Wing et Gomez-

N. Laminar shape ovate, margin entire, base is either cordate or sagittate, naked basal veins,

midrib multistranded, two intramarginal veins running near the margin, secondary veins

eucamptodromous, intersecondary veins present and parallel to secondaries, most tertiary veins

mixed opposite/alternate percurrent, but exmedially they merge perpendicularly with the

innermost intramarginal vein, higher order veins irregular polygonal to orthogonal reticulate.

Holotype. ING-NMNH-0902, only specimen. Plate 59-4-5 and 6.

Species description. Holotype incomplete; laminar shape ovate, length 6.2 cm, width 3.8

cm (Plate 59-4 and 6); margin entire; posterior division of the blade not completely preserved but

the presence of laminar tissue proximal to the petiole attachment shows the base is cordate or

sagittate; naked basal veins form part of the leaf margin near the petiole attachment; anterior









division of the blade longer than posterior division; midrib multistranded; the holotype preserves

6 pairs of secondary veins on the right side of the blade. The most basal secondary vein curves in

a proximal direction and feeds the basal lobe. The next most distal secondary is oriented almost

perpendicular to the midrib for much of its course, forms the primary intramarginal vein (Plate

59-5), and displaces toward the margin a minor intramarginal vein that arises from the posterior

division of the blade. The third secondary vein forms the secondary intramarginal vein, and the

rest of the secondary veins are strongly eucamptodromous and ascend from the midrib at angles

between 290 to 350; one or two intersecondary veins are present between each pair of

secondaries and have courses almost parallel to the secondaries. Both secondary and

intersecondary veins are strongly decurrent on the midrib and follow a course that curves toward

the margin distally. The tertiary veins are mixed opposite/alternate percurrent, but exmedially

most tertiaries are opposite percurrent and are oriented perpendicularly to the midrib (Plate 59-

5), except for those developed in the basal lobe which tend to be parallel to the midrib. Higher

order veins are irregular polygonal to orthogonal reticulate.

Derivation of specific epithet. From the Wayuu, an Amerindian group of the La Guajira

Peninsula, Rancheria Basin.

Source, age and stratum. Colombia, Rancheria Basin, Cerrej6n Formation, Cerrejon coal-

mine, locality 0315, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late

Paleocene, coordinates 11.13N, 72.57W.

Species diagnosis. Anthurioidea cerrejonense Herrera, Jaramillo, Dilcher, Wing et

Gomez-N. Margin entire, leaf apex acuminate, midrib stout and multistranded, secondary veins

eucamptodromous with dichotomies and holding minor agrophic veins, one intramarginal vein









running very close to margin, intersecondary veins occur; tertiary and higher order veins

reticulate, fimbrial vein present.

Holotype. ING-0804, only one specimen found. Plate 60-7-8-9-10, and 11.

Species description. Holotype incomplete, blade is >9.5 cm long and >24.3 cm wide

(Plate 60-7 and 11); margin entire; leaf apex acuminate; midrib stout and multistranded; five

pairs of secondary veins (preserved on the anterior division of the blade) which depart the midrib

at about -500 proximally, shifting to <200 near the apex; secondary veins branch at -450 and

arch apically (Plate 60-8); one or two pairs of intersecondary veins are present for each pair of

secondaries, and their course are more or less parallel to the them; secondary and intersecondary

veins are strongly decurrent on midrib; minor agrophic veins present, an intramarginal vein runs

very close (-0.5 mm) to margin (Plate 60-9); tertiary veins form polygons of different shapes and

sizes (Plate 60-8-9 and 10); higher order veins also form irregular polygons measuring between 5

mm and 8 mm in length and width; a fimbrial vein runs along the margin (Plate 60-9).

Derivation of specific epithet. From the Cerrej6n Formation, where the fossil was found.

Source, age and stratum. Colombia, Rancheria Basin, Cerrej6n Formation, Cerrej6n coal

mine, locality 0315, Pit Tabacol, locality placed between coal beds 100 and 102, middle to late

Paleocene, coordinates 11.13N, 72.57W.

Leaf morphology comparison within monocots. The family Araceae is placed in the

Alismatales, an order of fourteen families (APG-II, 2003) of predominantly herbs of aquatic

habitat. Most Alismatales have basic monocot leaves: oblong, and entire-margined with parallel

major veins and short, perpendicular cross veins. Araceae are unusual in the order for having a

strong similarity to dicot leaves (Keating, 2003b). Although some Alismataceae,

Limnocharitaceae, and Potamogetonaceae and outside of Alismatales, Dioscoreaceae









(Dioscoreales) have ovate to elliptic blades with cordate to sagittate bases and acrodromous or

campylodromous venation that resemble Araceae (see Riley and Stockey (2004) for a discussion

of leaf morphology in these families), unlike these four families, neither fossil of Anthurioidea

has primary veins running in strong convergent arches toward the leaf apex typical of

acrodromous or campylodromous venation. Furthermore, many species within Alismataceae,

Limnocharitaceae, Potamogetonaceae and Dioscoreaceae have dominantly opposite percurrent

tertiary veins unlike those in Anthurioidea.

Systematic affinity within Araceae. Anthurioidea wayuuorum and Anthurioidea

cerrejonense closely resemble species in the extant Neotropical genus Anthurium Schott. This

genus is placed in the tribe Pothoeae Engler (Keating, 2003; 2004) with Pothos (-70 spp., from

South-Southeast Asia, and Madagascar), Pedicellarum (1 sp., from Indonesia and Malaysia), and

Pothoidium (1 sp., Southeast Asia). Keating (2003b) recognized different types of leaf

morphologies within the tribe ranging from primary and secondary veins as acrodromous,

brochidodromous, and eucamptodromous, and higher orders reticulate. Clearly, Anthurium is

extremely more diverse and variable in leaf shape, size and venation types compared to the other

genera within the tribe. A. wayuuorum and A. cerrejonense do not resemble any of the typical

leaf shapes and characters present in these three genera: linear-lanceolate to ovate or narrowly

elliptic blades, furthermore the three genera lack either cordate or sagittate bases as seem in

extant Anthurium and the two fossil taxa.

Anthurium is restricted to tropical-subtropical America and the West Indies and together

with Philodendron and Monstera are perhaps the most variable genera in terms of leaf

morphology (Grayum, 1990; Ray, 1990). However, the combination of leaf characters such as

secondary veins forming intramarginal veins and cordate or sagittate bases and higher order









veins variables seem to be typical and diagnostic for the genus Anthurium (Croat and Bunting,

1979). A. wayuuorum and A. cerrejonense have similar venation patterns to many modern

species of Anthurium in terms of intramarginal venation, secondary and higher order veins types

(Table 2-1). A. wayuuorum is comparable to modern species that have two intramarginal veins

(Plate 61-12-13-14, and Table 2-1). In contrast, A. cerrejonense is similar to extant Anthurium

species that possess only a single intramarginal vein (Plate 61-15-16, and Table 2-1) running

about -0.5 mm from the leaf margin (Plate 60-9). Although, these differences in submarginal

vein patterns are distinctive characters used to discriminate between species, in some very rare

cases we found both types of submarginal veins present in extant species of Anthurium (Plate 61-

14), although this morphological variation changes only a small part of the margin and involves

one secondary vein simply. The specimens presented here are thought to represent two distinct

fossil species based on differences in: tertiary vein patterns, presence and absence of dichotomy,

agrophic and fimbrial veins, and finally the dissimilarity between angles of the secondary veins

(Table 2-1).

Although, there are several good leaf characters (secondary veins forming intramarginal

veins and cordate or sagittate bases and higher order veins variables) that are recognizable and

useful for many members in Anthurium, those characters are not unique for the genus. Keating

(2003b) distinguished the reticulate high order veins as those found in the tribe Pothoideae

(Keating, 2003b) and the subfamily Lasioideae as useful systematically. However, there is no

way to differentiate members of this tribe and subfamily based only on the higher order veins.

Both species reported here share similar higher order veins to those found in the tribe Lasioideae.

Also A. cerrejonense has comparable secondary vein types that end in one single intramarginal

vein. However, extant members of Lasioideae have other leaf venation patterns that are unique









for that subfamily and that are not present in the Cerrej6n fossils neither in the genus Anthurium.

For example in the genus Urospatha (Table 2-1) and all genera within the subfamily, generally

the leaves with pinnate venation have the posterior division of the blades more longer than the

anterior division, this is reflected in having deeply sagittate bases, few numbers of apical

secondary veins (usually 1 to 3), secondary veins angle very low, and their courses run in strong

convergent arches toward the leaf apex. None of the fossils reported here shows all these leaf

venation patterns, for this reason the fossil taxa are most closely related to Anthurium than to

another genus within the family by the combination of blade morphology and the venation

patterns, the assignation of the generic name was discussed previously on the Method section.

Subfamily. Philodendroideae Engler.

Tribe. Philodendreae Schott.

Genus. Montrichardia Cruger.

Generic leaf diagnosis. Leaf apex rounded, margin entire, cordate or sagittate base, each

basal lobe fed by a secondary vein that branches several times, naked basal veins, midrib and

secondary veins stout and multistranded, secondary veins crowded basally and attached to the

midrib at very obtuse angles basally (-90-100) and decreasing apically to -450, secondaries

merging at several intramarginal veins in a complex brochidodromous pattern, tertiary veins

attached to the midrib and secondaries at angles between 200 and 450 to form a network of

intersecondary venation, higher order veins are strongly irregular polygonal reticulate that have

wandering courses and connections at variable angles.

Fossil species. Montrichardia aquatica Herrera, Jaramillo, Dilcher, Wing et Gomez-N sp.

nov.

Fossil holotype. ING-0904. Fig. 23, 24, 25, and 26.









Fossil paratypes. ING-0808, Fig. 27, 28, 29, and 30; ING-NMNH-0903, Fig. 31, 32, and

33; ING-0905 and ING-0906, Fig. S5 (see Supplemental Data accompanying online version of

this article).

Species diagnosis. Montrichardia aquatica Herrera, Jaramillo, Dilcher, Wing et Gomez-N.

Same as for the genus Montrichardia.

Species description. Maximum length measured on the holotype (Plate 62-23) is -56 cm,

and maximum width measured at the posterior division -26 cm, the paratype ING-0808 (Plate

63-27) is -36 cm long and -26 cm wide measured from the midrib to the margin, ING-NMNH-

0903 (Plate 63-31) correspond to a portion of leaf margin that reaches a width of 7.3 cm and a

length of 6.8 cm, ING-0905 reaches a maximum length of -41 cm measured marginally and a

maximum width of -21 cm, ING-0906 is a fragment of the posterior division that reaches a

maximum length of about 11 cm and maximum width of 22 cm; margin entire; leaf apex

rounded; midrib multistranded and up to 1.5 cm wide; leaf base cordate or sagittate (Plate 62-24,

and 25), the basal lobes are fed by the most basal secondary veins, these secondary veins are

very well-developed and form part of the leaf margin at the base as naked basal veins (Plate 62-

25), the two basal secondary veins also dichotomize within the lobes several times to form minor

secondaries that have perpendicular courses with respect to the midrib and very obtuse angles

basally (Plate 62-24 and 25); secondary veins are multistranded, very thick and decurrent on the

midrib, the holotype and the paratypes ING-0808 and ING-0905 (Plate 62-26) preserve up to 10

and 11 secondary veins on the anterior division of the blade, secondary veins crowded basally

and four veins depart from the petiole insertion, secondary veins branch in a complex

brochidodromous pattern, never reaching the margin (Plate 63-31); secondary veins on the

anterior division are attached to the midrib at very obtuse angles, basally -90-100 but









decreasing apically to -450, the course of the secondary veins is straight to slightly up curved

exmedially; tertiary veins are very decurrent on the midrib and secondary veins at angles

between 200 and 450 (Plate 63-29 and 30), tertiary vein spacing is 1-6 mm (Plate 63-30 and 32);

tertiary veins join to form an intersecondary vein network, and are joined exmedially to the

secondaries by higher order veins, forming a system of intramarginal veins very close to margin

(Plate 63-31); the course and spacing in between the intramarginal veins are very irregular;

higher order veins are irregular polygonal reticulate with wandering courses (sinuous, angular, or

straight) and connections at variable angles (Plate 63-32 and 33).

Derivation of specific epithet. From the Latin aquaticus, "living in or near water.".

Source, age and stratum. Colombia, Rancheria Basin, Cerrej6n Formation, Cerrej6n coal

mine, ING-0904, ING-0905 and ING-906 were found at Pit Tabaco Extension, locality 0708 and

0705, localities placed between coal beds 165 and 170, coordinates 11.07N, 72.34W; ING-0808

and ING-NMNH-0903 were found at pit Tabaco 1, locality placed between coal beds 100 and

102, coordinates 11.15N, 72.55W, age middle to late Paleocene.

Leaf morphology comparison within angiosperms. Outside of Araceae, the only

families that share characters to Montrichardia aquatica are Nymphaeaceae and

Nelumbonaceae. These two families can be compared to the fossil exclusively in terms of size

and base shape, see Upchurch et al. (1994), and Wang and Dilcher (2006) for a complete

revision of the leaf morphology in these families. Nymphaeaceae and Nelumbonaceae lack

intramarginal and intersecondary venation, naked basal veins, and higher order veins that

characterize the Cerrej6n fossil and any araceous leaf venation. There are no extant monocot

families other than to Araceae that have all the characters seen in M. aquatica.









Systematic affinity within Araceae. The leaf form and venation patterns of the fossil

leaves are extremely similar to those found in the extant genus Montrichardia Cruger (Plate 61-

17 and 18) which has two species M. arborescens and M linifera. This genus is placed in the

tribe Philodendreae Schott (Keating, 2003; 2004) with Anubias (8 ssp., from tropical Africa),

Furtadoa (2 spp., from Indonesia and Malaysia), Philodendron (-500 ssp., from Mexico to

Argentina), and Homalomena (-110 spp. from tropical America and Southeast Asia). Keating

(2003b) recognized several leaf venation patterns for the tribe Philodendreae which range from

primary and secondary veins as acrodromous, brochidodromous, and eucamptodromous, and for

having mostly many closely spaced intersecondary veins parallel each other. The DNA data and

the anatomical characters situate this subfamily as the most varied and rich in the entire Araceae

(Keating, 2004). In addition, it has been indicated that Montrichardia has some genetic isolation

in respect of the other four genera within the tribe Philodendreae (Cabrera et al., 2003; Keating,

2004). This "isolation" is also seen in terms of leaf venation, Montrichardia is easily separated

from the other four genera for having the "colocasioid venation", and no species within those

four genera have this peculiar venation character.

The genus Montrichardia is restricted to tropical America and the West Indies. M.

aquatica has "colocasioid venation," defined as "a type of higher order leaf venation in which the

finer veins branch almost at right angles from the primary lateral veins and then arch strongly

towards the leaf margin, often fusing along the way to form a more-or-less sinuous interprimary

collective vein between the primary lateral veins, and finally joining within the margin to form a

intramarginal collective vein" (Mayo et al., 1997, page 310). Colocasioid venation is known in

the extant tribes Caladieae (New World, except for Hapaline from Southeast Asia and Brunei)

and Colocasieae (Old World), both tribes are part of the subfamily Aroideae, and seem to be very









well separated genetically each other (Keating, 2004). Colocasioid venation has been reported in

fossil leaves from the Neogene of Trinidad and the Eocene of Germany (Berry, 1925; Wilde et

al., 2005, Table 2-1). M. aquatica has a similar leaf shape and intersecondary venation to

Caladieae and Colocasieae (Plate 61-19), but different intramarginal vein and higher order vein

patterns. In Caladieae and Colocasieae the secondary veins (secondary veins are called primary

lateral veins by Mayo et al., 1997) merge bluntly with one to three straightened intramarginal

veins (Plate 61-19, Table 2-1), while the secondary veins in the fossil and extant Montrichardia

branch very close to the margin subsidized by higher order veins that finally form several

intramarginal veins. Also the spacing and course of these intramarginal veins are very irregular

compared to tribes Caladieae and Colocasieae in which the intramarginal veins run very parallel

to the margin and each other, and the vein spacing is constant (Plate 61-18, 20, and Plate 63-31).

Higher order veins and areolation in Colocasieae and Caladieae are dominated by cross venation

and elongated polygons of similar shape and size, while living Montrichardia and the fossil

leaves show higher order veins that anastomose irregularly with wandering courses, connections

at variable angles, and do not form well-developed areolation (Plate 61-21 and 22, Plate 63-30,

32, 33 and Table 2-1). Because the intramarginal venation pattern and higher order veins

characterize the extant genus Montrichardia and the fossil M. aquatica, we do not relate the

Cerrej6n fossil leaves to the fossil morphogenus Caladiosoma Berry, nor to its living relative

tribes Caladieae and Colocasieae.

Discussion

Sedimentary Deposition and Paleoclimate

Anthurioidea wayuuorum and Anthurioidea cerrejonense were recovered from a lens-

shaped deposit of laminated, gray-black, fine-grained sandstone approximately 50 cm thick and

-6 meters in lateral extent (locality 0315). A total of 239 leaf specimens have been found in this









locality. Most leaf specimens are complete and well preserved; sizes range from microphyll to

macrophyll with mesophyll leaves being dominant. Montrichardia aquatica (ING-0808) was

recovered approximately 3 meters below locality 0315; the rock is a brown, fine-grained lithic

sandstone with inclined heterolithic strata. These araceous fossils were deposited in fluvial

channel sediments. The fine-grained sandstones with laminations and heterolithic stratification

suggest overbank deposits, such as crevasse splays and forebanks (Bayona et al., 2004). These

deposits grow by accumulation of fine-grained sediment during episodes of flooding (Reading

and Levell, 1996)

The specimen ING-NMNH-0903 ofM. aquatica was found at the same stratigraphic level

as locality 0315, between coal beds 100 and 102, in a dark grey siltstone. The holotype of M

aquatica (ING-0904) and paratypes ING-0905 and 0906 were recovered from an extensive gray

siltstone bed approximately 60-90 cm thick (localities 0705 and 0708). The gray siltstone

underlies coal bed 170, one of the thickest coal beds at the top of the Cerrej6n Formation (-7 m).

These siltstones were deposited in a swampy-lacustrine environment.

Leaf margin and leaf area analysis of the entire associated flora suggest a mean annual

precipitation of 4 meters and a mean annual temperature > 24 C (Herrera et al., 2005). The

most abundant families and orders recognized in the Cerrej6n flora are Leguminosae, Arecaceae,

Malvales, Lauraceae, Menispermaceae, and Zingiberales. The Cerrej6n paleoclimate and its

floristic composition suggest the presence of a Neotropical rainforest during the middle-late

Paleocene in Northern South America (Wing et al., 2004; Herrera et al., 2005).

Paleoecology

Lithofacies, sedimentary structures, and the preservation of large leaves within the

associated paleoflora suggest that Anthurioidea wayuuorum and Anthurioidea cerrejonense were

part of the local flora, rather than having being transported from far away (Burnham, 1989). A









reconstruction of the life forms of A. wayuuorum and A. cerrejonense must be based upon what

is known of the closest living genus. Most Anthurium species are epiphytes (Croat, 1990). It has

been estimated that in some Amazonian floodplain forests -90% percent of trees carry epiphytic

aroids (Leimbeck and Balslev, 2001); such common epiphytes might have been preserved as

fossils. The sedimentary environment also raises the possibility that the fossils were rheophytes,

but this life form is rarely found in tropical American Araceae (Croat, 1990).

Montrichardia possesses a strict helophytic life form (Croat, 1990; Mayo et al., 1997).

Montrichardia grows in freshwater habitats, especially fluvial to coastal plain areas, along river

margins and adjacent swamps subject to seasonal changes in water level (Croat, 1990; Bown,

2000). The foliage generally remains above the water level, forming dense colonies that trap

sediment; it is a primary colonist in swamp forest areas (Bown, 2000). Montrichardia is found

throughout tropical America, but it is dominant along large rivers of central Amazonia and

deltaic zones where it grows even under tidal influence (Mayo et al., 1997; Bown, 2000).

Lithofacies and sedimentary structures suggest that M. aquatica grew along river margins

(forebanks), and swamps, where it coexisted in palm-dominated environments similar to the

ecology of extant Montrichardia.

Paleobiogeography and Palynology

The early appearance of the Araceae family in the fossil record (early Cretaceous), even

before the more diverse families seen today, is remarkable. However, the aroid fossil record is

sparse and most araceous fossils belong to taxa that inhabited ponds, lakes or flooded areas (e.g.

Dilcher and Daghlian, 1977; Kvacek, 2003). A complete revision of the Araceae macrofossil

record was recently made by Wilde et al. (2005); previous revisions include Grayum (1990),

Mayo et al. (1997), and Keating (2003a). It seems clear that the geographic distribution of

Araceae in the past was driven largely by the global climate (Plate 58-1, 2 and 3). From the early









Cretaceous (-120 Mya) to the early Paleogene (-65-40 Mya) the climate was warmer than today

during a period of about 60 million years (Zachos et al., 2001; Bralower et al., 2006). Thus

permitted migration and evolution of megathermal plants into current temperate latitudes

(Morley, 2003). By the end of the Cretaceous, three or four Araceous subfamilies were present,

according to the most recent classification by Keating (2003b; 2004) (Plate 58-1 and 2;

Pothoideae, Lemnoideae, Orontioideae, and Aroideae (?)), and during the early Paleogene three

other subfamilies and consequently a more diverse record of new tribes appear in the fossil

record, (Plate 58-3; Philodendroideae, Aroideae, and Lasioideae).

The fossil pollen record also shows interesting aspects of the paleobiogeography of

Araceae. Recently Hesse and Zetter (2007) made a complete review of the family's pollen

record, recognizing three pollen types attributable to Araceae: Limnobiophyllum (subfamily

Lemnoideae), Spathiphyllum (Pothoideae), and the almost globally distributed Proxapertites

operculatus (Philodendroideae/Pothoideae). The Cerrej6n palynoflora has high abundance and

in some samples dominance of araceous pollen. Jaramillo et al. (2007) studied the palynoflora

through the entire formation, comparing coal (associated with lacustrine-swampy environments)

and non-coal samples (associated with fluvial and estuarine plains). The three araceous pollen

grains present in the Cerrej6n palynoflora are Proxapertites operculatus (Van der Hammen,

1954) Van der Hammen, 1956, Proxapertites cursus Van Hoeken-Klinkenberg, 1966, and,

Spathiphyllum vanegensis (Van der Hammen & Garcia, 1966) Hesse & Zetter 2007.

