P~leocene Flora of the 

Rocky Mountains and 

Great Plains 

By ROLAND W. BROWN 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 

A study of IJO kinds of plants ant! the strata 

that yield them,. showing how they apply in 

the delimitation of the P aleocef!e series 

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1962 
UNITED STATES DEPARTMENT OF THE INTERIOR 

STEWART L. UDALL, Secretary 

GEOLOGICAL SURVEY 

Thomas B. Nolan, Director 

The U.S. Geological Survey Library has cataloged this publication as follows: 

Brown, Roland Wilbur, 1893-1961. 

. ~Paleocene flora of the Rocky Mountains a;nd Great Plains. 

Washington, U.S. Govt. Print. Off., 1961. 

iii, 119 p. illus., diagrs. 

Professional paper 375) 

Bibliography: p. 97-106. 

29 em. (U.S. Geological Survey. 

1. Paleobotany-Paleocelle. 2.· Paleobotany-Rocky Mountains. 

3. Paleobotany-Great Plains. I. Title. (Series) 

For sale by the Superintendent of Documents, U.S. Government Printing Office 

Washington 25, D.C. 
CONTENTS 

Abstract __________________________________________ _ 

Introduction-------------------~------------------- Geological background _________________________ _ 

Acknowledgments ______________________________ _ 

Historical review of the subdivision of the "Great Lig- nite" _______________________________________ _ 

Early observations _____________________________ _ 

The lignitic strata from 1890 to 1912 _____________ _ 

Effect of the discovery of mammals in the lignitic 

strata---------------~-----------------------

The Cannonball marine strata ___________________ _ 

Formula for locating the Cretaceous-Paleocene con- tact ________________________________________ _ 

Paleocene-Eocene contact _______________________ _ 

Summary of Paleocene stratigraphy __________________ _ 

Limits of the Paleocene in specific areas _____________ ~ _ 11ontana _____________________________________ _ 

Canada, North Dakota, a·nd South Dakota _______ _ 

VVyoming--------------------------~-----------

Colorado and New Mexico ______________________ _ Arizona _______________________________________ _ 

Utah _________________________________________ _ 

Gulf Coast _____________________________ - _____ --

Atlantic Coast __________________________ - _____ --

Pacific Coast __________________________________ _ 

Page 

1 

1 

1 

3 

3 

3 

5 

7 

8 

9 

11 

11 

14 

14 

18 

20 

23 

27 

27 

27 

28 

28 

Limits of the Paleocene in specific areas-Continued 

Arctic regions _________________________________ _ 

Europe _______________________________________ _ 

Asia __________________________________________ _ 

South America ________________________________ _ 

Africa ________________________________________ _ 

Paleocene localities _________________________________ _ 

Methods applied in the study of the plants ____________ _ 

The Paleocene flora ________________________________ _ 

Systematic description of the megascopic flora _________ _ 

Seedless plants ________________________________ _ 

Seedplants------------------~-----------------

Objects of uncertain classification ___________ . _____ _ 

Miscellaneous leaves, fruits, seeds, flowers, roots, and 

probable animal remains ______________ ..:. _______ _ 

Other objects of uncertain or mistaken identity ____ _ 

The microscopic plants _____________________________ _ 

Fossil wood _______________ · ________________________ _ 

Suggested rejection of some names ____________________ _ 

Composition and ecologic significance of the Paleocene 

flora ___________________________________________ _ 

Paleocene animals of the Rocky Mountains and Great 

Plains __________________________________________ _ 

References cited ___________________________________ _ 

Index ____________________________________________ _ 

ILLUSTRATIONS 

PLATES 1-3. Views of Paleocene strata. 

4-68. 1iega;_scopic Paleoce~e plants. 

69. Megascopic Paleocene fossils. 

[Plates 1-69 follow index] 

,. 

FIGURE 1. Diagrammatic cross. section from Sheridan, 

· VVyo., 'to Mandan,- N. Dak_, showing correlation of Cr.etaceous and Tertiary formaPage 

28 

29 

29 

29 

30 

30 

35 

38 

39 

39 

47 

88 

90 

92 

92 

93 

94 

95 

96 

97 

107 

tions as interpreted by the writer_ _ _ _ _ _ page IV 

m 
AGE 

LLJ 

z 

LLJ 

u 

0 

LLJ 

LLJ 

z 

LLJ 

u 

0 

LLJ 

.....J 

< 

a.. 

en 

::::l 

0 

LLJ 

u 

< 

1-

LLJ 

0::: 

·.• 

~~ 

SHERIDAN AND 

BUFFALO 

WYOMING 

~f~~~~:~~·: f. ~ ........ Uim coal group 

~:: .... .. •• tiD 

~.=: Wasatch 

.=:.::.o:·.·.·.······ 

~··· De Smet (part) oo.:o·o·.o.-:.-....... . 

Fort Union 

BROADUS AND· 

POWDERVILLE 

MONTANA 

.·.·:·::-·.-· ......... . 

ROSEBUD AND 

MILES CITY 

MONTANA 

SENTINEL BUTTE 

AND MARMARTH 

NORTH DAKOTA 

OLIGOCENE-White River 

GRASSY BUTTE 

AND HEBRON 

NORTH DAKOTA 

STANTON 

AND MANDAN 

NORTH DAKOTA 

FmURE i.-Diagrammatic cross section from Sheridan, Wyo., to Mandan, N. Dak., showing correlation of Cretaceous and Tertiary formations as interpreted by the writer. The Broadus and Rosebud 

columns· are interchanged to show closest relations to adjacent columns. Stippling indicates dark wnes of sttata of all kinds. Unstippled parts are light-colored wnes. (After Brown, 1948a.) 
PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

By RoLAND W. BRoWN 

ABSTRACT 

The description of the Paleocene series as a whole and particularly the determination of its lower and upper limits in the 

Rocky Mountains and the adjacent Great Plains have been 

matters of practical interest since the late 1850's. As this 

series contains much of the coal in that area, its mapping for 

the purpose of classifying the public lands has necessarily involved consideration of its boundaries; but, until recently, no 

general agreement prevailed about the position of these limits. 

Since 1929 a reexamination of the Paleocene strata has resulted 

in the collection of much additional and more precisely located 

fossil material and in the conclusion that the CretaceousPaleocene boundary is at the base of the first coal zone not 

very far above the latest remains of dinosaurs. Representative collections of fossil plants from above and below this level 

can usually be readily distinguished, but fragmentary collections may not always be distinctive. More species crossed the 

boundary than was formerly supposed and indicate that no 

radical break in the floral sequence' occurred, just as, at many 

localities, the stratigraphic succession continued unbroken from 

the Cretaceous into the Paleocene and beyond. 

The upper limit of the Paleocene, the Paleocene-Eocene 

boundary, can be detected locally by the presence of an obvious 

unconformity and of distinctive mammalian fossils. The flora 

of the early Eocene, however, except for a species of the fern 

Salvinia and perhaps a few others, does not seem to be greatly 

different in general composition from that of the late Paleocene. 

The restudy of the Paleocene flora has revealed many synonyms and has thus reduced the variously published estimates 

of its size to a tentative list of 170 recognizable species or 

forms. The presence of palms and breadfruit on the one hand 

and of birches, hazels, maples, and ashes on the other suggests 

either that remains from different ecologic habitats in a warm 

temperate to temperate environment mingled or that some 

Paleocene species were adapted to environmental niches different from their comparable living descendants or relatives. 

INTRODUCTION 

GEOLOGICAL BACKGROUND 

This paper is the outgrowth o£ an investigation 

I was asked to make when I joined the staff of the 

U.S. Geological Survey in 1929. W. C. Mendenhall, 

then chief geologist, specifically instructed me as follows: "Your job, in part, will be to solve the longdebated problem of the Cretaceous-Tertiary contact 

in the western interior. In doing this you will be 

concerned with the "somber beds" and their fossils, 

principally in the Dakotas, Montana, Wyoming, and 

Colorado." That summer, and many succeeding field 

seasons, often and for long intervals interrupted by 

other work, found me mapping and exploring the 

strata involved in this problem. 

Eastward for distances of as much as 500 miles 

from the foothills of the Rocky Mountains, the Great 

Plains area, extending from central-western Canada 

to New Mexico, is underlain beneath its Quaternary 

or other cover by Cretaceous and Tertiary coal-bearing strata. ·Erosion has carved this area into low, rolling hills, wide and narrow valleys, and in some localities, fantastic and colorful badlands (pis. 1-3). 

The marine and nonmarine strata thus exposed, and 

once collectively called the "Great Lignite," aggregate thousands of feet in thickness and comprise soft 

or indurated, sometimes conspicuously crossbedded 

sandstones; carbonaceous, calcareous, and· sandy shailes; 

clays; and coal beds, in more or less repetitive sequence. A diagrammatic cross section showing the 

stratigraphic units involved is given in figure 1. 

Comparable correlative strata occur, in somewhat 

isolated patches, high in the Rockies close to the con-. 

tin ental divide in North, Middle, and South Parks, 

Colo.; in the Bighorn Basin of Wyoming and its 

northwestward extension in Montana; in the Wind 

River Basin of Wyoming; and on the west side o£ the 

Rockies from :M:exico northward through the San Juan 

Basin of New Mexico and Colorado, and from southwestern to eastern Utah, northwestern Colorado, and 

western Wyoming. 

Seen close at hand on a drab or rainy day, most 

exposures of the nearly horizontal lignitic strata above 

the Fox Hills sandstone (Upper Cretaceous) in eastern Montana, eastern Wyoming, and adjacent areas 

are monotonously similar to one another. Their general color tone is dull, gray or even dark, from which 

they have been characterized as "somber beds." The 

subdued or dark colors of the carbonaceous shales 

and coals contrast with the lighter colored, yellowish 

sandstones and, here and there, with the pinks or brilliant reds of the baked and sometimes fused rock 

(Collier and Smith, 1909, p. 45; Rogers, 1918) above 

burned coal beds. The coal outcrops were ignited for 

the most part spontaneously, but sometimes perhaps 

1 
2 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

by lightning or by fires resulting from human negligence. The uninitiated often mistake this clinker or 

scoria for volcanic rock.1 

From a distance, however, and on a clear, bright 

day, thick, well-exposed sequences above the Fox Hills 

sandstone in any part of the area, but particularly in 

the eastern part of the Yellowstone Valley, can be 

divided roughly by eye into a number of contrasting 

light and dark zones beginning with a dark zone at 

the bottom. This lowest zone is coal barren except at 

the very base and is chiefly brownish sandstone or 

sandy clay characterized by a tinge of greenish rustiness and by the fact that it contains the last, and 

locally abundant, remains of the horned dinosaurs. 

Above this dark zone is a light-colored, yellowish sequence consisting principally of sandstones, shales, and 

coal beds. This is overlain by a dark zone, whose 

duskiness can best be described as livid, or grayish 

black. Near its base is a thick coal bed or carbonaceous shale-the Big Dirty-which is continuous over 

many square miles and serves well locally as a stratigraphic 1narker. This dark sequence is capped by a 

notable light-colored, yellowish group of sandstones, 

clays, and coals, making a zone that is conspicuous 

on all the higher hills and peaks along the Yellowstone and its tributaries from Forsyth to Sidney, 

Mont. Overlying this light-colored zone is a somewhat darker lignitic sequence exposed typically at 

Sentinel Butte, ·N. Dak., and at the original type 

section of the Fort Union formation on the north 

side of the Missouri opposite the mouth of the Y ellowstone. Perhaps no better description of all these 

conditions and circumstances has been given than that 

by Leonard ( 1908; 1911). For further details on speeific areas, the reader is referred to the U.S. Geol. 

Survey bulletins on the coal fields of the western 

interior, the latest list of references being by W. W. 

Olive (1957, p. 5), and W. C. vVarren (1960, p. 563). 

To the west in the direction of the source of the 

sedin1ents that became these strata, the older light and 

1 In his lively and entertaining book, "A Ghost Town on the Yellowstone," Elliot H. Paul described th~ extinct town of Trembles, in 

existence for about 15 years (1907-22) on the left bank of the Yellowstone River, 6 miles southwest of Sidney, Mont. He painted a stirring 

word-picture of the landscape from Glendive to the mouth of the 

Yellowstone but repeated two commonly held errors, namely, that the 

red rocks of the badlands are volcanic (p. 32) and that the lower end 

of the valley, in relatively recent prehistoric days, was a favorite 

haunt of dinosaurs (p. 6). The red rock, as explained, is clinkered 

sedimentary shale and sandstone. The uppermost Cretaceous strata 

that contain remains of the last dinosaurs-trachodon, triceratops, and 

tyrannosaurus-crop out in the badlands on the flanks of the Cedar 

Creek anticline near Gll'lndive but dip eastward to hundreds of feet 

below the surface at Sidney and the mouth of the Yellowstone where 

they are covered by Paleocene rocks. Therefore, the dinosaurs once 

existing in that area wandered around on a terrain antedating the 

present surface of the Yellowstone Valley by at least 60 or 70 million 

years. 

dark zones for the most part lose their distinguishing 

colors and· merge into a greenish or gray sandy sequence with little or no coal. In the easternmost 

areas farthest from the source of the sediments, color 

distinction between these zones is also either impossible or very unsatisfactory. 

Except that they are lignitic and may be sombercolored, the correlative strata in some adjacent regions cannot be so readily characterized. Each area 

has variant lithologic and structural features peculiar 

to itself. 

The mapping and classification of these lignitic 

strata were accompanied by much speculation and 

discussion about the respective limits of the several 

postulated large geologic units and particularly about 

the position of the Mesozoic-Cenozoic boundary, which 

was suspected early of being present somewhere in the 

sequence. Four reasons for the warm debate about 

these matters can be cited: 1, Among the earlier investigators there was a tendency to generalize without 

completely understanding the significance of the limited data in hand. 2, There was confusion, pointed out 

by Marvin in 187 4, in considering lithologically similar 

strata, although geographically distant from one another, as contemporaneous and correlative. 3, The 

third reason was the dogma that unconformities of 

considerable magnitude are necessary to mark systemic and series boundaries. 4, No good formula, applicable in the field, as to the probable position of 

the ~fesozoic-Cenozoic boundary, was proposed until 

1912. This was not tried out deductively and unanimously by all mappers in the pertinent areas but was 

more or less neglected until recently. 

On the map by Brown (1949) both the lower and 

the upper limits of the Paleocene strata of the Rocky 

Mountains and Great Plains are shown in the light 

of reconnaissance studies to that date. Since then, 

more detailed mapping by others indicates that in 

some localities the boundary lines need minor readjustment. 

The principal assumption in this study is that the 

last dinosaurs in the Rocky ~fountains and Great 

Plains region vanished some time before the extreme 

end of the Cretaceous and that some species of plants, 

now regarded as strictly Paleocene in age, left no 

apparent antecedent records in that region in the Cretaceous. Nevertheless, I admit the possibilities that 

fragmentary dinosaur bones may have been reworked 

into Paleocene deposits and that antecedents of the 

Paleocene plants may be found in the Cretaceous. 

Unless the principal assumption be gran~ed as a factual, working hypothesis I can at present see no 

other satisfactory, concrete basis for agreement on a 
HISTORICAL REVIEW OF THE SUBDIVISION OF THE "GREAT LIGNITE" 3 

solution of the Cretaceous-Paleocene (or ~fesozoicCenozoic) dating problem. 

ACKNOWLEDGMENTS 

The help of my field assistants is gratefully acknowledged, as follows: 1{. J. Murata, 1931; F. S. 

MacNeil, 1936; C. vV. Mumm, 1938; T. D. Lance, 

1939; C. E. Staudte, 1940; and H. R. Christner, 1952. 

T. W. Stanton visited localities in North Dakota 

with me in 1931. C. E. Dobbin accompanied me to 

localities in Wyoming, ~fontana, and North 'Dakota 

in 1936 · in Colorado in 1938 and 1939; and again 

with Ge~rge Horn, in northeastern Wyoming in 1948 

and 1950. F. ~f. Van Tuyl and J. H. Johnson also 

auided me to localities in Colorado in 1938 and 1939. 

Bryan Patterson visited localities in the vicinity of 

Rifle and DeBeque, Colo., with me in 1939 and C. L. 

Gazin examined Colorado localities with me in 1940 

and 1941. Chiefs of U.S. Geological Survey. field 

parties who helped me in one capacity or another 

·were: W. G. Pierce, Rosebud coal field, Montana, 

1929 and Cody, Wyoming, 1940; C. B. Hunt, Mt. 

Taylor coal field, New Mexico, 1930; F. W. Parker, 

Mizpah coal field, Montana, 1931 ; D. A. Andrews, 

Lovell, Wyo., 1936; vV. H. Bradley, Rock Springs, 

Wyo., 1941; R. P. Bryson, southern Montana, 1940, 

1941, 1946; C. H. Dane, Durango, Colo., and Cuba, 

N. Mex., 1940; ,V. C. Warren, Broadus, Mont., 1940; 

T. A. Hendricks, Billings, 1\font., 1944; C. E. Erdmann, Great Falls, Mont., 1944, 1947, 1948, 1949; W. E. 

Benson, Hebron, N. Dak., 1947, 1951; R. W. Lemke, 

Minot, N. Dak., 1947, 1955; R. _C. Townsend, Crosby, 

N.Dak., 1947; Garland Gott, Culbertson, Mont., 1947; 

Fred Jensen, Fort Peck Dam, Mont., 1948, 1950; J. D. 

Love and Raymond ~1. Thompson, ·Lander, Wyo., 

1944-51; ,V. T. Pecora, Havre, 1\font., 1948-55; W. W .. 

Olive, Spotted Horse coal field, Wyo., 1948; W. J. 

Mapel, Buffalo, Wyo., 1949; N. M Denson, Buffalo, 

S. Dak., 1950; P. W. Richards, Billings, Mont., 1950; 

Roger Colton, vVolf Point, Mont., 1949-51; R. M. 

Lindvall, Big Sandy, Mont., 1949-53; N. W. Bass, 

Glenwood Springs, Colo., 1951; George Pipiringos and 

Harold Masursky, Wamsutter, Wyo., 1952-53; M. R. 

J{lepper, Boulder, Mont., 1953-54; and D. M. Kinney 

and ,V. J. Hail, North Park, Colo., 1956. 

Local residents were uniformly cordial and helpful, 

but in particular the following: George Fountain, 

Circle, Mont.; Dell and Edwin Lewis, Glendive, 

Mont.; George Newlin, Ashland and Billings, Mont.; 

,V. H. Peck, Ekalaka, ~font.,. and A. C. Silberling, 

Harlowton, Mont. In the office and elsewhere many 

colleagues have contributed valuable criticism during 

this investigation. The photographs of specimens were 

made chiefly by N. W. Shupe. 

. Unless otherwise stated all the photographed specimens here reproduced are deposited in the U.S. N ationa! Museum. 

HISTORICAL REVIEW OF THE SUBDIVISION OF THE 

"GREAT LIGNITE" 

EARLY OBSERVATIONS 

The evolution of ideas about the appropriate division of this great group of lignitic strata into identifiable, mappable, and datable parts, according to the 

general geologic column, is a long story involving 

many actors and the interplay of much fact, fancy, 

and opinion. Accounts of this development may be 

found in the writings of G. ~1:. Dawson (1875, p. 182-

202), L. F. Ward (1886, p. 406-436), G. P. Merrill 

(1906, p. 64~~658), F. H. Knowlton (1922 a, p. 3-81), 

L. S. Russell (1932 a, p. 121-156; 1953, p. 106-113); 

R. L. Nace (1936, p. 66-87), and E. M. Spieker (1946, 

p. 135, 142-160). As my immediate concern here is 

to produce and discuss the evidence that purports to 

establish the position of the boundary limits of the 

Paleocene series only, I shall not attempt to recount 

all the details of the progress in classifying the entire lignitic sequence. I shall not c~te all the literature, but shall confine myself to a statement of only 

those opinions and decisions that re~ulted in positive 

action or that had a distinct influepce toward clarification in delimiting the Paleocene series. 

In 1854 and the immediately following years, F. V. 

Hayden a~d his colleagues explored large parts of the 

coal-bearing terrain of the Rocky Mountains and 

Great Plains, particularly the regiop accessible from 

the Missouri and Yellowstone Rivers in Montana 

(Hayden, 1861; Raynolds, 1868). They accumulated 

fundamental data about the genera~ stratigraphy and 

made collections of fossil vertebrates, invertebrates, 

and plants. Because of the coal ~earns, }Iayden at 

first called these strata the "Great Lignite" group, 

but later he and his chief collaborator, F. B. Meek, 

substituted for it the more specific name, Fort Union 

group (Meek and Hayden, 1862, p. 433). More particularly, this term applied to outcrops, now known 

to be only a part of the "Great Li~ite," on the north 

side of the Missouri River between ·the present Snowden, Mont., and Buford, N. Dak. (pl. 1, fig. 1) .

2 

2 Fort Union, located on what is now the Montana side of the line, 

was erected by the Northwest Fur Co. in the 1820's. In 1867 when 

the fort was razed, some of the material was ,Purchased by th~ Fe?-

eral Government and used in building Fort Btiford across the hne Ill 

North Dakota. See "Tales from Buffalo land. The story of Fort 

Buford," by Usher L. Burdick, 1940, Wirth Brothers, Baltimore, Md. 
4 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Among Hayden's collections in 1854 were dinosaurian remains from the Judith River region of 

Montana; and in. 1872 Meek found part of a saurian 

skeleton ( Agathaumas sylrvestris Cope) in lignitic 

strata near Black Buttes Station, Wyo. In consequence there was general agreement that the older 

parts of the "Great Lignite" are Cretaceous in age. 

The fossil plants taken from the upper part of 

the "Great Lignite" by Meek and Hayden were identified by J. S. Newberry in 1868 as Tertiary in age. 

Thus, at an early stage of exploration and discovery 

in the western interior States and adjacent parts of 

Canada, geologists recognized that the Mesozoic-Cenozoic boundary lay somewhere between the older dinosaur-bearing strata and younger plant-bearing strata 

of the great lignitic sequence. 

In 1876, Clarence King, without designating a specific type locality, introduced the term "Laramie" for 

lignitic beds on the Laramie Plains of Wyoming and 

thus, although denying identity of the Laramie and 

Fort Union strata, unwittingly complicated the Cretaceous-Tertiary boundary discussion because shortly 

the "Great Lignite" of Hayden (including the Fort 

Union group) was being referred to loosely as the 

Laramie group. Not until 1910 were the terms Laramie and Fort Union divorced, and Laramie was restricted to strata conformably overlying the Fox Hills 

sandstone in the Denver Basin of Colorado. This 

Laramie was conceded by all to be Late Cretaceous in 

age. 

The paleobotanist Leo Lesquereux had no clear idea · 

about the age of the "Great Lignite," for his publication, "The Tertiary flora" (1878), included notice 

of many plants now known to have come from Cretaceous strata. Indeed, there was a vigorous exchange 

between Lesquereux and Newberry concerning the 

age of some lignitic strata in New Mexico and adjoining portions of Colorado, Newberry insisting upon 

Cretaceous and Lesquereux upon Tertiary age. Newberry (1888, p. 329) strongly maintained however, 

that "The For~ Union group * * * in my judgment, 

should be· considered the basal member of the Tertiary." 

As late as 1883, C. A. White, on the evidence of 

nonmarine invertebrates, regarded the "Great Lignite" as transitional between the Cretaceous and Tertiary and, therefore, partaking of both. This idea was 

accepted by the paleobotanist Lester Ward ( 1886, p. 

435) who, after examining the opinions of Newberry, 

Lesquereux, Cope, White, and others, commented : · 

The discussions with regard to the age of the Laramie group 

which have been rapidly passed in review have, perhaps, sufficiently shown that it is impossible to refer that group either 

to the Cretaceous or to the Tertiary and in so doing harmonize all the facts that the group presents with those in conformity with which other deposits in other countries of the 

world have been so referred; but they have also sufficiently 

shown that this is not the fault of the investigators, but so to 

speak, of the facts, and the real disagreement is in the organic 

forms and the nature of the deposits, so that omniscience itself 

could never harmonize them with all kinds of forms and deposits in all parts of the world. It is, therefore, futile and 

indeed puerile, longer to discuss this question, and we can well 

afford to dismiss it altogether and settle down to the more 

serious study of the real problems which still lie before us. 

A~t~r sounding this discouraging note, Ward, not 

realizing the age significance of the compositional 

differences he recognized between the floras of the 

"Wyoming and Colorado Laramie" and the "Fort 

Union Laramie" (1886, p. 438, 439, 1887, p. 1; Newberry, 1890, p. 529), but still persuaded that "Laramie" was synonymous with "Great Lignite" and indivisible into formations of different ages, proceeded 

to con~use the situation still further hy lumping plant 

colle?twns from many localities and widely spaced 

stratigraphic levels in his "Types of the Laramie 

flora" ( 1887). This "Laramie" flora, as we now know . . . ' IS a composite of Items representing floras from Mesaverde (Upper Cretaceous) to Wasatch (Lower Eocene). Newberry ( 1888, p. 329), who was probably 

the first to perceive a marked difference between the 

floras of the lower and upper parts of the "Great 

Lignite," referring to Ward's paper, remarked that: 

It [the "Laramie" flora] contains few of the Lower or true 

Laramie plants, and is chiefly descriptive of the Upper Laramie 

or Fort Union flora, of which it affords the most important 

review yet published. The flora of the Lower or true Laramie 

has been made known to me by collections from * * * Black 

Buttes, Walsenburg, Canon City * * *, etc. These show that 

there is almost nothing in common between the Fort Union 

and Laramie floras, and that the two divisions of the so-called 

Laramie group, judging from the fossil plants, which are very 

numerous, must be regarded as distinct formations. 

This opinion was reiterated in. 1890 (p. 525). In 

1940 a~d 1942 Erling Dorf published a preliminary 

comparison of the Lance and Fort Union floras showing that, with increased knowledge of these floras, 

~ ewberry's original opinion is in large part still pertinent. 

The reasons for the general mixed state of ·opinion 

about the "Great Lignite" in 1890 lay, contrary to 

Ward's statement, not with the facts, but with the 

interpreters a~d a ppliers of the facts. The paleo-

. botanists, for example, had not worked out or agreed 

upon a satisfactory American standard of comparison 

to cover the floras in question, hut, with the exception 

of Newberry, as just stated, per~isted in assuming 

that the flora of the "Great Lignite" is an indivisible 

unit. Furthermore, no geologist or paleontologist proposed and applied a good working formula for iden-

,J 
HISTORICAL REVIEW OF THE SUBDIVISION OF THE "GREAT LIGNITE" 5 

tifying the probable position of the Mesozoic-Cenozoic boundary in the western interior States and 

Canada. Some of the critical stratigraphic sections 

also were not carefully measured and compared. Here, 

to cite an example, 'Vard erred in regard to the geologic situation on the west flank of the Cedar Creek 

anticline, 12 miles southwest of Glendive, Mont. In a 

discussion of a paper by Newberry (1890, p. 531) 

he said: 

I have one fact of my own observations which may be worth 

stating and which may not be known to all. About 15 miles 

above the town of Glendive, on the right bank of the lower 

Yellowstone river, there is a cliff, known as Iron bluff, which 

is colored very bright red from having the carbonaceous matter burned out, and· which is full of fossil plants ... This bluff 

is right on the bank of the Yellowstone river, and the railroad 

cuts through it, which makes the cliff conspicuous. Immediately below (east) there is a short anticline (Cedar Creek 

anticline), and apparently a little island about a mile in extent, 

filled with characteristic Fox Hills Cretaceous fossils. I have 

been on the ground and collected large numbers of them, and 

everywhere we meet with them: the wheels of the wagon as 

one drives over them crush the shells, so abundant are they; 

and there is no doubt that this is a typical Fox Hills bed, in 

Dr. White's understanding of the term "Fox Hills." Now, so 

far as I can tell, and so far as he could tell from a careful 

study of the ground, this Iron bluff deposit-this Laramie or 

Fort Union leaf-bed-rests directly and immediately upon the 

Fox Hills bed. If there is any difference of age there is no 

indication at that point * * *. 

The sequence of strata at Iron Bluff, as measured 

by me in 1936, is as follows: 

Section on the west flank of Cedar Creek anticline, 

12 miles southwest of Glendive, Mont. 

4. Fort Union formation : 

Thickness 

(feet) 

Light-colored shale and sandstone, coal, plants___ 50+ 

Somber carbonaceous shale, sandstone, coal, 

plants -------------------------------------- 224 

Gray shale, light-colored sandstone, coal, lowest 

bed locally clinkered, plants, fresh-water shells_ 170 

3. Hell Creek formation : 

Rusty green to gray carbonaceous shales and 

sandstones, three thin coal seams in basal part, 

dinosaur bones, fragmentary plants __________ 385 

2. Fox Hills sandstone : 

Gray to light-colored shales and sandstones, 

capped by the white Colgate member, apparently barren of fossils at this locality ________ 140 

1. Pierre shale : 

Dark-colored marine shales, with concretions full 

of fossils ------------------------------------ 75+ 

Base of section. 

Ward's "Fox Hills" with Cretaceous fossils is thus 

seen to be what.is now called the Pierre shale, and the 

lowest burned coal bed with plants is near the base 

of the Fort Union formation. Therefore, instead of 

resting directly upon the fossiliferous marine Pierre 

shale, as averred by Ward, this clinkered bed is separated from it by at least 525 feet of Fox Hills and 

Hell Creek strata, the latter containing diagnostic 

dinosaurian remains. 

THE LIGNITIC STRATA FROM 1890 TO 1912 

F. H. Knowlton succeeded Ward on the U.S. Geo- · 

logical Survey staff as paleobotanist in the study of 

Mesozoic and Cenozoic plants, and from the early 

1890's until his death . in 1926 greatly influenced the 

opinions pertaining to the age of the "Great Lignite." 

He, like his Survey predecessors, made a number of 

field trips into the disputed areas and obtained numerous collections. Stanton and he ( 1897), after a study 

of the terrain and fossils on Lance Creek, · in eastern 

Wyoming, differentiated the Upper Cretacequs Lance 

formation, with its primitive mammals, dinosaurs, nonmarine invertebrates, and plants, from the overlying 

Fort Union formation with its invertebrates and flora, 

but no dinosaurs. Had this initial example been followed in. subsequent years during the mapping and 

correlation of strata in adjacent areas, there might 

have been .little difficulty about the Cretaceous-Tertiary contact; but in areas to the north, particularly 

in northeastern Wyoming, Montana, and the Dakotas, 

strata that looked lithologically similar to those in 

the upper part of the Lance, except that they contained coal seams, a Fort Union flora, and lacked 

dinosaurs, were assumed to be Lance. In many papers 

they were called Lance or Upper Lance (Eocene~ ) , 

with much resultant confusion. This condition was 

aggravated by Stanton's misidentification of the Cannonball marine strata in the Dakotas as Lance (Upper Cretaceous) in age (Lloyd, 1914, p. 250; Stanton, 

1921, p. 18). A correlation sketch that illustrates these 

opinions and practices was published by Winchester 

and others (1916, p. 15, fig. 2). A similar sketch by 

me (1948a, p. 1266, fig. 2) reproduced here as figure 

1, shows the present conception of the same situation. 

The mappers in Montana and the Dakotas had by 

this time begun to recognize and name the local 

zones of light-colored and dark-colored strata in the 

upper part of the "Great Lignite." The designation 

of these zones, beginning with the lowest and oldest, 

resulted eventually as follows: Hell Creek formation 

(dark) ; Tullock member (light) ; Lebo member 

(dark); Tongue River member (light), which for a 

long time was itself erroneously considered synonymous with Fort Union or upper Fort Union; and 

Sentinel Butte member (dark). This nomenclature 

was found convenient in local mapping, but most observers discovered that color contacts between some 

of the contrasting zones are evanescent and migra-
6 PALEOCENE l!..,LORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

tory, both verticaUy and horizontally (Rogers, 1913b, 

p. 719-721; Pierce, 1936, p. 60; Brown 1948a, p. 

1270-1271). This is particularly noticeable from the 

vicinity of Miles City toward Glendive and from 

Miles City toward Ekalaka and Baker, Mont., and 

Marmarth, N. Dak. In those directions, zones recognized as more or less distinct west of Miles City 

become increasingly difficult or impossible to separate. 

For this reason th~ Lebo and Tullock in the eastern 

part of the area w_ere lumped as the Ludlow lignitic 

member. In the vicinity of Glendive· the base of the 

otherwise light-colored Tullock equivalent appears 

darker than the underlying Hell Creek formation because its strata consist of closely spaced dark carbonaceous shales and coal seams. 

In 1909 l{now ltqn and Stanton separately published 

preliminary opiniOJlS on these strata. Knowlton (1909, 

p. 180-193, 237) divided the sequence above the Fox 

Hills sandstone i~to an "upper" and "lower" Fort 

Union. His "upp~r" part, a light-colored unit, corresponded to that now called the Tongue River member. His "lower" ·_part comprised the "somber beds" 

underlying the T~;mgue River member; that is, the 

Lebo member, Tullock member, and the "Ceratops 

beds" or Hell Creek formation, as now known. In 

thus lumping the dinosaur-bearing Hell Creek with 

the remainder of the non-dinosaur-bearing "somber 

beds" he originated an error that led to and was 

comparable with run other made by the mappers, namely, 

in applying the tt:~rm "Upper Lance" to the "somber 

beds" (chiefly the Tullock member) overlying· the Hell 

Creek. Iri consequence of all this, Knowlton did not 

distinguish the flora of the Hell Creek and of the 

correlative Lance in Wyoming from that in the remainder of the ."somber beds" and "upper" Fort 

Union, but regarded all as a unit, Tertiary in age. 

This position, so far as I know, he never changed 

(Knowlton 1911, :P· 358; 1914, p. 325; 1922a, p. 31). 

Thus, the confusion of floras remained much as Ward 

left it in 1887. . . 

It is now evident that Knowlton's conception of the 

stratigraphic sitl~~tion along the lower reaches of the 

Yellowstone V all~y was not sound in that he did not 

differentiate the iwo dark zones of his "lower" Fort 

Union,- the Lebo ~nd Hell Creek, and did not realize 

that strata now called Tullock lie bet"·een the Lebo 

and Hell Creek. Thus, concerning the outcrops at 

Miles City and Glendive, respectively, he said (1909, 

p. 186) : 

With the excepti~m of some unimportant alluvial deposits in 

the valleys, all the· :rocks of this region [around Miles City] 

were found to belo,ng to the Fort Union formation, which is 

comprised of two members. The lower member, which corresponds to the "Hell Creek beds," is about 500 feet in thickness, 

but the base is not exposed ; however, from the comparison of 

the Miles City section with that at Glendive, it is inferred that 

the base is not very deeply bul'ied. 

What are the facts about the correlation of the sections at Miles City and Glendive? The strata exposed in the bluffs on the left bank of the Yellowstone River at Miles City belong to the so-called Tullock and Lebo members of the Fort Union sequence. 

The base of this sequence, and hence the top of the 

Hell Creek formation, is about 100 feet below river 

level. The Hell Creek, according to a well log taken 

12 miles west of Miles City approximates 500 feet in 

thickness beneath the Fort Union formation under 

Miles City. At Glendive the lowest strata exposed 

in the bluffs south of the city contain distinctive Upper Cretaceous plants and abundant ceratopsian bones 

and thus belong to the Hell Creek formation. Only 

about 100 feet of the top of the bluffs belong to the 

Tullock member, which lacks dinosaur bones but contains Fort Union plants (pl. 3, fig. 2). These definitive stratigraphic facts, however, were never, so far 

as I have been able to learn, recognized by l{nowlton, 

but T. W. Stanton (1909, p. 286) remarked: 

Doctor Knowlton's statistics show a close relation between 

the flora of the lower Fort Union and the upper Fort Union; 

if, however, comparison were made between the Ceratops beds 

that have actually yielded dinosaurs and the typical or upper 

Fort Union [Tongue River member] the figures would possibly 

be different, because in many sections such as those of the 

Bighorn Basin, Red Lodge, Sheridan, Fish Creek, and probably along the Yellowstone, he has included more than the 

Ceratops beds in his lower Fort Union. 

Had Knowlton acted on this cautious suggestion 

and reevaluruted the fossil and stratigraphic situation, 

particularly along the Yellowstone River between Miles 

City and Glendive, the Laramie-Fort Union or Cretaceous-Tertiary problem might have been solved at 

that time. 

The Hell Creek beds '_Vere named and described by 

Barnum Brown (1907) from the type section on Hell 

Creek, north of Jordan, Mont., and were unequivocally 

correlated with the Lance formation of Wyoming (B. 

Brown, 1914, p. 356). Brown was particularly careful to point out, by italicizing the statement, that the 

latest remains of dinosaurs in that area were confined to these strata, because after 5 years of fieldwork 

he and his party failed to find any evidence of dinosaurs in the overlying lignitic beds that contain the 

Fort Union flora. The Cretaceous-Paleocene boundary 

at the contact between the Hell Creek and Tullock 

in the region immediately east of Hell Creek was 

shown on the map accompanying the description of 

the coal resources of McCone County, Mont., by Collier and Knechtel ( 1939), although the ceratopsian 
HISTORICAL REVIEW OF THE SUBDIVISION OF THE "GREAT LIGNITE" 7 

I-Iell Creek and lignitic Tullock, or basal Fort Union, 

were, according to prevailing custom, referred to the 

Eocene(~). 

EFFECT OF THE DISCOVERY OF MAMMALS IN THE 

LIGNITIC STRATA 

Perhaps the first illuminating clue to the position of 

the Cretaceous-Paleocene contact in the western interior came from the San J nan Basin of Colorado and 

New lVIexico. There in 1880 and subsequent years, 

distinctive Tertiary mammals (Matthew, 1937) were 

found in the Puerco and Torrejon formations, which 

overlie strata containing dinosaurian remains. By contrast the lignitic strata of the Fort Union sequence 

in ~1:ontana yielded no mammals until Earl Douglass 

in 1901 found the first specimens on Fish Creek just 

east of the Crazy Mountains. In the following years 

this area was thoroughly explored, chiefly by A. C. 

Silberling of Harlowton, Mont., under the inspiration of J. "'\V. Gidley, who began a study of the fossils that was completed later by G. G. Simpson 

( 1937b). That part of the geologic section pertinent 

to this discussion, as generalized by Simpson and 

amended by me, is as follows : 

Generalized section east of the Crazy Mountains, JI,;Jont. 

[Adapted from Simpson, 1937b, p, 15] 

No.3 Melville Dark shales, greenish or gray with numerous gray to yellow sand- stones. Mammals, plants. 

No.2 Soft somber greenish, sandy shales 

and gray sandstones. Mammals, plants. 

Fort Union -- Lebo 

No.1 Somber shales with some hard 

brown sandstones. Mammals, 

plants. 

Alternating pale crossbedded sand- stones and shale. Rare turtle 

Bear bones, etc. No dinosaurs (except possibly reworked !ragments) or mammals. Plants. 

Hell Creek Pale, variegated clays with some gray sandstone. Dinosaurs. 

.'i,OOO 

+ft. 

850ft. 

500ft. 

500-600 

ft. 

2,000 

ft. 

Simpson concluded that the paleontologic evidence 

from that part of the Fort Union he called No. 1 

Lebo was inadequate for exact correlation, but tentatively assigned these strata to the middle Paleocene 

or Torrejonian of the vertebrate paleontologists. Concerning the next immediately underlying beds, which he 

called Bear, he pointed out that this sequence overlies 

the dinosaur-bearing Hell Creek formation at a relatively sharp and observable contact, that it contains 

no dinosaurs except possibly reworked fragments in 

its basal part, that the invertebrates are Paleocene 

forms, and that, therefore, it is likely the Bear is 

Tertiary rather than Cretaceous, and probabiy in part 

equivalent to the Tullock or basal Fort Union farther 

east. Accordingly, on his chart Simpson assigned the 

Bear to Fort Union or Lance. Iri 1936, I obtained 

a satisfactory collection of plants from the Bear in 

SWl~ sec. 29, T. 6 N., R. 16 E. Of the 10 species 

present, few seem referable to the Cretaceous, but 

nearly all are present in the basal Fort Union in the 

vicinity of Miles City and Glendive. On this and the 

stratigraphic evidence I am convinced that the Bear 

belongs with the Tertiary sequence and that the contact between it and the Hell Creek marks the Cretaceous-Paleocene boundary. 

The scarcity or lack of mammals in the Bear of the 

Crazy Mountain region is matched by a similar condition in the Paleocene sequence of extreme eastern 

Montana and the adjacent Dakotas. So far as known, 

the eastern outcrops have yielded only a few specimens, representing six genera: Titanoides primae,ous 

Gidley from Buford, N. Dak. (Gidley, 1917. Near 

Vernon Bailey's original find, Glenn Jepsen and Theodore White recovered additional parts of skulls of T. 

primaevus); Tetraclaenodon sp. and Pantolambda sp. 

from southwestern North Dakota (Lloyd and Hares, 

1915, p. 538); Aphronorus sp., an unpublished insectivore tooth from Donnybrook, North Dakota; and 

Olaenodon sp., Tricentes sp. from Medicine Lake, 

Mont. (pl. 69, figs. 9, 10). 

In 1926, Dr. J. C. F. Siegfriedt, of Red Lodge, 

Mont., discovered another locality that has yielded 

significant mammalian remains. It is the roof of the 

Eagle coal mine, 1 mile south of Bear Creek, Mont., 

at a level about 1,500 feet above the base of the Fort 

Union formation. These fossils, according to Simpson ( 1928; 1929), indicate a fauna that inhabited a 

heavily forested and swampy region and that was different in facies from the fauna on Fish Creek, no 

genus being common to the two. He conjectured its 

·age to be late Paleocene and approximately that of 

the Tiffany fauna of southwestern Colorado. 

The first mammals from the lignitic strata of the Big 

Horn Basin were discovered in 1881. Since that time, 

active exploration has resulted in th~ accumulation of 

much paleontologic and· stratigraphic information, giving a fairly clear picture of the geologic situation in 

that basin. All this has been summarized and discussed by G. L. Jepsen (1930; 1940) and is indicated 

on the correlation chart compiled by ;H. E. Wood and 

others (1941). Jepsen gave a new mime to the Paleocene section in the Polecat Bench area-Polecat Bench 

formation. At that locality he found no angular unconformity between the dinosaur-bearing Lance and 

the Polecat Bench formation, but the contact may be 

locally undulatory. Elsewhere in the Big Horn Basin, 

however, an unconformity marks the Cretaceous-Paleocene boundary, as for example, on the south side of 
8 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Dry Creek, 5 miles west of Greybull, Wyo. Similarly, 

Hewett ( 1926, p. 30-40) found an angular unconformity separating the Cretaceous and Tertiary beds 

on the southwestern side of the basin. 

In 1933 Bryan Patterson reported the finding of 

large titanoid mammaJs in strata southwest of De 

Beque, Colo., which in 1936 he proposed to call the 

Plateau Valley beds. These beds, according to the 

fauna, are of late Paleocene age .. 

My finding of mammalian remains in 1939 and 1940 

at several strategic localities in the Denver Basin, 

Colorado, helped materially to restrict the position of 

the Cretaceous-Paleocene boundary in that area ( Gazin, 

1941a; Brown, 1943a,). Gazin ( 1942; 1956c) reported 

late Paleocene mammals from the Almy formation in 

southwestern Wyoming, (1956a) a large fauna from late 

Paleocene strata in Bison Basin, on the north border 

of the Red Desert, Wyo., and (1956b) a Tiffanian 

fauna from Fossil Basin, west of Kemmerer, Wyo. In 

the vicinity of Hoback Canyon; south of Jackson, 

Wyo., Dorr (1952; 1958) found significant mammalian 

remains in what he and Eardley call the Hoback formation, a possible northward correlative of the Evanston and Almy formations near Evanston. During the 

summer of 1958 A. E. Roberts, of the U.S. Geological 

Survey, found a portion of the jaw, with three teeth, 

of a Paleocene condylarth, Tetraclaenodon symbolicus 

Gidley, in strata on Willow Creek, 9 miles north of 

Livingston, Mont. This find is stratigraphically close 

to the tentative Cretaceous-Paleocene boundary I drew 

(1949) in the Livingston area and narrows materially 

the search for the exact line. The recent discovery of 

fragmentary vertebrate remains in the Medicine Rocks 

area north of Ekalaka, Mont., by members of the 

Carter County Geological Society, has led G. L. Jepsen, of Princeton University, to further prospecting 

in those Paleocene strata. In a personal communication he states that the unpublished material now includes fish scales and teeth, amphibian and reptilian 

vertebrae, and mammalian teeth and jaws, indicating 

a probable middle Paleocene age. 

In Canada the Paskapoo formation, which at some 

localities in Alberta is clearly unconformable on the 

Edmonton formation (Upper Cretaceous), contains 

fragmentary remains of Paleocene mammals (Russell, 

1929, 1932a, 1932b, p. 137), associated with fishes, 

amphibians, and reptiles. 

A general survey of Paleocene vertebrate faunas 

was published by Simpson (1936; 1937a); and Van 

Houten ( 1945b) has reviewed the "latest Paleocene and 

early Eocene mammalian faunas, with suggestions 

about the associated climatic conditions and depositional facies. 

Thus, the revelation of the mammalian evidence has 

helped greatly in dating parts of the "Great Lignite" 

and in narrowing the quest for the lower and upper 

limits of the Paleocene series. 

THE CANNONBALL MARINE STRATA 

Not long after the finding of mammals in the Paleocene rocks of the Crazy Mountain region came the unexpected discovery of fossiliferous marine strata intercalated in the lignitic sequence near Mandan and 

southwestward in North and South Dakota. Speculations about the age of these strata soon produced a 

name that caused much discussion-the Cannonball 

marine member of the Lance. In 1907, before the main 

discovery in 1912 by Lloyd and Hares ( 1915, p. 523), 

A. G. Leonard, exploring the steep bluffs along the 

Little Missouri River south of Yule, N. Dak., found 

an oyster bed ( Ostrea glabra Meek and Hayden) lying 

between two coal seams about 150 feet above the river 

(pl. 2, fig. 2). This was later interpreted as a brackishwater tongue of the Cannonball member. In 1931 I 

discovered a second and lower brackish-water tongue 

at river level one-half a mile south of Leonard's 

oyster bed locality (pl. 2, fig. 3). It contains species 

of Oorbula and Oorbwula but apparently no oysters. 

These tongues of the Cannonball, which were probably 

deposits in estuarine channels or embayments of the 

Cannonball sea, interfinger with the nonmarine strata 

of the Ludlow member, which has never yielded any 

authentic indigenous dinosaurian remains but contains 

abundant specimens of the typical Fort Union flora. 

T. W. Stanton concluded that the Cannonball fauna 

has a general Tertiary aspect because of the absence 

of the exclusively Mesozoic forms and the presence of 

many long-lived modern forms; but that it has a 

closer relation with the Cretaceous because 24 of its 

species . ( 40 percent of its known molluscan fossils) 

were identified with species found in the Late Cretaceous formations of the same general region (Stanton, 1914, p. 351; 1921, p. 13). He, therefore, regarded the Cannonball strata as Cretaceous in age. 

The element of doubt that this fauna might after all 

not be Cretaceous in age lay not only in the absence 

of ammonites but in the presence of the mollusks 

Oalyptrap!Wrus and Oylichnella, the former occurring 

in North America only in strata ranging from Midway 

(Paleocene) to Jackson (Eocene) in age and the latter with no definite record older than Tertiary. 

Soon after Stanton's paper on the Cannonball appeared, Charles Schuchert ( 1921, p. 45, 4 7) reviewed 

it and forthwith advocated that the line between the 

Cretaceous and Tertiary be drawn at the contact of 

the Fort Union and overlying Wasatch. Knowlton, 
HISTORICAL REVIEW OF THE SUBDIVISION OF THE "GREAT LIGNITE" 9 

after reading Schuchert's proposal, remarked that "If 

the Cannonball marine member is Cretaceous ·then 

both Lance and Fort Union are Cretaceous, for there 

is no stopping point short of the top of the Fort 

Union." 

The reason for this statement was Knowlton's belief in the existence of an unconformity of great proportions at the contact of the Fox Hills sandstone and 

Hell Creek formation, and this being. the only such 

purported, available unconformity known to him in the 

"Great ·Lignite" he picked that contact as the Cretaceous-Tertiary boundary. Investigation of this alleged unconformity, mainly in eastern Montana and 

the Dakotas, resulted in the publication by Dobbin 

and Reeside (1929, p. 25) of observations tending to 

explain the phenomena at the localities they examined 

as local channeling, minor faulting, crossbedding, or 

slumping. This conclusion harmonized with the previous opinion expressed by Barnum Brown (1914, p. 

357) about stratigraphic relations along Dry Creek, 

north of Van Norman, Mont. However, Dobbin and 

Reeside cited Bauer ( 1925) as reporting that he had 

traced around Freedom Dome, south of Jordan, Mont., 

a stratum of coarse material consisting of quartzitic 

and other pebbles, at the base of the Hell Creek formation. Only Dobbin checked this report, finding 

some evidence of channeling, but both Dobbin and Reeside (p. 17) concluded: "As the conglomerate occurs 

only in this small area, its presence affords little proof 

of the existence of a major unconformity at the contact of the Fox Hills and Lance." Fieldwork by Fred 

Jensen, Roger Colton, and me in 1950 and subsequently (Colton, 1955) indicates that this channeled 

contact cannot be dismissed as a local condition. It is 

much more widespread than was known to Barnum 

Brown, Dobbin, Reeside, and Bauer. We found it at 

numerous localities over hundreds of square miles of 

the terrain north and south of the Missouri River, east, 

west, and south of Fort Peck Dam, and southward as 

far as Forsyth on the Yellowstone River. The conglomerate consists mainly of quartzitic and porphyritic pebbles, cobbles, and boulders, occasional worn 

dinosaur bones, shark's teeth, and rounded pellets of 

wood and lignite. Moreover, the interstitial sand of 

this conglomerate and that of the immediately overlying beds is somewhat yellowish brown with a liberal 

sprinkling of black grains, so that once seen at a 

typical outcrop it can be readily recognized elsewhere. 

This characteristic, friable, salt-and-pepper Hell Creek 

sand appears abruptly above the contrasting grayishwhite upper Fox Hills strata. Thus, although the 

transition from Fox Hills to Hell Creek is physically 

and lithologically sharp over a large area, the debate 

about the nature, magnitude, distribution, and correlation of the unconformity may go on. On one point, 

however, there is certainty-this unconformity, however interpreted, did not mark the end of the Cretaceous and the beginning of the Tertiary. 

Fieldwork by Laird and Mitchell (1942, pl. 1) and 

by me since the publication of Stanton's paper shows 

that the Cannonball member at most localities rests 

upon a tongue of the lowest coal-bearing zone of the 

Ludlow member, exposures of which can be seen beneath the lower brackish-water tongue of Cannonball 

deposits on the Little Missouri River, south of Yule; 

along the upper reaches of Cedar Creek, a tributary 

of Cannonball River; and north of Cannonball River 

across the southern part of Morton County, N. Dak. 

This 50-foot zone has yielded collections of typical 

Fort Union plants but no dinosaurs. 

Finally, S. K. Fox, Jr., and R. J. Ross, Jr., reported 

that "an analysis of 64 species of Foraminifera indicates Midway (Paleocene) age for the Cannonball 

beds of North Dakota" (1940, p. 1970; 1942). This 

study apparently climaxes the search for evidence 

necessary to settle the question of the age of the 

Cannonball member, and also that of the Ludlow member and equivalent beds with which the Cannonball 

brackish-water tongues interfinger. As early as 1931, 

I had become convinced of the Paleocene age of the 

Cannonball member and its correlative nonmarine 

equivalent but first expounded this opinion in detail 

before the Geological Society of Washington on March 

9, 1938 ( 1938, p. 421). 

FORMULA FOR LOCATING THE CRETACEOUSpALEOCENE CONTACT 

Just as the phrase "somber beds" occurs repeatedly 

in this boundary discussion, so another is also much 

used, namely, "the lowest persistent bed of lignite," 

invented by W. R. Calvert (1912a, p. 196, 197). Calvert observed that in the vicinity of Glendive, Mont., 

the latest dinosaurian remains are found just below 

(he says "above," but this is erroneous, as explained 

in what follows) a "persistent bed of lignite," or perhaps better, a persistent lignite zone. Coinciding with 

this level locally, a marked change in color takes 

place, from the dark Lance (Hell Creek) to the yellowish basal Fort Union. This is the first clear hint 

of a practical formula for locating the CretaceousPaleocene boundary in the lignitic sequence east of 

the Rocky Mountains. 

This lowest lignite zone, indicating the renewal of 

coal formation after a long coal-barren interval, represents a widespread, simultaneous phenomenon 

throughout eastern Montana, Wyoming, and the Da-. 
10 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

kotas (Winchester and others, 1916, p. 19; Laird and 

Mitchell, 1942, p. 12; Denson and others, 1959, p. 16). 

Can the base of this lowest coal zone be mapped and, 

therefore, be accepted as a close approximation of the 

Mesozoic-Cenozoic or Cretaceous-Paleocene contact~ 

To answer this question several conflicting published 

statements must ·first be considered. Calvert ( 1912a, 

p. 197~ 198) reported dinosaurian remains from above 

this lignite zone, as follows : 

One collection from a horizon just u !Jove the lowest persistent lignite bed [sec. 36, T. 13 N., R. 59 E., 11 miles south of 

Wilbaux, Mont.] repres~nted the following forms as recognized 

by C. W. Gilmore. · [Here. follows a list of turtles, crocodiles, 

Champosaurus, and a caudal vertebra of an undescribed 

dinosaur.] 

He also ventured the opin,ion that such remains may 

occur through perhaps 500 feet more of overlying 

strata. Were these statements correct, the detection of 

another usable contact would be extremely difficult, .if 

not impossible, because the color and lithologic composition of the strata above the basal coal do not permit satisfactory distinction for mapping purposes. 

The field notes of neither Calvert nor his assistant, 

Hance, yielded any further clues about the dinosaur 

vertebra. My examination of the area in 1941 showed 

that with the exception of a small patch of Lebo in 

the southwest corner, all the strata in sec. 36 belong 

to the Tongue River member. Therefore, the dinosaur 

vertebra could not have come from "just above the 

lowest persistent lignite," for that lignite is not present in sec. 36 but occurs 500 feet lower stratigraphically 

than the lowest stratum cropping out in that section. 

Some mixup in locality citation or in the fossil collections probably accounts for this erroneous record. 

Another report that dinosaurian remains occur above 

the basal coal zone is that cited by Rogers and Lee 

(1923, p. 34), who mapped the Tullock Creek coal 

field, Montana: 

The single bone fragment found was identified by C. W. Gilmore as a section of a supraorbital horn core of a ceratopsian, 

probably Triceratops. This fossil which was collected [by W. C. 

Mansfield] in sec. 22, T. 5 N., R. 35 E., was found associated 

with other bone fragments on the surface of a steep slope 

about 50 feet above the base of the Tullock member. 

The base of the Tullock here was considered to be at a 

coal seam called coal A. In 1941, I visited this locality 

and found that Rogers and Lee had stated the field relations correctly and that the steep slope mentioned 

in the locality citation does indeed yield dinosaurian 

bones. Coal A, however, crops out near the top of the 

hill, and in the strata above it. I found only turtle 

remains. Subsequent examination of Rogers' .field 

notes covering this locality revealed on page 1028 of 

notebook 27 45 the clarifying and saving statement that 

the horn core fragment was picked up 50 feet below 

coal A and that other bone fragments, presumably of 

turtles similar to those I also saw, were found above 

coal A. 

These dubious or erroneous records have had misleading consequences. Thus, in describing a new species 

of Triceratops from what he called the Torrington 

member of the Lance in Goshen Hole in eastern 

Wyoming, Schlaikjer (1935) said that the skull was 

found in the upper strata of a 240 to 425-foot sequence 

of shales, sandstones, and a brackish-water bed; which 

overlies a lignite seam at or near the contact with the 

Pierre shale or perhaps Fox Hills sandstone. Influenced by the geologic similarity of the upper part 

of this section to that in the bluffs of the Little Missouri River near Yule, N. Dak., and unaware that a 

second brackish-water tongue of the Cannonball member occurs 150 feet below the oyster bed originally 

reported at Yule, he concluded that his Triceratops 

postdated the Cannonball deposits. In support of this 

conclusion he cited the dinosaur bone reported by Calvert but misinterpreted Calvert's statement to mean 

· that "* * * in eastern l\1ontana ceratopsian bones were 

found in be9,s which are supposed to be stratigraphically higher than the Cannonball." Calvert, however, 

stated specifically that the bone came from "a horizon 

just above [below] the lowest persistent lignite bed." 

This, as has been shown, is a position just above the 

top of the Hell Creek formation and represents a level 

below, not above, the base of the Cannonball member. Schlaikjer (p. 54), it is true, recognized the possibility that the Goshen Hole tongue of brackishwater deposits may represent an earlier extension of 

the sea than the Cannonball tongue on the Little Missouri River. As the Lance of Goshen Hole lies (unconformably according to Schlaikjer) upon Pierre or 

Fox Hills strata, it invites comparison with the basal 

rather than the upper part of the true Lance in neighboring areas. In the Lance Creek area, 100 miles to 

the northwest, a lignitic zone and brackish-water bed 

occur near the base of the Lance or top of the Fox Hills 

(Stanton, 1909, p. 242; 1910, p. 187), and this lignitic 

zone appears at approximately the same level in 

other parts ~f eastern Wyoming and western North 

and South .I)akota (Shaw, 1909, p. 158; Wegemann, 

1912, p. 446 ;· Laird and Mitchell, 1942, p. 8). In the 

Cannonball _.:River region of North Dakota, a thin 

marine tong'Ue, called the Breien member, in the basal 

part of the Hell Creek formation may be the equivalent of the .Goshen Hole brackish-water stratum. This 

correspondence in lithologic composition and stratigraphic position suggests that the Lance in Goshen 
SUMMARY OF PALEOCENE STRATIGRAPHY 11 

I-Iole is basal Lance and that Schlaikjer's dinosaur is 

an early rather than a late form of Triceratops. 

No other published reports of the occurrence of indigenous dinosaurian remains above Calvert's "lowest persistent bed of lignite" have appeared, so far 

as I am aware. That reworked fragments may, however, occur in the basal Tertiary strata is quite possible, 

but diligent search by myself and field parties I have 

visited during the past 20 years has brought none such 

to light. 

My conclusion is that the last dinosaurs disappeared, 

for causes unknown, from the western interior area 

at or somewhat before the time when the latest sediments of the Hell Creek and Lance formations were 

deposited, and new environmental conditions were inaugurated. The depositional area began to subside, 

and the low flood plains of the Late Cretaceous 

changed to widespread Tertiary sloughs and swamps 

in which vegetation accumulated to form lignite. Species of a characteristically new flora and fauna appeared, so that when good representations of their 

fossilized remains are now found they can be easily 

distinguished from Late Cretaceous assemblages. 

Calvert's basal coal zone, as proved at all localities 

east of the present Continental Divide and north of 

the Colorado-Wyoming line to the Canadian border, 

can, with a litde experience, be detected readily and 

mapped satisfactorily. When the paleontologic evidence from above and below this zone is compared 

with that from strata in adjacent or distant areas 

where the coal zone does not occur, those strata can 

in accordance with their fossil content be identified as 

Tertiary or Late Cretaceous, respectively, and the contact between Mesozoic and Cenozoic beds can usually 

be fairly closely determined. In general, the horizon 

of the contact is nearly everywhere marked by changes, 

either subtle or obvious, in lithologic composition and 

color features that become more distinct as familiarity 

with the strata grows. 

PALEOCENE-EOCENE CONTACT 

Before use of the term Paleocene became common, 

the earliest Tertiary of the western interior was called 

Eocene, but the strata were regarded by many geologists as transitional between· Cretaceous and Tertiary, 

and the upper limit of this transitional series was 

considered to be the true Cretaceous-Tertiary boundary. 

The oldest strata overlying the Paleocene, conformably 

or unconformably, in many parts of the Rocky Mountains and the Great Plains, are conglomerates, sandstones, clays, shales, and coal seams, sometimes resembling the underlying .. beds in general aspect but 

more often being coal-barren, sandy, and varicolored. 

These strata are commonly called the "Wasatch" formation, and pending a restudy and possibly a renaming 

of much that is now called Wasatch, I shall here use 

the term only in a general identifying sense. For a 

review of the mixed situation involving the term 

Wasatch, the reader is referred to publications by N ace 

(1936, p. 120-146), Jepsen (1940, p. 223, 224), and 

Van Houten (1944, p. 172-174). 

The upper limits of the Paleocene have been determined in part by lithologic changes and in part by 

mammalian and floral remains. Thus, when examining a given terrain for this contact, the geologist, finding relics of the early horse, Hyracotherirum ( eohippus), and of the floating fern, Sal!vinia preauriculata 

Berry, will know that the upper limit of the Paleocene 

has been passed and that the containing strata are 

Eocene. Specific references to this contact will be 

found in the discussions of the Hanna Basin, Powder 

River Basin, western North Dakota, etc. 

SUMMARY OF PALEOCENE STRATIGRAPHY 

Because my chief purpose in this paper is to discuss 

the plants of the Paleocene and the evidence pertaining to detection of its boundaries, I shall not here 

review Paleocene stratigraphy in more than necessary 

detail. Readers interested in the minutiae of thicknesses and lithologic variations of strata should consult the numerqus pertinent coal and other reports. 

As now recognized, the Paleocene units or formations and their members in the Rocky Mountains and 

Great Plains are: 

Canada: Paskapoo, Ravenscrag, Willow Creek (part). 

Montana, the Dakotas: Fort Union (Tullock, Lebo, Ludlow, 

Cannonball, Tongue River, Sentinel Butte members), Willow Creek ( ? part), Livingston (part) , Mel ville of Simpson 

(1937), Bear of Simpson (1937). 

Wyoming: Fort Union, Polecat Bench of Jepsen (1940), Ferris, 

Evanston, Almy, Hoback of Horberg, Nelson and Church 

(1949). 

Colorado: Denver (part), Dawson (part), Coalmont, Middle 

Park, Plateau Valley of Patterson (1936), Raton, Poison 

Canyon, Animas (part). 

New Mexico: Raton, Animas (part), Puerco, Torrejon, Tiffany 

of Granger ( 1917). 

Utah: North Horn, Dragon of Kugler (1953), Henefer. 

In particular, the Fort Union of the northern areas, 

having been the subject of so much dispute and because the type section is not typical, has been redefined as follows and is thus virtually synonymous with 

Paleocene series in that region: The Fort Union formation in eastern Montana, western North and South 

Dakota, and eastern Wyoming is typified by the sequence of lignitic strata exposed in the right bank of 

the Yellowstone River from the top of the Cretaceous 

dinosaur-bearing Hell Creek formation at Glendive to 
12 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

and including the original type section (pl. 1, fig. 1) 

in the badlands on the left bank of the Missouri River, 

opposite the . mouth of the Yellowstone River. The 

top of the formation is the base of the Golden Valley ("Wasatch") formation (Eocene) in isolated outcrops in western North Dakota. 

The Paleocene series of the western interior includes coal seams, sandstones, conglomerates, shales, 

clays, thin impure limestones, and in some areas, igneous rocks or the products of igneous activity. The 

series attains a thickness of 5,000 feet or more in the 

Hanna Basin, Wyo., the Crazy Mountains, Mont., and 

North Park, Colo., but in most areas less than 2,000 

feet remain. Eastward, or seaward, from the mountains the strata thin and decrease in coarseness. Where 

the strata now lap against the mountains the sandstones may be coarse to conglomeratic. Some channel 

deposits within the sequence contain fairly coarse material at a considerable distance from the assumed 

source (Lloyd and Hares, 1915, p. 538, 539). Hewett 

(1926, p. 30-40) reported conglomerates in the Fort 

Union of the west side of the Big Horn Basin in 

Wyoming. 

The terrain that received the Paleocene sediments 

was physiographically a vast low-lying and intermittently subsiding flood plain over which flowed 

meandering and braided streams through poorly 

drained inland swamps and marshes, with shallow 

bogs, sloughs, low moors, ponds, lakes, and lagoons, 

where, during sedimenlt-free intervals, peat-forming 

processes were active. The piedmont portions of the 

flood plains nearest the sources of the silts and sands 

that buried the subsiding accumulations of vegetable 

remains, lapped against the flanks of the mountains 

and received the coarser detritus. Crossbedding of the 

sa~dstones, channeling, overlap, and interfingering suggest intermittent diversion and change of the inflowing sediment-bearing currents that buried the subsiding, incipient coal beds. With the exception of the 

coal, all these sedimentary deposits may be said to be 

of fluviatile origin (Davis, W. M., 1900). Generally, 

but not always, the coal seams thin out and disappear 

by lateral g~adation into carbonaceous shales. The 

coals may be underlain by light-colored or bluish underclays that were in part the soils or substrates for 

the coal-forming plants. Besides this peaty material, 

. the abundant plant remains in the sandstones, clays, 

and shales show that vegetation was more or less 

luxuriant on the higher parts of the flood plains and 

in the highlands. Van Houten ( 1945a, p. 444; 1948, 

p. 2121) , discussing depositional conditions in this 

general area during the early Cenozoic, suggests that 

the gray strata, because they contain small arboreal 

animals, probably accumulated in heavily wooded lowlands or swamps, whereas the variegated strata, containing remains of ungulates and carnivores, probably 

accumulate& on flood plains, piedmont areas, and valley flats, and ·reflect a grassland or savannah environment. 

Today, in few localities do these Paleocene strata 

retain their original more or less horizontal attitudes. 

All have been raised vertically, and in many localities 

they are now tilted, the steepest inclinations being on 

the flanks of anticlines or where they lap against the 

mountains that had intermittent uplifts d~ring or after the deposition of the sediments. Viewed in large 

perspective, most of the Paleocene strata in an~ around 

the Rocky Mountains lie in synclinal basins such as 

the Williston Basin in North Dakota, Powder River 

Basin, Bighorn Basin, Wind River Basin, and Hannah Basin in Wyoming, Denver Basin and North Park 

Basin in Colorado, and the San Juan Basin in Colorado 

and New Mexico. Such folds as occur in these large 

basins are,· for the most part, fairly gentle, such as 

Porcupine dome (Bowen, 1916) and Cedar Creek anticline (Erdmann and Larsen, 1934; Dobbin and Larsen, 1934) in Montana; Nesson anticline (Collier, 

1918) and Keene dome (Nevin, 1946) in North Dakota. Many minor unnamed swells are found in nearly 

all the basins. Other deformations of not clearly explained origin also occur (Townsend, 1950). Over 

large areas on the Great Plains east of the Rocky 

Mountains the general attitude of the Paleocene strata 

superficially seems to be that of horizontality, but in 

reality there is, particularly in Montana and North 

Dakota, a low dip to the northeast. 

Faulting is comparatively rare, but slumping and 

landslipping are common, particularly where thick 

coal beds have burned. Some sediments were covered, 

intruded, or otherwise affected by igneous activity, as 

for example, at Golden, Castle Rock, and Walsenburg, 

Colo.; in the Crazy Mountains northeast of Livingston, 

Bearpaw Mountains south of Havre, and Smoky Butte 

west of Jordan, Mont. Some of this igneous action, 

particularly in Colorado, occurred during the Paleocene. The age of that in the Bearpa ws is Eocene, but 

of that in the Crazy Mountains and near Jordan is 

indeterminate. 

On the Little Missouri River, near Yule in southwestern North Dakota, the early nonmarine Paleocene 

sediments interfinger with brackish-water strata that 

probably represent former estuarine channels of the 

Cannonball sea in which, near Mandan, about 300 feet 

of marine sediment comprising the Cannonball member was deposited. The fact that brackish-water tongues of the Cannonball member (pl. 2, figs. 2, 3) lie 
SUMMARY OF PALEOCENE STRATIGRAPHY 13 

in the midst of thick coal seams indicates that those 

and most likely all the Paleocene coals were paralic, 

that is, they were deposited at or near sea level in 

extensive coastal swamps and marshes that may at 

times have been considerably inland from the open 

coast. These marshy basins subsided intermittently 

and. locally for a total of thousands of feet in the 

course of millions of years, but the subsidences were 

interrupted by periods of relative stability permitting 

·vegetation to grow and its remains to accumulate as 

peat that was eventually transformed into lignite beds 

(Thorn, 1929, p. 19; Stutzer, 1940, p. 162) after burial 

under a cover of sand and mud. The repetitive occurrence of coal beds through the PaleoceJle sequence 

into the Eocene suggests the inference that not only 

before and during Cannonball time but for a long 

time afterward an open sea, perhaps extending from 

the Gulf of Mexico to_ the Arctic Ocean, occupied, 

more or less continuously, an unknown area not far to 

the east or northeast, where, however, there are now 

no marine strata left to testify to its former presence. 

There is perhaps no exactly comparable existing situation on the same scale. The Dismal Swamp area 

of Virginia and North Carolina (Osbon, 1920) may 

be cited, but it is small compared with the Tertiary 

lignitic area of the Rocky Mountains and Great 

Plains. I gather that conditions of sedimentation in 

the basins of the Amazon and Parana Rivers of South 

America may approximate those of the early Tertiary 

in the Rocky Mountains and Great Plains, for large 

areas on ·both sides of these rivers may be inundated 

during flood stages. Peat, I am also told, is accumulating in many of the swampy localities there. 

I£ the inference that all the Paleocene coals were 

formed at sea level is correct, the difference in level 

between the average base of the Paleocene (about 5,000 

feet above the sea) in the plains just east and west 

of the Rockies and its average base in North, Middle, 

and South Parks, now high (about 8,000 above the 

sea) in the Rockies near the Continental Divide, indicates a differential uplift by folding and faulting 

of the park -areas above the plains of at least 3,000 

feet and a total uplift for some outcrops of at least 

8,000 feet since Paleocene time. The general similarity 

in the composition of the floras from several basins 

now separated by mountain ranges also suggests that 

the basins were once more or less continuous or were 

at least not separated by barriers that prevented interchange of plants. W. T. Lee (1915, p. 27, 56) 

claimed there is lithologic and physiographic ~vidence 

that all the Upper Cretaceous formations from the 

Dakota sandstone through the Laramie once covered 

the Rocky Mounta,in region, the source of the sediments being an ancient landmass occupying the present site of the Great Basin and portions of the Sierra 

Nevada. On the other hand, Lovering (1929, p. 88 ff.) 

states that "there are a few places in the Front Range 

where the evidence strongly suggests that Dakota sandstone was never deposited," and that "some areas of 

the Front Range highland were above water at the 

close of Dakota time." The inference is that they 

remained above water and were never again completely covered by sedimentary formations. I am inclined to the opinion, after seeing the coarse andesitic 

and arkosic conglomeratic material in the Paleocene 

of the Denver Basin and Middle Park, Colo., that 

during most of Paleocene time the present site of the 

Rocky Mountains was occupied by an emerging chain 

of large hilly islands separated by shallow channels 

and fringed by very broad unstable coastal areas and 

that these landmasses contributed a fair percentage of 

clastic sediments to the making of the Paleocene for.:. 

mations. Some sediments, however, very likely originated in the ancient landmass still farther west and 

southwest. 

Intermittent orogenic uplifts of the ancestral Rocky 

Mountains core during the continuing so-called Laramide Revolution were reflected in the clastic sediments deposited near the source of supply by resulting 

unconformable relations between the strata. Localities 

or regions where the Paleocene strata· overlie older 

strata unconformably may be seen on Dry Creek, 5 

miles west of Greybull, Wyo. ; 2 miles southeast of 

Black Buttes Station, Wyo. (pl. 1, fig. 3); in Rifle Gap, 

6 miles north of Rifle, Colo.; in North and Middle 

Parks, Colo. (Beekly, 1915, p. 20; Grout and others, 

1913, p. 36, 37) ; and on Cimarron River,. northeast 

of Ute Park, N. Mex. (Lee, 1917, p. 69-74). 

At the close of Paleocene time, uplift in the Bighorn 

Mountains in Wyoming resulted in the deposition of 

conglomerate on both flanks of the mountains. This 

basal Eocene conglomerate, called the Kingsbury conglomerate member of the vV asatch formation on the 

east flank, is unconformable on Paleocene lignitic 

strata. 

Although locally, as in ea;stern Montana and adjacent areas, the Paleocene series has been divided into 

members of the Fort Union formation, largely for 

rna pping purposes, these divisions (Tullock, Lebo, 

Ludlow, Cannonball, Tongue River, Sentinel Butte) 

cannot be extended as such into all areas by either 

lithologic or paleontologic means. The fossil floras, for 

example, are notably different in the basal and upper 

strata-a condition that should not be considered unusual because the long time interval separating them 

made changes in the composition of the flora at any 
14 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

given locality inevitable. Intergrading floras, however, connect the two extremes. Moreover, collections 

from widely separated localities, latitudinally, although 

from the same general stratigraphic level, also differ 

notably, as might be expected if climatic zones prevailed in the Paleocene as they do today. 

No sharp distinctions can be made stratigraphically 

within the Paleocene at any given locality on the 

basis of the floras. Such distinctions as have been 

made on the basis of mammalian remains in the Crazy 

Mountains, Polecat Bench, San Juan Basin, and Bison 

Basin are to the point and may be useful some day 

in many other areas that have not as yet yielded such 

fossils. 