Proxapertites operculatus is one of the most abundant pollen types in both coal and non-coal

samples. On the other hand, Proxapertites cursus and Spathiphyllum vanegensis are present only

in non-coal samples. This distribution by depositional environment suggests that Araceae may

have been ecological diversified by the Paleocene.









Comments on Aroid Evolution

In the modem Neotropics, Araceae is diverse and shows high endemism. Anthurium and

Philodendron contain approximately 50% of the total of species for the family (Croat, 1983).

Anthurium is extremely diverse along the wet, Pacific side of the Andes in Colombia and

Ecuador (Croat, 1992b; Vargas et al., 2004; Mora et al., 2006). If the fossils reported here as

Anthurioidea really have affinities with the extant Neotropical genus Anthurium, it is interesting

to think that related forms of this genus have been present in the Neotropics since the Paleocene.

How Anthurium reached its modem high level of diversity is still a matter of discussion, but

probably its high diversity and endemism is related to the Mio-Pliocene Andean orogeny

(Gentry, 1982) and the resulting biogeographic and ecological factors in Andean and lowland

populations, respectively (Leimbeck and Balslev, 2001; Jacome et al., 2004). Other genera along

the Andean foothills seem to share similar diversity patterns (Richardson et al., 2001; Pirie et al.,

2006; Quijano et al., 2006).

Montrichardia has only two species today with widespread distributions in the Neotropics

(Mayo et al., 2007; Bown, 2000). Montrichardia has been present in the Neotropics for at least

60-58 Mya, but it did not diversify to the extent seen in Anthurium, Philodendron or Monstera.

Alternatively it could have diversified and then lost its diversity. Perhaps Montrichardia is more

influenced by long-distance dispersal by being near moving water, thus maintaining constant

genetic interchange over long distances and reducing the probability of speciation.

Montrichardia also inhabits a stable aquatic environment and it has not changed since the

Paleocene. The same water life forms and stable environments have also been seen to preserve

the integrity of another fossil angiosperm to the species level in Ceratophyllum (Herendeen et al.,

1990). This hypothesis might be tested by integrating the fossil record together with molecular

analysis in future studies. The long history of araceous Neotropical genera and their current









contrasting differences in species diversity constitutes a good case study to investigate the causes

of species diversity in the Neotropic Rainforests today.

The presence of abundant fossil leaves and pollen grains in the Cerrej6n Formation

establishes the fact that Araceae was already an important component in floodplains of coastal

tropical humid forests during the early Cenozoic. The Cerrej6n fossils are related to at least three

modern tribes which may have had strong habitat preferences as modem aroids.













Table 2-1. Morphological features of fossil leaf taxa and extant Araceae.


Naked basal 20 vein
Taxon Blade Apex Base vei 2 vein type Cowin
veins -Crowding


Caladium bicolor (Aiton) Vent
Xanthosoma mexicanum Liebm
Xanthosoma sagattifohum Schott
Xanthosoma robustum Schott
X. undipes (K.Koch & C.D.Bouche) K.Koch
Xanthosoma violaceum Schott
Colocasia esculenta (L.) Schott
S Ilanoense Croat
S. schottzanum H.Wendl. ex Schott
Montrichardia arborescens (L.) Schott
Montrichardia hnifera (Arruda) Schott
Montrichardia aquatica"
Caladiosoma messelense Wilde, Z. Kvacek & Bognerb
Caladiosoma miocenicum Berryb
Anthurum oxybelium Schott
Anthurum variegatum Sodiro
Anthurum subsagittatum (Kunth) Kunth
S Anthurium nigrescens Engl
Anthurum marmoratum Sodiro
Anthurum karstenzanum Engl
Anthurum concinnatum Schott
Anthurum browniu Mast
Anthurum rzedowski Croat
Anthurum coripatense N.E.Br. ex Engl
Anthurnm ** Schott
Anthurum cotobrusu Croat & R.Baker
Anthurum caperatum Croat & R.Baker
A. obtuslfolum (W.T.Aiton) G.Don in R.Sweet
Anthurum ochranthum K.Koch
Anthurum. ravens Croat & R.Baker
Anthurum roseospadix Croat
Anthurum sanctifidense Croat
Urospatha fredrzchsthahl Schott
Urospatha sagzttifola (Rudge) Schott
Urospatha somnolenta R.E.Schult
Anthurioidea wayuuorum"
Anthurioidea cerreionense"


Ovate Convex Cordate-peltate
Ovate Acute Cordate
Elliptic Acute Cordate
Ovate Obtuse Cordate
Ovate Acute Cordate
Elliptic Acute Cordate
Ovate Acute Cordate-peltate
Elliptic Acute Cordate
Elliptic Acute Hastate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Rounded Cordate-sagittate
Ovate Acute Cordate
e A e
Ovate Acute Lobate
Ovate Acute Lobate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Hastate
Ovate Acute Lobate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Lobate
Ovate Acute Lobate
Ovate Acute Lobate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Cordate
Ovate Acute Deeply sagittate
Ovate Acute Deeply sagittate
Ovate Acute Deeply sagittate
Ovate ? Cordate/sagittate
? Acute ?


Absent
Present
Present
Present
Present
Absent
Absent
Absent
Absent
Present
Present
Present
?9

Present
Present
Present
Present
Present
Absent
Present
Present
Absent
Present
Present
Present
Present
Present
Present
Present
Present
Present
Present
Present
Present
Present


Straightened I.V Present
Straightened I.V Present
curved I.V Present
Straightened I.V Present
Straightened I.V Present
Straightened I.V Absent
Straightened I.V Present
Straightened I.V Present
Straightened I.V Present
Brochidodromous Present
Brochidodromous Present
Brochidodromous Present
Straightened I.V Present
? ?
Eucamp-brochido Absent
Eucamptodromous Absent
Eucamptodromous Absent
Eucamp-brochido Absent
Eucamptodromous Absent
Eucamp-brochido Absent
Eucamp-brochido Absent
Eucamptodromous Absent
Eucamptodromous Present
Eucamptodromous Absent
Eucamptodromous Absent
Eucamptodromous Present
Eucamptodromous Absent
Eucamptodromous Absent
Eucamptodromous Absent
Eucamptodromous Absent
Eucamptodromous Absent
Eucamptodromous Absent
Eucamptodromous Present
Eucamptodromous Present
Eucamptodromous Present
Eucamptodromous Absent
Eucamotodromous ?


No. posterior No. anterior
2os 2os
4 6-8
4 4-5
4 5-6
4 7
4 6-7
4 7
8 5-6
6 15-17
6 11-13
4 6-11
4 6-11
4 10-11
? 8-9
? ?
6 7-9
4 7-9
4 7-8
6 8-9
6 7-9
4 6-7
4 8-9
4 8
8 6-7
4 10-11
6 5-8
4 7-8
6 9-10
6 8-9
6 9-10
4 8-10
6 11-12
7 9-10
4 1-2
6 1-3
6 1
4 >4
? >5












3 vein
No. intra-
2o veins angle No. intersecondaries Fimbrial vein 3 vein type Exmedial Higher order veins Glands
marginalsangle
angle


900-600
900-300
900-300
900-450
900-600
900-600
900-300
900-300
900-300
900-400
900-400
900-450
800-400

45o-400
450
45o-30
40o-300
45o-400
450
50o-450
450
S 400-300
S 450-400
400-300
600-400
450-400
500-450
500-450
450-300
400-300
450-300
30-150
350-200
350-200
350-300
500-200


2
2
2
2
2
1
1-2
3
3
1-4
1-4
1-4
Several
?
2
2
2
2
2
2
2
2
2
1
1-2
1
2
1
2
2
2
2
1
1
1
2
1


Colocasioid
Colocasioid
Colocasioid
Colocasioid
Colocasioid
Colocasioid
Colocasioid
Colocasioid
Colocasioid
"Colocasioid"
"Colocasioid"
"Colocasioid"
Colocasioid
Colocasioid
Mixed opp/alt
Mixed opp/alt
Mixed opp/alt
Mixed opp/alt
Mixed opp/alt
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Mixed opp/alt
Regular poly. ret.
Mixed opp/alt
Regular poly. ret.
Regular poly. ret.
Regular poly. ret.
Mixed opp/alt
Regular poly. ret.


Regular polygonal ret.
Regular polygonal ret. squarish
Squarish
Regular polygonal ret.
Regular polygonal ret.
Several
Regular polygonal ret. -squarish
Irregular polygonal ret.
Irregular polygonal ret.
Irregular polygonal ret.
Irregular polygonal ret.
Irregular polygonal ret.
Regular polygonal ret. -elongate
Irregular polygonal ret.-elongate
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-90 to midrib Regular polygonal ret.
-90 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-90 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
Regular polygonal ret.
-900 to midrib Regular polygonal ret.
Regular polygonal ret.
Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-90 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.
-900 to midrib Regular polygonal ret.


Notes
Taxon in bold text correspond to araceous fossil leaves.
aCerrej6n Fossils.
bWilde et al., 2005.

I.V. (intramarginal veins); Mixed opp/alt (M. opposite/alternate percurrent); Regular poly. ret. (R. polygonal reticulate); Eucamp-

brochido. (eucamptodromous-brochidodromous).


Absent
Hairs
Hairs
Hairs
Absent
Present
Absent
Present
Present
Present
Present
?
?

Absent
Absent
Present
Present
Absent
Present?
Present
Present
Absent
Present
Present
Absent
Present
Absent
Present
Absent
Present
Present
Present
Present
Present
Absent
Absent









APPENDIX A
PHOTOGRAPHIC PLATES


Plate 1 1. Exemplar, whole leaf.




































Plate 2 1. Exemplar, whole leaf. 2. Exemplar, toothed margin, close up. 3. Exemplar,
areolation, close up.








































Plate 3 1 Exempl hole leaf
a, whole leaf










































Plate 4 1. Exemplar, whole leaf 2. Exemplar, secondary veins joining mid vein, close up
areolation.




































Plate 5 1. Exemplar. 2. Leaf 2.













































Plate 6 1. Exemplar, sterile shoot with leaves. 2. Exemplar, close up showing leaves





































Plate 7 1. Exemplar, whole leaf. 2. Leaf 2. 3. Leaf 2, Secondary veins
4. Exemplar petiole, close up.


at the base, close up.















----





-I
--1
























1 2





Plate 8 1. Exemplar, whole leaf, asymmetrical blade. 2. Leaf 2, falcate apex.





























87




































Plate 9 1. Exemplar, leaf fragment. 2. Secondary junctions with primary vein, and toothed
margin 3. Leaf 2, decurrent base.






































Plate 10 1. Exemplar, leaf fragment. 2. Exemplar, tertiary veins and laminar resin dots, close
up. 3. Whole leaf 2.





























Plate 11 1. Exemplar, whole leaf. 2. Whole leaf 2. 3. Leaf 2, tertiary veins and fourth order
veins near lateral primary. 4. Leaf 2, higher order veins, close up.


e~st~8~11 ~I




































Plate 12 1. Exemplar, whole leaf 2. Exemplar, tertiary veins and fourth order veins near
primary vein, close up.



































Plate 13 1. Exemplar, whole leaf. 2. Primary vein being deflected at each secondary juncture
near apex. 3. Margin, close up. 4. Tertiary veins and higher order veins near mid vein, close up.



































Plate 14 1. Exemplar, three lobed leaf. 2. Secondary veins on lateral primary.







































Plate 15 1. Exemplar, whole leaf. 2. Secondary vein junction with mid vein and composite
intersecondary. 3. Admedially ramified tertiary veins near mid vein, close up.





































Plate 16 1. Exemplar, whole leaf. 2. Whole leaf 2. 3. Leaf 2, tertiary veins and fourth order
veins near mid vein. 4. Whole leaf 3.








































Plate 17 1. Exemplar, whole leaf. 2. Whole leaf 2.









































Plate 18 1. Exemplar, whole leaf. 2. Secondary veins at the base. 3. Closely spaced tertiary
veins near mid vein, close up.






















*7







,I










Plate 19 1. Exemplar, whole leaf 2. Secondary and tertiary veins near mid vein. 3. Higher order
veins, close up.









































Plate 20 1. Exemplar, whole leaf. 2. Secondary and tertiary veins near mid vein.













































Plate 21 1. Exemplar, whole leaf 2. Tertiary veins near the margin. 3. Secondary and tertiary
veins decurrent on mid vein.








































Plate 22 1. Exemplar. 2. Secondary and tertiary veins near mid vein.
























101

























2 2 -


Plate 23 1. Exemplar, whole leaf. 2. Closely spaced tertiary veins near mid vein, close up.











































Plate 24 1. Exemplar, whole leaf. 2. Secondary and tertiary veins near mid vein. 3. Higher order
veins and laminar resin glands, close up.






















103


!.. 1 ..

o". ., ..





































Plate 25 1. Exemplar, whole leaf 2. Leaf 2, long petiole. 3. Leaf 3, toothed margin.







































Plate 26 1. Exemplar, whole leaf. 2. Exemplar, toothed margin, close up 3. Whole leaf 2.

























4.
r 3A

r


2

3 It. --


Plate 27 1. Exemplar, whole leaf 2. Leaf 2. 3. Leaf 2, toothed margin, close up.


J~tam.








































Plate 28 1. Exemplar, whole leaf. 2. Exemplar, toothed margin. 3. Exemplar base, close up.








































Plate 29 1. Exemplar, whole leaf 2. Exemplar, exmedial tertiary veins. 3. Exemplar, higher
order veins and areoles close up.







































Plate 30 1. Exemplar, whole leaf. 2. Exemplar, secondary and tertiary veins near mid vein.
















































Plate 31 1. Exemplar, whole leaf 2. Exemplar, toothed margin. 3. Exemplar, tertiary and fourth
order veins.




































Plate 32 1. Exemplar, whole leaf 2. Exemplar, secondary and tertiary veins near mid vein.







































Plate 33 1. Exemplar, whole leaf. 2. Exemplar, toothed margin. 3. Whole leaf 2. 4. Leaf 3,
secondary and tertiary veins near mid vein, close up.






































Plate 34 1. Exemplar, whole leaf. 2. Exemplar, tertiary and higher order veins near mid vein.











































Plate 35 1. Exemplar, whole leaf. 2. Exemplar, tertiary and higher order veins near mid vein.











&l~s&~.A' rji

'Xa


Plate 36 1. Exemplar, whole leaf 2. Exemplar, secondary veins at the base and pulvinate
petiole. 3. Whole leaf 2.




















72






















Plate 37- 1. Exemplar, whole leaf 2. Exemplar, tertiary and higher order veins, close up. 3. Leaf
2






























116
.;'2 ',. r




































Plate 38 1. Exemplar, whole leaf. 2. Exemplar, tertiary veins decurrent on mid vein. 3. Leaf 2.









2

M^[


Plate 39 1. Exemplar, whole leaf. 2. Leaf 2, emarginate tooth, close up. 3. Leaf 2, agrophic
veins near toothed margin. 4. Whole leaf 3.












A2
IMFr
i .^. 41'i
isy~1 (r^ "' .- *


Plate 40 1. Exemplar, whole leaf 2. Exemplar, toothed margin, close up.




































Plate 41 1. Exemplar, whole leaf. 2. Leaf 2. 3. Leaf 2, tertiary and higher order veins, close up.









































Plate 42 1. Exemplar, whole leaf. 2. Exemplar, tertiary veins at junction with lateral primary
vein. 3. Exemplar, freely ending veins and laminar resin dots, close up.













































Plate 43 1. Exemplar, whole leaf. 2. Leaf 2, missing the base.















































Plate 44 1. Exemplar. 2. Attached leaflets. 3. Specimen 3.

















123
















5r
ad1


Plate 45 1. Exemplar, cordate base. 2. Leaf 2, toothed margin close up. 3. Leaf 2,
craspedodromous secondary veins.















124




































Plate 46 1. Exemplar, whole leaf. 2. Exemplar, secondary and tertiary veins decurrent on mid
vein. 3. Exemplar, higher order and freely ending veins.













































Plate 47 1. Exemplar, whole leaf



































Plate 48 1. Exemplar, leaf fragment. 2. Tertiary veins. 3. Higher order veins.





































Plate 49 1. Exemplar, whole leaf. 2. Tertiary and higher order veins, close up.





































Plate 50 1. Exemplar. 2. Exemplar, submarginal vein, close up. 3. Exemplar, secondary veins
and laminar resin dots, close up.




































Plate 51 1. Exemplar, whole leaf. 2. Exemplar, higher order veins, close up. 3. Leaf 2.







































Plate 52 1. Exemplar, whole leaf. 2. Exemplar, tertiary and fourth order veins joining mid vein.
3. Exemplar, higher order veins and laminar resin dots.






































Plate 53 1. Exemplar, whole leaf 2. Exemplar, tertiary veins joining secondary vein. 3.
Exemplar, higher order veins, close up.









































Plate 54 1. Exemplar, whole leaf. 2. Exemplar, secondary veins joining midrib.





































Plate 55 1. Exemplar, whole leaf 2. Exemplar,
midrib.


secondary and tertiary veins decurrent on




































Plate 56 1. Exemplar, fragment of palmate leaf. 2. Leaf 2, fragment.































Plate 57 1. Exemplar, leaf fragment. 2. Leaf 2, fragment.










































* Infructescence & Seeds Early Paleogene 65-40 Ma









Plate 58 1-3. Paleogeographic distribution of Araceae, maps based on Scotese (2001). 1.

Barremian-Aptian in age, Pothoideae subfamily suggested (this is according to the most recent

classification of Araceae by Keating (2003b; 2004)) (Friis et al., 2004). 2. Coniacian,

Lemnoideae (Krassilov, 1973). Campanian, Orontioideae (Kvacek and Herman, 2004; Bogner et

al., 2005), Aroideae? (Stockey et al.,2007). Maastrichtian, Pothoideae (Herngreen, 1974;

Sarmiento, 1994; Bonde, 2000; Hesse and Zetter, 2007). 3. Paleocene, Philodendroideae and/or

Pothoideae (Van der Hammen, 1956; Van der Hammen and Garcia de Mutis, 1966; Jaramillo

and Dilcher, 2001; Hesse and Zetter, 2007); Pothoideae and Philodendroideae (fossil leaves, this

paper); Lemnoideae (Mclver and Basinger 1993; Stockey et al., 1997); Unknown subfamily

(Van Hoeken-Klinkenberg, 1966; Jaramillo et al., in press). Paleocene-Eocene, Aroideae

(Fedotov, 1975). Eocene, Aroideae (Hickey, 1977; Dilcher and Daghlian, 1977);

Philodendroideae and/or Pothoideae (Bogner, 1976; Zetter et al., 2001; Hesse and Zetter, 2007).

Middle Eocene, Lasioideae (Ceballos-Ferriz and Stockey 1988; Smith and Stockey, 2003);

Pothoideae and Aroideae (Wilde et al., 2005); Lemnoideae (Hesse and Zetter, 2007); Late

Eocene, Aroideae, (Wilde et al., 2005).







































1 1


1- .- ,


























veins, scale 3 cm.
veins, scale 3 cm.









Plate 60 7-11. Anthurioidea cerrejonense gen. nov. sp. nov. Holotype (ING-0804). 7. Fossil

leaf preserves part of the middle and the apex of the leaf; scale 10 cm. 8. Right side of the blade,

secondary veins running toward margin; tertiary and higher orders veins showing reticulate

patterns; the arrow shows the region where the margin is preserved and the location of Fig. 9. 9.

The diagonal arrow shows a secondary vein merging with the intramarginal vein, which runs

very close to margin; the vertical arrow shows the fimbrial vein running along the margin; scale

5 mm. 10. Close up of reticulate higher order veins; scale 5 mm. 11. Drawing of veins, dashed

line indicates that apex and middle fragments were not attached so their positions relative to one

another are uncertain.









Plate 61 12-22. Extant aroid leaves. 12. Anthurium lucens Standl (National Herbarium,

#2980121), the species shows two intramarginal veins. 13. A. rotundistigmatum Croat (NH

#3113554), the arrow shows a secondary vein merging with the inner intramarginal vein. 14. A.

ravenii Croat & R.Baker (NH #1793970), the arrow shows a secondary vein ending on the main

intramarginal vein and then merging on a secondary intramarginal vein very close to margin. 15.

A. rubrinervium (Sims) G.Don in R. Sweet (NH #2025269), secondary veins merging on an

intramarginal vein that runs very close to margin. 16. A. cerropelonense Matuda (NH #3093115),

the arrows show secondary veins merging with an intramarginal vein that runs very close to

margin. Notice dominance of opposite percurrent secondary veins exmedially. 17. Montrichardia

arborescens (L.) Schott (NH #3307839). 18. M. linifera (Arruda) Schott (NH #2174078), arrows

on the right side show two secondary veins and arrow on the left side shows the intramarginal

venation pattern. Notice that the secondary veins branch toward margin and form several

intramarginal veins. 19. Colocasia esculenta (L.) Schott (NH #3290213), notice that the

secondary veins merge bluntly with a straight intramarginal vein. 20, 21 and 22. Cleared leaf of

M. arborescens (L.) Schott (NH #3307839). 20. Secondary veins branch toward margin and

form several intramarginal veins. 21. Tertiary veins decurrent on a secondary vein, higher order

veins strongly irregular polygonal reticulate that have wandering courses and connections at

variable angles, bar scale 5 mm. 22. Close up of higher order veins, scale 1 mm.





















































Plate 62 23-26. Montrichardia aquatica gen. nov. sp. nov. Holotype (ING-0904. Figs. 23-26).

23. Fossil leaf preserves partially the base and the anterior division, scale 10 cm. 24. Close up of

leaf base, scale 5 cm. 25. Close up of right lobule, notice naked basal veins, scale 3 cm. 26. Close

up of anterior division, margin preserved on the left side of the blade, scale 5 cm.



144









Plate 63 27-33. trichardia aquatica gen. nov. sp. nov. Paratypes (ING-0808, Figs. 27-30; ING-

NMNH-0903, Figs. 31-33). 27. Fossil leaf incomplete, scale 10 cm. 28. Drawing of veins. 29.

The arrow shows a secondary vein, notice intersecondary venation, scale 3 cm. 30. Tertiary veins

merging with a multistranded secondary vein, notice higher order veins strongly irregular

polygonal reticulate that have wandering courses and connections at variable angles, scale 5mm.