The differentiation of the several members of the 

Paleocene by comparison of their mineral constituents 

has made little progress, except perhaps, locally, as, 

for example, in its application to the Lebo shale member. Stone and Calvert (1910, p. 753) described the 

Lebo in the Crazy Mountain area of Mon,tana as andesitic in character and as passing westward into the 

Livingston formation. Eastward, Woolsey and others 

(1917, p. 25) found the Lebo still andesitic in the 

Bull Mountain coal field, and Rogers and Lee ( 1923, 

p. 36-40) detected a similar condition in the Tullock 

Creek coal field. North of the Yellowstone River in 

the Sheep Mountain coal field, between Miles City 

and Terry, Rogers (1913a, p. 171; 1913b, p. 723) reported what he considered to be andesitic material in 

the Lebo. On the other hand, chiefly south of the 

Yellowstone River, between Forsyth and Hathaway, 

Renick ( 1929, p. 17, 19, 21) found no essential difference in mineral constituents between the Hell Creek 

formation and Tullock member of the Fort Union 

formation. These formations, besides being very arkosic, contain quartz and volcanic debris. He found 

that the Lebo also contains considerable amounts of 

arkosic material and some volcanic debris, but did 

not consider the term "andesitic" as applicable. In 

short, he found that the Lebo is not markedly different, mineralogically, from either the Tullock or the 

Tongue River member. Andrews (1936, p. 387) and 

others concur in Renick's opinion that analysis of their 

mineral constituents seems at present ineffectual for 

differentiating these several formations in extreme 

eastern Montana and adjacent terrain. 

Although Stow (1938) made numerous heavy-mineral analyses in an attempt to distinguish the Cretaceous from the Tertiary and the several divisions of 

the Tertiary from one another in the vicinity of Red 

Lodge, Mont. the results were not definitely conclusive. Marie Lindberg's more recent study (1944) of 

the Cannonball area southwest of Mandan, N. Dak., 

divides the section from the Fox Hills sandstone 

through the Cannonball member into nine mineral 

zones and finds that the Fox Hills and Cannonball 

contain a high percentage of green amphibole, whereas 

the Hell Creek formation differs by its characteristic 

epidote, garnet, and sphene content. Whether these 

conclusions are o£ more than local application remains to be tested by a similar investigation of the 

correlative strata farther west in the valley of the 

Little Missouri River in western North Dakota and 

elsewhere. 

LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 

Inasmuch as this paper is concerned principally 

with the strata at the boundaries of the Paleocene 

series, I shall here center attention on that phase of 

the stratigraphy. It seems appropriate to begin wi.th 

the areas at first largely involved in the debate about 

these limits and then develop the subject from these 

focal points. 

MONTANA 

Glendive.-Mention has already been made of the 

fact that Calvert considered the Cretaceous-Tertiary 

boundary as at or just above the level of the highest 

dinosaurian remains, which are found below the "lowest persistent bed of lignite" in the vicinity of Glendive. This horizon is just above the level of the city 

water tank, about 100 feet below the top of the bluffs 

south of the city. If, on a clear day and preferably in 

the afternoon when the color of the bluffs is not modified by shadows, one views this line of bluffs from the 

north side of the Yellowstone River at a distance of 

approximately 1 mile, one can pick out this horizon 

by eye and follow it around the crests of the highest 

badland hills. On the outcrop itself, one can follow 

the "lowest persistent lignite zone" southeastward 

around the tops of the hills at Graveyard Coulee, 

thence around the hilltops on the west side of Glendive Creek to a point 5 miles out of Glendive on U.S. 

Highway 10 toward Wibaux, where, by reason of the 

gentle northeastward dip of the strata away from the 

Cedar Creek anticline, the lignite zone crosses the 

highway. In the badlands on the east side of the 

highway, this zone is burned at a number of exposures. 

Looking southwest from this point toward the badlands and mesas in the escarpment on the left bank 

of Glendive Creek, one can readily pick out the same 

lignite zone. Plate 3, figure 2 shows part of this view. 

It will be noted that the strata (Tullock) in the upper 

fourth of the exposure are darker, that is, more coaly, 

above a fairly sharp line, the "lowest persistent lignite zone" in this area, below which the strata (Hell 

Creek) are distinctly lighter in color, a local reversal 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 15 

of the usual color distinctions. I collected dinosaur 

bones from beds about 50 feet below this level and 

Fort Union plan1s from the carbonaceous shale just 

above, not only a,t this locality but everywhere in the 

vicinity of Glendive; and I conclude, therefore, that 

this level is the closest mappable approximation of 

the Cretaceous-Paleocene. boundary that can be found 

in that area. 

The boundary follows the base of the hills on the 

right bank to a point 4 miles down ~he Yellowstone 

River from Glendive, where it crosses. the river and 

reappears in the low bluffs just northwest of a point 

on State Highway 14 to Sidney, 3 miles from the 

Yellowstone bridge. The horizon is marked by the 

red clinker of the "lowest persistent lignite bed." 

Here I found triceratopsian bones 75 feet below the 

coal, and Fort Union plants in the clinkered shale. 

This zone can be traced around the hills to the northwest. At a point 8.5 miles along State Highway 18 

from the Yellowstone bridge toward Circle, a clinkered 

coal can be seen in the ridges one-half mile to the 

west. As these strata are on the east flank of the 

Cedar Creek anticline, the coal bed can be seen dipping across the highway just one-half mile ahead and 

from thence can be followed at the base of the low 

hills to the east. Here, however, another thin coal 

seam occurs about 25 feet below the clinkered bed; 

and in the carbonaceous shale just above the coal, I 

found typical Fort Union plants, but no dinosaurian 

remains, from which I infer that the Cretaceous-Paleocene boundary is at the level of the lower coal seam. 

Graveyard Coulee, extending southeastward from 

Glendive, is the location of the city dump. The surrounding eerie, somber badlands are, however, notable 

for the even more ancient relics that may be found 

in and on the almost bare outcrops. Dinosaur, turtle, 

and crocodile fragments are plentiful in the lower 

exposures of Hell Creek strata. These beds also yield 

such typical Cretaceous plants as Sequoia dakotensis, 

Brown, Ginkgo laramiensis Ward, Ficus ceratops 

Knowlton, Vitis stantoni (Knowlton) Brown, Dryophyll!um sub falcatum Lesquereux, and Oissus marginata (Lesquereux) Brown. From the same strata on 

Glendive Creek, southeast of Graveyard Coulee, Dell 

and Edwin Lewis of Glendive, collected several multituberculate mammalian jaws and the problematical 

plant, Paleoaster iJnquirenda l{nowlton; but in the 

overlying basal Tullock in Graveyard Coulee I collected the following Fort Union plants: Thuja iJnterrupta Newberry, Paranymphaea crassifolia (Newberry) Berry, and Viburnum asperum Newberry. 

I have already ( p. 5) discussed the section and 

situation on the west fl·ank of the Cedar Creek anticline, 12 miles southwest of Glendive. Northeastward 

from Glendive the average dip of the strata of about 

15 :feet to the mile, about six times the gradient of 

the Yellowstone River, brings higher and higher strata 

to the level of ·the river, so that at the original type 

locality of the Fort Union formation in the hills on the 

north side of the Missouri River opposite the mouth of 

the Y ellow.stone only the highest beds of the formation can be seen and all or nearly all of these belong 

to the Sentinel Butte member. 

Tullock Oreek coal field.-G. S. Rogers and Wallace 

Lee in 1923 divided the enlarged but a1nbiguous Lance 

into a lower part (now the dinosaur-bearing Hell 

Creek formation) and the Tullock member. The latter, a light-colored, yellowish zone, with coal beds, was 

separated from the Hell Creek at the level of the 

lowest persistent coal seam-bed A. This, from what 

has just been said about the Glendive area, is the 

level of the Cretaceous-Paleocene boundary. 

Baker and Ekalaka.--From Glendive, on the east 

flank of the Cedar Creek anticline, the CretaceousPaleocene boundary strikes southeastward toward 

Marmarth, N. Dak., and Buffalo, S. Dak. On the 

west flank of that anticline it runs southeastward 

toward Baker and Ekalaka. Ten miles north of 

Baker it crosses State Highway 7 (pl. 3, fig. 3). Here, 

as at the locality 5 miles southeast of Glendive, the 

lowest Tullock strata are highly carbonaceous and 

coaly and, therefore, appear darker than the underlying Hell Creek formation, in which, a quarter of a 

mile west of the highway and 25 feet below the base 

of the Tullock, I found a leg bone and part of the 

skull of a Trieeratop8. 

Along the border of Montana and South Dakota, 

east of Ekalaka, there was, until the dinosaur-lignite 

contact formula was applied, considerable uncertainty 

as to the position of the Cretaceous-Paleocene boundary. Thus, much of the area along Coal Bank Creek 

formerly mapped as Hell Creek (Bauer, 1924, pl. 33) 

has extensive outcrops of coal or clinker, said, however, by Bauer (1924, p. 239) to be in the upper 100 

feet of that formation. Search of the area by me 

through the sequence of strata from the base of this 

lignitic zone upward, and interviews with the ranchers 

in the vicinity, proved negative in the finding of dinosaurian remains, which, however, can be had at nearly 

every good outcrop beneath that zone, showing that the 

true Hell Creek does not include anything above that 

contact. South and southeast of Ekalaka are high 

forested buttes of Paleocene capped by later-thanPaleocene strata. Southwest of Ekalaka the Cretaceous-Paleocene contact swings around the rim of the 
16 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

escarpment facing Spring Creek and the Powder 

River valley. 

Powder River re1gion.-Northeast of Powderville on 

the east side of Powder River the Cretaceous-Paleocene boundary follows the rim of the badlands. east 

of Spring Creek and eventually descends to and crosses 

Powder River about 6 miles southeast of Mizpah. 

Thence it passes in and out along the hills and coulees 

on the left bank (pl. 3, fig. 1) to a point about 14 

miles southwest of Powderville, where it crosses to 

the east, following a zigzag course almost directly 

south and keeping to the east side of the Little Powder 

River. About 4 miles east of Biddle it crosses Ranch 

Creek, where, in the Hell Creek formation on the 

south bank, I found a large dinosaur vertebra, and 

. in the overlying Tullock cropping out in the hills to 

the north I found specimens of the Paleocene flora. 

The map accompanying the description of the Mizpah coal field by Parker and Andrews ( 1939, pl. 16) 

correctly depicts the Cretaceous-Paleocene boundary, 

although the legend to the map does not name it so. 

The contact as mapped is between Tls (sandstone 

member of Lance formation) and Tlsh (somber-colored beds). The latter strata, according to the map 

legend and text description, are said to include at their 

base about 300 feet of beds equivalent to the upper 

strata of the Hell Creek formation as mapped in the 

adjacent Rosebud coal field (Pierce, 1936) to the west. 

In 1929, I assisted in mapping the Rosebud field and 

have reexamined both areas a number of times since. 

No part of the basal coal-bearing strata of the Rosebud field is equivalent to any dinosaur-bearing part 

of the Hell Creek formation. No dinosaur remains 

have ever been found in the 300 feet of beds in question, but on the contrary, these strata have yielded 

satisfactory collections of Fort Union plants. The 

upper strata of the Hell Creek formation, however, do 

crop out along Rosebud Creek in the northwestern 

part of the field. 

West and· southwest of Powderville toward Coalwood, Broadus, and on to the Montana-Wyoming line, 

the altitude of the terrain increases and younger strata 

of the Fort Union formation appear (Bryson, 1952; 

Warren, 1960). 

Red Lodge and LiJvitngston.-The folded and faulted 

coal-bearing strata in the vicinity of Red Lodge and 

Bear Creek are in the upper part of the Fort Union 

formation. Just west of Red Lodge the upper beds 

of these strata, perhaps in part equi'Valent to the 

Sentinel Butte shale (Thorn and Dobbin, 1924, p. 496; 

Simpson, 1929, p. 8), are cut by the Beartooth fault 

that strikes northwestward. Northeast and east of 

Red Lodge and Bear Creek, the lower limit of the 

Paleocene, because of the folding, faulting, and lack 

of fossil evidence, is not as closely located as is desired 

in that area. From such reconnaissance studies as I 

have been able to make and with the few collections 

of fossil plants that have been gathered, I infer that 

this eastern boundary enters Montana from Wyoming 

in sec. 34, T. 9 S., R. 23 E., Mont., passes northwestward through a much-faulted area (Wilson, 1936, fig. 

2) about 4 miles west of Bridger and about 3 miles 

northeast of Absarokee (Stow, 1938, pl. 1) but loops 

around about 4 miles east of Reed Point without crossing the Yellowstone. North of the river the contact 

encircles a large patch of Paleocene and in the Crazy 

Mountain region becomes the Hell Creek-Bear contact. · 

Gradually along the western boundary of the Paleocene in the direction of Livingston and westward 

against the Bridger Range (McMannis, 1955) lower 

and lower strata appear. In 1908, about 3,700 feet of 

older beds· underlying the productive coal strata in 

the vicinity of Livingston, and originally assigned to 

the Colorado and Montana groups, were definitely determined to be of Colorado age on the basis of marine 

invertebrates. This sequence, for convenience of discussion, was divided by Richards (1957) into 10 units, 

which were tentatively correlated with similar Colorado units elsewhere in Montana, South Dakota, and 

Wyoming. The 800 to 1,000 feet of conformably overlying coal measures, originally called Laramie, were 

found to include marine strata containing invertebrates 

identified by T. W. Stanton as a lower Montana fauna, 

indicating probable equivalence to the Eagle sandstone or to the Mesa verde formation as developed in 

Colorado and Wyoming (Stone and Calvert, 1910, p. 

660, 738; Calvert, 1912b, p. 388). 

The dark-colored remainder of the section above the 

coal measures, west and south of the Crazy Mountains, is an apparently conformable sequence and comprises from 5,000 to 7,000 feet of greenish- or sombercolored andesitic and tufface~us sandstone and flinty 

shale (Weed, 1893, p.· 21; Stone and Calvert, 1910, 

p. 662; Calvert, 1916, p. 202; Wilson, 1936, p. 1165-

1166), including a lens of conglomerate 2,000 feet thick 

where it is cut by the Stillw~ter River. These andesitic beds were called Livingston beds and later the 

Livingston formation (Richards and Prichard, 1950). 

The rocks of more strictly volcanic derivation included 

in this sequence have been studied by Parsons (1942). 

Conformably overlying the Livingston formation are 

4,000 feet, more or less, of light-colored sandstones 

and gray shales acknowledged to be of Fort Union 

age and, from the contained flora, are equivalent in 

part to the light-colored beds east of the Crazy Moun~ 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 17 

tains called "Melville" by Simpson ( 1937b) and of the 

Tongue River member of the Fort Union farther east. 

If, therefore, the several thousand feet of sombercolored andesitic beds between the lignitic Eagle and 

the light-colored Fort Union are in conformable sequence, they must represent the time equivalents of 

the Claggett, Judith River, Bearpaw, Fox Hills, Hell 

Creek, and early Paleocen~ units. A diagram purporting to represent the probable relations of these strata 

was published by Woolsey, Richards, and Lupton 

(1917, p. 26, fig. 2). The resolution of this apparently 

homogeneous lithologic sequence into its component 

parts constitutes the "Livingston problem." 

From a number of scattered localities about 1,200 

to 1,500 feet above the ·Eagle sandstone, Stone and 

Calvert gathered invertebrates and plants. identified 

by Stanton and Knowlton, respectively, as of Judith 

River age. The fossils, it should be stated, came 

principally from strata in the northern part of the 

area that are laterally equivalent to a level at or beneath the horizon of the lenticular agglomerate that 

is said to rest unconformably upon underlying beds 

in the southern part of the area. A good section of 

this agglomerate, about 50 feet thick can be seen 

along U.S. Highway 10 on the south side of the 

Yellowstone River just west of Springdale. I have 

found only a few fragmentary plants (Araucarites 

sp.) in it, and they appear to be Cretaceous forms. 

The beds overlying the agglomerate, although still 

somewhat greenish, gradually become brownish to somber up to the transition into the lighter-colored, grayish beds of the Fort Union formation. 

From the highest loc~lity yielding marine invertebrates, about 2,500 to 3,000 feet ( accor¢ling to Stone 

and Calvert) above the Eagle, came a collection identified by Stanton as being near the top of the Bearpaw. At this locality, sec. 25, T. 4 N., R. 8 E., about 

5 miles north of Wilsall, there is an anticlinal structure exposing Bearpaw (Pierre) shale and higher, 

sandy beds in the top of which in 1949 I found dinosaurian bones. Such remains were also found by 

McMannis and party (1955, p. 1408) in outcrops 3,200 

feet above the base of t~e Livingston formation, 13 to 

16 miles east of Bozeman, and were tentatively considered to be Lancian in age. My saurian locality is 

3 ·miles west of the outcrop of the base of the lightcolored Fort Union. The 4,000 feet, more or less, of 

strata in this 3-mile interval are shaly and sandy, but 

they evidently include time equivalents of Fox Hills, 

Hell Creek, and early Paleocene beds. 

Knowlton, Berry, a~d Stanton in 1913 collected 

fossil plants on Brackett Creek in sec. 9, T. 1 N., 

R. 7 E., southwest of Wilsall. I made similar collections in that vicinity in 1940. These collections, 

however, are not strictly diagnostic but contain spe- · 

cies found in the Bear and Lebo on the east side of 

the Crazy Mountains. My tentative conclusion, pending receipt of more informative collections, therefore, is that about 1,500 feet of the upper part of the 

andesitic Livingston unit are basal strata of early 

Paleocene age, but are not now clearly separable 

from the underlying Cretaceous beds by recognizable 

lithologic differences and well-distributed unequivocal 

fossils. If this be true, the Cretaceous-Paleocene 

boundary roughly parallels the Livingston-Fort 

Union contact as mapped by Stone and Calvert (1910, 

pl. 7) but at some indefinite distance west of their 

line. The boundary probably crosses the Yellowstone 

River at or just a little west of Big Timber, undulates 

almost directly westward to the northeast corner of 

T. 1 N., R. 7 E., and then curves northward and 

northeastward around · the north end of the Crazy 

Mountains. Confirmation of this assumption and the 

fact that the upper part of the Livingston formation is Paleocene in age was made during the summer of 1958 when A. E. Roberts, of the U.S. Geological Survey, found the jaw of a Paleocene condylarth, Tetraclaenodon 8Ymbolicus Gidley, in strata on 

Willow Creek in the NW'I4 sec. 4, T. 1 S., R. 9 E., 

9 miles north of Livingston. 

The thick Fort Union sequence in the Crazy Mountains is preserved by intrusive igneous rocks. Simpson ( 1937b, p. 15) reports a thickness of more than 

5,000 feet for the Melville or his Fort Union No. 3. 

If to this be added the Lebo and the Bear, a total of 

more than 6,800 feet is indicated for the entire series. 

As mammalian remains have been found in only the 

lower 3,000 feet of the Melville, the age of the upper 

strata of this unit is conjectural. Whether. or not 

any of these upper strata are Wasatch equivalents 

remains to be seen. 

Bull Mountain coal field.-The authors (Woolsey, 

Richard, Lupton, 1917) of the bulletin dealing with 

this coal field say little about the stratigraphy of 

what they call the "Lance" formation but report that 

coal seams occur in its upper portion. This fact, 

judging by analogy with the situation in the coal 

fields farther east, suggests that the Cretaceous-Paleocene boundary at a number of localities is slightly 

below the contact of the "Lance" and Fort Union as 

mapped by those authors. In 1936, I studied the 

section from Big Timber to Roundup, and found that 

the outcrops of the strata between the Bearpaw and 

Fort Union-that is, of the "Lance" formation-are 

poor in every respect. Consequently, no satisfactory 
18 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

evidence seems to be available now for locating the 

boundary precisely in parts of this field. 

Bearpaw Mountai!M.-In small, faulted patches on 

the northeast, west, and south sides of the Bearpaw 

Mountains are outcrops of lignitic strata of Fort 

Union age (Pepperberg, 1912; Bowen, 1914; Reeves, 

1925; Brown and Pecora, 1949). Collections of fossil 

plants from a number of mines and prospects, particularly from the vicinity of the Mackton mine, 6 

miles east of Big Sandy, contain Glyptostrobus dakotensis Brown, Metasequoia occidentalis (Newberry) 

Chaney, Oereidiphyllum arcticum (Heer) Brown, 

Platanus nobilis Newberry, Viburnum antiquurn Newberry, and Viburnum asperum Newberry. Beneath 

Fort Union outcrops east of Big Sandy are several hundred feet of Hell Creek equivalents (Brown 

and Lindvall, 1953), and faulted against Fort Union 

in some localities are remnants of the variegated 

Wasatch formation with diagnostic plants and animals. The preservation of these relics of Late Cretaceous and early Tertiary formations now exposed 

at or near the level of the Great Plains suggests that 

a stratigraphic seq~ence probably exceeding 5,000 feet 

in thickness has been eroded from north central Montana since Eocene time. 

BTownilng.-A narrow synclinal strip of variegated 

beds, c·alled the Willow Creek formation (Stebinger, 

1916, pl. 15) extends from a point about 10 miles 

north of Browning deep into Alberta, Canada. As the 

underlying beds, the St. Mary River formation, contains dinosaurian remains, they are reliably regarded 

as of Late Cretaceous age. The Willow Creek formation, at least in its upper part, has yielded no dinosaurs but an essentially Fort Union invertebrate fauna; 

and the strata, according to Williams and Dyer ( 1930, 

p. 61), L. S. Russell (1932, p. 140), and E. T. Tozer 

(1956, p. 27) are in part, at least, equivalent to the 

Paskapoo formation. The flora from the latter at 

Calgary, Cochrane, and Red Deer River, Alberta, as 

listed and reviewed by Penhallow ( 1908, p. 13-15) 

and Berry (1926) is clearly a Fort Union ·flora, and 

the mammalian remains (Russell, 1932, p. 137-138) 

are considered to be undoubtedly Paleocene. The Cretaceous-Paleocene boundary, therefore, would seem to 

be at the contact of the St. Mary River or its Edmonton equivalent and the Paskapoo or its Willow 

Creek equivalent in Alberta. However, according to 

Rutherford (1947) who has studied the CretaceousTertiary boundary question in Alberta, there remain 

unreconciled lithologic and paleontologic anomalies. 

Tozer ( 1953) and Thompson and Axford ( 1953) think 

that the lower part is Cretaceous and the upper part 

is Paleocene. 

Scobey.-Collier described the geology of northeastern Montana in 1919 (p. 17-39), and in 1925 (p. 

165) he reported the geologic section in the glaciated 

Scobey lignite field to consist in part of Hell Creek, 

150-180 feet; Tullock, 30-40 feet; Lebo, 200-275 feet; 

and Tongue River, 600+ feet. His Hell Creek includes lignite bed A in its upper part, and bed C 

marks the base of his overlying Tullock. He did not 

state why bed C was chosen as the level for separating Hell Creek from Tullock, and in particular, he 

reported no dinosaur remains from the strata between 

bed A and bed C. 

Bauer (1914, p. 298) drew the boundary at the contact of somber-colored beds ("Lance") and yellow beds 

(Fort Union), but this, from what has already been 

said about the unreliability of colors in these strata, 

is not a satisfactory solution. From Penhallow's 

(1908, p. 9-13) review of the fossil plants contained 

in the lignitic strata of southern Saskatchewan adjacent to the Scobey lignite field, I conclude that most 

of those plants are Fort Union species. Further, the 

absence of any authentic dinosaurian remains strongly 

indicates that all these lignite-bearing strata both in 

Saskatchewan and in the vicinity of Scobey are Paleocene in age. Southwestward toward Wolf Point and 

Fort Peck, Hell Creek strata appear in the valleys of 

Wolf Creek and Poplar River (Colton, 1955; Colton 

and Bateman, 1956). 

CANADA, NORTH DAKOTA, AND SOUTH DAKOTA 

The strata of the Cannonball and Tongue River 

members that may be seen along the bluffs of the 

Missouri River west of Bismarck, N.Dak., pass under 

a cover of glacial debris east and northeast of that 

city. In 1947 Richard W. Lemke and I (Brown and 

Lemke, 1948) discovered isolated outcrops of the Cannonball member near Sawyer and Velva, 55 miles 

northeast of the nearest previously reported outcrops 

at Washburn, N. Dak. From the well log records accumulated by Virginia Kline (1942) it appears that 

any Paleocene rocks, if ever present, were removed 

before the Pleistocene from the terrain east of the 

100th meridian in the central part of the State; but 

the Turtle Mountains of Bottineau and Rolette Counties in the northern part are an elevated, more or less 

flat, isolated area in whose south-facing escarpment 

the bedrock contains lignite of Fort Union age (Hendricks and Laird, 1943, p. 593) underlain by brownish 

Cannonball strata containing foraminifers (Lemke, 

1960, p. 28, 31). Thus, it is probable that subsurface 

records may reveal the presence of Cannonball deposits 

even farther north in adjacent parts of Canada. 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 19 

The Cretaceous-Paleocene boundary, so far as can 

be determined in the glaciated area, is an undulating 

line running north westward to the Canadian border 

and- thence into Saskatchewan and Alberta. The 

Canadian strata chiefly involved are the St. Mary 

River, Edmonto:p., 'Vhitemud, Paskapoo, and Ravenscrag formations, succinctly reviewed by E. W. Berry 

( 1930b), W. A. Bell ( 1949), and L. S. Russell ( 1953). 

The Ravenscrag has been divided into a Lower and 

Upper Ravenscrag. The Whitemud yields a characteristic flora of Fox Hills age, whereas the Lower 

Ravenscrag is coal-barren (Fraser, and others, 1935, 

p. 39) but contains plant and dinosaurian remains. 

The Upper Ravenscrag is lignitic, lacks dinosaurian 

remains, but yields the typical Fort Union flora. Its 

base is marked by a coal seam called the Ferris or 

No. 1 (Fraser and others, 1935, fig. 1). 

The earliest account of these differences in stratigraphy and paleontology appears to be by G. M. Dawson ( 1875), who described the geology along the 49th 

Parallel for the British---North American Boundary 

Commission. On page 104 he gives a section which 

he illustrates in plate 7, figure 2. The B portion of 

the section is the critical part and consists of an'upper lignitic sequence with Fort Union plants, and a 

lower nonlignitic arenaceous sequence with dinosaurs, 

turtles, garpike scales, and a fruit which he called 

Aesculus antiqwus (his pl. 16, figs. 8, 9). The latter, 

although not identifiable as a species of Aesculus, 

is an index fruit that occurs abundantly with dinosaurian remains in the Hell Creek formation 3 miles 

southwest of Glendive, 1\tfont., and elsewhere, but not 

in any Paleocene strata so far as I am· aware. This 

contrast between the lower and upper parts of Dawson's B division clearly indicates the CretaceousPaleocene boundary, which here, in modern terminology separates the Lower and Upper Ravenscrag. This, 

however, was not Dawson's conclusion (p. 183-202). 

He located the boundary between what would now 

be called Eastend and mitemud plus Lower Ravenscrag (Hell Creek). Fraser and others (1935, p. 37-

55, and map, fig. 1, opp. p. 24) accept the top of the 

Lower Ravenscrag and base of the Upper Ravenscrag 

as the Cretaceous-Paleocene boundary. 

In northwestern South Dakota, in a gentle synclinal basin 12 to 15 miles north of Camp Crook, the 

Lance as mapped by Winchester and others (1916, pl. 

1) contains coal seams in its upper strata. My examination of this area in 1950 showed that no dinosaur bones occur above the base of the lowest lignite zone but that such remains are abundant beneath that zone. Further, the Fort Union flora occurs in the shales above the base of the coal zone. 

The Cretaceous-Paleocene contact, therefore, is at the 

base of this zone, and none of the sequence above 

this contact is Lance. 

The upper limits of the Paleocene are present in 

the high buttes in the contiguous areas of Montana 

and North Dakota and at a few localities farther east, 

near Dickinson, Hebron, and Beulah (Benson, 1951; 

Johnson and l{unkel, 1954; 1959). In western North 

Dakota is the conspicuous mesa called Sentinel Butte, 

whose lignitic strata are the type Sentinel Butte member of the Fort Union formation. This unit overlies 

the Tongue River member and underlies a massive 

sandstone that may be Eocene in age, and a thin bed 

of fish-bearing shale of White River (Oligocene) age, 

that caps the butte. The further distribution of the 

Sentinel Butte member in western North Dakota is 

described by 0. A. Seager and others (1942, p. 1417), 

and Collier and Knechtel (1939, p. 13) thought that 

50 feet of shale on Antelope Mountain in T. 18 N., 

R. 50 E., 10 miles southeast of Circle, Mont., might 

be Sentinel Butte. 

If the coal bed at the base of the Sentinel Butte 

member be traced northwestward to the Yellowstone 

and Missouri River valleys, its gentle northeastward 

dip of about 15 feet to the mile brings it or its 

equivalent horizon into the low hills north of Buford, N. Dak. Consequently, some of the st!,ata exposed there in the bluffs on the north side of the Mis-

.souri River (pl. 1, fig. 1), that is, at part of Meek 

and Hayden's original type section of the Fort Union 

group, are laterally equivalent to the. Sentinel Butte 

member. One Paleocene mammal,. Titanoides primaevus Gidley, was found in this type section. 

0. A. Seager and others .(i942, p. 1416) and Laird 

( 1944, p. 7) were inclined to. accept a Wasatch age 

for the Sentinel Butte shale on the basis of their areal 

studies in western· North Dakota and adjacent terrain, where they found Leonard's ( 1911, p. 535) 

additional but. undifferentiated light-colored unit, having an approximate maximum thickness of 100 feet, 

lying above the Sentinel Butte shale. This unit they 

regarded as the younger member of the Wasatch. An 

attempt to reinforce the Seager classification and 

clarify the position of these stratigraphic units was 

. made by R. V. I-Iennen (1943) who compiled a series of 

columnar sections across western North Dakota which 

he correlated by means of a so-called marker bed of 

grayish-white quartzitic sandstone containing a zone of 

silicified wood. At the west end of Hennen's group of 

sections, near Sentinel Butte, this bed is about 90 feet 

below the large coal seam that was considered to be the 

top of the Fort Union formation. Unfortunately, this 

"marker bed" was mistaken for another about 100 feet 

higher in the sequence near Sully Springs station 
20 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

southeast of Medora. This mistake invalidates the 

correlation and the structural deductions based on it. 

As similar quartzitic beds have been reported from 

both lower and higher horizons in the Fort Union 

formation in southwestern North Dakota and northwestern South Dakota (Winchester and others, 1916, 

p. 30; Bauer and Herald, 1922, p. 115, and sec. 5, 

pl. 16); Johnson and Kunkel, 1959, p. 12), and as 

they indicate repetition of similar silicifying conditions at intervals ·over a long period of time, correlations based on them across long distances of covered strata must be received with caution. For ·a 

full discussion of these siliceous beds, see Hares ( 1928, 

p. 34-36). The present stratigraphic and paleontologic evidence indicates that the Sentinel Butte shale 

belongs with the Paleocene sequence (Brown, 1948a). 

The beginning of the Eocene is marked by the strata 

of the Golden Valley formation containing the floating fern Salmimia preauriculata Berry, not hitherto 

found in the Paleocene of the Rocky Mountains and 

Great Plains. 

WYOMING 

Eastern Wyoming.-The broad basin of northeastern Wyoming, drained by the Tongue, Powder, Belle 

Fourche, and Cheyenne Rivers, is a continuation of 

·the Plains region of southeastern Montana and displays thick outcrops of the same sedimentary formations (Brown, 1958a) . The Lance Creek area northwest of Lusk, on the discovery of dinosaurs and oil, 

was the first part of the basin to receive close geologic 

attention. Hatcher ( 1896) , and Stanton and Know 1-

ton ( 1897) made close determinations of the limits and 

distribution o:f the :formations, naming the 1,650 :feet, 

more or less, o:f dinosaur-bearing strata first Ceratops beds, then Lance Creek beds, and finally Lance 

:formation. The immediately overlying strata, with 

diagnostic plants were referred to the Fort Union 

:formation. In :following years when coal-mapping 

parties investigated the contiguous areas to the north 

toward the Wyoming-Montana line, the Lance unfortunately was made to . include much of the sombercolored sequence o:f strata (called Fort Union on 

Lance Creek and Tullock in the Gillette coal field) 

overlying the typical dinosaur-bearing Lance. However, plates 9 and 10 accompanying the report on the 

Gillette coal field by C. E. Dobbin and V. H. Barnett 

(1928) show the position of the "lowest persistent 

coal beds," a zone, the base o:f which marks the position of ~he Cretaceous-Paleocene boundary, continuous with that originally determined by Stanton and 

Knowlton in the Lance Creek area. In 1938 and 1948, 

I studied profile sections at spaced intervals :from 

Lance Creek to Moorcroft and found that dinosaurian 

remains and typical Cretaceous plants are present in 

the Lance up to the lowest coal zone of the overlying 

strata. Dorf ( 1942) has studied the Lance flora at 

length. No dinosaur bones were found above the 

Lance, but the Fort Union flora could be had at every 

favorable locality. Thus, the boundary runs irregularly northward from Lance Creek through points 

about 4 miles west o:f Clareton and Moorcroft, thence 

northwestward, keeping to the east side of Little 

Powder River to the Wyoming-Montana line (Love, 

Weitz, and Hose, 1955; Dobbin, Kramer, and Horn, 

1957). 

The Cretaceous-Paleocene contact strikes southwest 

on the west side of Lightning Creek, western tributary o:f Lance Creek in eastern Wyoming, and passes 

through the town of Douglas, thence westward, crossing the North Platte River about 2 miles east o:f Careyhurst. For several miles east of this point, the basal 

500 :feet of the Paleocene are reddish as though discolored by particles derived from erosion of a red 

Chugwater (Triassic) source. Seven miles north of 

Glenrock the basal Paleocene is marked by openings 

on a large coal seam. At the M-C mine on Sand 

Creek I obtained Fort Union plants. Northwestward 

the strata adjacent to the boundary were mapped and 

described by Wegemann (1912; 1917). In his opinion 

the Lance comprised about 3,000 feet o:f gray shale 

and buff sandstone with coal beds in the basal and 

uppermost parts; and the Fort Union included 2,000 

:feet of white sandstone and interbedded gray shale 

topographically expressed as a prominent pine-covered ridge ("Great Pine Ridge") which is transected 

by the Po~der River east o:f Kaycee. Of W egemann's 3,200 feet of Lance, only the lower 2,000 feet 

(about 2,400, according to Horn, 1955) contains dinosaurian remains. The upper 1,200 :feet has yielded no 

ve_rt'ebrate and only a :few plant fossils. One collection o:f plants, from the north side of the highway 7 

miles £rom Kaycee toward ·sussex and 200 feet below 

the white sandstone, contains Paranymphaea erassifolia (Newberry) Berry, which is a characteristic 

species in early Paleocene rocks. Thus it is clear 

that Paleocene time began earlier than the deposition 

of the white sandstone and that the Cretaceous-Paleocene contact is s~veral hundred feet lower in the section; that is, at the level where coal :formation was 

resumed just above the top of the dinosaur-bearing 

coal-barren sequence. Thus the upper 1,200 feet of 

Wegemann's Lance, with coal beds, must be added to 

the Tertiary _sequence, making the Fort Union approximately 2,900 :feet thick in that area. 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 21 

For about 25 miles from a point southwest of Buffalo northward to a point west of Banner, the Fort 

Union, except in the Mowry Basin (Mapel, 1959, p. 61), 

does not crop out, being either absent or widely overlapped by 'Vasatch strata, thus breaking the otherwise 

continuous outcrop around the Powder River Basin. 

The upper limit of the Paleocene on the west side 

of the Powder River Basin between Sussex and Buffalo 

is marked by a slight angular unconformity with the 

overlying coarse conglomeratic IGngsbury conglomerate 

(Gale and Wegemann, 1910, p. 143-146). This basal 

conglomerate of the "Wasatch" formation does not 

occur on the east side of the basin, but at numerous 

localities on the west side, tongues of it can be seen 

interfingering eastward into lignitic strata. Notable 

among such localities are the right bank of Powder 

River at Sussex, hilltops 5 miles east of the point 

where U.S. Highway 87 crosses the south fork of 

Crazy Woman Creek, and hilltops and road cuts south 

and east of Banner. 

That the Kingsbury conglomerate is definitely of 

Eocene age was first suggested by 'Vegemann .(1917), 

who reviewed the fossil-mammal evidence as of that 

date. Jepsen (1940, p. 240, 241) concurred in this 

assignment. In 1941, I found additional Ooryphodon 

material as well as a fragmentary jaw of H yracotherium (eohippus) in the IGngsbury conglomerate 

member on a hilltop 4.5 miles southwest of Buffalo 

(Brown, 1948b) and a jaw of Hyracotherium about 

200 feet above the· conglomeratic base of the recognized varicolored Wasatch on the east side of the Carl 

Volkner ranch, 5 miles north of Sussex. 

At localities on the west side of the basin where the 

Kingsbury conglomerate is absent and lignitic· strata of 

the Fort Union and Wasatch are apparently conformable, the Paleocene-Eocene contact was tentatively considered to be (Baker, 1929, p. 28; Thorn, 1935, p. 66) 

at the top of the Roland coal bed. This bed, named by 

Taff (1909, p. 100) from a mine 2 miles northeast of 

Dietz, has been traced more or less confidently around 

most of the Powder River Basin and some of the 

higher parts of adjoining Montana on the divides between Little Big Horn and Tongue River and between 

Tongue River and Powder River. In areas of the 

Spotted Horse coal field, Olive (1957, p. 10-20) found 

that the Roland coal is discontinuous by reason of an 

erosion interval and, therefore, chose as the PaleoceneEocene contact the top of a limy . shale and sandstone 

containing many fresh-water invertebrates, from 5 to 

65 feet above that coal. Yen ( 1946) identified the 

mollusks as Paleocene in age. The most notable 

change in the fresh-water invertebrate fauna appears 

593121 0 - 62 - 2 

about 200 feet above this level, and there, just below 

the Felix coal, I found the first evidence of Salminia 

preauriculata Berry in the sequence. It may be that 

the strata once or now present between the Roland 

coal and the Felix coal represent the time equivalent 

of the Sentinel Butte member in western North 

Dakota. The boundary, it would thus seem, is a little 

higher in the section than the R.oland coal, the level 

. previously assumed. 

Hanna and Carbon Basirn.s.-In 1918, C. F. Bowen 

reviewed the stratigraphy of this area, the map and 

description of which accompany the fuller report by 

Dobbin, Bowen, and Hoots (1929). Bowen's particular purpose was to show that A. C. Veatch's great unconformity between the Medicine Bow formation 

("Lower" Laramie) and Ferris formation ("Upper" 

Laramie) is actually 6,000 feet higher in the section 

and separates the Ferris and Hanna formations. The 

Medicine Bow formation plus the lower coal-barren 

1,000 feet of the Ferris formation comprise a dinosaur-bearing unit, with Cretaceous plants (Dorf, 1938; 

Brown, 1943a), between the Fox Hills sandstone and 

the base of the Paleocene, which here, as in eastern 

Montana, is marked by the appearance of lignite beds 

after a sequence of coal-barren sandy and conglomeratic strata. In 1938 and 1940, I found typical Fort 

Union plants immediately above these basal lignitic 

beds. The Cretaceous-Paleocene boundary, therefore, 

cuts through the basal part of the Ferris formation in 

the Hanna and Carbon Basins. Only a small triangular patch of what in my opinion is Paleocene Ferris 

remains in the Carbon Basin just south of Como. The 

remainder of the F~rris in that basin appears to be 

the equivalent of the coal-barreri Cretaceous Ferris in 

the western part of the Hanna Basin. One collection 

of apparently brackish-water invertebrates I found in 

the old railroad cut in NW14 sec. 30, T. 22 N., R. 80 

W., 4 miles west of Carbon, is, however, not diagnostic. 

As stated, an unconformity representing erosion 

that truncated the edges of 20,000 feet of gently inclined strata separates the Ferris formation from the 

overlying Hanna formation (Bowen, 1918, p. 232) in 

the Hanna and Carbon Basins. Most of the present 

productive coal mines in these basins are in the Hanna 

formation, now believed to be of Wasatch age. 

In the Rock Creek oil field, southeast of the Carbon 

Basin, Dobbin, Hoots, Dane, and Hancock ( 1929) 

mapped areas of the Hanna formation but found no 

definite evidence of the presence of the Ferris formation. 
22 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Great Dimide Basin.-Ten miles west of Rawlins 

westward-dipping Cretaceous strata are succeeded by 

the coal-bearing Tertiary of the Great Divide Basin. 

The Cretaceous-Paleocene contact is just east of Knobs 

Station on the Union Pacific Railroad, at the base of 

conglomeratic beds just below the first lignites of the 

Tertiary sequence. E. E. Smith (1909, p. 233), discussing the lignitic zone above this horizon, called 

it "undifferentiated Tertiary," and found that it was 

mappable for a considerable distance because there 

occurs at its top a distinctive conglomerate with 

granite pebbles which he regarded as the base of the 

coal-bearing Wasatch. 

On the west side of this basin, that is, on the east 

flank of the Rock Springs uplift, the CretaceousPaleocene boundary crosses U.S. Highwa.y 30 at a 

point 7 miles east of Point of Rocks. There the transition from the somber-colored Cretaceous to the 

lighter colored Paleocene is plainly visible in the ridge 

north of the highway and is at the contact of the 

Black Buttes coal group with the Black Rock coal 

group as mapped by Schultz (1909). This contact, 

as noted by Schultz (1909, p. 264) and Stanton (1909, 

p. 273), is similarly clear about 2 miles southeast of 

Black Buttes Station (pl. 1, fig. 3), where the somewhat conglomeratic basal white sandstone of the 

Black Rock coal group overlaps unconformably the 

shales of the Black Buttes coal group (Cretaceous). 

No diagnostic mammalian remains have been found in 

the Cretaceous and Paleocene strata of this area. 

North of Wamsu~ter, part of the Great Divide. 

Basin is called the Red Desert. Its uranium-bearing 

strata (Masursky and Pipiringos, 1959) are Eocene 

in age, but on its northern border is Bison Basin, an 

eroded anticlinal structure exposing late Paleocene 

strata overlain by younger rocks. In 1952 I was present at and in part responsible for the discovery of a 

rich mammalian fauna in the Paleocene strata ( Gazin, 

1956a). A few plants were found in surrounding correlative beds. 

Rock Springs.-The city of Rock Springs is underlain by the Mesaverde formation on the west flank of 

the Rock Springs dome. In 1952, on Killpecker 

Creek, about 3 miles northwest of Rock Springs, I 

found fossil plants indicating that 1,000 feet, more or 

less, of the nonmarine lignitic strata above a brackishwater sequence of the Lance formation are Paleocene 

in age-the first time strata of ·this age were recognized on the west flank of the uplift. Outcrops of 

these Paleocene strata appear northward from IGllpecker Creek to about 3 miles south of the Boars 

Tusk, from whence they strike northeastward and 

pass under a cover of younger sediments. To the 

south the unconformable contact of l\t[esa verde and 

Tertiary can be seen (pl. 3, fig. 4) just east of Little 

Bitter Creek, at a point 2.5 miles above its mouth. 

These Tertiary strata may be divided into a lower, 

somber-colored lignitic series and an upper, coal-barren, variegated sequence. The lower strata here, 'as 

well as on the southeast side of the Rock Springs 

dome just north of Vermillion Creek, yield fossil 

plants of Fort Union aspect, but the only mammalian 

remains I have found in them are fragmentary teeth 

of Ooryphodon? sp., which are not diagnostic. From 

the overlying varicolored sandy nonlignitic strata, I 

collected in 1941 a skull of 11! eniscotherium as well as 

other mammalian remains diagnostic of Wasatch age 

(Gazin, 1952, p. 13, 62). Until further evidence is 

found, the probability that the Paleocene-Eocene 

boundary separates these two lithologically different 

zones of strata must remain conjectural. 

West of Oasper.-With C. E. Dobbin and F. S. 

MacNeil, I examined the Cretaceous-Paleocene section 

on Poison Spider Creek in 1936 and found Paleocene 

plants in the strata .previously identified as Fort 

Union. In the 'Vind River Basin, isolated patches of 

Paleocene rocks make an irregular elliptic pattern along 

the border of the Wind River formation (Eocene). 

At Twin Buttes, 30 miles northwest of Riverton, the 

outcrops yield shark teeth, indicating an origin comparable to that of the Cannonball member in North 

Dakota (Keefer, 1961, p. 1315, 1322). 

Southwestern Wyoming.-The Paleocene of southwestern Wyoming includes the Evanston and Almy 

formations described by A. C. Veatch ( 1907) and 

Tracey and Oriel ( 1959). The· Evanston yields diagnostic plants in the vicinity of the coal mines near 

Almy and in a Union Pacific· Railroad cut about 1 

mile east of the Evanston station. Strata in Fossil 

Basin probably equivalent to the upper part of the 

type Evanston contain Ti:ffanian mammals ( Gazin, 

1956a), and the Almy formation on La Barge Creek 

near La Barge yields manu;n~tJian remains said by 

Gazin ( 1942; 1956b) to be · Clarkforkian in age. 

Northward in Hoback Canyon a thick sequence of 

sandstones, conglomerates, and shales, called by Eardley and others the Hoback formation, contains only 

fragmentary plant remains; ~ut Dorr (1952; 1958) 

has found a considerable mammalian fauna indicative 

of Paleocene age. It seems:: iikely that these strata 

are in part, at least, correlative with the Evanston 

and Almy formations to the south and with part of 

the Pinyon conglomerate just south of Yellowstone 

National Park. In the latter at vVhetstone Falls on 

a tributary of Pacific Creek, J. D. Love and I found 

well-preserved Paleocene plants. 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 23 

Overlying the Almy formation is a unit composed 

of light-colored rhyolitic ash and thin fresh-water 

limestones, which Veatch called the Fowkes forma- · 

tion. The limestones contain invertebrates, fish remains, reptiles, and a few pl~nts. One of the latter 

is a fern, Acro'8tichum~ hesperi.Jum Newberry, a species 

unreported from pre-Eocene strata. This occurrence 

suggests strongly that the Fowkes is of Eocene age 

and that the Paleocene-Eocene boundary separates the 

Almy and Fowkes formations. 

Bighorn Basirn.-Jepsen lu~,s called the Paleocene 

strata in the north·west part .of the Big Horn Basin 

the Polecat Bench formation. Of its boundaries he 

says (1940, p. 232) : 

No angular unconformity between the Cretaceous and the 

Paleocene sediments has been detected at Polecat Bench, although the fossils suggest a discoriformity or hiatus. At the 

top of the Polecat Bench formation there may be, locally, angular unconformities between it and the overlying Gray Bull 

beds, but in most places where the contact has been determined the units are conformable. Conglome·rates and thick 

beds of coarse bluff sandstones mark the base of the Gray Bull 

in several localities. 

On the other hand, Hewett (1926, p. 38) reports the 

following conditions in the southwest part of the 

Basin: 

The upper limit of the Fort Union is the basal sandstone, 

generally conglomeratic, of the Wasatch formation. This sandstone is unconformable with the Fort Union, for it may be 

traced from the northeast corner of T. 48 N., R. 100 W., where 

it overlaps the base of the Fort Union formation, both northward and eastward to points whe~·e the Fort Union is about 

2,000 feet thick. Here and there the surface of unconformity 

is readily recognizable * * *. [See Hewett, pl. 16.] 

More recently, Van Houten (1944, pl. 7) has 

mapped the Eocene formations above the Polecat 

Bench formation in the Bighorn Basin as the Willwood and Tatman formations, the base of the Willwood being regarded, at least locally, as the PaleoceneEocene boundary. The upper part of Van Houten's 

vVillwood is equivalent to those strata generally understood to be Eocene "Wasatch," but the basal strata 

contain Clark Fork or upper Paleocene mammals. 

Van Houten's comparisons with other areas, however, 

do not include any reference -to the Wasatch east of 

the Bighorn Mountains in the Tongue River and 

Powder River Basins, but this 'Vasatch appears to be 

stratigraphically, lithologically, and paleontologically 

of the same age as that part of the Willwood above 

the zone of Clark Fork mammals. 

COLORADO AND NEW MEXICO 

Denver B asirn.-The strata (table 1) involved in the 

lo.cation of the Cretaceous-Paleocene boundary in this 

area crop out in a roughly oval, synclinal basin extending from Brighton to Colorado Springs and from 

the Rocky Mountain front to points 50 miles eastwa-rd 

on the Plains. 

TABLE I.-Latest Upper Cretaceous and earliest Tertiary 

formations in the Denver Basin, Colorado 

[Dotted line indicates the position of the Cretaceous-Paleocene contact, according to 'Brown (1943a, table 2, p. 78)] 

North 

Denver formation (400-1,200 ft): Arenaceous clays, shales, 

basalt flows, 

andesitic sandstones. 

Arapahoe formation (500±ft): 

Conglomerate, arenaceous shales. 

Laramie formation (600-1,200 ft): 

Sandstones, shales, clays, coals. 

Fox Hills sandstone (800 ft): 

Arenaceous marine shales, 

sandstones. 

South 

Dawson arkose (2,000±ft): 

Arenaceous clays, shales, coals, 

rhyolite flows, 

andesitic and arkosic sandstones, 

conglomerate, arenaceous shales. 

Laramie formation (300±ft): Sandstones, shales, clays, coals. 

Fox Hills sandstone (200-300 ft): 

Arenaceous marine shales, 

sandstones. 

Along the western side of the basin, there lies upon 

the Laramie formation a conglomerate, called the 

Arapahoe conglomerate, whose vertical to gently dipping outcrops can be seen at intervals from Golden 

to Jimmy Camp Creek, east of Colorado Springs. It 

is inferred that this conglomerate reflects diastrophic 

movements to the west and represents a change from 

the lignitic conditions of the Laramie, but it is not 

clear how much erosion of the Laramie occurred before the conglomerate was deposited. Emmons, Cross, 

and Eldridge ( 1896, p. 29, 31) . estimated that probably 600 feet may have been removed locally. Because this erosional contact offered the first visible 

evidence of discordance after the deposition of the 

Laramie, it was made the ·upper boundary of that 

formation in accord with King's definition that the 

Laramie comprises the conformable lignitic strata 

above the Fox Hills sandstone; and it also became to 

many geologists the Mesozoic-Cenozoic boundary, despite the fact that remains of ceratopsian dinosaurs 

were known to occur indigenously in the base of the 

Denver formation several hundred feet higher in the 

section. The fossil plants found in the strata between 

the conglomerate and those containing the latest dinosaurs were originally referred to the Tertiary flora, 

whose remains, however, in this area are now known 

only from the Denver formation and Dawson arkose 

above the dinosaur-bearing strata. 

The paleozoologists were reluctant to include any 

dinosaur-bearing beds in the Tertiary, and consequently the question of the location of the CretaceousPaleocene boundary remained under active discussion 

and investigation. In my fieldwork in the Denver 

Basin, I applied the same methods I had used successfully in Montana and 'Vyoming, namely, a con-
24 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

centrated study of the zone in the vicinity of the latest 

dinosaurian remains. On June 13, 1939, on the bare 

slope on the southeastern flank of South Table Mountain, in the SW1!t,NW!4 sec. 31, T. 3 S., R. 69 W., 

about 3 miles east of Golden, I found some mammal 

jaws and teeth, together with fragments. of turtles 

and crocodiles, and impressions of Tertiary leaves in 

light-colored sandy clay just 50 feet above an outcrop 

of drab-green sandy beds containing the highest dinosaurian remains in the Denver formation. In 1940, I 

found additional mammal specimens at the same spot. 

These, together with others, found at ·approximately 

the same stratigraphic level, but in the upper part of 

the Dawson arkose, in SW!4NE!4 sec. 6, T. 13 S., R. 

64 W., on Corral Bluffs, 11 miles east of Colorado 

Springs, were identified and described by Gazin 

(1941b) as Paleocene forms, indicating an age equivalent to. the Puercan or oldest division of the Tertiary 

in the paleozoological standard. 

From this evidence, checked by that of the fossil 

plants, it is possible to locate the Cretaceous-Paleocene 

boundary in the Denver Basin fairly closely (Brown, 

1943a, fig. 1), but this contact is not marked by an 

evident unconformity nor always by distinguishable 

differences in lithologic composition. 

The youngest indurated strata of the Denver Basin 

occur in the vicinity of Castle Rock and were originally called the Monument Creek group. Subsequently, the upper part of this group was differentiated as the Castle Rock conglomerate of Oligocene 

age, whereas the lower part was included in the upper 

part of the Dawson ar~ose. This phase of the Dawson arkose has been described as unconformably overlying Laramie, Arapahoe, and Denver strata (Emmons, Cross, Eldridge 1896, p. 195-199). No mammals have been taken from these beds, and such plants 

as have been found are equivocal, although they indicate that the strata are not much younger than the 

und.erlying mammal and plant-bearing Dawson and 

Denver. The base of these strata may mark the 

Paleocene-Eocene boundary, but tentatively I regard 

the Paleocene as extending upward to the base of the 

Castle Rock conglomerate. 

Walsenburg, Trinidad, arnd· Raton.-Along the 

Rocky Mountain front west of Walsenburg and Trinidad, Colo., and northwest of Raton and Cimarron, 

N. Mex., is a dissected elevated area that includes the 

Raton Mesa and adjacent coal fields. The sequence of 

sedimentary strata in this region, pertinent ·to the 

present discussion, has been classified as follows, beginning at the bottom : Pierre shale, Trinidad sandstone ( 100 ft +) , V ermej o formation ( 425 ft +) , 

Raton formation (1,800 ft+), Poison Canyon formation (2,000 ft+), Cuchara formation ( 500 ft+), and 

Huerfano formation (3,500 ft+). The chief concern 

here is with the coal-bearing Vermejo and Raton for~ 

mations. Between these formations W. T. Lee (1917, 

p. 55, 56) recognized an erosional boundary that was 

magnified into a, considerable unconformity and stated 

to be the line separating the Mesozoic and Cenozoic. 

From several weeks' field study of Lee's sections, I 

confirm the presence of the unconformity and the distinctness of the basal conglomerate of the Raton formation as compared with any other coarse sandstone 

or conglomerate in either the Vermejo or Raton. 

There is an indefinable "sugary" texture and aspect 

about the matrix of this conglomerate that, once seen, 

may be readily identified wherever the conglomerate 

occurs throughout the area. Depending upon Knowlton's study of the floras, Lee held that the Vermejo 

formation is of Montana and the Raton of Tertiary 

age. So far as I am a ware, no mammalian remains 

have been found in these strata. Additional collections and a restudy of the floras indicate that Knowlton's conclusions need amendment and that, although 

some of the earliest strata now assigned to the V ermejo may be of Montana age, the bulk of the formation is of Laramie age (Brown, 1943a, p. 82). Immediately above the Vermejo-Raton boundary is a sequence of relatively barren deposits, but overlying this 

zone are the productive coal measures of the Raton 

formation from which the greater part of the Raton 

flora was described. No one questions that this is a 

Tertiary flbra, but doubts arise concerning the age of 

the relatively barren basal strata. In .these, at the 

Bowen mine, locality 109 (Lee, 1917, p. 128, 279), and 

at locality 30 (Lee, 1917, p. 90), about 50 feet above 

the basal conglomerate of the Raton, was found the 

problematic Upper Cretaceous plant, Paleoaster inquirenda Knowlton. As this plant has never been 

collected elsewhere from strata proved to be Tertiary, 

it may indicate here that an indefinite thickness of the 

basal Raton is Late Cretaceous in age. If so, the erosion interval between the Vermejo and Raton formations, while appreciable, does not mark the end of the 

Cretaceous but represents only an episode in Late 

Cretaceous history and is apparently correlative with 

the similar interval that preceded the deposition of 

the Arapahoe conglomerate (Cretaceous) in the Denver Basin. 

In the Raton Mesa and adjacent coal fields, the 

Raton formation is overlain unconformably, according 

to Johnson (1958; 1959, p. 101), by the coal-barren 

massive coarse-textured sandstones and conglomerates 

of the Poison Canyon formation. However, except 

for the lack of coal in the Poison Canyon, the two 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 25 

formations resemble one another in general color and 

lithologic composition. No mammalian fossils have 

been taken from the Poison Canyon formation, and 

the few collections of fossil plants are similar to those 

from the Raton, or can be duplicated from the Paleocene in adjacent regions. The probabilities favor the 

top of the Poison Canyon formation as the PaleoceneEocene boundary. The Cuchara formation (Johnson 

and Stephens, 1954, 1955), unconformably overlying 

the Poison Canyon formation, is generally regarded 

as of early Eocene age, because the next overlying 

beds, the Huerfano formation, contain Bridger or 

middle Eocene mammals. 

Oanon Oity coal field.-This field occupies an isolated Cretaceous-Tertiary (?) area midway between 

the Denver Basin and the Raton Mesa coal field. The 

critical physical feature here, as in the Raton field, is 

an unconformity between the Vermejo (Laramie) £ormation and overlying conglomerate (Washburn, 1910, 

p. 348-350), which ·can be seen clearly in the vicinity 

of Alkali Gap south of Canon City. The conglomerate occupies the same stratigraphic position as that at 

the base of the Raton formation in the Raton field, 

but its lithologic composition is notably different. On 

the. other hand, it also occupies the same stratigraphic 

position as the Arapahoe conglomerate in the Denver 

Basin and is remarkably like it in lithologic coinposition. It would seem, therefore, that orogenic movements on the west side of these basins of deposition in 

the later part of Vermejo and Laramie time were reflected in erosional activity on the upturned edges of 

the sediments and in the subsequent deposition o£ conglomerate on the eroded surface. 

The small patches of strata overlying the conglomerate in the Canon City field were called Denver(~), but as I found no identifiable fossils in them, 

I am uncertain of their age. 

North, Middle, and South Park.-In North Park, 

according to Beekly ( 1915), £rom 4,000 to 5,000 feet 

of fresh-water beds, called the Coalmont formation; 

lie unconformably on the marine Pierre shale (Cretaceous). In some parts of the area, they underlie variegated sandy beds believed to be Wasatch (Eocene); 

at others they underlie whitish beds that may be 

White River (Oligocene) ; and at still other localities 

they underlie the sandy tuffaceous North Park formation (Pliocene n. The lower 3,000 to 4,000 feet of 

the Coalmont are dark colored and coal bearing, 

whereas the overlying 2,000 feet are light-colored barren sandstones and conglomerates. Beekly suggested 

that these lithologic differences might serve to distinguish two formations, but confirmatory paleontologic 

evidence for such ~ division is lacking. The fossil 

plants on both sides of this tentative boundary are 

.identical Paleocene forms. No diagnostic animal 

remains have been taken from the upper beds. This 

suggests another interpretation of the twofold character of the formation, namely, that the total 5,000 

feet, more or less, of the Coalmont may be equivalent 

to the Paleocene part of the Ferris formation in the 

Hanna and Carbon Basins across the ColoradoWyoming boundary to the north. The Paleocene Ferris approximates 5,500 feet in thickness and also has 

a lower coaly part overlain by an upper part that, 

with the exception of one or two relatively small 

seams, is nearly coal barren. Tentatively I am regarding the entire Coalmont formation as of Paleocene age, but the discovery of some species of pollen 

in the upper part of the formation may, on further 

confirmation, indicate that that part of the Coalmont 

is Eocene in age (Hail and Leopold, 1960) . 

Middle Park is separated from North Park by the 

Continental Divide, the former being west and the 

latter east of the Divide. About 5,000 feet of strata 

composed of a basal breccia and conglomerate, andesitic shales and sandstones, and thin coal seams are 

called the Middle Park formation. These beds, at 

least in part, are lithologically similar to the Denver 

and Coalmont formations and contain a similar flora 

(Cross, 1892) . They are of Paleocene age and lie 

unconformably on marine strata of Montana age. 

Narrow strips of Paleocene outcrops may be seen 

between Jefferson and Hartsel on the east side of 

South Park, which lies east of the Continental Divide. 

These are dark, sandy, and conglomeratic strata that 

overlie lignitic Laramie and underlie light-colored 

tuffaceous beds that may be somewhat younger than 

Paleocene. Chiefly through the efforts of J. H. Johnson and C. H. Behre, Jr., collections of fossil plants 

have been made that indicate close similarity to the 

flora of the Denver formation (Stark and others, 

1949). 

Rifle and DeBeque.-The Grand Hogback extending from New Castle on the Colorado River, northwestward to Meeker, is largely composed of the Mesaverde and related coal-bearing formations (Gale, 

1910). Unconformably upon these strata, lie approximately 1,200 feet of drab and varicolored beds, which 

in turn are overlain by the more distinctly yellow, red, 

and green beds of the typical 'Vasatch. A small collection of fossil plants was obtained by T. W. Stanton 

in 1907 in Rifle Gap, 6 miles north of Rifle, in gray 

sandy shales at a level 170 feet above the top of the 

Mesaverde formation; and a larger collection was obtained at the same locality in 1937 by Bryan Patterson, of the Chicago Natural History Museum. In 
26 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

1939 and 1940, I obtained additional material at the 

same locality. Many species in these collections also 

occur in the flora of the Denver formation, indicating 

Paleocene age. These and especially the immediately 

overlying varicolored beds are apparently the stratigraphic equivalents of similar beds (Plateau Valley 

beds of Patterson) in the Plateau Valley southwest of 

DeBeque, that have yielded upper Paleocene mammals. Patterson's Plateau· Valley beds are the Paleocene portion of Whitman Cross's Ruby formation the 

base of which may be, in part at least, equivalent to 

the Ohio Creek conglomerate. Along U.S. Highway 

24 at the first hill east of DeBeque is an outcrop of 

carbonaceous shale about 100 :feet beneath yelow sandstones o:f the Wasatch :formation. The small collection o:f :fossil plants obtained there in 1939 proved 

not to be diagnostic. 

From DeBeque westward to the Colorado-Utah line, 

the Paleocene is apparently represented by an un:fossili:ferous sandstone-shale unit, 155-370 :feet thick, 

lying uncon:formrubly between the· Hunter Canyon :formation o:f the l\{esaverde group and the varicolored 

Wasatch. This unit was described as Tertiary ( ~) 

sandstone by Erdmann ( 1934, p. 53). 

I conjecture, but have no fossils to substantiate it, 

that the Paleocene beds of the Rifle Gap section, or 

their equivalents, may be present, although perhaps 

intermittently, along the west side o:f the Grand Hogback to Meeker, thence to the west o:f the Danforth 

Hills and around to Rangely west o:f Cathedral Bluffs. 

The Ohio Creek conglomerate in the Anthracite, 

Crested Butte, and Grand Mesa regions (Lee, 1912, 

p. 48) is a conglomeratic sandstone 100 to 200 :feet 

thick, occurring in isolated patches and said to rest 

unconformably upon Upper Cretaceous strata and to 

underlie the Wasatch :formation. It has been assigned 

to the Tertiary, but the only :fossils :found in it have 

been fragments o:f carbonized wood. Consequently, its 

equivalence to the Paleocene o:f Rifle Gap, to Erdmann's Tertiary ( ~) sandstone, and to Fisher's Tuscher 

:formation o:f the Book Cliffs in Utah, though stratigraphically indicated, if3 not proved. 

Northwestern Oolorado.-In the region extending 

east and west o:f Cedar Mountain, 5 miles northwest 

o:f Craig, is a 2- to 3-mile-wide belt o:f coal-bearing 

strata, about 800 :feet thick, assigned originally to the 

"post-Laramie," now Fort Union (Gale, 1910, pl. 16; 

Sears, 1924, pl. 35). The basal strata o:f this sequence 

are coarse sandstones or conglomerates that lie uncon-

:formably on Upper Cretaceous rocks o:f Laramie (now 

called Lance) age. No mammalian remains have been 

:found in these strata, but such vetebrates, invertebrates, and plants as have been collected, particularly 

in the sandstones, dark sandy shales, and clays cropping out on the hillsides in sees. 28 and 29, T. 8 N., 

R. 93 ,V., indicate Paleocene age. Overlying the Fort 

Union o:f this region are the variegated sandstones 

and shales o:f the 'Vasatch (Sears, 1924, p. 292). 

Southwestern Colorado wnd New 11/exico.-In 1874, 

E. D. Cope discovered and named the mammal-bearing 

Puerco :formation in the San Juan Basin o:f Colorado 

and New Mexico. The strata were described by Gardner ( 1910b) and Reeside ( 1924), and the mammals 

by Matthew (1937). Reeside (1924, p. 34) suggested 

that the Puerco formation is possibly equivalent to 

the upper beds o:f the Animas :formation, a conclusion 

that appears to be confirmed by the fieldwork o:f C. H. 

Dane ( 1946) and also by the evidence o:f the :fossil 

plants :from both :formations, the bulk o:f those described as Tertiary by l{nowlton (1924) :from the 

Animas :formation having come :from the upper beds. 

Dinosaurian remains have been taken :from the lower 

beds o:f the Animas and their lateral equivalent, the 

Ojo Alamo sandstone; but only a :few :fragmentary, 

indeterminable plants have been found in these basal 

beds. It seems most probable, therefore, that the Cretaceous-Paleocene boundary passes through the basal 

part o:f the Animas :formation and is the lower limit 

o:f the Puerco :formation. 

The ~ariegated clays and sands o:f the Puerco and 

Torrejon :formations are not readily separable lithologically, and together the two :formations are sometimes called the Nacimiento group. This group is 

overlain by the "Tiffany beds" now assigned to the 

upper Paleocene, and these are covered by Eocene 

beds that Simpson (1948) proposed to call the San 

Jose :formation. 

In the Cerillos and Hagan coal fields, New Mexico, 

the Mesa verde :formation is unconformably overlain by 

varicolored, sandy, and conglomeratic strata called the 

Galisteo sandstone. According to Lee (1913, p. 288, 

1917, p. 184) some petrified wood and :fossil leaves 

were :found in these beds. Similarly, a small area on 

the southeast side o:f Elephant Butte Dam displays 

an exposure o:f reddish beds overlying coal-bearing 

strata and containing large quantities o:f fossil wood 

and some dinosaur bones (Lee, 1907, p. 57). Because 

of this resemblance Lee was disposed to regard the two 

occurrences as of the ·same age, namely Late Cretaceous. Recently (Stearns, 1943, p. 310; 1953, p. 467), 

however, mammalian remains, particularly o:f the titanothere Teleodus and the carnivore Uinta.cyon, have 

been :found in the uppermost beds o:f the Galisteo sandstone. They indicate a probable late Eocene age :for 

those strata. As the Galisteo at some localities is 4,000 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 27 

feet thick, its lowermost beds may be considerably 

older. 

Southwest of Gallup, Winchester mapped a number 

of outcrops which Sears ( 1925) has designated Tertiary undifferentiated. Whether any of these strata 

may be Paleocene has not been determined. 

ARIZONA 

From the Navajo country of Arizona and New Mexico, Gregory (1917) reported outcrops of unfossiliferous sandstone and shale, which he called Chuska 

sandstone and Tohachi shale, the former being regarded as of Wasatch and the latter of Puerco and 

Torrejon age on stratigraphic and lithologic grounds. 

Until fossils are found in the remarkably barren 

Chuska sandstone, its age will continue to remain 

doubtful. The Tohachi shale, however, is now known 

to be Late Cretacem~s 1n age (Repenning, 1954). 

UTAH 

The Cretaceous and Tertiary formations of the 

Wasatch Plateau in central Utah were described by 

Spieker and Reeside in 1925 and by Spieker in 1931. 

At that time, a hiatus was recognized between the 

Cretaceous Price River formation and overlying beds 

mapped as Wasatch. Since then, the basal part of this 

"Wasatch," beneath the Flagstaff limestone, averaging 

1,500 feet in thickness, has been called the North 

Horn formation. The lower 800 feet of the North 

Horn has been found to contain dinosaurian remains 

indicative of Cretaceous age (Spieker, 1946, p. 134). 

The upper 600 feet has yielded mammalian remains 

described by Gazin (1941a) as the Dragon fauna intermediate in development between the Puerco and 

Torrejon faunas. According to Gazin, "the variegated 

clays of the Paleocene series [called by Gazin the 

J oes Valley member] resemble those in the lower portion of the North Horn formation but are usually 

not so thick and appear to be more gaudily colored 

and with conspicuous white channel sands." The only 

plants found in the North Horn are macerated, unidentifiable fragments of dicotyledonous leaves. 

Eastward in the Book Cliffs toward the Utah-Colorado boundary, Fisher (1936) mapped intermittent exposures of a light-colored unfossiliferous sandstone 

and shale unit, 130 to 600 feet thick, apparently disconformable between the Cretaceous Price River formation and the varicolored Tertiary Wasatch. He 

called this unit the Tuscher formation and suggested 

that it might .be the equivalent of the Ohio Creek 

conglomerate in the Anthracite-Crested Butte region 

of Colorado. Tentatively, I am regarding the Tuscher 

formation as probably of Paleocene age and perhaps 

equivalent to the similarly situated unfossiliferous unit 

in Colorado immediately east of Fisher's area which 

Erdmann ( 1934, p. 53) called Tertiary ( ~) sandstone. 

It should be stated, however, that Erdmann (p. 51) 

considered the Tuscher formation, in whole or in part, 

to be equivalent to a portion of the Hunter Canyon 

formation ·of the Mesaverde group. 

West ward from the outcrops of the Wasatch formation described by Spieker and on the west side of 

the Sevier River valley in the escarpment of the Gunnison Plateau are outcrops assigned to the Tertiary. 

Richardson ( 1906, p. 281) described these beds as 

light-colored, gray, and red shales with interbedded 

limestones and sandstones, approximating 2,000 feet 

in thickness. How much, if any, of this sequence is 

Paleocene remains to be determined. 

In the vicinity of Henefer and Coalville, Summit 

County, are outcrops of variegated shale, sandstone, 

and conglomerate that Eardley ( 1944, p. 840) called 

the Henefer formation and correlated tentatively, on 

stratigraphic grounds, with the Evanston formation 

(early Paleocene) of southwestern Wyoming. Above 

the Henefer is a conglomeratic sequence with shales 

and coal seams that he assigned to the Almy formation (late Paleocene). 

GULF COAST 

The lower limits of the Tertiary on the gulf coast 

have been described chiefly by Vaughan ( 1900), Stephenson (1915), Trowbridge (1932), and Gardner 

(1935). The Paleocene there is comprised in the Midway formation which is reported as unconformably 

overlying the Cretaceous, and also at most localities 

as unconformably underlying the succeeding Eocene 

deposits of the Wilcox group. The Midway is almost 

entirely marine, and has yielded few fossil plants. 

Simpson (1932) has reported the almost miraculous 

recovery of a Paleocene man?.mal, A11Ai8onchus fortunatus Simpson, from. probab~e l\fidway strata in a 

core from a depth of 2,460 feet in the Junior Oil Co. 

Beard No. 1 well, sec. 9, T. lS~ N., R. 16 W., Caddo 

Parish, La. The only correlation, therefore, between 

the Paleocene of the Gulf Coast and that of the western interior is in the faunas of the l\fidway and Cannonball which are said to have a number of genera 

of mollusks and species of foraminifers in common. 

In the trans-Pecos region of Texas, particularly in 

the vicinity of Fort Davis and the Barilla Mountains, 

Upper Cretaceous sandy and shaly limestones identified as Taylor marl are unco~formably overlain by 

variegated deposits of calcareous clay and volcanic 

ash interbedded with other volcanic rocks (Jones, 

1938, p. 1432; Ives, 1941, p. 343). The basal tuffs of 
28 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

the latter strata contain fossil plants in the area of 

the Barilla Mountains. Six species of these were described by Berry (1919), who concluded from them 

"that the basal tuffs in the Barilla Mountains are postCretaceous and pre-'Vilcox in age and that they and 

the volcanic activity which they represent were probably contemporaneous with the floras and similar volcanic activity reflected in the Raton and Denver formations and elsewhere in the Rocky Mountain Region." As the plant specimens are fragmentary and 

the species indicated few, comparisons with the floras 

of the Raton and Denver formations are dubious and 

inconclusive. 

ATLANTIC COAST 

The age of the Vincentown sand of New Jersey 

was long in dispute. Greacen ( 1941) reviewed the 

whole problem and, although inclined to regard both 

the Vincentown and the underlying Hornerstown marl 

as Tertiary, did not decide whether they should be 

called Paleocene or Eocene. Cooke and Stephenson 

(1928), however, assigned them to the Eocene without 

qualification. Spangler and Peterson ( 1950) inclined 

toward allocating the Hornerstown to the Paleo~~me. 

Near Brightseat in Mary land, strata overlying Upper Cretaceous beds contain foraminifers and megafossils said to indicate Paleocene age (Bennett and 

Collins, 1952). Previous studies by Shifflett (1948) 

and Cushman of drill cuttings from a water well near 

Upper Marlboro also revealed foraminifers probably 

of Paleocene age. However, none of these writers suggested correlations of the Maryland deposits with the 

Hornerstown marl to the north or the Midway formation to the south. Johnson and Richards (1952) suggested that the lower part of the Hornerstown is Midway (Paleocene) and the upper part is Wilcox (Eocene) in age. Loeblich and Tappan (1957) referred 

the Hornerstown and Vincentown to the Paleocene, 

but Miller (1960, text fig. 2) concluded that the bulk 

of the Hornerstown is Paleocene and the uppermost 

part is Eocene in age. 

PACIFIC COAST 

Resting upon a basement of igneous and metamorphic rocks, the Swank is apparently the earliest 

Tertiary. formation of Washington. At some localities 

it contains leaves that inclined Knowlton to regard 

the formation as probably of the same age as the 

Denver and Fort Union formations (Weaver, 1937, 

p. 47-49). An Eocene age appears to be a more likely 

assignment. 

In 1939 the U.S. Geological· Survey recognized the 

Martinez formation of California as Paleocene in age. 

It is probable that nonmarine deposits containing 

leaves near Elsinore, Riverside County, may also be 

Paleocene. 

ARCTIC REGIONS 

Greenland, Alaska.-On the west coast of Greenland, particularly the south coastal region of the N ugsuak peninsula, which is just north of Disko Island 

and well within the Arctic Circle, there are outcrops 

of a Cretaceous-Tertiary section that have been examined by a number of explorers and geologists 

(Berry, 1916b, p. 186). Accounts of the geology and 

paleontology of this area are to be found in the publications of Oswald Heer (1868-83), White and Schuchert (1898), Seward (1926), Seward and Conway 

(1935), Imlay and Reeside (1954), and others. The 

generalized section for the N ugsuak peninsula includes 

the following strata, beginning with the oldest Cretaceous : l{ome (said to be Lower Cretaceous) , Atane 

(approximately middle Upper Cretaceous), Patoot 

(Upper Cretaceous), lower Atanekerdluk (Upper Cretaceous), and upper Atanekerdluk (Paleocene). This 

sequence, comprising shales, lignites, sandstones, ironstones, and thick beds of basalt, lies unconformably 

upon an old crystalline metamorphic basement and is 

intruded by many. dikes. Fossil plants have been 

collected :from all the named divisions of the sequence. 

The Atane beds, or their equivalents, have also yielded 

marine invertebrates that Stanton identified as belonging to the Senonian of Europe, approximately 

equivalent to the Montana group of the 'V estern 

United States (White and Schuchert, 1898, p. 356). 

The Patoot beds likewise have yielded marine invertebrates, but Stanton considered their evidence somewhat equivocal (mite and Schuchert, 1898, p. 362). 

Heer, however, assigned these beds to the uppermost 

Cretaceous. Their flora is relatively rich in species 

and bears considerable resemblance to that from the 

Laramie, Lance, and Hell Creek formations of . the 

Western United States and to the Late Cretaceous of 

Alaska. The bulk of the "Miocene" plants described 

by Heer from Atanekerdluk came from the strata not 

far above a 125-foot conglomeratic sandstone that 

crops out 1,000 feet above sea level and overlies the 

Patoot beds. mite and Schuchert assumed that this 

sandstone probably marks the base of the Tertiary. 

As pointed out by Saporta, the flora of these Tertiary beds resembles that from Sezanne and Gelinden 

in the Paris Basin and is of Paleocene rather than 

"Miocene" age, a term used by Heer apparently in the 

sense of early Tertiary rather than in its modern 

specialized meaning. Except for a few insects, no 

animal remains have been reported from these beds. 

That some of the highest Tertiary 3trata of this series 
LIMITS OF THE PALEOCENE IN SPECIFIC AREAS 29 

1nay be of Eocene age is a probability that should be 

considered by future collectors. 

Cretaceous and Tertiary deposits occur on the east 

coast of Greenland. Mathiesen (1932) contributes references to descriptions of this terrain and a report 

on some of the fossil plants found in those strata. He 

concludes that the plants with which he was concerned indicate an early Tertiary age. 

The Cretaceous and Tertiary floras of Alaska have 

been described chiefly by Hollick (1930, 1936), although a number of other paleobotanists and paleozoologists have contributed identifications of Alaskan 

fossils. P. S. Smith (Hollick, 1936, p. 24; Smith, 

1939) has reviewed the areal geology. Concerning the 

Cretaceous-Tertiary boundary G. C. Martin (Hollick, 

1930, p. 36) concluded: 

Upper Cretaceous time ended with the complete withdrawal 

of the sea from the Alaskan area and probably was closely 

followed by the folding and erosion of the Cretaceous rocks. 

The Cretaceous rocks of Alaska are highly folded everywhere, 

and many of them are cut by intrusive rocks and by metalliferous veins. In many places it is not possible to determine 

the exact date of the folding, intrusion, and mineralization, 

especially as some of the Tertiary rocks have been similarly 

affected. It is believed, however, that at least part of the 

folding, intrusion, and mineralization dates from about the 

end of Cretaceous time. The earliest post-Cretaceous rocks in 

most of Alaska are the widespread Tertiary coal-bearing beds. 

Although these rocks are highly folded in some places and 

have been cut by dikes and veins, they are in general notably 

less indurated, folded, and altered than the Cretaceous rocks. 

In some places there is clear proof of an unconformity at the 

base of the Tertiary rocks, and the writer believes that the 

Cretaceous rocks of Alaska were uplifted and eroded, if not 

folded, immediately at the end of Cretaceous time in all parts 

of the Territory. 

The Cretaceous plants described by Hollick are said 

to be from two distinct horizons (1930, p. 34) the 

upper of which, according to the correlation chart 

facing page 34, is considered equivalent to the Lance 

and Laramie formations of the Western United States. 

The flora of this division comprises many species identical with or at least closely related to species in the 

Lance and Laramie floras. 

The Tertiary flora described by Hollick and generally considered Eocene in age is a composite of a 

number of floras whose separation has not yet been 

attempted, and may not be possible until additional 

fieldwork supplies further stratigraphic and paleontologic information, p·articularly mammalian remains. 

EUROPE 

The Paris and London Basins, as well as a few 

other areas in Europe, display contiguous outcrops of 

Cretaceous and Tertiary strata. Unfortunately, but 

quite naturally, different parts of the sequence are 

present at different localities, and these are diversely 

fossiliferous, so that matching of the several sections 

paleontologically is not altogether satisfactory. Consequently, Europeans have had a Mesozoic-Cenozoic 

boundary problem known as the Danian-Montian question. The strata and fossils involved in this problem have been discussed by Saporta and Marion 

(1873, 1878), 1\::ayser (1908, p. 531-539), Berry (1917, 

p. 175-185), Simpson (1937a, p. 6), Wells (1938, p. 

193-202), Tromp (1949, p. 673-676), Hofker (1960, 

p. 584-5<88), J eletsky ( 1960, p. 1896), Brown ( 1961, p. 

632-A), Chandler (1961, p. 16-23), and others. The 

consensus now seems to be that the Danian, in whole 

or large part, is early Paleocene in age. 

Although Kayser considers the Sparnacian to be 

the upper stage of the Paleocene, Simpson, supported 

by Wells, assigns it unqualifiedly to the early Eocene. In the London Basin the Sparnacian includes 

the Woolwich and Reading beds. Beneath these are 

less than 100 feet of the marine Thanet sands, which 

Kayser assigns to the middle Paleocene, but more 

recent correlators to the upper Paleocene because the 

Cernay conglomerate in France contains a very late 

Paleocene mammalian fauna. At Ardtun Head, 

Island of Mull, Scotland, plant remains have been 

found in the sediments intercalated between lava 

flows and have been interpreted as Thanetian in age 

but are probably Eocene or Miocene. Whatever the 

final conclusion about the limits of the European Paleocene may be, there are fossil-bearing beds near the 

middle of the sequence that contain plants and animals sufficiently different from those of the Cretaceous and Eocene to warrant the term Paleocene series. 

ASIA 

No extended study of the Cretaceous-Tertiary 

boundary in Siberia has been made. 1\::ryshtofovich 

(1918, p. 7-9) reports that on Sakhalin Upper Cretaceous strata are conformably overlain by Tertiary 

conglomerates, sandstones, shales and lignites. Simpson (1937a, p. 9) reports that a Mongolian formation called Gashato has yielded 11 genera and 12 

species of fossil mammals that seem to be definitely 

of Paleocene age. Young ( 1934) discussed the uncertainties about the early Tertiary of China. 

The Cretaceous-Tertiary boundary is involved in the 

history of the Deccan traps of India. Sahni (1940; 

1943) has reviewed the situation and concludes that 

the evidence points toward an early Tertiary age for 

the first outpourings of the Deccan lavas. 

SOUTH AMERICA 

The most likely Paleocene strata in South America 

are those called the Rio Chico formation in Argentina 
30 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

(Simpson, 1937a, p. 10). This formation has yielded 

a variety of mammals described by Simpson and 11 

species of plants described by Berry (1937). 

AFRICA 

N akkady ( 1957) has attempted a correlation of the 

late Cretaceous and early Paleocene deposits of Egypt 

with those in other parts of the world. He concluded that the Danian and ~1ontian of Egypt are 

early Paleocene and are comparable to the lower and 

upper parts, respectively, of the ~1idway formation 

of the Gulf Coast of t}:le United States. This paper 

contains an extensive and helpful bibliography. 

PALEOCENE LOCALITIES, EXCEPT AS NOTED 

[i\:Iany locality numbers in th~· text are not listed : some are under a prior number, as &661) under 317, 81196 under 2416] 

229. 2 miles west of Wa~~s, San Pete County, Utah. 

317 (320, 321, 322, 324, 8663). South Table Mountain, Golden, 

Colo. 

318. Northwest side of Green Mountain, Golden, Colo. 

323. Bluff one-fourth of 8: mile south of Reform School and 

two-thirds of a miJe southwest of South Table Mountain, Golden, Colo.· (Probably Cretaceous.) 

325. 3,000 ft east of D~~glas (Lehigh) coal mine, 3 miles 

west of Sedalia, Colo. 

326. Sandstone at coal seam in Laramie at. Morrison, Colo. 

(Cretaceous.) 

331. 1,900 ft east of Dqnglas (Lehigh) coal mine, Sedalia, 

Colo. 

333 (334, 335, 8788). North side of Potato Hill Gap, 2 miles 

east of Hot Sulphljr Springs, Colo. 

336. Near Sheriff's coal shaft on Sheriff Creek east of Hot 

Sulphur Springs, Colo. 

337 (338, 8787) .. Mount Bross,. Hot Sulphur Springs, Colo. 

436. Red Lodge, Mont. 

541. At head of Buck Creek, Niobrara County, Wyo. 

607. On Sweetgrass Creek, west of Porcupine Butte, Mont. 

750. Fossil Forest Ridg~, Yellowstone National Park, Wyo. 

(Eocene.) 

1468. On Lightning Creek, 2 miles above mouth of Walker 

Creek, Lance Cre~k, Wyo. 

1471. Coal mines at Almy, Wyo. 

1474. At mine No. 5, Alriiy, Wyo. 

1502. 21h miles southeast of Black Buttes, Wyo. 

2414. 24 miles northwest of Glendive, Mont. 

2416 (2426, 8196). 6 miles above the mouth of Sevenmile 

Creek, northeast q

1

f Glendive, Mont. 

2417. On Crackerbox Creek, 18 miles northwest of Glendive, 

Mont. 

2420. Left bank of Yellowstone River at Burns, about 30 miles 

below Glendive, ~font. 

2421. Iron Bluff, 12 mile~ southwest of Glendive, Mont. 

2422. 2 miles east of Gladstone, N. Dak., on north side of 

Heart River. ' 

2423 ( 4984). Bull Mountains, 40 miles north of Billings, Mont. 

2424. Near Medora, N. bak. 

3653 ( 5538, 5542). Unio'n Pacific Railroad cut, 1 mile east of 

station, Evanston1 Wyo. 

3658 ( 5539). At mine No. 7, Almy, Wyo. 

3661. SElA,SWlA, sec. 19, T. 16 N., R. 120 W., Almy, Wyo. 

3667. SW14 sec. 33, T. 16 N., R. 120 W., near Evanston, W~'O. 

3728. Williston, N. Dak. 

3852. On Meeyero Creek, 8 miles southwest of Winchester 

Post Office, Wyo. 

3963. Southeast corner sec. 24, T. 22 N., R. 83 W., Wyo. 

3979. Signal Butte, 5 miles southeast of Miles City, Mont. 

3980. Left bank of Yellowstone River at Miles City, Mont. 

4005. West flank of Cedar Creek anticline, 12 miles southwest 

of Glendive, Mont. 

4010. 4 miles southwest of Winchester Post Office, Wyo. 

4028. On Turner ranch, 20 miles northwest of Jordan, Mont. 

4029. 13 miles north of Jordan, Mont. 

4031. At Kern ranch, 24 miles northwest of Jordan, Mont. 

4035. Abo'ut T. 14 N., R 42 E., Mont. 

4050. Three-fourths of a mile east of Florida River crossing, 

east of Durango, Colo. 

4256. 1% miles northwest of Coleharbor, N. Dak. 

4264 ( 4262). Bluffs on west side of Little Missouri River, 

2 miles south of Medora, N. Dak. 

4265. Bluffs on Little Missouri River, three-fourths of a mile 

south of Custer Trail ranch, on Davis Creek, 4 miles 

south of Medora, N. Dak. 

4268. Face of Picket Butte, facing Little Missouri River, 4 

miles south of Medora, N. Dak. 

4272. One-fourth of a mile southeast of mouth of Davis Creek, 

south of Medora, N. Dak. 

4273. On Davis Creek, south of Medora, N. Dak. 

4280. Bank of Yellowstone River, 10 miles northeast of Glendive, Mont. 

4293. Hedges coal mine, left bank of Yellowstone River at 

Miles City, Mont. 

4310. 3 miles north of Clyde Park, on east side of Shields 

River, Mont. 

4311. 2 miles north of Clyde Park, Mont. 

4315. Sec. 24, T. 8 N., R. 49 E., 18 miles east of Miles City, 

l\Iont. 

4323. NE:tA, sec. 20, T. 8 N., R. 48 E., Mont. 

4325. Signal Butte, south of Miles City, Mont. 

4334. Sentinel Butte, N. Dak. 

4368. NE14 sec. 32, T. 8 N., R. 93 W., 1 mile north of Emerson 

ranch, 6 miles north of Lay, Colo. 

4369. Sec. 5, T. 19 N., R. 90 W., north side of SPparation 

Creek, Wyo. 

4395. 20 miles southwest ,of Rawlins, Wyo. 

4404. NW1A,SE14 sec. 27, T. 33 N., R. 72 W., Wyo. 

4421 (4882, 8666). Sec. 16, T. 33 N., R. 73 w., at mine southwest of Inez, Wyo. 

44G8. SE14 sec. 31, T. 33 N., R. 2 W., Colo. (Cretaceous.) 

4474. SWlA, sec. 24, T. 51 N., R. 99 '\V., Wyo 

4571. Sec. 36, T. 6 N., R. 26 E., southeast of Roundup, l\Iont. 

4514. Sec. 21, T. 7 N., R. 10 E., 3 miles south of Lennep, Mont. 

4515. Ridge crossing near the mouth of Shields River below 

Livingston, Mont. 

4570. Sec. 22, T. 7 N., R. 26 E., 3 miles from head of Razor 

Creek, Mont. 

4582. Sec. 34, T. 6 N., R. 27 E., 6 miles east of Buckey, Mont. 

4617. 3 miles southeast of Lennep, Mont. 

4618. Lebo Creek, Mont. 

4620. On Lebo Creek, east of Crazy Mountains, Mont. 

4625. 9 miles west of Miles City, Mont. 

4626. SW'lA, sec. 22, T. 8 N., R. 47 E., Miles City, Mont. 

4661 ( 6666). SW14 sec. 5, T. 45 N., R. 97 W., near Ilo Post 

Office, Wyo. 

4665. 2 miles north of Ilo Post Office, Wyo. 
PALEOCENE LOCALITIES 31 

4674. 

4676. 

4694. 

4696. 

4699. 

4725. 

4843. 

4860. 

4870. 

4871. 

4874. 

On Shoshone River, Cody, Wyo. 

East side of divide between Red Lodge and Bear Creek, 

Mont. 

Hoof of Black Diamond mine, 3 miles north of Meeteetse, 

Wyo. 

9 miles southwest of Bridger, Mont. 

Fort Keogh Military Reservation, Miles City, Mont. 

South center sec. 25, T. 19 N., R. 100 W., Wyo. 

SW%, sec. 23, T. 22 N., R. 104 W., Wyo. 

Sec. 6, T. 55 N., R. 85 W, three-fourths of a mile northwest of Be<!kton, Wyo. 

SW1,4 sec. 12, T. 57 N., R. 85 W., Wyo. 

Sec. 2, T. 57 N., R. 85 W, Wyo. 

Sec. 13, T. 57 N., R. 85 W., about 51h miles east of Ranchester, Wyo. 

4875. Sec. 13, T. 57 N., R. 85 W., one-half of a mile west of 

4874. 

4876. 

4877. 

4878. 

4881. 

4892. 

4896. 

4897. 

4898. 

4908 .. 

Sec. 10, T. 57 N., R. 85 W., 3 miles northwest of Monarch, Wyo. 

Sec. 14, T. 57 N., R. 85 W., 21,4 miles northwest of 

Monarch, Wyo. 

Sec. 19, T. 57 N., R. 84 W., near Monarch, Wyo. 

South side of sec. 20, T. 57 N., R. 84 W., 11h miles 

southeast of Monarch, ·wyo. 

2 miles southeast of Carneyville, Wyo. 

1 mile southeast of Dietz, ·wyo. 

SW1,4 sec. 2, T. 57 N., R. 84 W., on west side of Tongue 

River, Wyo. 

Sec. 13, T. 57 N., R. 84 W., 3 miles east of Carneyville, 

Wyo. 

Southeast corner sec. 15, T. 57 N., R. 83 W., one-half 

of a mile east of Prairie Dog Creek, Wyo. 

4909. Three-fourths of a mile east of Prairie Dog Creek in 

southeast corner sec. 15, T. 57 N., R. 83 W., Wyo. 

4910. Near Decker, Mont. 

497 4 ( 4976, 8563). Sec. 33, T. 6 N., R. 16 E., in sandstone 

capping Bear Butte, near Melville, Mont. 

4975 (8567). Sec. 29, T. 6 N., R. 16 E., northeast of Widdecombe ranchhouse·; Wyo. 

4977. Sec. 25, T. 5 N., R. 15 E., Mont. 

4981. Sec. 23, T. 8 N., R. 25 E, one-half of a mile northeast 

of Roundup, Mont 

4984 (4985, 8568). NE1,4SW1,4 sec. 23, T. 5 N., R. 26 E., 2 

miles south of Buckey, Mont. 

5025.:: ._2 miles east of Black Buttes, Wyo. 

5029. 35 miles southwest of Black Buttes, Wyo. 

5030. Sec. 35, T. 15 N., R. 102 W., Wyo. 

5043. One-fourth of a mile north of center sec. 32, T. 19 N., 

R. 100 W., Wyo. 

504.6 ( 5122). 5 miles above Aguilar on Gonzales Creek, Colo. 

5063. At Bud Kimball mine in SW1,4 sec. 33, T. 46 N., R. 89 

W., Wyo. 

5094. At Bowen mine, 200 ft above lowest coal, Bowen, Colo. 

5099. Floor of Fishers Peak mine, 3 miles southeast of Trinidad, Colo. 

5121. NE1,4 T. 32 S., R. 66 W., Colo. 

5132. Norman ranch, 12 miles east of Raton, N. Mex. 

5133. Sec. 30, T. 32 N., R. 25 E., N. Mex. 

5134. Yankee mine No. 5, near top of section, Raton, N. Mex. 

5137. Near Yankee, N. Mex. 

5139. On ridge· east of Yankee mine, Raton, N. Mex. 

5140. Honeyfield~ine, Yankee, N. Mex. 

5142. Sec. 29, T. 32 N., R. 25 E., N. Mex. 

5144. In Tin Pan canyon, northwest of Raton, N. Mex. 

5147. West side of Dillon canyon, 1 mile north of mouth of 

Coal canyon, N. Mex. 

5151. South end of Raton tunnel, Raton, N. Mex. 

5194. Sec. 30, T. 6 N., R. 2 E., Wyo. 

5236. Spring canyon, Vermejo Park, N. Mex. 

5255. Sec. 12, T. 16 N., R. 102 W., Wyo. 

5259. NE1,4NE1,4 sec. 27, T. 16 N., R. 102 W., Wyo. 

5291. About sec. 14, T. 31 N., R. 22 E., Red River canyon, 

1 mile east of Jones Canyon, N. Mex. 

5300. Sec. 9, T. 6 N., R. 28 E., near head of Cow Gulch, about 

7 miles southeast of Fattig Post Office, ·wyo. 

5321. Sec. 16, T. 16 N., R. 105 W., Wyo. 

5322. North of road across creek at big bend in road, sec. 35, 

T. 16 N., R. 105 W., Wyo. 

5332. 4 miles east of Parkman, Wyo. 

5374. NW1,4 sec. 10, T. 14 S., R. 65 W., east bank of Jimmy 

Camp Creek, Colo. 

5385. River bluffs, right bank of the Missouri River, southeast of Brockton, Mont. 

5387. Near mouth of Smoke Creek, sec. 3, T. 29 N., R. 54 E., 

Mont. 

5388. 2 miles east of Ralston ranch, south of Brockton, Mont. 

5389. Right bank of Missouri River, south of Brockton, Mont. 

5437 (5438). NW1,4 sec. 5, T. 129 N., R. 88 W., N. Dak. 

5455. Sec. 31, T. 35 N., R. 6 W., Colo. 

5456. Sec. 15, T. 35 N., R. 6 W., Colo. 

5458. NW1,4 sec. 30, T. 35 N., R. 6 W., Colo. 

5460. Southeast from center of sec. 19, T. 35 N., R. 8 W., Colo. 

5461. 110 ft below 5460. 

5469. At Bowen mine, 5 miles north of Trinidad, Colo. 

5480. Minturn, Wyo. 

5495. Sec. 18, T. 22 N., R. 80 W., 5 miles east of Hanna, Wyo. 

5509. 8 miles northeast of Parkman, Wyo. 

5512. 7 miles northeast of Parkman, Wyo. 

5526 (9112). NE1,4SE1,4 sec. 22, T. 1 N., R. 2 E., east of 

Ethete, Wyo. 

5551. 150 ft above coal at mine No. 9, Almy, Wyo. 

5555. Mine No. 8, Almy, Wyo. 

5557. Ridge above coal mines No. 1 and No. 4, Almy, Wyo. 

5578. 4 miles southeast of Black Buttes, Wyo. 

5579. 1% miles east of Black Buttes, .Wyo. 

5582. 2 miles east of Black Buttes, Wyo. 

5594 (5595, 5596). SW1,4NW1,4 sec. 28, T. 31 N., R. 19E., south 

of Chinook, Mont. 

5609. 2 miles north of Musselshell, Mont. 

5612. NW1,4NW1,4 sec. 13, T. 9 N., R. 29 E., Mont. 

5613. SE1,4SW1,4 sec. 15, T. 8 N., R. 31 E., Mont. 

5618. Sec. 15, T. 52 N., R. 72 W., Wyo. 

· 5619. At mouth of Elk Creek on Little Powder River, Mont. 

5667. Top of Animas beds, Ignacio quadrangle, Colorado. 

5678. South side of Cucharas Hiver, 3 miles southwest of 

Walsenburg, Colo. 

5679. One-eighth of a mile northeast of Hockland mine, 3 miles 

southwest of Walsenburg, Colo. 

5683. Canyon west of Old Rouse, Colo. 

5684. Dump of Green Canyon mine near Aguilar, Colo. 

5686. One-half of a mile east of Abeton, Colo. 

5687. 1 mile north of Trujillo, Colo. 

5688. Apishapa Canyon, 3 miles northeast of Abeton, Colo. 

5689. 1 mile northeast of 5688. 

5690. At Delagua mine west of Hastings, Colo. 

5695. 3 miles west of Bowen, Colo. 

5697. Sec. 9, T. 33 S., R. 64 W., Colo. 
32 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

5699. Riley canyon, one-half of a mile north of Cokedale, 

Colo. 

5701. Riley canyon, near Cokedale, Colo. 

5711. One-half of a mile north of Wooton, Colo. 

5712. Wooton, Colo. 

5714. At Turner mine, 11,.~ miles north of Wooton, Colo. 

5716. 14 miles east of Douglas, 2 miles north of junction of 

western and middle forks of Shawnee Creek, Wyo. 

5720. Sec .. 27, T. 34 N., R. 68 W., Wyo. 

5721. Sec. 35, T. 34 N., R. 68 W.,. at Sunset mine, Wyoming. 

5738 ( 8656). In clinker above burned coal at top of a conical 

hill, three-fourths of a mile south of a dry hole in 

NW1,4 sec. 33, T. 9 S., R. 61 W., 6 miles northwest of 

Ramah, Colo. 

5760. llh miles east of Old Washakie Station on north side 

of Muddy Creek, Wyo. 

5796. llh miles south of Starkville, Colo. 

5798. Dump of mine at Primero, Colo. 

5799. 5 miles northwest of Weston, Colo. 

5803. Tercio, Colo. 

5826. Sec. 36, T. 30 N., R. 19 E., on Vermejo Creek, N. Mex. 

5831. Six-tenths of a mile north of Richfield Springs ranch, 

9 miles east of Colorado Springs, Colo. 

5835. 1 mile east of Richfield Springs ranch on Jimmy Camp 

Creek, Colo. 

5836 (8655). Railroad cut three-fourths of a mile west o! 

Falcon, Colo. 

5837. 1 mile southwest of Calhan, Colo. 

5838. Dump of Mosby coal mine, Mosby, Colo. 

5839. Sec. 27, T. 11 S., R. 61 W., one-fourth of a mile east 

of Purdon mine, Colorado. 

5840. 4 miles south of Ramah, Colo. 

5842. Sec. 2, T. 5 N., R. 58 E., 16 miles southwest of Baker, 

Mont. 

5863. Center sec. 9, T. 47 N., R. 70 W., Wyo. 

5882. Sec. 36, T. 12 N., R. 52 E., on O'Fallon Creek, east of 

Terry, Mont... 

5885. Sec. 14, T. 10 N~, R. 58 E., near Terry, Mont. 

5886. NE1,4 sec. 17, T. 10 N., R. 59 E., Mont. 

5889. Sec. 32, T. 11 N., R. 59 E., near Terry, Mont. 

5905. Sec. 2, T. 53 N., R. 71 . W., Wyo. 

5911. Sec. ·29, T. 47 N., R. 81 W., Wyo. 

5917. T. 42 N., R. 78 W., Wyo. 

5987. On west side of Illinois River, southwest of Walden, 

Colo. 

5994. SW1,4NW1,4 sec. 2, T. 10 N., R. 80 W., Colo. 

5997. SE1,4 sec. 31, T. 10 N., R. 81 W., in North Park, Colo. 

6000. SE1,4NW1,4 sec. 28, T. 9 N., R. 80 W., 7 miles west of 

Walden, Colo. 

6005. NW1,4NW1,4 sec. 19, T. 9 N., R. 80 W., in bluff on north 

fork of Platte River, Colo. 

6006. NE1,4 sec. 14, T. 7 N., R. 81 W., Colo. 

6050. SW1,4 sec. 7, T. 49 N., R. 68 W., Wyo. 

6051. NE1,4SE1,4 sec. 6, T. 49 N., R. 68 W., 6 miles west of 

Moorcroft, Wyo. 

6057. NE1,4 sec. 13, T. 46 N., R. 70 W., 20 miles southwest of 

Moorcroft, Wyo. 

6083. SE1,4 sec. 20, T. 45 N., R. 69 W., Wyo. 

6084. Sec. 3, T. 42 N., R. 67 W., 20 miles above Hampshire 

on Black Thunder Creek, Wyo. 

6099. NE1,4SE1,4 sec. 9, T. 7 N., R. 81 W., in bluff 3 miles 

northwest of Coalmont, Colo. 

6102. SE1,4NE1,4 sec. 30, T. 6 N., R. 80 W., Colo. 

6103. NE1,4NE1,4 sec. 19, T. 6 N., R. 79 W., Colo. 

6105. NE1,4 sec. 25, T. 7 N., R. 80 W., about 3 miles north of 

Riding ranch, in North Park, Colo. 

6107. SW1,4NE1,4 sec. 36, T. 8 N., R. 78 W., Colo. 

6110. SE1,4SE1,4 sec. 29, T. 9 N., R. 78 W., near Capron mine, 

Colorado. 

6113. Head of Wolf Draw in about sec. 24, T. 17 N., R. 9 E., 

S.Dak. 

6114. Sec. 7, T. 17 N., R. 11 E., S. Dak. 

6116. Sec. 17, T. 17 N., R. 13 E., S. Dak. 

6117. Sec. 17, T. 21 N., R. 16 E., S. Dak. 

6131. NE1,4 sec. 20, T. 12 N., R. 50 E., near Terry, Mont. 

6142. 21,4 miles southeast of Monument, Colo. 

6154. Sec. 1, T. 17 N., R. 7 E., S. Dak. 

6155. Center sec. 1, T. 17 N., R. 9 E., S. Dak. 

6156. Sec. 12, T. 17 N., R. 7 E., S. Dak. 

6161. Sec. 25, T. 19 N., R. 7 E., S. Dak. 

6171. North half of sec. 24, T. 50 N., R. 100 W., Wyo. 

G173. Sec. 31, T. 50 N., R. 99 W., Wyo . 

6176. SE1,4 sec. 6, T. 49 N., R. 99 W., Wyo. 

6215. Sec. 36, T. 136 N., R. 105 W., south of Yule, N. Dak. 

6219. Sec. 6, T. 137 N., R. 102 W., N. Dak. 

6225. SW1,4 sec. 7, T. 134 N., R. 102 W., N. Dak. 

6297. 3 miles north of Craig, Colo. 

6299. 7 miles northwest of Craig, Colo. 

6309. One-half of a mile downstream from station at Pagosa 

Junction, Colo. 

6344. SW1,4NW1,4 sec. 22, T. 33 N., R. 73 W., 2 miles southeast 

of Inez, Wyo. 

6359. Near center sec. 14, T. 51 N., R. 100 W., Wyo. 

6360. Sec. 24, T. 50 N., R. 100 W., Wyo. 

6376. SW1,4SE1,4 sec. 33, T. 132 N., R. 91 W., 10 miles south 

of Bentley, N. Dak. 

6377. Sec. 6, T. 133 N., R. 88 W., 1 mile south of l{ayser, 

N.Dak. 

6382. 3 miles east of Scranton, N. Dak. 

6384 (9125). NW1,4 sec. 34, T. 22 N., R. 9 E., at Anarchist 

Butte, S. Dak. 

6416. NW1,4SW1,4 sec. 24, T. 23 N., R. 84 W., Wyo. 

6417. SE1,4SE1,4 sec. 23, T. 23 N., R. 84 W., in north bank of 

Big Ditch, Wyo. 

6419. NW1,4SW1,4 sec. 26, T. 23 N., R. 84 W., Wyo. 

· 6420. NW1,4NW1,4 sec. 16, T. 22 N., R. 83 W., Wyo. 

6428. SE1,4SW1,4 sec. 11, T. 23 N., R. 84 W., Wyo. 

6431. SW1,4NE1,4 sec. 27, T. 23 N., R. 84 W., Wyo. 

6437 (5994). SW1,4NW1,4. sec. 2, T. 10 N., R. 80 W., Qolo. 

6440. NE1,4SW1,4 sec. 24, T. 7 N., R. 81 W., Colo. .-.. 

6443. 1 mile northeast of Pagosa Junction, Colo. 

6444. 1 mile northwest of Pagosa Junction, Colo. 

6525. South bank of Heart River, one-half of a mile south of 

Mandan, N. Dak. 

6592. Sec. 4, T. 7 N., R. 94 W., Colo. 

6594. Sec. 2, '1'. 7 N., R. 94 W., Colo. 

6598. SW1,4 sec. 3, T. 131 N., R. 85 W., N. Dak. 

6625. SEt,4 sec. 29, T. 23 N., R. 83 W., Wyo. 

6630. NE1,4NW1,4 sec. 12, T. 22 N., R. 84 W., Wyo. 

6652. NE1,4 sec. 30, T. 136 ~., R. 82 W., N. Dak. 

6667. North of Ilo Post Office, Wyo. 

6668. 200 ft above base of Fort Union, llo quadrangle, Wyoming. 

.. 6669. Near Ilo Post Office, Wyo. 

6738. SW1,4 sec. 9, T. 35 N., R. 75 W., 'Vyo. 

6765. Railroad cut three-fourths of a miJ:e south of Wilsall, 

Mont. 

6765-A. Three-fourths of a mile north of Wilsall, Mont. 

··~ 
PALEOCENE LOCALITIES 33 

6767. On Brackett Creek, 12 miles west of Clyde Park, Mont. 

6845. Sec. 1, T. 27 N., R. 90 W., on Coal Creek, northwest of 

Rawlins, Wyo. 

6892. SW% sec. 14, T. 139 N., H. 84 W., N. Dak. 

6905. SW~4 sec. 22, T. 138 N., R. 85 W., N. Dak. 

6943. One-half of a mile west of Ramah, Colo. 

6944. Coal gulch 2 miles northwest of Ramah, Colo. 

6971. NW% sec. 34, T. 22 N., R. 82 W., east of Dana, 'Vyo. 

7004. Sec. 33, T. 37 N., R. 47 E., Mont. 

7005. One-half of a mile north of northwest corner of sec. 4, 

T. 37 N., R 49 E., Mont. 

7371. 3 miles southwest of Farmington, N. Mex. 

7480. Top of hill west of Talian mine, 5 miles north of Pagosa 

Junction, Colo. 

7481. Above Talian mine, 5 miles north of Pagosa Junction, 

Colo. 

7483. 41h miles north of Pagosa Junction, Colo. 

7 484. Near 7 483. 

7485. Near 7483. 

7495. l mile northwest of Ojo Alamo, N.Mex. 

7496. One-half of a mile west of Pagosa Junction, Colo. 

7498. On Cat Creek, 1 mile north of Pagosa Junction, Colo. 

7538. SE1,4 sec. 22, T. 16 N., R 33 E., Mont. 

7547. NW%SE1,4 sec. 22, T. 3 N., R 16 E., 14 miles northeast 

of Big Timber, Mont. 

7548. NW%NE14 sec. 4, T. 2 N., R i6 E., 14 miles northeast 

of Big Timber, Mont. 

7552 (8540). At clay pit, 1 mile south of Dickinson, N. Dak. 

(Eocene.) 

7623. Sec. 35, T. 9 N., R. 89 W., at Pugh's place, northeast of 

Craig, Colo. 

7659. South half sec. 7, T. 1 N., R 40 E., 10 miles west of 

Colstrip, Mont. 

7662. SE1,4 sec. 36, T. 1 N., R. 39 E., Mont. 

7663. 4 miles west of Camp Crook, S. Dak. 

7685. NE% sec. 35, T. 6 S., R 43 E., Mont. 

7688. Sec. 20, T. 6 S., R. 43 E., Mont. 

7695. Sec. 20, T. 6 S., R 43 E., Mont. Near 7688. 

7i76. 2 miles southeast of coal mine at BlackButtes, Wyo. 

7839 (8249). 2 miles southwest of Edwards, Mont. 

7989. Weldon, McCone County, Mont. 

8163. SW1,4 sec. 12, T. 5 N., R 43 E., Mont. 

8164. Sec. 23, T. 3 N., R 45 E., Mont. 

8165. SW1,4 sec. 15, T. 5 N., R. 47 E., Mont. 

8166. Southeast of Colstrip, Mont. 

8167. Southwest corner of sec. 35, T. 3 N., R 44 E., Mont. 

8187. NW1,4 sec. 30, T. 11 S., R. 60 W., Colo. 

8188. NE% sec. 25, T. 11 S., R 61 W., south of Ramah, Colo. 

8190 ( 8542). 1 mile north of In take, Mont. 

8191. On left bank of Yellowstone River, north of Burns, 

Mont. 

8193. Right bank of Yellowstone River, east of Sidney, Mont. 

8199. 4 miles north of Stipek, along Sevenmile Creek, Mont. 

8200. 3 miles up Clear Creek, west of Glendive, Mont. 

8202. In left bank of Missouri River, 10 miles north of Sanish, 

N.Dak. 

8203. Between Sanish and Hoflund, N. Dak. 

8204. Near 8203. 

8205. 18 miles along river road from Sanish to Hoflund, 

N.Dak. 

8206. 6 miles north of Armstrong, N. Dak. 

8212. Near Elbowoods, N. Dak. 

8213. 5 miles southeast of Elbowoods, N. Dak. 

8215. 9 miles north of Sixmile Creek, Armstrong, N.Dak. 

~220. At Hoffman, west of Sanish, N. Dak. 

8222. Blue Buttes, near Keene, N. Dak. 

8224. At Brittian mine, 6 miles southeast of Mott, N. Dak. 

8225. At Hoffman mine, right bank of Missouri River, across 

from Sanish, N. Dak. 

8227. In North Dakota, 3 miles north of Watauga, S. Dak. 

8230. Near Grassy Butte, N. Dak. 

8231 ( 8232, 8233). Near Bicycle Post Office, N. Dak. 

8234. West of Grassy Butte, N. Dak. 

8238. In clinkered sandstone and shale near top of Sentinel 

Butte, N. Dak. 

8239. Left bank of Little Missouri River, three-fourths o·f a 

mile south of Yule, N. Dak. 

8210. On Little Missouri River, 3 miles southwest of Yule, 

N.Dak. 

8245. One-half of a mile north of Ekalaka, Mont. 

8246. Left bank of Yellowstone River at Terry, Mont. 

8247. Left bank of Yellowstone River near 8246. 

8248. At Roberts coal mine, 3 miles west of Amidon, N. Dak. 

8250. Signal Butte, southeast of Miles City, Mont. 

8252. 8 miles south of the mouth of Armell Creek, west of 

Forsyth, Mont. 

8253. 2 miles east of Powder River bridge, Mizpah, on road 

to Ekalaka and Ismay, Mont. 

8255 ( 8520). Colstrip, Mont. 

8256. 8 miles south of the mouth ot Armell Creek, west of 

Forsyth, Mont. 

8257. 5 miles northeast of Kinsey, Mont. 

8258 (8259, 8549). 1 mile east of Hathaway, Mont. 

8261. 2 miles west of Cap Rock, 18 miles northeast of Miles 

City, Mont. 

8262. At Cap Rock, on north side of the Yellowstone River, 

20 miles east of Miles City, Mont. 

8307. NW1,4 sec. 7, T. 12 S., R. 61 W., southeast of Calhan, 

Colo. 

8426. 6 miles south of Como, in South Park, Colo. 

8447. 6 miles south of Como, n~ar 8426. 

8493. 6 miles south of Como, near· 8426. 

8516. 1 mile east· of filling station· near Dana, Wyo. 

8517. Noi'th side of u.s.· Highway 10, in bank of Moon Creek, 

10 miles west of Miles City, Mont. 

8519. Hillside south of U.S. Highway 10, 25 miles east of 

Miles City, Mont. 

8521. Near top of conical .hill, sec. 23, T. 2 S., R. 44 E., Mont. 

8522. On Dry Creek, 5 miles west of Greybull, Wyo. 

8523. Sec. 19, T. 5 N., R. 13 E., on Sweetgrass Creek, Mont. 

8526. 600 ft south of .Jepsen mammal quarry on Polecat bench, 

west of Frannie, Wyo. 

8529. 10 miles east of Roundup, Mont. 

8535. On Poison Spider Creek, southwest of Casper, Wyo. 

8545. Sec. 23, T. 4 N., R. 40 E., east of highway Forsyth to 

Colstrip, Mont. 

8547. Near top of "Bear" formation on south side of Lebo 

Creek, Mont. 

8548. Near center of sec. 10, T. 22 N., R. 82 W., near Hanna, 

Wyo. (Eocene. ) 

8550. 18 miles northeast of Miles Dity, Mont. 

8551. On Sand Creek, 7 miles north of Glenrock, Wyo. 

8552. 3 miles west of Marsh station on Northern Pacific Railroad, west of Glendive, Mont. 

8553. West of Cap Rock, about 18 miles northeast of Miles 

City, Mont. 

8554. 25 ft above Big Dirty coal, on north side of Yellowstone 

River at Terry, Mont. 
34 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

8556. 10 miles north of Terry, Mont., in sec. 19, T. 13 N., 

R. 50 E. 

8558. Near center of sec. 21, T. 6 N., R. 13 E., Mont. 

8566. In clinker near Wyodak coal mine, 4% miles east of 

Gillette, Wyo. 

8652. 2 miles southeast of Black Buttes, Wyo. 

8654. South side of U.S. Highway 30, 7% miles east of Point 

of Rocks, Wyo. 

8660. Above coal at mine 1 mile northeast of U.S. Highway 30 

at Hadsell, 10 miles west of Rawlins, Wyo. 

8662. In bluff on north side of Big Ditch, in sec. 32, T. 23 N., 

R. 83 W., Wyo. 

8668. 10 miles south of Rozet, Wyo. 

8669. 4 miles west of Bayfield, Colo., in dump from large 

road cut. 

8670. At mine No. 1, Alroy, Wyo. 

8672. Sec. 36, T. 2 S., R. 66 W., in bank of Second Creek, 

10 miles east of Derby, Colo. 

8673. Sec. 3, T. 35 N., R. 66 W., west of Lance Creek, Wyo. 

8677. 8 miles west of Clareton, Wyo. 

8678. Near 8677. 

877 4 ( 4370, 4457). In almost vertical beds 170 ft above top 

of Mesaverde formation, on south side of creek in Rifle 

Gap, 6 miles north of Rifle, Colo. 

8775. Nigger Hill, 1 mile east of DeBeque, Colo., along U.S. 

Highway 6. 

8776. 20 ft above level of Cucharas River, 1% miles west of 

Walsenburg, Colo. (Cretaceous.) 

8777. West side of Platte River, near Colfax Pressed Brick 

Co., Denver, Colo. 

8779. Three-fourths of a mile east of ranch on Jimmy Camp 

Creek, 9 miles east of Colorado Springs, Colo. 

8780. South side of Beebe-Powderville road, 4 miles west of 

Powderville, Mont. 

8781. 2 miles up Little Powder River from junction with 

Powder River, northeast of Broadus, Mont,. 

8786. SW1,4 sec. 27, T. 5 S., R. 50 E., about 6 miles southwest 

of Broadus, Mont. 

8881. One-half of a mile west to northwest of Ramah, Colo. 

8882. One-half of a mile east of Purdon mine, 5 miles southwest of Ramah, Colo. 

8884. 2 miles east of Ohlman, Wyo. 

8885 ( 6342). Makton coal mine, 7 miles northeast of Big 

Sandy, Mont. 

8886. NW1,4 sec. 23, T. 7 S., R. 45 E., west of mouth of Cub 

Creek, on Otter Creek, Mont. 

8887. Sec. 33, T. 8 S., R. 46 E., Mont. 

8888. 5 miles northeast of Kinsey, Mont. 

8893. North of center of T. 46 N., R. 81 W., Wyo. 

8896. North side of Yellowstone River, at Big Timber, Mont. 

8897. 81h miles north of Glendive, Mont. 

8898. 2 miles west of Douglas, Wyo. 

8899. South of highway at Gwynn ranch on Grass Creek, Wyo. 

8901. East side of Graveyard Coulee, south of Glendive, Mont. 

8910. NW:tA, sec. 11, T. 8 S., R. 49 E., 11h miles north of 

Traub ranch on Bayhorse Creek, Mont. 

8913. Above coal in small hill to north of road leading up 

to Sentinel Butte, Sentinel Butte, N. Dak. 

8917. 3 miles east of Black Buttes, Wyo. 

8920. Sec. 13, T. 15 N., R. 105 W., one-fourth of a mile north 

of gate on road, 20 miles south of Rock Springs, Wyo. 

8921. Sec. 14, T. 16 N., R. 105 W., southwest of Antelope 

Butte, Wyo. 

8922. Sec. 26, T. 16 N., R. 105 W., east of Bacon ranch, east 

side of Bitter Creek, Wyo. 

8928. Sec. 2, T. 7 N., R. 92 W., northwest of Craig, Colo. 

8930. 7 miles east of Point of Rocks and 1 mile north of U.S. 

Highway 30, Wyoming. 

9056. About 17 miles southwest of Musselshell, Mont. 

9071. Sec. 16, T. 41 N., R. 110 ·w., about one-fourth of a mile 

east of mouth of Buck Creek, Wyo. 

9072. NE1,4 sec. 32, T. 1 S., R. 50 E., 1% miles southeast of 

Coalwood, Mont. 

9084. NW1,4SW1,4 sec. 16, T. 142 N., R. 91 W., N. Dak. 

9085. Near Young Man's Butte, 2% miles east of Richardson, 

N.Dak. 

9092. SE1,4 sec. 15, T. 55 N., R. 75 W., Wyo. About 50 ft 

above Roland coal. 

9104. At coal mine in NW% sec. 4, T. 36 N., R. 52 E., Mont. 

9109. NW1,4SWI,4 sec. 29, T. 1 S., R. 6 E., 13 miles east of 

Riverton, Wyo. 

9111. East of Little Dome, Carter Oil site 2, north of Lander, 

Wyo. 

9129. SE:tA,SEIA, sec. 36, T. 35 N., R. 91 W., Wyo. 

9130. SW1,4 sec. 31, T. 34 N., R. 92 W., Wyo. 

9131. NE:tA, sec. 20, T. 6 N., R. 1 E., Wyo. 

9132. Center of south half SE1,4NW:tA, sec. 35, T. 6 N., R. 1 E., 

on southwest side of Shotgun Butte, Wyo. 

9134. At Blue Pony coal mine in SW1,4SW1,4 sec. 25, T. 29 N., 

R. 14 E., Mont. 

9141. SW:tA, sec. 30, .T. 11 N., R. 1 E., on Cedar Creek a tributary of Salmon Creek, Wash. (Miocene.) 

9180. In carbonaceous shale west of Killpecker Creek, 3 miles 

northwest of Rock Springs, Wyo. 

9193. Whetstone Falls, on a tributary of Pacific Creek, northeast of Moran, Wyo. 

9194. NW1,4NW1,4 see. 1, T. 17 N., R. 7 E., at Mendenhall 

mine, South Dakota. 

9196. NEI,4SE1,4 sec. 17, T. 6 N., R. 1 E., Wyo. 

9198. SE:tA, sec. 8, T. 20 N., R. 90 W., Wyo. 

9199. Sec .. 33, T. 1 N., R. 47 E., Custer County, Mont. 

9200. 8 miles south of Taylor, N. Dak. 

9201. Sec. 24, T. 28 N., R. 14 E., about 4 miles east of Mackton coal mine, east of Big Sandy, Mont. 

9202. 3 miles east of Fort Stevenson on north side of Missouri 

'River, 71 miles north of Bismarck, N. Dak. 

9203. 1 imile south of Como, Colo. 

9204. s.ec. 14, T. 4 s., R. 41 E., Mont. 

9205. NE1,4 sec. 18, T. 142 N., R. 90 W., N. Dak. 

9206. ~14 sec. 31, T. 33 N., R. 55 E., Mont. 

9207. Near 9109, but stratigraphically below. 

9208. South half of sec. 15, T. 34 N., R. 91 w., Wyo. 

9210. S;E1,4 sec. 26, T. 18 N., R. 100 ,V., Black Buttes, Wyo. 

9235. About 2 miles east of Viola, Wyo., on north side of road 

, along LaBarge Creek. 

9236. NW:tA,NW1,4 sec. 22, T. 12 N., R. 100 W., Colo. Hiawatha 

Unit well 2. 

9237. A-bove white sandstone of "Castle Gardens" on Muskrat 

· ' Creek, south of Moneta, Wyo. 

9239. NE% sec. 10, T. 9 S., R. 47 E., Mont. 

9248 (9334). East side of Smoky Butte Creek, 14 miles northwest of Jordan, Mont. 

9249. Southeast corner of sec. 10, T. 16 N., R. 38 E., Mont. 

9252. 1 mile above mouth of Powder River, 6 miles southwest 

of Terry, Mont. 

9253. West side of Little Powder River, near Biddle, Mont. 

9322. At Eagle coal mine, 1 mile southwest of Bear Creek, 

Mont. 
METHODS APPLIED IN THE STUDY OF THE PLANTS 35 

9342. Sec. 28, T. 8 S., R. 93 ,V., one-half of a mile south of 

Blevin mine, Colo. 

9398. SW1,4 sec. 20, T. 142 N., R. 90 W., N. Dak. 

9399. NE14SW1,4 sec. 15, T. 6 N., R. 1 E., Wyo. . 

9400. About 2 miles southwest of Black Rock, which is northeast of Point of Rocks, Wyo. 

9401. Sec. 2, T. 9 S., R. 62 W., about 5 miles northeast of 

Fondis, Colo. 

9402. About 5 miles southwest of Bridger, Mont. 

9403. North side of highway, 5 miles northwest of Rock 

Springs, Wyo. 

9404. About 3 miles west of Elk Basin, Wyo. 

9405. Center of sec. 32, T. 6 N., R. 3 E., Wyo. 

9421 (9180). NE14 sec. 22, T. 19 N., R. 105 W., about llt{2 

miles north of Rock Springs, Wyo. 

9445. Sec. 9, T. 4 N., R. 81 W., about 11h miles northeast of 

Whitely Peak, Colo. 

9446. NWJ4NE1,4 sec. 35, T. 9 N., R. 78 W., at Marr coal mine, 

Colorado. 

9476. At Nygard coal mine, about 5 miles northeast of Big 

Sandy, Mont. 

9477. NE1,4NW1,4 sec. 33, T. 27 N., R. 95 W., Bison Basin, 

Wyo. 

9478. SE1,4 sec. 11, T. 27 N., R. 95 W., Bison Basin, Wyo. 

9479. Center of sec. 14, T. 27 N., R. 95 W., Bison Basin, Wyo. 

9482. NW14SW14 sec. 10, T. 34 N., R. 90 W., Wyo. 

9488. NE14NW1,4 sec. 19, T. 27 N., R. 15 E., Mont. 

9489. SW14 sec. 18, T. 27 N., R. 15 E., northeast of Ryan 

Butte, Bearpaw Mountains, M~nt. 

9490. NW14SW14 sec. 12, T. 27 N., R. 14 E., Mont. 

9491. NW14 sec. 26, T. 28 N., R. 14 E., Mont. 

9492. On west side of highway, 3% miles north of Pagosa 

Junction, Colo. 

9501. NW14 sec. 16, T. 9 S., R. 97 W., along highway on left 

bank of Colorado River, 4 miles southwest of DeBeque, 

Colo. 

9502. Sec. 15, T. 50 N., R. 68 W., 4 miles northwest of Moorcroft, Wyo. 

9532. Center of SW1,4 sec. 27, T. 33 N., R. 95 W., Wyo. 

(Eocene.) 

9533. Sec. 10, T. 42 N., R. 108 W., Wyo. 

9540. Northeast of Muskrat Creek camp at cabin, on road to 

Riverton, Wyo. 

9554. 2 miles north of Highway 94, 3 miles east of bridge over 

Jimmy Camp Creek, 9 miles east of Colorado Springs, 

Colo. 

9558. Center of NE14 sec. 36, T. 21 N., R. 117 ·w., Wyo. 

9564. Sec. 2, T. 130 N., R. 104 W., 8 miles south of Rhame, 

N.Dak. 

9565. Sec. 7, T. 30 N., R. 7 W., 1 mile above confluence of 

Los Pinos and San Juan Rivers, N. Mex. 

9566. 3 miles west of Douglas, just north of U.S. Highway 20, 

Wyoming. 

METHODS APPLIED IN THE STUDY OF THE PLANTS 

The numerous collections of Paleocene plants used 

in this study differed greatly in quantity and quality 

of specimens. Some comprised a large assortment of 

well-preserved leaves, flowers, fruits, seeds, pollen, 

spores, and wood; but many consisted of only a few 

specimens, often very poorly preserved. Nevertheless, 

whatever the condition of the collection, the first step 

in its study was the trimming and preliminary sorting of the specimens and the assignment of a locality 

number, which was put on each specimen. After a 

general survey of the collection to get a tentative idea 

of its contents and of its approximate place in the 

geologic column, I began concentrated examination of 

the specimens to fulfill the threefold -purpose, as I 

see it, of studying fossil plants : first, to segregate and 

identify the kinds as accurately as possible in order 

to establish their place in the evolution of the plant 

kingdom; second, to find items of restricted geologic 

range for the determination of the. relative age and 

correlation of the containing strata; and third, to 

infer from the composition of the flora the probable 

ecologic conditions that prevailed when the plants were 

alive. 

The process of identifying fossil plants is fraught 

with many difficulties. Chief among these is the comparative scarcity of satisfactory specimens in many 

collections; and second to this is the often poor quality 

of types and their illustrations, if these are still a vailable. Not all fragmentary specimens, however, are 

useless, for some may retain the very features necessary for their identification. Sometimes the fragments 

have proved to be worse than useless, for species 

founded on them have caused endless quibbling and 

frustration about their identities and the priority of 

the names given to them. In my opinion many paleobotanical names are invalid because they apply to fragments that were not and cannot be adequately characterized for practical use. Instead of scarcity there 

may, on· the other hand, be a surplus of good but 

variant specimens, and this may compel the identifier 

and describer to be a "lumper" or a "splitter." 

Fortunately, for this study of the Paleocene plants 

of the Rocky Mountains and Great Plains, I had at 

hand for comparison in the U.S. National ~fuse11m 

collections most of the type specimens described by 

previous writers. To these, as a result of many field 

trips during which I became intimately acquainted 

with the pertinent fossiliferous strata, I was able to 

add a large quantity of supplementary material. 

Unfortunately, I have not had an opportunity to 

see some types now located at other museums but 

have had to depend on the published descriptions and 

illustrations. Particularly is this true of all foreign 

Paleocene collections, more especially those described 

by Heer from Greenland, W atelet from the Paris 

Basin, France, and Saporta and Marion from Gelinden, 

Belgium. This foreign material is illustrated chiefly 

by drawings, for the most part beautifully done, but 

how faithful to the specimens I do not know. For 

this reason I have refrained from adopting the names 
36 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

of some of those types for American specimens even 

though the latter may seem, superficially at least, indistinguishable from them. A considerable number of 

the foreign types, as identified and illustrated, seem 

not to be represented by similar specimens in the 

American Paleocene flora. 

In any case, identifications will depend largely 

upon the identifier's knowledge of the morphology, 

classification, nomenclature, geographic distribution, 

and ecology of living plants, and his familiarity with 

paleobotanical literature. At the heart of all this will 

be his native botanical capacity. and his philosophy 

about species, living and fossil. 

On theoretic grounds I have difficulty in accepting 

the proposition that species, considered to be groups 

of closely related individuals having in common a 

multiplicity of features, are discrete, natural entities. 

However discrete such species may seem to be now, 

their discreteness vanishes when they are viewed in 

the perspective of their ancestral history. If the 

parade of organic life-every individual plant and animal that has ever lived-were passed in review from 

the beginning to the present before a competent biologist, could he, except arbitrarily, classify the colossal 

array into species, genera, and families? I doubt it 

and I deny that any of his arbitrary species would in 

actuality be a natural entity. If only one individual 

of Ginkgo biloba Linnaeus or one individual of the 

passenger pigeon, Ectopistes migrator-ius (Linnaeus) 

Swainson, remained, those specimens would simply be 

the last of many millions that preceded them in an uninterrupted line yesterday and ad infinitum into the 

limbo of the past. In the fossil record of Ginkgo 

many "species" .appear, but it is now manifestly impossible to circumscribe those "species" exactly and 

to decide upon the precise genetic moment when one 

evolved into a new "species." As I conceive it, the 

organic world from the beginning to now has been like 

a continuous tapestry, a fabric woven without observable natural breaks and obvious seams. The only 

separations are the purely arbitrary ones that have 

been invented and applied for. practical, talking purposes. 

Repeatedly, therefore, in the following systematics 

I point out that the species I recognize are not necessarily natural but are in many instances dubious and 

are chiefly form or practical species. For example, 

some such aggregates are: Ginkgo a.diantoides (Unger) Heer, Oercidiphyll!um arctioum (Heer) Brown, 

Ampelopsis acerifolia (Newberry) Brown, Platanus 

raynoldsi Newberry, and Viburnum antiquwm (Newberry) Hollick. These species, as l treat them, include a large variety of leaf forms. Within this variety, however, in each species there is a mode or norm 

that I visualize as the average mature leaf of that 

species. My synonymies, consequently, indicate a 

tendency to "lump" rather than to "split," and my 

species names are merely labels for identifying recognizable, usable combinations of form and organization. I practice this method because, after long experience with the publications and collections of 

"splitters," I conclude that upon the latter rests the 

greater burden of proof that they have erected valid, 

usable species. I find it much easier to justify a norm, 

with its variants, as typical of a species or a geographic race than to define many similar though 

slightly different leaves (or other parts of plants) as 

separate species because they have somewhat different 

shapes and some divergence, more or less, in the number of marginal teeth. Over-refined species in an 

abundance of material seem difficult to define, comprehend, and use, even by their describer. On the 

other hand, a common but sometimes deceptive practice in paleontological work seems to be: the fewer 

specimens a student has the more species he is likely 

to make! 

In many instances named species are considered dubiously identified, but question marks are not used. 

This practice may be condemned on the ground that 

all such species should be assigned to Phyllites, Oarpites, or similar catchall form genera. Nevertheless, 

it seems to me that a usable distinction in the degree 

of doubt can be made. For example, a leaf may have 

nearly all the features of Quercus, and Quercus could 

be an item of the flora, but the student may still have 

reservations because no acorns or other remains of 

Quercus have been found in the same strata as the 

leaf. Why should he relegate this leaf to Phyllites? 

If, however, the leaf has no features definitely suggestive of Q?MR(JUS or any other genus known to him, 

he should not assign it to Quercus even with a question 

mark. I hold that binomials, especially when used 

as center headings in publications and in systematic 

lists, should be kept as free as possible of question 

marks and parenthetical or other material that properly belongs in the discussion of the species. 

The descriptions of species sometimes leave much to 

be desired. Some students think they have done justice to a specimen when they have described it minutely 

to the last millimeter, marginal tooth, and surface 

hair. Although for some specimens this may be appropriate, I consider that it is more important to 

be told concisely by what particular feature the species differs from those with which it might be confused. The detection and clear description of specific 

distinguishing features is the nub and hardest part of 
METHODS APPLIED IN THE STUDY OF THE PLANTS 37 

paleobotanical work, but it emphasizes the fact that 

publication is intended for the benefit of others rather 

than as an outlet for the author. 

Casting about for help in identifying his material, 

the paleobotanist consults type collections, botanical 

manuals, herbarium sheets, seed collections, and experts. In my studies I have been privileged with the 

opportunity of using the U.S. National Herbarium and 

the U.S. Department of Agriculture seed collection, to 

whose courteous and helpful staffs I am greatly indebted. Despite all this help I have been disappointed many times, and my failures to find satisfactory comparative living material together with the possibility that many fossil specimens may represent extinct taxa, have caused me regretfully to relegate 

numerous items to the catchall genera Oarpites, Phyllites, etc., and to leave many unnamed. 

Another source of frustration is the not infrequent 

wrong citation of the locality of the type specimens 

in the older paleobotanical publications. Nevertheless, 

specimens of different ages and localities were thus 

mixed, some so much so that they have not been and 

perhaps never can be_ correctly placed geographically 

and stratigraphically. 

Still another annoyance derives from the fact that 

the drawings in many paleobotanical publications are 

of doubtful merit and are often misleading in that 

they do not show the outlines, margins, and venationa! details of the specimens accurately. Sometimes 

the specimens were not adequately cleaned and prepared before being handed to the artist, who, although professionally skillful, was most likely not a 

trained botanist and, consequently, could not be expected to recognize some significant details. Thus, for 

example, the figure of Comus newbe1'1'yi Hollick (Newberry, 1898, p. 124, pl. 37, fig. 4) illustrates a specimen as having an entire margin. However, when I 

cleaned the specimen, well-defined marginal teeth 

emerged (pl. 28, fig. 1), necessitating its reallocation. 

In the figures of Las-trea ( Goniopter.fS) polypodioides 

Ettingshausen (Lesquereux, 1878, p. 57, p. 4, figs. 11, 

12), the venational details are altogether misleading. 

Although the specimens are poorly preserved, careful 

inspection with a good lens in just the right light 

shows that they are examples of the same species of 

W oodwardia as illustrated by Lesquereux in his plate 

3, figure 1a. Numerous other examples could be cited, 

if it were necessary to emphasize this point. Practitioners of paleobotany, it would seem; may learn 

several lessons from such mistakes : specimens and 

especially types should be studied first before relying 

on illustrations. If an artist does the drawing or retouching, the figures should be checked minutely 

against the specimens before publication. In lieu of 

the best photographs, the most faithful drawings possible should be made. Even with the best photographic 

technique, satisfactory illustrations cannot be made 

from fossils, and recourse must be had to drawings. 

Because the purpose of illustrations is primarily for 

the reader's benefit and not to display the author's 

photographic skill, I have little patience with those 

paleontologists who stubbornly insist upon no retouching of photography. True, if retouching is necessary, 

it should be done accurately. To say that reasonably 

accurate and satisfactory retouching is impossible is 

sophistry based on the platitude that no one is perfect. 

Should retouching be done with a steady hand and 

keen eye for morphological details, such slight deviation from the truth as may occur is likely to be immaterial as compared with the benefit to the reader 

from having legible and informative illustrations. 

The treatment of the names of some species calls 

for a statement of my policy in this respect. When 

species based on obscure, atypical, and perhaps now 

lost fragments, have been given names that are inappropriate and misleading, and if better specimens 

have been described subsequently under more appropriate names, I have sometimes waived priority in 

favor of the better specimens and names. If the facts 

are stated clearly in each instance, I deny that this 

practice leads to confusion. 

For consistency in typography I have decapitalized 

all specific terms, no matter what their source, and 

have reduced all personal, specific terms ending in ii 

to i, except where Vi is absolutely required grammatically. 

A fossil flora from a given area was preceded there 

yesterday by a flora only slightly, if any, different; 

but could one examine the continuous record backward 

for many millions of years great differences in floral 

composition would be apparent. These differences may 

be inferred to have been caused by the evolution, emigration, immigration, and extinction of species, climatic conditions, and physical avenues or barriers. 

Consequently, in the study of a collection I have begun by comparing its composition first with that of 

the flora living in the area. Such comparisons, however, often prove futile, and one must look for clues 

in exotic floras, perhaps in distant parts of the world. 

The composition of the distant flora or floras may include species that give clues to other unknowns in the 

fossil flora under examination. By inference, all the 

satisfactory identifications together permit the formulation of a reasonably probable concept about the -
38 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

ecologic and climatic conditions that prevailed at the 

time the fossil flora lived. 

Some species may seem to be restricted to the fossil 

flora and its correlatives, thus serving as indices to 

relative age. These "index species," however, must be 

accepted with reservations for they or their tentatively 

indistinguishable ancestors or descendants may subsequently be found in earlier or later strata. Further, 

the absence of a species or species from a given flora 

in which they apparently should be present must likewise be accepted with caution. In brief, to determine 

the relative geologic age of a flora, not only the "index species" but the whole flora must be evaluated, 

particularly in the light of its immediately surrounding relatives, predecessors, and successors. 

The terms lower, middle, and upper following the 

formation name under each species description refer 

to general approximations within the Paleocene series 

and do not indicate definite stratigraphic limits within 

the Paleocene. 

THE PALEOCENE FLORA 

Distribution of the species is indicated by initials 

following their names, as Almy, A Y; Animas, A; 

Canada, CA; Coalmont, CT; Denver and Dawson, D; 

Evanston, E; Ferris, F; Fort Union, FU; Livingston, 

L; Middle Park, MP; Puerco, P; Raton, R. See page 

11 for names and distribution of formations and members. The Paleocene flora, for practical reasons, is here 

treated as a composite unit instead of by florules taken 

from the separately identified but fairly closely equivalent formations. 

SEEDLESS PLANTS 

Ohara spp ___________________________________________ FU 

Fungi -----------------------------------------------FU 

Marchantia lignitica (Ward) Brown, n. comb ____________ FU 

pealei Knowlton --~-------------------------------FU Preissites wardi Knowlton _____________________________ FU 

Hypnum coloradense Brown, n. sp _______________________ D 

Mnium montanense Brown, n. sp _______________________ FU 

Hymenophyllum contusum Lesquereux ___________________ D 

Allantodiopsis erosa (Lessuereux) Knowlton 

and Maxon ________________________ A, CT, D, FU, MP, R 

Blechnum anceps (Lessuereux) Brown, n. comb _____ D, FU, R 

Dennstaedtia americana Knowlton _______________ CA, D, FU 

Dryopteris lakesi ( Lesquereux) Knowlton __________ D, FU, R 

meeteetseana Brown, n. sp _________________________ FU 

serrata Brown, n. sp ____________________________ D, FU 

Last rea goldiana ( Lesquereux) 

Lesquereux ________________________ A, CT, D, FU, MP, R 

Onoclea hesperia Brown, n. name ___________________ F, FU 

Saccoloma gardneri (Lesquereux) Knowlton ______ AY, D, FU 

Woodwardia arctica (Beer) Brown, n. comb ____ D, F, FU, MP 

Gleicher/Jia hesperia Brown, n. sp ______ .:... ________________ FU 

Anemia elongata (Newberry) Knowlton ___ A, D, FU, L, MP, R 

Lygodi'ltm coloradense Knowlton __________________ CT, D, FU 

Osmunda greenlandica· (Heer) Brown, n. comb ________ D, FU 

mac1·ophylla Penhallow ________________________ CA, FU 

Eqttisetum spp _________________________________ D, FU, MP 

Isoetites horrid1.ts (Dawson) Brown _____________ D, F, FU, R 

Selaginella berthoudi Lesquereux _________________________ D 

collieri Knowlton _________________________________ FU 

monstrosa (Hollick) Brown, n. comb ________________ FU 

SEED PLANTS 

Gymnosperms 

Zmnia colo1·adensis (Knowlton) Brown, n. comb ______ FU, MP 

~vyomingensis Brown, n. sp _________________________ FU 

Ginkgo adi(wtoides (Unger) Heer ____________________ FU, L 

Amentotaxus camp belli (Gardner) Florin ________________ FU 

Araucnria longifolia (Lesquereux) Brown, n. comb ________ FU 

Glyptostrobus nordenskioldi (Beer) Brown, 

n. comb __________________________ CT, D, E, F, FU, L, MP 

Metasequoia occidentalis (Newberry) Chaney ____ CT, E, F, FU 

Taxodium olriki (Heer) Brown, n. comb ________ , _______ FU, L 

Fokienia catenulata (Bell) Brown, n. comb __________ CA, FU 

Thuja inte1·rupta Newberry ________________________ D, F, FU 

Monocotyledons 

SparganiJum antiquum (Newberry) Berry ____________ CT, FU 

Alismaphyllites grandifolius ( Penhallow) Brown, 

n. comb _________________________________________ FU, R 

Sagittaria megasperma Brown, n. sp ____________________ FU 

Hydromystr~a expansa (Beer) Hantke _________ CT, D, F, FU 

Grasslike monocotyledons 

Chamaedorea danae (Lesquereux) Berry __________________ R 

Paloreodoxites plicatus (Lesquereux) Knowlton ________ D, R 

Sabal grayana Lesquereux ______________________ D, F, FU, R 

imperialis Dawson _____________________________ D, FU 

powelli Newberry ______________________________ FU, R 

Thrinax dorji Brown, n. sp _____________________________ FU 

Canna? magnifolia Knowlton ___________________________ FU 

Zingiberites dubius Lesquereux ___________________________ D 

Dicotyledons 

Salix aquilina Brown, n. sp ____________________________ FU 

Carya antiquorum Newberry ______ A, CT, D, E, F, FU, MP, R 

Juglandicarya spp ____________________________________ FU 

Juglans berryana (Knowlton) Brown, n. comb __________ A, R 

taurina Brown, n. sp ______________________________ FU 

Pte1·ocarya glabra Brown, n. sp _________________________ FU 

hispida Brown, n. sp __________________________ CT, FU 

BePula stevensoni Lesquereux __________________ CT, E, FU, L 

Coryl1ts i.nsignis Beer _______________________________ FU, L 

Castanea intermedia Lesquereux _________ A, D, F, FU, MP, R 

Quercus asymmetrica Trelease __________________________ FU 

greenlandica Beer __________________________________ A 

macneili Brown, n. sp ______________________________ FU 

sullyi Newberry ______________________________ CT, FU 

yulensis Brown, n. sp ______________________________ FU 

Celtis newberryi Knowlton and Cockerell_ _______________ FU 

peracuminata Brown, n. sp _________________________ FU 

Planera microphylla Newberry _______________________ D, FU 

Ulm'lts rhamnifolia Ward _______________________________ FU 

Zelkova planeroides (Ward) Brown, n. comb _____________ FU 

Artoca1·p'lts lessigiana (Lesquereux) Knowlton __ A, D, FU, P, R 

Ji'imts affinis (Lesquereux) Brown ____________ A, D, E, FU, R 

artocarpoides Lesquereux ____________ E, F, FU, L, MP, R 

1n·imttidens Knowlton _____________________________ A, R 

planicostata Lesquereux..:. ___________ A, D, E, F, FU, P, R 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 39 

Ficu,s affinis (Lesquereux) Brown-Continued 

sttbtTuncata Lesquereux _______________________ D, F, FU 

ttncata Lesquereux _________________________________ R 

M m·us montanens·is Brown, n. sp ________________________ FU 

Platanus nobilis Newberry ______________ CT, E, F, FU, MP, R 

Taynoldsi Newberry __________ CT, D, F, FU, L, MP, P, R 

Credneria? daturaefolia Ward _______________________ .:_ __ FU 

Cinnarnornmn sezannense Watelet_ ___________________ D, FU 

Lattrophyll1t,rn cauda tum (Knowlton) Brown, n. comb ____________________________________ E, F,. FU, R 

pe1·seanurn Brown, n. sp __________________ CT, E, FU, R 

Lam·tts social·is Lesquereux ______________ A, CT, D, E, FU, R 

Lindem obtusata (Ward) Brown, n. comb __________ A, F, FU 

Persea. brossiana Lesquereux __ ~ _________ A, CT, FU, L, MP, R 

SassafTas therrnale ( Lesquereux) Brown, 

n. comb _______________________________ CT, E, F, FU, MP 

Cabornba inerrnis (Newberry) Hollick ___________________ FU 

N elurnbiurn rnontanmn Brown, n. sp _____________________ FU 

tenuifolittrn Lesquereux ___________________________ A, D 

Nyrnphaea leei (Knowlton) Brown, n. comb _______________ R 

pulchella (Knowlton) Brown, n. comb ________________ D 

Paleonelurnbo rnacroloba Knowlton ______________________ FU 

Paleomtpha1· hesperiurn Brown, n. sp ____________________ FU 

Paranyrnphaea crassifolia (NeWberry) Berry ____________ FU 

Cercidiphyllnm arcti.c'u.m (Heer) 

Brown ________________________ CT, D, E, F, FU, L, MP, R 

Magnolia berryi (Knowlton) Brown, n. comb ____________ A, R 

borealis Brown, n. name _________________________ FU, L 

rnagn1:folia Knowlton _______________________ A, D, FU, R 

regalis Heer _____________________________________ D, R 

rotttndifolia Newberry ______________________________ R 

Harnarnelites inaequalis (Newberry) Brown, n. comb ______ FU 

Liquidambar dakotense Bro-v.rn, n. sp ____________________ FU 

E1wornrnia sen·ata (Newberry) Brown, 

n. comb ____________________________ CA, CT, D, F, FU, R 

Hyd1·angea antica Brown, n. sp _____________________ FU, MP 

Asi1nina vesperalis Brown, n. sp ________________________ FU 

Ce1·cocarpus ravensm·agensis Berry----------------------CA 

Prttnus care-yhttrstia Brown, n. sp _______________________ FU 

colm·adensis Knowlton ____________________________ D, R 

corrugis Brown, n. sp _________________ AY, A, CT, F, FU 

rnclearni Berry -----------------------------------CA 

perita Brown, n. sp ________________________________ FU 

Bauhinia wyorningana Brown ___________________________ FU 

Legttrnim,osites coloradensis Knowlton ________________ FU, MP 

wilUanLSi Berry ___________________________________ CA 

M·irnosites coloradensis Knowlton ________________________ CT 

Robinia wa1·di (Knowlton) Brown, n. comb __________ CT, FU 

Staphylea n11inutidens (Knowlton) Brown, n. comb _________ R 

Acm· tra,qile Knowlton __________________________________ R' 

newberryi Brown, n. name __________________________ FU 

silberlingi Brown, n. sp ____________________________ FU 

spp. _______________________ _: _____________________ FU 

Sapindu.s Affin:is (Newberry) Brown ___________________ D, FU 

Koelreuteria annosa Brown ____________________________ FU 

!lex artocarpid·ioides (Lesquereux) Brown, n. comb ________ D 

Palittnts? sp. Berry------------------------------------CA 

Rhamnus clebttrni Lesquereux ____________________ D, FU, R 

goldiana Lesquereux ___________________________ A, D, R 

hi1·suta Brown, n. sp _______________________________ FU 

Zizyphus fibrillostt.s ( Lesquereux) Lesquereux ____ A, D, FU, R 

Z1:zyphoides mackayi BelL _______________________________ CA 

Arnpelopsis aceTifolia (Newberry) Brown, 

n. comb ____________________________ CT, D, E, FU, L, MP 

Cisstts nw1·ginata ( Lesquereux) Brown, 

n. comb ___________________________ A, CT, D, F, FU, P, R 

C·issites rocklandens·is Brown, n. sp _______________________ R 

Pa1·thenocissu.c; ttrsina Brown, n. sp ______________________ FU 

Vitis lobata (Knowlton) Brown, n. comb ______________ A, FU 

olriki Heer ___________________________ D, E, FU, MP, R 

sp ______________________________________________ FU 

Dornbeyopsis rna.r;nifica Knowlton _________________________ D 

Pte1·ospennites cordattts Ward _____________________ D, FU, R 

DWenites {j(W/ieldensis Brown, n. sp ____________________ FU 

My1·tophyllttm torreyi (Lesquereux) Dorf_ _________ A, D, FU 

Trapa angulata (Newberry) Brown, n. comb ________ ..:._CA, FU 

Trapa pattlula (Bell) Brown, n. comb ________________ CA, FU 

M elastornites rnontanensis Brown, n. sp ________________ F, FU 

Cornus hyperborea Heer ___________________________ FU, MP 

nebrascensis Schimper _________________________ FU, MP 

Nyssa alata (Ward) Brown, n. comb ____________ D, F, FU, R 

borealis Brown, n. sp ____________________________ F, FU 

Nyssa? obovata Knowlton _______________________________ D 

Kalmia elliptica Brown, n. sp ________________________ D, FU 

Fraxin1.ts eocenica Lesquereux _____________ CT, D, E, FU, MP 

Apocynophyllurn lesqttereuxi Ettingshausen ________________ R 

Vib'urmtm antiquttm (Newberry) Hollick ______________ F, FU 

asperum Newberry ________________________________ FU 

cupanioides (Newberry) Brown, n. comb _________ CA, FU 

tilioides Ward __________________________________ A, FU 

Macclintockia kanei (Heer) Seward and Conway ___ Greenland 

Phyllites dernoresi Brown, n. sp ________________________ FU 

disturbans Brown, n. sp ___________________________ ...FU 

pagosensis Knowlton __________________________ A, D, R 

Calycites hea;aphJJlla Lesquereux ______________________ E, FU 

polysepala Newberry ______________________________ FU 

Carpolithes spinosus Newberry ___________________________ R 

N ordenskioldia borealis Heer -----------------------CT, FU 

Palrnocarpon cornrnttne Lesquereux _____________________ D, R 

cornpositurn Lesquereux ____________ .;.. ________________ R 

lineaiurn Lesquereux. _______________________________ D 

subcylindricum Lesquereux _____________________ D, FU 

truncatttrn Lesquereux ______________________________ D 

Viburnum goldianurn Lesquereux _________________________ D 

solitai'ittrn Lesquereux ______________________________ D 

Ophiornorpha nodosa Lundgren __________________________ FU 

Fo,ssil wood and miscellaneous objects of uncertain affinity. 

SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC 

FLORA 

SEEDLESS PLANTS 

ALGAE 

Chara spp. 

Plate 5, figures 1-6 

Characeous oogonia, probably representing several 

distinct species, have been found in the Paleocene strata 

of the Rocky Mountains and Great Plains. One collection, made by Donald Duncan, of the U.S. Geological 

Survey, contains spherical oogonia (pl. 5, figs. 2, 3) 

averaging 1 mm in diameter. These resemble in size 

and number of sheathing cells or spiral ridges those 

(pl. 5, fig. 1) of Ohara convpressa l{nowlton (1888, p. 

156, 157), now called Acli8tochara con1pressa (l{nowlton) Peck and Reker (1948, p. 87, pl. 21, figs. 31-33), 
40 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

from the Flagstaff limestone near Wales, Utah, and the 

Willow Creek formation in Canada (Bell, 1949, p. 36, 

pl. 22, figs 1, 2, 3, 5, 7, 8). They ~lso resemble those 

described by W atelet ( 1866, p. 50-56) from the Tertiary of the Paris Basin. 

Somewhat similar specimens (pl. 5, figs. 4-6) were 

collected from about the same stratigraphic level in 

the Fort Union formation in North Dakota by W. E. 

Benson, of the U.S. Geological Survey; and by R. B. 

Colton, also of the Survey, from a slightly lower 

stratigraphic level in NE:tt4 sec. 31, T. 33 N., R. 55 E., 

Sheridan County, Mont. These smaller specimens appear to me to represent a different species than those 

in the Duncan collection and are perhaps close to 

A.clistochara mitella Peck and Reker ( 1948, p. 89, pl. 

21, figs. 16-20) from the Evanston (Hoback) formation in Hoback Canyon, Wyo. Another species, A.. cor-

()'lULta Peck and Reker also from the Evanston formation, is intermediate in size between A. compressa and 

A. mitella. I have seen none such in the Fort Union 

formation. 

Occurrence: Fort Union formation (lower) , 9206; 

(upper), 229 (fig. 1), 9204 (figs. 2, 3), 9205 (figs. 4-6). 

FUNGI 

Round or irregular spots occur on many species of 

Paleocene leaves. ·Oswald Heer described the following from the Paleocene strata of Greenland : Depooea 

gronlamdica, Rhytisma boreale, S clerotiwm dmmJJII'Mmi, 

S. populicola, S,plweria amtnulifera, S. arctica, and S. 

imterpwngens. Those called Sclerotiwm rubelltwm Lesquereux (1874, p. 35, pl. 1,. fig. 2) occur on the leaves 

of a palm; and those called S clerotites ~ cypericola 

Knowlton ( 1930, p. 16, pl. 1, fig. 7) and S. ~ hesperius 

Knowlton (idem, pl. 1, fig. 6) on sedgelike leaves. 

Doubtless some of these spots were caused by fungi or 

bacteria, but others may have been caused by insects, 

worms, or mechanical wear and tear. I have made no 

e1fort to study the spots microscopically by peel or 

other methods and have found no satisfactory criteria 

for djstinguishing them megascopically. Consequently 

I offer no identifications of them. On the contrary I 

consider it best that such objects be not named unless 

characterizable spores are found. Notice of them may 

well be confined to the description of the foliage on 

which they occur. 

What purports to be the context of a bracket-fungus 

was described by Heer as Polyporites sequoiae (1874, 

p. 48, pl. 83, fig. 10). This dubious object also might 

better have remained unnamed. 

Spores of fungi occur in the Paleocene coals, as 

found by Wilson and Webster (1946, p. 271, fig. 1) 

who described Brachyspo~m sp. from near Red 

Lodge, Mont. 

HEPATICAE 

Marchantia lignitica (Ward) Brown, n. comb. 

Plate 4, figures 4, 6 

Fucus li,qnitum Lesquereux. Ward, 1886, p. 549, pl. 31, fig. 2; 

1887, p. 13, pl. 1, fig. 2 [not fig. 1, which is Trapa angulata (Newberry) Brown]. 

Ward's illustration (his fig. 2) of this species (fig. 4) 

does not show a feature present on the specimen when 

cleaned up, namely, the distinct areolation, similar 

to that on the upper surface of the thallus of a living 

liverwort of the family Marchantiaceae. Reexamination of the specimens illustrated in Ward's figures 1 

and 2 shows that they do not represent the same spe,. 

cies of plant. Specimen 1 was collected from Upper 

Cretaceous strata at Point of Rocks, Wyo., and is the 

dissected, submerged foliage of Trapa~ angulata (Newberry) Brown, whose typical floating rosettes are also 

found at the same locality. Specimen 2 is from Fort 

Union strata at Burns Ranch, on the left bank of the 

Yellowstone River, 30 miles northeast of Glendive, 

Mont. As these are fresh-water deposits, the specimen 

cannot be a species of Fucus, which is a genus of 

marine algae. The size of the thallus, surface markings, and general habit resemble those of the living 

M archantia polymorpha Linnaeus. The areolations of 

the related Oonoceplwl;us are considerably larger. 

Occurrence: Fort Union formation (middle), 2420 

(fig. 4, 6). 

Marchantia pealei Knowlton 

Plate 4, figures 1, 10 

Marchantia pealei Knowlton, 1908, p. 157-159, pl. 25. 

M archantites pealei (Knowlton) Steere, 1946, p. 305. 

This species resembles M aroha!ntia lignitica (Ward) 

Brown but apparently, lacks the areolations of the 

latter. Hollick (1930, p. 37, pl. 1, fig. 2) described 

M. yukonensis from Cretaceous strata in Alaska. It 

also lacks distinguishing surface features and may be 

related to M. pealei. The transfer to M archantites was 

a futile gesture. 

Occurrence: Fort Union formation (lower), 4293 

(fig. 10, a reproduction of Knowlton's type), 8517 

(fig. 1), 8258; (upper), 8202. 

Preissites wardi Knowlton 

Plate 4, figure 7 

Preissites wardii Knowlton, 1894, p. 458-460, pl. 219, figs. 1-3. 

Jfm·cha,ntites wardii (Knowlton) Steere, 1946, p. 305. 

The thalli of this species were represented as having 

"lines" simulating secondary veins branching from the 

"midrib." These "lines" are folds or wrinkles in the 

troughs of which some carbonaceous matter is preserved giving the appearance of veinlets. As no fruit-
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 41 

ing bodies have been found on these specimens, there 

remains some doubt as to the affinities imp1ied in the 

generic name. The transfer to M archatntites by Steere, 

in my opinion, was needless. 

Occurrence: Fort. Union formation (lower), 8252; 

(middle), 2420 (fig. 7); (lower), 8252; (upper), 9125. 

MUSCI 

Hypn um coloradense Brown, n. sp. 

Plate 5, figure 7 

This is a leafy sterile shoot, of probable procumbent 

habit. The leaves are narrow ovate-lanceolate in outline, sharp-pointed, with costae. Sporophyte lacking. 

Occurrence: Dawson arkose, 5738 (fig. 7). 

Mnium montanense Brown, n. sp. 

Plate 5, figure 12 

Leaves ovate, acute, entire, in two rows. Costa 

fairly strong ending in the apex. In the absence of 

spore capsules and other distinguishing features, it 

seems futile to compare this species with any living 

species of M nium. Resemblances to species of Fusidens 

are suggested. 

Occurrence: Fort Union formation (lower), 8519 

(fig. 12). 

FERNS 

HYMENOPHYLLACEAE 

Hymenophyllum confusum Lesquereux 

Plate 6, figure 9 

Rymenophyllum confusum Lesquereux, 1878, p. 51, pl. 2, figs. 

6-6a. 

Knowlton, 1930, p. 17, pl. 1, fig. 4. 

Knowlton's figure of this species is perhaps a little 

clearer than Lesquereux's, but all the specimens are 

poorly preserved, so that the species is difficult to 

characterize. 

Occurrence: Denver formation, 317 (fig. 9). 

POLYPODIACEAE 

Allantodiopsis erosa (Lesquereux) Knowlton and Maxon 

Plate 5, figures 13-16 

Allantodiopsis et·osa (Lesquereux) Knowlton and Maxon. 

Knowlton, 1919, p. 61. 

Knowlton, 1924, p. 78, pl. 6, fig. 1 ; 1930, p. 24, pis. 4, 5. 

See synonymy. 

Bell, 1957, p. 21, pl. 3, figs. 3, · 6, 9. 

Asplenium remotidens Knowlton, 1899, p. 669, pl. 80, fig. 7. 

OsmJunda af]inis . ( Lesquereux) Lesquereux, 1878, p. 60, pl. 4, 

fig. 1. 

Pteris pseudopenna·etormis Lesquereux. Hollick, 1936, p. 39, 

pl. 75, fig. lb. 

Saccoloma gat·dneri (Lesquereux) Knowlton. Knowlton, 1930, 

p. 26, pl. 3, fig. 4. [not other figs.] 

Not much can be added to the description of this 

species except the record of new localities. Apparently 

only one fruiting specimen-that illustrated by Knowlton (1930, pl. 4, fig. 1)-has ever been found in Paleocene strata. 

The features that distinguish ·this species are : pinnules large, coarse, serrate with cuneate bases more or 

less petioled; secondary veins almost at right angles 

to the midrib, widely spaced, not anastomosing, branching once near the relatively slender midrib and sometimes again near the margin, with occasional unbranched intermediaries; vein terminations in the margin not conspicuously swollen; sori large, elliptic, 

astride the distal branch of a secondary vein near the 

midpoint between midrib and margin. 

This or a very closely related species occurs in Upper 

Cretaceous strata in the Rocky Mountain region; and 

possibly the same or another species occurs in Eocene 

strata near Mitchell, Clarno, and Comstock, Oreg.; on 

Skookumchuck River, Swauk Creek, Fletts Creek (near 

Wilkeson), and other localities in Washington. 

Occurrence: Fort Union formation (lower), 7663, 

8551, 8884; (upper) , 877 4 ; Animas formation, 7 496, 

8447; Coalmont formation, 6105 (fig. 14); Denver formation, 317 (fig. 15), 318 (fig. 13), 325 (fig. 16, 

Knowlton's fruiting type reproduced), 8777; Middle 

Park formation, 326; Raton formation, 5826. 

Blechnum anceps (Lesquereux) Brown, n. comb. 

Plate 6, figures 3, 4 

Salpichlaena anceps (Lesquereux) Knowlton, 1930, p. 27, pl. 7, 

figs. 1-3. [Omit all but last ite~ of synonymy, for they 

are Allantodiopsis erosa (Lesquereux) Knowlton and 

Maxon.] 

Pteris linearis Knowlton, 1917, p. 284, pl. 54, fig. 3. 

From such fragments of this fern as are .preserved, 

it would appear that the frond was branched and 

relatively open, like the living OsrnA.Ifflila regalu, with 

the pinnules widely spaced and approximately at right 

angles to the rachis. The pinnules have entire or subdenticulate margins, and rounded, cordate, or auriculate bases. The midvein is slender, and the secondary 

veins, although simulating those of Allamtodiopsis 

erosa, are more closely spaced, the distal portions being directed almost at right angles to the midvein, 

and the vein terminations in the margin are conspicuously enlarged or club shaped. No fertile remains of this species have been recognized. Blechtntum 

is now the accepted name for these ferns. 

Occurrence: Fort Union formation (lower), 8666, 

8678 (figs. 3, 4), 8928; Denver formation, 322, 6142; 

Raton formation, 5099. 
42 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Dennstaedtia americana Knowlton 

Plate 6, figures 1, 2, 5-7 

Dennstaedtia americana Knowlton, 1910, p. 492, pl. 63, fig. 4; 

pl. 64, figs. 3-5. 

Berry, 1935, p. 16. 

Dennstaedtia crossiana Knowlton, 1930, p. 21, pl. 2, figs. 7-9. 

Dennstaedtia blomstrandi ( Heer) Hollick. Bell, 1949, p. 39, 

pl. 24, figs. 1, 4 ; pl. 26, fig. 5. 

Knowlton's types from North Dakota include both 

fertile and sterile fronds, and similar material, as 

illustrated, from other localities confirms the characterization of the species. The venation of the fertile 

pinnules consists of veins to the sori only, whereas 

that of the sterile pinnules is considerably more diffuse and forked. 

The relationship of this species to the sterile foliage 

from the Tertiary of Greenland and Alaska called 

De'TIInStaedtia blom.gtratndi (Heer) Hollick, and to that 

from the Upper Cretaceous of western Alberta, called 

Sphenopteris (Dennstaedtia?) burlirngi Bell, is problematical. Many of those specimens are fragmentary, 

and their characterization is difficult. This, or a close 

ancestral species occurs in the Cretaceous of northern 

Alaska. 

Occurrence: Fort Union formation (upper), 4273 

(fig. 1, Knowlton's type reproduced), 4334, 5029 (fig. 

7), 6084, 8222 (figs. 2, 5), 8231 (fig. 6), 8913; Dawson 

arkose, 325; Paskapoo and Ravenscrag formations, 

Canada. 

Dryopteris lakesi (Lesquereux) Knowlton 

Pia te 8, figure 15 

Dryopteris lakesii (Lesquereux)· Knowlton, 1930, p. 18, pl. 1, 

figs. 1, 2. 

Asplenium? primero Knowlton, 1917, p. 285, pl. 54,:ftg. 4. 

Knowlton's figure 2, from the Denver formation, 

shows a fruiting specimen, the numerous sori being 

small, round, and evenly spaced on both sides of the 

ultimate veins. l(nowlton's figure 1, the specimen of 

which I cannot now locate, appears to be misleading 

in showing excessively forked venation. In most specimens the ultimate veinlets are simple, and enter the 

apexes of the lobes. 

Occurrence: Fort Union formation (lower), 7538, 

8899; Denver formation, 317 (fig. 15); Raton formation, 5798. 

Dryopteris meeteetseana Brown, n. sp. 

Plate 5, figure 17 

Frond at least twice divided, anadromic, that is, with 

the first pinnule closest to the main rachis on the 

upper side of the pinna axis. Proximal pinnules free 

or almost free, but distal pinnules more or less united. 

Pinnules with rounded lobes each of which receives a 

vein. Astride the veins are round sori, but no spores 

are preserved. 

Occurrence: Fort Union formation, 4694 (fig. 17). 

Dryopteris serrata Brown, n. sp. 

Pia te 8, figure 17 

Frond at least once divided; lowermost pinnae 

petioled, parted to the midvein into rounded obtuse 

divisions; uppermost pinnae becoming more and more 

sessile and less divided. Margins serrate. Lowest 

secondaries of the almost free pinnules thrice forked, 

the next, twice forked, and the rest once forked. Similarly for the undivided pinnae. 

No comparison with a living species is sugested, 

and the reference to Dryopteris is entirely nominal. 

The species may be the same as H emitelites torelli 

Heer ( 1871, p. 462, pl. 49, figs. 1-5a; pl. 55, fig. 2) 

from Atanekerdluk, Greenland. 

Occurence: Fort Union formation (lower) 85H11 

8678 (fig. 17); Dawson arkose, 5836. 

Lastrea goldiana (Lesquereux) Lesquereux 

Plate 5, figures 9-11, 18 

Aspidium goldianJum Lesquereux, 187 4, p. 293. 

Last rea ( Goniopteris) goldiana ( Lesquereux) Lesquereux, 1878, 

p. 56, pl. 4, fig. 13. 

Lastrea ( Goniopteris) fischeri Heer. Lesquereux, 1883, p. 239, 

pl. 50, figs. 1, 1a. 

Newberry, 1898, p. 10, pl. 48, fig. 6. 

Lastrea (Goniopteris) knightiwna Newberry, 1883, p. 503. 

Asplenium·? penhallowi Bell, 1949, p. 41, pl. 26, figs. 1, 3, 4. 

Dryopteris? cladophleboides Knowlton, 1917, p. 284, pl. 54, 

fig. 1; 1924, p. 78, pl. 6, fig. 2. 

Dryopteris integra Knowlton, 1930, p. 17, pl. 1, fig. 5. 

Dryopteris lesquereua:ii Knowlton, 1922a, p. 109. 

Dryopteris richardsoniana Knowlton, 1930, p. 20, pl. 2, figs. 3-5. 

The type specimen of this species, originally called 

Aspidium goldianwm, is now poorly preserved but may 

have been in better condition when first studied and 

figured. In 1922 Knowlton (p. 109) recorded this 

specimen with the Laramie flora of the Denver Basin, 

but in 1930 (p. 20) spoke of it as being from the 

Denver formation although he did not list it as a species of the Denver (Tertiary) flora. Examination of 

the matrix, however, confirms the latter assignment. 

The few to many equally spaced gracefully forwardcurving, rarely forked, secondary veins of the entiremargined slightly falcate pinnules, cha.racterize the 

specimens assigned to this species. Fertile pinnae are 

scarce (figs. 9, 11; and l(nowlton's Dryopteris integra), but when present have small round sori on the 

secondary veins midway between the midrib and margin. 

The relationship of this species to D1·yopteris laramiensis Knowlton ( 1922a, p. 109, pl. 1, fig. 5) from 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 43 

the Laramie formation in the Denver Basin; to Heer's 

Paleocene specimens from Greenland called Aspidium 

esch.eri, A. heeri, and A. meyeri; to Saporta's Alsophila thelypteroides ( 1868, p. 324, pl. 3, fig. 1) ; to 

Knowlton's Eocene specimens from Yellowstone Park, 

called Aspleniu1n iddingsi, Dryopteris weedi, D. W(J;'t1-

tholithensi8; to Berry's Eocene species, Goniopteris lesquereuwi, from the 'Vind River Basin in Wyoming; to 

Hollick's Tertiary species, Asplenium alaska;nru;m and 

Dryopteris meyeri, from Alaska; to Heer's Lastrea 

fisch.eri from Miocene ]akebeds at Oeningen, Germany; 

and to specimens from many Eocene localities in 

Washington and Oregon, is difficult to evaluate, for 

in many instances the figured specimens involved are 

poorly preserved and provide only tantalizing resemblances. The Eocene specimens, however, appear 

to be somewhat larger, with more secondary veins 

in each pinnule. 

Occurrence: Fort Union formation (lower)) 4661, 

8258 (fig. 18); Animas formation, 7484; Coalmont 

formation, 6105 (fig. 11); Dawson arkose, 325 (figs. 

9, 10), 5738; Denver formation, 317; Middle Park 

formation, 333; Raton formation, 5803. 

Onoclea hesperia Brown, n. name 

Plate 7, figures 1, 4 

Onodea sensibilis fossilis Newberry, 1868, p. 39; 1898, p. 8, 

pl. 23, fig. 3. 

Knowlton, 1902, p. 705, pl. 26, figs. 1-4. 

Berry, 1935, p. 16, pl. 1, figs. 2, 3; pl. 2. [From Berry's 

synonymy delete the items W oodwardia Zatiloba and 

W. latiloba minor, which are W. arctica (Heer) Brown.] 

Onoclea hebridica (Forbes) Ettingshausen and Gardner. Bell, 

1949, p. 40, pl. 20, fig. 5 ; pl. 24, figs. 3, 5 ; pl. 25, fig. 2. 

This species is represented by characteristic sterile 

foliage at many Paleocene localities, but fertile fronds 

have been reported from only one locality, along 

Sweetgrass Creek, west of Porcupine Butte, on the east 

side of the Crazy 1\tiountains, Mont. (Knowton, 1902, 

p. 705). Similar fertile fronds, originally called Oaulimites fecwnda, were described by Lesquereux from 

the Laramie formation at Erie, Colo. 

The sterile fronds most likely to be confused with 

Onoclea are those of Woodrwardia. However, if the 

margins are well enough preserved, those of Onoclea 

will be entire, whereas those of W oodwardia will be 

minutely serrulate or serrate. If the venation is clearly 

preserved, the veins of Onoclea make a more uniform 

pattern of anastomosis, even close to the margins, than 

those of W oodwardia, which, after one or two anastomoses, tend to be forked t9 the margins. 

Thjs species is distinguished from the Cretaceous 

species, Onoclea fecwnda Knowlton, in having more 

elongated, pointed, marginal lobes. It may be equivalent to 0. h.ebridica (Forbes) Gardner and Ettingshansen (1879, p. 68) from the Eocene Ardtun beds, 

Isle of Mull. 

A new name is hereby proposed for this American 

species, because the original trinomial is misleading, 

unnecessary, and inappropriate. First, there is no assurance that this species is identical with the living 0. 

sensibilis Linnaeus, however closely they may resemble 

each other; nor that it is a variety or subspecies of 0. 

sensibilis, as indicated by the third term fossili8. 

Moreover, every fossil species of Onoclea deserves the 

adjective fossilis. Finally, trinomials are unnecessary 

in paleobotany, for they suggest a degree of refinement 

in identification of fossil plant remains that is neither 

achieved nor even possible of achievement. 

Occurrence: Fort Union formation (lower), 4029, 

5842, 7004 (fig. 4), 8897, exact loc. not stated (fig. 1, 

Newberry's fig. 3 reproduced); (upper), 607, 5322, 

8212, 8917, 8921, 8922, 9072;. Ferris formation, 6419. 

Saccoloma gardneri (Lesquereux) Knowlton 

Plate 5, figure 19; plate 6, figures 12, 13 

Sa,ccoloma gardneri ( Lesquereux) Knowlton, 1919, p. 560; 

1930, p. 26, pl. 3, figs. 3, 5-8 [not fig. 4, which is Allantodiopsis erosa (Lesquereux) Knowlton and Maxon]. 

Bell, 1957, p. 20, pl. 1, figs. 6, 8. 

Phanerophlebites pea~ei Knowlton, 1922a, p. 110, pl. 3, fig. 5. 

Pteris goldma'flli Knowlton, 1922a, p. 111, pl. 2, fig. 3. 

Saccoloma sp. Knowlton, 1930, p. 27, pl. 3, figs. 1, 2. 

The distinguishing features of this species are : 

pinnules sometimes longer and larger than those of 

Allantodiopsi8 erosa and Blechnum aJJUJeps,- margins 

entire or slightly undulating; bases rounded or cuneate; midrib stout; secondary venation somewhat resembling that of Alla!ntodiopsis erosa, except for frequent anastomoses; in sterile pinnules the veins terminate in the margin without conspicuous thickening, 

but in fertile pinnules the veins terminate in round 

to squarish sori that unite to form a marginal border. 

Occurrence: Fort Union formation (lower), 4029 

(pl. 6, fig. 13)' 6845, 7538, 8678 (pl. 6, fig. 12)' 8897 

(pl. 5, fig. 19) ; (upper) 5030; Almy formation, 3667; 

Dawson arkose, 325, 5738) 5831, 6142. 

Woodwardia arctica (Heer) Brown, n. comb. 

Plate 7, figures 2, 3, 5 

Woodwardites arcticus Heer, 1868, p. 86, pl. 1, figs. 16, 16b; 

pl. 45, fig. 2c ; pl. 48, fig. 9 ; 1871, p. 462, pl. 40, fig. 6. 

Woodwat·dia latiloba Lesquereux, 1874, p. 391; 1878, p. 54, pl. 3, 

figs. 1, 1a. . 

Woodwardia latiloba mimor Lesquereux, 187 4, p. 391 ; 1878, 

p. 54, pl. 4, figs. 9, 9a. 

Woodwardia latiloba serrata Knowlton, 1930, p. 22, pl. 2, figs. 

1, 2. 

Woodwardia maxoni Knowlton, 1910, p. 489, pl. 63, fig. 3; 

pl. 64, figs. 1, 2. 

Berry, 1926, p. 95, pl. 10, figs. 2, 3. 

Dryopteris polypodioides (Ettingshausen) Knowlton, 1930, p. 20. 
44 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Last rea ( Goniopteris) polypodioides ( Ettingshausen) Lesquereux, 1878, p. 57, pl. 4, figs. 11, 12. 

Onoclea sensibilis Linnaeus. Hollick, 1936, p. 35, pl. 2, figs. 2-4. 

Onoclea sensibiUs [ arctica] Heer, 1883, p. 48, pl. 70, fig. 6. 

Pteris pseudopmvnaetormis Lesquereux, 1878, p. 52, pl. 4, figs. 

3, 4. 

Sterile leaves of Onoclea and Wood/wardia may resemble one another closely, a circumstance that renders 

fragmentary fossil material of these genera difficult to 

distinguish. In general, however, the leaf-margins of 

Onoclea are entire, whereas those of · W oodwardia, 

except for some fertile fronds, are either minutely 

serrulate or conspicuously serrate. Thus, Hollick's figures of Onoclea sensibili8 from the Tertiary of Alaska 

give the impression of having entire margins and 

therefore of being assignable to Onoclea. His specimens, however, particularly that of his figure 4, show 

clearly serrate margins, and therefore, they are transferred to W oodwardia. In Lesquereux's figures of 

Pteris pseudopemnae formis the venation is incorrectly 

illustrated. Practically, the misidentification of one 

for the other of these genera would probably make 

little or no difference for ecological or climatic interpretations. 

The relationship of this species to W oodwardia erenata Knowlton ( 1900, p. 22, pl. 3, fig. 3) and other 

Cretaceous and post-Paleocene species is conjectural. 

Occurrence: Fort Union formation (lower), 4661, 

6116, 7005, 8519 (figs. 3, 5), 8852, 8666, 8897; (upper), 

5030 (fig. 2), 6420, 8652, 8913; Denver formation, 318, 

844 7; Ferris formation, 6420; Middle Park formation, 

333. 

GLEICHENIACEAE 

Gleichenia hesperia Brown, n. sp. 

Plate 8, figures 11, 12, 16 

Pinnules long, relatively narrow, squarely sessile on 

the rachis at right angles or nearly so, with crenate 

or rounded marginal lobes pointing slightly distad. 

The notches between the lobes do not reach the rachis, 

as in most fossil species of Gleiehenia. The venation 

of each marginal lobe consists of one somewhat undulant vein and its two or three branches. No sori 

have been found. 

These specimens are only nominally assigned to 

Gleiehenia. Somewhat similar material from the Cretaceous of Greenland was called Phegopteris jorgen.seni 

Heer (1883, p. 32, pl. 35, figs. 1-3), and later Sphent!pteris jorgenseni ( Heer) Seward. Sa porta and Marion 

(1878, p. 16, pl. 1, figs. 2-3), de'scribed a comparable 

fragment as B enitzia minima from the Paleocene of 

Gelinden, Belgium. 

Occurrence: Fort Union formation (lower), 8519 

(figs. 12, 16), 8897 (fig. 11). 

SCHIZAEACEAE 

Anemia elongata (Newberry) Knowlton 

Plate 6, figure 8 

Sphenopteris (Aspleni7tm) elongatum Newberry, 1863, p. 511. 

A111emia elongata (Newberry) Knowlton, 1922a, p. 112, pl. 2, 

fig. 2. [In the synonymy, under Anemia perplexa Hollick, 

omit pl. 63, figs. 1-4.] 

_1neimia eocenica Berry, 1916a, p. 164, pl. 9, fig. 7 ; pl. 10, fig. 2 ; 

pl. 11, figs. 1, 2; 1930, pl. 47, fig. 1. 

Anemia grandifolia Knowlton, 1924, p. 78, pl. 5. 

A:nmnia lnnceolata Knowlton, 1930, p. 29, pl. 8, fig. 10. 

Ancmin mosbyensis Knowlton, 1930, p. 28, pl. 8, fig. 9. 

A.·nemia occidentalis Knowlton, 1917, p. 285, pl. 54, fig. 2. 

.. Anemia supercretacea Hollick. Knowlton, 1917, p. 248, pl. 30, 

fig. 5. 

Anemia sp. Knowlton, 1916, p. 87, pl. 15, figs. 6, 7. 

Anemia sp. Knowlton, 1916, p. 333, pl. 84, fig. 4. 

Anemia sp. Knowlton, 1922a, p. 113, pl. 2, fig. 1. 

A.ne1nia sp. Knowlton, 1930, p. 30, pl. 8, fig. 7. 

A.neimia sp. Dorf, p. 44, pl. 1, fig. 8. 

Aspleni1.tm eoligniticum Berry, 1961a, p. 167, pl. 11, fig. 3; 

1930c, p. 62, pl. 8, figs. 2-4. [Not .1. eoligniticnm described by Brown, 1934, p. 52, pl. 8, figs. 3, 4. This is 

probably A. hurleyensis Berry.] 

Diplazium crossii (Knowlton) Knowlton, 1930, p. 23, fig. 6. 

[See synonymy.] 

Lygodium? compactum Lesquereux. Knowlton, 1922a, p. 113, 

pl. 1, fig. 1. 

Pteris pennaeformis Heer. Newberry, 1898, p. 7, pl. 48, fig. 5. 

Pteris pseudope111naetormi8 Lesquereux. Knowlton, 1902, p. 22. 

[Lesquereux's original specimens are Woodwardia. arctica (Heer) .Brown.] 

Berry, 1916, p. 168, pl. 9, fig. 6. 

Pteris russellii Newberry, 1883, p. 503; 1898, p. 7, pl. 61, figs. 

1, la. 

Anemia is sometimes spelled A neimia. Originally 

Swartz spelled it Anemia, but in a footnote gave the 

derivation as from the Greek a;neimon, unclad, naked. 

The correct ultimate transliteration, therefore, would 

be Animia. 

The reference of these Cretaceous and Tertiary ferns 

to Anemia seems to be justified, following the discovery by Andrews and Pearsall (1941, p. 168) of what 

purport to be the fertile parts of Anemia fremonti 

Knowlton from the Frontier formation (Cretaceous) 

in western Wyoming. No such fructifications have 

yet been found or reported from later formations containing sterile fronds. Collectors, however, should 

be on the alert :for such remains. 

The synonymy given here is the writer's opinion only 

and rests on no definite characters that ally all these 

specimens as one species. That a number of species 

may here be mixed is quite probable inasmuch as the 

range of specimens extends from uppermost Cretaceous 

in to Eocene. 

It will be noted that in general only fragments, 

usually the tips of fronds, are figured. Knowlton's 

Anemia grandijolia, from the Animas formation, is 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 45 

the largest figured specimen, and inspection of it reveals the great variation in degree of dissection of 

the pinnules. It is for this reason that the long .synonymy includes species with pinnules that are practically entire, and others that are dissected almost 

if. 11-ot entirely, to the rachis. ' 

The relationship of Anemia elongata to A. fremonti 

Knowlton of the Colorado group and to A. delicatula 

Brown of the Green River and related formations is ' obscure. Gradations from one to the other seem plausible but hardly demonstrable. 

Occurrence: Fort Union formation (lower), 2421; 

4028; 7538, 8928; (upper) 1502, 2416, 4369, 6084 (fig. 

8), 8678, 8775, 9532; Animas formation, 7481; Dawson arkose, 5738, 5836, 8188; Livingston formation, 

8896; Middle Park formation, 333, 335, 337; Raton 

formation, 5697. 

Lygodium coloradense Knowlton 

Pia te 8, figures 9, 10 

Lygodium coloradense Knowlton, 1930, p. 30, pl. 8, fig. 8. 

This species was based upon a single specimen whose 

base was deeply scalloped and venation widely spaced. 

Material collected subsequently fro"m other Paleocene 

localities indicates that not all the specimens have 

bases like that of the type but that some may be 

cuneate. The relatively widely spaced and generally 

less forked venation, however, seems to be a persistent 

and distinguishing feature. The American Eoce~e 

species, Lygodium neuropteroides Lesquereux [ L. kaulfussi Heer of authors], has more closely spaced venation, giving the pinnules a less coarse appearance. 

Fertile pinnules (fig. 9) of Lygodium coloradense 

occur in the. Coalmont formation of North Park, Colo., 

but these do not appear to be significantly different 

from those of Eocene species. No authentic remains 

of Lygodium have been reported from Laramie, Lance, 

and Hell Creek strata, the specimens from those fo~­

mations alleged to belong to Lygodium being fragments assignable to other genera. The earliest known 

American climbing fern is L. pwmil!wm Brown ( 1943b, 

p. 141, figs 1-5), with delicate, small pinnules, from 

the Montana group near Casper, Wyo. 

Occurrence: Fort Union formation (lower), 8678 

(fig. 10); Coalmont formation, 6105 (fig. 9); Dawson 

arkose, 5738 (type) . 

OSMUNDACEAE 

Osmunda greenlandica (Heer) Brown, n. comb. 

Plate 5, figure 8; plate 6, figures 14, 15 

Pteris gronlamdica Heer, 1883, p. 49, pl. 70, figs. 1-5; pl. 107, 

fig. 1. 

593121 0 - 62 - 3 

Pteris oeningensis Unger. Heer, 1880, p. 9, pl. 3, figs. 2, 3, 4, 8b. 

Cladophlebis groenlandica ( Heer) Bell, 1949, p. 38, pl. 26, fig. 2. 

Pinnules 2 em long, somewhat broad falcate in outline, entire or only slightly crenulate, pointed, and 

separated from one another by narrow, rounded sinuses 

reaching to within 1 or 2 mm of the rachis. Secondary 

veins openly spaced, once forked near the midvein ' and rarely forked again near the margin. No fertile 

fronds assignable to this species have been found. It 

can be compared with Osmunda claytoniana Linnaeus, 

of the Eastern United States. 

The specimens described by Heer as Pteris gronlatndica from Paleocene strata at Atanekerdluk, Greenland, have only slightly smaller pinnules than those 

of the Montana specimens here figured as the same 

species. The relationship of OsmJIJJ'I1.da greenlandica to 

species described by Hollick (1936) from the Tertiary 

of Alaska as Pteris inaequilateralis and 0 smwnda dubiosa, is problematical. 

Occurrence: Fort Union formation (lower), 8519 

(figs. 14, 15), 9253; (upper), 8774, 9238; Denver formation, 317 (fig. 8). 

Osmunda macrophylla Penhallow 

Plate 6, figures 10, 11; plate 8, figures 7, 8, 13, 14 

Osmunda macrophylla Penhallow, 1908, p. 65, text fig. 15. 

Bell, 1949, p. 38, pl. 24, fig. 2 ; pl. 25, figs. 1, 3, 4. 

Osnwnda heerii Gaudin. Heer, 1868, p. 88, pl. 1, figs. 6-11; 

pl. 8, fig. 15b. 

Pinnules oblong, broad at the somewhat auriculate 

base, sessile, rounded at the apex. Margin crenulate. 

Secondary veins twice or thrice forked, undulant. No 

fertile fronds assignable to this species have been 

·found. The species may be compared favorably with 

the royal fern, Osmunda regalis Linnaeus, of widespread occurrence. 

This species apparently lived into Eocene time and 

perhaps later. It may be the same as Osmunda eocenica Saporta and Marion (1873, p. 30, pl. 1, fig. 2; 

1878, p. 18, pl. 1, fig. 1) from Paleocene strata at 

Gelinden, Belgium. 

Occurrence: Fort Union formation (lower), 8519 

(figs. 10, 11); (middle and upper), 8220, 8248 (fig. 

13); 9084 (fig. 8); 9085 (figs. 7, 14). 

EQUISETACEAE 

Equisetum spp. 

Plate 8, figures 1-6 

Remains of Equisetum in the Fort Union and related 

Paleocene formations include portions of sterile aerial 

shoots with sheaths, underground stems with tubers, 

and cross sections of both aerial and subterranean 
46 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

stems. One object that may be the fruiting cone of an 

E quisetum has been found (pl. 67, fig. 43) . 

The following species have been described: Equisetum alexoensis Bell ( 1949, p. 36, pl. 22, fig. 4; pl. 23, 

figs. 5, 7, 9: pl. 67, fig. 5) ; E. arcticum Heer (Bell, 

1957, p. 37, pl. 23, figs. 2, 4, 8) ; E. boreale Heer (1868, 

p. 89, pl. 1, fig. 17; pl. 45, figs. 10, 13e, f) ; E. coloraden8e J{nowlton (1930, p. 31, pl. 8, figs. 1-6); and 

E. globulosum Lesquereux ( 1883, p. 222, pl. 48, fig. 3). 

When, however, these species are examined in the light 

of the material now on hand, they do . not seem to 

be readily distinguishable. Size of stem, number of 

teeth in the sheaths, shape of the tubers-all vary so 

greatly, with gradations from one to another, and 

with no apparent constancy in any set of features, that 

I shall not attempt to define the species. 

Occurence: Fort Union formation (lower), 7538 (fig. 

3), 8678; (middle), 2414 (figs. 2, 5), 2417 (fig. 1), 

4474; (upper), 8164 (fig. 6), 8204 (fig. 4), 8774; 

Dawson arkose, 5838; l\1iddle Park formation, 333. 

ISOETACEAE 

lsoetites horridus (Dawson) Brown 

Plate 9, figures 1-8 

I soetites horrid1ts (Dawson) Brown, 1939c, p. 268, figs. 3, 6. 

[See synonymy, and correct Xantholithes to Xantholithus.] 

This species was characterized as having leaves 

(sporophylls) whose spatulate terminations had entire margins. A few specimens have since been found 

that have minute obscure teeth which are not nearly 

as conspicuous as those of I soetites serratu8 Brown 

from Upper Cretaceous strata. 

Additional collections also now permit illustration 

of the complete leaf, with its undulant margins and 

the ligule (figs. 1, 2) above the basal part containing the sporangium filled with spores. Some of this 

ne'v material is from Eocene strata. Figure 4 shows 

the rosette of leaves around a center that is a mass 

of megaspores. 

The genus Isoetites was already in existence in the 

Triassic as shown by a species· called /. circula.ris 

(Emmons) Brown (1958b, p. 358, figs. 5, 8, 9, 11, 13), 

represented by specimens from the Pekin formation 

in the Deep River coal field, North Carolina, and the 

Chinle formation in the Petrified Forest National 

Monument, Ariz. 

Occurrence: Fort Union formation (lower), 2432 

(fig. 8), 5437, 8517; (middle), 2420 (figs. 4-6), 8535 

(fig. 3); Dawson arkose, 8881 (fig. 2); Ferris formation, 5255 (fig. 7) ; Hanna formation (Eocene), 8548 

(fig. 1); Raton formation, 5144. 

SELAGINELLACEAE 

Selaginella berthoudi Lesquereux 

Pia te 4, figure 9 

Selaghl-ella berthoudi Lesquereux, 1878, p. 46, pl. 5, figs. 12, 12a. 

Knowlton, 1930, p. 33. 

This species has larger, more ovate leaves, and a 

more spreading branching habit than Selagilnella collieri Knowlton. 

Occurrence: Dawson arkose, 8779 (fig. 9); Denver 

formation, 317. 

Selaginella collieri Knowlton 

Plate 4, figures 2, 3, 8 

Selaginella collieri Knowlton, 1916, p. 201, pl. 1, figs. 1-6. 

There is nothing new to add to l{nowlton's description of this species except to cite localities where it 

has since been found. This species may be related to 

S. berthoudi Lesquereux from the Denver formation, 

but differs in having smaller, more acute leaves, and 

apparently in the habit of branching. 

Occurrence: Fort Union formation (lower), 6215, 

7004 (figs. 3, 8) ; (middle) 2420 (fig. 2) . 

Selaginella monstrosa (Hollick) Brown, n. comb. 

Plate 4, figures 12, 13 

Anomalofilicites monstrosus Hollick, 1916, p. 474, pl. 31, figs. 1-3. 

Hollick described this species from sterile fronds 

as a sport or monstrosity which he likened to some 

illustrated specimens of N ephrolepis ewaltata (Linnaeus) Schott. Apparently the feature of these specimens that caused him to think of them as monsters 

is the fact that the side branches ·differ greatly in 

length. · 

In general appearance the sterile shoots are not unlike some mosses, but unfortunately: the ultimate details are not clearly preserved and :under magnifications of more than 5 or 6 the leaf-1 details fade into 

the background of the matrix. However, the leaves 

appear to be sharp pointed. 

'Vith these sterile shoots are here associated fertile 

shoots that come from the same., strata in Garfield 

County, Mont., but not :from the. :same locality. The 

belief that the sterile and ferti~~ shoots belong together is based upon the resemblance in general habit, 

both kinds of shoots having branches differing greatly 

in length. 

The short ultimate branches carry sporangia bearing numerous megaspores (10-15), but no details· of 

the megaspores can be :seen clearly. Scattered along 

the main rachis are a few small ovate, hairy ( ~ ) , 

pointed leaves. 

Comparison of this species with any living species 

of S ela.ginella is not suggested. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 47 

Occurrence: Fort Union formation (lower), 4031 

(fig. 13), 8249 (fig. 12) .. 

SEED PLANTS 

GYMNOSPERMS 

·CYCl\DACEAE_ 

Zarnia coloradensis (Knowlton)' Brown, n. comb. 

Plate 10, figm·es 6, 9, '12, 13 

Chamaedorea? colo1·adensis Knowlton, 1930, p. 40, pl. 10, fig. 3. 

Podozamites latipennis Heer. Brown, 1939b, p. 246, pl. 49, 

figs. 1-4. 

Since my first report on material of this species, I 

have found additional specimens at other localities 

and have concluded that the fossils do not represent 

Podozamites la.tipennis l:Ieer, described from Cretaceous strata in Greenland, but are more likely a species of Zamia. 

The leaflets are linear, somewhat curved in outline, 

pointed, entire and are attached by either a broad or 

narrow base, depending ·upon their position on the 

main rachis, which is broad and stout. There is no 

midvein. The thin parallel veins arising from the 

base of the leaflet fork n~ar the base and anastomose 

several times thereafter on their course through the 

leaflet. The palms of the genus Ohamaedorea have 

neither forked nor anastomosing veins. 

fleer described a fragment from strata at Puilasok 

on Disko Island, that he called Poacites nielseni 

(1874, p. 19, pl. 4, fig. l) in honor of Jens Nielsen, 

a Dane who lived at Atanekerdluk and collected the 

first fossil plants in that area. Of him Heer said 

simply "Er verungliickte auf der J agel und liegt in 

Atanekerdluk begraben." (He met misfortune while 

hunting and lies buried at Atanekerdluk). This fragment is from a blunt-pointed leaf with parallel veins 

that, according to the illustration, fork and anastomose 

-a feature not found in grasses. I suspect, therefore, that this specimen represents a cycad, but cannot 

identify it as definitely belonging to Zamia coloradensis. 

I have no reason to abandon my hypothesis that the 

objects (figs. 12, 13) originally figured as the fruits 

of this species are properly allocated, for similar 

fruits have been found associated with the foliage of 

this species at three additional localities in the basal 

strata of the Paleocene series in Wyoming. 

Occurrence: Fort 1Jni0n formation (lower) 8551 

(figs. 6, 9, 12, 13), 8884, 8893, 9130; (upper) 9195; 

l\iiddle Park formation, 333 (type). 

Zarnia wyorningensis Brown, n. sp. 

Plate 10, figure 1 

Rachis stout, pinnules abruptly contracted near the 

base, but attached to the rachis by a broad insertion. 

Size of pinnules unknown. Veins widely spaced, forking near the insertion, and again, in son1e instances, 

farther along toward the apex. Anastomoses not present in the specimen. 

This species differs from Zamia coloradensis 

(Knowlton) Brown in apparent lack of anastomosing veins and in having broad-based insertions of the 

pinnules. It resembles 0 eratozamia wrig hti Hollick 

( 1936, p. 42, pl. 12, fig. 6) in general appearance, but 

in 0. wrighti the veins do not fork at the base of 

the pinnule, and the pinnules may have scattered, 

marginal teeth distad. 

Occurrence: Fort Union formation (upper) 5911 

(fig. 1). 

GINKGOACEAE 

Ginkgo adiantoides (Unger) Heer 

Plate 10, figures 2-5, 8 

Ginkgo adiantoides (Unger) Heer. Ward, 1887, p. 15, pl. 1, 

figs. 5, 6. 

Berry, 1926, p. 190. 

Ginkgoites adiantoides (Unger) Seward. Bell, 1949, p. 43, 

pl. 32, figs. 1, 3. 

In previous papers (Brown, 1939a, p. 246; 1943b, 

p. 862) discussing Ginkgo laran1iensis Ward, I expressed my view's on the distinction of fossil species 

of Ginkgo and concluded that the fossil record of 

Ginkgo indicates an evolution in leaf form from the 

deeply lobed, digitate cuneate in early l\1esozoic strata, 

through entire or bilobecl cuneate in late Mesozoic, 

to entire or bilobecl reniform leaves in the Cenozoic. 

As is well known, the only living species, Ginkgo biloba Linnaeus, may produce all these leaf-forms on a 

single tree, or given trees may specialize in any one 

of the three kinds. Ii1 short, at any stage in the 

history of GinJe:go that stage may produce leaf forms 

occurring in its ancestors .but will have few hints concerning the shapes of leaves to come, and thus, a good 

collection of Ginkgo leaves frmn a given geological 

level may provide a useful clue to stratigraphic dating. 

As the first well-developed reniform leaves of Ginkgo appear in the Paleocene, I favor restriction of G. 

adiantoides to the Cenozoic. This species, however, 

like G. bilobn, may include leaves having the form 

of the Cretaceous G. l(l.rcwnien~is and the .Jurassic G. 

digitata. When, therefore, these kinds only are found 

in a Paleocene or later collection, they can, nevertheless, be identified as G. adiarntoides, on the presumption that they are such variants. Certainly less harm, 

if any, is clone by this practice than, for example, to 

identify simple, undivided cuneate Ginkgo leaves 

wherever found, whether in Mesozoic or Cenozoic 

strata, or even on the living tree, as G. lara;miensis! 
48 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Except apparently during Eocene time, species of 

Ginkgo graced the landscape of the Northwestern 

States from the Jurassic to the late Miocene, but G. 

adiantoides was present in Alaska and probably British Columbia during the Eocene. Hence its apparent 

absence from the States during that epoch invites 

speculation about the possible reasons for this absence. Is the fossil record incomplete or was Ginkgo 

actually missing for climatic or other causes? 

During the Paleocene G. adiantoides lived in the 

Dakotas, Montana, Wyoming, and perhaps Colorado. 

At a locality on the south side of Dry Creek, 5 miles 

west of Greybull, Wyo., the leaves are thickly matted 

and mixed with those of many other species of trees. 

Oval bodies (figs. 3, 4) chiefly internal molds of nuts 

or pits, are found among these le~ves and presumably 

are the seeds of Ginkgo. Similar seeds associated with 

Gimkgo leaves have been found on Sevenmile Creek . ' east of Glendive, Mont., and at other localities including Alberta, Canada, from which Bell re~orts one as Carpolithus (Gitnkgoitesn sel!wyni (1949, p. 

45, pl. 66, fig. 1). These objects may on the other 

hand, be the stone fruits of other plants, such as 

Prunus, etc. The specimen repDrted from the Denver 

:formation at Golden, Colo., as Ginkgo? tuncata (Lesquereux) Knowlton (1930, p. 34, pl. 9, fig. 3) is very 

suggestive of Ginkgo, but no leaves of Ginkgo have 

been taken from that formation. 

Occurrence: Fort Union formation (lower), 4661, 

5526,5721,8256 (fig. 2), 9196,9208, 9334; (middle and 

upper), 2416 (figs. 4, 5), 6219, 8522, 8786 (fig. 3), 

9109 (fig. 8), 9125; Livingston formation, 6765, 6767. 

PINACEAE 

No foliage, seeds, or wood, attributable to the genus 

PinUB, have been found in the Paleocene strata of 

the Rocky Mountains and Great Plains. Oswald Heer, 

however, reported needles, purporting to be those of 

Pitnus, from the Paleocene strata at Atanekerdluk and 

vicinity, Greenland: Pinus hayesiarna Heer (1883, p, 

66, pl. 71, figs. 13a, 14, 14b), P. hyperborea Heer 

(1868, p. 94, pl. 17, fig. 5f; 1871, p. 465, pl. 44, fig. 5; 

pl. 56, fig. · 9c; 187 4, p. 16, pl. 2, fig. 12). As undoubted remains of Pim.U8 have been found in the 

Upper Cretaceous and Eocene of the Rocky Mountains 

and Plains region, the apparent absence of the genus 

from the Paleocene is, like the similar absence of 

SalllJilnia, cause for speculation. 

CEPHALOTAXACEAE 

Amentotaxus campbelli (Gardner) Florin 

Plate 14, figures 7, 8 

Podocarpus campbelli Gardner, 1886, p. 97, pl. 26. 

Amentotaxus campbelli (Gardner) Florin, 1940, p. 163, text 

fig. 1. 

Bell, 1957, p. 30, pl. 13, figs. 3, 4. 

A.mentotaxus florini Krause!, 1935, p. 137, text figs. 1, 2, 5-8. 

Cephalotaxus californioa Potbury, 1935, p. 61, pl. 1, fig. 2. 

Needles elliptic-lanceolate in outline, 3 to 6 em long, 

4 to 10 mm wide, with sharp-pointed apexes and with 

bases narrowed to a short, stubby petiole. Stomata in 

two broad bands. 

These needles resemble closely those of the living 

A'I'Mntota::cus argo,taenia (Hance) Pilger, from southeastern China. Comparisons, however, may also· be 

made with those from some of the larger leaved species of Oephalotamus. As only needles are represented, 

the foregoing synonymy may have to be changed 

when and if seeds, wood, and other material are found. 

Occurrence: Fort Union :formation (lower) 4696 

(fig. 8); (upper) 9109 (fig. 7'). 

ARAUCARIACEAE 

Araucaria longifolia (Lesquereux) Brown, n. comb. 

Plate 13, figures 1-7 

Sequoia longifolia Lesquereux, 1878, p. 79, pl. 7, figs. 14, 14a; 

pl. 61, figs. 28, 29. 

Sequoia acuminata Lesquereux, 1878, p. 80, pl. 7, figs. 15, 16, 

16a. 

Dorf, .1938, p. 45, pl. 2, fig. 1. 

Sequoiites artus Bell, 1949, p. 47, pl. 4, figs. 7, 8; pl. 16, fig. 2. 

Abietites tyrellii Dawson, 1887, p. 17. 

AraJucaria hatcheri Wieland, 1910, p. 80, fig. 2. 

Araucarites longifolia (Lesquereux) Dorf, ·1942, p. 130, pl. 4, 

fig. 9. 

Cunninghamites? sp. Knowlton, 1900, p. 29, pl. 5, fig. 3. 

Cunninghamites eZegans (Corda) Endlicher. Knowlton, 1905, 

p. 135, pl. 15, fig. 1. 

Cunninghamites recurvatus Hosius and von der Marek. Knowlton, 1905, p. 136, pl. 16, fig. 6. 

Torreya borealis Heer, 1883, p. 56, pl. 70, fig. 7a. 

Torreyites tytTellii (Dawson) Bell, 1949, p. 45, pl. 7, figs. 3, 4; 

pl. 8, figs. 1, 4. 

Branches with spreading, recurved lanceolate . ' ' pOinted, apparently veinless, sometimes wrinkled 

needles, 2 em or more long. Stom~tal lines, about 

equally spaced and diffused over the surface of the 

needle. Base of the needle expanded, so that on its 

fall there remains a large obovate, smooth or striated 

scar. Cones 6 to 8 em long, 3 em wide, with many 

scales, apparently without spines but with rhombic, 

outer faces of apophyses. No detached scales or seeds 

were found. 

In the Cretaceous of the Atlantic Coastal Plain and 

the western interior occur foliage, cones, and twigs 

that, at least superfiicially, appear to be related to 

those here described. These remains have been vari~ 

ously referred to Abietites, Arauoaria, Arauamrvte8, 

Oephalotamopsis, Oephalota::cus, Ounnilnqhamites, Dam-
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 49 

m(JJJ'a, Elatocladus, Geinitzia, Podocarpus, Sequoia, 

Sequoiites, Torreya, and perhaps others of which I am 

not aware. Even casual inspection· makes it apparent 

that several genera are represented by these fossils, but 

distinguishing features, such as stomatal lines on the 

needles and cones attached to twigs, are rarely found. 

At least three species and perhaps genera, however, 

can be predicated provisionally: ( 1) that here identified as Araucaria longifolia, having lanceolate fairly 

flexible needles with diffused stomatal lines, and cones 

without apparent spines on the persistent scales; (2) 

that having long, fairly straight, lanceolate, flexible 

needles with two stomatal lines in broad zones near 

the margins of the needles, exemplified by Sequ,oia 

m.agnifolia Knowlton, and probably Oephalotaxus colo~-­

radensis l{now lton ; and ( 3) that having crowded, 

stiff, arcuate needles tapering to a broad base, and 

cones with readily deciduous spine-tipped scales, exemplified by Geinitzia formosa Heer, Sequoia reichenbOXJhi (Geinitz) Heer, and Dammara acicu,lari-8 Knowlton. 

The foliage and cone features of Ararucaria longifolia 

are closely comparable to those in living araucarias, 

and there seems to be little doubt that the species is an 

araucarian. It ranged across the Cretaceous-Paleocene boundary. 

Occurrence: Fort Union formation (lower), 588~ 

(figs. 6, 7), 6738,8519 (figs. 1-5), 8678. 

TAXODIACEAE 

Glyptostrobus nordenskioldi (Heer) Brown, n. comb. 

Plate 11, figures 3, 7-22 

Sequoia nordenskioldi Heer, 1871, p. 36, pl. 2, fig. 13b; pl. 4, 

figs. 1a, 1b, 4-38 ; 187 4, p. 9, pl. 1, fig. 30. 

Newberry, 1898, p. 20, pl. 26, fig. 4. 

Sequoia atfinis Lesquereux. Knowlton, 1930, p. 33. 

Sequoia couttsiae Heer, 1883, p. 63, pl. 68, fig. 6b. 

Sequoia langsdorfii (Brongniart) Heer, 1883, p. 61, pl. 68, figs. 

6b, 6c ; pl. 70, fig. 12 ; pl. 86, fig. 2b. 

Sequoia sternbergi Goeppert. Heer, 1883, p. 63, pl. 70, fig. 13. 

Oryptomerites lambii Bell, 1949, p. 49, pl. 29, figs. 2, 4 ; pl. 30, 

figs. 1, 3, 4, 5; pl. 31, fig. 4; pl. 32, figs. 2, 4. 

Elatocla~us ( Oryptomerites?) nordenskioldi (Beer) Bell, 1949, 

p. 50, pl. 31, figs. 2, 3, 5. 

Glyptostrobus dakotensis Brown, 1936, p. 355, figs. 2-4. 

Glyptostrobus e1tropaeus (Brongniart) Heer. Newberry, 1898, 

p. 24, pl. 26, figs. 6-8a ; pl. 55, figs. 3, 4. 

Hollick, 1936, p. 51, pl. 10, fig. 3b; pl. 17, fig. 1 (part), 

2, 3, 4, 6; pl. 18, figs. 1-5a; pl. 19, figs. 1, 2; pl. 104, 

fig~ 7b ; pl. 109, fig. 11. 

Glyptostrobus ungeri Beer, 1883, p. 61, pl. 70, figs. 9, 10; pl. 85, 

figs. 6-8. 

Taxodium distichum miocenum Heer, 1871, p. 32, pl. 3, figs. 19, 

20, 26 ; pl. 4, fig. 13b. 

Occurring with the three kinds of foliage-cryptomeroid, cupressoid, and taxodioid-commonly seen 

on living Glyptostrobus, are fossil cones, cone scales, 

and seeds, the latter here identified for the first time 

as belonging to Glyptrostrobus. Heer assigned seeds 

of this kind and perhaps some cone scales, associated 

with Glyptostrobus foliage (called Sequoia nordenskioldi) to Tamoditum distichum miocenum from Spitzbergen and to Pterospermites vagruns from Oeningen. 

In 1936 I suggested several reasons for regarding the 

American species as distinct from G. europaeus, which 

is typically developed in the European Miocene. 

The winged seeds are shaped somewhat like a boomerang and, except for size, resemble those of species 

of Sagittaria. The nutlet occupies a large, elliptic 

area from which a short, pointed wing extends at an 

oblique angle. In the only living species, G. pensili8 

Koch, of southeastern China, the wing of the seed is 

only slightly oblique to the nutlet. By contrast, the 

seeds of Tamodium, a close relative, are irregular, unwinged, somewhat triangular objects, in which the 

nutlet is not clearly defined. 

Staminate catkins, not identifiable as those of Metasequoia or Tamodium but associated with Glypto<Strobus 

foliage at Elbowoods, N. Dak., are here assigned to 

this species. 

Remains similar to those here assigned to G. nordenskioldi occur at many Eocene localities in Oregon, 

Washington, and Alaska. The American Miocene species is 0. oregonensi8 Brown. 

Occurrence: Fort Union formation (lower) 4571, 

5063, 5579, 6154, 7005, 8245 (fig. 17), 8249 (fig. 20), 

8549 (fig. 12), 8552 (fig. 3), 8677 (fig·. 14), 8781, 9104, 

9180, 9192 (fig. 9), 9334; (upper), exact loc. unknown (fig. 22, reproduction of Newberry, 1898, pl. 

26, fig. 4), 2416, 4625, 5194, 8193 (figs. 11, 16), 8203 

(fig. 8), 8206 (fig. 21), 8212 (figs. 15, 19), 8224 (figs. 

10, 18), 8521, 8529, 8786 (figs. 7, 13), 888.5, 8917, 8928, 

9125, 9135; Coalmont formation, 6105; Dawson arklose, 5836; Denver formation, 317, 849'3; Evanston 

formation, 8670; Ferris formation, 4661, 4694, 6971; 

Livingston formation, 6765; Middle Park formation, 

337. 

Metasequoia occidentalis (Newberry) Chaney 

Plate 12, figures 1-14 

Taxodium occidentale Newberry, 1863, p. 516; 1868, p. 45; 1898, 

p. 23, pl. 76, figs. 1-3 ; pl. 55, fig. 5 (part). 

Berry, 1935, p. 19. 

Metasequoia occidento,lis (Newberry) Chaney, 1951, p. 225, 

pl. 10, figs. 3-6. [See synonymy.] 

Elatocladus ( Taxodites?) tinajorum ( Heer) Bell, 1949, p. 51, 

pl. 32, fig. 5. 

Juniperus gracilis Beer, 1883, p. 57, pl. 70, figs. 18-20. 

Sequoia brevitolia Heer, 1874, p. 5, pl. 2, figs. 7, 8. 

Sequoia Zangsdorfii (Brongniart) Heer, 1868, p. 91, pl. 2, figs. 

2-22; pl. 47, figs. 15, 15b; 1871, p. 464, pl. 40, fig. 5b; 

pl. 43, figs. 1-3; pl. 46, figs. 1a, 7b; pl. 55, fig. 3a; 1883, 

p. 61, pl. 68, fig. 8. 
50 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Sequoiites langsdorfli (Brongniart) Heer. Bell, 1949, p. 46, 

pl. 29, fig. 5 ; ; pl. 30, fig. 2. 

Tarcites ol1·iki Heer, 1868, p. 95, pl. 1, figs. 24a, b; 1874, p. 16, 

pl. 1, fig. 9. 

Tarcites validus Heer, 1874, p. 13, pl. 1, fig. 11. 

Tarcodium distichum miocenum Heer, 1869, p. 463, pl. 43, fig. 

4c ; 187 4, p. 9, pl. 1, figs. 13c, d, 15b ; p. 19, pl. 4, fig. 5 ; 

1883, p. 60, pl. 70, fig. 11; pl. 87, fig. 7; pl. 88, fig. 2b; 

pl. 96, figs. 8, 9. 

Tarcodium dubium (Sternberg) Heer, 1868, p. 89, pl. 2, figs. 

24-27 ; pl. 12, fig. 1c ; p. 156, pl. 30, figs. 3, 4. 

The verification in 1946 of the occurrence of living 

trees called shuihsa, discovered in 1944 in a small area 

south of the Yangtse River in Szechuan and Hupeh 

provinces, China, and having foliage and cones similar 

to, if not identical with, fossil remains identified as 

Metasequoia, focussed attention on this genus and resulted during the next 5 years in many papers dealing 

with the living and fossil species. The data were 

summarized by Chu and Cooper (1950), who described 

the ecology of the living species, and by Chaney 

(1951), who restudied the fossil species. 

The distinctive feature of the living tree, Metasequoia glyptostroboides Hu and Cheng, also seen in 

the fossils, is oppositeness, the twigs, needles, floral 

organs, and cone scales being opposite one another in 

two ranks. Here, to keep the record straight, it must 

be stated that Metasequoia is not the only conifer with 

oppositeness. AmentotamU8 and OeplwiotamU8 also have 

opposite needles, although these are in general much 

stouter than those of Metasequoia. Nevertheless, confusion of the three genera is possible when dealing 

only with their fossil foliage. 

The needles of Metasequoia are lanceolate to elliptic 

in outline, variable in size, and slightly rounded at 

the base, which is twisted and decurrent diagonally 

on the twig. The cones are pedunculate, relatively 

small, averaging less than 2 em in length, with about 

24 sca~es that, because of their decussate arrangement, 

give the cone a squarish appearance when viewed from 

above. The peduncles may be several centimeters long 

with little or no trace of foliage or foliage scars. The 

seeds are winged, flat, and irregularly oval in outline, 

with wings as wide as the nutlet or wider. They resemble those of the Sierra redwood or Big Tree (Sequoia gitgamlea) closely, and are somewhat larger than 

those of the coast redwood (Sequoia sempervirens). 

It is difficult to distinguish the wood from that of 

8 equoia and T amodifum. 

The ecological studies by Chu and Cooper (1950) 

indicate many interesting associates of the living tree, 

including katsura ( Oercidiphyll!um) and others that, 

according to the fossil record, were companions of 

shuihsa in other parts of the world millions of years 

ago. Indeed, shuihsa and its associates today may 

represent a relict Pliocene flora. In its native habitat 

shuihsa lives at an altitude of about 3,000 feet and 

prefers a slightly acid soil along stream banks or wet 

places at the lower end of ravines and seepage areas 

at the foot of slopes. Climatic conditions there simulate those of the piedmont areas of the Southeastern 

United States. The planting of shuihsa in many places 

outside of China will, in the course of time, reveal the 

most suitable area for the tree and thus supply further data for speculation on the. altitudinal, zonal, 

physiographic, and climatic requirements of its fossil 

antecedents. 

Occurrence: Fort Union formation (lower), 4661, 

5063, 7005, 8240, 8249, 8551 (fig. 9)' 8899, 9193, 9334; 

(upper), 2420 (figs. 13, 14), 4264 (fig. 5), 4582 (fig. 6), 

4984 (fig. 2), 5595, 8165 (fig. 11), 8196, 8212 (fig. 12), 

8238 (fig. 3), 8521 (figs. 1, 8, 10), 8550 (figs. 4, 7) 8556, 

8786, 8885, 9101, 9135; Coalmont formation, 6000; 

Evanston formation, 3563; Ferris formation, 6173, 6176. 

Taxodium olriki (Heer) Brown, n. comb. 

Plate 10, figures 7, 11, 15; plate 11, figures 4-6 

Tarcites olriki Heer, 1868, p. 95, pl. 1, figs. 21-24 c; pl. 45, figs. 

la, 1b, 1c ; 187 4, pl. 15, pl. 1, fig. 10. 

Ela.tocladus ( Tarcodites?) tinajorum ( Heer) Bell, 1949, p. 51, 

pl. 16, fig. 3. 

These alternate, more or less pointed needles, inserted on the twig with their petioles decurrent parallel to the axis of the twig, make it appear that a 

species of Tawodium is represented. No seeds, however, have been found that confirm this suspicion. The 

twigs are somewhat rare, suggesting that the species 

had not yet become a dominant part of the swamp 

forests. 

The description. of Paratamodium wiggiln8i Arnold 

and Lowther (1955) from Upper Cretaceous strata in 

Alaska characterizes what may be a possible ancestral 

link between jJf etase(juoia and Tamodifum. 

Occurrence: Fort Union formation (lower), 4661, 

4984,7548 (pl.11, fig. 5), 8549 (pl.10, fig.15; pl.11, 

fig. 6); (upper), 2420 (pl. 10, figs. 7, 11), 2416 (pl. 

11, fig. 4), 8522, 8899, 9125; Livingston formation, 

6767. 

CUPRESSACEAE 

Fokienia catenulata (Bell) Brown, n. comb. 

Plate 11, figures 1, 2 

Androvettia catewulata Bell, 1949, p. 46, pl. 15, figs. 1-5; pl. 16, 

fig. 4; pl. 27, figs. 5, 7, 8. 

This species, well characterized by Bell, was identified by him as an Androvettia, similar to A. aarolinen8is Berry, A. elegans Berry, and A. statenensis 

Hollick and Jeffrey, from Upper Cretaceous strata of 

the Eastern and .Southeastern United States. Super-
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 51 

ficially, the comparison is apt, but on closer examination significant differences become apparent. Hollick 

and Jeffrey (1909, p. 22), describing A. statenensis, 

remarked about its fernlike appearance and might 

have assigned it to the ferns but for the fact that 

sections of the stems indicate close affinity with the 

conifers. 

The Paleocene specimens differ from the Cretaceous 

examples in being much less fernlike and in having a 

more pronounced jointed or segmented appearance. 

At some localities separate segments rather than twigs 

are the rule. For these reasons and their striking 

likeness to the foliage of Fokienia with at least two 

living species in China, I am inclined to refer them 

to the latter, although admitting the possibility that 

Androvettia may have been in their ancestry. Similar 

segments and twigs have been referred to Libocedr'U8, 

and Watelet (1866, p. 108, pl. 32) ascribed some to 

0 allitrites. 

Occurrence: Fort Union formation (lower) , 7548 

(figs. 1, 2), 8256, 9300; (upper); 9135, 9532; Evanston 

formation 9558; Paskapoo and Willow Creek formation, Alberta, Canada. 

Thuja interrupta Newberry 

Plate 10, figures 10, 14, 16 

Thuja intm·rupta Newberry, 1868, p. 42 ; 1898, p. 25, pl. 26, 

figs. 5-5d. 

Berry, 1935, p. 21, pl. 3, fig. 1. [See synonymy and discussion.] 

Thuites interruptus (Newberry) Bell, 1949, p. 52, pl. 27, figs. 

1-3. 

Thujopsis eur.opaea Saporta. Heer, 1868, p. 1, pl. 50, figs. 

11 a-c. 

Biota borealis Heer, 1874, p. 7, 13, pl. 1, figs. 13-29; 1880, p. 9, 

pl. 3, figs. 5, 5b. 

Libocedrus sabiniana Heer, 1871, p. 34, pl. 2, figs. 6-15; pl. 4, 

fig. 4d; 1883, p. 58, pl. 70, fig. 17; pl. 86, figs. 1, 2; pl. 87, 

fig. 8. 

There is considerable variation among the specimens 

of this species taken from different localities in the 

Paleocene strata of the Rocky Mountains and Great 

Plains. One specimen from the basal Fort Union 

north of Glendive, Mont., has branchlets whose leaves 

appear not as sharp pointed as those of most specimens and resemble those of Cretaceous species of 

M orkonia and Brachyphyll!um. A species, Tlvuja 

colgatensis Brown (1939, p. 247, pl. 48, figs. 2-4) from 

the Upper Cretaceous Colgate member of the Fox 

Hills sandstone, southwest of Glendive, Mont.., is most 

likely in the ancestral line of 1'. interrupta and differs 

chiefly in having longer internodes between leaf pairs, 

but intergrading specimens are likely to be found. 

No cones or seeds attributable to this species have 

yet been recognized in the collections from the Rocky 

Mountains and Great Plains. Heer, however, illustrates a branch and cone from Greenland. There 

may here be a mixture of Thuja, Ohamaecypam, Androvettia, and other genera. Some examples may be 

the cupressoid foliage of Glyptostrobus. 

Occurrence: Fort Union formation (lower), 4028, 

8239, 8258, 8897 (fig. 14), 9194; (upper), 4395, 5030 

(fig. 10), 7989 (fig. 16), 8222, 9180; Dawson arkose, 

8882; Ferris formation, 6420. 

MONOCOTYLEDONS 

SP ARGANI.&CEAE 

Sparganium antiquum (Newberry) Berry 

Plate 14, figures 4, 5 

Brasenia? antiqua Newberry, 1883, p. 514; 1898, p. 93, pl. 68, 

fig. 7. 

Sparganium antiquum (Newberry) Berry, 1924, p. 342-348, 

7 text figs. ; 1930c, p. 64, pl. 8, fig. 5, text fig. 5. 

Sparganium stygium Heer, 1868, p. 97, pl. 45, fig. 2a [probably 

not fig. 13d]; 1871, p. 467, pl. 42, figs. 4b, 5, 5b. 

Ward, 1887, p. 18, pl. 3, figs. 6, 7. 

Berry, 1935, p. 22. 

Platanus basilobata Ward. Bell, 1957, p. 58, pl. 43, fig. 1 only. 

The identity of the specimens of Tertiary age now 

assigned to Spargwnilum has become somewhat mixed. 

Heer described several species, among them S. stygium 

and S. valdense from the middle Tertiary of southern 

Germany, but all may represent a sjngle species. He 

identified somewhat similar material, including separate foliage and fruit, from the Paleocene of Greenland as S. stygiwm to which Ward atld Berry also referred specimens from the Fort Union formation of 

Montana and the Ravenscrag formatjon of Saskatchewan. All the Heer material from Greenland except 

the fruits ( 1871, pl. 42, fig. 5) is fra~mentary and indecisive. There is little question, however, that the 

fruits are identical with those figured by Ward and 

Bell, but that all these should be identified with S. 

stygium of Europe may well be que~tioned. 

The fruits are born in racemes of pedunculate heads 

composed of numerous ovate to lanceolate long-beaked 

nutlets. Often the only part left i~· the small receptacle showing the attachment scars ~f the nutlets. 

The Eocene specimens described oy Berry ( 1930c) 

as Sparganium amtiquum (Newberry) Berry from 

western 'Vyoming have sessile fruit heads with obovate angular unbeaked nutlets, borne on stouter 

branches of a fruiting stalk. It is. perhaps possible 

that Berry's specimens represent only the uppermost 

fruit heads of the fruiting stalk or that they may be 

immature, and that the lower fruits may have been 

peduncled as they sometimes are in living species. 

The peduncled heads of Sparqaniwm antiquwm resemble superficially the heads of Platanus, Liquidambar, EchinodorU8, and others, but both the receptacles 
52 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

and the nutlets have features that seem not to be 

matched even remotely in those living genera. 

This species ranged into the Eocene at numerous 

localities in Wyoming and in Yellowstone National 

Park. 

Occurrence: Fort Union formation (lower) 9197; 

(upper) 2416 (fig. 5) 5594, 8230 (fig. 4), 8556, 9109, 

9125; Coalmont formation, 5987. 

ALISl'riACEAE 

Alismaphyllites grandifolius (Penhallow) Brown, n. comb. 

Pia te 15, figures 1, 4, 6 

Maianthemophyllum grandifoZium Penhallow, 1902, p. 54, text 

fig. 5. 

Bell, 1949, p. 80, pl. 63, fig. 2 ; pl. 64, figs. 1, 3 ; pl. 66, 

figs. 2, 3. 

Alismaphyllites crassifolilum Knowlton, 1917, p. 286, pl. 55, fig. 1. 

Olintonia oblongifolia Penhallow, 1902, p. 55, text fig. 6. 

Ovate leaves with entire margins, probably reaching 

a length of 30 em and width of 10 em. Midvein at 

base of leaf a composite of closely spaced veins giving 

it a stout appearance, but these veins separate in the 

body of the leaf and become 10 or 12 widely spaced, 

parallel veins that converge toward the apex. Closely 

spaced forked veinlets connect the main parallel veins. 

These leaves may be compared with those of some 

species of Alisma, Aponogeton, Echilnodorus, and 

Potamogeton, but as no fruits assignable clearly to 

any of these genera have been found at the same localities as the leaves, there is some uncertainty in the 

assignment. The regular, p~rallel, tertiary venation 

eliminates them from the liliaceous genera 11/ aianthemophyll!wm and Olitntonia. 

This species is apparently closely related to Alisma 

maer'ophyl/;um Heer (1877, p. 66, pl. 26; pl. 27, figs. 

3b, 3c, 4-7) and Potamogeton nordenskioldi Heer 

(1868, p. 157, pl. 30, figs. 1b, 5c, 5d, 6-8; 1871, p. 52, 

pl. 4, figs. 18b, 19; pl. 8, figs. 9, 10; pl. 15, fig. 51h; 

1877, p. 66, pl. 27, figs. 1-3a), but these are considerably younger and may well be different species. 

Occurrence: Fort Union formation (lower) 8567, 

8227 (fig. 6); (upper) 7004, 8206 (figs. 1, 4), 8922; 

Raton formation, 5099. 

Sagittaria megasperma Brown, n. sp. 