31. Arrow on the left side marks a secondary vein branching toward margin; arrow on the right

side shows several intramarginal veins, scale 2 cm. 32. Close up of tertiary veins and higher

order veins, scale 5 mm. 33. Close up of higher order veins, scale 3 mm.









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BIOGRAPHICAL SKETCH

Fabiany Herrera was born in 1978 in Bucaramanga, Colombia. The youngest of three

children, he grew up in the city of his origin graduating from the public high school "INEM" in

1995. After one year of public service from 1996 to 1997 he started his B.S. program in geology

with emphasis in paleontology in 1998 from the Industrial University of Santander, and obtained

his degree in 2005. He was an awardwinner with the distinction "Thesis of the Year" from the

Colombian Geological Society in 2004. The topic of his thesis was the discovery of the world's

oldest Neotropical rainforest.

Upon graduating in May 2005, Fabiany worked as research assistant in biostratigraphy for

the Colombian Institute of Petroleum in Bucaramanga. Then, he moved to the Republic of

Panama in Central America to work for an internship at the Smithsonian Tropical Research

Institute and later at the National Museum of Natural History, Smithsonian Institution in

Washington DC where he worked on paleobotany topics. During that time, he spent several

months collecting fossils in Colombia, Panama and Wyoming.

In 2006 Fabiany moved to Gainesville, Fl in order to attend to the University of Florida to

get his master's degree in science with major in geology. Upon completion of his masters

program, Fabiany wants to continue in graduate school as a PhD student, and aspires to continue

his contributions to the scientific world.





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1 NEOTROPICAL RAINFOREST ORIGIN S: PALEOFLORA AND PALEOCLIMATE By FABIANY ALBERTO HERRERA-TOLOSA A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2008

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2 2008 Fabiany Alberto Herrera-Tolosa

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3 To my family, my mom, my dad, my nephew Jose ph and my niece Sofia for their happiness. To my friends

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4 ACKNOWLEDGMENTS This research was supported by grants from Carbones del Cerrejn, a graduate research fellowship of the University of Florida, N SF Deep Time grant DEB-0090283 to DLD, NSF grant DEB-0733725, Smithsonian Paleobiology Endowment Fund, the Unrestricted Endowments SI Grants, the Fondo para la Investigacin de Cienci a y Tecnologa del Banco de la Repblica de Colombia, The Explorers Club, Colombian Petr oleum Institute, and Corporacin Geolgica ARES. I deeply thank David Dilcher, Carlos Ja ramillo, Scott Wing and Carolina Gomez for their helpful discussions and contributions. Steven Ma nchester, Ellen Martin and Mark Brenner for comments improving the manuscript. Fernando Chav ez, Camilo Montes, and the geology team at the Cerrejn mine for their help and support during field trips. I thank Dan Nicolson for helping at the National Herbarium and Stanley Yankow ski for his assistance clearing the extant Montrichardia leaf. I thank Josef Bogner, Ma rcela Mora, and Thomas Croat for helpful discussions about the systematics of the fossils and Giovanni Bedoya for his suggestions about the nomenclature. I thank everybody in the Paleobotany lab for their support, finally special thanks go to Edwin Cadena for his friendship and Benjamin Himschoot for reading part of this thesis and such great happiness.

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5 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES…………………………………………………………………………...........7 LIST OF FIGURES................................................................................................................ .........8 ABSTRACT....................................................................................................................... ..............9 CHAPTER 1 MONOGRAPH OF THE CERREJ"N FLORA....................................................................10 Abstract....................................................................................................................... ............10 Introduction................................................................................................................... ..........10 Materials and Methods.......................................................................................................... .11 Fossil Leaf Descriptions....................................................................................................... ..15 Ferns.......................................................................................................................... ......15 Cycads......................................................................................................................... ....17 Conifers....................................................................................................................... ....17 Dicots......................................................................................................................... ......18 Monocots....................................................................................................................... ..48 Paleoclimate................................................................................................................... .........55 Floristic Composition of the Cerrejn Flora...........................................................................56 Conclusions.................................................................................................................... .........57 2 FOSSIL ARACEAE FROM A PALEO CENE NEOTROPICAL RAINFOREST IN COLOMBIA................................................................................................................ ...........59 Abstract....................................................................................................................... ............59 Introduction................................................................................................................... ..........59 Materials and Methods.......................................................................................................... .61 Systematics.................................................................................................................... .........62 Discussion..................................................................................................................... ..........72 Sedimentary Deposition and Paleoclimate......................................................................72 Paleoecology................................................................................................................... .73 Paleobiogeography and Palynology................................................................................74 Comments on Aroid Evolution........................................................................................76 APPENDIX A PHOTOGRAPHIC PLATES..................................................................................................80 LIST OF REFERENCES............................................................................................................. 147

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6 BIOGRAPHICAL SKETCH.......................................................................................................155

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7 LIST OF TABLES Table page 2-1 Morphological features of foss il leaf taxa and extant Araceae..........................................78

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8 LIST OF FIGURES Figure page 1-1 Map and location of Cerrejn mine a nd Cerrejn flora localities (stars)..........................13 1-2 Stratigraphic column of the Cerrej n Formation showing lithology and stratigraphic positions of the plant megafossil localities. Modified from Bayona et al (2004).............14

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9 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science NEOTROPICAL RAINFOREST ORIGIN S: PALEOFLORA AND PALEOCLIMATE By Fabiany Alberto Herrera-Tolosa May 2008 Chair: David Dilcher Major: Geology Little is known about the early history of Ne otropical rainforests, although this vegetation types are extremely diverse, they have a di stinctive spectrum of leaf size and shape, a characteristic floral composition at the family level, no fossil flor as with these unique features have been reported from low latitudes yet. Here I compile a monograph on a middle to late Paleocene (58-60 Ma) flora from the Cerrejn Form ation of northeastern Colombia that is the oldest record of Neotr opical rainforest. Fifty seven fossil mo rphospecies were described; they include 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. The Cerrejn assemblage has a high proportion of large, entire-mar gined, thin-textured leaves indi cating a moist tr opical climate with mean annual temperature ~24 C and rainfall 2.3 m/y. Family-level taxonomic composition is similar to that of extant Neotropical rainforests. Three monocot morphospecies were indentified at the genus level. The fossil leaves belong to the family Araceae, and they are the oldest re port of this family from the Paleogene in northern South America. The three leaf taxa are related to two ex tant Neotropical genera, two of them closely related to Anthurium and the other fossil taxa was pl aced within the extant genus Montrichardia The fossil leaves provide new inform ation about patterns of angiosperm diversification in the Neotropics.

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10 CHAPTER 1 MONOGRAPH OF THE CERREJ"N FLORA Abstract Little is known about the early history of Ne otropical rainforests, although this vegetation types are extremely diverse, they have a di stinctive spectrum of leaf size and shape, a characteristic floral composition at the family level, no fossil flor as with these unique features have been reported from low latitudes yet. Here I compile a monograph on a middle to late Paleocene (58-60 Ma) flora from the Cerrejn Form ation of northeastern Colombia that is the oldest record of Neotr opical rainforest. Fifty seven fossil mo rphospecies were described; they include 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. The Cerrejn assemblage has a high proportion of large, entire-mar gined, thin-textured leaves indi cating a moist tr opical climate with mean annual temperature ~24 C and rainfall 2.3 m/y. Family-level taxonomic composition is similar to that of extant Neotropical rainforests. Introduction Rainforests in tropical America have the highest plant diversity of any region in the world (Morley, 2000), and much of this plant diversity occurs in the floweri ng plants. However, the paleobotanical fossil record from the tropics have remained scarce and nearly unexplored, leaving the origin of the Neotr opical rainforest as one the biggest mysteries in plant evolution and paleoecology. An exhaustive review of the fossil record in norther n South America (Burnham and Graham, 1999; Dilcher, 2000; Burnham and J ohnson, 2004) suggests th at the Neotropical rainforests could have appeared during the early Cenozoic (~50-65 million years ago). This evidence comes mainly from th e fossil pollen record (Jaramillo et al., 2006). But this fragmentary fossil record still has not addres sed many questions: when did the Neotropical

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11 rainforest appear in South America? Under what climatic conditions did those ancient rainforests grow? Did ancient rainforests have the leaf morphologic characters seen in modern rainforests? What were the first families that inhabited the Neotropical rainforest? Did the families that are dominant in the Neotropical rain forest today originate in Sout h America or did they immigrate from other latitudes? Here, I report on a middle-late Paleocen e flora from Northern Colombia: Cerrejn paleoflora that is the oldest reco rd of a Neotropical rainforest. A complete description of the leaf fossils, a paleoclimatic reconstruction based on the fossil leaf morphology and their implications on the Paleocene climate, and finally a review of the floristic composition are presented in this monograph. Materials and Methods Fossil plants from the Cerrejn mine were first described by Doubinger and Pons in 1970 which consisted of a report on the remains of fo ssil leaf cuticle. After this report there was no additional research on the Cerrejn fossil plants until they were collected during 2003-2007. The Cerrejn Formation is exposed in the Cerrejn mine, located in Northern Colombia (Guajira Peninsula, Fig. 1-1). The Paleogene age of the Cerrejn flora is based upon the palynological assemblages, which demonstrates th at the pollen flora belong to the Foveotricolpites perforatus zone and to zone Cu-02 (Bayona et al., 2004; Ja ramillo et al., 2007) based on the presence of Foveotricolpites perforatus and Bombacacidites annae throughout the interval. These zones are considered to be middle to late Paleocene, base d on their co-occurrence with marine microfossils in shallow marine rocks and st able carbon isotop e stratigraphy. The ~800-m-thick stratigraphic sequence of Cerrejn Formation is composed of thick coals, fluvially deposited sandstones, and lacustri ne siltstones that were deposited in a rapidly subsiding basin (Bayona et al., 2004). Most of the Cerrejn Formation consists of a depositional

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12 cycle of three parts (Bayona et al., 2004). The lowest part is fossiliferous black shale and laminated black mudstone with thin lenticular laminae of sandstone (interpreted as anoxic lagoonal, or flooded coastal-plain environments). The middle part of the type section includes slightly bioturbated mudstones and sandstones with flaser and heterolithic lamination and dispersed plant remains, interprete d as deposits of subtidal and tid al flats. The upper part of the type section is dominated by fine-grained, ma ssive to lenticularly-laminated, bioturbated mudstones and siltstones with a bundant plant remains, which ar e cut by thick to very thick massive to cross-bedded sandst ones (interpreted as coastal pl ains crossed by channels). The thickest and extensive coal be ds in the Cerrejn Formation o ccur in the middle part of the section, these coal deposits have low-ash and low-sulfur contents. The megafloral localities occur from 100-750 meters above the base of the Cerrejn Formation (Fig. 1-2). In total, I recovered 1347 megafossil specimens. The collections of fossil plants were made from six lens-shaped, siltston e/sandstone bodies with inclined heterolithic strata that suggest deposition in low-energy channels (Fig. 1-2; localities 0410, 0317, 0315, 0322, 0319, and 0323). Two other localities preserve l eaves in thin but tabular beds of flatlaminated siltstone that indicat e deposition in small lakes (Fig 1-2; localities 0318 and 0324). The classification and descri ption of fossil leaf mor photypes is based upon leaf architectural features such as the characters presented in th e Leaf Architecture Working Group Manual (LAWG, 1999). The botanical affinities s uggested for some of the leaf morphotypes were determined after extensive comparisons with cleared leaves of living plants in the National Cleared Leaf Collection held in the Department of Paleobiol ogy, National Museum of Natural History, Smithsonian Institution, the modern refe rence Leaf Collection of Florida Museum of Natural History, and the Herbarium of the Smithsonian Tropical Research Institute.

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13 Figure 1-1. Map and location of Cerrejn mine and Cerrejn flora localities (stars).

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14 Figure 1-2. Stratigraphic column of the Cerr ejn Formation showing lithology and stratigraphic positions of the plant megafossil localities. Modified from Bayona et al (2004).

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15 Fossil Leaf Descriptions Ferns CJ42 (Plate 1) Diagnosis Blade elliptic and symmetrical, emar ginated apex. Tertiaries forming quadrangular areolation. Entire margin. Description The only specimen found reaches ~2 cm of length and width of 1.7 cm; the specimen is missing part of the leaf base, a midvein is visible with a curved course that ends in an emarginated apex. Tertiary veins forming a very tiny quadrangular areolation. The margin is entire. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, two specimens. Systematic affinity suggested The leaf closely resembles th e pantropical, floating-aquatic fern Salvinia (Salviniaceae) CJ57 (Plate 2) Diagnosis Pinnate leaf, secondary veins are cr aspedodromous, decurrent on midvein, and closely spaced. Irregular polygonal re ticulate tertiaries. Serrate marg in, cc/st and cc/cv teeth with long basal sides. Description The specimen is not complete but r eaches ~16 cm in length and a maximum width ~6 cm. Leaf texture coriaceous. Lami nar shape oblong. Secondary veins are numerous, very closely spaced (0.5 mm), decurrent on mi dvein, and craspedodromous. The margin is serrate, teeth have long basal sides and the mor phology varies between apical sides concave and basal sides straight and convex. Tertiary vein s are the highest orde r veins present on the

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16 specimen, these veins are irregu lar polygonal reticulate aerolat ions with elongate and rounded shapes. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0322, Pit La Puente, locality placed between coal beds 103 and 105, middle to late Paleocene, coordinates 11.9 N, 72.3 W, one specimen. Systematic affinity suggested The specimen closely resembles members of the family Stenochlaenaceae, common climbing ferns in Old World tropical swamps. CJ61 (Plate 3) Diagnosis Lobate leaf with many veins of equal gauge branching toward apex. Description The only specimen found has a maximum length of 2 cm and a maximum width of 3 cm (measured apically); leaf shape re sembles a palmate leaf with four asymmetrical lobes preserved, their basal sinus de parts either very to close to the base of the blade or from more apical sides; the lobes miss prominent midv eins. Many thin veins depart from the base and branch in random patterns toward apex. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0322, Pit La Puente, locality placed between coal beds 100 and 115, middle to late Paleocene, one specimen. Systematic affinity suggested The specimen closely resembles members of the cosmopolitan genus Lygodium CJ66 (Plate 4) Diagnosis Pinnate leaf. Secondaries anastomose near the midvein to form relatively short cells, and then these cells become more elongated toward the margin of the leaf. Secondaries are weekly decurrent on mid-rib.

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17 Description The specimen is about 8 cm in length and the maximum width is about 4 cm; laminar shape elliptic; margin entire; primary vein thick; secondary veins thin, weekly decurrent on midvein and anastomose near midvein to form relatively short cells, then the cells become more elongated toward the margin of the leaf. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0322, Pit La Puente, locality placed between coal beds 103 and 105, middle to late Paleocene, c oordinates 11.9 N, 72.3 W, two specimens. Systematic affinity suggested The leaf closely looks like Acrostichum a pantropical fern common in freshwater and brackish mangrove swamps today. Cycads CJ46 (Plate 5) Diagnosis Rachis stout; leaflets no n-articulate, decurrent at th e base, opposite; a midrib is absent from the leaflets. Sec ondary veins are thin and paralle l each other. Entire margined. Description Fragment of a leaves that reach ~15 in length and ~18 in width. Rachis is about 3 cm width. The maximum number of leafle ts preserved on the same specimen are sixteen; leaflets more than 22 mm wide, opposite, non-artic ulate, decurrent at the base and lack a midvein. Secondary veins are numerous, closel y spaced, and parallel each other. Entire margined. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0319, P it Tabaco High Dip, locality placed below coal bed 110, middle to late Paleocene, coor dinates 11.66 N, 73.31W, three specimens. Systematic affinity suggested The cycad is similar to extant Dioon Conifers CJ60 (Plate 6)

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18 Diagnosis Shoot of conifer with scale leaves. Description Length ~11 cm and maximum width re aches ~0.8 cm; scales leaves are crowded and overlapped through the axes; scales arranged in a sp iraled course. Maximum length of the scales reach ~5-7 mm and maximum width measured ~4-5 mm. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, c oordinates 11.13N, 72.57W, three specimens. Systematic affinity suggested The shoot closely resembles members of living Taxodiaceae and Araucariaceae. Dicots CJ1 (Plate 7) Diagnosis Petiole transversely striated, short an d pulvinate. Secondary angle to primary increases basally, and secondary ve ins fork at low angles near ma rgin. Tertiaries closely spaced and mostly opposite percurrent. Description Laminar size range from notophyll to macrophyll; leaves elliptic, oblong and obovate; Petiole transversely striated, short and pulvinate; base shapes from convex, concave and weekly asymmetrical; apex rounded and acuminate ; margin entire; unlo bed; leaves pinnate; secondary veins brochidodromous, between 19-22 pa ir of 2’s; Secondary vein angle to primary increases basally, and secondary veins fork at lo w angles near margin; tertiary veins mostly opposite percurrent with courses straight and convex and their angles respect to the primary range from obtuse to perpendicula r; fourth order veins mostly a lternate percurrent, higher order veins regular polygona l reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locali ties 0315; 0317, 0318, 0319, 0322, 0323, and 0410; Pits

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19 Tabaco 1, Tabaco High Dip, La Puente, Expanded West, localities placed between coal beds 40 and 161, middle to late Paleocene, 173 specimens. Systematic affinity suggested Based on the leaf morphology a nd the presence of a petiole transversely striated, short and pulvinate the morphotype closely resembles the family Leguminosae. CJ2 (Plate 8) Diagnosis Strongly asymmetrical blade. Second ary veins are weak brochidodromous at the base and brochidodromous apically. Secondary vein angle on primary increases apically to almost 90. Falcate apex. Thick margin. Description Laminar size range from notophyll to mesophyll, laminar shape elliptic; strongly asymmetrical blades; base shape complex; apex falcate; margin entire; leaves pinnate; secondary veins are weak brochidodromous at the base and brochidodromous apically, between 23 and 18 pairs; the angle of the 2’s on primary in crease apically to almost 90; inter-secondary veins are week; tertiary veins ar e alternate percurrent; the angle of the 3’s on primary veins range from perpendicular to acute; a th ick margin is visible easily. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317 and 0318; Pits Tabaco 1 and Tabaco High Dip; localities placed between coal beds 100 and 161, middle to late Paleocene, coordinates 11.13N, 72.57W and 11.12 N, 73.55 W, 8 specimens. Systematic affinity suggested The fossil leaves seem similar to living species within the family Meliaceae. CJ4 (Plate 9)

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20 Diagnosis Decurrent base. Leaf shape narrow er than CJ32. Semicraspedodromous secondaries, they become acute basally more th an in CJ32. Tertiary veins opposite percurrent and about 90 degrees to primary, epimedial tert iaries forming intersec ondaries. Fourth order veins mostly irregular polygonal and alternate percurrent. Teeth closely spaced and have long basal sides. Description Long petioles; laminar shapes from microphyll to mesophyll; laminar shape elliptic to oblong; base shape strongly decurren t; margin entire; leaves pinnate; secondary veins are semicraspedodromous, between 23 to 24 pairs of 2s; the secondaries become acute basally, secondary vein spacing decreases toward base; intersecondary veins are weak on the specimens; tertiary veins are opposite percurrent with sinuous to straight courses, the angle of the 3’s on primary is mainly perpendicula r to it; higher order veins range from irregular to regular polygonal reticulate; areolation is well developed; margin type cr enate, teeth are closely spaced and have short convex apical side s and long convex basal sides; leaf texture is chartaceous. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317 and 0318; Pits Tabaco 1 and Tabaco High Dip; localities placed between coal beds 100 and 161, middle to late Paleocene, coordinates 11.13N, 72.57W and 11.12 N, 73.55 W, 24 specimens. Systematic affinity suggested No extant affinity identified. CJ5 (Plate 10) Diagnosis Suprabasal or basal acrodromous, two lateral primaries running in convergent arches toward the leaf apex. Eucamptodromous 2 s. One secondary vein running just inside the margin. Two specimens covered with small laminar resin dots.

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21 Description Laminar size range from microphyll to macrophyll; laminar shape elliptic; base shape from concavo-convex to convex; apex acuminate; margin entire; primary veins are either suprabasal or basal acrodromous (t hree veins only); costal secondary veins are eucamptodromous, at about 10 pairs of 2’s, one s econdary vein runs from the base parallel and close to the margin as an intramarginal veins wh ich collects the costal 2’s, the spacing of the costal secondaries decreases towa rd base; tertiary veins are mixe d opposite/alternate percurrent, the course of the opposite ones range from convex to straight and their angles on primaries are obtuse; the variability of the angl e of 3’s increase basally (underco stals) and increase exmedially (epimedial); fourth order veins are alternate pe rcurrent; a fimbrial vein is present along the margin, only two specimens are covere d with small laminar resin dots. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317, 0319, and 0410; Pits Tabaco 1, Tabaco High Dip, and Expanded West, localities placed betw een coal beds 40 and 107, middle to late Paleocene, 22 specimens. Systematic affinity suggested Leaves with suprabasal or basal acrodromous venation and covered with resin glands are very common in the family Lauraceae. The fossil leaves closely resemble this family. CJ6 (Plate 11) Diagnosis Deeply cordate or truncate base. Brochi dodromous 2s that loop just inside the margin both costal and minor secondaries. Opposite percurrent tertiaries concentric. Fifth and sixth order veins are regular pol ygonal reticulate. Tiny and squarish areoles. Thick fimbrial vein. Description Petiole with wiry thickening; l eaves size range from microphyll to macrophyll; laminar shape ovate; base shape are d eeply cordate, cordate and truncate; apex

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22 acuminate, margin entire; primary veins are ba sal actinodromous with 5 to 7 veins; agrophic veins compound; intersecondary veins are absent; te rtiary veins mostly opposite percurrent with straight to strongly convex course s, the angle of the 3’s on primar ies is mostly perpendicular to it, the tertiary veins angle variability decreases exmedially and basally; fourth order veins are strongly alternate percurrent; highe r order veins (5th and 6th orde rs) regular pol ygonal reticulate; areolation is well developed, tiny, and squarish (3-4 sided); a th ick fimbrial veins is present along the margin. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localitie s 0315; 0317, 0318, 0319, 0322, and 0323; Pits Tabaco 1, Tabaco High Dip, and La Puente, localities pl aced between coal beds 100 and 161, middle to late Paleocene, 68 specimens. Systematic affinity suggested Based on the actinodromous leaf venation, blade shape, higher order veins and the presence of a thick fi mbrial vein the morphot ype closely resembles the family Menispermaceae. CJ8 (Plate 12) Diagnosis Primary vein and petiole very thick. Secondaries b ecome more obtuse basally (almost 90), and 3-4 pairs crowded, secondary vein spacing decreases towa rd base but apically increases. Tertiaries thin in mid-course. Description Petiole very thick, sometimes this reaches at about 2 cm in width; laminar size range from notophyll to macrophyll; laminar sh ape between elliptic but mostly obovate leaves; base shape concave; apex acuminate to rounded; margin entire; leaves pinnate, the midvein is stout; secondary veins are eucam ptodromous; secondary vein spacing crowded basally or decreases toward base but apically increases; secondary veins also become more

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23 obtuse basally (almost 90); tertiary veins mos tly opposite percurrent w ith straight to sinuous courses, their angles on primaries are perpendicular to acute; the tertiary vein angle variability on the rest of the blade increases exmedially and basally; tertiary veins are as well thin compared to secondaries in mid-course; higher order veins are regular polygonal retic ulate and not strongly visible. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localitie s 0315; 0317, 0318, 0319, and 0323; Pits Tabaco 1, Tabaco High Dip, and La Puente, localities place d between coal beds 100 and 161, middle to late Paleocene, 36 specimens.. Systematic affinity suggested Leaves with these characters occur in several families such as Sapotaceae, Bombacaceae, and Annonaceae. CJ10 (Plate 13) Diagnosis Primary vein is very thick at the base and thins distally and develops a zig-zag course with the primary being deflected at each secondary juncture. Brochidodromous 2s forming prominent arches almost festooned br ochidodromous. Poorly organized tertiaries. Fimbrial vein. Description Petiole is very long, reaches at about 5 cm in lenght; laminar size mesophyll; laminar shape elliptic; base shape varies between concave-convex and convex; apex shape either retuse or convex; margin entire; leaves pinnate, midvein very thick at the base and thins distally and develops a zig-zag course with the primar y being deflected at each secondary juncture; secondary veins are brochidodromous, these vein s form prominent arches almost like festooned brochidodromous venations, 6 to 7 pairs of 2’s, sometimes two or three of these veins depart from the base of the leaf; secondary vein spaci ng decreases toward base and the angle of these

PAGE 24

24 veins smoothly increases toward base; tertiary veins are mixed opposite/alte rnate percurrent but poorly organized; tertiary vein angle variability d ecreases apically and more acute relative to the midvein; higher order veins are irregular a nd regular polygonal reticu late; areolation well developed (3-4 sided); a fimbrial ve in runs parallel to margin. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317, 0318, and 0323; Pits Tabaco 1 and Tabaco High Dip, localities placed between coal beds 100 a nd 161, middle to late Paleocene, 13 specimens. Systematic affinity suggested No extant affinity identified. CJ11 (Plate 14) Diagnosis Distinctly lobate leaf thick primary veins. Brochidodromous 2s closely spaced, and forming loops at the margin. Description Laminar size occurs from mesophyll to macrophyll; laminar shape elliptic; apex angle odd lobed acute; apex acuminate; margin entire; primary veins are basal actinodromous, at least 7 veins are recognized in one of the specimens; at least three lobes are preserved; agrophic veins are simple; costal secondary vein spacing decreases toward base; tertiary veins are mixed oppos ite/alternate percurrent. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, c oordinates 11.13N, 72.57W, seven specimens. Systematic affinity suggested Three-lobate leaves with en tire margin are common in the order Malvales. CJ12 (Plate 15)

PAGE 25

25 Diagnosis Tertiaries are alternate percurrent and form composite intersecondaries, Tertiaries admedially ramified on the midvein. Specimens covered with irregular black dots. Description Petiole with base swollen; lamina r size mesophyll; laminar shape elliptic; base shape convex; margin entire; leaves pi nnate; secondary veins are eucamptodromous; secondary vein spacing is weekly irregular through the blade, 2’s vein angle on primary smoothly increases towards apex; tertiary veins are strongly a lternate percurrent and form composite intersecondaries, then th e tertiaries ramify admedially on the midvein, the angle of the 3’s respect to the midvein varies between perpen dicular and acute; higher order veins are regular polygonal reticulate; the specimens are covered with irregular black dots. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315 a nd 0323; Pit Tabaco 1 and Tabaco High Dip, localities placed between coal beds 100 and 120, middle to late Paleocene, seven specimens. Systematic affinity suggested Some characters such as t ype of tertiary veins and its courses are similar to those seen in the families Moraceae and Ulmaceae, but more characters are necessary to prove this affinity. CJ13 (Plate 16) Diagnosis Petiole with base swollen and short. 13-14 pairs of weakly brochidodromous 2s, secondaries diverge from one another exmedi ally. Tertiaries are oppos ite percurrent, mostly convex or sinuous and wider spaced than in CJ1. Description Petiole short with a swollen base; la minar size varies between mesophyll and macrophyll; laminar and base shape constantly elli ptic and convex respectively; apex shape from acuminate to convex; margin entire; leaves pi nnate; secondary veins are weak brochidodromous looping very close to margin, 10 to 13 pairs of secondaries that diverge from one another

PAGE 26

26 exmedially with a course almost sinuous; secondary vein spacing is uniform, and secondary vein angle smoothly increases towards base; tertiary veins are opposite percurrent with strong convexsinuous course, the angles of the 3’s on primaries go from obtuse to perpendi cular, tertiary vein angle variability increases exmedi ally; higher order veins regular polygonal reticulate; areolation moderately developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315, 0317, 0323 and 0410; Pit Tabaco 1, Tabaco High Dip and Expanded West, localities placed be tween coal beds 40 and 120, middle to late Paleocene, 16 specimens. Systematic affinity suggested No extant affinity identified. CJ15 (Plate 17) Diagnosis Similar to CJ8 but secondaries are no t perpendicular near base of primary, secondary veins spacing increases toward (2/3) of blade and then decreases. Fourth order veins are regular polygonal reticulate. Description Petiole thick and short; laminar si ze from mesophyll to megaphyll; laminar shape obovate to elliptic; base shape cuneate; apex shape is either rounded or acuminate; margin entire; leaves pinnate; secondary veins are eucamptodromous, be tween 20 to 25 pairs of 2’s; secondary vein spacing increases toward (2/3) of blade and then decreases, and secondary vein angle smoothly decreases towards base; tertiaries mostly opposite percurre nt with straight to sinuous courses and their angles on primary range from perpendicular to obtuse; tertiary vein angle variability decreases exmedially; highe r order veins regular polygonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localitie s 0315; 0317, 0318, 0323, and 0410; Pits Tabaco 1,

PAGE 27

27 Tabaco High Dip, Expanded West, localities placed between coal beds 40 and 161, middle to late Paleocene, 28 specimens. Systematic affinity suggested Leaves with these characters occur in several families such as Anacardiaceae, Apocynaceae, and Sapotaceae. CJ16 (Plate 18) Diagnosis Pinnate with cordate base. Secondaries (3-4 pairs) crowded at the base, secondary angle at the base is more obtuse th an CJ28. More closely sp aced opposite percurrent tertiaries than CJ28. Description Petiole long, reaches at about 6 cm; laminar size mesophyll; laminar shape constantly ovate; base shape cordate; apex sh ape acuminate; margin entire; leaves pinnate; secondary veins are eucamptodromous (13-15 pairs of 2’s) having five to six of those veins crowded and departing from the base; secondary vein spacing decreases toward base, and their angles abruptly increase toward base; tertiary veins are opposite percurrent, closely spaced each other and having straight to sinuous courses, their angles on primary are obtuse; the tertiary vein angle variability decreases exmedially and apically; higher order veins probably regular polygonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0318 a nd 0319, Pit Tabaco High Dip, localities placed between coal beds 105 and 161, middle to late Paleocene, five specimens. Systematic affinity suggested No extant affinity identified. CJ17 (Plate 19) Diagnosis Pinnate with slightly cordate base with 5 basal veins. Compound agrophic veins. Tertiaries mixed opp/alt percurre nt. Areolation very small and squarish.

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28 Description Long petiole, reaches at about 5.5 cm ; laminar size and shape mesophyll and elliptic respectively; base shape slightly cordat e; apex shape acuminate; margin entire; leaves pinnate; secondary veins weak brochidodromous and looping ve ry close to margin, five secondaries are crowded at the ba se; secondary vein spacing appa rently decreases toward base, and the angle seems to increase toward base; agrophic veins are compound; tertiary veins are mixed opposite/alternate percurrent with courses sinuous to straight and their angles on primary vein are perpendicular to obtuse; fourth order veins are alternate percurrent; higher order veins regular polygonal reticulate; areolation well develope d, small, and squarish. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315 a nd 0318; Pit Tabaco 1 and Tabaco High Dip, localities placed between coal beds 100 and 61, middle to late Paleocene, three specimens. Systematic affinity suggested Pinnate leaves with cordate bases common in the order of Malvales. CJ18 (Plate 20) Diagnosis Like CJ8 or CJ15 but about half as many secondarie s ( 8-10 pairs), widely spaced secondaries, and the angle of 2s smoothly increasing toward base, one pair acute basal secondaries. Percur rent tertiaries. Description Petiole long (5.3 cm) and thin (0.5 cm); laminar size and shape mesophyll and elliptic respectively; base shape concaveconvex and decurrent; margin entire; leaves pinnate; secondary veins are eu camptodromous, at least eleven pair of 2’s are preserved; secondary vein spacing is widely spaced, a nd the secondary vein angle smoothly increases toward base; tertiary veins are opposite percurrent with straight to sinuous courses; 4th and 5th order veins are alternate percurrent and opposite percurrent respectively.

PAGE 29

29 Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, c oordinates 11.13N, 72.57W, seven specimens. Systematic affinity suggested No extant affinity identified. CJ19 (Plate 21) Diagnosis Pulvinate petiole. Very large number of closely spaced secondaries that are decurrent on the primary and arise from it at ab out 90 degrees. Tertiaries are random reticulate and also decurrent on primary. Description Petiole pulvinate; laminar size from microphyll to mesophyll; laminar shape elliptic; base shape convex and rounded; apex acuminate; margin entire; leaves pinnate; secondary veins are eucamptodromous, numerous, closely spaced, and decurrent on the primary and they arise from it at about 90 degrees; seconda ry vein angle increases toward base, and their angles smoothly increase toward base; inters econdary veins are common, usually one to two pairs per secondaries; tertiary ve ins are random reticulate and as well decurrent on primary; their angles on primary range from perpendicular to acute; tertiary vein angle variability is inconsistent’ 4th and 5th order veins ar e irregular and regul ar polygonal reticulate correspondingly; areolation is we ll to moderately developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localitie s 0315; 0317, 0318, 0319, 0323 and 0324; Pits Tabaco 1, Tabaco High Dip, La Puente, localities placed between coal beds 100 and 161, middle to late Paleocene, 22 specimens. Systematic affinity suggested Based on the secondary and tertiary venation and the presence of a pulvinate petio le the morphotype closely rese mbles the family Leguminosae.

PAGE 30

30 CJ20 (Plate 22) Diagnosis Like CJ22 in major venation, but fourth and fifth order veins less distinct than in CJ22. Secondary spacing mostly irregular. Tertia ries with minutely irre gular course, leaf rank 2r. Description Laminar size from mesophyll to macrophyll; laminar shape elliptic; base and apex shape convex; margin entire; leaves pinna te; secondary veins are eucamptodromous, 10-15 pairs of 2’s; secondary veins spacing mostly irre gular, and their angles smoothly increase toward base; intersecondary veins occur but they are weak; tertiary veins mostly mixed opposite/alternate percurrent with minutely irregular course, and their angles re spect of the primary oscillate between perpe ndicular to acute; tertiary vein angle variability increases exmedially and apically; 4th and 5th order vein s are irregular and regul ar polygonal reticulate; areolation well developed, leaf rank 2r. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315; 0318, 0319; Pits Tabaco 1and Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, nine specimens. Systematic affinity suggested No extant affinity identified. CJ21 (Plate 23) Diagnosis Decurrent base. Very closely spaced (~1 mm) opposite percurrent tertiaries, course mostly sinuous, angle to primary str ongly perpendicular and increases exmedially. Description Laminar size and shape mesophyll and elli ptic correspondingly; base angle at almost 90 degrees and base shape very decurrent; apex acuminate; margin entire; leaves pinnate; secondary veins are eucamptodromous, at least 12 pairs of 2’s are preserved on the specimens; secondary vein angle increases from base until 2/ 3 length of leaf and their angles are uniform;

PAGE 31

31 tertiary veins are consta ntly opposite percurrent and very cl osely spaced (~1mm), with course sinuous and the angle of the 3’s on primary vein is strongly perpendicular; tertiary vein angle variability increases exmediall y, 4th vein order are irregular polygonal reticulate; areolation is moderately developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315; 0318, 0323; Pits Tabaco 1and Tabaco High Dip, localities placed betw een coal beds 100 and 161, middle to late Paleocene, 10 specimens. Systematic affinity suggested Some species in Chrysobalanaceae and Annonaceae have similar characters as the fossil leaf. CJ22 (Plate 24) Diagnosis Between 7 and 14 pairs of secondari es, crowded basal secondaries are decurrent and more acute than apical secondaries. Fourth and fifth order veins are alternate percurrent and regular polygona l reticulate unlike CJ20. Most sp ecimens covered with laminar resin glands (<0.1mm). Leaf rank 3r. Description Laminar size vary from notophyll to macrophyll leaves; laminar shape principally elliptic; base shape from convex, concave to weakly decurrent; apex shape acuminate; margin entire; leaves pinnate; s econdary veins are eucamptodromous, between 7 to 14 pairs of 2’s were counted on the specimens, ge nerally a few pair of 2’s are crowded at the base and they are decurrent on primary veins and more acute than apical secondaries; one pair acute basal secondaries is common within th e morphotype; intersecondaries are strong and common; tertiary veins are mixed alternate/oppos ite percurrent, the opposite percurrents having courses straight to sinuous and their angles are mostly perpendi cular on primary; tertiary vein angle variability is perpendicu lar for both on midvein and ending; 4th order veins are alternate

PAGE 32

32 percurrent; 5th order veins are regular polygona l reticulate; most sp ecimens covered with laminar resin glands (<0.1mm); leaf rank 3r. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localitie s 0315; 0317, 0318, 0319, and 0323; Pits Tabaco 1, Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 72 specimens. Systematic affinity suggested The fossil leaf closely rese mbles the typical characters seen in Lauraceae, pinnate leaves, eucamptodromous venation and resin glands. CJ24 (Plate 25) Diagnosis Long petiole. One pair of strictly agro phic lateral primaries, sometimes it may be difficult to distinguish from costal seconda ries. Brochidodromous looping arches close to margin. Teeth are low closely spaced cr enations mostly fed by agrophic veins. Description Maximum length of the petiole measured reached at about 9 cm; laminar size from notophyll to macrophyll; laminar shape ellip tic; base shape oscillates from truncate to cordate; apex shape acuminate; primary veins basal actinodromous; secondary veins are brochidodromous, 12-14 pairs were counted on the specimens; secondary vein spacing increases toward base and their angles remain uniform; ag rophic veins are simple; intersecondary veins are weak; tertiary veins are opposite pe rcurrent with courses straight to sinuous; tertiary vein angle variability decreases exmedially; 4th order veins are a lternate percurrent; margin toothed, teeth are closely spaced with crenations mostly fed by agrophic veins; the spacing is irregular and the sinus of the teeth are rounded. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317, 0318, and 0323; Pits Tabaco 1, and Tabaco

PAGE 33

33 High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 30 specimens. Systematic affinity suggested The fossil leaf closely rese mbles the typical characters seen in many species of Euphorbiaceae, leaves with primary veins basal actinodromous, long petioles, and margin toothed. CJ25 (Plate 26) Diagnosis Mucronate or rounded apex, base sh ape highly variable (concavo-convex, cordate, truncate), sometimes almost are lobate, 3rd and 4th order veins are mostly opposite percurrent. Teeth distantly spa ced, and formed by simple extension of the vein beyond the margin of the lamina, teeth sometimes retuse. Description Petiole has a base swollen, and it is long and thick; laminar size from microphyll to macrophyll; laminar shape varies from e lliptic to ovate; base shape highly variable (concavo-convex, cordate, truncate); apex shap e highly variable acuminate, mucronate and rounded; primary veins are basal actinodromous, be tween 5 to 7 veins from the base; agrophic veins compound; secondary veins ar e craspedodromous; and the costal secondaries have a vein spacing and angle uniform; tertiary veins mainly opposite percurrent, their courses vary from straight to sinuous, angles of tert iary veins on primaries are perpendi cular to obtuse; tertiary vein angle variability increases exmedially and basa lly; 4th order veins mostly opposite percurrent, 5th order veins are regular polygona l reticulate; a fimbrial vein runs parallel to margin. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localitie s 0315; 0317, 0318, 0319, 0323 and 0324; Pits Tabaco 1, Tabaco High Dip, and la Puente; localities pl aced between coal beds 100 and 161, middle to late Paleocene, 67 specimens.

PAGE 34

34 Systematic affinity suggested The fossil leaf closely rese mbles the typical characters seen in the group of Malvales. Leaves actinodr omous, with long petioles, and margin toothed. CJ26 (Plate 27) Diagnosis Almost lobate, primaries branching near margin. Fourth and fifth order veins orthogonal reticulate. Teeth closely spaced and convex/convex and almost symmetrical. Description Petiole long and having a swolle n base; laminar size from notophyll to macrophyll; laminar shape constantly ovate; base shape cordate; apex shape convex; primary veins are basal actinodromous, between 8 to 9 veins depart from the base; agrophic veins compound; secondary veins are craspedodromous and their angles are irre gular; tertiary veins mainly opposite percurrent with straight to sinuou s courses, their angles on primaries vary from perpendicular to obtuse; tertia ry vein angle variability incr eases exmedially and basally; distinctly 4th and 5th order vein s are orthogonal reticulate; fimbri al vein parallel to margin; margin type crenate; the teet h are closely spaced and convex/ convex and almost symmetrical, 6 to 7 teeth per cm, teeth sinuous rounded with simple apex. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, c oordinates 11.13N, 72.57W, six specimens. Systematic affinity suggested The fossil leaf closely rese mbles the typical characters seen in the group of Malvales. Leaves actinodr omous, with long petioles, and margin toothed. CJ27 (Plate 28) Diagnosis Naked basal lateral pimaries, 7-8 pa irs of semicraspedodromous secondaries, teeth closely spaced and convex/conv ex with elongate basal sides.

PAGE 35

35 Description Laminar size oscillates from notophyll to macrophyll; laminar shape ovate; base shape cordate; apex shape acuminate; prim ary veins are basal actinodromous with 5 basal veins; agrophic veins are compound; secondary veins are semicraspedodromous (7-8 pairs of 2’s); the costal secondary veins increases toward base; tertiary veins are opposite percurrent with a constant straight course and an angle acute; tert iary vein angle variability increases exmedially and basally; 4th vein order are orthogonal reticulate; 5th order vein regular polygonal reticulate, areolation is well developed, margin toothed, teeth closely spaced and convex/convex with elongate basal sides. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317 and 0319 Pit Tabaco 1 and Tabaco High Dip; locality placed between coal beds 100 and 115, middle to late Paleocene, six specimens. Systematic affinity suggested The fossil leaf closely rese mbles the typical characters seen in the group of Malvales. Leaves actinodr omous, with long petioles, and margin toothed. CJ28 (Plate 29) Diagnosis Pinnate with cordate base. Secondary vein angle at the base is more acute than in CJ16. Tertiary veins are opposit e percurent and are less closely spaced than in CJ16. Fourth order veins are mostly mixe d opp/alter and very thin. Description Laminar size mesophyll; laminar shape elliptic; base shape varies from rounded to slightly cordate; margin entire; leav es pinnate with a very thick midvein; secondary veins are eucamptodromous and five of these veins depart from the base; secondary vein spacing decreases toward base and thei r angles smoothly increases towa rd base; tertiary veins are opposite percurrent with straight courses and th eir angles on primary mainly perpendicular; tertiary vein angle variability are perpendicular for both vein s on the midvein and margin; 4th

PAGE 36

36 vein category mostly opposite/alter nate percurrent, veins very thin ; 5th order type from regular polygonal reticulate to dichotomizing. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, c oordinates 11.13N, 72.57W, seven specimens. Systematic affinity suggested No extant affinity identified. CJ30 (Plate 30) Diagnosis Asymmetrical blade. Secondary vein spacing and angles are irregular; however the angle abruptly increases towa rd base. Tertiaries opposite perc urrent, very sinuous and widely spaced, epimedial tertiaries form intersecondaries. Description Laminar size mesophyll; blade asymmetrical; laminar shape obovate; base is incomplete; margin entire; leaf pinnate; secondary veins are eucamptodromous; secondary vein spacing irregular; secondary vein angle abruptly increases toward base; intersecondary veins are strong and common; tertiary veins are opposite perc urrent with very sinuous course and widely spaced, their angles on primary are mostly perpendicu lar; tertiary vein angl e variability increases exmedially; 4th order vein are mainly opposite percurrent; 5th orde r veins are irregular polygonal reticulate, areolati on moderately developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen. Systematic affinity suggested No extant affinity identified. CJ32 (Plate 31)

PAGE 37

37 Diagnosis Shape wider than CJ4. Semicraspedodromous 2s decurrent on primary. Tertiaries opposite percurrent but do not form in tersecondaries, tertiaries more closely spaced and obtuse than CJ4. Fourth order veins mostly opposite percurrent. Fimbrial vein. Teeth similar to CJ4. Description Long petioles, reaches up to 10 cm ; laminar size from microphyll to macrophyll; laminar shape elliptic; base shape st rongly decurrent; apex shape acuminate; laves pinnate; secondary veins are se micraspedodromous, at least 15 pa irs of 2’s; secondary vein spacing uniform and secondary vein angle smoothl y increases toward base; tertiary veins are opposite percurrent with courses st raight to sinuous, their angles on primary are mostly obtuse; tertiary veins angle vari ability decreases exmedially; distinctly opposite percurrent; 5th order veins are regular polygonal retic ulate; areolation well developed; fimbrial veins parallel to margin; margin toothed, teeth similar to CJ4, but spacing is irregular or closely spaced. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315, 0317, 0318, and 0319; Pits Tabaco 1, and Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 34 specimens. Systematic affinity suggested Some species in Violaceae has the characters seen on the morphotype. CJ33 (Plate 32) Diagnosis Two pairs of long acrodromous secondari es in basal half of leaf, no apical secondaries (hemieucamptodromous).