Plate 68, figures 14-16 

Achenes 2 em long, asymmetric, winged, with beaks 

4 mm long turned to one side, in dense heads borne 

on long pedicles. Wings with a few conspicuous 

parallel veins reticulated near the .margin. IDtimate 

surface pattern minutely papillate. 

No leaves identifiable as those of SagUtruria were 

found with the seeds; but foliage assigned to AliMnaphyllites occurs at a number of other Paleocene localities. The relation if any, between these leaves and 

the seeds here called S agittaria remains undemonstrated. 

Occurrence : Fort Union formation (upper) 4268 

(fig. 14), 9129 (figs. 15, 16), 9400. 

HYDROCHARITACEAE 

Hydromystria expansa (Heer) Hantke 

Plate 16, figures 1, 3, 8-11 

Hiraea ea:pansa Heer, 1859, p. 65, pl. 121, figs. 16, 16b. 

H ydromystria ea:pansa ( Heer) Hantke, 1954, p. 81, pl. 14, 

figs. 9-12. 

Lernna (Spirodela) scutata Dawson, 1875, p. 329, pl. 16, figs. 5, 

6, 7a ; 1887, p. 23, pl. 1, fig. 16. 

Ward, 1887, p. 17, pl. 3, figs. 4, 5. [Exclude Lesquereux 

citation in Ward's synonymy.] 

Spirodela scutata Dawson. Berry, 1935, p. 182. 

Bell, 1949, p. 82, pl. 63, figs. 1, 3; pl. 67, fig. 1. 

Nymphaeites browni Dorf, 1942, p. 142, pl. 10, fig. 9. 

Carpites verrucosus Lesquereux, 1878, p. 305, pl. 60, fig. 3. 

Dorf, 1942, p. 157, pl. 17, fig. 7. 

_ This was a creeping or floating aquatic with more 

or less asymmetrical, orbicular to reniform, cordate, 

fleshy, hairy, entire leaves, sometimes in pairs, one 

often smaller than the other, sessile or nearly so at 

nodes on a succulent prostrate stem or runner. Feathery rootlets may sometimes be seen at the nodes. The 

venation consists of 10 or more parallel veins that 

arise from the top of the petiole and converge toward 

the apex. Between the veins is a network of relatively 

large uniform areoles that probably were air chambers. In fossilization these spaces filled with mud 

that hardened and, after the residue of carbon was 

removed, caused the surface to be papillated, tuberculated or verrucose, deceptively like the surface of some 

seeds or fruits, whence the erroneous identification as 

0 a:rpites verru<Josus by Lesquereux. 

Oswald Heer also identified as fruits some similar 

specimens he found in the Miocene strata of southern 

Germany. Hantke, after restudy of this and additional material, assigned the species to H ydromystria, 

a genus of the Hydrocha.ritaceae, which is a family 

of aquatic monocotyledons. Hydiromystria includes 

only a few species that are distributed from Mexico 

to Paraguay. 

I do not regard the assignment to H ydrowystria as 

completely appropriate._ Neither does Tilo Notzold 

(1957, p .. 95, 96). One objection is the fact that the 

leaves of living species of Hydromystria are long 

petioled, whereas those of the fossils are sessile or 

nearly so. An assignment to LimJIU)biwm, also in the 

Hydrocharitaceae, would be open to the same objection. Pistia, an aquatic in the Araceae, almost meets 

the requirement of matching the fossils, but its leaves 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 53 

differ notably in shape. . Lewna, in the Lemnaceae, 

is too small. Thus, no closely comparable living counterpart seems to exist. The assignment to H ydromystria may be retained as a working hypothesis until 

further enlightening evidence accumulates. The fossil 

species ranged from the Late Cretaceous into the early 

Eocene in the United States and Canada and to the 

Miocene in Europe. 

Occurrence: Fort Union formation (lower) 2421, 

4031, 5512 (fig. 10), 6652, 6667, 8519, 9402, 9403, 9404; 

(upper) 436, 2420 (fig. 8), 4897 (fig. 1), 4974, 5321, 

6376, 6985, 8191 (figs. 9, 11), 8196, 8212 (fig. 3), 82244 

8556, 8786, 8920, 8922, 9109, 9405; Coalmont formation, 6005; Dawson arkose 8881; Denver formation, 

9401; Ferris formation, 6417. 

Grasslike monocotyledons 

Plate 14, figure 3; plate 16, figure 7 

Fragments of linear strap-shaped leaves wit:P, closely 

spaced parallel venation have been described from 

Paleocene localities as species of Oarew, Oyperaeites, 

Oyperus, Iris ( cf. Iris sp. Newberry, 1898, p. 33, pl. 

22, fig. 6), Phragmites ( cf. P. alaskmna Heer in Ward, 

1887, p. 17, pl. 3, figs. 1-3), and Poaeites. Associated 

with this foliage (pl. 14, fig. 3) at the same localities 

are rootstocks (pl. 16, fig. 7) or rhizomes identified as 

OaruliJnites and Oulmites. At present there seems to 

be no reliable method for distinguishing such remains 

as grass, sedge, iris, or something else. 

Occurrence: Fort Union formation (lower) 8519, 

8678 (pl. fig. 7); (upper), 2417 (pl.14, fig. 3). 

PALMACEAE 

Students of fossil palms recognize the difficulties 

in identifying such remains in terms of living species.· 

Consequently, arbitrary working classifications are 

necessary and herewith is such a key to the foliage 

of Cretaceous and Tertiary palms of the Rocky Mountains and Great Plains region. 

Leaves fan shaped, with radiate venation: 

Apex (acumen or hastula) of upper and under side of the 

petiole wedge-shaped, very stout, much prolonged, with 

fairly straight margins _______________ SabaZ imperialis. 

Apex of upper side of the petiole short, wedge-shaped, angular, with inside angle of less than 100°; apex of 

under side moderately prolonged with concave mar- gins ____________________________________ Sabal poweUt 

Apex of upper side of the petiole irregularly rounded or 

only slightly angular, with inside angle of 100° or more; 

apex of under side moderately prolonged, with concave margins _____________________________ SabaZ grayana. 

Apex of both sides of the petiole very short, 

rounded ----~-------------------------Thrina~ dorft. 

Leaves pinnate or feather-shaped, with very gradually tapering rachis: 

Pinnules linear, narrow, not 

decurrent ___________________ Phoenicites integrifolius. 

Pinnules linear, broad, decurrent_ ____ Ohamaedorea danae. 

Pinnules elliptic _________________ Paloreodomites plicatus. 

A number of Paleocene seeds (Palmoearpon) have 

been described as those of palms, but most of these are 

doubtfully so assigned. See the discussion (p. 89) 

under "Objects of uncertain classification." Palm 

wood (Palmowylon) is present at a number of Paleocene localities but is relatively rare, except in the Denver Basin, Colo. On the southeast-facing slope of 

South Table Mountain near Golden, I have seen a 

small palm stump in place, with adventitious roots 

running down into what was once the soil in which 

tbe tree grew. 

The fragmentary foliage described by Heer ( 1883, 

p. 69, pl. 68, figs. 5, 5b, 6, 7; .P· 70, pls. 140-106, fig. 1) 

as Flabellaria gronlandiea and F. johnstrupi, from the 

Paleocene of Greenland, is indefinite, though perhaps 

not improbably palmaceous. No seeds or wood have 

been reported that might confirm the identification. 

Rejected palm names 

The binomials attached to the following fossil palm 

remains from the Rocky Mountains and Great Plains 

region are here rejected for naming purposes because 

the specimens described were mere fragments or were 

not figured. 

FZabeZZaria communis LesqU:ereux, 1876, p. 385. 

FZabeZZaria zinckeni Heer. Lesquereux, 1878, p. 110, pl. 9, 

figs. 6-8. 

Geonomites goZdianus ( Lesquereux) Lesquereux, 1878, p. 115, 

pl. 9, figs. 9, 9b, 9c. 

Knowlton, 1930, p. 39, pl. 10, fig. 12. 

Geonomites graminifolius Lesquereux, 1888, p. 44. 

Geonomites tenuirachis Lesquereux, 1878, p. 117, pl. 11, fig. 1. 

Knowlton, 1917, p. 291, pl. 62; 1930, p. 38, pl. 6; pl. 9, 

figs. 1, 2, 6. 

Geonomites schimperi Lesquereux, 1878, p. 116, pl. 10, fig. 1. 

.ilfanicaria haydeni Newberry, 1898, p. 31, pl. 64, fig. 3. 

PaZmacites goldianus Lesquereux, 1876, p. 385. 

Sabal rigida Hatcher, 1901, p. 263. 

Sabal communis Lesquereux, 1876, p. 385. 

Chamaedorea danae (Lesquereux) Berry 

Oalamopsis danai Lesquereux, 1863, in Dana, p. 513, fig. 795. 

Lesquereux, 1869, p. 411, pl. 14, figs. 1-3. 

Ohamaedorea danai (Lesquereux) Berry, 1916a, p. 179, pl. 12, 

fig. 4 ; pl. 13, figs. 1-3. 

"Geonoma" gigantea Knowlton, 1917, p. 291, pl. 61. 

Geonomites haydenii (Newberry) Knowlton, 1923, p. 152. 

The remains assigned to this species are fragments 

of more or less pinnate leaves. Characteristic features 

of the leaflets are their somewhat decurrent attachment to the rachis, and the comparatively slight de-
54 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

velopment of the~r midribs as compared with the 

flanking veinlets. -. 

Compared with the leaves called Phoenwites integrifolilus Ball (1931, p. 35, pis. 42-; 47, fig. 1) from the 

Indio formation (Eocene) of Texas, the latter have 

narrower, linear leaflets, attached by conspicuously 

constricted bases, not noticeably decurrent on the 

rachis. 

Occurrence: Raton formation, 5679. 

Paloreodoxite~ plicatus (Lesquereux) ·Knowlton 

:Plate 15, figures 3, 7 

Oreodoxites pUoat1ts J"esquereux, 1883, p. 122, pl. 18, figs. 1-4. 

Knowlton, 1917, p. 287, pl. 63, fig. 1. 

Paloreodomites plicatus (Lesquereux) Knowlton, 1930, p. 41, 

pl. 11, figs. 1-4: 

Brown, 1956, p. f08, pl. 33, fig. 5. 

The relation of . this species to living palms is uncertain because it is not known definitely whether or 

not the imprints are leaves or leaflets. The presumption is that they are leaflets of a pinnate palm, but 

they may represent some other monocotyledon than a 

palm. 

Occurrence: D~~son arkose, 8188 (fig. 3) ; Denver 

formation, 317 ( f\g. 7) ; Raton formation, 5803. 

Sabal grayana Lesquereux. 

Plate 15, figure 5; plate 16, figure 4 

Sabal grayana Lesquereux, 1869, p. 412, pl. 14, figs. 4-6. 

Sabalites grayanus ~(Lesquereux) Lesquereux. Berry, 1916a, 

p. 177, pl. 13, .figs. 1-3; pl. 14, fig. 1. 

Knowlton, 1900, p. 32, pl. 6, fig. 5 ; 1930, p. 36, pl. 9, fig. 5. 

SabaZ campbelli Newberry, 1989, p. 27, pl. 21, figs. 1, 2. 

Dawson, 1985, p. 142, pl. 5, fig. 7. 

Sabalites campbelli · __(Newberry) Lesquereux, 1878, p. 113. 

SabaZ grandifolia N~wberry, 1898, p. 28, pl. 25; pl. 64, figs. 2, 

2a. · 

SabaZ inquirenda K:gowlton, 1917, p. 288, pl. 56. 

Sabal powelli Newberry, 1898, p. 30, pl. 64, figs. 1, 1a, only. 

Sabal? rugosa Knowlton, 1917, p. 288, pl. 58. 

Sabal? ungeri (Lesq~ereux) Knowlton, 1917, p. 254, 289, pl. 59. 

Sabalites eocenica (Lesquereux) Dorf, 1938, p. 48, pi. 2, fig. 6; 

pl. 3, fig. 3; 1942, p. 131, pl. 7, fig. 1. 

Sabalites florissant( (Lesquereux) Berry, 1930c, p. 66, pl. 9. 

Flabellaria alaskat¥£ Hollick, 1936, p. 57, pl. 22, figs. 2a, 3; 

pl. 111 ; pl. 1~2. 

FZabellaria eocenic4 Lesquereux, 1878, p. 111, pl. 13, figs. 1-3. 

FZabellaria ftorissar~:ti Lesquereux, 1878, p. 144, pl. 24, figs. 1, 

2, 2a. 

Hollick, 1936, p. 56, pl. 21. 

This fan palm with large many-rayed leaves differs 

from others dis_pussed here as follows : The apex 

(acumen or hastula) of the petiole on the upper side 

is short, more or'Jess rounded to slightly angular, with 

an inside angle of more than 100°. On the under side 

the hastula is moderately prolonged or attenuated, 

with concave m~rgins. The type is from the Eocene 

of Mississippi. 

To this species probably belong the Greenland specimens named Flabellaria gronl~iJea Heer ( 1883, p. 

69, pl. 68, figs. 5, 5b, 6, 7) and F. johnstrupi Heer 

(1883, p. 70, pl. 104; 105; 106, fig. 1). However, a 

few paleobotanists, including Potonie and Gothan 

(1921, p. 14), Kryshtofovich (1929, p. 306), and Berry 

(1930d, p. 13), have expressed doubts concerning the 

identity of these specimens, as also of those from the 

Jurassic called Propalnwphyllwm liasinum by Lignier ( 1908, p. 121-152, figs. 1-6), some holding that 

they are not plants but rill or ripple marks. If 

Heer's specimens do not represent palms, one frequently used argument for a relatively warm climate 

in Paleocene time within the Arctic Circle in western 

Greenland, loses its force. On the other hand, the 

undoubted palms reported by Hollick from the Tertiary of Alaska came from localities only a few hundred miles south of the Arctic Circle. 

The on] y pa]ms so far reported from Canada are 

Sabal imperialis from the Upper Cretaceous at 

N anaimo on Vancouver Island and S. gray{]JfUJ; (called 

S. campbelli by J. W. Dawson) from the Tertiary on 

Burrard Inlet near Vancouver, British Columbia. 

Thus it appears that inland from the Pacific coast the 

northernmost limit for palms was in the latitude of 

the Yellowstone River in Montana. 

Occurrence: Fort Union formation (lower), 6051, 

8526, 8545, 8567; (middle) 2416; Animas formation 

6443, 6444, 7480, 9565; Denver formation, 317 (pl. 16, 

fig. 4), 8672 (pl. 15, fig 5) ; Ferris formation, 8662; 

Raton formation, 5094, 5139, 5142, 5467, 5469, 5712, 

5827. 

Sabal imperialis Dawson 

Plate 14, figures 2, 6 

Sabal imperialis Dawson, 1883, p. 26, pl. 6, figs. 23, 23b; 1894, 

p. 57' pl. 14, fig. 61. 

Newberry, 1898, p. 30, figs. 6, 6a. 

Sabal montana Knowlton, 1916, p. 335, pl. 85, fig. 2; 1917, 

p. 253, pl. 32, fig. 3 ; 1922, p. 119, pl. 3, fig. 4. 

Sabal grandifolia Newberry, 1898, p. 28, pl. 63, fig. 5 only. 

Sabal? leei Knowlton, 1917, p. 289, pl. 60. 

Sabal sp. Knowlton, 1916, p. 336, pl. 85, fig. 1. 

Sabalites gray anus ( Lesquereux) Lesquereux, 1878, p. 112, 

pl. 12, fig. 1 (fig. 2 of doubtful nature). 

Geonomites imperialis (Dawson) Bell, 1957, p. 37,pl. 22, fig. 5; 

pl. 23, fig. 2 ; pl. 24, fig. 3. 

Geonomites 1ungeri Lesquereux, 1878, p. 118, pl. 11, fig. 2. 

This fan palm with large, many-rayed leaves differs 

from others discussed here as follows: The apex 

(acumen or hastula) of the petiole on both the upper 

and the under side is unusually stout, greatly prolonged, and with relatively straight margins. 

The species was most abundant in the late Cretaceous, but at some localities it ranged into the early 

Paleocene. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 55 

Occurrence: Fort Union formation (lower), 5886 

(fig. 2); Dawson arkose, 8188 (fig. 6); Raton formation, 5146, 5147, 5679, 5826. 

Sabal powelli Newberry 

Plate 16, figures 2, 5, 6 

SabaZ powelli Newberry, 1883, p. 504; 1898, p. 30, pl. 63, fig. 6 

only. 

Sabalites powelli (Newberry) Berry, 1930c, p. 67, pl. 10, figs. 

6, 7. 

Sabal? unger·i (Lesquereux) Knowlton, 1917, p. 289, pl. 57. 

This fan palm with large many-rayed leaves differs 

from others discussed here as follows : The apex 

(acumen or hastula) of the petiole on the upper side 

is short and wedge shaped, with slightly concave margins and an inside angle of about 75° at the tip. On 

the under side the hastula is moderately prolonged 

with concave margins. 

The species was first described from the Green 

River formation (Eocene) of Wyoming. 

Occurrence: Fort Union formation (lower) 8519 

(figs. 2, 5, 6), 8673; Raton formation 5112, 5142. 

Thrinax dorft Brown, n. sp. 

Plate 14, figure 1 

Small fan palm with orbicular leaves, approximately 30 em in diameter. Rays 30, radiating from 

the short rounded apex of the slender unarmed petiole, 

which is 5 mm in diameter. 

This is apparently a smaller leaved species than 

Thrirnam eocenwa Berry (1914, p. 136, pl. 25) from 

the Eocene of Georgia, but the leaf has more rays. 

The genus Thrilnam includes a numbe.r of living species 

growing near the coast from southern Florida through 

the West Indies to Central America. 

The species is named for Dr. Erling Dorf, of 

Princeton University. 

Occurrence: Fort Union formation (lower) 8256 

(fig. 1). 

CANNACEAE 

Canna1 magnifolia Knowlton 

Plate 15, figure 2 

Oanna? magnifolia Knowlton, 1917, p. 254, pl. 36, fig. 3. 

Oanna? sp., Knowlton, 1917, p. 255. 

Oyperacites? tesselatus Knowlton, 1922a, p. 117, pl. 3, figs. 1, 2. 

Oanna cf. 0.? rnagnifolia Knowlton. Dorf, 1938, p. 49, pl. 2, 

fig. 5. 

Oannophyllites rnagnifolia (Knowlton) Bell, 1949, p. 81, pl. 64, 

fig. 2 ; pl. 65. 

There is little, if anything, to add to Knowlton's 

description of these monocotyledonous lea;ves. I can 

detect no constant differences between the specimens 

from the Vermejo, Medicine Bow, Laramie, and Fort 

Union formations. The type specimen from the Vermejo formation (Upper Cretaceous) is not well illustrated, much of the blade having been erased from 

the photograph. Furthermore, the heavier widely 

spaced veins appear somewhat exaggerated in the 

photograph, but in reality are comparatively subdued, 

and in some parts of the blade, as also in the Paleocene specimens, are not readily distinguishable. The 

thin veins are closely spaced and may number six or 

more between the slightly heavier veins. 

The genus Ownnophyllites was instituted by Brongniart in 1828 for a Paleozoic plant, 0. virletti, from 

the Stephanian of the Loire Basin in France, and 

considered by him to have been a monocotyledon. 

Later it was found to be a fern. The term, therefore, 

cannot with propriety be applied to the monocotyledons described here. 

Occurrence: Fort Union formation (middle and 

upper), 9134, 9200 (fig. 2). 

ZINGIBERACEAE 

Zingiberites dubius Lesquereux 

Zingiberites dubius Lesquereux, 1878, p. 95, pl. 16, fig. 1. 

Dorf, 1938, p. 49, pl. 2; fig. 4. 

The upper half of Lesquereux's specimen is not as 

illustrated but appears to represent a fragment of 

wrinkled bark. The lower half is u:n.doubtedly a portion of a monocotyledonous leaf. This specimen is 

from the Paleocene Denver formation. Dorf's small 

specimen from the Medicine Bow formation apparently resembles the lower half of the Lesquereux 

specimen. 

Occurrence: Denver formation, 317. 

DICOTYLEDONS 

SALICACEAE 

Salix aquilina Brown, n. sp. 

Plate 61, figures 1-3 

Leaves linear lanceolate, acute at apex and base. 

Margin with numerous crenate, gland-tipped teeth. 

Secondary veins widely spaced with a tendency to 

fonn closed loops. Intersecondaries obscure. 

These fragmentary leaves are referred to Saliro with 

some hesitation. If this plausible assignment holds, 

they are the first representatives of the genus and 

family to be recorded from the Paleocene of the 

Rocky ::tv.Iountains and Great Plains .. 

Occurrence: Fort Union (lower), 6592, 8678; 

(upper) , 9322 (figs. 1-3) . 

JUGLANDACEAE 

Carya antiquorum Newberry 

Plate 17, figures 1-7; plate 18, figure 4 

Oarya antiquorurn Newberry, 1868, p. 72; 1898, p. 35, pl. 31, 

figs. 1-4. 

Lesquereux, 1878, p. 289, pl. 57, figs. 1-5; pl. 58, fig. 2. 

Ward, 1887, p. 34, pl. 15, fig. 2. 
56 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Oelastrinites insignis (Heer) Bell, 1949, p. 71, pl. 57, fig. 4; 

pl. 58, fig. 2. 

Dryophylbum aquamarum Ward. Knowlton, 1917, p. 299, pl. 70, 

fig. 2. 

Hicoria antiquora (Newberry) Knowlton, 1919, p. 319. 

Juglans leconteana Lesquereux. Bell, 1949, p. 54, pl. 57, fig. 1. 

Juglans nigella Heer. Lesquereux, 1883, p. 235, pl. 46A, fig. 11. 

Bell, 1949, p. 55, pl. 57, fig. 5. 

Juglans schimperi Lesquere·ux, 1878, p. 287, pl. 56, fig. 10 [not 

other figures]. 

Prunus dakotensis Lesquereux, 1883, p. 237, pl. 46A, fig. 8. 

Quercus valdensis Heer. Lesquereux, 1878, p. 153, pl. 19, fig. 8. 

So far as I am a ware there is no reliable method 

for distinguishing between the many fossil leaflets 

assigned to Oarya and Juglams. Consequently, those 

here· called Oarya may represent several species of 

Oavrya, or a mixture of Oarya, Juglan.s, and other 

juglandaceous genera such as Engelhardtia, Platycarya, and R hoiptelea. In general, however the leaflets of 0 arya antiquorum, especially the terminals, 

have an elliptic rather than a spatulate outline, and 

the marginal teeth appear to be· more evenly sized 

without a marked tendency toward doubling. 

The leaf called Juglans sohimperi by Lesquereux, 

from the Evanston formation, contrary to the illustration, has a serrate margin, but the teeth are not 

well preserved. The leaf, PruJfUUS drikotensis Lesquereux, as illustrated, has too many teeth. The teeth are 

fewer, coarser, and more unequal in size, as here illustrated (pl. 17, fig. 7). This dentition and a suggestion of asymmetry indicate that the specimen is a 

small leaflet of Oarya a;ntiquorum. Knowlton's illustration (1917, pl. 70, fig. 2) of Dryophyllum aquamarum shows many coarse teeth. ·Actually the specimen is poorly preserved and no teeth are clearly discernible. 

The difficulty in distinguishing the leaflets of 0 arya 

and Juglans is duplicated for the nuts also. The 

latter are here assigned to the form genus J uglarndioarya. 

Occurrence: Fort Union formation (lower), 541 

(pl. 17, fig. 1), 2432,4688, 5259, 5526, 6299, 6738, 8551, 

8660, 8884, 9112; (upper) exact loc. unknown (pl. 17, 

fig. 4, reproduction of Newberry's pl. 31, fig. 4), exact 

loc. unknown (pl. 17, fig. 7, reproduction of Lesquereux's Prunus dakotensi8), 4262, 4898, 5905, 8195, 8206 

(pl. 17, fig. 2), 8222, 8224 (pl. 17, figs. 3, 6), 8234, 

8523, (pl. 17, fig. 5)' 8566 (pl. 17, fig. 5)' 9125, 9501; 

Animas formation, 9565 ; Coalmont formation, 5987; 

Dawson arkose, 5836, 8307; Evanston formation, Lesquereux's figures; Ferris formation, 6625, 6971 ; 

Middle Park formation, 3361; Raton formation, 5711 

(pl. 18, fig. 4)' 5712. 

Juglandicarya spp. 

Plate 19, figures 5, 7-11 

Molds of that part of the cavity of a nut, exposed 

when the nut dehisces naturally into its two halves 

and loses the meat, have been found at a number of 

Paleocene localities. These molds represent the yoked 

halves of the two cotyledons. The cotyledons themselves would be at right angles to them. 

As no husks or exteriors of shells seem to have been 

preserved, definite identification of these nuts, other 

than they belong to the J uglandaceae, does not seem 

warranted at this time. The different sizes and shapes 

suggest that different species are involved, but whether 

or not the differences are really specific or represent 

degrees of maturity is conjectural. Reid and Chandler (1933, p. 140) proposed a form genus, Jugla!ndioarya, to receive similar remains from the London 

Clay (Eocen·e), and Kirchheimer (1935, p. 82) proposed Oaryojuglans for equally unassignable nuts 

:from the German brown coals (Oligocene). The 

status o:f Juglandioarya was discussed by Scott (1935, 

p. 667; 1954, 'p. 73) . 

Occurrence: Fort Union formation (lower), 4876, 

55~6, 5917 (figs. 5, 10), 6738, 9112 (figs. 9, 11); 

(upper), 8910 (fig. 7), 9198 (fig. 8). 

Juglans berryana (Knowlton) Brown, n. comb. 

Dryophyllum berryaria Knowlton, 1924, p. 81, pl. 8, fig. 2. 

Juglans rugosa Lesquereux. Knowlton, 1917, p. 293, pl. 112, 

fig. 4. 

Magnolia angustijolia Newberry. Knowlton, 1924, p. 88, pl. 10, 

fig. 3 [counterpart of Dryophyllum berryana, pl. 8, fig. 2]. 

Nectandra lanoifolia (Lesquereux) Berry. Knowlton, 1924, 

p. 87, pl. 13, fig. 1. 

Rhus ooloradensis Knowlton, 1924, p. 90, pl. 10, fig. 4. 

These are long elliptic to oblanceolate leaflets with 

inconspicuously toothed margins. The secondary 

veins are widely spaced, parallel, looping near the 

margin. The items Dryophyllum berryam.a and 11/ agnolia arngustifolia were based inadvertently by Knowlton on counterparts o:f the same specimen. 

Occurrence: Animas :formation, 7 496; Raton :formation, 5679, 5714, 5797. 

Juglans taurina Brown, n. sp. 

Plate 56, figures 4, 8, 9 

Berohemia multinervis (Braun) Heer. Ward, 1887, p. 73, pl. 

33, fig. 1. 

Broad ovate entire asymmetric leaflets with blunt 

to acuminate apexes and cuneate-rounded bases. 

Petioles short. Secondary veins numerous, sometimes 

branched, looping near the margin, with a tendency 

to branch :from the midrib at right angles near the 

base o:f the blade. Tertiary connecting veinlets numerous, diagonal, conspicuous. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 57 

Occurrence: Fort Union formation (upper), 2423 

(figs. 4, 9), 4262 (fig: 8), 5595, 6342. 

Pterocarya glabra Brown, n. sp. 

Plate 18, figures 1-3, 5-9, 13 

Leaflets elliptic spatulate, smooth, serrate with 

small, moderately sharp teeth. Petioles rather long, 

unless all the leafllets seen are terminals. Secondary 

veins numerous, closely spaced. Tertiary veins very 

numerous. 

Some of these leaflets are scarcely distinguishable 

from those of Pterocarya hispida Brown, except that 

under the magnifying glass they appear to lack the 

glandular dots of that species. Associated with these 

leaflets are nutlets (figs. 6-9) whose wings ·are 8 to 

10-lobed. These are tentatively assigned to this 

species. 

Occurrence: Fort Union formation (lower), 8930, 

6594, (upper) 8921 (figs. 1-3, 5-9, 13). 

Pterocarya hispida Brown, n. sp. 

Plate 18, figures 11, 12'; plate 19, figure 1; plate 68, 

figures 12, 13 

Leaflets ovate to. ovate spatulate, covered with 

minute, glandular dots, serrate with rather small, 

blunt teeth. Terminal leaflet long petioled, side leaflets short petioled. Secondary veins numerous, closely 

spaced. Tertiary veins very numerous. 

Some of these leaflets, on the basis of shape, serration, and venation alone, apparently are indistinguishable from those of Oarya antiquoT1J!m Newberry and 

Pterocarya glabra Brown. Associated with them, 

however, are entire-winged fruits (pl. 68, figs. 12, 13) 

similar to those of the living Pterocarya paliurus 

Batalin, and these are tentatively assigned to the same 

species. They are gland dotted, like the leaves. 

Occurrence: Fort Union formation (lower), 6839. 

(pl. 68, figs. 12, 13); (upper) 8910 (pl. 19, fig. 1) 

8920 (pl. 18, figs. 11, 12) ; Coalmont formation, 6005. 

BETULACEAE 

Betula stevensoni Lesquereux 

Plate 20, figures 1-5, 7-9 

Behtla stevensoni Lesquereux, 1878, p. 139, pl. 18, figs. 1-5; 

pl. 34, fig. lb. 

Betula gopperti Lesquereux, 1878, p. 138, pl. 17, figs. 21-23. 

Behtla vogdesi Lesquereux, 1878; p. 137, pl. 17, figs. 18, 19. 

Alnus ame1·icana Ettingshausen, 1883, p. 115. 

Al1Vus keferste-inii Goppert. Lesquereux, 1878, p. 140, pl. 18, 

figs. 6-8. 

Cassia concinna Heer. Lesquereux, 1878, p. 299, pl. 59, figs. 8, 

Sa. 

Cassia evanstonensis Knowlton and Cockerell. Knowlton, 

1919, p. 146. 

001·ylus macquan·ii (Forbes) Heer. Lesquereux, 1878, p. 144, 

pl. 18, fig. 10. 

Framinus denticulata Heer. Lesquereux, 1878, p. 228, pl. 40, 

fig. 1 [probably not fig. 2]. 

Rhus evansii Lesquereux, 1878, p. 291, pl. 58, figs. 5-9 [not 

pl. 50, fig. 4, which is Oastanaea intermedia Lesquereux]. 

The foregoing synonymy may exceed the bounds of 

probability that all belong to a single species. However, many of the types are based on battered specimens that scarcely permit typification of a speCies, but 

if more than one species is represented the distinction 

must await the finding of better material at those 

localities. 

These betulaceous leaves differ from those assigned 

to Oorylus insignis Heer in having basal secondaries 

with less pronounced tertiary branching and margins 

with more numerous and less coarse teeth. No fruits 

and seeds of Alnus, Betula, or Oarpitnus have been 

found in association with the leaves, but catkins were 

found at localities 4661, 8551, and 9109. 

Occurrence: Fort Union formation (lower), 4661 

(figs. 1, 3), 8551, 8893 (figs. 2, 4) 9104, 9180; (upper), 

2423, 4984 (fig. 8), 5905, 8522, 8917, 9109 (fig. 7), 

9199 (fig. 9) ; Coalmont formation, 6110; Evanston 

formation, 3658, 3661 (fig. 5), Livingston formation, 

6767, 8896. 

Corylus insignis Heer 

Plate 21, figures 1-11 

0or1}lus insignis Heer, 1871, p. 469, pl. 49, fig. 5. 

Oo1·1}lus macq•uarrii (Forbes) Heer, 1868, p. 104, pl. 8, figs. 

9-12; pl. 9, figs. 1-8; pl. 17, fig. 5d; pl. 19, fig. 7c; 1871, 

p. 469, pl. 44, fig. 11a ; pl. 45, fig. 6b. 

Lesquereux, 1878, p. 144, pl. 18, fig. 11 [not fig. 9, which 

is V·iburnum asperum Newberry;. not fig. 10, which is 

Bet·ula stevensoni Lesquereux]. 

Ward, 1887, p. 30, pl. 13, fig._ 7 .. 

Newberry, 1898, p. 61, pl. 32, fig. 5 [not pl. 48, fig. 4, which 

is TUia oregona LaMotte]. 

Oorylus fosteri Ward, 1887, p. 29, pl. 13, figs. 5, 6. 

Oorylits americana Walter. Ward, 1887, p. 28, p. 11, figs. 3-5; 

· pl. 12, figs. 1, 2. 

Ooryl1ts americana tossilis Newberry, 1898, p. 60, pl. 29, figs. 

8-10. 

Co1·ylus orbiculata Newberry, 1898, p. 62, pl. 32, fig. 4. 

Cor1}lus rostrata Aiton. Ward, 1887, p. 29, pl. 13, figs. 1-4. 

Oorylus rostrata tossil·is Newberry, 1898, p. 63, figs. 1-3. 

Corylites tosteri (Ward) Bell, 1949, p. 53, pl. 33, figs. 1-5, 7. 

Betula basiserrata Ward, 1887, p. 32, pl. 14, fig. 4. 

Betiula coryloides Ward, 1887, p. 31, pl. 14, fig. 3. 

Betula prisca Ettingshausen. Ward, 1887, p. 31, pl. 14, fig. 2. 

Pterospennites whitei Ward. Bell, 1949, p. 69, pl. 47, figs. 1, 5. 

The leaf originally identified by Forbes from the 

Ardtun beds (?Eocene) of the Isle of Mull as A.l!nites 

ma.cq'U£lff7'ii is fragmentary, but upon it Heer established Ooryl!us ma.cquarrii represented by equally fragmentary specimens from Atanekerdluk in Greenland. 

On the basis of a leaf from the same strata, Oorylus 
58 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

insignis Heer is a less controversial name for these 

American specim.ens. 

Although this species is represented by an a bundane~ and variety of leaves in the Paleocene of the 

Western States, only one nut (fig. 6) has been taken. 

This is small, being only 5 mm in diameter. Perhaps 

some shrinkage . occurred before fossilization. One 

male catkin (fig. 9) has also been found. 

The relationship of this species to those identified 

by Hollick as Ooryltus in the Tertiary floras of Alaska 

is not clear. Some Alaskan Tertiary species of Aln!U8 

and Betula may also be confused with Ooryltus. 

Occurrence: Fort Union formation (lower), 8224, 

8519 (figs. 9, 10), 8551, 8552, 8897, 8930, 9104; (upper), 2414 (fig. 5), 2416 (figs. 1, 6, 11), 3979, 4974, 

8231, 8255 (fig. 4)' 8921, 9072 (figs. 2, 3, 8)' 9202 (fig. 

7) ; Livingston formation, 4310, 6765. 

FAGACEAE 

Castanea intermedia Lesquereux 

Plate 22, figures 3, 4, 6, 7-10 

Castanea intermedia Lesquereux, 1878, p. 164, pl. 21, fig. 7. 

Knowlton, 1917, p. 297, pl. 68, fig. 2; 1924, p. 81, pl. 8, 

fig. 1. 1930, p. 50. 

Dryophyllum moori (Lesquereux) Berry. Knowlton, 1917, 

p. 299, pl. 70, fig. 1. 

Rhus evansi Lesquereux, 1878, p. 291, pl. 50, fig. 4 [not other 

figures]. 

The type of this species, it should be noted, is from 

the Paleocene of Middle Park, Colo., and not from 

the Oligocene at Florissant, Colo., as listed in 

LaMotte's catalog ( 1952, p. 106). 

In general the leaves here assigned to 0 astanea are 

relatively long and narrow, with fairly sharp, slightly 

hooked marginal teeth. Notably, the sinuses between 

the teeth are deep and angular below. The specimen 

described but not figured by l{nowlton (1930, p. 50) 

is somewhat atypical in having one secondary vein 

that is forked near the midrib, and a number of secondaries that are forked near the margin, sending 

short branches into teeth. 

No fruits or nuts recognizable as those of Castanea 

have thus far been found in American Paleocene 

strata. One specimen of 0. wngeri Heer ( 1883, p. 84, 

pl. 69, fig. 3), from Atanekerdluk, Greenland, may 

belong here. 

Occurrence: Fort Union formation (lower), 541 

(fig. 7), 6344, 8928 (figs. 3, 4, 6); (upper) 8521, 8774; 

Animas formation, 7485; Dawson arkose, 5836 (fig. 9), 

5839; Ferris formation, 5495 (fig. 10), 6428, 9203 

(fig. 8); Middle Park formation, 333; Raton formation, 5712. 

Quercus asymmetrica Trelease 

Plate 22, figure 1 

Querctts dubia Newberry, 1898, p. 73, pl. 37, fig. 5. [Homonym.] 

Quercus asymmetrica Trelease, 1924, p. 27. 

These fragments have the appearance of belonging 

to the Fagaceae but may, when adequate material is 

found, be assigned elsewhere. 

Occurrence: Fort Union formation, exact locality unknown (fig. 1, reproduction of Newberry's type). 

Quercus greenlandica Heer 

Plate 19, figures 3, 6, 12; plate 22, figure 5; plate 40, 

figures 1, 2 

Quercus groenlandica Heer, 1868, p. 108, pl. 8, fig. 8 ; pl. 10, 

figs. 3, 4; pl. 11, fig. 4; pl. 47, fig. 1; 1871, p. 56, pl. 12, 

figs: 1-4; p. 471, pl. 45, figs. 4, 4b; 1883, p. 89, pl. 69, 

fig. 4 ; pl. 89, figs. 1, 2 ; pl. 91, figs. 1, 2a. 

Quercus furcinervis Rossmassler, Heer, 1868, p. 107, pl. 7, figs. 

6a, 7a; pl. 45, fig. 1d; pl. 46, fig. 6. 

Quer·cus steenstrupiana Heer, 1868, p. 109, pl. 11, fig. 5; pl. 46, 

figs. 8, 9 ; 1883, p. 92, pl. 69, fig. 5. 

Castanea ungeri Heer, 1883, p. 84, pl. 69, fig. 3; pl. 73, fig. 14; 

pl. 88, fig. 3 ; pl. 89, fig. 4. 

Fagus castaneaefolia Unger. Beer, 1868, p. 106, pl. 10, figs. 

7a, 8; pl. 46, figs. 1-3. 

Fagus dentata Unger. Beer, 1868, p. 106, pl. 10, figs. 1, 2, 7b, 9. 

Fagus deucalionis Unger. Heer, 1868, p. 105, pl. 8, figs. 1-4; 

pl. 10, ,fig. 6; pl. 46, fig. 4; 1871, p. 470, pl. 46, fig. 9; 

1875, p. 5, pl. 3, figs. _11, 12. 

The numerous specimens here referred to Quercus 

and hitherto identified variously as 0Mtanea, Fagus, 

and Quercus, are characterized particularly by large, 

more or less rounded teeth between which the sinuses, 

although somewhat scalloped, are nevertheless slightly 

or noticeably angular. Several species and genera 

may here be! confused, but, in the absence of confirmatory seeds or fruits, I do not see how they can 

be separated satisfactorily. Probably the fragmentary 

leaves identified by Heer from the Paleocene strata 

at Atankerdluk, Greenland, as Fagus macrophylla 

Unger, Quercus? at(J!Va Heer, and Quercus drymeia 

Unger, also .. bel~:mg here. 

Occurrence~: Fort Union formation (lower), 4010 

(pl. 19, fig. :3), 5526 (pl. 40, figs. 1, 2), 6171 (pl. 19, 

fig. 12), 9111, 9208, 9482; (upper), 4878 (pl. 19, fig. 

6), 9132, 9196; Animas formation, 9549; Coalmont 

formation, 6000 (pl. 22, fig. 5) ; Evanston formation, 

5551. 

Quercus macneili Brown, n. sp. 

Plate 19, figure 2 

Leaves elliptic in outline, with cuneate bases. Margin with coarse toothlike, sharp-pointed lobes, separated by angular sinuses. Secondary venation somewhat irregular, the veins entering the marginal lobes. 

Surface pattern a quadrangular meshwork. 

This species is a representative of the blackoak 

group. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 59 

Named for F. Stearns MacNeil, of the U.S. Geological Survey. 

Occurrence: Fort Union formation (lower), 8239 

(fig. 2). 

Quercus sullyi Newberry 

Plate 23, figures 1-7; plate 27, figure 9; plate 57, figures 6, 7 

Que1·c1ts sullyi Newberry, 1883, p. 506; 1898, p. 79, pl. 60, fig. 2. 

QtteTctts aTtocaTpites Ettingshausen. Hollick, 1936, p. 102, 

pl. 43, fig. 1 ; pl. 56, fig. 6. 

QueTctts chaTpenteri Heer, 1883, p. 93, pl. 73, figs. 11-13; pl. 74, 

fig. 9. 

QtteTcus conjunctiva Hollick, 1936, p. 101, pl. 42, figs. 3, 4a. 

Q1teTmts /1tTC'ineTV'iS Rossmassler. Heer, 1883, p. 89, pl. 74, 

fig. 8. 

Que1·cus jttglandina Heer, 1883, p. 89, pl. 71, fig. 19; pl. 74, 

figs. 4-7; pl. 76, fig. 12; pl. 102, fig. 9a. 

Hollick, 1936, p. 101, pl. 42, figs. la, 2 ; pl. 43, fig. 2. 

QtteTcns lahaTpi Gaudin. Heer, 1883, p. 92, pl. 74, figs. 1-3. 

Quercus la•uTifoUa Newberry, 1898, p. 76, pl. 59, fig. 4; pl. 60, 

fig. 3. 

Q1te1·mts lyelli Heer, 1883, p. 87, pl. 72, figs. 1-10; pl. 73, figs. 

1-6. 

Q1tm·cus me1·iani Heer. Hollick, 1936, p. 102, pl. 44, fig. 1. 

QtteTmts penhallowi Trelease, 1918, p. 499. 

.AndTomeda dentimtlata Heer, 1868, p. 116, pl. 50, figs. 11d, 11e. 

[Specific term preoccupied by another fossil oak.] 

.Amlia ta1winensis (Ward) Sanborn, 1937, p. 27, pl. 10, figs. 

1, 2, 4. 

Dryophyllum aquiloni1tm Hollick, 1936, p. 104, pl. 43, fig. 6. 

Fmxin~ts juglandina Saporta. Hollick, 1936, p. 163, pl. 42, 

fig. 4b only. 

Ilea; lOngifol!ia Heer, 1868, p. 124, pl. 48, figs. 3-6; 1871, p. 481, 

pl. 50, fig. 17; pl. 56, t1g. 1. [Name given to a dubiolis 

Miocene fragment from Italy.] 

!lex? retioulata Heer, 1868, p. 124, pl. 48, fig. 7. [Name preoccupied by a living oak.] 

!lex tTiboleti Heer, 1883, p. 129, pl. 72, fig. lOb ; pl. 73, figs. 

7-10. 

Juglans bilinica Unger. Heer, 1883, p. 100, pl. 69, fig. 8. 

Juglans CTossi Knowlton. Hollick, 1936, p. 80, pl. 44, figs. 3, 4. 

Juglans denUculata Heer, 1871, p. 483, pl. 56, figs. 6-9a. 

J1tglans heeTi Ettingshausen. Heer, 1883, p. 102, pl. 76, figs. 

2-11. 

Juglans sp. Hollick, 1926, p. 42, fig. lb. 

MyTica ac1t1ninata Unger. Heer, 1883, p. 78, pl. 71, figs. 6, 7. 

M1JTica langeana Heer, 1883, p. 78, pl. 71, figs. 1-5; pl. 86, fig. 5. 

M y1·sine g1·onlandica Heer, 1.883, p. 11.1, pl. 81, figs. 4-8 ; pl. 85, 

fig. 4. 

Popnl1ts meedsU Knowlton, 1893,. p. 34, pl. 1, figs. 1, 2. 

PTotoficns 'inaequaUs Newberry, 1898, p. 89, pl. 60, fig. 1 only. 

Pnm1ts scotti Heer, 1871, p. 483, pl. 55, fig. 5; 1883, p. 137, 

pl. 84, fig. 13. 

PteToca1·ya denUculata Heer, 1883, p. 102, pl. 76, fig. 1. 

Ptm·oca1·ya septentTionale Hollick, 1936, p. 84, pl. 40, figs. 5-7. 

Sapind1ts <undulatus Braun. Heer, 1883, p. 127, pl. 84, figs. 1-3. 

Well-preserved impressions of these elliptic-lanceolate entire to toothed leaves display several characteristic features that render identification easy. The 

sharp, somewhat dentate teeth, when present, are 

separated by scalloped sinuses; the prominent forks 

of the undulant secondary veins connect to form loops 

near the margin; and the minute venational surface 

pattern makes a conspicuous, quadrangular network. 

Although these leaves have been variously identified, as shown by the synonymy, there is no certainty 

now that they represent Quercus. The species resembles variants of the Late Cretaceous Dryophyllum. 

sub falcatum Lesquereux and may be descended from 

it. The resemblance to fossil species of Oe~trus and 

Viburnum is more remote. The latter have rounded 

or sometimes cordate bases and more numerous teeth 

which are serrate, not dentate. The species left 

abundant remains at many localities, but particularly 

in the late Paleocene and early Eocene. 

Occurrence: Fort Union formation (lower), 4289, 

6602, 8239 (pl. 23, fig. 4; pl. 57, figs. 6, 7) 8253 (pl. 

27, fig. 9), 8552 (pl. 23, fig. 3); (upper), exact locality unknown (pl. 23, fig. 2, reproduction of Newberry's 

type, pl. 60, fig. 2), 436 (pl. 23, fig. 1), 2414 (pl. 23, 

fig. 5, reproduction of l{nowlton's pl. 1, fig. 1 of 

Populus meedsii), 5582, 6985, 8257, 8885, 8886 (pl. 23, 

figs. 6, 7), 8913, 8920; Coalmont formation, 6005 . 

Quercus yulensis Brown, n. sp . 

Plate 19, figure 4 

Leaf elliptic to obovate in outline with blunt apex, 

large rounded teeth and narrow, deep sinuses between 

the teeth. Secondary veins few, widely spaced, subparallel, and entering the teeth. 

The fewer secondaries and larger teeth readily distinguish this species from Quercus greenlandica Heer, 

another member of the white oak group. 

Occurrence: Fort Union formation (lower) 8239 

(fig. 4). 

ULMACEAE 

Celtis newberryi Knowlton and Cockerell 

Plate 27, figures 3, 4 

Celtis newbeTTyi Knowlton and Cockerell. Knowlton, 1919, 

p. 160. 

Celtis paTvifolia Newberry, 1898, p. 84, pl. 53, fig. 6. 

CoTnus denveTensis Knowlton. Bell, 1949, p. 76, pl. 56, fig. 5 

(lower leaf). 

Leaves more or less asymmetric, with slightly 

rounded bases and acute apexes. Margins entire or 

with coarse, serrate teeth. Venation incipiently palmate, with a minor. pair of subdued veins between the 

primary pair and the top of the petiole. Primary and 

secondary veins enter marginal teeth and emit 

branches to subsidiary teeth. 

Occurrence: Fort Union formation (middle), exact 

locality unknown (fig. 3, reproduction of Newberry's 

type); (upper), 9109 (fig. 4). 
60 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Celtis peracuminata Brown, n. sp. 

Plate 20, figure 6 

Leaf ovate lanceolate in outline, with rounded base 

and attenuate apex. Margin with .short serrate teeth. 

Venation palmate. Basal primaries curve narrowly 

upward toward the apex, their numerous branches 

forming conspicuous loops from which minute 

branches run into the teeth. V einlets connecting the 

major venation relatively coarsely spaced. 

Occurrence : Fort Union formation (upper), 7688 

(fig. 6). 

Planera microphylla Newberry 

Plate 24, figures 1-11, 13, 15, 16 

Planera microphylla Newberry, 1868, p. 55; 1898, p. 81, pl. 33, 

figs. 3, 4. 

Rhus? nervosa Newberry, 1898, p. 114, pl. 33, figs. 5, 6. 

Rhus unitus Knowlton and Cockerell. Knowlton, 1919, p. 553. 

U~mus antecedens Lesquereux. Knowlton, 1930, p. 62, pl. 23, 

fig. 7. 

Most of these leaves are symmetric or only slightly 

asymmetric, with cordate bases. The marO'inal teeth • b 

are Single, double, or mixed. The striking feature 

of the best leaves is the notable forking of many 

secondary veins at some distance inward from the 

margin. Because no characteristic fruits of Pla.nera 

have been found with these leaves the identification 

should be regarded as tentative. The leaves may 

represent Ohaetoptelea, Ulmus, Zelkova, or some other 

ulmaceous genus. A suggestion of an ancestral form 

may be Phyll#es sp. (Dorf, 1942, p. 156, pl. 17, figs. 

5, 13), from the Lance formation (Upper Cretaceous) 

in eastern vVyoming. 

Occurrence: Fort Union formation (lower), 5385, 

5387, 5885 (fig. 10); (upper), 4264 (figs. 3-5, 8, 9, 

13), exact locality unknown (figs. 6, 16, reproduction of 

Newberry's type of Rhus~ '1Wrvosa), 4898, 5300, 6225, 

8206 (figs. 7, 11), 8166 (fig. 15), 8255, 8523, 8885 

(figs. 1, 2) 9398; Denver formation, 317. 

Ulmus rhamnifolia Ward 

Plate 24, figures 17, 18, 21-23 

Ulmus 1'hamnifolia Ward, 1887, p. 45, pl. 23, fig. 5. 

Ulm1ts orbimtlaris Ward, 1887, p. 46, pl. 23, fig. 6. 

Leaves relatively large, more or less asymmetric 

with rounded double marginal teeth. Secondary vena~ tion curving, equidistant, as in living species of Ulmrus. 

Fruit obovoid, with the seed about the width of the 

wing, resembling that of the living Ul1nus japonica 

Sargent of eastern Asia. 

The present correlation of leaves and fruit seems 

reasonably certain because no other elmlike leaves or 

leaves of any other family having elmlike fruits 

occur at these Paleocene localities. Heer's Sorbus 

grandifolia (1883, p. 483, pl. 54, fig. 4) is fragmentary, but it and his Ulmrus fruit ( 1883, p. 94, pl. 75, 

fig. 12), from Atanekerdluk, Greenland, probably belong with this species. Newberry ( 1883, p. 508) described but did not illustrate some large leaves from 

the Yellowstone River region of Montana, as Ulmus 

grandifolia. These were probably the same as 

U. rha.wn.ifolia but were never again alluded to, and 

their whereabouts is now unknown. As their identity 

cannot be established, the name is rejected. 

Occurrence: Fort Union formation (lower), 4620; 

(upper), 2414 (fig. 21, reproduction of vVard's type 

of Ulmrus rham;nifolia), 8910 (figs. 18, 23), 9109 

(fig. 22), 9207 (fig. 17). 

Zelkova planeroides (Ward) Brown, n. comb. 

Plate 22, figure 2; pl. 24, figures 12, 14, 19, 20 

Ulmus planeroides ·ward, 1887, p. 44, pl. 23, figs. 1, 2. 

Ulmus minima Ward, 1887, p. 45, pl. 23, figs. 3, 4. 

Ulmus -ward-ii Knowlton and Cockerell. Knowlton, 1919, p. 636. 

Planera ere-nata Newberry, 1898, p. 81, pl. 67, fig. 3. 

Planera Zingualis Knowlton and Cockerell. Knowlton, 1919, 

p. 464. 

Quermts castanopsis Newberry, 1898, p. 71, pl. 56, fig. 4. 

Quercus praegroenlandica Berry, 1935, p. 26, pl. 3, figs. 3-7. 

Bell, 1949, p. 53. 

The best preserved of these leaves are comparatively 

long and narrow, with cuneate or rounded, not cordate, 

bases. The teeth are rounded, ge~1erally single, but 

may occasionally be double. 

Occurrence: Fort Union formation (lower), 4035 

(pl. 24, fig. 12), 4620, 6154, 8163, 9104; (upper), 2414 

(pl. 24, figs. 14, 20, counterpart of vVard's type), 

3979, 5613, 7662, 8196, 8255, 8521, exact locality unknown (pl. 24, fig. 19, reproduction of Newberry's 

type of Quermts castarnopsis), exact locality unknown 

(pl. 22, fig. 2, reproduction of Newberry's type of 

Planera crenata) . 

MORACEAE 

Artocarpus lessigiana (Lesquereux) Knowlton 

Plate 25, figure 7 

Myrica? lessigii Lesquereux, 1878, p. 136, pl. 64, fig. 1. 

A1·tocarp1ts lessigiana (Lesquereux) Knowlton, 1893a, p. 24; 

1922a, p. 128, pl. 12, fig. 1 ; pl. 22, fig. 4. 

Berry, 1916a, p. 194, pl. 26, fig. 1. 

Ball, 1931, p. 52, pl. 37, figs. 1-3; pl. 38, figs. 1-8; pl. 39, 

figs. 1, 2. 

MacGinitie, 1941, l). 109, pl. 15; pl. 16, fig. 1. 

A_rtocarp1ts dicksoni Nathorst, 1890, p. 1-10, pl. 1, figs. 1-4. 

A.1·tocarpns dissecta Knowlton, 1917, p. 67, pl. 42, fig. 6. 

Artocarpu,s? gigantea Knowlton, 1930, p. 75. 

Artocarpus liriodendroides Knowlton, 1922a, p. 129, pl. 21, fig. 2. 

A1·tocarpus pungens (Lesquereux) Hollick, 1899, p. 281, pl. 38, 

figs. 1, 2. 

Berry, 1916a, p. 195, pl. 25, fig. 1; pl. 27, fig. 1; pl. 29, 

fig. 1. 

Knowlton, 1924, p. 85, pl. 12; 1930, p. 73, pl. 30, fig. 2; 

pl. 32, figs. 1, 2. 

,_j 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 61 

A.rtocarpus similis Knowlton, 1917, p. 306, pl. 77; pl. 78, figs. 

1, 2 ; 1930, p. 75, pl. 32, fig. 6. 

A.rtocarpophyll1t1n occidentale Dawson, 1894, p. 60, pl. 12, fig. 

51 ; pl. 13, fig. 52. 

Aralia p1mgens Lesquereux, 1883, p. 123, pl. 19, figs. 3, 4. 

QuerC1ts an,qustiloba Braun. Lesquereux, 1878, p. 161, pl. 21, 

figs. 4, 5. 

Quercus praean,qustiloba Knowlton, 1922a, p. 126, pl. 5, figs. 6, 7. 

Sterculia coriacea Knowlton, 1917, p. 272, pl. 48, fig. 1. 

Phyllites sp. Knowlton, 1924, p. 98, pl. 19, fig. 1. 

I have found no satisfactory method for distinguishing species among the materials here synonymize~ as 

Artocarpus lessigiair~a (Lesquereux) 1\::nowlton. It,· 

however, seems somewhat improbable that only one 

species prevailed over the range of time and space 

indicated by these specimens. Nevertheless, I can 

·duplicate almost exactly Nathorst's A. dicksoni from 

the Upper Cretaceous of Greenland with a specimen 

from the Denver formation (Paleocene) on South 

Table Mountain, Colo. No fruits comparable to those 

illustrated by Nathorst have yet been found in the 

United States. Thus far, also, no leaves recognizable 

as those of Artocarpus have been reported from the 

Paleocene of any locality in the Rocky Mountain 

region north of latitude 43'0 • This or a similar species 

was present in the Eocene of the Gulf Coastal Plain 

and of Yellowstone National Park. 

Occurrence:· Fort Union formation (low~r), 9237; 

Animas formation, 7 496 ; Dawson arkose, 8779; Denver formation, 317 (fig. 7), 8777; Puerco formation, 

7371; Raton formation, 5799. 

Ficus affi.nis (Lesquereux) Brown, n. comb. 

Plate 27, figures 1, 2, 5 ; plate 43, figures 7, 8 

Oinnamomum a:ffine Lesquereux, 1878, p. 219, pl. 37, fig. 5- [not 

other figures except the unnumbered one below fig. 1]. 

Oinnamonutm? ficitolium Knowlton, 1917, p. 318, pl. 90, fig. 3. 

Oinnamomum la1"tceolatium (Unger) Heer. Lesquereux, 1878, 

p. 219, pl. 36, fig. 12. 

Oinnamomum linifoli1tm Knowlton, 1917, p. 319, pl. 88, figs. 

3-7; 1930, p. 86, pl. 59, fig. 3. 

Dorf, 1938, p. 59, pl. 9, fig. 1. 

Oinnamom1tm polymorphum Braun. Lesquereux, 1878, p. 221, 

pl. 37, fig. 10. [Same as 0. ellipticum Knowlton, 1893, 

p. 54.] 

Oinnamomum salicoides Knowlton, 1924, p. 88, pl. 15, fig. 2. 

Oinnamomum? sp. Knowlton, 1930, p. 87, pl. 36, fig. 4. 

Ficus cann01vi Knowlton, 1922a, p. 136, pl. 6, fig. 3 ; pl. 10, fig. 1. 

Ficus dalmatica Ettingshausen. Knowlton, 1922a, pl. 21, fig. 9; 

pl. 22, fig. 5. 

Ficus haddeni Knowlton, 1917, p. 260, pl. 38, figs. 6, 7. 

Fimts neoplanicostata Knowlton, 1930, p. 69, pl. 29, figs. 3, 4. 

Ficus planicostata problematica Knowlton, 1930, p. 71, pl. 28, 

fig. 2. 

Ficus pop,uloides Knowlton, 1900, p. 44, pl. 8, fig. 3. 

Ficus posttrinervis Knowlton, 1922a, p. 136, pl. 6, figs. 1, 2. 

Ficus praetrine1·vis Knowlton, 1917, p. 263, 304, pl. 41, figs. 1-3 

[not fig. 4, which is F. planicostata Lesquereux] ; 1930, 

p. 71, pl. 28, figs. 8-10. 

Ficus pseudopopulu~ Lesquereux. Knowlton, 1917, p. 304, pl. 

72, fig. 2 [not figs. 3, 4, which are F. planicostata Lesquereux] ; pl. 73, figs. 1, 2 ; pl. 112, fig. 3 ; 1924, p. 83, 

pl. 9, fig. 3. 

Ficus trinervis Knowlton, 1900, p. 42. 

Lee, 1912, pl. 20, fig. 1. 

Dorf, 1938, p. 56, pl. 6, figs. 1, 4. 

Ficus sp. Knowlton, 1930, p. 72, pl. 30, fig. 1. 

Malapoenna louisvillensis Knowlton, 1922a, p. 144, pl. 7, fig. 5. 

Phyllites herbacea Knowlton, 1924, p. 97, pl. 15, fig. 4. 

Platanus platanoides (Lesquereux) Knowlton, 1930, p. 82, 

pl. 36, fig. 5. 

Populus arctica Heer. Lesquereux, 1878, p. 178, pl. 23, fig. 4. 

[Locality dubious, but not Spring Canyon, Mont.] 

Pop1tlus? distorta Knowlton, 1922a, p. 126, pl. 4, fig. 6. 

Zizyphus lanceolatus Knowlton, 1924, p. 93, pl. 15, fig. 3. 

Zizyphus minutus Knowlton, 1922a, p. 158, pl. 18, fig. 1. 

These triveined leaves, hesitantly assigned to Ficus, 

differ from those called Ficus planicostata Lesquereux 

chiefly in tl;le conspicuous decurrency of the lateral 

primaries into the petiole. Both species however, as 

now set up, include specimens having almost similar 

venation, so that choice of allocation is sometimes difficult and the result arbitrary. Both species range 

from the Cretaceous into the Paleocene and are associated at the same or nearby localities in correlative 

strata. 

Some extremely narrow specimens, formerly called 

Oi'flln(J,momwm linifoluim Knowlton, are here reassigned to FiOU8 atfinis, because they seem to be variants 

of this species a.nd are found at the same localities. 

As this species was based by Knowlton on one of 

Lesquereux's types of Oinnamomum affine, its proper 

name, instead of FiOU8 trinervis Knowlton, should 

have been Ficus affinis (Lesquereux) Knowlton, n. 

comb~ 

Occurrence: Fort Union formation (lower) 4625 

(pl. 43, fig. 8), 5194, 8899; Animas formation, 7481, 

(pl. 27, fig. 5), 7483; Dawson arkose, 5374, 5831, 6943, 

8779, 9554; Denver formation, 317 (pl. 27, figs. 1, 2), 

325, 8672, 8777; Evanston formation, exact locality 

unknown; Raton formation, 5683, 5684, 5686, 5699, 

5796 (pl. 43, fig. 7)' 5826, 5827. 

Ficus artocarpoides Lesq uereux 

Plate 28, figures 1-7 

Ficus artocarpoides Lesquereux, 1878, p. 47, pl. 47, figs. 1-5. 

Knowlton, 1917, p. 300, pl. 71, fig. 3. 

Oornus? tosteri Ward, 1887, p. 54, pl. 25, fig. 5. 

Oornus newberryi Hollick. Newberry, 1898, p. 124, pl. 37, fig. 

4 only. 

Meliosma cuneata (Newberry) Berry, 1939b, p. 377 [not text 

fig. 1]. 

Parrotia cuneata (Newberry) Berry, 1916a, p. · 219; 1930a, 

p. 71 [not pl. 12, fig. 10]. 

Phyllites retusoides Knowlton, 1917, p. 349, pl. 102, fig. 5. 

Protophyllum canadensis Berry, 1935, p. 31, pl. 4B. 

Protophyllum sp. Berry, 1926, p. 194, fig. 1. 
62 l.)ALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Pterospermites dawsoni (Knowlton) Bell, .1949, p. 70, pl. 47, 

figs. 2, 4 ; pl. 49, figs. 1, 2. 

Pterospermites penhallowi Berry, 1935, p. 49, pl. 13. 

Quercus bicontis Ward, 1887, p. 24, pl. 9, fig. 3. 

Quercus dawsoni Knowlton, 1898, p. 191. 

Quercus platania Heer. Dawson, 1889, p. 72, pl. 11. 

Rhamnus cannoni Knowlton, 1930, p. 107, pl. 46, fig. 5. 

Rhamnus cleburni Lesquereux. Knowlton, 1930, p. 104, pl. 46, 

figs. io, 11 only. 

Rhamnus goldianus Lesquereux. Knowlton, 1917, p. 332, pl. 

101, fig. 4. 

Viburnum cuneatum Newberry, 1883, p. 511; 1898, p. 130, pl. 57, . 

fig. 2. 

The types of this species, unfortunately, are fragmentary, and their present location is unknown to me. 

They are depicted as oval, blunt at the apex, cuneate, 

rounded, or cordate at the base, with entire margins, 

and long, stout petioles. The venation is pinnate, the 

secondaries evenly spaced, camptodrom~, and connected by numerous parallel diagonal veinlets. The 

lower secondaries may be branched near the margin. 

Collections from other localities in the Fort Union 

of Montana, particularly locality 8910, contain leaves 

that would be identified with the original FicU8 artocarpoides, but they also include toothed leaves that 

have been identified as Viburnum cuneatwm, Quercus 

da.wsoni, Oornus newberryi, and Pterospermites· penh.allo'toi. As all these differently shaped specimens, 

however, have identical venation and as they occur 

together at a number of localities, I am inclined to 

regard them as representing a single species; but, although their features are suggestive, I am not certain 

that they represent a species of Ficus. 

Except that these leaves are less coarse in every 

respect-size, venation, and dentition-they seem to 

be clearly descendant from Ficus preartocarpoides 

Brown (1939b, p. 249, pl. 53, figs. 3-5) and its synonym, Araliaephyllum artocarpoides (Lesquereux) 

Dorf (1942, p. 147, pl. 13, fig. 4) from the Hell Creek 

and Lance formations, respectively. Indeed, some of 

the Paleocene leaves may, by atavism, match those of 

the Late Cretaceous. Most of the Late Cretaceous 

leaves display an indefinable irregularity not found in 

the Paleocene specimens. 

The original illustration of Oornus newberryi Hollick should be compared with that here (fig. 1) made 

of the same specimen after the margin was cleaned, 

showing that the leaf is toothed, not entire. 

Occurrence: Fort Union. formation (lower), 4570, 

4625 (fig. 6)' 4860, 4876, 4984, 5839, 6117' 6668, 8246, 

8551, 8552; (upper), 2416 (fig. 5,. reproduction o:f 

Ward's type of Quer(]U8 bicornis), 4369 (fig. 3), 4676, 

5760, 8261 (fig. 4), 8774, 8910 (fig. 2), 8922, exact 

locality unknown (fig. 1, reproduction of Hollick's 

type of Oornu.s newberryi), exact locality unknown 

(fiO'. 7, reproduction of Newberry's type of Viburnum b . 

cwneatum) ; Evanston formation, 14 7 4, 3653 ; Ferris 

formation, 6630, 6971, 8516; Livingston formation 

4311; Middle Park formation, 337; Raton formation, 

5711. 

Ficus minutidens Knowlton 

Plate 48, figure 8 

Ficus minutidens Knowlton, 1917, p. 305, pl. 71, fig. 2. 

The distinctive feature of these figlike leaves is 

their dentate margin, with scalloped or angular sinuses 

between the teeth. The venation is essentially pinnate but with a suggestion of a tendency toward 

palmateness. Acceptance of these leaves as Ficus is 

made witn reservations, because they also resemble 

somewhat the leaves of some species of Hydrangea, 

M orus, Populus, Tilw, Viburnurm, and Vitis. 

Occurrence: Animas formation, 7496 (fig. 8); Raton 

formation, 5711. 

Ficus planicostata Lesquereux 

Plate 26, figures 1-8 

Ficus planicostata Lesquereux, 1878, p. 201, pl. 31, figs. 1-8, 

1Q-12. 

Lee, 1912, pl. 20, fig. 2. 

Knowlton, 1924, p. 82, pl. 9, fig. 2. 

Dorf, 1938, p. 53, pl. 5, figs. 3-5, 7 ; 1942, p. 136. 

Fious planicostata clintoni (Lesquereux) Knowlton, 1917, p. 303, 

pl. 76, fig. 3. 

Ficus planicostata goldiana Lesquereux, 1878, p. 202, pl. 33, 

figs. 1-3. 

Knowlton, 1930, p. 70, pl. 28, fig. 5. 

Ficus planicostata magnifolia Knowlton, 1922a, p. 133, pl. 10, 

fig. 3. 

Ficus berr'Yana Knowlton, 1922a, p. 139, pl. 11, fig. 11. 

Ficus cockerelli Knowlton, 1922a, p. 132, pl. 12, fig. 2 ; pl. 23, 

figs. 1, 2. 

Dorf, 1938, p. 55, pl. 7, fig. 2. 

Ficus dawsonensis Knowlton, 1930, p. 67, pl. 26, fig. 1. 

Ficus denveriana Cockerell. Knowlton, 1917, p. 302, pl. 75, 

fig. 2 only. 

Ficus impressa Knowlton, 1922a, p. 134, pl. 7, figs. 1-3. 

Ficus leei Knowlton, 1916, p. 338, pl. 90, fig. 2; 1917, p. 261, 

pl. 39, figs. 1-6 ; pl. 40, figs. 1, 2. 

Ficus neodalmatica Knowlton, 1922a, p. 135, pl. 7, fig. 6. 

Ficus neoplanicostata Knowlton, 1922a, p. 303, pl. 73, fig. 4; 

pl. 7 4, figs. 2, 3 ; pl. 76, fig. 4 ; 1924, p. 82, pl. 9, fig. 4 ; 

1930, p. 69, pl. 28, figs. 3, 4, 6, 7. 

Ficus occidentalis ( Lesquereux) Lesquereux, 1878, p. 200, pl. 

32, fig. 4. 

Knowlton, 1917, p. 302, pl. 72, fig. 1; 1924, p. 82, pl. 8, fig. 5 ; 

1930, p. 68, pl. 26, fig .. 5. 

li'icus pagosensis Knowlton, 1924, p. 84, pl. 11. 

Fious praeplanicostata Knowlton, 1922a, p. 133, pl. 22, fig. 2. 

F·icus praetrinervis Knowlton, 1917, p. 263, pl. 41, fig. 4. 

Ficus problematica Knowlton, 1900, p. 46, pl. 9, fig. 3. 

Ficus pseudopopulus Lesquereux, 1878, p. 204, pl. 34, fig. 1a 

only. 

Knowlton, 1917, p. 304, pl. 72, figs. 3, 4 only; 1930, p. 66, 

pl. 25, figs. 3-5; pl. 26, figs. 2, 4 only. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 63 

Ficu,s richardsoni Knowlton, 1917, p. 304, pl. 76, fig. 1. 

Fictts schimper'i Lesquereux. Knowlton, 1917, p. 304, pl. 75, 

figs. 3, 4. 

Ficus speciosissima Ward, 1887, p. 39, pl. 21, fig. 3. 

Lee, 1912, pl. 19. 

Knowlton, 1916, p. 90, pl. 16, fig. 3. 

Ficus squarrosa Knowlton, 1900, p. 45, pl. 48, fig. 2. 

Ficus tiliaefolia Braun. Lesquereux, 1878, p. 203, pl. 32, figs. 

1-3; pl. 63, fig. 8. 

Knowlton, 1930, p. 67. 

Ficus wardi Knowlton, 1900, p. 48, pl. 9, fig. 1. 

F'icu,s cf. mississipJJiensis (Lesquereux) Berry. Bell, 1957, p. 44, 

pl. 33, figs. 1, 5. 

Ficus sp. Knowlton, 1924, p. 84, pl. 6, fig. 7. 

Cinnamontum ?n-issiSS'ippiense Lesquereux. Knowlton, 1917, 

p. 320, pl. 89, fig. 2. 

Cissus coloradensis Knowlton and Cockerell. Knowlton, 1930, 

p. 112, pl. 47, fig. 6; pl. 50, fig. 2. 

Ciss1ts laevigatn Lesquereux, 1878, p. 238, pl. 40, fig. 13 [not 

fig. 12, which is Platanus raynoldsi Newberry]. 

Knowlton, 1917, p. 340, pl. 103, fig. 1. 

?Corntts stude1·i Heer. Knowlton, 1917, p. 342, pl. 109, fig. 2. 

Dombeyopsis obtusa, Lesq'uereux. Knowlton, 1922, p. 162; 

1930, p. 124, for reference to USNM 281d. 

Hedera rotundi/oUn Knowlton, 1917, p. 272, pl. 47, fig. 8. 

Jttglans thermnlis Lesquereux, 1878, p. 287, pl. 56, fig. 4 [not 

fig. 3, which is Sassafras thermale (Lesquereux) 

Brown]. 

Launts tttahensis Lesquereux. Knowlton, 1917, p. 318, pl. 70, 

fig. 4. 

Phaseolites colo1·adensis Knowlton, 1930, p. 97, pl. 45, fig. 1. 

Phyllites eocenica; Knowlton, 1924, p. 96, pl. 18, fig. 4. 

Phyllites sp. Knowlton, 1930, p. 130, pl. 56, fig. 3. 

'Vith the exception of the first and a few others, 

the citations in the foregoing synonymy concern 

specimens from known Paleocene localities. The first 

cites Lesquereux's types of this species from the 

Upper Cretaceous at Black Buttes, Wyo. Other citations for Cretaceous localities have been published 

by Dorf ( 1938, p. 53; 1942, p. 136). These synonymies 

indicate that this species, if it is a single species, 

comprises leaf specimens illustrating great variety 

of outline, but with virtually the same venation. The 

latter consists of a pair of strong lateral veins arising 

from or near the top of the petiole and curving upward toward the apex. These primary veins have 

relatively numerous, closely spaced, parallel side 

branches. Between the midvein and secondaries and 

between the secondaries are numerous, more or less 

closely spaced, parallel connecting veinlets. The 

apexes of these entire ovate to ovate-lanceolate leaves 

may be blunt rounded to long acuminate, and the 

bases may be strongly cordate to cuneate. The leaves 

from the Paleocene seem to average somewhat larger 

and are perhaps more acuminate than those from the 

Cretaceous. However, it seems that this species 

crossed the Cretaceous-Paleocene contact with only 

slight change, if any. 

The Cretaceous specimens called Ficus leei and 

F. speciosissima have strongly cordate bases but are 

otherwise like typical F. planicostata. l\{oreover, they 

occur in association with the latter and, I suspect, are 

merely extreme variants of it. 

Many items such as Hedera rotundifolia, Jur;law 

thermalis, and Phyllites sp. are fragments that might 

better have been left unnamed, or at least they should 

have received more careful scrutiny before being 

named. 

The relationship of Ficus planwostata to the Eocene specimens called F. wyomingiatna Lesquereux and 

F. mississippiensis (Lesquereux) Berry is not clear 

but may be close. 

The assignment of all these figlike leaves to Fie~, 

although plausible, is not confirmed by authentic associated fruits. The Cretaceous specimens named Ficus 

ceratops Knowlton are still controversial. Hence, 

there is an open possibility that F. plarnicostata, as 

well as F. affinis, may belong to some other genus and 

family. 

The following statement of occurrence indicates 

that, so far as known, this species was absent from 

Paleocene areas north of the latitude of Terry, Mont. 

Occurrence: Fort Union formation (lower), 4876 

(fig. 6), 6057, 8246, 8666; (upper), 8774 (figs. 2, 5, 7, 

8; fig. 2 is in Chicago Nat ural History Museum) ; 

9501; Animas formation, 5456, 7481 (fig. 3); Dawson 

arkose, 8188 (fig. 4); Denver formation, 317 (fig. 1), 

325, 8777; Evanston formation, exact locality unknown; Ferris formation, 8516; Puerco formation, 

7371, 7495; Raton formation, 5046, 5132, 5147, 5236, 

5678, 5679, 5684, 5687, 5688, 5689, 5711 5799. 

Ficus subtruncata Lesquereux 

Plate 25, figures 1-6 

Ficus subtruncata Lesquereux, 1878, p. 205, pl. 30, figs. 7-9. 

Knowlton, 1919, p. 289 ; 1924, p. 83, pl. 8, fig. 3 ; 1930, p. 63. 