PAGE 38

38 Description Laminar size microphyll; laminar shape elliptic; apex shape rounded; margin entire; leaf pinnate; s econdary veins are suprabasal acrodr omous with no apical secondaries (hemieucamptodromous); secondary vein angle is low relative to primary. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen. Systematic affinity suggested No extant affinity identified. CJ34 (Plate 33) Diagnosis Craspedodromous secondaries at almost 90 degrees to primary and spacing decreasing toward base. Perpendicular and thin te rtiaries at the midvein. Teeth have long basal sides. Description Laminar size from notophyll to mes ophyll; laminar shape from elliptic to ovate; base shape concavo-convex to truncate; apex shape convex; leav es pinnate; secondary veins are craspedodromous, with no or two basal veins; secondary vein spacing decreases toward base; secondary vein angle smoothly increases to ward base; intersecondari es are weak; tertiary veins mainly opposite percurrent with sinuous to straight courses, thei r angles on primary are perpendicular; 4th order vein are alternate pe rcurrent; 5th order vein are regular polygonal reticulate, areolation well developed; margin t oothed, teeth have long basal sides, 1-2 teeth per cm and spacing regular. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, two specimens.

PAGE 39

39 Systematic affinity suggested Leaves pinnate, craspedod romous secondaries and tertiaries perpendicular and thin at the midvein are very common in the family Anacardiaceae. CJ36 (Plate 34) Diagnosis Actinodromous, three primary veins, lateral primaries forming loops. Weak brochidodromous 2s that loop just inside the margin both, costal and minor secondaries. Fourth order veins alternate percurrent a nd leaf texture different from CJ25. Description Long petiole; laminar size notophyll to mesophyll; laminar shape elliptic; base shape varies from truncate to convex; marg in entire; primary veins are basal actinodromous with three veins at the base; agrophic ve ins are simple; secondary veins are weak brochidodromous, the spacing of cost al 2’s increases toward base and their angles are acute; tertiary veins are opposit e percurrent having sinuous to c onvex courses, their angles are perpendicular on primaries; tertia ry vein angle variabil ity uniform; 4th orde r veins are alternate percurrent; 5th order veins are regular polygona l reticulate; areolati on well developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315, 0318, and 0319; Pits Tabaco 1, and Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, nine specimens. Systematic affinity suggested The fossil leaf closely rese mbles the typical characters seen in the group of Malvales, leaves actinodromous with long petioles. CJ37 (Plate 35) Diagnosis Few secondary veins (5 or 6 pairs), secondary spacing decreasing toward base but not crowded basally like CJ22. Secondary an gles more acute than CJ20 and CJ22. Lacking laminar resin dots.

PAGE 40

40 Description Petiole short; laminar size mesophyll; laminar shape elliptic; base shape decurrent; margin entire ; leaf pinnate; secondary veins are eucamptodromous, their spacing decreases toward base; secondary vein angle uniform; tertiary veins are mixed opposite/alternate percurrent, their angles on primary are perpendi cular; tertiary vein angle variability mostly uniform; 4th order vein alternat e percurrent; 5th order vein ar e regular polygonal reticulate; areolation well developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal beds 160 and 161, middle to late Paleocene, coordinates 11.12 N, 73.55 W, one specimen. Systematic affinity suggested No extant affinity identified. CJ38 (Plate 36) Diagnosis Pulvinate and striated petiole. Seco ndaries are perpendicular basally on midvein, intersecondary veins are common. Tertiaries alternate percurrent and wider spaced than CJ8. Tertiary angle increases basally on mi dvein, tertiaries d ecurrent on midvein. Description Petiole pulvinate and striated; la minar size from notophyll to mesophyll; laminar shape elliptic; base shape rounded; ma rgin entire; leaves pinnate; secondary veins eucamptodromous, sometimes having three basa l veins; secondary vein spacing decreases toward base, and their angles smoothly increases toward base; intersecondary veins are weak to strong; tertiary veins are alternat e percurrent with their angles perpendicular to base; tertiary veins angle variability increases basally on midve in; 4th order vein regular polygonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317 a nd 0318; Pits Tabaco 1 and Tabaco High Dip, localities placed betw een coal beds 100 and 161, middle to late Paleocene, 20 specimens.

PAGE 41

41 Systematic affinity suggested Based on the leaf morphology a nd the presence of a petiole transversely striated, short and pulvinate the morphotype closely resembles the family Leguminosae. CJ40 (Plate 37) Diagnosis Pinnate with cordate to rounded base. Costals 2s vein s appear near base, one pair of agrophic lateral seconda ries. Higher order veins are regul ar polygonal reticulate with areoles of rounded shape and tiny size. Description Long petiole (~6 cm); laminar size from microphyll to macrophyll; laminar shape is elliptic to ovate; base shape cordate to rounded; apex shape acuminate; margin entire; leaves pinnate; agrophic veins compound ;second ary veins are brochidodromous, 3 to 5 veins depart at the base; costal secondary vein spac ing decreases toward ba se; tertiary veins are opposite percurrent with sinuous to convex courses, their angles on primary are perpendicular to obtuse; tertiary vein angle va riability increases basally; 4th and 5th vein category regular polygonal reticulate; areolation we ll developed, tiny and rounded. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315 ; 0317, 0318; 0319 and 0323; Pits Tabaco 1 and Tabaco High Dip, localities placed between coal beds 100 and 161, middle to late Paleocene, 37 specimens. Systematic affinity suggested No extant affinity identified. CJ41 (Plate 38) Diagnosis Six or seven pairs of cost als 2s, crowded basal sec ondaries with relative long space between basal 2-3 and more apical s econdaries, secondaries curved up smoothly.

PAGE 42

42 Secondaries and tertiaries are decurrent on midvein, opposite percur rent tertiaries perpendicular to primary and the angle increases basally. Description Long petiole (~6 cm); laminar size from microphyll to macrophyll; laminar size from elliptic to ovate; base shape either co rdate or rounded; apex shape acuminate; margin entire; leaves pinnate; agrophic veins compound; secondary veins are brochidodromous with 3 to 5 veins at the base; costal seconda ry vein spacing decreases toward base; tertiary veins distinctly opposite percurrent, their courses vary from sinuous to convex; the angles of 3’s on primary are perpendicular to obtuse; tertiary vein angle variability increa ses basally; 4th and 5th regular polygonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0318 a nd 0324; Pit Tabaco High Dip; localities placed between coal beds 125 and 161, middle to late Paleocene, seven specimens. Systematic affinity suggested No extant affinity identified. CJ43 (Plate 39) Diagnosis Asymmetrical blade. Craspedodromous 2 s with agrophic branches on mid-leaf secondaries, 2s become more obtuse basa lly. Teeth broad conve x/convex and almost symmetrical, apical teeth are simple or foraminate. Fimbrial vein. Description Laminar size from mesophyll to macrophyl l; laminar shape elliptic to ovate; laminar symmetry strongly asymmetrical; base shape concave; apex shape concave; leaves pinnate; agrophic veins compound; secondary ve ins are craspedodromou s; secondary veins spacing increases toward base and their angles sm oothly increases toward base as well; tertiary veins mostly opposite percurrent with sinuous and straight courses, thei r angles on primary are only perpendicular; tertiary ve in angle variability increases exmedially; 4th order veins are

PAGE 43

43 mixed opposite/alternate percurrent; 5th order veins are regular polygonal reticulate; areolation moderately developed; fimbrial ve in parallel to margin; margin t oothed, two orders of teeth with irregular spacing, teeth are convex-convex and c oncave-convex, teeth apex are foraminate or simple. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0319, 0322; 0323, and 0324; Pits Tabaco High Dip and La Puente ; localities placed between coal beds 103 and 130, middle to late Paleocene, 62 specimens. Systematic affinity suggested No extant affinity identified. CJ48 (Plate 40) Diagnosis Actinodromous with closely spaced nippl e-shaped teeth with darkened apices Description Laminar size mesophyll; laminar shape ovate; base angle obtuse and base shape probably cordate; primary veins are basa l actinodromous; agrophic veins are simple; tertiary veins are opposite percu rrent with mainly sinuous course s; teeth are closely spaced nipple-shaped with darkened apices. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315 a nd 0319; Pits Tabaco 1 and Tabaco High Dip, localities placed between coal beds 100 and 107, middle to la te Paleocene, three specimens. Systematic affinity suggested The leaf and teeth morphol ogy may resemble species in the order Malvales. CJ50 (Plate 41) Diagnosis Pinnate and cordate base. Percurrent tertiaries, mostly convex and widely spaced with straight course on the midvein a nd sinuous on secondaries. F ourth order veins are

PAGE 44

44 alternate percurrent and thick veins which is diffe rent from CJ28. Areolation is not squarish as in CJ17. Description Laminar size and shape from mesophyll and elliptic respectively; base shape is closely cordate to cordate; margin entire; l eaves pinnate, sometimes having three basal veins; agrophic veins are compound; secondary veins are eucamptodromous; secondary vein spacing decreases toward base and their angles smoothly increases toward base; tertiary veins mainly opposite percurrent with courses straight to si nuous/convex; the angle of 3’s on primary is mainly perpendicular; tertiary vein angle vari ability increases exmedially; 4th vein type are alternate percurrent; 5th orde r veins are regular polygonal re ticulate; areolation is well developed. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0315 and 0322; Pits Tabaco 1 and La Puente, localities placed between coal beds 100 and 105, middle to late Paleocene, three specimens. Systematic affinity suggested No extant affinity identified. CJ53 (Plate 42) Diagnosis Actinodromous. Distinct higher orde r veins with moderately and well developed areoles, 1-branched F.E.V.s. covered with small laminar resin dots. Description Laminar size mesophyll, laminar shape ovate; base shape cordate; margin entire; primary veins are basal actinodromous, at least three veins are observed; agrophic veins are compound; tertiary veins are mainly opposite percurrent and their courses are sinuous; the angles of the 3’s on primaries is perpendicula r; 4th vein types are mixed opposite/alternate percurrent; 5th order veins are regular polygonal reticulate; areolation is moderately to well developed; 1-branched F.E.V.s. cove red with small laminar resin dots.

PAGE 45

45 Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0318, Pit Tabaco High Dip, locality placed between coal beds 160 and 161, middle to late Paleocene, co ordinates 11.12 N, 73.55 W, three specimens. Systematic affinity suggested No extant affinity identified. CJ54 (Plate 43) Diagnosis Decurrent base. Acrodromous, three or five primary veins. Secondaries are mostly opposite percurrent a nd perpendicular on midvein, marg inal secondaries are weak brochidodromous. Description Petiole has a swollen base; laminar size mesophyll; laminar shape distinctly oblong; base shape decurrent; apex shape rounded; margin entire; primary veins are parallelodromous, 3 to 5 veins; secondary veins are opposite perc urrent and their angles are perpendicular on primaries, but marginal second aries are weak brochido dromous; tertiary veins are regular polygonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0319 and 0410, Pits Tabaco High Dip and Expanded West, localities placed between coal beds 40 and 107, middle to late Paleocene, six specimens. Systematic affinity suggested No extant affinity identified. CJ55 (Plate 44) Diagnosis Leaflets. Short and pulvin ate petiole. Asymmetrical ba se. Four to six crowded basal veins, on either side of the midvein, 2 or three secondary veins runn ing with acute angle. Description Petiole pulvinate and short; laminar si ze of the leaflets from microphyll to mesophyll; laminar shape elliptic; laminar symmetr y strongly base asymmetrical; base shape complex; apex shape acuminate; margin enti re; pinnate leaflets; secondary veins are

PAGE 46

46 brochidodromous, four to six of these veins ar e crowded at the base; secondary vein spacing decreases toward base and their angles abruptly decrease toward base as well; intersecondaries are weak; tertiary veins are random reticulate and their angle variab ility is inconsistent; 4th order types are regular po lygonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, localities 0317 and 0410, Pits Tabaco 1 and Expanded West, localities placed betw een coal beds 40 and 102, middle to late Paleocene, 21 specimens. Systematic affinity suggested Based on the leaf morphology a nd the presence of a short and pulvinate petiole the morphotype closely resembles the family Leguminosae. CJ56 (Plate 45) Diagnosis Pinnate with cordate base. Thick prim ary vein. Four basal veins, compound agrophic veins. Craspedodromous 2s with strai ght course and branching at the margin. Teeth concave/concave. One specimen covered with peltate scales or laminar resin dots. Description Laminar size from mesophyll to megaphyll; laminar shape elliptic; base shape from cordate to convex; leaves pinnate; agrophic veins are compound; secondary veins are craspedodromous; tertiary veins constantly opposite percurrent with strong sinuous courses, the angles of 3’s on primary are perpendicular; tertia ry vein angle variability increases exmedially; 4th order veins are mostly opposite percurrent; teeth concave/concave. One specimen covered with peltate scales or laminar resin dots. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0324, Pit Tabaco High Dip; locality placed between coal beds 125 and 130, middle to late Paleocene, coordinates 11.62N, 73.32 W, 11 specimens.

PAGE 47

47 Systematic affinity suggested Some species in Flacourtiaceae have similar leaf characters as those seen in the fossil. CJ58 (Plate 46) Diagnosis Thick and slightly curved midvein. S econdaries and inters econdaries strongly decurrent on midvein. Tertiaries very thin and mostly random reti culate on midve in and mostly opposite percurrent exmedially. Tertiaries are decurrent on primary and secondaries. 1-2 branched F.E.V.s. Description Laminar size mesophyll; laminar shape el liptic; base angle shape acute and probably decurrent; leaf pinnate; intersecondary ve ins are thin and weak ; tertiary veins are opposite percurrent and their angles on primary are perpendicular; tertiaries are decurrent on primary and secondaries; tertiary vein angle variability increase s exmedially; 4th and 5th order types are distinctly irregular po lygonal reticulate; areolation is poorly developed; 1-2 branched F.E.V.s. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen. Systematic affinity suggested No extant affinity identified. CJ59 (Plate 47) Diagnosis Primaries are flabellate. Higher or der veins mostly random reticulate. Description Laminar size notophyll; laminar shape elliptic; base angle obtuse and probably base shape rounded; margin entire; pr imary veins are flabellate; tertiary veins are random reticulate; coriaceous texture.

PAGE 48

48 Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0317, Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, c oordinates 11.13N, 72.57W, three specimens. Systematic affinity suggested No extant affinity identified. Monocots CJ3 (Plate 48) Diagnosis Leaf apex rounded, margin en tire, cordate or sagittate ba se each basal lobe fed by a secondary vein that branches several times, naked basal veins, midrib and secondary veins stout and multistranded, secondary veins crowded basally and attached to the midrib at very obtuse angles basally (~90-100) and decreasing api cally to ~45, secondaries merging at several intramarginal veins in a complex brochidodromous pa ttern, tertiary veins a ttached to the midrib and secondaries at angles between 20 and 45 to form a network of intersecondary venation, higher order veins are strongly irregular polygonal re ticulate that have wandering courses and connections at variable angles. Description Maximum length measured is ~56 cm, and maximum width measured at the posterior division ~26 cm, margin entire; leaf apex rounded; midrib multistranded and up to 1.5 cm wide; leaf base cordate or sagittate, the basa l lobes are fed by the most basal secondary veins, these secondary veins are very we ll-developed and form part of th e leaf margin at the base as naked basal veins, the two basal secondary veins also dichotomize within the lobes several times to form minor secondaries that have perpendicula r courses with respect to the midrib and very obtuse angles basally; secondary veins are multistranded, very thick and decurrent on the midrib, secondary veins crowded basally and four veins depart from the petiole insertion, secondary veins branch in a complex brochidodromous patte rn, never reaching the ma rgin; the course and spacing in between the intramarginal veins are ve ry irregular; higher order veins are irregular

PAGE 49

49 polygonal reticulate with wandering courses (sinuous, angular, or st raight) and connections at variable angles. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0322, Pit La Puente, locality placed between coal beds 103 and 105, middle to late Paleocene, six specimens. Systematic affinity suggested The leaf form and venation pa tterns of the fossil leaves are extremely similar to those found in the family Araceae and the extant genus Montrichardi a. CJ23 (Plate 49) Diagnosis Leaves ovate, margin entire, apex acumin ate, base cordate or sagittate, naked basal veins, midrib multistranded, secondary ve ins eucamptodromous, one or two intramarginal veins running close to th e margin, one or two intersecondary veins running parallel in between secondaries, tertiary veins mixed opposite / alternate percurre nt and reticulate. Description Laminar shape ovate, length 6.2 cm, wi dth 3.8 cm; margin entire; posterior division of the blade not completely preserved bu t the presence of laminar tissue proximal to the petiole attachment shows the base is cordate or sagittate; naked ba sal veins form part of the leaf margin near the petiole attachment; anterior divi sion of the blade longer than posterior division; midrib multistranded; the most basal secondary vein curves in a proximal direction and feeds the basal lobe. The next most distal secondary is oriented almost perpendicular to the midrib for much of its course, forms the primary intramar ginal vein, and displaces toward the margin a minor intramarginal vein that arises from the pos terior division of the blade. The third secondary vein forms the secondary intramarginal vein, an d the rest of the secondary veins are strongly eucamptodromous and ascend from the midrib at angles between 29 to 35; one or two intersecondary veins are present between each pa ir of secondaries and have courses almost

PAGE 50

50 parallel to the secondaries. Both secondary a nd intersecondary veins are strongly decurrent on the midrib and follow a course that curves toward the margin distally. The tertiary veins are mixed opposite/alternate percurrent but exmedially most tertiari es are opposite percurrent and are oriented perpendicularly to the midrib, excep t for those developed in the basal lobe which tend to be parallel to the mi drib. Higher order veins are ir regular polygonal to orthogonal reticulate. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, two specimens. Systematic affinity suggested The leaf form and venation pa tterns of the fossil leaves are extremely similar to those found in the family Araceae and the extant genus Anthurium CJ47 (Plate 50) Diagnosis Multistranded midrib. Submarginal vein. Numerous parallelodromous secondaries, closely spaced and decurrent on midvein. Thick and darkened fimbrial vein. Specimen covered with small resin dots. Description Fossil fragment; laminar size mesophyll; the fossil is missing the base; margin entire; primary vein multistranded and it may be pinnate, numerous parallelodromous secondaries, closely spaced and decurrent on midvein ; thick and darkened intramarginal vein that collects the secondary veins; thick fimbrial vein runs parallel to margin; the specimen is covered with small resin dots. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0315; Pit Tabaco 1, locality placed between coal beds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W, one specimen.

PAGE 51

51 Systematic affinity suggested The leaf form and venation patterns of the fossil leaves seem similar to those found in the family Araceae, however a more detailed comparison with other families is necessary. CJ49 (Plate 51) Diagnosis Blade decurrent on petiole. Numerous parallelodromous secondaries, closely spaced with perpendiculary cross veins. Description Laminar size from mesophyll to macr ophyll; margin entire, primary vein pinnate; secondary veins are numerous and parallelodromous, closely spaced with perpendiculary cross veins. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0322; Pit La Puente, locality placed between coal beds 103 and 105, middle to late Paleocene, seven specimens. Systematic affinity suggested The leaf form and venation patterns of the fossil leaves seem similar to those found in the order Zinge berales and the family Araceae, however a more detailed comparison with other families is necessary. CJ62 (Plate 52) Diagnosis Primary and/or secondaries decurrent and multistranded. Tertiaries are mostly opposite percurrent and strongly pe rpendicular to primary and/or to secondaries. Higher order veins are regular polygonal reticulate. Specimen covered with resin dots. Description Laminar size mesophyll; margin entire; primary and/or secondaries decurrent and multistranded; tertiaries are mostly opposit e percurrent and strongly perpendicular to primary and/or to secondaries; higher order ve ins are regular polygonal re ticulate; the specimen is covered with resin dots.

PAGE 52

52 Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0324; Hi gh Dip, locality placed between coal beds 125 and 130, middle to late Paleocene, one specimen. Systematic affinity suggested No extant affinity identified. CJ63 (Plate 53) Diagnosis Stout midrib. Secondaries multistrande d and decurrent on midvein. Tertiaries very thin, decurrents, i rregular paths and with acute angle on secondaries. Description Laminar size megaphyll; the fossil misses the base and the margin; midrib stout and multistranded; secondary veins are mu ltistranded and decurrent on midvein; tertiaries very are thin, decurrent, and have irregular paths with very acute angle on secondaries. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0322; Pit La Puente, locality placed between coal beds 103 and 105, middle to late Paleocene, one specimen. Systematic affinity suggested The leaf form and venation patterns of the fossil leaves seem similar to those found in the order the family Araceae, however a more complete specimen is required and a more detailed comparison with other families is necessary as well. CJ64 (Plate 54) Diagnosis Pinnate, multistranded midrib. Secondar ies strongly decurrent, secondaries widely spaced and without fimbrial vein like CJ47. Description Laminar size mesophyll; leaf pinnate and multistranded; secondary veins are strongly decurrent on midrib, and th ey widely spaced, the specimen misses a fimbrial vein like seen in CJ47.