Ficus auriculata Lesquereux, 1878, p. 206, pl. 30, figs. 4-6. 

l!'icus martini Knowlton, 1930, p. 69, pl. 40, fig. 5. 

The assignment of these specimens to Ficu.s is unsatisfactory, but I have no better suggestion. 

Occurrence: Fort Union formation (lower), 6598, 

8227 (figs. 1, 4), 8551 (fig. 2), 8901 (fig. 5); Animas 

formation, 7483; Denver formation, 317 (fig. 6); Ferris formation, 8516 (fig. 3). 

Ficus uncata Lesquereux 

Ficus uncata Lesquereux, 1878, p. 197, pl. 35, figs. 1, 1a only. 

Ficu,s duplicata Knowlton, 1917, p. 302, pl. 74, fig. 1. 

Populus mooodon Lesquereux, 1878, p. 180, pl. 24, figs. 1, 2. 

Quercus? neomexicana Knowlton, 1917, p. 298, pl. 70, fig. 5. 

M nunolia cordi folia Lesquereux. Knowlton, 1917, p. 315, pl. 

86 ; pl. 88, fig. 1. 

Magnolia rotundifolia Newberry. Knowlton, 1917, p. 314, pl. 83. 
64 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

These leaves resemble those of some species of Cornbret1tm and 11/ agnolia, and their present allocation IS 

dubious. 

Occurrence: Raton formation, 5142, 5684, 5714. 

:Morus montanensis Brown, n. sp. 

Plate 20, figure 10 

Leaf ovate, with cordate base and abruptly attenuate 

apex. Margin with numerous small rounded slightly 

crenate teeth, except in the region of the apex, which 

is smooth. Venation palmate with one pair of curved 

strong basal primaries, which have numerous side 

branches that connect near the margin to form loops. 

The first strong secondary branches from the midvein 

well below the middle of the leaf. Also branching from 

the midvein are short intermediates between the secondaries. 

I have found no lobed leaves like those from the living species of M or1t8. 

Occurrence: Fort Union formation (upper), 4032 

(fig. 10)' 8920. 

PLATANACEAE 

Platanus nobilis Newberry 

Plate 29, figures 1, 3-6 

Platanus nobilis Newberry, 1868, p. 67; 1898, p. 106, pl. 34; 

pl. 50, fig. 1 only. 

Dawson, 1886, p. 24, pl. 1, fig. 7. 

Platanus basilobata Ward, 1887, p. 35, pl. 17, fig. 1; pl. 18, 

figs. 1-3; pl. 19, fig. 1. 

Berry, 1935, p. 32, pl. 6, fig. 7. 

Bell, 1949, p. 58, pl. 42; pl. 43, figs. 1-3. 

Acer trilobatum tricuspidatum Heer. Ward, 1887, p. 66, pl. 29, 

fig. 3. 

Aralia acerifolia Lesquereux, 1883, p. 232, pl. 49, fig. 5 only. 

Aralia dakotana Knowlton and Cockerell. Knowlton, 1919, 

p. 82. 

Aralia, digitata Ward, 1887, p. 62, pl. 27, figs. 3-5; pl. 28, fig. 1. 

Aralia? gracilis Lesquereux, 1878, p. 236, pl. 39, fig. 1. 

Aralia looz·iana Saporta and Marion. Ward, 1887, p. 61, pl. 27, 

fig. 2. 

Aralia notata Lesquereux:, 1878, p. 237, pl. 39, figs. 2-4. 

Ward, 1887, p. 60, pl. 27, fig. 1. 

Aralia wardiana Knowlton and Cockerell. Knowlton, 1919, 

p. 87. 

Sassafras seltoyni Dawson, 1886, p. 28, pl. 2, fig. 13. 

Vibu,rnum oxucoccoides Dawson, 1886, p. 29, pl. 2, fig. 15. 

The consensus seems to be that these leaves represent Platanu.s rather than Aratia, despite the fact that 

in minute surface features and general outline they 

differ somewhat from those of living species of Platanus. Their gross architecture, however, is strikingly 

like that of the living leaves. In particular the forking of the lateral primary veins at a short distance 

above their emergence from the top of the petiole is 

matched in living leaves having five or more lobes. 

Some of the fossil leaves, like the living (fig. 2), have 

conspicuous basilar lobes that sometimes cover the top 

of the petiole. Lester 'Vard (1888; 1890) illustrated 

these features and speculated that they pointed toward the origin of stipules. That stipules arose in 

this way seems doubtful because it is anomalous that 

living leaves with basilar lobes have stipules in addition. 

When marginal teeth are present the sinuses between them are scalloped, as in living leaves. This 

combination of features is not, so far as I am aware, 

matched in any other living genus. A number of fossil 

species, somewhat like P. nobilis, have been described 

from Eocene and Oligocene strata in the Southern and 

Western United States. 

No authentic seed balls or seeds of Platanus have 

ever been reported from the same strata that contain 

these Paleocene leaves. The seed balls identified by 

Bell ( 1949, p. 58, pl. 43, fig. 1) as belonging to P. 

basilobata are instead the fruiting heads of Sparganium antiquum (Newberry) Berry. 

Occurrence: Fort Union formation (lower), 8567; 

(upper), 241'4 (fig. 3, reproduction of Ward's type of 

Aralia digitata), 2416 (figs. 1, reproduction of Ward's 

type of P. basilobata, 4) , 4984, ~594 (figs. 5, 6) , 7695, 

9109, 9125, 9322; Coalmont formation, 6103, 6107; 

Evanston formation, 5539; Ferris formation, 7548, 

9531; Middle Park formation, 337; Raton formation, 

5134. 

Platanus raynoldsi Newberry 

Plate 30, figures 1-4; plate 31, figures 1-6; plate 66, figure 8 

Platamts raynoldsi Newberry, 1868, p. 69; 1898, p. 109, pl. 35. 

Lesquereux, 1878, p. 185, pl. 27, figs. 1-3. 

Ward, 1887, p. 37, pl. 20, figs. 2, 3. 

Knowlton, 1917, p. 324, pl. 95, fig. 1 ; 1930, p. 77, pl. 35. 

Bell, 1949, p. 59, pl. 37, fig. 2 ; pls. 38-41; pl. 34, fig. 3 ; 

pl. 60, fig. 3. 

Platanus raynoldsi var. integrifolia Lesquereux, 1878, p. 185, 

pl. 26, figs. 4, 5. 

Platanus aceroides Goppert. Heer, 1868, p. 111, pl. 12, figs. 

1-8; pl. 32, figs. 1, 2; pl. 47, fig. 3; 1883, p. 96, pl. 97, 

fig. 1. 

Lesquereux, 1878, p. 184, pl. 25, figs. 4-6; 1883, p. 227, 

pl. 49, fig. 1. 

Knowlton, 1917, p. 321, pl. 63, fig. 4; pl. 97, figs. 2, 3. 

Platanus ace1·oides cuneata Knowlton, 1917, p. 321, pl. 113, 

fig. 1. 

Platan1ts ace1·oides latifoUa Knowlton, 1917, p. 321, pl. 92; 

pl. 93, fig. 3 ; pl. 94. 

Platanus coloradensis Knowlton, 1930, p. 82, pl. 37, figs. 1, 2; 

pl. 38, fig. 1. 

Platawus ,r;uillelrnae Goppert. Lesquereux, 1878, p. 183, pl. 25, 

figs. 1-3. 

Ward, 1887, p. 37, pl. 20, fig. 1. 

Knowlton, 1930, p. 76, pl. 34, fig. 1. 

Platanus haydeni Newberry, 1868, p. 70; 1898, p. 103, pl. 36; 

pl. 38; pl. 56, fig. 3. 

Knowlton, 1930, p. 79, pl. 36, fig. 1. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 65 

P~atawus nobilis Newberry, 1898, p. 106, pl. 37, fig. 1 [not 

pl. 34; pl. 50, fig. 1, which are Platamts nob.·ilis New-. 

berry]. 

Platan1ts regularis Knowlton, 1917, p. 325, pl. 113, fig. 4. 

Acer· ar·ct·icurn Heer. Lesquereux, 1883, p. 234, pl. 49, figs. 7, 9. 

Acer· graoilens Lesquereux, 1883, p. 234, pl. 49, fig. 7 only. 

Acer· indivisurn Weber. Ward, 1887, p. 66, pl. 29, fig. 5. 

Acer tr·ilobaturn trimtspidaturn (Braun) Heer. Ward, 1887, 

p. 66, pl. 2~. fig. 6. 

Acer· sp. Knowlton, 1930, p. 101, pl. 45, fig. 6. 

Ampelops·is montanensis Cockerell, 1908, p. 103, for Vitis cuspidata Ward, 1887, p. 71, pl. 32, fig. 8 only. · 

Ar·alia coloradens·is Knowlton, 1917, p. 241, pl. 107, fig. 2. 

A.r·alia reesidei Knowlton, 1924, p. 94, pl. 17, fig. 3. 

Aral·ia? serrata Knowlton, 1917, p. 341, pl. 108, fig. 4. 

Oelastnts tam·ir~Jensis Ward, 1887, p. 79, pl. 34, fig. 5 only. 

Oissius gr·ossedentata Knowlton, 1917, p. 340, pl. 104, fig. 1. 

Oiss1tS laevigata Lesquereux, 1883, p. 238, pl. 40, fig. 12 only. 

Oissus parrotiaetoUa Lesquereux, 1878, p. 239, pl. 42, fig. 1 only. 

Fimts tiliaetoUa (Braun) Heer. Ward, 1887, p. 40, pl. 22, fig. 1. 

Grewiopsis platanifoUa Ward, 1887, p. 89, pl. 40, fig. 1. 

Grew·iopsis popuUfoUa Ward, 1887, p. 90, pl. 40, figs. 3-5. 

Gr·ew·iops·is vib1wni!olia Ward, 1887, p. 89, pl. 40, fig. 2. 

Liqu.i(lantbar? C1tcha:ras Knowlton, 1917, p. 320, pl. 91, fig. 6. 

PopttMts balsarnoides exirnia ( Goppert) Lesquereux, 1883, p. 226, 

pl. 46A, fig. 10. 

Pop'ltl1ts nervosa Newberry, 1898, p. 48, pl. 27, figs. 2, 3. 

Popiul·us S1tbr·otmtdata Lesquereux, 1878,. p. 173, pl. 24, figs. 6-8. 

Knowlton, 1930, p. 59, pl. 22, fig. 9. 

Q'lterous negundoides Lesquereux, 1878, p. 161, pl. 21, fig. 2. 

Quer·c'lts platania Heer, 1868, p. 109, pl. 46, fig. 7; 1871, p. 472, 

pl. 46, fig. 5 ; pl. 55, fig. 3c ; 1883, p. 91, pl. 68, fig. 1. 

Lesquereux, 1878, p. 160, pl. 21, fig. 1. 

Rhus? vibttrnoid·es Knowlton, 1917, p. 328, pl. 98, fig. 5. 

Stermtlia berryana Knowlton, 1917, p. 337, pl. 102, figs. 3, 4. 

Ster·cuUa? heterodont a Knowlton, 1930, p. 117, pl. 50, fig. 5 ; 

pl. 51, figs. 4, 8. 

Vib1tr'1Mtrn antiqu1trn (Newberry) Hollick. Berry, 1925, p. 58, 

pl. 17, figs. 1, 2 only ; pl. 18, figs. A, B. 

Vibttr'1Vu.rn lakesi Lesquereux. Knowlton, 1917, p. 348, pl. 110, 

fig. 4. 

Bell, 1949, p. 78, pl. 14, fig. 3 ; pl. 37, fig. 1 ; pl. 58, fig. 1. 

Phyllites pellucidus Knowlton, 1930, p. 130, pl. 39, fig. 2; pl. 56, 

fig. 4. 

The somewhat arbitrary synonymy here proposed 

may seem too inclusive for one species of plane·tree 

or sycamore. However, after observing the great variation shown by leaves from living species of Plataruus, 

I am amazed that paleobotanists have had the temerity 

to describe as many fossil species as they have. Ward 

(1888), Berry (1923, p. 157-164), and MacGinitie 

(1941, p. 126-128) have discussed the frustrating difficulties attending the unraveling of this tangled situation. The synonymy includes the names of many oddshaped and abnormal specimens, as well as readily 

recognizable ones, but with some peculiarities. Comment on all or many of these seems unprofitable at 

this juncture. 

The illustration of the type specimen (Newberry, 

1898, pl. 35) shows too many teeth and the sinuses 

between the teeth as angular, whereas they are typically scalloped. The leaves of P. raynoldsi are in general not strongly lobed, and they resemble the terminal 

leaflets of Cis sus marginata (Lesquereux) Brown so 

closely that separation of the two species, especially 

when the specimens are poorly preserved, is virtually 

impossible. When well preserved and well developed 

however, both the lobed and the unlobed leaves have 

strong, basal, lateral veins that usually branch at wide 

angles from the midrib at a noticeable distance above 

the base of the blade and top of the petiole, but in 

Obssus the corresponding veins are conspicuously de· 

current into the petiole by relatively narrow angles. 

The surface of Platanus leaves also, when well preserved, is distinctly roughened by a minute but prominent quadrangular meshwork pattern. In Oissus this 

feature is subdued or absent. It is not unusual to find 

characteristic, finely corrugated impressions of the inside layer o£ Platanus bark, but so far no definite seed 

balls or seeds have been found in the Paleocene strata. 

However, wood purporting to be that of Platanus was 

collected by Ross Johnson from the Raton formation 

in SEI4 sec. 24, T. 34 S., R. 68 W., near the mouth o£ 

Leon Canyon on the South Fork of Purgatoire River, 

Colo. 

This species doubtless evolved from a Cretaceous 

ancestor in an as yet unresolved complex of numerous 

identified forms. It may have been in the ancestral 

line leading to P. appendiculata Lesquereux, in the 

Eocene of the Sierra Nevada in California, the Cascades of Oregon, and the petrified forests of Yell owstone National Park. 

Occurrence: Fort Union formation (lower), 4032 

(pl. 31, figs. 1, 5, 6), 4404, 4625, 4661, 5609, 6057, 6113, 

6297, 7538, 8199, 8556, 9180 (pl. 66, fig. 8) 9334; (up· 

per), 2414, 2420 (pl. 30, fig. 2, reproduction of Ward's 

type of Grewiopsis viburnifolia/ 3, reproduction o£ 

"\Vard's illustration of Platanus guillelmae), 2424 (pl. 

31, figs. 3, 4, reproduction of Ward's illustration o£ 

Acer trilobatum tricuspidatum), 4369, 4897, 4910 (pl. 

31, fig. 2)' 4974, 4979, 8257 (pl. 30, fig. 4)' 8774, 8888, 

8913, 9109, 9125, 9501, exact locality unknown (pl. 30, 

fig. 1, reproduction of N e'Yberry's type of Platanus raynoldsi) ; Animas formation 644:), 7463; Coalmont formation, 5994, 6004; Dawson arkose, 5835; Denver 

format·ion, 317; Ferris formn;tiou, 6417, 6625, 6971; 

Livingston formation, 8896; ~!fiddle Park formation, 

3H7; Puerco formn:tion, 7371; Raton formation, 5134, 

5141, 5679, 5700, 5701, 5711, 5713, 5715, 5798, 6535. 
66 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Credneria 1 daturaefolia Ward 

Plate 18, figure 14; plate 32, figures 1-5 

Credneria? daturaefolia Ward, 1887, p. '97, pl. 42, fig. 4; pl. 43, 

figs. 1-3; pl. 44, figs. 1-3; pl. 45, figs. 1-3. 

Knowlton, 1919, p. 200. 

Pop1tlus? daturaefolia (Ward) Cockerell. Bell, 1949, p. 511, 

pl. 24, figs. 2-6. 

The identity of these. leaves remains as doubtful 

as when Ward described them. Some or all of those 

identified by Bell as belonging to this species may 

be variants of Platanus raynoldsi Nev{berry. The 

species ranged into the Eocene (pl. 32, fig. 2). 

Occurrence: Fort Union formation (lower), 4975 

(pl. 32, fig. 4); (upper), 2416 (pl. 32, fig. 1, reproduction of Ward's pl. 44, fig. 3), 8164 (pl. 32, fig. 5), 

8205, 8920 (pl. 18, fig. 14). Eocene, 7552, at clay pit, 

1 mile south of Dickinson, N. Dak. (pl. 32, fig. 2). 

LAURACEAE 

Cinnamomum sezannense Watelet 

Plate 66, figures 1-3, 6, 9 

Cinnamom1tm sezannense Watelet, 1866, p. 175, pl. 50, fig. 2. 

Saporta and Marion, 1878, p. 60, pl. 9, figs. 2-6. 

Knowlton, 1930, p. 86, pl. 38, figs. 7 [same as pl. 59, fig. 

2], 8. 

Cinnamomum, affine Lesquereux, 1878, p. 219, pl. 37, figs. 1-4, 7 

[not fig. 5, and probably not the lower left-hand leaf on 

the block with fig. 1, which are Ficus ajJinis ( Lesquereux) Brown]. 

Knowlton, 1922a, p. 145, pl. 8, fig. 4; pl. 17, fig. 6. 

Dorf, 1938, p. 60, pl. 9, figs. 3, 4 .. 

Brown, 1939a, p. 250, pl. 53, figs. 1, 2. 

Cinnamommn d;ubimn Watelet, 1866, p. 176, pl. 50, fig. 4. 

Minnamomum ellipsoideum Saporta and Marion, 1878, p. 61, 

pl. 9, figs. 7-9. 

Cinnamomum tm·mos1tm Watelet, 1866, p. 175, pl. 50, fig. 5. 

Cinnamomum heeri Lesquereux. · Newberry, 1898, p. 100, pl. 17, 

figs. 1-3. 

Berry, 1914, p. 118, pl. 21, fig. 8. 

Cinnamom1tm inaequale Watelet, 1866, p. 174, pl. 50, fig. 1. 

Cinnamomum lanceolatum (Unger) Heer. Ward, 1887, p. 49, 

pl. 24, fig. 2. 

Cinnamomum larteti Watelet, 1866, p. 173, pl. 49, figs. 9-13. 

Cinnamomum middendortensis Berry, 1914, p. 55, pl. 8, fig. 14; 

pl. 9, fig. 1. 

Cinnamomum newberryi Berry, 1911, p. 150, pl. 16, fig. 3; 

1914, p. 54, 117, pl. 9, figs. 12, 13; pl. 21, figs. 9-11; 

1916a, p. 860, pl. 71, fig. 6 ; 1919, p. US, pl. 21, figs. 6-9 ; 

1925, p. 75, pl. 16, fig. 5. 

C. newberryi ellipticum Berry, 1925, p. 77, pl. 16, fig. 7. 

C. n. lanceolatum Berry, 1925, p. 76, pl. 16, fig. 4. 

C. n. minimum Berry, 1925, p. 77, pl. 16, fig. 6. 

Cinnamornn.tm paucinervum Watelet, 1866, p. 176, pl. 50, fig. 3. 

Cinnamomum polymorph1tm (Braun) Heer. Lesquereux, 1878, 

p. 221, pl. 37, fig. 6 [not fig. 10, which is Ficus ajJinis 

( Lesquereux) Brown.] 

Cinnamomum scheuchzeri Heer. Lesquereux, 1878, p. 220, pl. 

37, fig. 8. 

Cinnamomum wardi Knowlton, 1898, p. 69. 

Cinnamomum sp. Knowlton, 1930, p. 87, pl. 39, fig. 1. 

Cinnamomoides buckhami Bell, 1957, p. 491, pl. 35, fig. 1. 

F'imtS dalmatica Ettingshausen. Lesquereux, 1878, p. 199, pl. 

63, figs. 3-5. 

Ficus e1tealyptifolia Knowlton, 1916, p. 340, pl. 87, fig. 2 only. 

Laurus asiminoides Berry. Bell, 1957, p. 50, pl. 36, fig. 4. 

Lnu1·ophyllum insigne Dawson, 1894, p. 61, pl. 7, figs. 24, 25. 

Bell, 1957, p. 51, pl. 26, figs. 1, 2; pl. 37, fig. 1; pl. 65, 

fig. 3 (upper leaf). 

Smilax inq1tirenda Knowlton, 1922a, p. 118, pl. 4, fig. 5. 

In these ovate to elliptic entire leaves the large primary veins normally branch from the n1idrib at an 

appreciable distance above the top of the petiole. The 

few secondaries above the primaries are generally confined to the uppermost part of the leaf. Short branches 

from the primaries along the margin are numerous and 

evenly spaced. Thus, these leaves are much like those 

of Ficus planicostata Lesquereux, Ficus affinis· Lesquereux, and the unlobed leaves of Sassafras thermale 

(Lesquereux) Brown. Distinction between some of 

these closely similar leaves is practically impossible. 

Reference of these leaves to Oinnamomum is more 

than conjectural, for it is now known definitely that 

one or more species of cinnamon were present during 

the Cretaceous and Tertiary of the southern and western United States. Berry (1925, p. 77, pl. 16, fig. 7), 

for example, reported a twig with opposite leaves 

from the Ripley formation (Upper Cretaceous) in 

Tennessee; and in 1955, in company with C. A. Repenning, I had the good fortune to find in the Mesaverde 

formation (Upper Cretaceous) at Black Mountain 

(Mesa) about 40 miles southeast of J{ayenta, Ariz., 

several twigs of a species of Oinna1nomum having 

opposite leaves. One of these is here illustrated (pl. 

55, fig. 5). The identification of these leaves as Ficus 

is not indicated because oppositeness in figs is extremely rare and in the extant species, F. hispida Linnaeus, the opposite leaves are toothed ! On the other 

hand, in most living cinnamons the leaves are opposite and all are entire. No fruits authentically assignable to cinnamon have been found with the Cretaceous and Paleocene leaves here referred to Oinnamomum. 

The long synonymy of varied leaves from strata of 

different ages may be criticized as unrealistic. Does 

the speciation exemplied by W atelet in the Paleocene 

of the Paris Basin and by· Berry in the Upper Cretaceous of Tennessee, inspire confidence in the validity 

of their many species, particularly as these came from 

the same localities~ That specific differences between 

these many species might have been recognized, had 

one seen the living trees, is granted, but until a workable method, not solely based on segregated areas or 

strata, appears for detecting recognizable, constant 

differences in the fossils, it seems more feasible, for 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 67 

descriptive purposes, to regard the specimens as variants of a single species. The species sezanneme is selected as the name bearer because it is the best known 

and is geographicaJly descriptive. 

The relationship of Oi'fllrl.,(lmomwm sezannense Watelet to the ubiquitous 0. polyrnorphum (Braun) Heer 

of Europe, and to 0. dilleri Knowlton in the Eocene 

of the Western United States, is unknown. 

Occurrence : Fort Union formation (lower), 4571 

(fig. 2) , 4 725 (figs. 6, 9), 8652; Dawson arkose, 5835, 

9554; Denver formation, 8426 (fig. 1), 8777 (fig. 3). 

Laurophyllum caudatum (Knowlton) Brown, n. comb. 

Launts? caudata Knowlton, 1917, p. 316, pl. 89, fig. 1. 

La'ztrus socialis Lesquereux. Knowlton, 1917, p. 317, pl. 91, 

fig. 5. 

Andromeda? lanceolata Knowlton, 1917, p. 344, pl. 110, fig. 1 

only. 

Campa eolignitica Berry. Knowlton, 1917, p. 327, pl. 69, fig. 2. 

Oass·ia lancifolia (Lesquereux) Berry. Knowlton, 1917, p. 327, 

pl. 96, fig. 1. 

Magnol·ia ang1tstifolia Newberry, 1883, p. 513. 

Knowlton, 1917, p. 309, pl. 79, fig. 1; pl. 80, figs. 1, 2; 

pl. 81, fig. 1. 

Magnolia attenuata Weber. Lesquereux, 1878, p. 250, pl. 45, 

ftg. 6. 

Magnol·ia la1wifolia Lesquereux. Knowlton, 1917, p. 309, pl. 

106, fig. 2 only. 

Nectandr-a lancitoUa (Lesquereux) Berry. Knowlton, 1917, 

p. 318, pl. 90, fig. 2 only. 

Q1teTC'US? 1·atonensis Knowlton, 1917, p. 298, pl. 69, fig. 6. 

Sapind1ts caudatus Lesquereux. Knowlton, 1917, p. 330, pl. 100, 

fig. 2. 

As the name anguslifolium, which this species should 

take, is preoccupied, caudatwm seems next in order. 

The leaves have attenuate apexes, strongly cuneate 

bases, and somewhat irregular secondary venation, 

thus simulating those of some species of Magnolia. 

The species apparently did not reach the northernmost 

areas of the Paleocene terrain. Inclusion of specimens from Eocene strata of the gulf coast may be 

unjustified. 

Occurrence: Fort Union formation (lower) 8677; 

Evanston formation, 3658; Ferris formation, 8516; 

Raton formation, 5140, 5142, 5690, 5712, 5714, 5798, 

5826, 5830. 

Laurophyllum perseanum Brown, n. sp. 

Plate 34, figures 3, 5 

As·im,ina eocen·ica Lesquereux. Knowlton, 1930, p. 94, pl. 40, 

fig. 4; pl. 43, figs. 1, 2 only. 

Nectand1·a lancifolia (Lesquereux) Berry. Knowlton, 1917, 

p. 318, pl. 90, fig. 1 only. 

Lanceolate leaves with blunt to acute apexes, cuneate bases, entire margins. Petioles 1 to 2 em long. 

. Secondary veins evenly spaced, branching from the 

midrib at about 30° and somewhat decurrent on it. 

Occurrence: Fort Union formation (lower), 3979, 

8567; (upper), 2416 (fig. 5), 4661; Coalmont formation, 5994, 6102; Dawson arkose, 5738; Evanston formation, 3661 (fig. 3) ; Raton formation, 5236, 5826. 

Laurus socialis Lesquereux 

Plate 27, figures 6, 7 

Laurus socialis Lesquereux, 1878, p. 213, pl. 36, figs. 1-4, 7. 

L(tUnts primigen·ia Unger. Heer, 1880, p. 12, pl. 3, figs. 8-13; 

1883, p. 104, pl. 77., figs. 8-13. 

Lesquereux, 1878, p. 214, pl. 36, figs. 5, 6, 8. 

Knowlton, 1924, p. 86. 

La,m·us mtonensis Knowlton, 1917, p. 316, pl. 91, figs. 1-4. 

Be1·chemia mulUnervis (Braun) Heer. Knowlton, 1917, p. 333, 

pl. 101, fig. 5. 

Oass·ia pu1·yea1·ensis Berry. Knowlton, 1924, p. 90, pl. 14, figs. 

1, 2. 

Ficus? smithsoniana Lesquereux, 1878, p. 200, pl. 32, fig. 5. 

Quercus simplex Newberry. Knowlton, 1917, p. 298, pl. 70, 

fig. 3. 

Lesquereux likened the type of this species to the 

living Persea borbonia (Linnaeus) Sprengel, of the 

Southeastern United States, but perhaps the best that 

can be said is that it has a general lauraceous aspect. 

Occurrence: Fort Union formation (lower), 4325 

(fig. 7), 4696 (fig. 6), 4877, 8551, 8567; (upper), 4974, 

8922; Animas formation, 7 496; Coalmont formation, 

5993 ; Dawson arkose, 8307; Evanston formation, 3658, 

3661, 5555; Raton formation, 5134, 5140, 5711, 5826. 

Lindera obtusata (Ward) Brown, n. comb. 

Plate 43, figure 1; plate 66, figure 10 

Diospyros? obtusata Ward, 1887, p. 105, pl. 49, fig. 5. 

D·iospyros? ficoidea Lesquereux. Ward, 1887, p. 105, pl. 49, 

fig. 3 [not fig. 4,. which is Viburnum antiquum Lesquereux]. 

Oh1·ysobalanus coloradens·is Knowlton, 1930, p. 95, pl. 43, figs. 

5, 7 only. 

Oon~us st1tder·i Heer. Knowlton, 1924, p. 94, pl. 13, fig. 2; 

pl. 15, fig. 1. 

Oreodaphne pagosensis Knowlton, 1924, p. 87, pl. 15, fig. 5. 

Leaves elliptic to ovate, relatively broad, with cuneate bases and attenuate apexes. Margin entire. Secondary veins few, branching from the midrib at about 

37°, curving sharply upward near the margin. Second pair of secondaries with conspicuous branches in 

the marginal area, a characteristic of some lauraceous 

genera, particularly Lindera and Persea. 

Occurrence: Fort Union formation, (lower), 5720 

(pl. 43, fig. 1), 6057, 8666; (upper), 2416, 2420, 4323 

(pl. 66, fig. 10), 5618; Animas formation, 5455, 6443; 

Ferris formation, 6971, 8516. 

Persea brossiana Lesquereux 

Pia te 34, figures 6-8 

Persea brossiana Lesquereux, 1874, p. 407. 

Knowlton, 1930, p. 85. 
68 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

La1wus bt·ossiana ( Lesquereux) Lesquereux, 1878, p. 216, pl. 

36, fig. 9. 

001·nus rhamnifolia I .. tsquereux, 1878, p. 244, pl. 42, fig. 6. 

Magnolia angustifoUa Newberry. Knowlton, 1924, p. 88, pl. 16, 

fig. 5. 

Nectandra lanci{olia (Lesquereux) Heer. Knowlton, 1917, 

p. 318, pl. 90, fig. 1 only. 

Rhamnius et·idani Unger. Heer, 1868, p. 123, 153, pl. 19, figs. 

5-7; pl. 49, fig. 10. 

These are long, oval or slightly obovate leaves with 

cuneate bases and abruptly acuminate apexes. The 

evenly spaced secondary veins curve upward near the 

margin to the secondaries above. 

Occurrence: Fort Union formation (lower), 4661 

(fig. 7); (upper) 4582, 5612 (fig. 8); Animas formation, 7 496 (fig. 6) ; Coalmont formation, 5987; Livingston formation, 6767; Middle Park formation, 337; 

Raton formation, 5236, 5826. 

Sassafras thermale (Lesquereux) Brown, n. comb. 

Plate 33, figures 4-7; plate 34, figures 1, 2, 4 

Juglans thermalis Lesquereux, 1878, p. 287, pl. 56, fig. 3 [not 

fig. 4, which is Ficus planicostata Lesquereux.] 

Litsea carbonensis Ward, 1887, p. 48, pl. 24, fig. 1. 

Malapoenna carbonensis (Ward) Knowlton, 1919, p. 378. 

M,ala.poenna praecursoria (Lesquereux) Knowlton, 1919, p. 379. 

Malapoenna sessiUflo'ra (Lesquereux) Knowlton, 1898, p. 142. 

Platanus raynoldsi var. integrifolia Lesquereux. Knowlton, 

1930, p. 78, pl. 34, fig. 2. 

Sassafras ferretianum Massalongo. Heer, 1871, p. 474, pl. 50, 

figs. 1, 2 ; 1883, p. 103, pl. 97, fig. 5. 

Tetranthera praecursoria Lesquereux, 1883, p. 288, pl. 48, fig. 2. 

Tetranthera sessilijlO?'a Lesquereux, 1878, p. 217, pl. 35, fig. 8 

only. 

These leaves display considerable variation in size, 

but internally they are fundamentally similar in having a system of venation in which the first strong pair 

of basal secondaries emerges sharply from the midvein usually at a point 1 em, more or less, above the 

top of the petiole. These secondaries are seldom decurrent along the sides of the midvein into the petiole 

as are the corresponding veins in M elrutomites montanensis Brown. Further, S. therm.ale has a weaker, 

sometimes almost invisible, pair of basal secondaries 

. generally close to the margin, as in nearly all Lauraceae, brit this pair is lacking in 111. montanesis, although the latter may have a faint, irregular vein 

formed from the union of the outer arcs of the 

branches from the primary veins. For some distance 

above the first strong secondaries, the midrib of S. 

therm.ale lacks branches, but at or below the middle of 

the blade secondaries reappear at regularly spaced 

intervals to the apex. All these, except those entering 

lobes, loop near the margin. 

The various objects identified by Lesquereux with 

the leaf, Tetranthera sessiliflora, on the block of his 

plate 35, figure 8, are probably not organically related 

to the leaf. The four-parted involucre on the petiole 

of the leaf is, on close inspection, seen not to be attached to the petiole. Moreover, the basal secondaries 

of the leaf are not accurately depicted, but are opposite one another. Thus, the leaf appears to be a small, 

entire leaf of Sassafras thermale. 

Because the margins of all the latest Cretaceous and 

Paleocene specimens here synonymized are unlobed, 

I lo~g regarded the species as probably referable to 

Oinnamomum, Laurus, Litsea, Ill alapo:.emna, Oreodaphne, Ocotea, or other lauraceous genera. In January 1944, however, I discovered in a collection from 

the Fort Union of the Big Horn Basin, a specimen 

(pl. 33, fig. 4) with two conspicuous lobes well toward 

the apex of the blade. Oddly, these lobes receive the 

first secondaries from near the middle of the blade 

instead of the strong basal secondaries, as is normal 

for the three-lobed leaves. However, this condition, 

though rare, sometimes appears in living leaves (pl. 33, 

fig. 1). I am inclined, therefore, to believe that the 

fossil leaf is an incipient three-lobed leaf of Sassafra.r;;, 

and that ~11 the unlobed leaves in the synonymy are 

to be referred to Sassafras. Lesquereux referred characteristically three-lobed leaves from the Dakota sandstone of Kansas to Lindera venusta and a leaf from 

the same Kansas locality, similar to the lobed specimens from the Fort Union of the Big Horn Basin, to 

Oinnamomum sezanrnense. The association of entire 

and lobed leaves of this kind in strata of different 

ages strengthens my conviction that they represent 

Sassafras or a primitive ancestor of Sassafras or 

Linderra. Berry ( 1902c) discussed the ancestry of 

Sassafras. 

Occurrence: Fort union formation (lower), 3852 

(pl. 33, fig. 6)' 4665 (pl. 33, fig. 4)' 4874 (pl. 33, 

fig. 5)' 4876, 5720, 6057, 6083, 8551 (pl. 33, fig. 7)' 

8928, 9111; (upper), 1502, 5578, 5618, 8887 (pl. 34, 

fig. 1); Coalmont formation, 5594, 6102, 6105; Evanston formation, 3653 (pl. 34, figs. 2, 4); Ferris formation, 6431; Middle Park formation, 336, 8787 . 

NYMPHEACEAE 

Cabomba inermis (Newberry) Hollick 

Plate 35, figures 5, 6 

Psilotu,m inerme Newberry, 1868, p. 38. 

Oabomba inermis (Newberry) Hollick. Newberry, 1898,_ p. 92, 

pl. 22, fig. 2 ; pl. 23, fig. 2. 

Oabomba grcwilis Newberry, 1883, p. 514; 1898, p. 91, pl. 22, 

fig. 1 ; pl. 23, fig. 1. 

Octbomba grandis Newberry, 1883, p. 514. 

Although these specimens suggest Oabomba, no entire, peltate leaves attached to the stems bearing these 

dissected leaves have been found. The foliage may 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 69 

belong to some other aquatic genus, such as Oeratophyllum. 

Occurrence: Fort Union formation, exact locality 

unknown (fig. 6, reproduction of Newberry's pl. 22, 

fig. 1)' 4272, 9072 (fig. 5). 

Nelumbium montanum Brown, n. sp. 

Pia te 35, figures 2-4 

Nelumbo temtifolia (Lesquereux) Knowlton. Berry, 193!'1, 

p. 37, pl. 7, figs. 2, 3. 

N elun,bites p1·otoluteus (Berry) Bell, 1949, p. 64, pl. 63, fig. 5. 

Orbicular leaves sometimes 10 em in diameter. 

Margin entire or slightly undulate. Radial veins 24 

or thereabouts, considerably forked. Fruit receptacle 

3 em or more in diameter. 

The folded condition in which most of these leaves 

are found suggests that in life they were somewhat 

cupped as is the habit of living species of this genus. 

Although assumed by Berry to be the same as those 

formerly called Nelumbo tenuifo.Ua (Lesquereux) 

l{nowlton and N. lakesiana (Lesquereux) Knowlton 

from the Denver Basin,. Colorado, this species differs 

in having twice as many radial veins, which are usually four-forked dichotomously, whereas those of the 

Denver species are few and appear irregularly 

branched rather than dichotomous. Some variation 

in number of veins is to be expected in any species, 

but the wide disparity here seems to indicate more 

than intraspecific variation. Berry's specimens from 

the Ravenscrag formation of southern Saskatchewan 

agree in all respects with N elunt.bian montanwm. fron1 

the Fort Union formation. The identity of these 

leaves with Nelun~bo protolutea Berry, from the 

Eocene of Mississippi, as averred by Bell, is doubtful. 

The fragmentary fruit receptacle found with the 

leaves at locality 8282 was evidently somewhat 

squeezed during fossilization, as the casts of the seed 

cavities are considerably flattened and distorted. 

Occurrence: Fort Union formation (lower), 3980, 

4315,6667,8227 (fig. 4), 8786 (fig. 2), 8897; (upper) 

8262 (fig. 3). 

Nelumbium tenuifolium Lesquereux 

N elmnbi1tm tenuifolium I .. esquereux, 187 4, p. 402 ; 1878, p. 253, 

. pl. 46, fig. 3. 

Nelmnbo temtifoUa (Lesquereux) Knowlton, 1922a, p. 141, 

p. 26, fig. 7 ; 1930, p. 92, pl. 41, fig. 2. 

Dorf, 1942, p. 141, pl. 10, fig. 10. 

Nelu,mbittm lakesianum~ Lesquereux, 1874, p. 403. 

NelU1nb·i1t1n lakesU Lesquereux, 1878, p. 252, pl. 46, figs. 1, 2. 

Nelmnbo lakesiana (Lesquereux) Knowlton, 1917, p. 308; 1924, 

p. 88; 1930, p. 91, pl. 41, fig. 1; pl. 42, fig. 2. 

N elmnbo c1·oss·i Knowlton, 1930, p. 93, pl. 41, fig. 3. 

The leaves of this species have 15, more or less, 

radial veins that are not conspicuously dichotomous 

593121 0 62 - 4 

but give rise to a few curved lateral branches. The 

margin may be entire, slightly undulate, or almost 

lobed. There appears to be a close relationship between this species and Paleonelum,bo nwcroloba Knowlton, and the former may be merely a variant of the 

latter. 

Occurrence: Animas formation, 6309; Denver formation, 317. 

Nymphaea leei (Knowlton) Brown, n. comb. 

Castalia leei Knowlton, 1917, p. 307, pl. 79, fig. 3. 

This fragmentary leaf with cordate base is the only 

one of its kind so far found. As the margin is largely 

missing, its character, whether entire, sinuate, or 

toothed, is unknown. 

Occurrence: Raton formation, 5679. 

Nymphaea pulchella (Knowlton) Brown, n. comb. 

Castalia pulchella Knowlton, 1930, p. 94, pl. 42, fig. 1, a restoration of pl. 57, fig. 3. 

This specimen is thus far the only one of its kind 

known from Paleocene deposits. It has a notched 

base and sinuate margin with moderately large teeth 

or lobes. 

Occurrence: Dawson arkose, 5831. 

Paleonelumbo macroloba Knowlton 

Plate 35, figure 1 

Paleonelumbo rrwc·rololJa Knowlton, 1930, p. 93, pl. 39, fig. 3; 

pl. 42, figs. 3, 4 [.which is the base of pl. 39, fig. 3]. 

The specimen figured here is only half the size of 

the types from the Denver formation but apparently 

agrees in all other essential respects with them. The 

number of radial veins varies from 9 to 11. 

Occurrence: Fort Union formation (lower), 8673 

(fig. 1). 

Paleonuphar hesperium Brown, n. sp. 

Plate 35, figure 7 

Leaves large, 20 em or more long, 15 em wide, 

cordate, with a deep sinus. Margin entire, slightly 

undulate. M:idvein stout. Secondaries widely spaced, 

once or twice forked toward the margin, the ultimate 

branches joining to form intramarginal loops. 

This species resembles Paleonuphar inopina Hollick 

(1930, p. 75, pl. 40, fig. 5) from .the Cretaceous of 

Alaska, in general outline but is larger. The secondary veins of P. inopVn.a fork and loop considerably 

farther in ward from the margin than those of P. hesperium. 

The living red-disked pond-lily, Nuphar rubrodiscum Morong, is comparable to P. hesperium. 

Occurrence: Fort Union formation (lower), 4877, 

7538 (fig. 7). 
70 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Paranymphaea crassifolia (Newberry) Berry 

Plate 36, figures 1-5 

Catalpa cmssifolia Newberry, 1868, p. 56. 

A1"istolochia cmssifoU(~ (Newberry) Cockerell, 1908, p. 90. 

Knowlton, 1919, p. 96. 

Berry,. 1930a, p. 20, pl. 6, figs. 1-3. 

Pamnyntphaea crassifolia (Newberry) Berry, 1935, p. 39, pl. 7, 

figs. 4, 5 ; pl. 9 ; pl. 10. 

Bell, . 1949, p. 68, pl. 50, fig. 3 ; pl. 52, fig. 5. 

Aristolochia cordifolia Newberry, 1898, p. 90, pl. 39; pl. 40, 

fig. 7 ; pl. 60, fig. 4. 

Cm·cis borealis Newberry, 1883, p. 162. 

There is little to add to previous discussions of these 

leaves except to note a feature not heretofore reported, 

namely, that these leaves were minutely hairy, particularly along the veins. This hairiness is best shmvn on 

leaves preserved in fine-grained rocks. 

Several of the leaves described by Heer ( 1868, p. 99, 

pl. 7, figs. 2-4; pl. 50, fig. 9) from Atanekerdluk, 

Greenland, as Populus gaudini Fischer, and that called 

AristoZochia pagei Hollick (1930, p. 74, pl. 40, fig. 3) 

from Alaska, may also belong with this species. 

North of the latitude of mid-vVyoming this species 

appears to be confined to basal Paleocene strata· but ' 

a fragment identified as 11/ enispermites knig htii 

Knowlton by Dorf (1942, p. 143, pl. 10, fig. 11) may 

belong here, and, if so, would indicate that the species 

was already in existence farther south in late Cretaceous time. 

The specimens identified as C occ~dus haydenianus 

Ward by l(now It on in collections made by Hewett 

(1926, p. 28, locs. 6667, 6669), and identified as coming 

from the Lance formation, 30 and 60 feet, respectively, 

below the base of the Fort Union formation as mapped 

by Hewett, were taken, at least at locality 6667, from 

blocks of talus that had dropped. down the slope from 

the Fort Union formation above, as I verified by a 

visit there in 1940. 

Occurrence : Fort. Union formation (lower), 2432 

(fig. 1), 4625, 6154, 6667, 6669, 8517, 8519 (figs. 3, 4), 

8553 (fig. 2), 8567, 8673, 8678, 8780, 8899, 9334, exact 

locality unknown (fig. 5, reproduction of Newberry's 

type of Aristolochia cordifolia, pl. 39). 

CERCIDIPHYLLACEAE 

Cercidiphyllum arcticum (Heer) Brown 

Plate 37, figures 1-24; plate 38, figures 1-17; plate 52, figure 9 

Cercidiphyllmn at·cticnm (Heer) Brown, 1939, p. 492, pl. 53; 

pl. 54, figs. 1-7, 12, 13, 17; pl. 56, figs. 1-6a, 10, 11. 

[Delete from the synonymy Populus newberryi Cockerell 

(for P. accrifolia Newberry), Popul'Lts smilacifolia Newberry, and Ziz-yplws tneelci Lesquereux. 'Vard, fig. 6 

.only.) 

Cercidlph-yllmn elliptimmt (Newberry) Brown, 1939a, p. 491, 

pl. 52, figs. 10, 11, 12, 14-16; pl. 54, figs. 8-11, 14-16. 

[Transfer from the synonymy to that of C. m·cticwn 

the following: Berr-ya mcenwsa Knowlton, Carpites 

lakesi Knowlton, Cercis colomdensis Knowlton, Leg'!lminosites? arachioide.s Lesquereux, Leg1.tminosites amcllioides 1nino1· Berry, Nyssa,? ra,cemosa Knowlton, Piper 

hee1·i Lesquereux, Popuhts arct-ica Heer, P. gl(tnd1tlifern 

Heer, Saba lites? fntet-ifer Lesquereux, and Zizypl11u~ 

hes1Jer·i1ts Knowlton]. 

Dolichites deiztsseni Berry, 1916a, p. 14, pl. 3, fig. 3. 

Jenkinsella a1·ctica (Heer) Bell, 1949, p. 57, pl. 44, fig. 1. 

Leguminosites? borenlis Dawson, 1889, p. 72, pl. 10, fig. 7. 

Orites sp. Chandler, 1926, p. 47, pl. 8, fig. 6. 

Pali1tnts? borealis Heer, 1868, p. 122, pl. 19, fig. 1. 

Popul1ts nebrascensis Newberry, 1898, p. 47, pl. 27, figs. 4, 5. 

Smilax gmndifolia Unger. Lesquereux, 1878, p. 94, pl. 9, fig. 5, 

Trochodendroides a1·ctica, (Heer) Berry. Bell, 1949, p. 56, 

pl. 20, fig. 3; pl. 44, fig. 2; pl. 45, figs. 1, 2; pl. 46. 

figs. 1-3 only. 

Z·izyphoides colombi (Heer) Seward and Conway. Bell 1949, 

p. 73, pl. 50, fig. 1 ; pl. 51, fig. 1 ; pl. 52, figs. 2-4. 

Since my publication on Oercidiphyll!urn in 1939. 

I have collected at many more localities in the lower 

Paleocene and have found that here and there deltoid 

leaf forms appear among the abundant elliptic forms 

generally characteristic of that time. As the emergence and predominance of the deltoid norm was considered indicative of Oerddiphyll1t1n arctimtrn I no~r 

find it dif!icult to justify the assignment of many 

Paleocene Items to 0. ellipticurn (Newberry) Brown. 

Therefore, as a workable arbitrary line must be drawn 

some~vhere, I propose that the name 0. ellipticum· be 

restncted to the elliptic specimens of t-he Upper Cretaceous and that it retain all of the original synonymy 

except the items transferred in the present synonymy. 

Cercidiphyllurn. a.rcticu.rn is characterized by having 

a leaf norm that is deltoid in outline, with a crenate 

margin, but, associated with. examples of this norm 

are variants suggesting the ·norms of C. ellipticurn 

(Newberry) Bro·wn and others in its ancestral line. 

Some of these var~ants, like those on the living 

C. japonicurn Siebo1d and -~ Zuccarini ( Swamy and 

Bailey, 1949, p. '188), are so astonishingly different 

from the norms that the possibility for misidentification of single given leaves is-very great. Some of the 

elliptic leaves resemble those of the living Tetracentron 

sinense Oliver, of western China. 

The fruits and seeds are little if any different from 

those of C. ellipticu.n1 and descendant species. 

No twigs with att.ached .leaves and fruits have yet 

been reported but in the lower Tertiary· strata of the 

London Basin, England, Chandler (1961, p. 70, pl. 6, 

figs. 4-7; p. 84, pl. 8, figs. 18-20; p. 85, pl. 9, figs. 1-5; 

p. 113, pl. 12, figs. 8, 9) found among the typical separate remains two fruits ( p. 298, pl. 30, figs. 1-7, text 

fig. 43) that, fort'unately and as ori()"inally r)ostulatecl . b ' contained the characteristic wino-eel seeds. In her • • .. . 0 

opnnon, however, these fossils do not represent OeJ'-

cidiphyll'l.wn but some genus in the Hamamelidaceae. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 71 

The item, Popul-us "'nebrasoensis Newberry, was 

founded on specimens collected by F. V. Hayden from 

"ferruginous shale, banks of the Yellowstone River." 

(Newberry, 1868, p. 63, 64.) Both the locality and 

matedal from the same collection examined by Newberry is, how~ver, clea-i'ly Oerc'idiphyllun~ arcticu1n 

and not the same as the many specimens from the 

Denver formation called PopuJus nebrascens·is (1\:nowlton, 1930, p. 55-57), which, in the present paper, are 

assigned to A1npelopsis iwerifolia (Newberry) B.rown. 

The item, Populus wibnatta,e Cockerell (1925, p. 3, 

pl. 2, fig. 8), included in my previous synonymy but 

from the Green River formation (Eocene), may indeed be a Populus, but further collections and study 

will be necessary to determine this point. 

The fossil species of Oercidiphyllrum have been compared with the living 0. japonicurr~, of eastern Asia, 

which in Shuihsa. Valley, Hupeh Province, China, is 

associated with 111 etasequoia glyptostroboides Hu and 

Cheng, at 3,000 feet aboii-e sea level, where, according 

to Chu and Cooper ( 1950, p. 267), frosts and snowfall are rare. As 0. arcticum. was associated with M. 

ocoidentalis (Newberry) Chaney in the Rocky Mountains and Great Plains :region during the early Tertiary, the probability seems strong that the latter region also had a mild climate at that time. 

Occurrence: Fort Union formation (lower), 1468, 

3852, 3981 (pl. 52, fig. 9)' 4625, 4661, 4674, 5063, 5437 

(pl. 37, figs. 4, 7, 14), 6131, 6154, 7547, 8188 (pl. 37, 

fig. 9), 8227 (pl. 37, fig. 5), 8517 (pl. 37, figs. 19, 20, 

22-24), 8519· (pl. 37, fig .. 15), 8545,8551,8567 (pl. 37, 

figs. 13, 18), 8673 (pl. :37, fig. 3), 8781, 8884, 8893, 

8896, 8899, 8928, 9130 (pl. 37, figs. 12, 16), 9334 (pl. 

37, figs. 8, 10, 11, 17), 9530; (upper), 2416 (pl. 38, 

fig. 1), 2420,· 4032, 4256 (pl. 38, figs.- 5, 6), 4264 (pl. 

38, figs. 7, 8, 10, 14, 16); 4881, 4892, 4908, 4909, 4974, 

4975, 8167 (pl. 38, fig. 11), 8234 (pl. 38, fig. 2), 8255, 

8556 (pl. 38, fig. 12)' 8774 (pl. 38, fig. 17)' 8885, 8910 

(pl. 38, fig. 3, left column), 8913, 8921, 8922, 9056, 

9072, 9109, 9125 (pl. 38,. figs. 3, 4, 9, 15) ; Coalmont 

formation, 5987, 6107, · 6110; Denver formation, 317 

(pl. 37, fig. 21), 8672 (pl. 37, fig. 6) ; Evanston formation, 3653 (pl. 37, figs. 1, 2), 3661, 5539, 5555; 

Ferris formation, 6415, 6420, 6971; Livingston formation, 333, 337; Poison C~nyon formation, 5121; Raton 

forma6on, 5137, 5140, 5:826. Eocene, 8540 (pl. 38, 

fig. 13). 

MAGNOLIACEAE 

Magnolia berryi (Kn.owlton) Brown, n. comb. 

Plate 39, figure 1 

Jtt{Jlans berryi Knowlton. Berry, 1916a, p. 183. 

Knowlton, 1917, p. 293, :v.L 63, fig. 3; pl. 64, fig. 3; pl. 73, 

fig. 3 ; 1924, p. 80, pl. '6, figs. 5, 6. 

F'agu.s crossi Knowlton, 1924, p. 81, pl. 19, fig. 2. 

These leaves are elliptic to obovate in outline, with 

long petioles, a feature not characteristic of living 

species of Juglans. Further, they do not seem to be 

leaflets of compound leaves, but in form and venation 

they suggest a species of 111 agnolia. 

Occurrence : Animas formation, 5455, 7 496; Raton 

formation, 5679 (fig. 1), 5799, 5826, 6535. 

Magnolia borealis Brown, n. name 

Plate 39, figure 3 ; plate 66, figure 11 

Jtt{Jlans unged Heer. Ward, 1887, p. 33, pl. 14, fig. 6. 

Ficus rnonodon (Lesquereux) Berry. Knowlton, 1924, p. 83, pl. -

10, fig. 1. 

These are broad ovate to elliptic leaves with cuneate or subcordate bases and acute apexes. Margin 

entire. Secondary veins few, evenly spaced, regularly 

curved, and forking well within the margin. 

Occurrence: Fort Union formation (middle), 2420, 

5579 (pl. 66, fig. 11), 8523 (pl. 39, fig. 3) ; Livingston 

formation, 6765. 

Magnolia magnifolia Knowlton 

Plate 39, figure 4 

Magnolin rnagwifolia Knowlton, 1917, p. 311, pl. 84. 

Ma.,qnolia ll:ilgardiana Lesquereux, 1878, p. 249, pl. 44, fig. 4. 

Knowlton, 1917, p. 310, pl. 79, fig. 2 ; pl. 85, fig. i; 1924, 

p. 89, pl. 16, fig. 4. 

MnguoUa lau.rifolia Lesquereux. Knowlton, 1917, p. 309, pl. 85, 

fig. 2. 

MagnoUa leei Knowlton, 1917, p. 313, pl. 64, fig. 2; pl. 65, 

fig. 2 ; pl. 81, fig. 2. 

1v!ngnoUa lesleyana Lesquei:eux, 1878, p. 248, pl. 44, figs. 1-3. 

Knowlton, 1917, p. 313, pl. 82, figs. 1, 2. 

·Magnolia teuiuine1·vis Lesquereux; 1878, p. ·249, pl. 45, fig. 2. 

OhionantlHts 1nembmnacetts Knowlton, 1917, p. 345, pl. 108, 

fig. 2. 

Ji'icu,s (L{Jttila.?· Knowlton, 1917, p. 300, pl. 71, fig. 1. 

Ficus denve1·iana Cockerell. Knowlton, 1917, p. 302, pl. 7fi, 

fig. 1. 

F'·icus puryearcusis Berry. Knowlton, 1930, p. 72, pl. 29, figs. 

1, 2. 

Ficus sp. Knowlton, 1924, p. 84, pl. 7, fig. 3. 

Jttglans? innmnina.ta Knowlton, _1924, p. 80, pl. 7, fig. 2. 

Juglnns schirnpcri Lesquereux. Knowlton, 1917, p. 296, pl. 64, 

fig. 1. 

Juglans sa.pindi[onnis Knowlton, 1917, p. 295, pl. 65, fig. 3. 

Quercus? mtouens·is Knowlton, 1917, p. 298, pl. 69, fig. 7. 

The species here synonymized are based on large 

leaves that appear to be magnolialike in form and 

venation. Many of the illustrations of these fossils 

look much better than the specimens themselves, for 

the latter, as a rule, are very poorly preserved, so that 

more than cursory identification is practically impossible. 

I\:nowlton's name 11! agnolia magnifolia is c.hosen for 

the species. It was instituted for JI;J. tenuinervis Les-
72 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

quereux, one original Paleocene specimen of which is 

preserved in the U.S. National Museum collections 

(Lesquereux, 1878, pl. 45, fig. 2, from South Table 

Mountain, Golden, Colo.). As Lesquereux included a 

mixture of diverse Cretaceous and Tertiary leaves under this. name, Knowlton proposed that the term 

tenuinervis be restricted to Lesquereux's plate 45, figure 4, from the Laramie formation at Golden, Colo. 

Occurrence: Fort Union formation (lower), 3852, 

4665, 4694, 9111; (upper), 4882 (fig. 4), 8196; Animas formation, 5455, 7481, 7496; Denver formation, 

317; Raton formation, 5046, 5133, 5137, 5142, 5684, 

5690, 5697, 5699, 5798, 5826, 5837. 

Magnolia regalis Heer 

Plate 39, figure 2 

Magnolia regalis Heer, 1877, p. 81, pl. 20. 

Knowlton, 1917, p. 314, pl. 87. 

Fimts ag'ltilar Knowlton. Knowlton, 1930, p. 65, pl. 24, fig. 4. 

Fivus 'ltncata Lesquereux, 1878 ,p. 197, pl. 35, fig. 1 only. 

Knowlton, 1917, p. 301, pl. 76, fig. 2. 

The leaf identified by Knowlton as Ficus aquilar 

from the Dawson arkose is apparently not conspecific 

with the type of that species (herein referred to 

11/aqnolia magnifolia Knowlton) from the Raton formation but seems to be only a small specimen of 

Magnolia regalis. Heer's specimen is fragmentary, 

thus leaving the original identification somewhat dubious. Perhaps assignment to Ficus would be more 

appropriate. 

Occurrence : Da w5on arkose, 5839 ; Denver formation, 317 (fig. 2)·; Raton formation, 5465, 5689, 5799. 

Magnolia rotundifolia Newberry 

Magnolia rot'ltnd·ifolia Newberry, 1898, p. 95, pl. 59, fig. 1. 

Oreodaphne? ratonensis Knowlton, 1917, p. 318, pl. 88, fig. 2. 

These leaves have features suggestive of some Lauraceae and Moraceae, but I cannot place them more 

definitely. 

Occurrence : Raton formation, Fisher's Peak, Colo. 

(type specimen) , 5687. 

HAMAMELIDACEAE 

Hamamelites inaequalis (Newberry) Brown, n. comb. 

Plate 40, figures 4-6 

Protojimts inaequalis Newberry, 1883, p. 512; 1898, p. 89, pl. 58, 

fig. 2. 

Hamamelites tothergilloides Saporta. Ward, 1887, p. 64, pl. 

29, fig. 1. 

These leaves resemble those of the witchhazel, 

Hamamelis virginiana Linnaeus, but neither fruits nor 

seeds have been found to confirm the identification. 

They compare well with those called I~. fotherqilloides 

Saporta from Sezanne in the Paris Basin, but differ 

in being somewhat broader with fewer, more widely 

spaced cross veinlets between the secondary veins. 

Occurrence: Fort Union formation (upper), 2416 

(fig. 5), exact locality unknown (fig. 4, reproduction 

of Newberry's pl. 58, fig. 2), 4977 (fig. 6). 

Liquidambar dakotense Brown, n. sp. 

Plate 68, figures 23, 24 

Winged seeds, 7 mm long, the seed itself about equal 

to the wing in length, pointed at the proximal end, 

rounded at the distal end. 

The seeds of Liquida1rubar are much like those of 

the closely related Exbucklandia, but no foliage or 

·fruits surely attributable to either genus have been 

found in association with the Paleocene seeds. Liquidambar, however, is found in the Eocene of the Rocky 

Mountains and Great Plains, and Exbucklo;n.dia in the 

Oligocene ·and Miocene of Oregon and Washington. 

That Liqudambar existed in the Paleocene of North 

America is further suggested by the presence of a Cretaceous species, L. fontanella Brown, from Fontanelle 

Creek northwest of Kemmerer, Wyo. (Brown, 1933, 

p. 8, pl. 2, fig. 3, text fig. 2) . 

Occurrence: Fort Union formation (upper), 9125 

(figs. 23, 24). 

EUCOMMIACEAE 

Eucommia serrata (Newberry) Brown, n. comb. 

Plate 44, figures 1-6; plate 45, figures 1-7 

Alnus sen·ata Newberry, 1868, p. 55; 1898, p. 66, pl. 33, fig. 11. 

Oelastrinites insignis ( Heer) Bell, 1949, p. 71, pl. 36, fig. 4 

[part] ; pl. 53; pl. 54, figs. 1-4; pl. 55, figs. 1, 2; pl. 56, 

figs. 1, 3, 4. 

Oelastrus curvinervis Ward, 1887, p. 82, pl. 36, figs. 3, 4. 

Oelastrus ferrugineus Ward, 1887, p. 78, pl. 34, figs. 1-4. 

Oelastrus grewiopsis Ward, 1887, p. 81, pl. 36, fig. 2. 

Oelastrus montane.nsis Knowlton and Cockerell. Knowlton, 

1919, p. 159, for 0. alnifolius Ward, 1887, p. 80, pl. 35, 

figs. 1, 2. 

Oelastnts pterospei·nwides Ward, 1887, p. 80, pl. 35, figs. 3-6. 

Oelastrus sm-ratizts Knowlton, 1917, p. 329, pl. 98, fig. 3; pl. 99, 

fig. 4 ; pl. 100, fig. 1. 

Oelastrus taurinmisis Ward, 1887, p. 79, pl. 34, fig. 6. 

Oelastrus wa1·dii Knowlton and Cockerell. Knowlton, 1919, 

p. 160, for 0. ovatus Ward, 1887, p. 81, pl. 36, fig. 1. 

Oelast1·us sp. Knowlton, 1917, p. 329, pl. 98, fig. 1. 

Elaeodendron polymorphum Ward, 1887, p. 84, pl. 38, figs. 1-7. 

Elaeodendron SC1'1'ulatum Ward, 1887, p. 83, pl. 37, figs. 3-5. 

Euonyntus splendens Berry. Knowlton, 1917, p. 329, pl. 69, 

fig. 1. 

Euonymus xantholithensis Ward, 1887, p. 82, pl. 37, figs. 1, 2. 

GTewia celast1·oides Ward, 1887, p. 86, pl. 39, fig. 2. 

Grewia pealei Ward, 1887, p. 87, pl. 39, figs. 3-5. 

Ilex 'ins·ignis Heer. Hollick, 1936, p. 131, pl. 73; pl. 74, fig. 1. 

J1tglans nigella Heer. Ward, 1887, p. 33, pl. 15, fig. 1. 

Magnolia elUptica Newberry, 1898, p. 94, pl. 12, fig. 1. 

Monhniopsis ambomefolia Saporta. Ward, 1887, p. 51, pl. 25, 

fig. 2. 

Oreopanax sp. Bell, 1949, p. 76, pl. 59, fig. 3. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 73 

Populus anomala Ward, 1887, p. 23, pl. 8, fig. 7. 

Pozmhus ·inaeqtwUs Ward, 1887, p. 24, pl. 9, fig. 2. 

Quen::n.'l leonis Knowlton, 1930, p. 53, pt. 18, fig. 3. 

Yib11.1'1HUn finale Ward, 1887, p. 115, pl. 57, fig. 5. 

Berry, 1930b, p. 27, pl. 5, fig. 4. 

Although these leaves resemble superficially those 

of some living species of Oela8trus and Prun'IM, they 

compare most strikingly with those of Eucommia ulmoides Oliver, a dioecious, monotypic tree of Central 

China. The characteristic fruits of Eucommia, although present in some American Eocene and Oligocene floras (Brown, 1940, p. 349), have, however, not 

yet been reported from the Paleocene. The fact that 

they occur in slightly younger strata in the Southern 

and 'Vestern States suggests strongly that a concentl·ated search for Paleocene examples could be successful. The presence of Eucommia in the Paleocene 

flora of America is not surprising in view of the fact 

that this flora also contains representatives of other 

Jiving Asiatic congeners: Ginkgo, Glyptostrobus, 

11/ etaseq1wia, Pterocarya, Zelkova, Oercidiphyllum, 

K oelreuteria, etc. 

The species ranged from New Mexico to Canada and 

probably Alaska during the Paleocene. It may have 

arisen from the Cretaceous species Oela8trinites alatus 

knowlton found in the Laramie formation of the 

Denver Basin. The leaves of the latter average somewhat sma1ler in size, with more numerous and more 

closely spaced secondary veins that emerge from the 

midrib at a smaller angle and are more conspicuously 

directed toward the apex of the blade. 

Occurrence: Fort Union formation (lower), 2421, 

(pl. 44, fig. 1)' 2423 (pl. 44, fig. 4)' 4005, 4565, 4617, 

4625, 4626 (pl. 44, fig. 5)' 4699, 5388, 5389, 5885, 5889 

(pl. 44, fig. 2; pl. 45, fig. 4)' 6598, 7004, 8190, 8227, 

8239, 8240, 8249, 8253, 8258, 8512, 8530, 8547, 8550, 

8552 (pl. 45, fig. 3), 8568, 8677, 9193, 9301; (upper), 

2416, 2420 (pl. 44, fig. 3; pl. 45, figs. 2, 5-7), 2422 

(pl. 44, fig. 6), exact locality unknown (pl. 45, fig. 1, 

reproduction from Newberry, 1898, pl. 12, fig. 1), 4369, 

4661, 4908, 5480, 5595, 5760, 6156, 6161, 7685, 8206, 

8224, 8888, 9239 ; Coalmont formation, 5994; Dawson 

arkose, 8881; Ferris formation, 6971; Raton for-?Iation, 5140, 5151, 5712, 5798; Ravenscrag formatwn, 

Alberta and Saskatchewan. 

SAXIFRAGACEAE 

Hydrangea antica Brown, n. sp. 

Plate 41, figures 3, 5 

Leaves lanceolate with rounded-cuneate bases and 

acute apexes. Margins .with low relatively few serrate teeth. Secondary veins widely spaced, camptodrome with short branches into the teeth. Connecting veinlets diagonal, parallel. Petiole 1 em long. 

No calyces identifiable as those of Hydrangea have 

yet been found in the Paleocene strata of the Rocky 

Mountains and Great Plains, but authentic leaves and 

calyces have been taken from Eocene and later strata 

in Alaska, California, Colorado, Oregon, and Washington. · 

Occurrence: Fort Union formation (lower), 4661 

(fig. 5) ; Middle Park formation, 337 (fig. 3). 

ANNONACEAE 

Asimina vesperalis Brown, n. sp. 

Plate 46, figure 4 

Leaves obovate, with entire margins and cuneate 

bases. Secondary veins widely spaced, slightly decurrent on the midrib and forming loops near the 

margin. Petiole short, thick. 

No seeds characteristic of pawpaw have been found 

in the strata containing these leaves. The assignment 

to Asimina, therefore, is tentative. Some species of 

111 agnolia have leaves that match these fairly closely. 

Occurrence: Fort Union formation (upper), 8774 

(fig. 4). 

ROSACEAE 

Cercocarpus ravenscragensis Berry 

Ce1'COCa1'P'U8 1'at'ensc1·ar;ensis Berry, 1930, p. 23, pl. 5, fig. n. 

This leaf, the only one of its kind so far seen in the 

Paleocene strata of the Rocky Mountains and Great 

· Plains, purports to be a species of 0 ercocarp'IM. 

Occurrence: Ravenscrag formation, SE:L4 sec. 22, T. 

7, R. 22 W., third meridian, on north branch of 

Frenchman River, Saskatchewan, Canada. 

Prunus careyhurstia Brown, n. sp. 

Plate 42, figures 2, 8 

Leaves lanceolate elliptic, but shape of the base and 

apex not definitely known. Margin finely dentate to 

serrate with scalloped to angular sinuses between the 

sharp teeth. Pinnate secondary veins strongly curved 

distad, sometimes branched near the margin. Connecting veinlets delicate, closely spaced, almost horizontal when leaf is viewed in upright position. 

Occurrence: Fort Union formation (lower), 8666 

(fig. 2), 8552 (fig. 8). 

Prunus coloradensis Knowlton 

Plate 42, figures 3, 4, 9 

Prunus coloradensis Knowlton, 1917, p. 326, pl. 96, fig. 2. 

Prunu.s denveren.<;is Knowlton, 1930, p. 97, pl. 44, figs. 3, 4, 6, 11. 

Juglan.s acumtin(t.ta Brown. Knowlton, 1917, p. 292, pl. 65, fig. 1. 

Lattnts prim.ir;enia. Unger. Knowlton, 1930, p. 83, pl. 38, figs. 

5, 6 [not fig. 3, which is Latu;u8 socia.l·is Lesquereux]. 

These leaves lack the acropetiolar glands usually 

but not always present on the deciduous, serrate leaves 
74 PALEOCENE FLORA OF THE ROCKY MOUNTAINS ANP GREAT PLAINS 

of Prunus. Their resemblance to the serrate leaves of 

other genera, particularly Oarya and Saliw, emphasizes caution in accepting them as Prunus. The secondaries of this species are less numerous, more widely 

spaced and irregular than those of Prunus careyhur8tia Brown. The intersecondaries, however, are 

more numerous and prominent than in the latter. The 

leaves identified by l{nowlton as Laurus primigenia 

are not entire, as figured, but have minutely serrate 

margins. 

Occurrence: Dawson arkose, 5738 (fig. 3, 4), 5836 

(fig. 9), 5838; Raton formation, 5714, 5799. 

Prun us corrugis Brown, n. sp. 

Plate 67, figures 1-4, 8 

Impressions of nuts, broadly elliptic, rounded at 

one end and rounded to blunt pointed at the other. 

Length, 1.5 to 4 em, width to 2.8 em, diameter about 

1 em. Outer surface slightly concave, pitted and irregularly furrowed; inner surface slightly convex, 

smooth or reflecting the features of the outer surface, 

with a smooth flange about 1 mm wide. 

Although in some respects these fossils suggest walnuts, their affinity seems more plainly to be with the 

Amygdalus or almond section of Prunus. ·They are 

somewhat more elliptic in outline than the specimen 

from the Eocene of Tennessee called Amygdalus wilcowiana Berry ( 1930a, p. 72, pl. 12, fig. 1). Just what 

Paleocene foliage belonged with this species is a moot 

question. 

Occurrence: Fort Union formation (lower), 541 

(fig. 8), 4984 (figs. 2, 3), 8660, 8668, 9540; (upper), 

9"132; Almy formation, 9235 ; Animas formation, 4050 

(fig. 4), 5460, 9492 (fig. 1); Coalmont formation, 6005; 

Ferris formation, 9209. 

Prun us mclearni Berry 

Pntnus nwleanti Berry, 1935, p. 41, pl. 11, figs. 1, 2. 

These lanceolate, acuminate leaves resemble those of 

Prunus carey hurstia in venational and marginal features, but are more lanceolate and long acuminate. 

Occurrence: Ravenscrag formation, sec. 29, T. 3, R. 

24 W., second meridian, Saskatchewan, Canada. 

Prunus perita Brown, n. sp. 

Plate 43, figures 2, 3 

Leaves 15 em long, 4.5 em wide, elliptic, narrowed 

at base and apex, which is somewhat attenuate. · Margin serrate, with numerous low, blunt regular teeth, 

separated by angular sinuses. Venation pinnate 

though obscure, the leaves apparently having had a 

thick texture. Secondaries evenly spaced, forming 

loops well within the margin. Some intersecondaries 

present. 

The features of these leaves suggest those of the 

evergreen section of Prunus that includes the laurelcherries. However, I find no clear evidence of glands 

within the blade near the midrib, as often seen in 

living species and in the Oligocene specimens from 

Oregon called· PTunus prist-ina Brown ( 1950, p. 323, 

text figs. 1-3). The relationship of P. perita to the 

species described by Heer (1880, p. 16, pl. 6, figs. 2, 3) 

as P. scotti is problematical. 

Occurrence: Fort Union formation (middle), 8552 

(figs. 2, 3). 

LEGUMINOSAE 

Ba uhinia wyomingana Brown 

Plate 43, figure 10 

Ba1thinia wymningnna Brown, 1956, p. 104, text fig. 1. 

The deep sinus and bilobed outline of this leaf, together with the venational details, strongly suggest 

the assignment to B auhinia. Since this species was 

published, large pods have been found in the same 

collection that yielded the leaf, but whether or not the 

pods, Leguminosites coloradensis Knowlton, belong 

with the 1 eaf is uncertain. 

Occurrence: Fort Union formation (lower), 4877 

(fig. 10). 

Leguminosites coloradensis Knowlton 

Plate 68, figures 1, 3-G 

ljegmninosites? colontdensis Knowlton, 1922a, p. 147, pl. 19, 

fig. 9. 

Impressions of the interiors of large capsules which 

were attached by broad bases to a stout petiole. Valves 

with broad flanges, blunt pointed at the apex and 

rounded at the base. Some specimens show cross 

wrinkles, folds, or striations. 

The type specimen, described by Knowlton from the 

Laramie formation at Marshall, Colo., is only the apical end of what evidently was a large pod. That these 

pods are leguminous and belong with some one of the 

described Paleocene legumes, or to some other family 

of plants such as the Bignoniaceae, is uncertain. 

Occurrence: Fort Union formation (lower) 3852, 

4877 (figs. 1, 6), 5526 (fig. 3), 5918, 8551 (fig. 4), 

8884, 9130, 9221, 9566; Middle Park formation, 336 

(fig. 5), 337. 

Leguminosites williamsi Berry 

Lc.r;n1ninosites williamsi Berry, 1930b, p. 23, pl. G, fig. 1. 

No further specimens assi~p1able to this species have 

been found, and no new light has been shed on its 

generic identity. 

Occurrence: Ravenscrag formation, SE1Jt sec. 22, 

T. 7, R. 22, vV., third meridian, on north branch of 

Frenchman River, Saskatch~wan, Canada. 
SYSTEMATIC DESCRiPTION OF THE MEGASCOPIC FLORA 75 

Mimosites coloradensis Knowlton 

Plate 43, figures 12, 13 

.Mimosites coloradens'is Knowlton, 1923, p. 166, pl. 40, figs. l-3. 

Brown, 1934, p. 59. 

As only two specimens were collected from the Coalmont· formation, near Coalmont, Colo., and as these 

can be matched by specimens from the large collections of 11/imosites coloradensis Knowlton from the 

Green River formation, it would seem that they should 

be assigned to this species. 

Occurrence: Coalmont formation, 6440 (figs. 12, 13). 

Robinia wardi (Knowlton) Brown, n. comb. 

Plate 43, figures 4-6 

Acacia wa1·di Knowlton, 1899, p. 730, pl. 98, fig. 7. 

Acacia lanwrens·is Knowlton, 1899, p. 730, pl. 98, fig. 6. 

Acac·ia 1nacrospennn Knowlton, 1899, p. 729, pl. 98, fig. 8. 

Robinia, nwso.zo·ica Cockerell, 1912, p. 32, text fig. 1. 