PAGE 53

53 Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine; Pit La Puente, locality placed between coal beds 103 and 105, middle to late Paleocene, one specimen. Systematic affinity suggested The leaf form and venation patterns of the fossil leaves seem similar to those found in the order Zinge berales and the family Araceae, however a more detailed comparison with other families is necessary. CJ65 (Plate 55) Diagnosis Midrib and secondaries are multistran ded. Secondaries and tertiaries are strongly decurrent on midvein and have parallel pa ths. Fourth order vein s with acute angle on tertiaries and i rregular course. Description Laminar size megaphyll; margin entire; leaf pinnate; the midrib and secondary veins are multistranded; secondaries a nd tertiaries are strongl y decurrent on midvein and have parallel paths; 4th order veins with acute angle on tertiaries and irregular course. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal-mine, locality 0323; Pit High Dip, locality placed between coal beds 115 and 120, middle to late Paleocene, two specimens. Systematic affinity suggested The leaf form and venation patterns of the fossil leaves resemble to those found in the order Zingeberales. CJ67 (Plate 56) Diagnosis Fragments of costapalmate or palmate l eaves. Segments with entire margins, a midvein, and four orders of pa rallel venation connected by irre gular transverse commisures. Description Rachis 23 mm wide in the base an d 7 17 mm wide in more apical fragments, with a longitudinal ri dge; leaflets are single fold, w ith entire margins, regularly

PAGE 54

54 arranged in one plane, opposite to semialternate, separated by 8 – 42 mm at their insertion; midrib prominent, 0.5 – 0.7 mm wide, two orders of parallel veins se parated by 1 – 2.3 mm, transverse veinlets not evident; the leaflets are joined together in fragments with the thinnest rachis. Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal mine, locality 0319; Pit Tabaco High Dip; locality placed between coal beds 105 and 107, middle to late Paleocene, three specimens. Systematic affinity suggested Arecaceae (palm family). CJ68 (Plate 57) Diagnosis Fragments of pinnate leaves. Leaflets si ngle fold, with entire margins, regularly arranged in one plane, opposite to semialternate, midrib prominent, two orders of parallel veins, transverse veinlets not evident. Description Fragments of costapalmate or palm ate leaves; segments 51.6 – 112 mm wide, with entire margins, and a midvein 1.5 mm wide; pa rallel veins of four or ders: first order veins 1.17 – 1.6 mm wide, separated by 38 38.29 mm; s econd order veins 0.8 1.1 mm wide; primary and secondary veins separated by 0.2 mm appr oximately; third order veins 0.1 0.2 mm wide, separated by 2.0 – 3.44 mm; tertiary veins or tertiary and sec ondary veins separated by 6 – 8 mm; fourth order veins very thin, almost indi scernible, separated by 0.2 mm and approximately seven in number between two third order veins; transversal veins 0.10 – 0.12 mm wide, with an irregular course, inserted to the middle or primary veins with an acute angle, and separated by 1 5 mm; after their inse rtion, they bend to cross the secondary veins with an almost straight angle; they can connect two primaries or a primary and a lesser order vein; transversal venation is not always visible, in some leaf fragments is onl y visible where a partia l decomposition took place.

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55 Source, age, number of specimens, and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal mine, localities 0317, 0318, and 0323; Pits Tabaco 1 and Tabaco High Dip; localities placed between coal beds 100 and 107, middle to late Paleocene, 61 specimens. Systematic affinity suggested Arecaceae (palm family). Paleoclimate I used modern correlations between dicot leaf margin t ype and temperature (Wilf, 1997) and leaf size and precipitation (Wilf et al., 1998) to estimate the climatic conditions under which the Cerrejn flora grew. Sevent y-four percent of the 41 dicot leaf types have entire margins, yielding a mean annual temperature (MAT) estimate of 23.8 2.3 C. It is pr obable that riparian and wetland floras like those represented by the fossils typically have a higher proportion of toothed species than adjacent te rra firme vegetation and thus can underestimate MAT by up to 57 C (Burnham et al., 2001; Kowalski and Dilcher, 2003). More than 50% of the dicot leaves in the Ce rrejn flora are mesophy ll or larger in size (>1900 mm2), which is striking given the preservationa l biases against large leaves (Roth and Dilcher, 1978). I infer a mean annual preci pitation (MAP) of 3240 mm (range 2260-4640 mm), which is likely an underestimate (Burnham et al., 2005). The abundant, thick beds of low-ash, low-sulfur coal also indicate a wet climate. The Cerrejn fossil leaf physiognomy and th e reconstructed paleoclimate prove the existence of a 58-60 Ma Neotropical Rainforest Modern rainforests ar e recognized for having assemblages with high proportions of large, en tire-margined, thin-textured leaves. Modern rainforests have a MAT that oscillates between 22 and 27 C, and MAP usually is higher than 2200 mm, with a dry season no longer than 2 m onths (Dolph and Dilcher, 1980; Burnham and Johnson, 2004).

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56 The Cerrejn MAT estimate is similar to modern tropical temperatures, and does not display very high MAT as do some high p-CO2 climate simulations and oxygen isotope estimates of Paleocene tropical sea surface temperature (Pearson et al., 2001; H uber et al., 2003; Shellito et al., 2003). The paleoclimatic simulations suggest a low-latitude MAT higher than 28 C with 500 ppm of CO2, and a maximum of 34 C with 2000 ppm of atmospheric CO2 (Shellito et al., 2003). If leaf margin analysis underestimates MAT at Cerrejn by 7 C, the equator-to-pole temperature contrast for the middle to late Paleocene w ould have been only ~20 C, requiring climate mechanisms that retained heat at high latitudes more effectively than in the present day (Sloan and Pollard., 1998; Huber et al., 2003; Sewall et al., 2004). The Cerrejn MAP estimate is congruent with the predictions by Shell ito et al (2003), but the small scarce differences between the predicti ons of paleoprecipitation under different levels of CO2 (from 500 to 2000 ppm) make in ferences about the possible CO2 levels reached during the early Paleogene based solely on precipitation estimates inconclusive. Floristic Composition of the Cerrejn Flora The floristic composition of the Cerrejn flora compared informally with the most important tropical families (Burnham and Johnson, 2004) shows that this middle-late Paleocene flora had a family-level taxonomic composition typi cal of extant Neotropical rainforests. The most diverse and abundant families in the Cerrej n florawere: Fabaceae (4 leaf morphotypes), Malvales (3), Araceae (3), Arecaceae (2), Laurac eae (2), Zingiberales (2), Anacardiaceae (1), Cycadaceae (1), and Menispermaceae (1). Also, the four ferns found include: the pantropical, floating-aquatic Salvinia ; Stenochlaena a common climbing fern in Old World tropical swamps; the cosmopolitan Lygodium ; and Acrostichum a pantropical fern common in freshwater and mangrove swamps.

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57 As well as fossil leaves, fruits, seeds and fl owers were found in the Cerrejn Formation; the most abundant fossil fruits belong to Leguminosae (4 types of pods) and Arecaceae (3 morphotypes) which are related to m oderns coconuts and the mangrove palm Nypa In addition, the pollen record obtained from the Cerrejn Form ation shows components of modern rainforests (Jaramillo et al., 2007). Some of the most abundant pollen grains are proxapertites operculatus (Araceae), Mauritiidites franciscoi var. pachyexinatus (Arecaceae), Psilatriletes types (ferns), and Momipites africanus (Moraceae). When the familial/ordinal taxonomic composition of the Cerrejn megaflora was compared statistically with that of extant lowland Neotr opical forests, the results showed a remarkable degree of similarity (Wing et al., in press). The comparison was done with 73 of Gentry’s 0.1 hectare transects (Phillips et al., 2002). The mode rn sites were tabulated for the proportions of species and individuals belonging to each family /order. The fossils were tabulated for the proportion of leaf morphotypes in each family/order for the whole flora a nd the number of leaves in each family/order for a random sample collected at one site. The results obtained by Wing et al. (in press) showed that the mean Spearman coefficient between the Cerrejn and all extant sites varied from 0.41 to 0.95. Thus, in terms of the rank-order diversity and abundance of plant families/orders, the Cerrejn flora is not distinguishable from livi ng Neotropical lowland forests. Conclusions I recovered 1347 megafossil specimens and se gregated them into 57 morphotypes using leaf architectural characters. Th e fossils include foliar organs of 41 dicots, 10 monocots, 4 ferns, 1 cycad, and 1 conifer. In additi on, numerous fruits, seeds and flowers have been found from the Cerrejn Formation, but they were not described in this monograph. Cerrejn plant fossils document the existence 58-60 million years ago of Neotropical Rainforest with modern family composition. The most important Neotropical families and orders

PAGE 58

58 were present in Cerrejn such as Fabaceae (the richest family in number of species in the Neotropics), Malvales, Lauraceae, Menispermace ae, Arecaceae, Araceae, Cycadaceae, ferns, and Zingiberales. The Cerrejn assemblage has a high proportion of large, entire-margined, thin-textured, leaves, indicating a tropical clim ate with mean annual temperature 24 C and rainfall 2.3 m/y. This paleoclimate and fossil leaf physiognomy is consistent with modern climatic conditions of tropical rainforests. Also, the abundan t, thick beds of low-ash, low-sulfur coal from the Cerrejn Formation indicate a wet climate. The estimated Cerrejn MAT is consistent with modern tropical temperatures; this result is much cooler than paleoclimatic simulations for the Paleocene. This paleotemperature inference should be considered tentatively. There are two f actors that may impact the result. First, the riparian and wetland effect may underestimate MAT by up to 5-7 C in tropical floras. Secondly, the empirical limitation in the equations availabl e for reconstructing MAT; nowadays there is not a single forest with a MAT higher than 26-27C, so based on this limitation, it is impossible to get temperatures as they have been predicte d by paleoclimatic simulations during the early Paleogene. It is necessary to obtain independent proxies of MAT from the same formation. The reconstructed MAP points to very humid cond itions during the low-latitude middle-late Paleocene.

PAGE 59

59 CHAPTER 2 FOSSIL ARACEAE FROM A PALEOCENE NEOT ROPICAL RAINFOREST IN COLOMBIA Abstract The family Araceae is very diverse in tropi cal America. Both the fossil record and molecular data support a long evolutionary hi story. Araceous fossils, however, have been recorded only from mid and high latitudes. Here we report the oldest fossil leaves of Araceae from the middle-late Paleocene Cerrejn flora in northern South America. The three fossil leaf taxa are related to two exta nt Neotropical genera, two of them closely related to Anthurium The other fossil taxons were placed within the extant genus Montrichardia The fossil aroids inhabited a coastal rainforest ~60-58 million years ago and provide new information about patterns of angiosperm divers ification in the Neotropics. Introduction Araceae is one of the most diverse monocotyl edonous families, comprising 9 subfamilies, 106 genera, and about 3300 species (Croat, 1979; Mayo et al., 1997; Govaerts and Frodin, 2002; Keating, 2003a; 2004). It is most diverse in the tropics, with the Pacific slopes of the Northern Andes home to the greatest number of species (C roat, 1992b; Vargas et al., 2004; Mora et al., 2006). Growth forms include herbs, vines, hemi epiphytes, epiphytes, l ithophytes, rheophytes, and helophytes (Croat 1990; 1992a; Mayo et al., 1997; Bown, 2000). Despite high species diversity, a large range of life forms, and a wide geographic distribution, th e origin and evolution of the Araceae are poorly known. Its fossil record is sparse and controversial, possibly because the herbaceous life form of most Araceae makes them unlikely to fossilize (Herendeen and Crane, 1995). The oldest reported fossils of Araceae are inaper turate, striate pollen grains from the late Barremian early Aptian (~124-117 Mya) of Po rtugal (Plate 58-1; Fr iis et al., 2004, 2006),

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60 although this age has been questioned and remain s problematic (Heimhofer et al., 2007). Those pollen grains were assigned to the fossil genus Mayoa and may be related to the subfamily Pothoideae, tribe Monstereae according to the mo st recent classification of Araceae by Keating (2003b; 2004). Fossils of Araceae have been repo rted from a number of Late Cretaceous localities (Plate 58-2), including several from th e Coniacian deposits of Eastern Asia (Krassilov, 1973), the Campanian of Central Europe (Kva ek and Herman, 2004), North America (Bogner et al., 2005), and from the Maastrichtian of India and South America (Her ngreen, 1974; Sarmiento, 1994; Bonde, 2000; Hesse and Zetter, 2007). The num ber of fossil reports increases during the Paleocene and Eocene records in both mid and high latitudes (Plate 58-3). Mid latitudes had a subtropical climate during the early Paleogene, e xplaining the prevalence of araceous taxa in present day temperate regions (Dilcher and Daghlian, 1977; Smith and Stockey, 2003; Wilde et al., 2005). The minimum age estimate for the origin of th e family, using molecular analysis calibrated with fossils, is between 105 and 128 million years (Bremer, 2000; Janssen and Bremer, 2004). The extant diversity and subfamilial structure of Araceae is complex and predominantly tropical (Croat, 1979; Mayo, 1993), suggesting that low latitu des should have been the area of origin and radiation. The fossil record of tropical aroids has remained nearly unexplored (Plate 58-1-2, and 3). In tropical South America, for example, th e only araceous records ar e dispersed pollen from the Maastrichtian and Paleocene stages of Spathiphyllum and Proxapertites operculatus types (Van der Hammen & Garcia, 1966; Sarmiento, 1994; Jaramillo and Dilcher, 2001; Hesse & Zetter 2007). Here, I describe three new species of Araceae based upon fossil leaves preserved in Paleocene sediments in northern South America. Two species are relate d to the extant genus

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61 Anthurium and the other is placed within the extant genus Montrichardia The araceous fossil leaves are part of the middle-late Paleocene Ce rrejn flora, which provides new data about the evolution of Neotropical Rainforests (W ing et al., 2004; Herrera et al., 2005). Materials and Methods The fossil leaves described in this study were collected from the Cerrejn Formation, exposed in the Cerrejn mine, a large open pit co al mine located in Northeastern Colombia (11N, 72W). The Cerrejn Formation has been dated by fossil pollen as middle to late Paleocene, making it about 60-58 million years old (Jaramillo et al., 2007). The 700-m-thick stratigraphic sequence, composed of thick coals, fluvially-depos ited sandstones, and lacustrine siltstones was deposited in a rapidly subsiding basin (Bayona et al., 2004 ). The seven specimens of fossil leaves reported here were found ~380 meters above the base of the formation. The fossil material is stored in the paleontological coll ections of the Colombian Geological Institute (INGEOMINAS) in Bogot. I performed an exhaustive comparison of the fo ssils with collections of extant Araceae in the United States National Herbarium, Washington DC, Smithsonian Tropical Research Institute, Republic of Panama, and University of Florida He rbarium, Gainesville, Fl. We studied 90% (95 of the 106) of the extant genera, as well as all the tribes and s ubfamilies. Araceae leaf morphology is extremely variable and diverse; it has not been used commonly as a taxonomic tool. Nevertheless, recent morphological work by Keating (2003b) considers leaf venation a potentially useful taxonomic characteristic. Here we follow Keati ng (2003b, 2004) in using venation to assign the fossil leaves to genera within subfamilies of Araceae. We used leaf terminology adapted from LAWG (1999), due to the confusion w ith the traditional terminology in aroid descriptions (see K eating, 2003b for a discussion). Consequently, some terms such as “primary lateral veins” (Croat and Bunting, 1979 ; Mayo et al., 1997) are called secondary veins,

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62 and submarginal collective vein s or collective veins (Croat and Bunting, 1979; Mayo et al., 1997) are called intramarginal veins. The term midrib was used to describe the primary vein or midvein; secondary veins are the ne xt narrower class of veins after the midrib (and they originate from it); and intramarginal veins closely paralleling the leaf margin (LAWG, 1999). Although two species described in this paper are similar to an extant genus we used form generic names rather than place them in an extant ge nus. This is because of the lack of floral and many leaf structural characters that are used by botanists to identify this extant genus. Although, the fossil material is good and contains many us eful characters, it lacks the total number of characters to place it closer than to subfamily and suggests a gene ric relationship at this time. The standard up until about 35 years ago was to place all fossil angiosperms in extant genera using a general statement that they “looked most similar” to a particular modern genus. After Dilcher (1974) pointed out that when examined carefully, many fossil leaves did not belong to the genera, or even families to which they had been assigned; angiosperm paleobotanists began to examine and use leaf characters with more car e. As a result, we have found that there are many examples of angiosperm evolution throug hout their history. The use of form genus, as done here, reflects this caution a nd honesty in establishing relations hips as a part of angiosperm paleobotany (Roth and Dilcher, 1979; Jones and Dilcher 1980). The leaves of fossil and extant Araceae were photographed for comparison using Nikon D70 and DXM1200F cameras. Leaves of Montrichardia arborescens Schott were cleared and mounted following the techniques of Hickey (1973) and Dilche r (1974). Fossil leaf reconstructions were made using Adobe Photoshop and Adobe Illustrator CS2. Systematics Family Araceae. Subfamily Pothoideae Engler.

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63 Tribe Pothoeae Bartl. Morphogenus Anthurioidea Herrera, Jaramillo, Dilcher, Wing et Gomez-N gen. nov. Generic diagnosis Leaves ovate, margin entire, apex acuminate, base cordate or sagittate, naked basal veins, midrib multistranded, s econdary veins eucamptodromous, one or two intramarginal veins of secondary gauge, one or two intersecondary veins per intercostal region with courses parallel to thos e of the secondaries, tertiary veins mixed opposite/alternate percurrent and reticulate. Derivation of generic name From the extant genus Anthurium Type species Anthurioidea wayuuorum Herrera, Jaramillo, Dilcher, Wing et Gomez-N sp. nov. Second species Anthurioidea cerrejonense Herrera, Jaramillo, Dilcher, Wing et Gomez-N sp. nov. Species diagnosis Anthurioidea wayuuorum Herrera, Jaramillo, Dilcher, Wing et GomezN. Laminar shape ovate, margin entire, base is either cordate or sagitta te, naked basal veins, midrib multistranded, two intramarginal vein s running near the margin, secondary veins eucamptodromous, intersecondary veins present and parallel to secondaries, most tertiary veins mixed opposite/alternate percur rent, but exmedially they me rge perpendicularly with the innermost intramarginal vein, higher order veins irregular po lygonal to orthogonal reticulate. Holotype ING-NMNH-0902, only specime n. Plate 59-4-5 and 6. Species description Holotype incomplete; laminar sh ape ovate, length 6.2 cm, width 3.8 cm (Plate 59-4 and 6); margin entire; posterior di vision of the blade not co mpletely preserved but the presence of laminar tissue proximal to the pe tiole attachment shows the base is cordate or sagittate; naked basal veins form part of the l eaf margin near the petiole attachment; anterior

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64 division of the blade longer than posterior divisi on; midrib multistranded; the holotype preserves 6 pairs of secondary veins on the right side of the blade. The most basal secondary vein curves in a proximal direction and feeds the basal lobe. The next most distal secondary is oriented almost perpendicular to the midrib for much of its cour se, forms the primary intramarginal vein (Plate 59-5), and displaces toward the margin a minor intr amarginal vein that arises from the posterior division of the blade. The third secondary vein forms the secondary intramarginal vein, and the rest of the secondary veins are strongly eucampt odromous and ascend from the midrib at angles between 29 to 35; one or two intersecondary veins are present between each pair of secondaries and have courses almost parallel to the secondaries. Both secondary and intersecondary veins are strongly de current on the midrib and follow a course that curves toward the margin distally. The tertia ry veins are mixed opposite/alterna te percurrent, but exmedially most tertiaries are opposite percur rent and are oriented perpendicu larly to the midr ib (Plate 595), except for those developed in the basal lobe which tend to be parallel to the midrib. Higher order veins are irregular pol ygonal to orthogonal reticulate. Derivation of specific epithet From the Wayuu, an Amerindian group of the La Guajira Peninsula, Rancheria Basin. Source, age and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coalmine, locality 0315, Pit Tabaco 1, locality placed betw een coal beds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W. Species diagnosis Anthurioidea cerrejonense Herrera, Jaramillo, Dilcher, Wing et Gomez-N. Margin entire, leaf apex acuminate, midrib stout and multistranded, secondary veins eucamptodromous with dichotomies and holding mi nor agrophic veins, one intramarginal vein

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65 running very close to margin, intersecondary ve ins occur; tertiary and higher order veins reticulate, fimbrial vein present. Holotype ING-0804, only one specimen f ound. Plate 60-7-8-9-10, and 11. Species description Holotype incomplete, blade is >9.5 cm long and >24.3 cm wide (Plate 60-7 and 11); margin entire; leaf apex acuminate; midrib stout and multistranded; five pairs of secondary veins (preserved on the anterior division of the blade) which depart the midrib at about ~50 proximally, shifting to <20 near the apex; secondary vein s branch at ~45 and arch apically (Plate 60-8); one or two pairs of intersecondary veins are present for each pair of secondaries, and their course are more or less parallel to the them; secondary and intersecondary veins are strongly decurrent on midrib; minor agrophi c veins present, an intramarginal vein runs very close (~0.5 mm) to margin (P late 60-9); tertiary veins form polygons of different shapes and sizes (Plate 60-8-9 and 10); high er order veins also form irregu lar polygons measuring between 5 mm and 8 mm in length and width; a fimbrial vein runs along the margin (Plate 60-9). Derivation of specific epithet From the Cerrejn Formation, where the fossil was found. Source, age and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal mine, locality 0315, Pit Tabaco1, lo cality placed between coal be ds 100 and 102, middle to late Paleocene, coordinates 11.13N, 72.57W. Leaf morphology comparison within monocots The family Araceae is placed in the Alismatales, an order of fourteen families (A PG-II, 2003) of predominantly herbs of aquatic habitat. Most Alismatales have basic monocot leaves: oblong, a nd entire-margined with parallel major veins and short, perpendicular cross vein s. Araceae are unusual in the order for having a strong similarity to dicot leaves (K eating, 2003b). Although some Alismataceae, Limnocharitaceae, and Potamogetonaceae and outside of Alismatales, Dioscoreaceae

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66 (Dioscoreales) have ovate to el liptic blades with cordate to sa gittate bases and acrodromous or campylodromous venation that resemble Araceae (s ee Riley and Stockey (2004) for a discussion of leaf morphology in these families), unlike thes e four families, neither fossil of Anthurioidea has primary veins running in strong convergent arches toward the leaf apex typical of acrodromous or campylodromous venation. Furthermore, many species within Alismataceae, Limnocharitaceae, Potamogetonaceae and Diosco reaceae have dominantly opposite percurrent tertiary veins unlike those in Anthurioidea. Systematic affinity within Araceae Anthurioidea wayuuorum and Anthurioidea cerrejonense closely resemble species in the extant Neotropical genus Anthurium Schott. This genus is placed in the tribe Pot hoeae Engler (Keating, 2003; 2004) with Pothos (~70 spp., from South-Southeast Asia and Madagascar), Pedicellarum (1 sp., from Indonesia and Malaysia), and Pothoidium (1 sp., Southeast Asia). Keating (2003b) recognized different types of leaf morphologies within the tribe ranging from primary and seco ndary veins as acrodromous, brochidodromous, and eucamptodromous, and higher orders reticulate. Clearly, Anthurium is extremely more diverse and variable in leaf shap e, size and venation types compared to the other genera within the tribe. A. wayuuorum and A. cerrejonense do not resemble any of the typical leaf shapes and characters present in these th ree genera: linear-lanceolate to ovate or narrowly elliptic blades, furthermore the th ree genera lack either cordate or sagittate bases as seem in extant Anthurium and the two fossil taxa. Anthurium is restricted to tropical-subtropical Am erica and the West Indies and together with Philodendron and Monstera are perhaps the most variable genera in terms of leaf morphology (Grayum, 1990; Ray, 1990). However, the combination of leaf characters such as secondary veins forming intramarginal veins a nd cordate or sagittate bases and higher order

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67 veins variables seem to be typi cal and diagnostic for the genus Anthurium (Croat and Bunting, 1979). A. wayuuorum and A. cerrejonense have similar venation patterns to many modern species of Anthurium in terms of intramarginal venation, secondary and higher order veins types (Table 2-1). A. wayuuorum is comparable to modern species that have two intramarginal veins (Plate 61-12-13-14, and Ta ble 2-1). In contrast, A. cerrejonense is similar to extant Anthurium species that possess only a singl e intramarginal vein (Plate 61-15-16, and Table 2-1) running about ~0.5 mm from the leaf margin (Plate 609). Although, these differences in submarginal vein patterns are distinctive characters used to di scriminate between species, in some very rare cases we found both types of submarginal veins present in extant species of Anthurium (Plate 6114), although this morphological va riation changes only a small part of the margin and involves one secondary vein simply. The specimens presen ted here are thought to represent two distinct fossil species based on differences in: tertiary ve in patterns, presence and absence of dichotomy, agrophic and fimbrial veins, and finally the dissimilarity between angles of the secondary veins (Table 2-1). Although, there are several good le af characters (secondary ve ins forming intramarginal veins and cordate or sagittate ba ses and higher order veins variab les) that are recognizable and useful for many members in Anthurium, those ch aracters are not unique for the genus. Keating (2003b) distinguished the reticu late high order veins as those found in the tribe Pothoideae (Keating, 2003b) and the subfamily Lasioideae as useful systematically. However, there is no way to differentiate members of this tribe a nd subfamily based only on the higher order veins. Both species reported here share similar higher or der veins to those found in the tribe Lasioideae. Also A. cerrejonense has comparable secondary vein types that end in one single intramarginal vein. However, extant members of Lasioideae ha ve other leaf venation patterns that are unique

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68 for that subfamily and that are not present in the Cerrejn fossils neither in the genus Anthurium For example in the genus Urospatha (Table 2-1) and all genera within the subfamily, generally the leaves with pinnate venation have the posteri or division of the blades more longer than the anterior division, this is reflect ed in having deeply sagittate bases, few numbers of apical secondary veins (usually 1 to 3) secondary veins angle very low, and their courses run in strong convergent arches toward the leaf apex. None of the fossils repor ted here shows all these leaf venation patterns, for this reason the fo ssil taxa are most closely related to Anthurium than to another genus within the family by the comb ination of blade morphology and the venation patterns, the assignation of the generic name was discussed pr eviously on the Method section. Subfamily Philodendroideae Engler. Tribe Philodendreae Schott. Genus Montrichardia Crger Generic leaf diagnosis Leaf apex rounded, margin entire, cordate or sagittate base each basal lobe fed by a secondary vein that branches several times, naked basal veins, midrib and secondary veins stout and multistranded, secondary veins crowded basally and attached to the midrib at very obtuse angles basally (~90-100) and decreasing ap ically to ~45, secondaries merging at several intramarginal veins in a co mplex brochidodromous pa ttern, tertiary veins attached to the midrib and secondaries at angl es between 20 and 45 to form a network of intersecondary venation, higher order veins are st rongly irregular polygonal reticulate that have wandering courses and connectio ns at variable angles. Fossil species Montrichardia aquatica Herrera, Jaramillo, Dilcher, Wing et Gomez-N sp. nov. Fossil holotype ING-0904. Fig. 23, 24, 25, and 26.

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69 Fossil paratypes ING-0808, Fig. 27, 28, 29, and 30; ING-NMNH-0903, Fig. 31, 32, and 33; ING-0905 and ING-0906, Fig. S5 (see Supplemen tal Data accompanying online version of this article). Species diagnosis Montrichardia aquatica Herrera, Jara millo, Dilcher, Wing et Gomez-N. Same as for the genus Montrichardia. Species description Maximum length measured on the holotype (Plate 62-23) is ~56 cm, and maximum width measured at the posterior division ~26 cm, the paratype ING-0808 (Plate 63-27) is ~36 cm long and ~26 cm wide measur ed from the midrib to the margin, ING-NMNH0903 (Plate 63-31) correspond to a portion of leaf margin that reaches a width of 7.3 cm and a length of 6.8 cm, ING-0905 reaches a maximum le ngth of ~41 cm measur ed marginally and a maximum width of ~21 cm, ING-0906 is a fragment of the posterior division that reaches a maximum length of about 11 cm and maximum wi dth of 22 cm; margin entire; leaf apex rounded; midrib multistranded and up to 1.5 cm wide ; leaf base cordate or sagittate (Plate 62-24, and 25), the basal lobes are fed by the most basal secondary vein s, these secondary veins are very well-developed and form part of the leaf margin at the base as naked basal veins (Plate 6225), the two basal secondary veins also dichotomize within the lobe s several times to form minor secondaries that have perpendicu lar courses with respect to the midrib and very obtuse angles basally (Plate 62-24 and 25); secondary veins ar e multistranded, very thick and decurrent on the midrib, the holotype and the paratypes ING-0808 and ING-0905 (Plate 62-26) preserve up to 10 and 11 secondary veins on the anterior division of the blade, secondary veins crowded basally and four veins depart from the petiole inse rtion, secondary veins branch in a complex brochidodromous pattern, never reaching the ma rgin (Plate 63-31); secondary veins on the anterior division are attached to the midrib at very obtus e angles, basally ~90-100 but

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70 decreasing apically to ~45, the course of the s econdary veins is straight to slightly up curved exmedially; tertiary veins are very decurrent on the midrib and secondary veins at angles between 20 and 45 (Plate 63-29 and 30), tertiary vein spacing is 1-6 mm (Plate 63-30 and 32); tertiary veins join to form an intersecondary vein network, and are joined exmedially to the secondaries by higher order veins, forming a system of intramarginal veins very close to margin (Plate 63-31); the course and sp acing in between the intramarginal veins are very irregular; higher order veins are irregular polygonal reticulat e with wandering course s (sinuous, angular, or straight) and connections at vari able angles (Plate 63-32 and 33). Derivation of specific epithet From the Latin aquaticus “living in or near water.”. Source, age and stratum Colombia, Rancheria Basin, Cerrejn Formation, Cerrejn coal mine, ING-0904, ING-0905 and ING-906 were found at Pit Tabaco Extension, locality 0708 and 0705, localities placed between coal beds 165 and 170, coordinates 11.07N, 72.34W; ING-0808 and ING-NMNH-0903 were found at pit Tabaco 1, locality placed between coal beds 100 and 102, coordinates 11.15N, 72.55W, age middle to late Paleocene. Leaf morphology comparison within angiosperms Outside of Araceae, the only families that share characters to Montrichardia aquatica are Nymphaeaceae and Nelumbonaceae. These two families can be compared to the fossil exclusively in terms of size and base shape, see Upchurch et al. (1994) and Wang and Dilcher (2006) for a complete revision of the leaf morphology in these fa milies. Nymphaeaceae and Nelumbonaceae lack intramarginal and intersecondary venation, nake d basal veins, and higher order veins that characterize the Cerrejn fossil and any araceous leaf venation. There are no extant monocot families other than to Araceae that ha ve all the characters seen in M. aquatica

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71 Systematic affinity within Araceae The leaf form and vena tion patterns of the fossil leaves are extremely similar to those found in the extant genus Montrichardia Crger (Plate 6117 and 18) which has two species M. arborescens and M. linifera This genus is placed in the tribe Philodendreae Schott (Keating, 2003; 2004) with Anubias (8 ssp., from tropical Africa), Furtadoa (2 spp., from Indonesia and Malaysia), Philodendron (~500 ssp., from Mexico to Argentina), and Homalomena (~110 spp. from tropical America and Southeast Asia). Keating (2003b) recognized several leaf venation patterns for the tribe Philodendreae which range from primary and secondary veins as acrodromous, brochidodromous, and eucamptodromous, and for having mostly many closely spaced intersecondary veins parallel each other. The DNA data and the anatomical characters situate this subfamily as the most varied and rich in the entire Araceae (Keating, 2004). In addition, it has been indicated that Montrichardia has some genetic isolation in respect of the other four genera within th e tribe Philodendreae (Cabre ra et al., 2003; Keating, 2004). This “isolation” is also seen in terms of leaf venation, Montrichar dia is easily separated from the other four genera for having the “co locasioid venation”, and no species within those four genera have this pe culiar venation character. The genus Montrichardia is restricted to tropical America and the West Indies. M. aquatica has “colocasioid venation,” defined as “a t ype of higher order leaf venation in which the finer veins branch almost at ri ght angles from the primary late ral veins and then arch strongly towards the leaf margin, often fusing along the wa y to form a more-or-le ss sinuous interprimary collective vein between the primary lateral veins, and finally joining within the margin to form a intramarginal collective vein” (Mayo et al., 1997, page 310). Colocasioid venation is known in the extant tribes Caladieae (New World, except for Hapaline from Southeast Asia and Brunei) and Colocasieae (Old World), both tribes are part of the subfamily Aroideae, and seem to be very

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72 well separated genetically each other (Keating, 2 004). Colocasioid venation has been reported in fossil leaves from the Neogene of Trinidad a nd the Eocene of Germany (Berry, 1925; Wilde et al., 2005, Table 2-1). M. aquatica has a similar leaf shape and intersecondary venation to Caladieae and Colocasieae (Plate 61-19), but diffe rent intramarginal vein and higher order vein patterns. In Caladieae and Colo casieae the secondary veins (seconda ry veins are called primary lateral veins by Mayo et al., 1997) merge bluntly with one to thr ee straightened intramarginal veins (Plate 61-19, Table 2-1), while the s econdary veins in th e fossil and extant Montrichardia branch very close to the margin subsidized by higher order veins that finally form several intramarginal veins. Also the spacing and course of these intramarginal veins are very irregular compared to tribes Caladieae and Colocasieae in which the intramarginal veins run very parallel to the margin and each other, and the vein spacing is constant (Plate 61-18, 20, and Plate 63-31). Higher order veins and areolati on in Colocasieae and Caladieae are dominated by cross venation and elongated polygons of similar shape and size, while living Montrichardia and the fossil leaves show higher order veins that anastomose irregularly with wandering courses, connections at variable angles, and do not form well-developed areolation (Plate 61-21 and 22, Plate 63-30, 32, 33 and Table 2-1). Because the intramargina l venation pattern and higher order veins characterize the extant genus Montrichardia and the fossil M. aquatica we do not relate the Cerrejn fossil leaves to the fossil morphogenus Caladiosoma Berry, nor to its living relative tribes Caladieae and Colocasieae. Discussion Sedimentary Deposition and Paleoclimate Anthurioidea wayuuorum and Anthurioidea cerrejonense were recovered from a lensshaped deposit of laminated, gray-black, fine-g rained sandstone approximately 50 cm thick and ~6 meters in lateral ex tent (locality 0315). A to tal of 239 leaf specimens have been found in this

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73 locality. Most leaf specimens ar e complete and well preserved; sizes range from microphyll to macrophyll with mesophyll leaves being dominant. Montrichardia aquatica (ING-0808) was recovered approximately 3 meters below locality 0315; the rock is a brown, fine-grained lithic sandstone with inclined heterolithic strata. These araceous fossils were deposited in fluvial channel sediments. The fine-grained sandstones w ith laminations and hete rolithic stratification suggest overbank deposits, such as crevasse spla ys and forebanks (Bayona et al., 2004). These deposits grow by accumulation of fine-grained sediment during episodes of flooding (Reading and Levell, 1996) The specimen ING-NMNH-0903 of M. aquatica was found at the same stratigraphic level as locality 0315, between coal beds 100 and 102, in a dark grey siltstone. The holotype of M. aquatica (ING-0904) and paratypes I NG-0905 and 0906 were recovered from an extensive gray siltstone bed approximately 60-90 cm thick (localities 0705 and 0708). The gray siltstone underlies coal bed 170, one of the th ickest coal beds at the top of the Cerrejn Formation (~7 m). These siltstones were de posited in a swampy-lacustrine environment. Leaf margin and leaf area an alysis of the entire associat ed flora suggest a mean annual precipitation of ~4 meters a nd a mean annual temperature 24 C (Herrera et al., 2005). The most abundant families and orders recognized in the Cerrejn flora ar e Leguminosae, Arecaceae, Malvales, Lauraceae, Menispermaceae, and Zingiberales. The Cerrejn paleoclimate and its floristic composition suggest the presence of a Neotropical rainforest during the middle-late Paleocene in Northern South America (W ing et al., 2004; Herr era et al., 2005). Paleoecology Lithofacies, sedimentary struct ures, and the preservation of large leaves within the associated paleoflora suggest that Anthurioidea wayuuorum and Anthurioidea cerrejonense were part of the local flora, rather than having being transported from far away (Burnham, 1989). A

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74 reconstruction of the life forms of A. wayuuorum and A. cerrejonense must be based upon what is known of the closest living genus. Most Anthurium species are epiphytes (Croat, 1990). It has been estimated that in some Amazonian floodplain forests ~90% percent of trees carry epiphytic aroids (Leimbeck and Balslev, 2001); such comm on epiphytes might have been preserved as fossils. The sedimentary environmen t also raises the possibility th at the fossils were rheophytes, but this life form is rarely found in tr opical American Araceae (Croat, 1990). Montrichardia possesses a strict helophytic life form (Croat, 1990; Mayo et al., 1997). Montrichardia grows in freshwater habitats, especially fluvial to coastal plain areas, along river margins and adjacent swamps subject to seasona l changes in water level (Croat, 1990; Bown, 2000). The foliage generally remains above the wate r level, forming dense colonies that trap sediment; it is a primary colonist in swamp forest areas (Bown, 2000). Montrichardia is found throughout tropical America, but it is dominant along large ri vers of central Amazonia and deltaic zones where it grows even under tida l influence (Mayo et al., 1997; Bown, 2000). Lithofacies and sedimentary structures suggest that M. aquatica grew along river margins (forebanks), and swamps, where it coexisted in palm-dominated environments similar to the ecology of extant Montrichardia Paleobiogeography and Palynology The early appearance of the Araceae family in the fossil record (early Cretaceous), even before the more diverse families seen today, is remarkable. However, the aroid fossil record is sparse and most araceous fossils belong to taxa that inhabited ponds, la kes or flooded areas (e.g. Dilcher and Daghlian, 1977; Kva ek, 2003). A complete revision of the Araceae macrofossil record was recently made by Wilde et al. (2005) ; previous revisions include Grayum (1990), Mayo et al. (1997), and Keating (2 003a). It seems clear that the geographic distribution of Araceae in the past was driven largely by the global climate (Plate 58-1, 2 and 3). From the early

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75 Cretaceous (~120 Mya) to the early Paleogene (~ 65-40 Mya) the climate was warmer than today during a period of about 60 million years (Zachos et al., 2001; Bralower et al., 2006). Thus permitted migration and evolution of megatherma l plants into current temperate latitudes (Morley, 2003). By the end of the Cretaceous, th ree or four Araceous subfamilies were present, according to the most recent classification by Keating (2003b; 2004) (Plate 58-1 and 2; Pothoideae, Lemnoideae, Orontioideae, and Aroid eae (?)), and during the early Paleogene three other subfamilies and consequently a more divers e record of new tribes appear in the fossil record, (Plate 58-3; Philodendroid eae, Aroideae, and Lasioideae). The fossil pollen record also shows intere sting aspects of the paleobiogeography of Araceae. Recently Hesse and Zetter (2007) made a complete review of the family’s pollen record, recognizing three pollen ty pes attributable to Araceae: Limnobiophyllum (subfamily Lemnoideae), Spathiphyllum (Pothoideae), and the almost globally distributed Proxapertites operculatus (Philodendroideae/Pothoideae). The Cerre jn palynoflora has high abundance and in some samples dominance of araceous pollen. Ja ramillo et al. (2007) studied the palynoflora through the entire formation, comparing coal (ass ociated with lacustrine-swampy environments) and non-coal samples (associated with fluvial an d estuarine plains). The three araceous pollen grains present in the Cerrejn palynoflora are Proxapertites operculatus (Van der Hammen, 1954) Van der Hammen, 1956, Proxapertites cursus Van Hoeken-Klinkenberg, 1966, and, Spathiphyllum vanegensis (Van der Hammen & Garcia 1966) Hesse & Zetter 2007. Proxapertites operculatus is one of the most abundant polle n types in both coal and non-coal samples. On the other hand, Proxapertites cursus and Spathiphyllum vanegensis are present only in non-coal samples. This distribution by depos itional environment suggests that Araceae may have been ecological dive rsified by the Paleocene.

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76 Comments on Aroid Evolution In the modern Neotropics, Araceae is diverse and shows high endemism. Anthurium and Philodendron contain approximately 50% of the total of species for the family (Croat, 1983). Anthurium is extremely diverse along the wet, Paci fic side of the Andes in Colombia and Ecuador (Croat, 1992b; Vargas et al., 2004; Mora et al., 2006). If the fossils reported here as Anthurioidea really have affinities with the extant Neotropical genus Anthurium it is interesting to think that related forms of th is genus have been present in th e Neotropics since the Paleocene. How Anthurium reached its modern high level of divers ity is still a matter of discussion, but probably its high diversity and endemism is related to the Mio-Pliocene Andean orogeny (Gentry, 1982) and the resulting biogeographic and ecological fa ctors in Andean and lowland populations, respectively (Leimbeck and Balslev, 2001; Jcome et al., 2004). Other genera along the Andean foothills seem to share similar divers ity patterns (Richardson et al., 2001; Pirie et al., 2006; Quijano et al., 2006). Montrichardia has only two species today with widesp read distributions in the Neotropics (Mayo et al., 2007; Bown, 2000). Montrichardia has been present in the Neotropics for at least 60-58 Mya, but it did not divers ify to the extent seen in Anthurium Philodendron or Monstera Alternatively it could have diversified and then lo st its diversity. Perhaps Montrichardia is more influenced by long-distance dispersal by being n ear moving water, thus maintaining constant genetic interchange over long distances a nd reducing the probab ility of speciation. Montrichardia also inhabits a stable aquatic environm ent and it has not changed since the Paleocene. The same water life forms and stable e nvironments have also been seen to preserve the integrity of another fossil angiosperm to the species level in Ceratophyllum (Herendeen et al., 1990). This hypothesis might be test ed by integrating the fossil reco rd together with molecular analysis in future studies. The long history of araceous Neotropical ge nera and their current

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77 contrasting differences in species diversity constitutes a good case study to investigate the causes of species diversity in the Neotropic Rainforests today. The presence of abundant fossil leaves a nd pollen grains in the Cerrejn Formation establishes the fact that Araceae was already an important component in floodplains of coastal tropical humid forests during the ear ly Cenozoic. The Cerrejn fossils are related to at least three modern tribes which may have had strong habitat preferences as modern aroids.