Of the pods described by Knowlton as Acacia from 

the Eocene of Yellowstone National Park, that called 

A. ttvardi is represented by the best specimen. Although the specimens here assigned to Robinia wardi 

differ somewhat in size and shape, they can be matched 

readily in every respect by pods from the living black 

locust, R. pseudo acacia Linnaeus. The leaves described by l(nowlton as Leguminosites lamarensis and 

L. lesquereuwiana from the same Yellowstone strata 

may be leguminous, but they are too fragmentary for 

satisfactory identification and definite assignment to 

Robinia wardi. Likely leaves that might belong to 

this species are scarce at Paleocene localities, but the 

fragments here illustrated (pl. 43, figs. 9, 11) may be 

such. · 

The specimens described by Cockerell as Robinia 

mesozoica and here refigured (fig. 5) are parts of two 

pods, slightly different in size, one representing the 

proximal and the other the distal half of the pods. 

The proximal half ends in an abruptly tapered symmetrical base somewhat extended to the point where· 

it was attached to the peduncle. The distal half ends 

more bluntly except that the wide flange running along 

the upper or placental margin of the pod curves downward at the end to make a sharp projection about on 

a median line length wise through the pod. In the 

pods of living redbuds, Oercis, with which those of 

fossil Robirn'ia might be confused, the upper margin 

runs almost straight into the apical tip, with little 

downward curvature, thus giving them the appearance of the blades of carving knives. The lower margin of the pods has a narrower flange. 

CockereU's remark that in his pods "the seeds are 

placed very obliquely" and that this distinguishes 

them from those described by l(nowlton from Yellowstone National Park, is not strictly correct. In all 

the pods here synonymized with Robinia wardi, the 

position of the seeds, so far as can be detected from 

their impressions, is only slightly oblique, but, nevertheless; normal for Robinia. 

Cockerell assumed that his specimen came from 

strata of Laramie age, whence the specific term me'8ozoica/ but as leaves of Platanus raynoldsi Newberry 

are on the same block and as the containing strata are 

similar to the Coalmont formation in North Park and 

the ~tfiddle Park formation in Middle Park, the age 

is Paleocene. 

Occurrence : Fort Union formation (upper), 9109 

(figs. 4, 6) ; Coalmont or Middle Park formation, 9445 

(fig. 5, Specimen in University of Colorado Museum); 

Eocene of Yellowstone National Park. 

STAPHYLEACEAE 

Staphylea minutidens (Knowlton) Brown, n. comb. 

Juglans m'inutidens Knowlton, 1917, p. 293, pl. 55, fig. 3. 

J11glans nigella. Heer. Knowlton, 1917, p. 292, v. ;'"iiJ, fig. 2; 

pl. 63, fig. 2. 

The long petioles of these leaflets are not harmonious 

with an assignment to Juglans, whose leaflets are generally sessile or nearly so. 

Occurrence: Raton formation, 5686, 5714, 5799. 

ACERACEAE 

Acer fragile Knowlton 

Ace1· frngilis Knowlton, 1917, p. 330, pl. 101, figs. 1, 2. 

Q1tercus jisher'ictna Knowlton, 1917, p. 297, pl. 68, figs. 3, 4. 

These fragmentary specimens, as stated by 1\:nowlton, are at least superficially aceroi~ in appearance. 

That they represent a species of Acer may be doubted. 

The reader after inspecting I\:nowlton's figures of 

Acer fragilis (pl. 101, figs. 1, 2) 1nay question the 

relationship of the two specimens. Figure 2, unfortunately, was retouched, so that it does not show the 

well-defined rounded sinus between two lobes in the 

uppermost portion of the blade to the left of the midrib. All the specimens here synonymized are from 

the same locality. 

Occurrence: Raton formation, 5099. 

Acer newberryi Brown, n. name 

Plate 46, figures 2, 3, G, 8 

Negundo t'riloba Newberry, 1898, p. 115, pl. 31, fig. 5 . 

.tiralia trilobn Newberry, 1898, p. 123, pl. 40, figs. 4, G. 

The specimens here assigned to a sp~cies of boxeldermaple are fragmentary and are not accompanied by 

appropriate samaras. As an earlier. described fossil 

maple is called trilobatun1 I give the present specimens a new name. 
76 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Occurrence: Fort Union formation (upper), 2420 

(figs. 2, 6, 8, reproductions from Newberry, 1898, pl. 

31, fig. 5; pl. 40, figs. 4, 5), 4871, 8910 (fig. 3). 

Acer silberlingi Brown, n. sp. 

Plate 46, figure 7 

Leaves palmately three~lobed. One specimen shows 

two pairs of primary veins indicating that there may 

have been an additional pair of existing or incipient 

basal lobes, but the margin of the specimen is broken 

so that this assumption cannot be ver1fied without 

further material. The lobes are attenuate pointed with 

margins having large, coarse unequal teeth. No 

samaras were found with these leaves. 

Without samaras it seems futile to suggest the modern affinities of these leaves. There may be some relationship between this species and Acer disputabilis 

Hollick (1936, p. 134, pl. 74, fig. 4) from the Tertiary 

of Alaska. No undoubted leaves and fruits of Acer 

are known from the Paleocene deposits of Greenland, 

but Heer described such remains from other areas and 

ages in the Arctic regions. 

The specific term honors A. C. Silberling of Harlowton, Mont., whose labors as a collector of fossil 

mammals in the Crazy Mountain region contributed 

greatly to knowledge of Paleocene vertebrates. In 

1936 he guided me to the plant locality where Acer 

silberlitngi was found. 

Occurrence: Fort Union formation (lower), 8567 

(fig. 7). 

Acer sp. 

Plate 46, figure 5 

Samara 3 em or more in length, with greatest width 

of the wing 7 mm. Only half of the seed portion is 

present, the point of att~chment not being preserved. 

The only maple leaves found at the locality that supplied this fruit are those assigned to a boxelder type, 

Acer newberryi, but the fruit may not belong to that 

species. 

Occurrence: Fort Union formation (upper) 8910 

(fig. 5)' 9532. 

Acer sp. 

Plate 46, figures 9, 10 

Samaras up to 5 em in length, with large elliptical 

seed heads 1.3 em long. Wing slightly constricted just 

behind the seed head. In life the twin samaras diverged at an angle of about 160°. No leaves were 

found with these seeds and no modern affinities are 

therefore suggested. 

Occurrence: Fort Union formation, 9109 (figs. 9, 10). 

SAPINDACEAE 

Sapindus a:ftinis (Newberry) Brown, n. comb. 

Plate 47, figures 1-8 

Sapindus ajfinis Newberry, 1868, p. 51; 1898, p. 116, pl. 30, 

fig. 1 ; pl. 40, fig. 2. . 

Ward, 1887, p. 67, pl. 30, figs. 1, 2. 

Sapindtts alattls Ward, 1887, p. 68, pl. 31, fig. 3 only. 

Sapind'lls angustifolius Lesguereux. 'Vard, 1887, p. 68, pl. 31, 

figs. 5-7. 

Sapindtts berryanws Knowlton, 1930, p. 101, pl. 46, figs. 1, 2. 

Sap·indus glendivensis Knowlton, 1919, p. 579. 

Sapindus gmndifoliolus Ward, 1887, p. 67, pl. 30, figs. 3-5; 

pl. 31, figs. 1, 2. 

Sapindizts? membranacet_ts Newberry, 1878, p. 117, pl. 30, fig. 2 

only. 

Sapindus obtusitolius Lesquereux, 1878, p. 266, pl. 49, figs. 8-11; 

1883, p. 235, pl. 48, figs. 5-7. 

Launts resu'rgens Saporta. Ward, 1887, p. 47, pl. 23, fig. 7. 

The large amount of variable n1aterial found at 

each locality yielding these leaflets makes the distinction of more than one species difficult or impossible. 

On such fairly complete compound leaves as have been 

found, the leaflets are alternate, opposite, or mixed. 

Despite all appearances it is still uncertain that this 

foliage represents Sapindus. Suggestive comparisons 

can be made with Bursera, Phellodendron, Rhus, and 

.LYanthoxyl!um. 

Occurrence: Fort Union formation (lower), 3728, 

4315, 8517, 8519, 8523 (figs. 2, 5), 8654, 8678 (figs. 

7, 8); (upper), 1502,2416 (figs. 4, 6), 2422,5578,8206, 

8920, 8921, 9180, exact locality unknown (figs. 1, 3, 

reproduction from Newberry, 1898, pl. 30, figs. 2, 1) ; 

Dawson arkose, 5839, 8188. 

Koelreu teria annosa Brown 

Plate 59, figures 8, 9 

Koel?'euteria annosa Brown, 1956, p. 107, text fig. 3. 

The leaflet (fig. 8) here assigned to this species was 

not found with the capsule (fig. 9), but at a locality 

on the Little Powder River, near Biddle, Mont. It, 

however, seems to be an authentic K oelreuteria leaflet. 

Occurrence: Fort Union formation (lower), 9253 

(fig. 8); (upper), 9344 (fig. 9). 

ILICACEAE 

Ilex artocarpidioides (Lesquereux) Brown, n. comb. 

Celastrinites artocarpidioides Lesquereux, 1878, p. 268, pl. 35, 

fig. 3. 

Celast1"inites popttUfol-ius Knowlton, 1930, p. 100, pl. 40, fig. 1 ; 

pl. 45, fig. 11. 

Celast1·tts gwud·in·i Lesquereux, 1888, p. 54. 

Knowlton, 1930, p. 99, pl. 45, fig. 3 ; pl. 45, figs. 3, 4 [not 

fig. 9, which is G-isstt8 rnarginata (Lesquereux) Brown]. 

flex? ovata Knowlton, 1930, p. 102, pl. 46, fig. 4. 

Q'uerctts celast1·ifol-ia Lesquereux. Knowlton, 1930, p. 54, pl. 

18, fig. 5. 

Quercus Zeonis Knowlton, 1930, p. 53, pl. 18, fig. 3 only. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 77 

These leaves resemble the smaller leaves of Euc01n1nia Se'n·ata (Ne,Yberry) Brown, but differ in having 

conspicuously cuneate bases and more open secondary 

venation which, in the better preserved specimens, has 

connecting loops between the secondaries well within 

the margin. 

Occurrence: Denver formation, 317, 318. 

RHAMNACEAE 

Paliurus 1 sp. Berry 

PaU'Itnts? sp. Berry, 1930a, p. 25, pl. 5, fig. 7. 

This specimen was described as a disklike fruit resembling that of some living species of Paliurus. If 

correctly· identified it is the earliest good evidence of 

Paliurus in the Tertiary strata of the Rocky Mountains and Great Plains. 

Occurrence: Ravenscrag formation (part of which 

was formerly called Estevan on the Souris River) in 

S""\Vl!.J, sec. 6, T. 8, R. 3, ""\V., fourth meridian, Alberta, 

Canada. 

Rhamnus cleburni Lesquereux 

Plate 42, figures 1, 5-7 

. Rhmnnu.s cleb'ltrni Lesquereux, 1878, p. 280, pl. 53, figs. 1-3. 

i:<nowlton, 1930, p. i04, pl. 40, fig. 6 only. 

Rha1nmts? d·iscolm· Lesquereux, 1878, p. 280, pl. 52, fig. 17. 

Rhantmts inaeq·uaUs Lesquereux, 1878, p. 279, pl. 52, fig. 16. 

. Cm·nus denverensis Knowlton, 1930, p. 119, pl. 13, fig. 1. 

Co1·1ws imp1·essa Lesquereux. Knowlton, 1930, p. 118, pl. 51, 

fig. 1 only. 

Cm·nus lakes'ii Knowlton, 1930, p. 119, pl. 50, fig. 1. 

Conws neomexicana Knowlton, 1917, p. 342, pl. 109, fig. 1. 

ConLtts stttde1··i Heer. Lesquereux, 1878, p. 244, pl. 42, fig. 5 

"[not fig. 4, which is Rhamnus goldiana Lesquereux]. 

Ward, 1887, p. 55, pl. 26, fig. 1. 

JJ'·icu.s coloradensis Cockerell, 1910, p. 223. 

Knowlton, 1922, p. 134, pl. 22, fig. 1. 

Ji'ic·us c·rossU Ward, 1887, p. 39, pl. 21, fig. 2. 

Dorf, 1938, p. 56, pl. 7, fig. 3. 

FiottS denveria.na Cockerell, 1910, p. 224. 

Knowlton, 1930, p. 64, pl. 24, figs. 2, 3 only. 

F'·i()zts ·i1-regula1·is Lesquereux, 1878, p. 196, pl. 34, figs. 4-7 ; 

pl. 63, fig. 9. 

Ward, 1887, p. 38, pl. 20, fig. 4 only. 

F'·iotts 1·1wmnoides Knowlton, 1916, p. 339, pl. 87, fig. 3. 

li'ic·ns spectabiUs Lesquereux, 1878, p. 199, pl. 33, figs. 4-6. 

Ward, 1887, p. 38, pl. 21, fig. 1. 

Ji'·ictts 1mcata Lesquereux, 1878, p. 197, pl. 35, fig. 2 only. 

Pomtltts 'Ungeri Lesquereux, 1878, p. 175, pl. 24, fig. 5. 

Lesquereux's types of this species, like those of R. 

goldiana, came from the Denver formation that crops 

out on the slopes of South Table Mountain near 

Golden, Colo. Except that they have a cuneate base, 

the leaves of R. cleburni are much like those of R. 

goldiana, and it is probable that both kinds may have 

come from. the same species of tree or shrub. For the 

present, however, I shall regard them as having originated from distinct species. That these were in fact 

species of Rhamnus is uncertain. The leaves of R. 

cleburni have some of the aspects of those borne by 

living species of Cornaceae, and particularly of 

Oamptotheca acumin.ata Decaisne, of western China. 

Included in the foregoing synonymy are some items 

representing Upper Cretaceous specimens. Dorf's 

synonymy ( 1938, p. 67; 1942, as Dillenites cleburni, 

p. 146) included other Upper Cretaceous items such 

as R. rectinervis Heer, R. brittoni Knowlton, and 

R. marshallensis ·Knowlton. These three, however, in 

my opinion, after comparison of the types with those 

of R. salicifolia Lesquereux, also from the Upper 

Cretaceous, should be referred to the latter species, 

for some of the specimens of all these species have 

definitely toothed margins. Thus far I have seen no 

leaf, authentically comparable to the types of R. cleburni, with a toothed margin. 

Occurrence: Fort Union formation (lower) 8901; 

(upper), 5863; Denver formation, 317 (figs. 5, 7), 

6145, 8672 (figs. 1, 6), 8777; Dawson arkose, 5838; 

Ferris formation, 6431; Raton formation, 5464, 5687, 

5826. 

Rhamnus goldiana Lesquereux 

Plate 48, figures 2, 5-7 

Rhamnus gold·ianus Lesquereux, 1878, p. 281, pl. 53, figs. 4-8. 

Knowlton, 1924, p. 91 . 

Rham?mts clebttrni Lesquereux. Knowlton, 1924, p. 90, pl. 17, 

fig. 4. 

Berchemia multinervis (Braun) Heer. Lesquereux, 1878, p. 277, 

pl. 52, figs. 9, 10. 

Ward, 1887, p. 73, pl. 33, fig. 2 [not fig. 1, which is Juglans 

taurina Brown]. 

Cornus studeri Heer. Lesquereux, 1878, p. 244, pl. 42, fig. 4 

[not fig. 5, which is Rhamnus cleburni Lesquereux]. 

F'icus irregularis Lesquereux. Ward, 1887, p. 38, pl. 20, fig. 5. 

The distinguishing ·feature of these leaves is the 

conspicuous cordate base: Of perhaps minor importance is the presence of short branches from . the first 

secondary veins above the base. In other respects, 

particularly the spacing, delicacy, and regularity of 

the veinlets connecting the secondaries, the leaves of 

R. gold·iana and R. cleburni are closely comparable. 

Living species of Rhamnus may have leaves with entire margins, but in general the margins are serrate or 

serrulate. No species, however, so far as I am aware, 

has leaves with strongly cordate bases and incipient 

palmate venation. 

Occurrence: Animas formation, 7481, 7496; Denver 

formation, 317 (fig. 5), 877 4 (figs. 2, 6, in Chicago 

Nat. History Mus., 7) ; Raton formation, 5291. 

Rhamnus hirsuta Brown, n. sp. 

Plate 49, figures 1-10 

Leaves elliptic in outline, long and narrow, or 

short and wider through the middle. Base cuneate, 
78 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

apex blunt pointed. Petiole short. Margin entire to 

coarsely serrate. : Surfaces rough hairy. Venation 

pinnate .with evenly spaced curved secondaries that 

may or may not be :forked near the margin. 

There is some ,doubt that these leaves represent 

Rhamn'lt8. However, they resemble in many respects 

those called Rhan?Jnus ma.rginatus var. apiculatus Berry 

( 1922, p. 15, pl. 1~, figs. 2, 3) from the Wilcox group 

in Louisiana, except that the latter are described as 

having entire margins and attenuate apexes. 

Occurrence: Fort Union formation (lower), 8519 

(figs. 1-10), 8677; 8897. 

Zizyphus :fi]?rillosus (Lesquereux) Lesquereux 

Plate 50, figures 5, 6, 10, 1l 

Oeanothus fibrillostts Lesquereux, 1873, p. 381. 

Zizyphtts fibrillostts (Lesquereux) Lesquereux, 1878, p. 276, 

pl. 52, figs. 1--'6. 

Knowlton, 1917,. p. 335, pl. 102, fig. 1; 1924, p. 91; 1930, 

p. 108, pl. 46, figs. 6, 9. 

Zizyphus beckwithii Lesquereux, 1883, p. 125, pl. 19, fig. 5. 

Knowlton, 1930, p. 110, pl. 47, fig. 3. 

Zizyphtts corrugattts Knowlton, 1922, p. 158, pl. 17, fig. 3. 

Zizyphtts daphnogenoides Knowlton, 1924, p. 91 ; 1930, p. 109, 

pl. 48, figs. 5; 6. 

Zizyphius d·istorttts Lesquereux, 1878, p. 275, pl. 51, fig. 7 only. 

Knowlton, 1930, p. 110, pl. 47, fig. 2; pl. 52, fig. 1. 

Zizyphus hendersoni Knowlton, 1922a, p. 157, pl. 15, figs. 1, 2. 

Fimts pseudopomtl1tf: Lesquereux. Knowlton, 1924, p. 83, pl. 7, 

fig. 4 ; 1930, P.; 66, pl. 26, fig. 3. 

Ficus wardi Knowlton, 1917, p. 266, pl. 42, fig. 2. 

PaUurus z·izyphoide~ Knowlton, 1917, p. 334, pl. 104, fig. 2. 

Phyllites trilUoides Dorf, 1938, p. 78, pl. 17, fig. 3. 

So :far as I can see, the unusually fine specimens I 

:found in 1939 in the upper strata of the Vermejo 

:formation (Upp_er Cretaceous) on the south side of 

the Cuchara Riyer, 1.2 miles west of Walsenburg, 

Colo., cannot be ·distinguishe¢1 :from those called Zizyphus fibrillosus Lesquereux, occurring in the Denver 

:formation (Pale<;>cene). Inasmuch as the Vermejo, is, 

in large part at least, equivalent to the Laramie :formation, it seems highly probable that the poorly preserved leaves from the Laramie assigned to Z. corrugatus, and z. hendersoni also belong with Z. 

fibrillosus. 

The identification of these leaves as ZizyphU8 is, in 

my estimation, ~omewhat dubious. 

Occurrence: Fort Union :formation (lower) 8928; 

Animas :formation, 7496; Denver :formation, 317 (fig. 

11), 877 4 (figs. 5, 6) ; Raton :formation, 5690, 5712; 

Vermejo :formation (Upper Cretaceous), 8776 (fig. 

10). . 

·Zizyphoides mackayi Bell 

Zizyphoides mackayi Bell, 1949, p. 7 4, pl. 19, figs. 2, 3 ; pl. 20, 

figs. 2, 4. 

These elliptic remotely dentate leaves, described by 

Bell as coming :from the Post-Brazeau Paleocene strata 

in Alberta, Canada, do not seem to be matched by any 

found in the Paleocene rocks of the United States. 

Some in the latter that bear a superficial resemblance 

to Zizyphoides maclcayi and were formerly referred to 

Pali'lurus or Zizyphus have been found to be variants 

of Oercidiphyllu,m arcticum (Heer) Brown. The closest comparable specimens from strata in the United 

States are those called Z.izyphus ci111nam01noides Lesquereux from the Eocene Green River :formation in 

Wyoming. Is it possible that the Canadian specimens 

are :from Eocene rather than Paleocene strata~ 

VITACEAE 

Ampelopsis acerifolia (Newberry) Brown, n. comb. 

Plate 51, figures 1-18; plate 52, figures 1-8, 10; plate 59, 

figures 6, 11; plate 66, figure 7 

Popultts acerifolia Newberry, 1868, p. 65; 1898, p. 37, pl. 28, 

figs. 5-8. [Cockerell in 1906 changed the name to 

P. newberryi, but ace1··ifolia is here retained because 

that term has not been used with AmpelO]Jsis.] 

Acer articum Heer. Bell, 1949, p. 71, pl. 48, fig. 2. 

Ampelopsis brune1··i C(L1'bonensis (Ward) Cockerell, 1908, p. 103. 

Ampelopsis mont(tnensis Cockerell, 1908, p. 103. 

Ampelopsis xa1itholithensis (Warcl) Cockerell, 1908, p. 103. 

OerC'id·iphyllum elUpUcu,m (Newberry) Brown, 1939a, p. 491, 

pl. 52, figs. 13, 17 only. 

Cissus? C(Lrmon·i Knowlton, 1930, p. 113, pl. 47, fig. 5. 

C·isws lesquere'uxi Knowlton, 1930, p. 114, pl. 48, figs. 3, 4. 

Oiss1ts lobato-orenatn Lesquereux, 1878, p. 240, pl. 41, figs. 1-3 

[fig. 1, from Upper Cretaceous strata at Black Buttes, 

Wyo.] 

Oissus trimtsp'idata Heer. Lesquereux, 1878, p. 240, pl. 41, 

figs, 4-7. [Fig. 4, Middle Park formation; figs. 5, 6, 

Denver formation; fig. 7, Upper Cretaceous, Black Buttes, 

Wyo.] 

Dombe1fopsis neb?'a8cen8is (Newberry) Bell, 1949, p. 62, pl. 2, 

figs. 1, 4; p1. 9, figs. 1-3; pl. 10, fig. 1; pl. 12, figs. 9, 10; 

pl. 19, fig. 1; pl. 20, fig. 1; pl. 22, fig. 6; pl. 46, fig. 4; 

pl. 48, fig. 1. 

Popttl1tS cmspedodr01na Ward, 1887, p. 21, pl. 8, fig. 3. 

Popttlu.s lacoeana Knowlton, 1930, p. 60, pl. 22, fig. 5. 

Popul1ts smilacifoUn Newberry, 1898, p. 53, pl. 29, fig. 5. 

Populus whUei Ward, 1887, p. 22, pl. 8, fig. 4. 

Pomtlus zaddachi Heer. Knowlton, 1930, p. 60, pl. 22, fig. 8. 

Populus sp. Knowlton, 1930, p. 57, pl. 21, fig. 9. 

Trochodendro'ides nebrascensis (Newberry) Dorf, 1938, p. 61, 

pl. 11, figs. 1, 4, 6, 7. [Include entire synonymy, except 

Populus nebrascensis Newberry, 1898, p. 47, pl. 27, figs. 

4, 5, which is Oercid·iphyllmn (L'J'Cticum (Heer) Brown.] 

VWunHtm anceps Lesquereux, 1878, p. 227, pl. 38, fig. 11. 

VIburnum lakesii Lesquereux. Bell, 1949, pl. 61, fig. 1 ; pl. 62, 

figs. 3, 4. 

Vibtu"num~ tr'ine1·vis Berry. Be1l, 1949, p1. 61, figs. 2, 3. 

Vitis bnmm·i Ward, 1887, p. 69, pl. 32, figs. 1, 2. 

Vitis carbonensis Ward, 1887, p. 70, pl. 32, fig. 3. 

Vitis mtspidatn Ward, 1887, p. 71, pl. 32, figs. 6, 7 [not fig. 8, 

which is Platanus raynoldsi Newberry.] 

Vitis xantholithensis Ward, 1887, p. 71, pl. 32, figs. 4, 5. 

Ziz-yph1ts distort-Its Lesquereux, 1878, pl. 51, figs. 8, 9 [not fig. 7, 

which is Z. jib1·illosus (Lesquereux) Lesquereux.] 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 79 

These leaves or leaflets, as the case may be in some 

instances, have large, rounded, sometimes notably unequal marginal teeth, separated by more or less concave to sljghtly angular sinuses. In appearance the 

better leaves are maplelike. (See Acer arcticum Heer, 

1876, p. 86, pis. 22, 23, 24, figs. 1, 2; pl. 25, figs. 1-3.) 

The pdmary veins are in general fairly straight but 

may, in the more elliptic specimens, be somewhat 

curved. As evolution proceeded, the tendency toward 

lobation became more marked, the later leaves being 

distinctly more maplelike than those from the early 

Paleocene. 

The variants among these leaves suggest those of 

the living Parthenoci8sus (A1npelopsi8) tricuspidata 

(Siebold and Zuccarini) Planchon and Ampelopsis 

brevipedunmtlata ICoehne, of China, and the early 

leaflets on seedlings of the Virginia creeper, Parthenoci8sus quinq1.tejolia (Linnaeus) Planchon. Some 

also resemble strongly those. of a number of species of 

Viburnum, particularly TT. acerifolium Linnaeus, a 

shrub of the Northeastern United States and Canada. 

Because some of these leaves are strikingly like 

variants of Oercidiphyllum arcticum (Heer) Brown, 

there is likelihood that they may be misidentified as 

the latter and vice versa, particularly when both are 

found together. Thus, a number of specimens formerly assigned to Oercidiphyllurn are here reassigned. 

Occurrence: Fort Union formation (lower), 4625, 

4699 (pl. 51, fig. 10; pl. 52, fig. 7)' 4877, 5063, 5437, 

5718, 6154, 8163, 8227 (pl. 52, fig. 1), 8240, 8517 (pl. 

52, figs. 2, 3, 6, 10), 8545, 8673, 8780, 8884, 8893, 8899, 

8928,9104,9130 (pl. 51, fig. 17), 9301,9334, 9502,9530; 

(upper), 2420 (pl. 51, figs. 8, 15, 18; pl. 52, fig. 4; pl. 

59, fig. 11), 2432, 4974 (pl. 51, figs. 3, 7, 12), 5333, 

6359, 7659, 8225, 8255, 8521 (pl. 51, figs. 2, 13), 8523 

(pl. 51, figs. 1, 16), 8552 (pl. 59, fig. 6), 8774 (pl. 51, 

figs. 4, 11), 8922 (pl. 51, figs. 5, 6, 9, 14), 9085, 9092, 

9477; Coalmont formation, 5994; Dawson arkose, 5837, 

8188; Denver formation, 317 (pl. 52, figs. 5, 8; pl. 66, 

fig. 7), 8447; Evanston formation, 3653, 3658; Livingston formation, 4311, 6767; Middle Park formation, 

333, 337. 

Cissus marginata (Lesquereux) Brown, n. comb. 

Plate 53, figures 1-4, 6; plate 54, figures 1-4; plate 55, 

figures 4, 6, 7 

T1ib~tnwm ma1·ginat'wm Lesquereux, 1873, p. 395; 1878, p. 223, 

pl. 37, fig. 11; pl. 38, figs. 1-4 [not fig. 5, which is a 

small leaf of F'imts pla,n;icostatn Lesquereux.] 

Dorf, 1938, p. 73, pl. 15, figs. 3, 5 ; pl. 17, figs. 4, 5 ; 1942, 

p. 150, pl. 14, fig. 3; pl. 15, figs. 1, 5. 

Brown, 1939b, p. 252, pl. 59, fig. 7. 

Aoer colUe1·i Hollick, 1930, p. 99, pl. 78, fig. 5. 

Almts a1t1'a1·ia Knowlton and Cockerell. Knowlton, 1930, p. 49, 

pl. 15, fig. 6. 

Aln1ts carpini,folia Lesquereux. Knowlton, 1930, p. 49, pl. 15, 

fig. 2. 

Betula beat1·icina conto1·mis Hollick, 1930, p. 68, pl. 35, fig. 4. 

Betula brongniarti Ettingshausen. Heer, 1883, p. 81, pl. 96, 

fig. 3 only. 

Bet1tla fallax Lesquereux. Knowlton, 1930, p. 50, pl. 15, figs. 

7-10; pl. 16, figs. 1, 2. 

Betttla schi1nperi Lesquereux. Knowlton, 1930, p. 49, pl. 15, 

figs. 3, 4. 

Celastt·tts gnudini Lesquereux. Knowlton, 1930, p. 99, pl. 45, 

fig. 9 only. 

C·iss~ts coryUfolia Lesquereux. Knowlton, 1930, p. 113, pl. 47, 

fig. 4; pl. 48, fig. 1, pl. 49, fig. 1. 

Cisstts grossedentata Knowlton, 1930, p. 112, pl. 48, fig. 2; 

pl. 51, fig. 10. 

C·issus hesperia Knowlton, 1930, p. 112, pl. 47, fig. 1. 

Cissu.s lobato-crenata Lesquereux. Knowlton, 1930, p. 114, 

pl. 49, figs. 3, 4. 

Dorf, 1942, p. 153, pl. 16, fig. 3. 

C1·ataegtts 1ny1·icoides Lesquereux. Knowlton, 1930, p. 96, pl. 

44, figs. 7, 10. 

Fngtts pnpymcea Knowlton, 1917, p. 297, pl. 68, fig. 1. 

F·ictts? lakesi Knowlton, 1930, p. 60, pl. 28, qg. 1. 

Fmximus sp. Knowlton, 1930, p. 124, pl. 58, ftg. 7. 

G1·ew'iopsis ten~t'ifoUa Lesquereux, 1878, p. 2.58, pl. 40, fig. 14. 

Knowlton, 1930, p. 116, pl. 50, fig. 3. 

Negundo dec'ltrrens Lesquereux. Knowlton, 1930, p. 100, pl. 45, 

fig. 10. 

PhyllUes denve1·ensis Knowlton, 1930, p. 130, pl. 59, fig. 5. 

Phyllites he1·bacea Knowlton, 1924, p. 97, pl. 15, fig. 4. 

Phyllites leei Knowlton, 1917, p. 280, pl. 49, fig. 8. 

Phyllites pell1tcidens Knowlton, 1930, p. 130, pl. 39, fig. 2; 

pl. 56, fig. 4. 

Platnmts aceroides lntifolia Knowlton, 1924, p. 86, pl. 13, fig. 3 ; 

pl. 14, fig. 4; 1930, p. 76, pl. 33, fig. 1. 

Platan1ts aj]inis Lesquereux. Bell, 1957, p. 52, pl. 37, fig. 5; 

pl. 39, figs. 1, 4, 5 ;. pl. 42, fig. 6. 

Plntanus tnt'illelmae Goppert. Heer, 1883, p. 96, pl. 97, fig. 6; 

pl. 98, figs. 1, 2 ; pl. 99, fig. 1 ; pl. 103, fig. 4. 

Knowlton, 19).7, p. 322, pl. 93, fig. 1; 1930, p. 76, pl. 33, 

fig. 2. 

Platantts q·uillelnwe hee1·i Knowlton, 1917, p. 323, pl. 96, fig. 5; 

pl. 97, fig. 1; pl. 98, fig. 2. 

Plntan/us heer·i Lesquereux. Ward, 1887, p. 37, pl. 15, figs. 3, 4. 

Platantts nw1·ginata, (Lesquereux) Heer, 1883, p. 97, pl. 98, 

figs. 3-5 ; pl. 99, figs. 2, 3 ; p1. 101, fig. 5. 

Knowlton, 1930, p. 81, pl. 36, figs. 2, 3. 

Platanus? newben·yann Heer. Hollick, 1930, p. 83, pl. 46 

figs. 2, 3; pl. 47, fig. 3. 

Platanus? newberry ana cond'iUonalis Hollick, 1930, p. 83, pl. 4 7 

fig. 4. 

Platanus platanoides (Lesquereux) Knowlton, 1917, p. 323 

pl. 95, fig. 4; 1922a, p. 146, pl. 13, fig. 1 ; 1930, p. 82 

pl. 36, fig. 5. 

Platanus rhomboidea Lesquereux, 1878, p. 186, pl. 26, figs. 6, 7. 

Knowlton, 1930, p. 75, pl. 32, fig. 3. 

Platanus sp. Knowlton, 1930, p. 83, pl. 38, fig. 2. 

Poprultts denverens·is Knowlton, 1930, p. 6~, pl. 23, fig. 5. 

Populus knowltoni Berry. Knowlton, 1930, p. 61, pl. 23, figs. 

1-4. 

Querctts celastrifolin Lesquereux. Knowlton, 1930, p. 54, pl. 18, 

figs. 4, 6, 7 [not fig. 5, which is !lex' artocarpo·ides (Lesquereux) Brown]. 

Quercus purdonensis Knowlton, 1930, p. 52, pl. 17, fig. 2. 
80 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Quercus viburnifolia Lesquereux, 1878, p. 159, pl. 20, figs. 11, 12. 

Knowlton, 1930, p. 50, pl. 16, figs. 3-8; pl. 17, fig. 1. 

Dorf, 1938, p. 53, pl. 6, figs. 3, 5, 7 ; 1942, p. 154, pl. 16, 

figs. 5, 7, 8. 

Quercus whitei Lesquereux. Knowlton, 1930, p. 53, pl. 17, 

fig. 3 ; pl. 18, fig. 1. 

Rhamnus cleburni Lesquereux. Knowlton, 1917, p. 332, pl. 1i3, 

fig. 3. 

Rhus? viburnoides Knowlton, 1917, p. 328, pl. 98, fig. 5. 

Sophora pm·yea1·ensis Berry. Knowlton, 1930, p. 98, pl. 45, 

fig .. 8. 

Ulmus quercifolia Unger. Knowlton, 1930, p. 62, pl. 23, fig. 10. 

Viburnum antiqutm~t (Newberry) Hollick. Knowlton, 1924, 

p. 96, pl. 18, figs. 1-3. 

Bell, 1949, p. 77, pl. 61, fig. 4. 

Viburnum contortum Lesquereux. Knowlton, 1917, p. 346, 

pl. 108, fig. 3 ; 1930, p. 127, pl. 55, fig. 1. 

Viburnum crassum Knowlton, 1917, p. 277, pl. 52, figs. 3, 4. 

Viburnum dichotomttm Lesquereux, 1878, p. 225, pl. 38, fig. 6. 

Viburnum lakesi Lesquereux, 1878, p. 226, pl. 37, fig. 13. 

Knowlton, 1917, p. 348, pl. 110, fig. 3 [not fig. 4, which is 

Platanus raynoldsi Newberry] ; 1930, p. 128, pl. 55, fig. 3. 

Vibuntum mag1vum Knowlton, 1917, p. 347, pl. 110, fig. 2. 

Viburmtm nw1·ginatmn ravenscragensis Berry, 1935, p. 58. 

Viburnmn melaemmlt Knowlton and Cockerell. Knowlton, 1930, 

p. 128, pl. 55, fig. 5. 

Viburnum montanttm Knowlton, 1900, p. 73, pl. 19, figs. 1, 2 ; 

1917, p. 276, pl. 52, fig. 2. 

Dorf, 1938, p. 75, pl. 17, fig. 2; 1942, p. 150, pl. 15, figs. 2, 3. 

Viburnum platanoides Lesquereux, 1878, p. 224, pl. 38, figs. 8, 9. 

Viburnum? problematicum Knowlton, 1900, p. 71, pl. 19, fig .. 4 ; 

1917, p. 276, pl. 49, fig. 9. 

Vib,ztrnttm richardsoni Knowlton, 1930, p. 126, pl. 52, fig. 8; 

pl. 53, figs. 1, 3-5 ; pl. 54, figs. 2, 3. 

Viburnum rotundifolium Lesquereux, 1878, p. 225, pl. 37, fig. 

12 ; pl. 38, fig. 10 ; pl. 61, fig. 22. 

Knowlton, 1924, p. 96, pl. 16, fig. 1. 

Viburnttm s·imile Knowlton, 1917, p. 277, pl. 49, fig. 3. 

Viburnum speciosmn Knowlton, 1917, p. 347, p. 111, figs. 1-5; 

1924, p. 96, pl. 16, fig. 3. 

Vibm·num whympe1··i Heer. Lesquereux, 1878, p. 225, pl. 38, 

fig. 7; pl. 61, fig. 23. 

Knowlton, 1900, p. 72, pl. 17, fig. 1; pl. 18, fig. 1; pl. 19, 

fig. 3. 

Viburnmn sp. Knowlton, 1917, p. 278, pl. 44, fig. 3. 

Vitis? platanifoUa Knowlton, 1917, p. 339, pl. 103, fig. 2. 

Vitis stantoni (Knowlton) Brown. Bell, 1949, p. 75, pl. 3, 

fig. 3 ; pl. 9, fig. 5. 

Winchellia triphylla Lesquereux, 1893, p. 209, pl. 8, fig. 1. 

The well-preserved ternate or trifoliate leaf (pl. 53, 

fig. 4), called by Lesquereux W inchellia triphylla in 

a posthumous publication, adds new information on 

the mixture of many-named Cretaceous and Paleocene leaves variously assigned, as seen from the 

synonymy, to the alders, birches, beeches, maples, 

grapes, sycamores, viburnums, and others. This fine 

compound leaf, it now appears, is not the only one of 

its kind known, others being N egurulo decurrens Lesquererix (ICnowlton, 1930, pl. 45, fig. 1), Platanus 

guillelmae Goppert (ICnowlton, 1930, pl. 33, fig. 2, in 

which the right-hand leaflet is obscure), and some not 

hitherto figured (pl. 54, figs. 3, 4). One unfigured 

specimen, collected by me in 1930 from the Aspen 

shale on Fontanelle Creek north of Kemmerer, Wyo., 

is also now in the U.S. National ~1useum collections. 

The normal terminal leaflets of this species have the 

appearance of partly opened fans and are incipiently 

to strongly three-lobed like some maples and viburnums. However, many of the figured single leaves, 

purporting to have been terminal leaflets, may in reality have been simple leaves, for they show no evidence of leaflet scars on their long petioles. Moreover, some of the assumed ancestral, simple, Cretaceous specimens (pl. 55, fig. 2) have remains of 

narrow acute stipules. It seems evident, therefore, 

that 0. 7narginata had both simple and compound 

leaves. 

Associated at all localities with the characteristic 

terminal leaflets are the more or less asymmetric, petioled, lateral leaflets, of great variety in shape, venation, and marginal features, causing much diversity 

of identification as illustrated by the extensive synonymy here proposed. Perhaps I have included in 

this species some aberrant specimens that properly 

belong elsewhere, for I have no infallible method of 

classifying such disputable material. 

The venation of these leaves is fundamentally pinnate, but the tendency toward palmateness is unmistakable, the first pair of basal secondaries being, in 

many specimens, especially strong, so tl1at they should 

probably be called primaries. These veins have numerous lateral branches from the lower side but sometimes give off a strong branch or two from the upper 

side. Most veins and their branches terminate in 

marginal teeth that differ greatly in size in different 

leaves. The prominent, fairly straight primaries are 

decurrent at acute angles into the top of the petiole, 

or, if they appear to 'branch from the mid vein a lit-

. tie higher, are decurrent on the mid vein into the 

petiole. This feature helps to distinguish these leaves 

from the normal leaves of Platanus raynoldsi Newberry in which the curved or bowed primaries branch 

from the midvein at greater angles and at a centimeter, more or less, within the blade above the top of 

the petiole. Otherwise, the general venational pattern, the overall outline, and the teeth, separated by 

smoothly scalloped sinuses, are very similar in both 

species. The teeth of this species, however, are, with 

few transitional exceptions, sharp pointed and are 

thus in contrast with the blunt, rounded teeth of an 

associated vitaceous species, Ampelopsis acerifolia 

(Newberry) Brown. 

A canvass of the possibilities among plant families 

having species with features comparable to those here 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 81 

described suggests that this species most likely belongs 

in the grape family, Vitaceae, rather than in the 

Caprifoliaceae or Platanaceae. Among living Vitaceae, Rhoicissus usambarensis Gilg, from South Africa, 

is a fair match for this species, but difficulty in explaining the wide breach in distribution makes an 

assignment to Rlwicissus seem like stretching a point. 

Choice of Ampelopsis or Vitis also seems less appropriate than Oissus. Further, although these leaves are 

here called Oissus, their resemblance in many particulars to those of Platanus prompts the needed caution 

of looking for examples of compound leaves in the 

ancestry of Platanus. So far, after examining many 

seedlings, suckers, and young shoots of living species 

of Platanus, I have found no compound specimens 

that could be regarded as possible atavistic throwbacks. 

Lesquereux saw a resemblance between W inchellia 

triphylla and the deerfoot or vanilla-leaf, Achlys triphylla DeCandolle, of the barberry family, found in 

the Pacific Coast region from California to British 

Columbia. Except for the trifoliateness, however, the 

two are in other features not closely comparable, and 

the assignment to the Berberidaceae is deemed farfetched. It seems remarkable that Lesquereux did not 

detect the striking similarity in every respect between 

the terminal leaflet of Winchellia triphylla and the 

Cretaceous leaves he had called Viburnum margilnatum. 

After exhaustive comparisons between Cretaceous 

and Paleocene assemblages of these leaves, I have 

found no unfailing features that clearly distinguish 

them. At some localities the Cretaceous leaves appear 

to average smaller in size and may be somewhat 

rounder in . outline, with the apical lobe less pronounced. The terminals of some Paleocene specimens show a stronger lobing or tendency to be aceroid or platanoid in outline. 

Closely similar to this species is the Upper Cretaceous Vitis stantoni (Knowlton) Brown, which differs chiefly in having leaves that are broad rounded, 

and usually with cordate bases. Perhaps this species 

should also be reassigned to Ois8us·, although no compound examples have been reported. 

The more distant ancestors or relatives of this species may possibly be found among the many described forms of Aspidiophyllum, Betulites, Oissites, 

Orataegus, Grettoiopsis, H edera, Platanus, Populites, 

and Viburnum. These occur in the Cretaceous of 

Greenland and Alaska, and notably in the Dakota 

sandstone of l{ansas, Nebraska, and Minnesota. A 

few are figured here (pl. 55, figs. 1-3) for comparison in shape, venation, and marginal dentition with 

the, corresponding features of the later species. The 

speCimen (pl. 55, fig. 7) from the Cretaceous at Parowan Gap, Utah, is noteworthy in showing a rounded 

base with a perfoliate petiole, thus simulating some 

leaves of fossil and living Platanus. 

Occurrence·: Fort Union formation (lower), 4280, 

4976, 6297, 8551, 9193, 9252 (pl. 53, fig .. 4), 9334; (upper), 8774; Animas formation, 5455, 6309, 6443, 7485, 

7498, 9565; Coalmont formation, 6107; Dawson arkose, 

331, 5783, 5835-5840, 8187, 8307; Denver formation, 

317 (pl. 53, fig. 1; pl. 54, fig. 3), 5738, 8672; Ferris 

formation, 3852, 4369, 5494, 6416 (pl. 53, fig. 6; pl. 54, 

fig. 2), 6431 (pl. 53, figs. 2, 3), 8516, 8528, 8660~ 

Puerco formation, 7495 (pl. 54, figs. 1, 4); Raton formation, 5154, 5679, 5683, 5701, 5711, 5714, 5802, 5826. 

Cretaceous localities: Pl. 55, fig. 4, Mesaverde formation, south of Gallup, N. Mex.; pl. 55, fig. 6, Hell 

Creek formation, east of Glendive, Mont.; pl. 55, fig. 

7, l{aiparowitz formation, Parowan Gap, Utah. 

Cissites rocklandensis Brown, n. sp. 

Plate 27, figure 8 

Large ovate three-lobed leaf, the two lateral lobes 

relatively small or incipient. Venation with two 

large primary veins that rise from the top of the 

petiole and run toward the lateral lobes; and secondary veins, widely spaced from the middle of the blade 

to the apex. Margin entire. Base somewhat cordate. 

Superficially this leaf resembles those called Ficus 

planicostata Lesquereux, but I have never seen any 

lobed specimens of the latter among the abundant 

material in the collections of the U.S. National Mu-

. seum. Cretaceous species that may have been ancestral to Oissites rockla!ndensis• are Oissites .colgatensis 

Brown, 0. lobatus Dor£, Dombeyopsis obtusa Lesquereux, D. trivialis Lesquereux, Phyllites dombeyopsoides l{nowlton, P. populoide8 Knowlton, and P. trinervis l{now lton. 

Occurrence: Raton formation, 5679 (fig. 8). 

Parthenocissus ursina Brown, n. sp. 

Plate 48, figures 1, 3, 4 

Leaflets 12 em or more long, 5 em wide, narrow 

elliptic in outline, blunt pointed at the apex, and cuneate, more or less asymmetric, at the base. Secondary veins relatively few, widely spaced, with some intersecondaries, slightly undulant and forming loops 

but with small branches into the marginal teeth. Margins serrate with large teeth and angular sinuses. 

Texture apparently thin. 

No complete leaflet or compound leaf was found. 

The fragments, however, suggest comparison with the 

leaves of the living Parthenocissus quinquefolia (Sie-
82 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

bold and Zuccarini) Planchon, of the Eastern United 

States. The species seemingly was larger and coarser 

than the delicate P. tertiaria (Lesquereux) Knowlton 

:from the Eocene Green River :formation in Wyoming. 

Occurrence: Early Paleocene, 854 7 (figs. 1, 3, 4), in 

the Bear of Simpson, Crazy Mountain area, Montana. 

Vitis loba.ta (Knowlton) Brown, n. comb. 

Plate 56, figures 3, 7, 10 

Aralia lobata Knowlton, 1924, p. 93, pl. 17, figs. 1, 2. 

Aralia sp. Knowlton, 1924, p. 94, pl. 6, fig. 4. 

The specimens from Montana figured here resemble closely those called Aralia by Knowlton from 

southern Colorado, except that the lobes of the latter 

are apparently not sharply toothed. This may be a 

local variation, but it may also mean that two species 

are involved. The Montana specimens, under magnification, show some evidence of having been glandular or at least somewhat hairy. There is a suggestion 

of relationship between this species and the Upper 

Cretaceous Cis sites panduratus Knowlton ( 1917, p. 

274, pl. 49, fig. 10) from the Vermejo formation in 

Colorado. Leaves of the living Vitis mbra Michaux 

[or V. palmata Vahl] match those of V. lobata fairly 

well. 

Occurrence: Fort Union formation (lower) 8519 

(figs. 3, 7, 10) ; Animas formation, 7483. 

Vitis olriki Heer 

Plate 27, figure 10; plate 59, figure 10 

Vitis olriki Heer, 1868, p. 120, pl. 48, fig. 1. 

Lesquereux, 1878, p. 241, pl. 41, fig. 8. 

Knowlton, 1930, p. 115, pl. 49, fig. 6; pl. 50, fig. 4. 

Vitis arctica Heer, 1868, p. 120, pl. 48, figs. 2, 2c; 1871, p. 478, 

pl. 55, fig. 1. 

Vitis inominata Knowlton, 1917, p. 339, p. 107, fig. 1. 

Vitis leei Knowlton, 1917, p. 338, pl. 66, fig. 4. 

Vitis sp. Heer, 1871, p. 478, pl. 55, figs. 5d, 6 (seed). 

Ficus denveriana Knowlton, 1930, p. 64, pl. 24, fig. 1. 

Popul1ts cordata Newberry, 1868, p. 60; 1898, p. 38, pl. 29, 

fig. 6. 

Popul'tt8 richardsoni Heer. Knowlton, 1930, p. 62, pl. 23, figs. 

6, 8. 

Tilia sveciosissima Knowlton, 1917, p. 336, pl. 67. 

These are relatively large leaves with cordate bases 

and entire or toothed margins. Between the rounded 

teeth are smooth concave sinuses. The seed Vitissp., 

figured by Heer and found associated with a leaf of 

that genus at Atanekerdluk, Greenland, probably· belongs with this species. 

Occurrence: Fort Union for~ation (lower) 8517 

(pl. 27, fig. 10), 8899,9104 (pl. 59, fig. 10); (upper) 

2414, 8921; Dawson arkose, 331, 5837, 5840, 8447; 

Evanston formation, 3653; Middle Park· formation, 

337; Raton formation, 5679, 5684. 

Vi.tis sp. 

Plate 53, figure 5 

This rounded seed with the two prominent adjacent cavities clearly belongs with the Vitaceae. As 

such it is the first to be reported from the Paleocene 

strata of the Rocky Mountains · and Great Plains. 

However, as no leaves were found with the specimen, it cannot be plausibly assigned to any one of 

the vitaceous leaf species here listed. 

Occurrence: Fort Union formation (upper) 9132 

(fig. 5). 

STERCULIACEAE 

Dombeyopsis magnifica Knowlton 

Dombeyopsis 1nagnijica Kno'\ylton, 1930, p. 125, pl. 54, fig. 4; 

pl. 55, fig. 4; pl. 57, fig. 1; pl. 58, fig. 8. 

Dombeyopsis grandifolia Unger. Lesquereux, 1878, p. 255, pl. 

47, fig. 6. 

Dombeyopsis? sedaliensis Knowlton, 1930, p. 126, pl. 56, fig. 5; 

pl. 59, fig. 1. 

Ficus sp. Knowlton, 1930, p. 72, pl. 29, fig. 5. 

Ph1Jllites aTistolochioides Knowlton, 1930, p. 129, pl. 57, fig. 2 

only. 

The implication that these leaves are related to living Dmnbeya seems to me farfetched. They are somewhat like Oissites 8teenstmpi Heer (1883, p. 118, pl. 

81, fig. 1) from Greenland. 

Occurrence: Dawson arkose, 325, 5831, 5839; Denver formation, 317. 

Pterospermites cordatus Ward 

Plate 41, figures 1, 2, 4 

Pterospermites cordaVus Ward, 1887, p. 93, pl. 41, fig. 4. 

Ficus berthoudi Lesquereux, 1888, p. 49. 

Knowlton, 1930, p. 65, pl. 25, figs. 1, 2, 0. 

Ficus r(ttonensis Knowlton, 1917, p. 306, pl. 74, fig. 4. 

These large, entire-margined, heart-shaped leaves resemble some described as Ficus, Paranymphaea, Ooccolobis, DombeyopsiB·, Catalpa, and others, but differ 

in venational and other features from all these. The 

regular parallel interconnecting tertiary veins are part~cularly coarse and widely spaced. I doubt that this 

species is allied with the genus Pterosperm;wm but have 

no better suggestion to make. 

Occurrence: Fort Union formation (lower), 4625 

(fig. 4), 6892, 8678 (fig. 1); (upper) 2416, 7776 (fig. 

2), 8190, 8774, 8921, 9109; Dawson arkose, 325; Denver formation, 317; Raton formation, 5140. 

DILLENIACEAE 

Dillenites garfieldensis Brown, n. sp. 

Plate 57, figures 2, 5 

Leaves large, elliptic, with cuneate to rounded bases 

and acute apexes. Length of petiole unknown. Margin beset with mammilately rounded to pointed, un-

" ,:!;.. 

~ .. AJ" 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 83 

equal, serrate teeth separated by angular sinuses. 

Venation pinnate, the numerous equally spaced secondaries curving gracefully away from the midvein 

into the larger marginal teeth and sending inconspicuous branches into the subsidiary teeth. Secondaries connected by closely spaced diagonal parallel 

veinlets. 

This species differs from Dillenites miorodentatus 

(Hollick) Berry in the Wilcox group of the Southeastern St~tes and in the Tertiary of Alaska by having subsidiary teeth in addition to the main teeth and 

in having the teeth rounded instead of acute. A further study of Hollick's four species of Dillenites from 

the Tertiary of Alaska may prove that they are only 

a single species, as he himself intimated. 

Occurrence: Fort Union formation (upper), 8774 

(figs. 2, 5; specimens in Chicago Nat. History Mus.). 

MYRTACEAE 

Myrtophyllu.m torreyi (Lesquereux) Dorf 

Plate 50, figures 1-4, 7-9 

Myrica to1-reyi Lesquereux. Dorf, 1938, p. 49, pl. 6, figs. 1-3. 

Myrtophyll1t1n torreyi (Lesquereux) Dorf, 1942, p. 146. [See 

synonymy.] 

Apocynophyllum wilcoxensis Berry. Knowlton, 1924, p. 95, 

pl. 14, fig. 5. 

Myrica sp. Kno\vlton, 1930, p. 44, pl. 11, fig. 7. 

Except that the intramarginal vein is more subdued and sometimes absent, these Paleocene leaves 

with eucalyptoid venation are scarcely distinguishable from their Cretaceous predecessors. Their reference to the family Myrtaceae may be appropriate in 

respect to venation, but the general absence of marginal teeth in leaves of that family is discordant, thus 

raising doubts about the present reference. 

I have not seen the specimens called Oelastrop'hyllu'ln benedini by Saporta and Marion (1873, p. 67, 

pl. 10, fig. 6; pl. 12,. figs. 1, 2; 1878, p. 86, pl. 14, fig. 2) 

from the Paleocene at Gelinden, Belgium, and am not 

certain that the venation is accurately drawn, but 

there is a strong suggestion of resemblance between 

them and the leaves of 11/yrtophyllum torreyi. The 

leaves described by Heer as Rhus lwlbolliana (1883, 

p. 134, pl. 59, fig. 7) from Atanekerdluk, Greenland, 

are also similar, and favorable comparison can be 

made with llfyrica toilooxensis Berry (1916a, p. 188, 

pl. 18, fig. 1) from the Eocene at Grenada, Miss. The 

illustration of the leaf called Apooynophyllttm wilooxensis Berry by Knowlton is retouched and is inaccurate in showing numerous regular, evenly spaced secondary veins; but these veins are irregular and unevenly spaced and they unite near the margin to form 

an obscure intramarginal vein. 

Occurrence: Fort Union formation (lower) 8196, 

8551 (figs. 2, 7), 8666, 8677 (figs. 3, 4), 8899, 9210 

(fig. 1); Animas formation 7483; Dawson arkose, 

5738, 8188 (fig. 8), 8881 (fig. 9). 

. TRAPACEAE 

Trapa angulata (Newberry) Brown, n. comb. 

Plate 58, figures 1-12 

Neuropteri.s angula.ta Newberry in Ives, 1861, p. 131, pl. 3, 

fig. 5. 

Trapa? cunea,ta Knowlton, 1900, p. 64, pl. 5, fig. 6. 

Trapn? mic1·ophylln Lesquereux, 1878, p. 295, pl. 61, figs. 16-17a. 

Dawson, 1887, p. 31, pl. 2, figs. 19, 19a. 

Ward, 1887, p. 64, pl. 28, figs. 2-5. 

Knowlton, 1899, p. 661, pl. 77, figs. 3, 4; 1900, p. 62, pl. 5, 

fig. 7; 1919, p. 627. 

Hollick, 1930, p. 109, pl. 84, fig. 4. 

Berry, 1935, p. 61, pl. 19, figs. 1-11. 

Brown and Houldsworth, 1939, p. 336-339, text figs. 1-9. 

F1tcus lignitmn Lesqnereux, 1878, p. 42, pl. 61, figs. 24, 24a. 

Ward, 1887, p. 549, pl. 1, fig. 1 only. 

Knowlton, 1900, p. 17, pl. 3, fig. 4. 

Nel1tntbites strinta Berry, 1935, p. 38, pl. 8, figs. 1-3. 

Nyn1-phaeUes angulatus (Newberry) Bell, 1949, p. 64, pl. 17, 

figs. 4, 7. 

Nymphaeites strintus (Berry) Bell, 1949, p. 67, pl. 17, figs. 

1-3, 5, 6. 

Berry (1935, p. 61) and Bell (1949, p. 64) have 

summarized the history and information concerning 

this species, but Bell did not cite the report by Brown 

and Houldsworth ( 1939) of fruit-bearing specimens 

from the Ravenscrag formation in southern Saskatchewan, Canada. These fruits have been found at nearly 

all localities that have also yielded the characteristic 

leaves. 

N elu1nbite8 striata Berry from the Whitemud formation (Upper Cretaceous) in Canada has all the 

features of the typical compound foliage of Trapa 

angulata except that the leaflets tend to be eccentrically peltate, the blade being attached to the petiole 

near the margin or some distance inward. Such leaflets, wherever reported, as at Point of Rocks, Wyo., 

are found associated with the typical floating foliage 

of Trapa angulata, thus, together with the similarities in anatomy, arousing the suspicion of close relationship," if not identity. 

The National Museum's collection of Trapa angulata includes many almost perfect rosettes of this plant 

both from Upper Cretaceous and Paleocene strata. 

Inspection of these rosettes shows that the leaflets near 

the center tend to be reniform, rounded, and peltate, 

and those farther out, spatulate, rounded, truncated, 

cuneate, and attached basally. The margins are serrate with sharp teeth. The secondary veins or their 

branches do not, as a superficial glance might indi-
84 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

cate, enter the marginal teeth but run into glands at 

the bases of the teeth at the angle of the sinus. 

The affinity of this species with living Trapa may 

well be questioned, and comparison with water-cress, 

although attractive so far as habit is concerned, is 

otherwise untenable. 

The fruits lack the conspicuous "horns" of the living species. 

Occurrence: Fort Union formation (lower), 2421, 

3980, 5438, 6360, 8229, 8673, 8678, 8677, 9180; (upper) 

2420 (figs. 3-5, 7-12) , 8550; Ravenscrag formation, 

sec. 4, T. 2, R. 22 W., second meridian, Saskatchewan, 

Canada (figs. 1, 2) . 

Trapa paulula (Bell) Brown, n. comb. 

Plate 58, figure 13 

Adiantum? pattlttlurn Bell, 1949, p. 41, pl. 1, figs. 2, 4, 6, 10. 

Legurninosites stagnurn Bell, 1949, p. 68, pl. 1, figs. 8, 9. 

This species differs from Trapa angulata (Newberry) Brown in being notably smaller, with floating 

rosettes not distinctly four-parted and with cuneate 

leaflets throughout. The submerged foliage is many 

times dichotomously forked into filiform divisions. 

No definite mid vein is discernible. Most of the veins 

arise from the top of the petiole and fork several 

times, thus simulating roughly the venation of some 

ferns. The marginal dentition is blunter than that 

in T. angulata and is confined to the a pi cal half of 

the leaflet. 

The figured specimen shows the thickened subsurface portion of th~ stem that bore the floating· rosette 

of leaves. Farther down the stem are remains of the 

filiform submerged :foliage. _ 

Occurrence: Fort Union :formation (lower), 7004 

(fig. 13); (upper), 4896. 

MELASTOMACEAE 

Melastomites montanensis Brown, n. sp. 

Plate 56, figures 1, 2, 5, 6 

Leaves ovate lanceolate, attaining a length o:f 15 em. 

Apex acute, base cuneate. Margin entire. Venation 

composed o:f a strong midvein flanked by one pair o:f 

strong primaries that arise from the top o:f the petiole 

or :from the midrib a short distance above the top of 

the petjole and upward, extending to the very apex 

o:f the blade. Branches :from the primaries coalesce 

near the margin to :form a minor pair o:f intramarginal veins. Midvein, primaries, and intramarginal 

vein connected by closely spaced, more or less horizontal veinlets and secondaries, as in SMsafras and 

Oinna1nonum. Surfaces of the better preserved leaves 

display minute dots, about 300 per sq em, that were 

probably glandular. Petioles 2 to 3 em long. 

These leaves are somewhat larger than but resemble those of M elastomites verus (Berry) Berry, originally Oinnammnum vera Berry, from the Eocene of 

the Southeastern States. They may be compared favorably with those of a number o:f species of Toco<Ja 

:from South America. 

Occurrence: Fort Union :formation (lower), 5716, 

7005 (fig. 6), 7662 (fig. 5), 8519 (figs. 1, 2), 8897; 

Ferris :formation, 6420. 

CORNACEAE 

Cornu's hyperborea Heer 

Plate 59, figure 1 

Corntts hyperborea Heer, 1871, p. 476, pl. 50, figs. 3, 4. 

Leaves ovate, with cuneate to rounded bases and 

acute to blunt apexes. Margin entire. Secondary 

veins usually four on each side of the midvein, curved 

strongly upward toward the apex. For more than 

half the length of the leaf, no secondaries branch 

:from the midvein, and there are no branches from 

the secondaries toward the margin. The tertiary venation is composed of more or less parallel veinlets, 1 or 

2 mm apart and at or nearly at right angles to the 

midvein and secondaries and practically horizontal as 

viewed by the observer when holding the leaf upright. 

These specimens have all the earmarks o:f being an 

authentic species o:f Comus. However, no flowers or 

seeds of Oornu8 have yet been found in the Paleocene 

strata o:f the Rocky Mountain region. The flower described by I\:nowlton as Om·nus speciosissima (1922a) 

from Converse County, Wyo., although superficially 

comparable, is very likely not that of a dogwood. 

Moreover, the locality is in the Lance :formation (Upper Cretaceous) and is in T. 34 N., not T. 33 N., as 

cited. 

Numerous :fossil species o:f 0 om us, based chiefly on 

leaves, have been described-some authentic but most 

o:f them probably spurious. As the colored involucra} 

bracts o:f the flower-heads :fall separately, it is most 

unlikely that the usual :four-parted flower head would 

be preserved intact. If it should be preserved, the 

center, composed o:f a number of flowers, would add a 

characteristic identifying :feature. Most so-called :fossil dogwood flowers lack this conspicuous center and 

hence seem unconvincing to me. 

This species resembles Comus platyphylla Saporta 

( 1868, p. 391, pl. 11, figs. 8, 9) from Sezanne in the 

Paris Basin. 

Occurrence: Fort Union formation (upper), 4871 

(fig. 1), 9109, and an inexact locality in Montana; 

Middle Park formation, 337. 
SYSTEMATIC DESCRI_PTION OF THE MEGASCOPIC FLORA 85 

·Cornus nebra.scensis Schimper 

Plate 59, figures 2-5, 7, 12 

Cm·mts cwuminata Newberry, 1868, p. 71. 

ConMts nebTascensis Schimper, 1874, v. 3, p. 54. 

Cm·nus hnpr·essa Lesquereux, 1878, p. 243, pl. 42, fig. 3. 

Cont~ts newben·yi Hollick. Newberry, 1898, p. 124, pl. 37, figs. 

. 2, 3 [not fig. 4, which after cleaning shows prominent 

teeth and is F'·icus m·tocaTpoides Lesquereux]. 

This species differs from Oornus hyperborea Heel'" 

in having somewhat smaller leaves with more secondaries in the apical halves. 

Occurrence: Fort ·union formation (lower), 3980, 

4626 (fig. 5), 5389 (fig. 12), 5720, 6765, 8567 (figs. 2, 

3, 7), 8781, exact locality unknown (Hollick's specimens of Oornus newberryi); (upper), 8195 (fig. 4), 

8920; Middle Park formation, 337. 

Nyssa alata (Ward) Brown, n. comb. 

Plate 60, figures 1-4, 6 

Sapindus alat~ts Ward, 1887, p. 68, pl. 31, fig. 4 [not fig. 3, 

which is Sapind~ts a[fin-is Newberry]. 

Andr·omeda scr·iptct Knowlton, 1917, p. 344, pl. 112, figs. 1, 2. 

Apeibops·is nemnex·icana Knowlton, 1917, p. 336, pl. 101, fig. 3. 

ChTysobalanus colomdensis Knowlton, 1930, p. 95, pl. 40, figs. 

2, 3; pl. 43, figs. 3, 8, 9 [not other figures] ; pl. 44, figs. 

1, 2. 

Clwysobalamts? lanceolatus Knowlton, 1930, p. 96, pl. 44, fig. 5. 

Cor·mts emrnonsi Ward, 1887, p. 55, pl. 26, fig. 2 only. 

Cor·nus impTessa Lesquereux. Knowlton, 1930, p. 118, pl. 51, 

fig. 2 only. 

Diospyr·os brachysepala Braun. Ward, 1887, p. 104, pl. 49, 

figs. 1, 2. 

Knowlton, 1930, p. 121, pl. 51, figs. 5-7, 9; pl. 52, fig. 3. 

F·ic~ts? alata Knowlton, 1930, p. 64, pl. 23, fig. 9. 

F·icus occidental-is (Lesquereux) Lesquereux. Knowlton, 1930, 

p. 68, pl. 27' fig. 4. 

F-imts oval·is Lesquereux, 1878, p. 198, pl. 30, fig. 2. 

Jnga heter·ophylla Knowlton, 1917, p. 327, pl. 54, fig. 5. 

La~tnts social·is Lesquereux. Knowlton, 1930, p. 85, pl. 38, fig. 4. 

Nyssa lanceolata Lesquereux. Knowlton, 1917, p. 343, pl. 108, 

fig. 1 ; pl. 113, fig. 2 ; 1930, p. 120, pl. 52, fig. 2. 

?Sapind~ts obt1tsifoUus Lesquereux. Knowlton, 1930, p. 102, 

pl. 46, fig. 3 [same specimen as Ficus occidentalis, pl. 

27, fig. 4]. 

Sophor-a TichaTdsoni Knowlton, 1930, p. 97, pl. 45, fig. 7. 

Phyllites calhanensis Knowlton, 1930, p. 130, pl. 56, fig. 1. 

The assignment of these leaves to Nyssa is unconfirmed by any authentic seeds suggestive of the genus. 

The leaf identified by Lesquereux ( 1878, p. 245, pl. 35, 

fig. 5) as Nyssa lanceolata, from the Denver formation at Golden, Colo., is a leaflet of Fraxinus eooeniaa 

Lesquereux; and the seed, identified with the leaf 

(renamed by Knowlton, 1930, p. 120, as Nyssa dernveriana), is so poorly preserved and the ·artist's drawing of it (Lesquereux, 1878, pl. 35, fig. 6) so idealized 

that only confusion can result from retaining it as a 

named object. 

The leaves of Nyssa alata can be matched by the 

smaller leaves of the black tupelo, Nyss·a sylvatica 

Marshall, of the Eastern United States. 

Occurrence: Fort Union formation (upper) 2416 

(figs. 2, 4, 6), 5388, 5609, 8224 (fig. 3), 8250, 8257, 

8542 (fig. 1) ,. 877 4; Dawson arkose 325, 5837, 5838, 

8672; Denver· formation, 317; Ferris formation, 5971; 

Raton formation, 5679, 5683, 5704. 

Nyssa borealis Brown, n. sp. 

Plate 61, figure 4 

Leaves elliptic, with entire margins. Secondary 

veins widely spaced, branching from :the midrib almost at right angles and joining the secondaries above 

in conspicuous loops well within the margin. 

These leaves resemble those of the water tupelo, 

Nyssa aquatiaa Marshall, of the Southeastern United 

States. 

Occurrence: Fort Union formation, 4981 (fig. 4); 

Ferris formation, 8516. 

Nyssa 1 obovata Knowlton 

Nyssa? obovata Knowlton, 1930, p. 121, pl. 54, fig. 1. 

No further light on the identity of this specimen 

has appeared. 

Occurrence : Denver formation, 317. 

ERICACEAE 

Kalmia elliptica Brown, n. sp. 

Plate 40, figure 3 

Leaves elliptic, with entire margins. Secondary 

veins conspicuously decurrent at the thick midrib, 

somewhat undulant or irregular, forking and looping 

near the margin. 

These leaves compare fairly well with those of the 

mountain laurel, Kalmia latifolia Linnaeus of the 

Eastern United States. 

Occurrence: Fort Union formation, 4661 (fig. 3); 

Denver formation, 317. 

OLEACEAE 

Fraxinus eocenica Lesquereux 

Plate 57, figure 4; plate 62, figures 1-7 

Frax·imts eocenica Lesquereux, 1878, p. 229 ; 1883, p. 123, pl. 

20, figs. 1-3. 

Knowlton, 1930, p. 123, pl. 52, figs. 4-6. 

Ar·alia alexoens-is Bell, 1949, p. 7G, pl. 5G, figs. 2, 5 [upper 

leaf] ; pl. 59, fig. 1. 

Celnst?"in-ites ·insignis ( Heer) Bell, 1949, p. 71, pl. 58, fig. 3; 

pl. 59, fig. 4. 

Ju.glans denver-ia.na Knowlton, 1930, p. 44, pl. 12, figs. 1, 2; 

pl. 13, figs. 2-4. 

Ju.glans rugosa Lesquereux, 1878, p. 28G, pl. 54, figs. 5, 14; 

pl. 55, figs. 1-9. 

Knowlton, 1930, p. 46, pl. 14, fig. 1. 
86 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Ju,qlan8 sp. Knowlton, 1930, p. 48, pl. 14, fig. 5; pl. 15, fig. 1. 

N 1J88a lanceolata Lesquereux, 1878, p. 245, pl. 35, fig. 5. 

Knowlton, 1930, p. 120, pl. 52, fig. 2. 

l'teroca:rya? retu8a Lesquereux. Knowlton, 1930, p. 48, pl. 14, 

figs. 2-4 ; pl. 15, fig. 5. 

Q'tterc'tt8 leonis Knowlton, 1930, p. 53, pl. 18, fig. 2 [not fig. 3, 

which is Eu,.convmia serra,ta (Newberry) Brown]. 

Quermts straminea Lesquereux, 1878, p. 151, pl. 19, fig. 6. 

Q'uerc1tS whUnwni Knowlton, 1930, p. 52, pl. 17, fig. 5. 

Sapindus caudatus Lesquereux, 1878, p. 264, pl. 48, fig. 6. 

Knowlton, 1930, p. 101. 

The leaflets of this species are entire or remotely 

toothed, asymmetric, ovate lanceolate, elliptic, or oblanceolate in outline, petioled, with acute or attenuated apexes and cuneate to rounded bases. Fragments of the compound leaves with three leaflets still 

attached have been found, but no seeds attributable to 

Fraxinus have been recovered. 

Some of the specimens here assigned to Frawinus 

eocenioa were originally identified by Lesquereux as 

Juglans rugosa and came from a locality doubtfully 

considered to be Evanston, Wyo. The complicated 

history of J. rugosa was discussed by Knowlton ( 1930, 

p. 46). Other correlative specimens, however, were 

identified simultaneously by Lesquereux as Frawinus 

eocenica, from the Denver formation at Golden, Colo. 

This seems to be a more satisfactory allocation. 

Occurrence: Fort Union formation (lower), 4626 

(pl. 62, fig. 7), 4661, 8165; (upper), 5760, 8234 (pl. 

62, fig. 1), 8774 (pl. 62, figs . .3-5; pl. 57, fig. 4) ; Coalmont formation, 6105 (pl. 62, fig. 6); Denver formation, 317 (pl. 62, fig. 2); Evanston formation, 3658; 

Middle Park formation, 337. 

APOCYN ACEAE 

Apocynophyllum lesquereuxi Ettingshausen 

Apocynophyllum lesque1·e1tW'i Ettingshausen, 1883, · p. 132. 

Knowlton, 1917, p. 345. 

Apocynoplt.yllum linijoUum Knowlton, 1917, p. 346, pl. 104, fig. 3. 

Apocynophyllum wilcowensis Berry. Knowlton, 1917, p. 345, 

pl. 103, fig. 3 ; pl. 105, figs. 1, 2 ; pl. 106, fig. 1. 

Quermts 1W1'i'ifoUa Brown. Lesquereux, 1878, p. 150, pl. 19, 

fig. 5 only. 

The published illustrations of these leaves are for 

the most part unsatisfactory and misleading. In actuality the secondary veins are numerous and fairly 

closely spaced but are unequally parallel, and with 

the intersecondaries, are quite irregular and variously 

branched or interconnected, and are united to form 

an intramarginal vein. As remarked by nearly all 

students dealing with these leaves, their affinity is 

more probably with the Myrtaceae than with the Apocynaceae, but, lacking suggestive fruits as evidence, 

doubt about their correct allocation will continue to 

be entertained. 

Occurrence: Raton formation, 5611, 5684, 5701, 5798. 

CAPRIFOLIACEAE 

Viburnum antiquum (Newberry) Hollick 

Plate 63, figures 1-8; plate 64, figure 4 

Tilia (tnt,iqlzut Newberry, 1868, p. 52. 

Viburn1t1n antiquu.m~ (Newberry) Hollick. Newberry, 1898, 

p. 128, pl. 33, figs. 1, 2. 

Berry, 1935, p. 58, pl. 17, fig. 3 only. 

Vibtwn;um betulaefolium Ward, 1887, p. 114, pl. 57, fig. 4. 

Viburnum dakotense Lesquereux, 1883, p. 231, pl. 46A, fig. 9. 

Viburnum dentoni Lesquereux, 1883, p. 231, pl. 49, figs. 2, 3. 

Viburnu,m elongaf;ium Ward, 1887, p. 112, pl. 54, figs. 4, 5. 

Viburnum limpidum Ward, 1887, p. 110, pl. 53, figs. 3-6. 

Viburnum mncrodontum Ward, 1887, p. 110, pl. 53, fig. 2. 

Viburnum nordenskioldi Hee1·. Lesquereux, 1883, p. 230, pl. 49, 

fig. 4. 

Viburnum oppo.siUne1·ve Ward, 1887, p. 112, pl. 55, figs. 1, 2. 

Viburnum perfectum Warcl, 1887, p. 109, pl. 52, figs. 3, 4; 

pl. 53, fig. 1. 

Vib1tr?Vum perplewum Ward, 1887, p. 111, pl. 54, figs. 2, 3. 

Viburnum whymper'i Heer, 1869, p. 475,. pl. Mi, fig. lb. 

Ward, 1887, p. 111, pl. 54, fig. 1. 

Alnus serrata Newberry. Bell, 1949, p. 52, pl. 33, fig. 6. 

Co-ryl1ts macquarrii (Forbes) Heer. Lesquereux, 1883, p. 223, 

pl. 49, fig. 4. 

Diospyros ficoidea Lesquereux. Ward, 1887, p. 105, pl. 49, 

fig. 4 only. 