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78Table 2-1. Morphological feat ures of fossil leaf taxa and extant Araceae. Taxon Blade Apex Base Naked basal veins 2 vein type 2 vein Crowding No. posterior 2s No. anterior 2s Caladium bicolor (Aiton) Vent Ovate Convex Cordate-peltate Absent Straightened I.V Present 4 6-8 Xanthosoma mexicanum Liebm Ovate Acute Cordate Pres ent Straightened I.V Present 4 4-5 Xanthosoma sagittifolium Schott Elliptic Acute Cordate Present curved I.V Present 4 5-6 Xanthosoma robustum Schott Ovate Obtuse Cordate Present Straightened I.V Present 4 7 X. undipes (K.Koch & C.D.Bouch) K.Koch Ovate Acute Cordate Present Straightened I.V Present 4 6-7 Xanthosoma violaceum Schott Elliptic Acute Cordate Absent Straightened I.V Absent 4 7 Colocasia esculenta (L.) Schott Ovate Acute Cordate-peltate Absent Straightened I.V Present 8 5-6 Syngonium llanoense Croat Elliptic Acute Cordate Absent Straightened I.V Present 6 15-17 S. schottianum H.Wendl. ex Schott Elliptic Acute Hastate Absent Straightened I.V Present 6 11-13 Montrichardia arborescens (L.) Schott Ovate Acute Cordate Present Brochidodromous Present 4 6-11 Montrichardia linifera (Arruda) Schott Ovate Acute Cordate Present Brochidodromous Present 4 6-11 Montrichardia aquaticaa Ovate Rounded Cordate-sagittate Present Brochidodromous Present 4 10-11 Caladiosoma messelense Wilde, Z. Kva ek & Bognerb Ovate Acute Cordate ? St raightened I.V Present ? 8-9 Caladiosoma miocenicum Berryb ? ? ? ? ? ? ? ? Anthurium oxybelium Schott Ovate Acute Lobate Present Eucamp-brochido Absent 6 7-9 Anthurium variegatum Sodiro Ovate Acute Lobate Present Eucamptodromous Absent 4 7-9 Anthurium subsagittatum (Kunth) Kunth Ovate Acute Cordate Present Eucamptodromous Absent 4 7-8 Anthurium nigrescens Engl Ovate Acute Cordate Present Eucamp-brochido Absent 6 8-9 Anthurium marmoratum Sodiro Ovate Acute Cordate Present Eucamptodromous Absent 6 7-9 Anthurium karstenianum Engl Ovate Acute Hastate Absent Eucamp-brochido Absent 4 6-7 Anthurium concinnatum Schott Ovate Acute Lobate Present Eucamp-brochido Absent 4 8-9 Anthurium brownii Mast Ovate Acute Cordate Present Eucamptodromous Absent 4 8 Anthurium rzedowskii Croat Ovate Acute Cordate Absent Eucamptodromous Present 8 6-7 Anthurium coripatense N.E.Br. ex Engl Ovate Acute Lobate Present Eucamptodromous Absent 4 10-11 Anthurium hoffmannii Schott Ovate Acute Lobate Present Eucamptodromous Absent 6 5-8 Anthurium cotobrusii Croat & R.Baker Ovate Acute Lobate Present Eucamptodromous Present 4 7-8 Anthurium caperatum Croat & R.Baker Ovate Acute Cordate Present Eucamptodromous Absent 6 9-10 A. obtusifolium (W.T.Aiton) G.Don in R.Sweet Ovate Acute Cordate Present Eucamptodromous Absent 6 8-9 Anthurium ochranthum K.Koch Ovate Acute Cordate Present Eucamptodromous Absent 6 9-10 Anthurium. ravenii Croat & R.Baker Ovate Acute Cordate Present Eucamptodromous Absent 4 8-10 Anthurium roseospadix Croat Ovate Acute Cordate Present Eucamptodromous Absent 6 11-12 Anthurium sanctifidense Croat Ovate Acute Cordate Present Eucamptodromous Absent 7 9-10 Urospatha friedrichsthalii Schott Ovate Acute Deeply sagittate Present Eucamptodromous Present 4 1-2 Urospatha sagittifolia (Rudge) Schott Ovate Acute Deeply sagittate Present Eucamptodromous Present 6 1-3 Urospatha somnolenta R.E.Schult Ovate Acute Deeply sagittate Present Eucamptodromous Present 6 1 Anthurioidea wayuuoruma Ovate ? Cordate/sagittate Present Eucamptodromous Absent 4 >4 Anthurioidea cerrejonensea ? Acute ? ? Eucamptodromous ? ? >5

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79 Notes Taxon in bold text correspond to araceous fossil leaves. aCerrejn Fossils. bWilde et al., 2005. I.V. (intramarginal veins); Mixed opp/ alt (M. opposite/alternate percurrent); Re gular poly. ret. (R. polygonal reticulate); Eu campbrochido. (eucamptodromous-brochidodromous). 2 veins angle No. intersecondaries Fimbrial vein No. intramarginals 3 vein type 3 vein Exmedial angle Higher order veins Glands 90-60 1 2 Colocasioid Regular polygonal ret. Absent 90-30 1 2 Colocasioid Regular polygonal ret. squarish Hairs 90-30 1 2 Colocasioid Squarish Hairs 90-45 1 2 Colocasioid Regular polygonal ret. Hairs 90-60 0 2 Colocasioid Regular polygonal ret. Absent 90-60 1 1 Colocasioid Several Present 90-30 1 1-2 Colocasioid Regular polygonal ret. – squarish Absent 90-30 1 3 Colocasioid Irre gular polygonal ret. Present 90-30 1 3 Colocasioid Irre gular polygonal ret. Present 90-40 0 1-4 “Colocasioid” Irregular polygonal ret. Present 90-40 0 1-4 “Colocasioid” Irregular polygonal ret. Present 90-45 0 1-4 “Colocasioid” Irregular polygonal ret. ? 80-40 ? Several Colocasioid Regular polygonal ret. –elongate ? ? ? ? Colocasioid Irregular polygonal ret.–elongate ? 45-40 1-2 0 2 Mixed opp/alt ~90 to midrib Regular polygonal ret. Absent 45 1-2 1 2 Mixed opp/alt ~90 to mi drib Regular polygonal ret. Absent 45-30 1-2 1 2 Mixed opp/alt ~90 to midrib Regular polygonal ret. Present 40-30 1-2 0 2 Mixed opp/alt ~90 to midrib Regular polygonal ret. Present 45-40 1-2 0 2 Mixed opp/alt ~90 to midrib Regular polygonal ret. Absent 45 1-2 1 2 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present? 50-45 1-2 0 2 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 45 2-3 1 2 Regular poly. ret. Regular polygonal ret. Present 40-30 1-2 1 2 Regular poly. ret. ~90 to midrib Regular polygonal ret. Absent 45-40 1-2 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 40-30 1-2 1 1-2 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 60-40 1-2 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Absent 45-40 1-2 0 2 Regular poly. ret. Regular polygonal ret. Present 50-45 1-2 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Absent 50-45 1-2 0 2 Regular poly. ret. Regular polygonal ret. Present 45-30 1-2 0 2 Mixed opp/alt Regular polygonal ret. Absent 40-30 1-2 1 2 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 45-30 1-2 0 2 Mixed opp/alt ~90 to midrib Regular polygonal ret. Present 30-15 0-1 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 35-20 0-1 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 35-20 0-1 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Present 35-30 1-2 0 2 Mixed opp/alt ~90 to midrib Regular polygonal ret. Absent 50-20 1-2 1 1 Regular poly. ret. ~90 to midrib Regular polygonal ret. Absent

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80 APPENDIX A PHOTOGRAPHIC PLATES Plate 1 1. Exemplar, whole leaf. 1

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81 Plate 2 1. Exemplar, whole leaf. 2. Exem plar, toothed margin, close up. 3. Exemplar, areolation, close up. 1 2 3

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82 Plate 3 1. Exemplar, whole leaf. 1

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83 Plate 4 1. Exemplar, whole leaf. 2. Exemplar secondary veins joining mid vein, close up areolation. 1 2

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84 Plate 5 1. Exemplar. 2. Leaf 2. 2 1

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85 Plate 6 1. Exemplar, sterile shoot with leaves. 2. Exempl ar, close up showing leaves 2 1

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86 Plate 7 1. Exemplar, whole leaf. 2. Leaf 2. 3. Leaf 2, Secondary veins at the base, close up. 4. Exemplar petiole, close up. 2 3 4 1

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87 Plate 8 1. Exemplar, whole leaf, asymmetr ical blade. 2. Leaf 2, falcate apex. 2 1

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88 Plate 9 1. Exemplar, leaf frag ment. 2. Secondary junctions w ith primary vein, and toothed margin 3. Leaf 2, decurrent base. 1 2 3

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89 Plate 10 1. Exemplar, leaf frag ment. 2. Exemplar, tertiary veins and laminar resin dots, close up. 3. Whole leaf 2. 1 2 3

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90 Plate 11 1. Exemplar, whole leaf 2. Whole leaf 2. 3. Leaf 2, ter tiary veins and fourth order veins near lateral primary. 4. L eaf 2, higher order veins, close up. 2 3 4 1

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91 Plate 12 1. Exemplar, whole leaf 2. Exemplar, tertiary veins and fourth order veins near primary vein, close up. 1 2

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92 Plate 13 1. Exemplar, whole leaf. 2. Primary vein being deflected at each secondary juncture near apex. 3. Margin, close up. 4. Tertiary vein s and higher order veins near mid vein, close up. 1 2 3 4

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93 Plate 14 1. Exemplar, three lobed leaf 2. Secondary veins on lateral primary. 1 2

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94 Plate 15 1. Exemplar, whole leaf. 2. Secondary vein junction with mid vein and composite intersecondary. 3. Admedially ramified te rtiary veins near mid vein, close up. 1 2 3

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95 Plate 16 1. Exemplar, whole leaf 2. Whole leaf 2. 3. Leaf 2, te rtiary veins and fourth order veins near mid vein. 4. Whole leaf 3. 1 2 3 4

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96 Plate 17 1. Exemplar, whol e leaf. 2. Whole leaf 2. 1 2

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97 Plate 18 1. Exemplar, whole leaf. 2. Secondary veins at the base. 3. Closely spaced tertiary veins near mid vein, close up. 1 2 3

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98 Plate 19 1. Exemplar, whole leaf 2. Secondary and tertiary veins near mid vein. 3. Higher order veins, close up. 1 2 3

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99 Plate 20 1. Exemplar, whole leaf. 2. Sec ondary and tertiary veins near mid vein. 1 2

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100 Plate 21 1. Exemplar, whole leaf 2. Tertiary veins near the ma rgin. 3. Secondary and tertiary veins decurrent on mid vein. 2 3 1

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101 Plate 22 1. Exemplar. 2. Secondary a nd tertiary veins near mid vein. 1 2

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102 Plate 23 1. Exemplar, whole le af. 2. Closely spaced tertiary veins near mid vein, close up. 1 2

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103 Plate 24 1. Exemplar, whole leaf 2. Secondary and tertiary veins near mid vein. 3. Higher order veins and laminar resin glands, close up. 2 3 1

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104 Plate 25 1. Exemplar, whole leaf. 2. Leaf 2, long petiole. 3. Leaf 3, toothed margin. 3 2 1

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105 Plate 26 1. Exemplar, whole leaf. 2. Exemplar toothed margin, close up 3. Whole leaf 2. 3 2 1

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106 Plate 27 1. Exemplar, whole leaf. 2. Leaf 2. 3. Leaf 2, toothed margin, close up. 3 1 2

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107 Plate 28 1. Exemplar, whole leaf. 2. Exemplar toothed margin. 3. Exemplar base, close up. 3 2 1

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108 Plate 29 1. Exemplar, whole leaf 2. Exemplar, exmedial tertiary veins. 3. Exemplar, higher order veins and areoles close up. 3 2 1

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109 Plate 30 1. Exemplar, whole leaf 2. Exemplar, secondary and tertiary veins near mid vein. 1 2

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110 Plate 31 1. Exemplar, whole leaf 2. Exemplar, toothed margin. 3. Exemplar, tertiary and fourth order veins. 3 2 1

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111 Plate 32 1. Exemplar, whole leaf 2. Exemplar, secondary and tertiary veins near mid vein. 1 2

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112 Plate 33 1. Exemplar, whole leaf 2. Exemplar, toothed margin. 3. Whole leaf 2. 4. Leaf 3, secondary and tertiary vein s near mid vein, close up. 1 2 3 4

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113 Plate 34 1. Exemplar, whole leaf. 2. Exemplar, te rtiary and higher order veins near mid vein. 1 2

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114 Plate 35 1. Exemplar, whole leaf. 2. Exemplar, te rtiary and higher order veins near mid vein. 1 2

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115 Plate 36 1. Exemplar, whole le af. 2. Exemplar, secondary veins at the base and pulvinate petiole. 3. Whole leaf 2. 1 2 3

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116 Plate 37 1. Exemplar, whole leaf. 2. Exemplar, tertiary and higher order veins, close up. 3. Leaf 2 1 2 3

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117 Plate 38 1. Exemplar, whole leaf. 2. Exemplar, tertiary veins decu rrent on mid ve in. 3. Leaf 2. 1 2 3

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118 Plate 39 1. Exemplar, whole le af. 2. Leaf 2, emarginate toot h, close up. 3. Leaf 2, agrophic veins near toothed margin. 4. Whole leaf 3. 1 2 3 4

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119 Plate 40 1. Exemplar, whole leaf. 2. Exemplar, toothed margin, close up. 1 2

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120 Plate 41 1. Exemplar, whole l eaf. 2. Leaf 2. 3. Leaf 2, tertiary and higher order veins, close up. 1 2 3

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121 Plate 42 1. Exemplar, whole leaf. 2. Exemplar, te rtiary veins at junction with lateral primary vein. 3. Exemplar, freely ending vein s and laminar resin dots, close up. 1 2 3

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122 Plate 43 1. Exemplar, whole le af. 2. Leaf 2, missing the base. 1 2

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123 Plate 44 1. Exemplar. 2. Att ached leaflets. 3. Specimen 3. 2 3 1

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124 Plate 45 1. Exemplar, cordate base. 2. Leaf 2, toothed margin close up. 3. Leaf 2, craspedodromous secondary veins. 1 2 3

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125 Plate 46 1. Exemplar, whole leaf 2. Exemplar, secondary and te rtiary veins decurrent on mid vein. 3. Exemplar, higher orde r and freely ending veins. 1 2 3

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126 Plate 47 1. Exemplar, whole leaf. 1

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127 Plate 48 1. Exemplar, leaf fragment. 2. Tertiary veins. 3. Higher order veins. 1 2 3

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128 Plate 49 1. Exemplar, whole leaf. 2. Tert iary and higher order veins, close up. 2 1

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129 Plate 50 1. Exemplar. 2. Exemplar, submargi nal vein, close up. 3. Exemplar, secondary veins and laminar resin dots, close up. 1 2 3

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130 Plate 51 1. Exemplar, whole leaf. 2. Exempl ar, higher order veins, close up. 3. Leaf 2. 1 2 3

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131 Plate 52 1. Exemplar, whole leaf. 2. Exemplar, tertiary and fourth order veins joining mid vein. 3. Exemplar, higher order veins and laminar resin dots. 1 2 3

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132 Plate 53 1. Exemplar, whole le af. 2. Exemplar, tertiary veins joining secondary vein. 3. Exemplar, higher order veins, close up. 1 2 3

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133 Plate 54 1. Exemplar, whole leaf. 2. Exem plar, secondary veins joining midrib. 1 2

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134 Plate 55 1. Exemplar, whole leaf. 2. Exemplar secondary and tertiary veins decurrent on midrib. 1 2

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135 Plate 56 1. Exemplar, fragment of palmate leaf. 2. Leaf 2, fragment. 1 2

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136 Plate 57 1. Exemplar, leaf fr agment. 2. Leaf 2, fragment. 1 2

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137

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138 Plate 58 1-3. Paleogeographic distribution of Araceae, maps based on Scotese (2001). 1. Barremian-Aptian in age, Pothoideae subfamily s uggested (this is accordi ng to the most recent classification of Araceae by Keating (2003b; 2004)) (Friis et al., 2004). 2. Coniacian, Lemnoideae (Krassilov, 1973). Campanian, Orontioideae (Kva ek and Herman, 2004; Bogner et al., 2005), Aroideae? (Stockey et al.,2007). M aastrichtian, Pothoideae (Herngreen, 1974; Sarmiento, 1994; Bonde, 2000; Hesse and Zetter, 2007). 3. Paleocene, Philodendroideae and/or Pothoideae (Van der Hammen, 1956; Van der Ha mmen and Garca de Mutis, 1966; Jaramillo and Dilcher, 2001; Hesse and Zetter, 2007); Pothoi deae and Philodendroideae (fossil leaves, this paper); Lemnoideae (McIver and Basinger 1993 ; Stockey et al., 1997); Unknown subfamily (Van Hoeken-Klinkenberg, 1966; Jaramillo et al ., in press). Paleocene-Eocene, Aroideae (Fedotov, 1975). Eocene, Aroideae (Hic key, 1977; Dilcher and Daghlian, 1977); Philodendroideae and/or Pothoideae (Bogner, 1976; Zetter et al., 2001; Hesse and Zetter, 2007). Middle Eocene, Lasioideae (Ceballos-Ferriz and Stockey 1988; Smith and Stockey, 2003); Pothoideae and Aroideae (Wilde et al., 2005); Lemnoideae (Hesse and Zetter, 2007); Late Eocene, Aroideae, (Wilde et al., 2005).

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139 Plate 59 4-7. Anthurioidea wayuuorum gen. nov. sp. nov. Holot ype (ING-NMNH-0902). 4. Whole leaf; the arrow indi cates the location of Fig. 5; scale 3 cm. 5. The arrows point out the two intramarginal veins; notice that the opposite pe rcurrent secondary veins dominate exmedially; higher order veins from irregular polygonal to orthogonal reticulate; scal e 5mm. 6. Drawing of veins, scale 3 cm.

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140

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141 Plate 60 7-11. Anthurioidea cerrejonense gen. nov. sp. nov. Holotype (ING-0804). 7. Fossil leaf preserves part of the middle and the apex of the leaf; scale 10 cm. 8. Ri ght side of the blade, secondary veins running toward margin; tertiary and higher orders veins showing reticulate patterns; the arrow shows the region where the ma rgin is preserved and the location of Fig. 9. 9. The diagonal arrow shows a secondary vein merg ing with the intramarginal vein, which runs very close to margin; th e vertical arrow shows the fimbrial vein running along the margin; scale 5 mm. 10. Close up of reticulate hi gher order veins; scale 5 mm. 11. Drawing of veins, dashed line indicates that apex and middle fragments were not attached so their positions relative to one another are uncertain.

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142

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143 Plate 61 12-22. Exta nt aroid leaves. 12. Anthurium lucens Standl (National Herbarium, #2980121), the species shows two intramarginal veins. 13. A rotundistigmatum Croat (NH #3113554), the arrow shows a secondary vein merg ing with the inner intramarginal vein. 14. A ravenii Croat & R.Baker (NH #1793970), the arrow shows a seconda ry vein ending on the main intramarginal vein and then merg ing on a secondary intramarginal vein very close to margin. 15. A rubrinervium (Sims) G.Don in R.Sweet (NH #2025269), secondary veins merging on an intramarginal vein that runs very close to margin. 16. A cerropelonense Matuda (NH #3093115), the arrows show secondary veins merging with an intramarginal vein that runs very close to margin. Notice dominance of opposite percurrent secondary veins exmedially. 17. Montrichardia arborescens (L.) Schott (NH #3307839). 18. M. linifera (Arruda) Schott (NH #2174078), arrows on the right side show two secondary veins and arrow on the left side shows the intramarginal venation pattern. Notice that the secondary ve ins branch toward margin and form several intramarginal veins. 19. Colocasia esculenta (L.) Schott (NH #3290213), notice that the secondary veins merge bluntly with a straight intramarginal vein. 20, 21 and 22. Cleared leaf of M. arborescens (L.) Schott (NH #3307839). 20. Secondary veins branch toward margin and form several intramarginal veins. 21. Tertiary ve ins decurrent on a secondary vein, higher order veins strongly irregular polygona l reticulate that have wanderi ng courses and connections at variable angles, bar scale 5 mm. 22. Close up of higher order veins, scale 1 mm.

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144 Plate 62 23-26. Montrichardia aquatica gen. nov. sp. nov. Holotype (ING-0904. Figs. 23-26). 23. Fossil leaf preserves partially the base and the anterior division, sc ale 10 cm. 24. Close up of leaf base, scale 5 cm. 25. Close up of right lobule, notice naked ba sal veins, scale 3 cm. 26. Close up of anterior division, margin preserved on th e left side of the blade, scale 5 cm.

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145

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146 Plate 63 27-33. trichardia aquatica gen. nov. sp. nov. Paratypes (ING-0808, Figs. 27-30; INGNMNH-0903, Figs. 31-33). 27. Fossil leaf incomp lete, scale 10 cm. 28. Drawing of veins. 29. The arrow shows a secondary vei n, notice intersecondary venation, scale 3 cm. 30. Tertiary veins merging with a multistranded secondary vein, notice higher order veins strongly irregular polygonal reticulate that have wande ring courses and connections at variable angles, scale 5mm. 31. Arrow on the left side marks a secondary vein branching toward margin; arrow on the right side shows several intramarginal veins, scale 2 cm. 32. Close up of tertiary veins and higher order veins, scale 5 mm. 33. Close up of higher order veins, scale 3 mm.

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147 LIST OF REFERENCES APG-II. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG-II. Botanical Journal of the Linnean Society : 399-436. BAYONA, G., C. JARAMILLO, M. RUEDA, A. PARDO, A. CHRISTIE, AND G. HERNANDEZ. 2004. Important Pale otectonics and chronostratig raphic considerations of the Late Paleocene in the northernmost Ande s as constrained by Pa leocene rocks in the Cerrejn coal mine. In Tercera Convencion Tcnica de la Asociacin Colombiana de Gelogos y Geofsicos de el Petrleo, Bogot. Memorias. BERRY, E. W. 1925. Miocene Araceae rela ted to Caladium from Trinidad. The Pan-American Geologist 44: 38-42. BOGNER, J. 1976. Die systematische Stellung v on Acoropsis Conwentz, einer fossilen Aracee aus dem Bernstein. Mitt Bayer Staatssamml Palaeontol. Hist. Geol 16: 95–98. BOGNER, J., G. L. HOFFMAN, AND K. R. AULENBACK. 2005. A fossilized aroid infructescence, Albertarum pueri gen.nov. et sp.nov., of La te Cretaceous (Late Campanian) age from the Horseshoe Canyon Fo rmation of southern Alberta, Canada. Canadian Journal of Botany 83: 591-598. BONDE, S. D. 2000. Rhodospathodendron tomlinsonii gen. et sp. nov., an araceous viny axis from the Nawargaon intertrappean beds of India. Palaeobotanist 49: 85-92. BOWN, D. 2000. Aroids, Plants of the Arum Fa mily. Timber Press, Portland, Oregon, USA. BRALOWER, T., I. PREMOLI SILVA, A ND M. MALONE. 2006. Leg 198 synthesis: A remarkable 120-m.y. record of climate and oceanography from Shatsky Rise, northwest Pacific Ocean. In T. J. Bralower., I. Premoli Silva., a nd M. J. Malone [eds.], Proc. ODP, Sci. Results, 198: 1-47. BREMER, K. 2000. Early Cretaceous linea ges of monocot flowering plants. Proceedings of the National Academy of Sciences, USA 97: 4707–4711. BURNHAM, R. J. 1989. Relationships between standing vegetation a nd leaf litter in a paratropical forest: implications for paleobotany. Review of Palaeobotany and Palynology 58: 5–32. BURNHAM, R. J. AND A. GRAHAM. 1999. The hi story of Neotropica l vegetation: new developments and status. Annals of the Missouri Botanical Garden 86: 546–589. BURNHAM, R. J., N. C. A. PITMAN, K. R. JOHNSON, AND P. WILF 2001. Habitat-related error in estimating temperatures from leaf margins in a humid tropical forest. American Journal of Botany 88: 1096-1102.

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BIOGRAPHICAL SKETCH Fabiany Herrera was born in 1978 in Bucaramanga, Colombia. The youngest of three children, he grew up in the city of his origin graduating from the public high school “INEM” in 1995. After one year of public service from 1996 to 1997 he started his B.S. program in geology with emphasis in paleontology in 1998 from the I ndustrial University of Santander, and obtained his degree in 2005. He was an awardwinner with the distinction “Thesis of the Year” from the Colombian Geological Society in 2004. The topic of his thesis was the discovery of the world’s oldest Neotropical rainforest. Upon graduating in May 2005, Fabiany worked as research assistant in biostratigraphy for the Colombian Institute of Petroleum in Bucar amanga. Then, he moved to the Republic of Panama in Central America to work for an internship at the Smithsonian Tropical Research Institute and later at the National Museum of Natural History, Smithsonian Institution in Washington DC where he worked on paleobotany topics. During that time, he spent several months collecting fossils in Co lombia, Panama and Wyoming. In 2006 Fabiany moved to Gainesville, Fl in orde r to attend to the Univ ersity of Florida to get his master’s degree in science with ma jor in geology. Upon completion of his masters program, Fabiany wants to continue in graduate school as a PhD st udent, and aspires to continue his contributions to the scientific world.