Ficus viburnifolia Ward, 1887, p. 42, pl. 22, figs. 4-8. 

G1·ewia obovnta Heer. Ward, 1887, p. 88, pl. 39, fig. 7 only. 

H edera pnrvula Ward, 1887, p. 57, pl. 26, fig. 4. 

Monhnio]J8is tmterna Saporta. Ward, 1887, p. 52, pl. 25, fig. :l. 

Popul1ts grewiop.sis Ward, 1887, p. 23, pl. 9, fig. 1. 

In this synonymy I have grouped those viburnumlike leaves having a broad-ovate, elliptic, or cordate 

outline; a margin with large, rounded, dentate to serrate teeth; and secondaries widely spaced, somewhat 

irregular and undulating and conspicuously branched. 

Exceptions to this general statement appear to be 

Diosypros ficoidea, Ficus viburn.ifolia, and H edera 

parvula. These I regard as extreme or abnormal variants from the general average. Diosypros ficoidea was 

figured as an entire-margined leaf. The type specimen, however, is not as perfect as the sketch, much of 

the margin being missing, but on that which is preserved, undulations may be seen which suggest rudimentary teeth. Fic'ltS vibttrnifolut (Ward's fig. 8) has 

an entire margin and resembles Il edera parvula. Both 

see1n to me to be abnormal or malformed leaves. The 

normal leaves of V. antiquu1n find matches among 

those of several living species. 

The seeds found at locality 4264 in association with 

typical leaves of this species appear to be 1nore clearly 

referable to V iburnun~ than those identified by Ward 

as V. tilioides. These seeds were flattened and had 

four longitudinal furrows. 

Occurrence: Fort Union formation (upper), 2414 

(pl. 63, figs. 1-7), 2416, 4264, 4974, 8166 (pl. 63, fig. 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 87 

8)' 8230, 8255, 8523, 8774, 9071 (pl. 64, fig. 4)' 9125; 

Ferris formation, 6971. 

Viburnum asperum Newberry 

Plate 64, figures 1-3, 15, 7-11 

Vibu·rnum. aspe?ium Newberry, 1868, p. 54; 1898, p: 129, pl. 33, 

fig. 9. 

Ward, 1887, p. 113, pl. 55, figs. 4-9. 

Berry, 1935, p. 56, pl. 16, figs. 1-4. 

Bell, 1949, p. 79, pl. 62, fig. 2; pl. 67, figs. 4, 6. 

Vib1trn1un ca.st·rae Knowlton and Cockerell. . Knowlton, 1919, 

p. 641, f.or V. lanceolatum Newberry, 1898, p. 131, pl. 33, 

fig. 10. 

Berry, 1935, p. 56, pl. 15, fig. 5. 

Vib1t?'1W?n e1·ect1tm Ward, 1887, p. 112, pl. 55, fig. 3. 

Vibun~um newberrianmn Ward, 1887, p. 113, pl. 56, figs. 1-6. 

Vib1trnmn no1·denskiold·i fleer. Ward, 1887, p. 114, pl. 57, figs. 

1-3. 

Vibm·nmn antiquum rout. trine1·v;mn Berry, 1935, p. 60, pl. 7, 

fig. 6 ; pl. 15, fig. 6. 

Alnns. nostmt1tm Unger. fleer, 1868, p. 103, pl. 47, figs. 12a, b. 

A·melanchie1· si?n·iUs Newberry, 1898, p. 111, pl. 40, fig. 6. 

C01·nus denve1·ensis Knowlton. Bell, 1949, p. 76, pl. 56, fig. 5 

[part.] 

Fimts li.?npida Ward, 1887, p. 42, pl. 22, fig. 3. 

Rham.nUes condnmts Newberry, 1898, p. 118, pl. 33, figs. 7, 8. 

Rham.nu.<J gold·ianns Le~quereux. Knowlton, 1917, p. 332, pl. 

112, fig. 5. 

Tilia weed·i Knowlton, 1902, p. 706, text fig. 1. 

UZ1n11..<; p·ra.eC'I.t1'S01' Dawson, 1886, p. 28, pl. 2, fig. 11. 

The striking features of these leaves are the remarkable smoothness and grace in the curvature of 

the many secondary veins; the numerous closely 

spaced connecting veinlets; the regular, rounded to 

pointed, crenate-serrate teeth; the tendency to have 

strong basal secondaries with many branches; and the 

lanceolate-pointed shape with rounded or cordate 

sometimes slightly asymmetric base. 

The leaf called An~ela.nchier simili.s Newberry is apparently somewhat abnormal, and this, together with 

the fact that it was incorrectly illustrated in critical 

details, almost masks its relationship to Viburnrum 

aszJerunu. Its venational and marginal features, however, on close examination are distinctive. The leaf 

described as Tilia weedi ICnowlton (fig. 11) is an unusually large example of V. asperun1. 

These leaves resmnble those of the hamamelidaceous species Oorylopsis sinensis Hemsley, but particularly those of the cornaceous Davidia involucrata 

Baillon, of western China. 

The species, with other associates, is a fairly reliable index for the Paleocene of the Rocky Mountains 

and Great Plains. At only one Upper Cretaceous locality, about 4 miles east of Reed Point, Mont., have I 

found probable ancestral specimens (pl. 64, fig. 6) that 

simulate the leaves of this species, except that they 

are smaller and somewhat more asymmetrical. 

Occurrence: Fort· Union formation (lower) 8519, 

8558, 8654, 8673, 8678 (figs. 7, 8), 9056 (figs. 9, 10) ; 

(upper), 607 (fig. 11), 2417 (figs. 1, 3), 2422, 4871, 

5194,5882,6377,8166,8192,8213,8222,8520 (fig. 5), 

8885 (fig. 2)' 8921, 8922, 9322. 

Viburnum cupanioides (Newberry) Brown, n. comb. 

Plate 46, figure 1; plate 57, figure 1; plate 65, figures 1-8 

Phyllites cu.panioides Newberry, 1868, p. 74; 1898, p. 135, pl. 41, 

figs. 3, 4. 

Pte'l'ospennites cupanioides (Newberry) Knowlton, 1893, p. 35, 

pl. 2. 

Celtis rugosa Newberry, 1883, p. 510. 

Celtis lingu.alis Knowlton and Cockerell. Knowlton, 1919, 

p. 160. 

Phyllites cameosus Newberry, 1898, p. 134, pl. 41, figs. 1, 2. 

Phyllites carneosa (Newberry) Bell, 1949, p. 55, pl. 35, figs. 

1-3 ; pl. 36, figs. 1-6. 

Phyllites venows Newberry, 1898, p. 136, pl. 30, fig. 4. 

Pterospermites minor Ward, 1887, p. 95, pl. 42, figs. 1-3. 

Berry, 1935, p. 48, pl. 11, fig. 4. 

Pterospermites whitei Ward, 1887, p. 94, pl. 41, figs. 5, 6. 

Bell, 1949, p. 69, pl. 4, fig.' 3. 

Rhmnnites nw1·ginatus (Lesquereux) Bell, 1949, p. 73, pl. 61, 

fig. 5. 

Sctpindu.s? membmnaceu.s Newberry, 1878, p. 117, pl. 30, fig. 3 

only. 

Viburnum pa'ltcidentatum Newberry, 1883, p. 511. 

The general outline, marginal dentition, and venation of these leaves can be matched most readily in 

V iburnu1n. Indeed, several specimens assigned to this 

species might serve as connecting links with Viburnum 

antiquu1n (Newberry) Hollick. 

The ~leaf described by Newberry as Oeltis rugosa, 

renan1ed 0. lingualis by Knowlton and Cockerell, and 

that called Viburnu1n paucidentatum were never illustrated or again referred to by Newberry; nor is the 

present location of the specimens known. The descriptions, however, suggest their assignment to this species. 

Occurrence: Fort Union formation (lower) 4514, 

4625 (pl. 65, fig. 6)' 5885 (pl. 57, fig. 1)' 5889 (pl. 65, 

figs. 5, 7), 6050 (pl. 65, fig. 2), 8519, 8521, 8627, exact 

locality unknown (pl. 65, fig. 1, reproduction of Newberry, 1898, pl. 30, fig. 4); (upper), 2420 (pl. 65, figs. 

3,4), 5324,8190 (pl. 46, fig. 1), 8774,8910,8920, 9322; 

Paskapoo formation, Alberta, Canada. 

Viburnum tilioides Ward 

Plate 67, figures 18, 19, 24, 25, 30, 31 

Vibiu1·num tilioides Ward, 1887, p. 107, pl. 51, figs. 4-8 [not 

other figures, which are V. ant'iqmun (Newberry) Hollick]. 

Carpites S'ltlcatu.s Knowlton, 1924, p. 96, pl. 19, figs. 3, 4. 

As at least two authentic species of Viburnum and 

one species of Nyssa are represented by leaves from 

the same strata at or near the same locality that 

yielded the types of these seeds, there is reasonable 
88 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

doubt concerning the relationship of the leaves and 

seeds. The seeds may not even be those of V iburnwm 

or Nyssa. Some may well be mistaken for those of 

Trapa angulata (Newberry) Brown. Others bear considerable resemblance to the cornaceous seeds assigned 

by IGrchheimer. to 111 astimia, from the brown coal deposits of Germany. 

Occurrence: Fort Union formation (lower) 4626, 

5509, 5619, 8519, 8554 (fig. 25); (upper), 2414 (fig. 

19), 2424, 8196 (figs. 18, 24), 8225 (fig. 31), 8774 (fig. 

30); Animas formation, 7496. 

OBJECTS OF UNCERTAIN CLASSIFICATION 

Macclintockia kanei (Heer) Seward and Conway 

Macclintockia kanei (Heer) Seward and Conway, 1935, p. 24, 

pl. 2, figs. 16, 17, 20, text fig. 9. 

Cooculus kanii (Heer) Saporta and Marion, 1873, p. 63, pl. 10, 

fig. 1. 

Daphnogene kanii Heer, 1868, p. 112, pl. 14; pl. 16, fig. 1. 

Saporta and Marion, 1873, p. 65, pl. 10, figs. 2, 3. 

Macclintockia dentata Heer, 1868, p. 115, figs. 3, 4. 

Macclintockia lyalUi Heer, 1868, p. 115, pl. 15, figs. 1a., 2; pl. 16, 

figs. 7a, 7b; pl. 17, figs. 2a, 2b; pl. 47, fig. 13; pl. 48, 

fig. 8. 

Macclintockia trinm·vis Heer, 1868, p. 115, pl. 15, figs. 7-9; 

1883, p. 95, pl. 77, fig. 8. 

The spelling kanei for this species is here used as 

being more appropriate to the dedicatee, Dr. Elisha 

Kane. 

Heer, in my opinion, was not justified in defining 

three species of dentate-leaved 11! cwclintookia from the 

same locality; and for defining entire leaves, having 

the same texture and internal architecture as the dentate leaves, as a species in a different genus and family. All these leaves appear to be variants of a single 

species. 

No authentic Paleocene 11! aoolintookia foliage has 

been reported from the mainland of North America. 

Daphnogene kanii Heer, report.ed from the Eocene of 

Alaska by Hollick (1936, p. 121, pl. 67, fig. 11), although somewhat like Heer's types, differs by having 

its pair of primary veins arise from the midrib at a 

considerable distance above the top of the petiole. 

Similar leaves occur in the Eocene lignitic strata of 

'V ashington. The 0 oooulus kanii specimen reported 

by Berry (1926, p. 111, pl. 16, fig. 2) from the Eocene 

at l{itsilano, British Columbia, appears to have the 

form of 111 aoolintookia kanei, but venational details 

are missing, so that no definite conclusion can be 

drawn. 

Heer, on the basis of fragments, reported two species of J,f aoolintookia from the Upper Cretaceous of 

Greenland at Igdlokunguak on Disko Island: M. oretacea (1880, p. 70, pl. 36, figs. 1, 2a; pl. 37, figs. 2-4), 

and M. appendioulata (1880, p. 71, pl. 37, fig. 1). If 

these fragments, as illustrated, are authentic, they may 

represent forerunners of the Paleocene M. kanei. 

111 aocl'intockia kanei differs little from 11/. heersiensis Sa porta and Marion ( 1878, p. 55, pl. 9, fig. 1) from 

the Paleocene at Gelinden, Belgium. 

No satisfactory conclusion has been reached about 

the botanic classification of this species. Comparisons 

often made with Ooooulus laurifolius DeCandolle, 

though suggestive, are not completely convincing .. 

Occurrence: Atanekerdluk, Greenland. 

Phyllites demoresi Brown, n. sp. 

Plate 61, figures 5-8 

Leaves ovate to ovate lanceolate, with rounded bases 

and acuminate apexes. Margin entire. Petiole slendder, more than 3 em long. Venation pinnate, with 

secondaries widely spaced, forming conspicuous, closed 

loops well within the margin. Intersecondaries common. 

Named for the Marquis DeMores, who founded the 

town of Medora, N. Dak. 

Occurrence: Fort Union formation (upper), 4264 

(figs. 5, 6, ~), 4879,4892, 8196 (fig. 7). 

Phyllites disturbans Brown, n. sp. 

Plate 60, figures 5, 7-10 

Leaves elliptic, with acute apexes and cuneate bases. 

Secondary veins evenly spaced, camptodrome. Margin entire or very obscurely, distantly, and minutely 

toothed. 

Occurrence: Fort Union formation (upper) 4262 

(fig. 7), 6342 (fig. 8), 7659 (fig. 5), 8563 (figs. 9, 10). 

Phyllites pagosensis Knowlton 

Plate 18, figure 10 

Phyllites pnfJosensis Knowlton, 1924, p. 97, pl. 16, fig. 2. 

Ul1nus sp. Knowlton, 1917, p. 300, pl. 70, fig. 4. 

The leaf fragments here cited and figured seem to 

be closely related morphologically. Their margins 

have large double teeth, and their slightly arched 

secondary veins enter the major marginal teeth. 

Branchlets from the secondaries enter the subsidiary 

teeth. 

Ulmaceous affinities may be indicated here, but some 

rosaceous genera also have leaves displaying somewhat 

similar features. Somewhat comparable leaves called 

Ulmlu8 sorbifolia G<:>ppert (Hollick, 1936, p. 106, pl. 

57, figs. 3-5) occur in the Tertiary floras of Alaska. 

Occurrence: Animas formation, 7 496; Denver formation, 317 (fig. 10) ; Raton formation, 5695. 

·Calycites hexaphylla Lesquereux 

Plate 4, figure 11 

Cnlycites hexaphylla Lesquereux, 1873, p. 402. 
SYSTEMATIC DESCRIP'riON OF THE MEGASCOPIC FLORA 89 

D·iospy1·os hercaphylla (Lesquereux) Knowlton, 1919, p. 238. 

D·iospy·ros wodani Unger. Lesquereux, 1878, p. 233, pl. 59, 

fig. 13. 

The lobes of Oalycites hexaphylla differ frmn those 

of 0. polysepala Newberry in being longer, slightly 

spatulate, and finely striate. The unlobed portion is 

smaller and has a central, crater like depression. No 

remains of stamens or pistils are present. 

Lesquereux's figured specimen of Diospyros wodani 

fortunately retaii1s a long peduncle. Excavation of 

the upper part of the peduncle and the bases of the six 

lobes shows a true six-parted calyx underneath, 8 mm 

in diameter, and having short pointed lobes that alternate with those of the "corolla" above. The specimens, therefore, are gamopetalous flowers, with the 6 

petals and 6 sepals, respectively, united near their 

bases. The affinity of these flowers is uncertain but 

may be with the Lauraceae, species of which are represented by leaves in the Paleocene flora. 

Occurrence: Fort Union formation (lower), 9248 

(fig. 11), 9249; Evanston formation, 1471. 

Calycites polysepala Newberry 

Plate· 4, figure 5 

Oalycites polysepala Newberry, 1898, p. 139, pl. 40, fig. 3. 

Newberry's type has seven lobes. Other specimens 

in the collection have six, seven, and eight lobes. The 

sinuses between the lobes are rounded, not sharply 

angular, but there are no discernible definite markings 

on the lobes. The unlobed central portion of the 

specimens shows a broad, shallow, saucerlike depression, with no apparent diagnostic features. No peduncle has been seen but a central pit or scar may 

represent the point of attachment of a peduncle. 

Superficially, except for the rounded sinuses, these 

specimens resemble · species of earthstar puffballs 

( Geaster) somewhat like the doubtful Geaster florissCflntemis Cockerell from Oligocene lakebeds at Florissant, Colo. Smaller specimens bearing the names 

Antholithes' brottoni Berry (1930, p. 77, pl. 15, fig. 3) 

and A. frernontensis Berry (1930, p. 78, pl. 15, fig. 2), 

which are clearly synonymous, are known from the 

middle Eocene of the Wind River Basin in Wyoming. Other and more suggestive comparisons could 

perhaps be made, but the true affinity of these "calyces" remains for the present unknown. 

Occurrence: Fort Union, exact locality unknown (fig. 

5). One specimen from an unspecified locality near 

Dickinson, N. Dak., was collected by R. L. Coville. 

Carpolithes spinosus Newberry 

Oa1·polithes spinostts Newberry, 1898, p. 138, pl. 68, figs. 2, 3. 

Occurrence: ~Raton formation, North Fork of the 

Purgatoire River, Colo. 

N ordenskioldia borealis Heer 

Pia te 67, figures 13, 45 

Nordenskioldia bo1·eaUs Heer, 1871, p. 65, pl. 7, figs. 1-13; 1880, 

p. 13, pl. 6, fig. 8. 

Bell, 1949, p. 80, pl. 18, fig. 2 ; pl. 21, fig. 3. 

Ca.rpites cocculoides (Heer) Schimpei'. Lesquereux, 1878, 

figs. 32-35. 

Oarpites coccttloides var. major Lesquereux, 1878, p. 307, pl. 60, 

p. 307, pJ. 60, figs. 38, 39. 

CarpolUhes cocculoides Heer, 1871, p. 484, pl. 52, figs. 9, 9b. 

Diospyros bmchysepala Alexander Braun. Heer, 1868, p. 117, 

pl. 47, figs. 4b, 5, 5f, 5g. 

The surface markings of these lunate impressions 

are veinlike, diagonal, and forked. Thus, there is a 

superficial resemblance between N ordemkioldia and 

the capsules of Oercidiphyllum, and inasmuch as both 

are found together at many localities, there is likelihood of confusion if the specimens are not carefully 

examined. In outline, N ordemkioldia impressions resemble seeds of persimmon, Diosypros, with one side 

practically straight and the remainder of the border 

arched. The seeds of N ordenskioldia, however, were 

packed around a central axis like those of Malva, and 

the fruits were sessile, alternate, and opposite (fig. 

45) , at intervals on a long rachis. There seems as yet 

no inkling as to the correct botanical affinity of 

N ordenskioldia. 

Occurrence: Fort Union formation (lower), 4661, 

5509, 5918, 8240, 8654, 9130; (upper) 2420, 4870, 4892, 

5618, 8167, 8234, 8236, 8887 (fig. 45)' 8910, 8913 (fig. 

13), 8917, 9125; Co~lmont formation, 6110. 

Palmocarpon commune Lesquereux 

Palmocarpon commune Lesquereux, 1878, p. 119, pl. 13, figs. 4-7. 

Palmocarpon mexicanmn (Lesquet·eux) Lesquereux, 1878, p. 

119, pl. 11, fig. 5. 

The identifying feature of these roundish flattened 

seeds is the somewhat narrow elliptic scar (perhaps a 

raphe) with a nipple in the middle of the broader 

end, well illustrated in Lesquereux's plate 13, figure 7. 

What species of living palm, if any, produces seeds of 

this kind~ 

Occurrence: Denver formation, 317; Raton formation, exact locality uncertain. 

Palmocarpon compositum Lesquereux 

Palnwca1·pon composit'lttn Lesquereux, 1878, p. 119, pl. 11, fig. 4. 

Euphorbocarpum, richardsoni Knowlton, 1917, p. 328, pl. 96, 

figs. 3, 4. 

The assignment of these specimens to either the 

Palmaceae or Euphorbiaceae is unconfirmed by convincing evidence. 

Occurrence: Raton formation, 5046, and an uncertain 

locality. 
90 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Palmocarpon lineatum Lesquereux 

Palmoca1·pon Uneatmn Lesquereux, 1888, p. 44. 

Knowlton, 1930, p. 42, pl. 11, fig. 5. 

Assignment of these seeds to. the Palmaceae is 

dubious. 

Occurrence: Denver formation, 317. 

Palmocarpon subcylindricum Lesquereux 

Plate 67, figure 37 

Palmocarpon s7tbcyUnd'l"icmn Lesquereux, 1878, p. 121, pl. 11, 

fig. 12. 

Knowlton, 1930, pl. 52, fig. 3b. 

Lesquereux said of his specimens that they were 

split in two at the top. Careful excavation of these 

objects, however, shows that the tops are three-parted 

and that the bottoms have a well-defined roundish 

attachment scar. J(nowlton's plate 52, figure 3b, illustrates a specimen with a broad cuplike base or a 

thick peduncle. Something about these objects suggests that they are empty calyces of some flower. 

That they belonged to palms seems doubtful. 

Occurrence: Fort Union formation, 4901, 5918; 

Denver formation, 317 (fig. 37). 

Palmocarpon truncatum Lesquereux 

Palmocarpon tnmcah11n Lesquereux, 1878, p. 120, pl. 11, figs. 

6-9. 

Gink{Jo? tnmoata (Lesquereux) Knowlton, 1930, p. 34, pl. 9, 

fig. 3. 

The long peduncle (see J(nowlton's pl. 9, fig. 3) of 

these specimens seems to rule out assignment to the 

palms; and the "fleshy cup" referred to by J(nowlton 

seems on examination of the specimen itself to be a 

remnant of a once larger enclosing sheath or calyx, 

thus casting doubt on the assignment to Ginkgo. Moreover, no foliage attributable to Ginlcgo has yet been 

found at the plant localities in the Denver formation 

on South Table Mountain near Golden, Colo. It is 

probable that these objects belong with Palmocarporn 

subcylindricum.- Lesquereux. 

Occurrence: Denver formation, 317, 325. 

Viburnum goldianum Lesquereux 

Viburmt,m gold·iam.nn Lesquerenx, 1878, p. 227, pl. 60, figs. 2, 

2a-c. 

Nothing new has been learned about the true identity 

of these seeds, but reference o:f them to Viburnum is 

questionable. 

Occurrence: Denver :formation, 317. 

Viburnum solitarium Lesquereux 

Plate G7, figures 33, 34 

Vib7trmun wlitari'lt'ln Lesquereux, 1878, p. 227, pl. 60, fig. 3. 

Carpites 1'1tomboidalis Lesquereux, 1878, p. 306, pl. 60, figs. 28, 

29. 

These small seeds had ovate kernels, apparently enclosed in a bivalved shell whose suture in fossilization 

made a 'vide flange, squarish at the top. The assignment to Yiburnun~ is dubious. 

Occurrence: Denver formation, 317 (figs. 33, 34); 

Laramie :formation (Upper Cretaceous) :for 0. rhomboidalis. 

Ophiomorpha nodosa Lundgren 

Ophimnorpha nodosa Lundgren. Hantzschel, 1952, p. 148, pis. 

13, 14. [See synonymy and discussion.] 

Ralymenites 1najo1· Lesquereux. Brown •. 1939b, p. 253, pl. 62, 

figs. 1-7; pl. 63, figs. 1, 2. 

Thus far the only specimens o:f this kind :found in 

the Paleocene o:f the Rocky ~fountains and Great 

Plains "·ere taken :from the sandy :facies o:f the Cannonball member o:f the Fort Union :formation by T. ,V. 

Stanton in 1913 in the right bank o:f Heart River, onehalf a mile south of l\1andan, N. Dak. They are 

:fragmentary but characteristic. In Europe, 'V atelet 

( 1866, p. 24, pl. 4, fig. 1) reported a specimen :from 

the upper Paleocene o:f the Paris Basin, which he 

called Phyn1a.toder1na dienvalii, now synonymized by 

H~intzschel, with Ophiomorpha nodosa. 

As stated in my study of H aly1nenites n1ajor in 1939 

and by Hantzschel in his review o:f Ophiomorpha in 

1952, there is strong evidence that this supposed alga 

represents the burrow o:f a marine animal, probably a 

crustacean. As yet, however, no remains o:f the purported burrmvers themselves have been :found in or 

directly connected with the burrows. Field geologists and collectors should keep this problem in mind 

and be on the lookout :for such conclusive evidence. 

Occurrence: Fort Union :formation (lower), 6525. 

MISCELLANEOUS LEAVES, FRUITS, SEEDS, FLOWERS, 

ROOTS, AND PRPBABLE ANIMAL REMAINS 

Plate 57, figure 3 

Leaves large, ovate, with cuneate bases and rounded 

or blunt apexes. ~Iargin with minute scattered sharppointed teeth. Venation pinnate, the secondaries 

evenly spaced, camptodrome. Intersecondaries common. V einlets connecting the secondaries subparallel, 

oblique. Fort Union formation, 8774. 

Plate 66, figure 4 

Palmately veined leaf, with cordate base, and a :few 

coarse marginal teeth. Fort Union :formation, 8920. 

Plate 66, figure 5 

Narrow elliptic leaves or leaflets with undulate to 

toothed margins, but teeth :few, blunt, and serrate. 

Venation obscure, but apparently palmate wit~1 the 

veins extending well toward the apex. These may 
SYSTEMATIC DESCRIPTION OF THE MEGASCOPIC FLORA 91 

be leaflets of TTapa angulata. (Newberry) Brown. 

Fort Union formation, 6215. 

Plate 67, figure 5 

A five-loculed fruit, pentagonal in outline, simulating the seed case of an apple after the flesh has 

been removed. A somewhat similar fruit was described by Reid and Chandler (1933, p. 469, pl. 26, 

figs. 31-38) as Sapoticarpun1 latum, from the Eocene 

London clay. Fort Union formation, 8247. 

Plate 67, figures (), 7 

vVinged, maplelike samaras, but with long peduncles, 

thus suggesting some genera in the soapberry family, 

Sapindaceae. · Fort Union formation, 9125. 

Plate 67, figures 9, 14, 15, 16, 20, 22, 23, 27 

Ovate seeds with a thin shell or skin whose outer 

surface was fairly smooth but whose inner surface 

was marked by closely packed papillae, squarish protuberances, or cavities as the case may be. Seeds of 

Actinidia might leave such impressions. Fort Union 

formation, 4031, 4320, 4896, 4901, 5438, 6215 (figs. 15, 

22, 23, 27), 8519, 8550 (figs. 9, 14) 8910, 9109, 9236 

(figs. 16, 20); Denver formation, 317. 

Plate 67, figure 10 

A flattened carbonized specimen, elliptical 111 outline and showing irregular angular divisions as though 

it represented a multiple fruit. The surface of each 

division is marked by two overlapping depressions, 

one smooth ~nd shallow, the other with a central pit. 

'Vhether this object was originally round before being flattened is unknown. Fort Union formation, 6382. 

Plate G7, figures 11, 12, 17 

Narrow w1~inkled specimens, blunt rounded at one 

end and squarish, fringed at the other. Possibly a 

fruit, topped by the remnants of a calyx. Fort Union 

formation, 4626 (fig. 11), 8547 (fig. 17), 8566 (fig. 12), 

8917. 

Plate 67, figure 21 

A spike or catkin of ·seedlike bodies that are somewhat lunate in shape 'vith two or three concentric 

folds. As these catkins were found with matted palm 

leaves, there is some suspicion that they may have 

been parts of a flowering palm spadix. Fort Union 

formation, 8519 (fig. 21) ; Evanston formation, exact 

locality unknown. 

Plate 67, figure 2G 

Ovate seeds with longitudinal rows of pits. Fort 

Union formatimi, 4618 (fig. 26), 8551. 

Plate G7, figures 28, 29 

Oval seeds with five or more conspicuous longitudinal ridges and furrows with scattered glandlike 

marks on the surface of the furrows. Fort Union 

formation, 4264. 

Plate 67, figure 32 

Flat heart-shaped apparently two-lobed fruits, occurring in clusters. Fort Union formation, 8519 (fig. 

32)' 8775. 

Plate m; figure 35 

An elliptic flat object, divided into two equal halves 

by a median, longitudinal line. Numerous fine hairs 

radiate from the margin. The specimen is somewhat 

like the Upper Cretaceous Carpolithus hirsutus Newberry. Fort Union formation, 4977~ 8522 (fig. 35). 

Plate G7, figure 36 

An oval fruit, which, according to the longitudinal 

ridges, may have been multilocular. Fort Union formation, 8167. 

Plate 67, figure 38 

Oval seeds with wide flanges, rounded at both ends. 

Animas formation, 5667. 

Plate Ei7, figures 39-42, 47 

These are apparently burs, with a few relatively 

stout, sometimes branched, sharp-pointed spines. In 

figure 4 7 the burs are definitely associated with round 

fruitlike bodies. They may be the fruits of a sandburgrass, Oenchr-us. Fort Union formation, 2427, 7538 

(figs. 39-42), 9109 (fig. 47). 

Plate G7, figure 43 

An elliptic body covered with roundish projections. 

This object may be the fertile cone of a species of 

E quisetun~;. Fort Union formation, 6905. 

Plate Ei7, figure 44 

This fragment appears to have been a cylindrical 

object originally, and is now studded with bilobed 

projections. Fort Union formation, 9130. 

Plate Ei7, figure 46 

Small calyces, with five blunt slightly unequal lobes. 

These resemble the calyces and corollas of elderberry, 

Sa1n-bucus. Fort Union formation, 5194, 8774 (fig. 

46). 

Plate G8, figures 2, 7-11 

Roots, with rootlet scar pits in three or more longitudinal rows. Specimens have been taken from nearly 

every formation from the Upper Cretaceous to the 

Eocene, but to what plant they belonged is unknown. 

Fort Union formation (lower), 5614; (upper) 8668 

(fig. 10); Animas formation, 5458, 5461, 8669 (fig. 2); 

Ferris formation, 3963; Livingston formation 6765 

(fig. 11) ; lVIiddle Park formation, 337; Raton forma-
92 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

tion, 5129. Upper Cretaceous, Fruitland formation, 

4468 (fig. 7); Eocene, ""\Vind River formation, 9533 

(fig. 9) ; Fossil Forest Ridge, Yellowstone .National 

Park, 750 (fig. 8). 

Pia te 68, figures 17-22 

Two-winged seeds, simulating those of catalpa and 

other Bignoniaceae. A seed similar to these was described for Gop pert by ""\Veber ( 1851, p. 233, pl. 25, 

fig. 5) and called Dipterospermum bignonioides, from 

Oligocene lignitic strata near Bonn in the Rhineland. 

Reid and Chandler ( 1926, p. 129, pl. 8, fig. 20) also 

described a similar seed, Raderrnachera pulchra, from 

the Bembridge beds (Oligocene) of the Isle of Wight. 

Fort Union formation, 8910. 

Plate 68, figure 25 

This small roundish square-beaked seed is probably 

a stone-fruit of some kind. Fort Union formation, 8913. 

Plate 68, figure 26 

This boat-shaped object resembles the glumes or 

bracts of some grasses. Fort Union formation, 8913. 

Plate 68, figures 27-29 

Small pedicelled fruit or flowerlike objects. Fort 

Union formation, 5595 (fig. 29), 9109 (fig. 27), 9125 

(fig. 28). 

Plate 69, figure 1 

Feather of an unknown bird. This feather is preserved in the pinkish baked shale from above a burned 

coal bed about 10 miles north 'of Glendive, ~1ont. It 

was found about 1930 by John S. Larimer, former 

jeweler at Glendive, and given to me by his son in 

1950. Bird feathers and down have sometimes been 

mistaken for plant remains, as for example Lesquereux's supposed moss, Fontinalis pristina (1883, 

p. 135, pl. 21, fig. 9), from Oligocene lake beds at 

Florissant, Colo., and Cziffery-Szilagyi's Donacites 

erdobenyensis (1955, p. 164, fig. 7, on p. 31) from the 

Miocene of Hungary. 

Plate 69, figures 15-19 

Ropy striated concretions blunt pointed at both 

ends when perfect. These are found with others that 

are more irregular, warty, and without striations. 

Figures 16-19 represent specimens from a clayey, silty 

stratum in the Fort Union formation, 8 miles south of 

Rhame, N. Dak. They are composed of limonite and 

are rusty brown on the surface. That they were 

chiefly siderite before they were oxidized to limonite 

is inferred from the fact that the comparable specimen, figure 15, from Miocene strata (Roberts, 1958, 

p. 35) on Cedar Creek, a tributary of Salmon Creek, 

Wash., is still largely siderite, without discernible 

organic matter, like most of its kind at that locality. 

These come from a bluish clay containing some woody 

material and, locally, well-preserved leaves and seeds. 

Speculation has ranged widely in attempts to account 

for these objects, but the most persistent hypothesis, 

based on their shape and their occurrence in fluvial 

deposits, is that they are coprolites, perhaps of large 

fishes, turtles or other reptiles. 

OTHER OBJECTS OF UNCERTAIN OR MISTAKEN 

IDENTITY 

Besides the foregoing illustrated objects of uncertain though most likely of organic affinity, the Paleocene collections contain many other seeds, fruits, flowers, and problematica not here described or illustrated. Many of these are not well preserved, are not 

good subjects for photography, and, although exciting curiosity and stimulating much thought without 

definite classificatory results, are here deemed best to 

remain temporarily nameless. 

Some objects hitherto thought to be seeds, fruits, or 

other parts of plants, are now known to be of inorganic origin. Thus, for example, Ward (1887, p. 14, 

pl. 1, fig. 3) described a specimen he identified as an 

alga, Spirraxis bivalvis. This is a concretion, showing 

the original bedding planes of the sandstone that gave 

Ward the mistaken impression that these were spirally 

alined. On revisiting the locality I found similar 

specimens and verified their identity as concretions. 

Stainbrook ( 1924) described an object .he thought 

might be a fossil fruit, from Chimney Butte in SWlh, 

sec. 29, T. 139 N., R. 102 W., North Dakota. At this 

locality on September 7, 1949, I found numerous similar specimens. All are concretions, the centers of 

which in many instances are somewhat septarian thus 

giving the deceptive appearance of being locular fruits. 

THE MICROSCOPIC PLANTS 

Although plant microfossils from Paleozoic coal 

seams and associated strata have received considerable attention, those from Mesozoic and Cenozoic strata 

. have, until recently, been more or less neglected. Led 

by the investigation of the pollens and spores in Pleistocene and Recent peat bogs and the microfossils of 

the middle Eocene Green River formation, the trend 

now seems strong toward· a comprehensive study of 

the Mesozoic and Cenozoic microscopic flora. E. L. 

Miner ( 1935) found numerous spores in the Cretaceous coals of Montana and one species, Selaginellites 

mirabilis (now Triletes mirabilis (Miner) Schopf, Wilson, Bentall), in the Paleocene coal at the Homer 

mine, Bear Creek, Mont. Subsequently, Wilson and 

Webster ( 1946) reported the following species from 
FOSSIL WOOD 93 

the IColarich coal mine (Paleocene) in sec. 25, T. 7 S., 

R. 20 E., ~ miles east of Red Lodge, Mont.: 

Alnus specUpites Woclehouse. Wilson and Webster (1946, 

p. 275, figs. 10, 11). 

Bet•ula, cla1·ipites Woclehouse. Wilson and Webster (p. 275, 

fig. 12). 

Bmchysporimn sp. Wilson and Webster (p. 271, fig. 1). 

Ccwya ve1·ipUes Wilson ancl Webster (p. 276, fig. 14). 

Cm·ya viridi/l1tminipUes (Woclehouse) Wilson and Webster 

(p. 276, fig. 13). 

Cycadop-ites follic1tlcwis Wilson and Webster (p. 274, fig. 7). 

Deltoidospo1·a, dlaphana Wilson and Webster ( p. 273, fig. 3). 

Laevigatospo1·ites gmciUs Wilson and Webster ( p. 273, fig. 4). 

Laevigatospo1·ites ovatus ·wilson and Webster ( p. 273, fig. 5). 

Momipites co1·ylo·ides Wodehouse. Wilson ancl Webster (p. 275, 

fig. 15). 

P.icea gmndivesc·ipites 'Vodehouse. Wilson and ·webster 

(p. 275, fig. 8). 

Querc1ts p1·iscipites Wilson ancl Webster (p. 276, fig. 16). 

Seq1wia la.1Jill-i1Jites Wilson and Webster (p. 275, fig. 9). 

S1Jargani'um globip-ites Wilson and Webster (p. 276, fig. 18). 

Sphagmt?n antiquasp01·Ues Wilson ancl Webster (p. 273, fig. 2). 

'l'axodi1tm ll:icttip-ites Wodehouse. Wilson and Webster (p. 275, 

fig. 6). 

'l'ypha lctUfol·i1Jites Wilson ancl Webster ( p. 276, fig. 17). 

Study of the l\1:esozoic and Cenozoic plant microfossils may prove helpful in checking the identification of the 1negafossils and in the correlation of strata, 

but the writer, for a number of cogent reasons, has 

not attempted this study. No diatoms have been reported from Paleocene strata in America. 

FOSSIL WOOD 

Silicified and carbonized woods are locally abundant 

in the Paleocene strata, the silicified material being 

found chiefly in the shales and sandstones and the 

carbonized material in or near coal seams. In the 

vicinity of Medora, N. Dak., in the badlands along 

the course of the Little Missouri River, and also in 

the right bank of the Missouri River between Elbowoods and Sanish, N. Dak., silicified stumps and logs 

are plentiful. (See illustrations on pl. 21 of U.S. 

Geol. Survey Bull. 726, 1922, and pl. 4 of North Dakota Geol. Survey Second Bienn. Rept., 1902.) Most 

of the Paleocene wood is in prostrate position, but at 

some localities, as in Roosevelt State Park, just east 

of l\1:edora, and near Sully Springs station on the 

Northern Pacific Railway, southeast of Medora, upright stumps of swamp forests may be seen in place. 

The uninitiated traveler may sometimes be deceived 

by objects that look like prostrate logs when seen from 

a distance. Thus, along and on the east side of U.S. 

Highway 85, south of the bridge over the Little Missouri River, south of Watford City, N.Dak., and also 

on the hillside east of Seven l\1:ile Creek about 6 miles 

north of Stipek, Mont., long gray loglike sandstone 

concretions, simulating fossil wood in appearance, 

593121 0 - 62 - 5 

may seem to the observer to resemble, except in color, 

the prostrate trunks in parts of the Petrified Forest 

National Monument near Holbrook, Ariz. 

Most of the Paleocene wood is coniferous, but occasionally pieces of palms and dicotyledons may be 

found. One undescribed piece of a sycamore (Platanus), for example, was found in the Raton formation on the South Fork of the Purgatoire River, Colo., 

by Ross Johnson in 1950. 

The following woods have been described, but it 

should be noted that some of their names have since 

been changed by Krausel ( 1949) : 

Oupres8inowylon breverni Mercklin. Cramer in 

He~r, 1868, p. 167, pl. 42, figs. 11-17. Sinikfik, on 

east coast of Disko Island. 

Oupressilnowylon da'wsoni Penhallow, 1903, p. 46, 

figs. 9-11. The describer concluded that this wood 

represents a species of cypress ( Oupr.essus) , probably 

related to 0. macrocarpa Hartweg. No cones suggesting the presence of Oupressus in the Paleocene 

strata have yet been found. Southern Saskatchewan, 

Canada. 

Oupressinowylon elongatu1n Knowlton, 1888, p. 7, 

pl. 3, figs. 1-4. ICnowlton cited the locality for this 

wood as "Tiger Buttes, Dawson County, Mont." and 

concluded that "its age is, without doubt Laramie, as 

it is not far from Glendive from which come typical 

Laramie plants." The term Laramie as used by· 

ICnowlton means Laramie group in the old sense of 

Great Lignite. As I have not been able to relocate this 

locality to determine its stratigraphic position, there 

must remain some doubt as to the source of this wood 

and its age, whether Cretaceous or Paleocene. 

Ottpressinowylon ucranicu1n Goppert. Cramer, in 

Heer, 1868, p. 168, pl. 34, fig. 5; pl. 38, figs. 7-12. 

Atanekerluk, Greenland. 

Palmowylon cannoni Stevens. ICnowlton, 1930, p. 

43. Denver formation, 317, South Table l\1:ountain, 

Colo. Much palm wood occurs in the Paleocene of 

the Denver and adjacent basins, but I have made no 

attempt to study and classify it. In 1939, near the 

mammal locality on South Table Mountain (Brown, 

1943a) I found a small palm stump in place, with 

roots penetrating the clay that was once the soil in 

which the tree grew. 

Pityowylon krausei Felix, 1886, p. 48, pl. 12, figs. 1, 2. 

A radial section shows two or three pits on the side 

walls of the ray tracheids. A tangential section shows 

rays nine or more cells high and one resin canal. The 

transverse section is not illustrated, but the description 

of it records the presence of numerous resin canals 

and · parenchyma cells. The name was changed to 

Ooniferowylon krausei (Felix) Beck (1945, p. 69, 94). 
94 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

Fort Union formation, northeast of Medora, N. Dak. 

Pityoxylon sp. Torrey, 1923, p. 65. The specimen 

was said not to be well preserved and was not illustrated. It was cited as coming "from Cretaceous Laramie deposits at Sentinel Butte, North Dakota." As 

there are no Cretaceous outcrops at Sentinel Butte, 

the wood if the locality citation is correct, must be 

Tertiary in age, and presumably Paleocene. 

Podooarpoxylon dakotense Torrey, 1923, p. 73. Not 

illustrated. Torrey is "inclined to believe that this 

twig may have belonged to one of the m~ny 'sequoias' " 

described from the Great Lignite. Tertiary (not 

Cretaceous Laramie, as cited), Sentinel Butte, N. Dak. 

Pseudotsuga miooena Penhallow, 1903, p. 47, figs. 

12-13. The describer regarded Taxoxylon sp. Dawson (1875, p. 331) as synonymous with this species. 

Southern Saskatchewan, Canada. 

Rhamnaoinium poroupiniamum Penhallow, 1903, p. 

48, figs. 14-16, 21, 22. The describer compared this 

wood with that of the Salicaceae but concluded that 

its relationship is with Rhamnus'. Southern Saskatchewan, Canada. 

Rhamnaoinium triseriatim Penhallow, 1903, p. 54, 

figs. 17-20. The describer separated this species from 

R. poroupinianum by alleged differences in the rays as 

seen in tangential section. Southern Saskatchewan, 

Canada. 

Sequoia langsdorfi (Brongniart) Heer. Penhallow, 

1903, p. 41, figs. 2-4. This wood was probably misidentified for that of llfetasequoia ocoidentalis (Newberry) 

Chaney, whose twigs, cones, and seeds are common 

at many Paleocene localities in Canada and Montana. 

Southern Saskatchewan, Canada. 

Sequoioxylon burgessi (Penhallow) Torrey, 1923, p. 

79, for Sequoia burgessi Penhallow, 1903, p. 42, figs. 

5-8. Penhallow, because the bordered pits in this 

wood are chiefly in two rows instead of one, concluded 

that its affinity "appears to be in the direction of 

Sequoia gigantea." The presence of fusiform rays, 

seen in the tangential section, raises doubt about the 

reference of this wood to Sequoia. Penhallow's material is from southern Saskatchewan, Canada; Torrey's 

is from a lignite mine, locality unspecified, near Colorado Springs, Colo. The latter may, therefore, be 

from Cretaceous rather than Paleocene strata. 

Sequoioxylon dakotense Torrey, 1923, p. 77, pl. 10, 

fig. 24; pl. 11, figs. 25-28. Fort Union formation 

(not Cretaceous Laramie, as cited), 'iVilton, N. Dak. 

Sequoioxylon lara.1nense Torrey, 1923, p. 78, pl. 11, 

figs. 29-31. This is said to differ from S. dakotense 

in having far more abundant wood parenchyma and 

in lacking the small pits that characterize the 

tracheids of darkoten.se. Fort Union formation (not 

Cretaceous Laramie, as cited), Plentywood, Mont. 

Sequoioxylon montanense Torrey, 1923, p. 74, pl. 10, 

figs. 19-23. This species was considered not to be identical with Sequoia langsdorfi (Brongniart) Heer. Fort 

Union formation (not Cretaceous Laramie, as cited), 

bank of the l\{issouri River, .Culbertson, Mont. 

Taxodiun~ distiohum (Linnaeus) Richard. Penhallow, 1903, p. 36, fig. 1. As both Taxodium and Glyptostrobus are represented in the Paleocene strata by 

twigs, cones, and seeds, respectively, there is some 

doubt about the correct assignment of this wood. 

Southern Saskatchewan, Canada. 

SUGGESTED REJECTION OF SOME NAMES 

Because they are poorly preserved and sometimes 

badly illustrated or because they were named but never 

illustrated or because they. are now probably lost, the 

following leaves, fruits, and seeds are herein disregarded as name bearers : 

Ficus eva.nstonensis K.nowlton and Cockerell. 

Knowlton, 1919, p. 276, for F. (Protofious) nervosa 

Ne·wberry, 1883, p. 512. Said to have come from 

Evanston, Wyo., but no specimen corresponding to 

Newberry's description is now in the Evanston collection of the U.S. National Museum. 

flex 1niorophyllina Cockerell, 1911, p. 264, for I. 

miorophylla Newberry, 1883, p. 510. No specimen. 

Laurus palaeophila J{nowlton and Cockerell. 

l{nowlton, 1919, p. 347, for L. aoumitnata Newberry, 

1883, p. 511. No specimen. 

Rhan~nus nMvberryi ICnowlton and Cockerell. 

ICnowlton, 1919, p. 548, for Rhamnus ]Jarvifolius Newberry, 1883, p. 511. No specimen. 

Ulm~us grandifolia Newberry, 1883, p. 508. No 

speCimen. 

Viburnum grandidentatu1n Newberry, 1883, p. 511. 

No specimen. 

Viburnum pauoidentatum Newberry, 1883, p. 511. 

No specimen. 

Oarpites ooffeaeformis Lesquereux, 1878 (p. 303, pl. 

60, figs. 6, 7). 

0. ooryloides Knowlton, 1930 (p. 132, pl. 58, figs. 

3-6). 

0. oostattts Lesquereux, 1878 ( p. 303, pl. 60, fig. 5) . 

0. lattrineus Lesquereux; 1878 (p: 304, pl. 60, figs. 

20, 21). 

~Oarpites lineatus Ne\vberry. (f...esquereux, 1878, 

p. 302, pl. 60, figs. 1h-:ld; Newberry, 1898, p. 138, 

pl. 40, fig. 1). 

0. 1ninutulus Lesquereux, 1878 (p. 305, pl. 60, fig. 

25). 

0. 1nyrioarum Lesquereux, 1878 (p. 303, pl. 60, figs. 

8-11). 

0. oviformis Lesquereux, 1878 (p. 302, pl. 30, fig. 6a). 
COMPOSiTION AND ECOLOGIC SIGNIFICANCE OF THE PALEOCENE FLORA 95 

0. rostellatus Lesquereux, 1878 ( p. 303, pl. 60, figs. 

12, 13). To this probably belongs Palmocarpum corrugatum Lesquereux, 1878, p. 121, pl. 11, fig. 1i. This 

irregular, angular object may be an inorganic effect. 

0. triangulosus Lesquereux, 1878 ( p. 302, pl. 60, 

fig. 4 [USNM 496] ). The numbered specimen does 

not seem to be that illustrated by Lesquereux, but is a 

fru.it of Oercidiphyllum arcticum (Heer) Brown from 

the Denver formation, .317, at Golden, Colo. The 

other specimens, plate 62, figures 19, 20, said by Lesquereux to have come from the Upper Cretaceous 

strata at Point of Rocks~ Wyo., are missing from the 

U.S. National Museum collections. 

0. utahensis Lesquereux, 1878 (p. 305, pl. 60, fig. 22). 

Nyssa lane eo lata Lesquereux, 1878 ( p. 245, pl. 35, 

fig. 6). 

Tetranthera sessilifiora Lesquereux, 1878 ( p. 217, 

pl. 34, figs. 1c, 1d only). 

See rejected names for _palms (p. 53). 

COMPOSITION AND ECOLOGIC SIGNIFICANCE OF THE 

PALEOCENE FLORA 

Some of the named and described items in the list 

( p. 38) and foregoing systematic treatment of about 

170 species of the Paleocene flora are not classified 

with certainty. The remainder are considered to be 

placed fairly closely, at least generically, and show 

that a respectable variety of seedless and seed plants 

was present in the Rocky Mountains and Great Plains 

region over the span of some 10 million years. During this interval there were noticeable changes in the 

flora, so that a good collection from a given locality in 

the early part of the epoch can be distinguished from 

a similarly satisfactory one in the later part. 

Formerly, in various publications, the number of 

species in the Paleocene (-that is, Fort Unio~ .as then 

understood) flora was given as about 500-a too-generous estimate based oil overspeciation and guesswork about the known specimens. Nevertheless, the 

present list of about 170 is doubtless too small, for, 

had a botanist been present to take a complete census 

of the actual flora, the ·:total would, by comparison 

with the floras of assumed similar areas and environments today, be very much larger. It is interesting, 

but perhaps futile, to speculate about what species 

were present or should have been present in the flora, 

though not now represented in the fossil record. 

Formerly, also, it was thought that there was an almost complete floral break at the close of the Late 

Cretaceous. Now it is apparent that many early 

Paleocene plants cannot,. with assurance, be distinguished from their Late Qretaceous predecessors; and 

that species thought to be restricted to the Paleocene 

must be cited with cautious reservations. Among the 

latter the following, not singly but in combinations of 

two or more, are characteristic and common enough 

to be useful for tentative age determinations: Lygodium coloradense, lsoetites horridus, Gilnkgo adiantoide8, Glyptostrobus 'lWrdenskioldi, M eta8equoia 

occidentalis, Tamoditum olriki, Tlvuja interrupta, Sabal 

grayana, Oarya antiquorum, Pterocarya glabra, Pterocarya hispida, B;etula stevensoni, Oorylus insignis, 

0 astanea intermedia, Quercus greenlatndica, QuerCU8 

sullyi, Ulmus rhamnifolia, Platanus nobilis, Sassafra8 

thermale, Paranymphaea crassifolia, Eucommia serrata, Sapirndus affinis, Rhamnus goldiana, Fraxinu'8 

eocenica, Viburn'll!m antiquum, ·Viburnum asperum, and 

Viburnum cupanioides. Some of these, it should be 

stated, ranged into the early Eocene, or at least had 

very close relatives in the flora of that epoch. 

Inspection of the items in the Paleocene list shows 

that the seedless plants include algae, fungi, liverworts, mosses, ferns, and fern allies; and that the 

seed plants include cycads, ginkgo, conifers, waterweeds, palms, willow, hickories, walnuts, birch, hazel, 

chestnut, oaks, hackberries, elm, zelkova, breadfruit, 

figs, mulberry, sycamores, cinnamon, laurels, sassafras, 

waterlilies, katsura, magnolias, witchhazel, sweetgum, 

hydrangea, cherries, legumes, maples, koelreuteria, 

buckthorns, grapes, dogwood, black tupelo, ash, and 

viburnums. Suggestions about the possible relationship or likeness to modern species have been made in 

the discussion of each item, but these must be taken 

with the customary grains of salt, for no human observer was there to note some of the significant and 

perhaps evanescent features of the now extinct plants 

that, if known, would be most helpful in making comparisons with the assumed comparable living species. 

Many of the species, it will be noted, have counterparts, or similar relatives, in the temperate floras of 

the Eastern~United States and eastern Asia, thus reflectingthe similar aspects of these modern floras. A 

glance at the dicotyledonous leaves of the Paleocene 

flora shows that those with toothed margins far outnumber those with entire margins-a significant indication of adaptation to a temperate climate. Intermingled with the temperate species, however, are some 

ferns, palms, breadfruit, and figs (if they are figs) 

that were relics or adaptations from a warmer, subtropical flora. 

The Paleocene flora was essentially a lowland flora 

that extended for hundreds of miles inland from the 

open sea to the foothills and mountains that were the 

sources of the sediments in which the plant debris 

was entombed. Most of the fossils, it shonld be noted, 

were collected from strata that were formed at or near 
96 PALEOCENE FLORA OF THE ROCKY MOUNTAINS AND GREAT PLAINS 

sea level; and there are very few, if any, that can be 

singled out as having been collected from strata 

formed at higher elevations. That such strata, however, may have been in existence for a time, but were 

removed by erosion, is quite probable. From the aspect 

of the flora it would appear that many species were 

adapted to drier hillsides at some distance back from 

the swampy lowlands· and that their leaves, fruits, 

seeds, and wood were borne by streams to the burial 

ground in the lower areas of deposition. Some of 

the trees, particularly the conifers, were of large size, 

as shown by the remains of their 4- to 6-foot silicified 

trunks and stumps that may be seen, for example, in 

and around Roosevelt State Park, east of Medora, 

N. bak. Some of the stumps are still upright and, 

from the fact that they rest on clays or carbonaceous 

strata associated with coal seams, are without much 

doubt relics of a swamp forest like that of the bald 

cypress areas around the Dismal Swamp in Virginia 

and North Carolina. 

Comparing the Paleocene plants from the several 

areas, north and south, one is struck by the fact that 

some climatic zoning is clearly evident. For example, the remains of breadfruit (Artocarpus) and cinnamon (tOinna1rwmu1n) are limited to central Wyoming and southward. Nevertheless, palms are present as far north as the Yellowstone River in Montana. 

On the other hand, the ginkgos, birches, hazels, maples, and viburnums are limited to the areas north of 

the Colorado-Wyoming border or to what were presumably high areas near the headwaters of the southern terrain. Taking a broad view of the situation, the 

flora may be said fairly to have been a mesophytic one 

in a warm temperate environment with a medium 

amount of precipitation well distributed through the 

year. 

This flora and its environment were similar to those 

that existed in the Paris Basin in France at the same 

time, if one may rely on the description of the geology and the reports on the fossils of that area. Comparison with the Paleocene situation in Greenland is 

somewhat obscured by the lack of detailed geologic 

information about the strata and by the numerous 

probable misidentifications of· species. Precise comparisons of the plants with those of the American 

Paleocene are not always possible, but in their larger 

aspects the two floras seem to be closely comparable. 

PALEOCENE ANIMALS OF THE ROCKY MOUNTAINS 

AND GREAT PLAINS 

The Paleocene sedimentary strata of the Rocky 

Mountains and Great Plains are mostly nonmarine, 

but about 400 feet-the Cannonball member of the 

Fort Union formation-is marine, with a few brackish tongues. The fauna of the C~nnonball, consisting 

of foraminifers, corals, mollusks, crabs, and sharks, 

has been described by Stanton ( 1921), Fox and Ross 

(1940, 1942), and Holland and. Cvancara (1958). It 

was a fauna of an epicontinental sea that was open 

ocean probably to the north or northeast, and perhaps 

also toward the southeast Midway invasion of the 

gl.11f coast. 

The fauna from the nonmarine strata includes mollusks, insects, crustaceans, fishes, amphibians, reptiles, 

hi rds, and mammals. 

The fresh-water mollusks have been described in 

part by Meek (1876), White (1883, 1886), Russell 

(1931), Henderson (1935), Yen (1946, 1948), and 

Tozer (1956). 

Remains of insects, usually not well preserved, have 

been found at many localities, chiefly in carbonaceous 

shales. Although amber is sometimes present in these 

shales, I have never found any that contained insects 

or plants. Of the Paleocene insects only the wing of 

a cockroach has been previously figured (Brown, 1957, 

p. 341, fig. 1). Those illustrated here are listed by 

common pames only (pl. 69, figs. 2, 3, 4). Wings of 

beetles are perhaps the most common insect remains 

found. 

Scattered occurrences of ostracodes, some not identified (Lemke, 1960, p. 30), are known. Other crustaceans include some crabs from the Cannonball marine 

member reported by Holland and Cvancara (1958). 

Fishes are represented by scales (pl. 69, figs. 7, 8, 

14),- teeth, separate bones, and partial skeletons. 

Amphibians, apparently, left only footprints, such 

as A1nnwbatrachus montanensis (Gilmore (1928). 

Reptiles such as turtles and the crocodilelike 

Ohampsosaurus left abundant remains (pl. 69, figs. 

11-13). 

One feather (pl. 69, fig. 1), not identified, is at present all that represents the birds. 

Locally, mammals left relatively numerous remains 

(seep. 7ff; and pl. 69, figs. 9, 10). These have been 

treated by Russell ( 1932b), Matthew ( 1937), Simpson ( 1937b), Patterson ( 1939), Jepsen ( 1940), G~zin 

( 1941a, b, 1942; 1956a, b, c), and Van Houten ( 1945b). 

Problematic objects (pl. 69, figs. 15-19) that may 

be coproljtes are the bizarre and perhaps only traces 

of some unidentified aquatic or land vertebrate. 

From this survey of Paleocene life in the Rocky 

:Mountains and Great Plains, one may draw several 

inferences. First, the number of species now known 

must be only a fraction of the total that actually existed. It is clearly not reasonable to conclude that 

because one feather 'vas found only one kind of bird 

existed at that time ! The great number and thick-
REFERENCES CITED 97 

ness of the coal beds, however, does not mean that the 

number of species of plants that· contributed to the 

formation of the coal must necessarily have been large. 

A few species may have been so luxuriant as to have 

supplied most of the material. Second, most of the 

species now known must have been adapted by individual qualities or by favorable habitats for getting 

into the fossil record. This perhaps is more particularly true of the plants than of the land animals. 

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A Page 

Abietites. _ -------- ___ ------------ ___ -. _ -- _ __ __ __ 48 

tyrelliL _______________ ---- _ ---- _ ----- __ __ __ _ 48 

Acacia ... _________ ._-- ____ ----.--_______________ 75 

lamarensis. _ -- _________ -- _______ -- _ _ _ __ __ __ _ 75 

macrosperma. _ _ _ __ _ _ _ _ _ _ _ __ __ _ _ __ __ _ _ _ _ _ __ _ 75 

wardL. __ __ __ _ __ _ _ _ _ __ _ _ _ __ _ _ _ _ __ _ __ _ __ __ _ _ _ 75 

Acer _ --------------- --------------------------~- 75 arcticum .. _______________________________ 65, 78, 79 

collierL. __ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ __ ___ _ _ _ _ _ __ _ _ _ __ _ 79 

disputabili.~- ___ ---- ____ -- __ __ __ ___ __ __ _ __ __ _ 76 

fragile. ___ -----------_--_- __________________ 39, 75 

tragilis ... __ --------- __ ----- _ _ _ _ __ __ _ _ _ _ __ _ _ _ 75 

gracilens .. __ _ __ _ __ __ _ __ _ _ _ __ _ _ __ _ __ _ _ _ __ _ _ _ _ 65 

indivisum___________________________________ 65 

newberrvL------------------------ 39, 75, 76; pl. 46 silberlingi__ __________________________ 39, 76; pl. 46 

trilobatum tricuspidatum ____________________ 64,65 

SP----------------------------------- 65, 76; pl. 46 spp ____ ----------- __ _ __ __ _ _ __ _ _ __ __ _ _ _ _ _ _ _ _ _ 39 

·Aceraceae. _________ -- ____ -- _ __ _ _ _ __ _ _ _ __ __ _ _ _ _ _ 75 

acerifolia, Ampelopsis .. 36, 39, 71, 78, 80; pls. 51, 52, 59,66 Aralia .. ____________________________________ 64 

Populus. ____ -- ___ -- ___ -____________________ 70, 78 

acerifolium, Viburnum__________________________ 79 

aceroides cuneata, Platanus______________________ 64 

latifolia, Platanus ___________________________ 64,79 

Platanus___________________________________ 64 

Achlys triphylla_________________________________ 81 

acicularis, Dam mara____________________________ 49 

Aclistochara compressa _________________ _________ 39,40 

coronata. ______________ -- _ _ _ ___ __ __ __ __ _ _ _ __ 40 

·mitella ___ ______ -- __ _ _ _ _ _ _ _ _ _ ____ _ _ __ __ __ _ _ _ _ 40 

Acrostichum hesperium__________________________ 23 Actinidia. _ _ _ __ _ __ _ _ _ __ _ _ _ _ _ _ __ __ __ __ __ __ __ _ _ _ _ _ 91 

acuminata, Camptotheca. _ ---------------------- 77 Corn us. _______________ --___________________ 85 

Juglans. _ ---------------------------------- . 73 Laurus ___ _ _ __ ___ _ _ __ _ _ _ _ _ _ _ __ __ ____ _ __ _ ___ _ 94 

Myrica.____________________________________ 59 

Sequoia ______ -- __________ --_________________ 48 

adiantoides, Ginkgo ______________ 36, 38, 47, 48, 95; pl. 10 

Ginkgoites _ _ _ __ __ __ __ _ _ __ _ _ _ __ __ _ _ __ ___ __ _ _ _ 47 

Adiantum paululum____________________________ 84 

Aesculus _______ ------- ____ -- __ __ _ _ _ _ _ __ __ _ _ __ __ 19 

antiquus. __ _ __ __ _ __ _ _ _ ___ __ __ _ _ _ _ _ _ _ __ __ _ __ _ 19 

affine, Cinnamomum ____________________________ 61,66 

a/finis, Ficus ___________________ 38, 61, 63, 66; pls. 27,43 

Osmunda. _______ ------ __ _ _ _ __ _ __ _ _ __ __ _ _ _ _ _ 41 

Platanus___________________________________ 79 

Sapindus _____________________ ._ 39, 76, 85, 95; pl. 47 

Sequoia _____ -------_________________________ 49 

Agathaumus sylvestris, Sauria __________________ _ 

aguilar, Ficus ___________________________________ 71, 72 

Alaska. ________________________________________ 28, 29 

alas kana, Flabellaria__ _ ____ __ __ _ _ _ _ __ __ __ __ __ _ __ 54 

Phragmites. _ _ ___ _ _ __ __ _ _ _ __ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ 53 

alaskanum, Asplenium·------------------~----- 43 

alata, Ficus.____________________________________ 85 

Nyssa _______________________________ 39, 85; pl. 60 

olatus, Celastrinites. _ _ ___ __ __ __ _ __ _ _ __ ___ _ _ __ _ _ _ 73 

Sapindus ____ _______________________________ 76, 85 

alexoensis, Aralia._----------------------------- 85 Equisetum .• _ _ _ _ _ __ __ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ 46 

Alisma. _ __ __ __ _ _ _ _ __ __ __ __ __ __ __ __ _ _ _ _ _ _ _ _ _ _ _ _ _ 52 

macrophyllum. _ _ _ _ _ _ ____ _ _ _ _ __ _ _ _ __ __ _ _ _ _ _ _ 52 

Alismaceae. _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __ __ __ __ _ __ _ _ ___ _ _ 52 

Alismaphyllites •• __ __ _ _ _ _ __ __ _ _ _ _ __ __ __ _ _ _ _ _ _ _ _ _ 52 

crassifolium ___ ____________________________ • _ 52 

grandifolius __________________________ 38, 5£; pl. 15 

INDEX 

Page 

Alkali Gap, Colorado___________________________ 25 

Allantodiopsis erosa ____________________ 38, 41, 43; pl. 5 

Almy formation __________________________ 8, 22, 23, 27 

alrJ,ijolius, Celastrus____ __ __ __ _ _ _ _ __ __ __ _ _ __ _ _ __ _ 72 

Alnites macquarrii. ---------------------- _______ 57 

Alnus .. ---------------------------------------- 57, 58 americana___________________________________ 57 

auraria. ------------------------------------ 79 carpinifolia. _ _ _ __ __ _ _ __ __ __ _ _ __ _ __ _ __ __ __ __ _ 79 

kefersteiniL _ _ __ __ _ _ __ _ _ _ _ __ _ _ _ _ _ _ __ __ _ _ _ _ _ _ _ 57 

nostratum. ___ ------------------------------ 87 serrata _____ _________________________________ 72, 86 

speciipites______ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ __ __ _ 93 

Alsophila thelypteroides__________________________ 43 

amboraefolia, Monimiopsis______________________ 72 

Amelanchier similis"---------------------------- 87 Amentotaxus ____ ___ -- _______ -- __ --- ____ -_ _ _ _ _ _ __ 50 

argotaenia ____ ------------------------------ 48 campbelli. ___________________________ 38, 48; pl. 14 

florini. ----- _ _ _ _ _ _ _ __ _ _ __ _ __ __ __ _ _ _ _ _ __ _ _ _ _ _ 48 

americana, Alnus_______________________________ 57 

Corylus ___ ________ -------------------------- 57 

Dennstaedtia __________________________ 38, 42; pl. 6 

fossilis, Corylus_ __ _ _ _ _ __ _ _ ___ __ _ _ __ _ _ _ _ _ _ _ _ _ 57 

A mmobatrachus montanensis _ _ _ _ _ _ _ __ __ _ _ _ _ _ __ _ _ 96 

A mpelopsis __ ___________________________________ . 78, 81 

acerifolia ________ 36, 39, 71, 78, 80; pls. 51, 52, 59, 66 

brevipedunculata __ -------------------------- 79 bruneri carbonensis___ _ _ __ _ _ _ _ _ ___ _ _ __ _ _ _ __ __ 78 

montanensis ________________________________ 65, 78 

triwsp idata, Parthenocissu.s _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 79 

xantholithensis. __ _____ _ _ __ __ _ _ _ ___ __ __ __ _ __ _ _ 78 

A mygdalns. __ ____ _ __ __ ____ __ _ _ __ _ _ _ __ __ _ _ _ _ _ _ _ _ 74 

wilcoxiana_ _ __ _ __ ____ _ ___ _ _ _ __ ____ __ __ _ __ _ _ _ 74 

anceps. Blechnum ______________________ 38, 41, 43; pl. 6 

Salpichlaena_ ___ __ __ ___ __ _ _ _____ __ _____ _ _ _ _ _ 41 

Viburnum._________________________________ 78 

Andromeda denticulata__________________________ 59 

lanceolata __ ________________________ ---- _ _ _ _ _ 67 

scripta .. ___________________________ --_- ___ -_ 85 

Androvettia. ________________ c ___________________ 50, 51 

carolinensis. __ _ _ __ __ __ __ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 50 

catenulata_____ _ ___ _ __ _ _ _ __ _ _ _ __ __ __ _ __ _ _ __ _ _ 50 

elegans______ _ _ _ __ __ _ __ __ __ _ _ _ _ __ _ __ _ _ _ _ _ _ _ _ _ 50 

statenensis _ _ _ __ _ _ _ _ _ __ __ _ _ __ __ ___ __ _ __ __ _ _ __ 50 

Aneimia____ _ _ _ _ _ ____ _ _ _ _ __ __ ____ _ _ _ _ __ __ __ __ _ _ _ 44 

eocenica __________ -------- __ __ _ _ _ _ _ _ ____ _ _ _ _ 44 

Anemia_________________________________________ 44 

delicatula_ _ _ ____ __ __ _ _ _ ____ __ __ __ __ _ ___ _ __ _ _ 45 

elongata ___________________________ 38, 44, 45; pl. 6 

fremonti _____________ ~ ------ ________________ 44, 45 

grandifolia_ _ __ ___ _ _ _ __ _ __ __ __ _ _ __ _ _ _ _ _ _ _ _ _ _ _ 44 

lanceolata____ ____ _ _ ___ _ __ __ __ __ __ _ _ _ _ _ _ _ _ _ _ _ 44 

mosbyensis.~ _ __ _ _ _ _ __ _ _ __ ___ _ __ __ __ __ _ _ _ _ _ _ _ 44 

occidentalis ___________________________ -_- __ _ 44 

perplexa_ _ _ _ _ __ __ _ _ __ _ _ _ _ ____ __ __ __ __ __ __ _ _ _ 44 

snpercretacea_ _ _ _ _ _ _ _ ___ _ _ _ _ _ ____ _ _ _ _ __ _ _ _ _ _ 44 

sp_ _ ____ _ _ _ _ __ __ __ __ _ ____ __ _ _ __ __ _ _ _ _ __ _ _ _ _ _ 44 

Animia •.. ____________________________ • __ __ __ _ _ _ 44 

angnlata, Neuropteris___________________________ 83 

Trapa_ -------------- 39, 40, 83, 84, 88, 91; pls. 58,66 

angulatus, Nymphaeites_ ------------------------ 83 angustifolia, Magnolia __ · ______________________ 56, 67,68 

Sapindus ______________________________ -- _ __ 76 

angustiloba, Quercus _______ -------------------- 61 

Animas formation______________________________ 26 

Anisonchus fortunatus_____ __ _ _ _ _ __ __ _ _ _ _ _ _ _ _ _ _ _ _ 27 

Annonaceae_ _ _ _ _ _ _ ____ _ _ _ _ ____ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ 73 

annosa, Koelreuteria ____________________ 39, 76; pl. 59 

Page 

annuli/era, Sphaeria. ___________________ ------- _ 40 

anomala, Populus._.-- ________ -- __ -- ____ ------- 73 

Anomalofilicites monstrosus______________________ 46 

antecedens, Ulmus______________________________ 60 

AntelopP. Mountain____________________________ 19 

Antholithes browni. ______________ -- ______ -- _- _- _ 89 

fremontensis _______________ ----- _____ ------- 89 

Anthracite region __ ---------------------------- 26,27 

antica, Hydrangea _______________________ 39, 73; pl. 41 

antiqua, Brasenia. _____________ -- ________ ------- 51 

Tilia. _______________________________ -- _ --- _ 86 

antiquasporites, Sphagnum______________________ 93 

antiquora, Hicoria ___ ____________ -- _ -- _ -- _ ------- 56 

antiquorum, Carya __________ 38, 55, 56, 57, 95; pls.17, 18 

antiquum, Sparganium ______________ 38, 50, 51, 64; p1.14 

trinervum, Viburnum_______________________ 87 

Viburnum __ 18, 36, 39, 65, 67, 80, 86, 87, 95; pls. 63,64 

antiquus, Aesculus______________________________ 19 

Apeibopsis neome:Licana_________________________ 85 

Aphronorus sp __________________ ------ ___ -------

apiculatus, Rhamnus marginatus________________ 78 

Apocynaceae. ___________________________ - ____ -- 86 

Apocynophyllum lesquereuxi__ ___________________ 39,86 

linijolium ___ _________________________ ------- 86 

wilcoxensis ___ ____________________________ --- 83, 86 

Aponogeton __ _______ ------ c _ -------------------- 52 

appendiculata, Macclintockia____________________ 88 Platanus. ________________________________ -- 65 

aquamarum, Dryophyllum______________________ 56 

aquatica, Nyssa._-- ______ ---- ____________ --_-.-- 85 

aquilina, Salix __________________________ 38, 55, pl. 61 

aquilonium, Dryophyllum_______________________ 59 

Araceae. ____ ------- _______ ------ _ _ _ _ __ __ _ _ _ _ __ _ 52 

arachioides, Leguminosites ______ __________ --.--.- 70 

minor, Leguminosites __ • ___________ --------- 70 

Aralia ___________________________________ --_---- 64, 82 

acerijolia ___ __________________________ --.-.-- 64 

alexoensis __ __________________________ -. --.-- 85 

coloradensis .. _________________________ ------ 65 

dakotana ___ __________________________ -.. -.-- 64 

· digitata .•• ____________________________ ------ 64 

gracilis ________________ ------_-- ______ ------- 64 

lobata ___ _______________________________ ----- 82 

looziana. _____________________________ ------ 64 

notata. ________________________ --- ___ ------- 64 

pungens.-------------------c ---------------

reesidei_ _____ ----- ______ -- __ -----------------

serrata .. _ ------ ____________ -----------------

taurinensis .. ___________ ---------------------

triloba. ________________ -_-------------------

wardiana. _________________ -------- __ -------

sp __ ----------------------------------------

61 

65 

65 

59 

75 

64 

82 

Araliaephyllum artocarpoides____________________ 62 

Arapahoe conglomerate ______________________ 23, 24,25 

Araucaria. __ ----------------------------------- 48 hatcheri ... _____________________________ ----- 48 

longifolia __________________________ 38, 48, 49; pl.13 

Araucariaceae. ________________ ----- _____ ------- 48 

Araucarites. _______ ----------------------------- 48 

longijolia ___________________ ---- ______ -- ----- 48 

sp __ ---------------------------------------- 17 

arctica, Jenkinsella _______ ------------- _____ ----- 70 

Populus. ____________________ -- ______ ------- 61, 70 

Onoclea sensibilis---------------------------- 44 

Sphaeria ___________ ------------ ___ ---------- 40 

Trochodendroides ______ ---------------------- 70 Vitia ____________________________ -------_---- 82 

Woodwardia .•. ---------------------38, 43, 44; pl. 7 

107 
108 

Page 

arcticum, Acer _ --------------------------------- 65,79 

Cercidiphyllum ___ -------------------------- 18, 

36, 39, 70, 71, 78, 79, 95; pls. 37, 38, 52 

Equisetum __ ------- ______ ---------- _ -------- 46 

arcticus, Acer ____________ ----------------- _ -- _-- 78 

Woodwardites __ _____ ------- __________ ------ _ 43 

Argentina ________________ ---.-- _____ -------_____ 29 

argotaenia, Amentotaxus_________________________ 48 

Aristolochia cordi[olia _____ ______ : ---------------- 70 

crassi[olia __________ _____________ -- ___ ------ _ 70 

pageL ________ ----- ____________ --------- _ __ _ 70 

aristolochioides, Phyllites ____ -------------------- 82 

Arizona, Navajo country_______________________ 27 

artocarpidioides, Celastrinites_ ------------------- 76 

!lex. ______ -------------_-------------------- 39, 76 

artocarpites, Quercus ________ -------------------- 59 

artocarpoides, Araliaephyllum___________________ 62 

Ficus _________________________ 38, 61, 62, 85; pl. 28 

!lex __________ ------------------------------- 79 

Artocarpophyllum occidentale____________________ 61 

Artocarpus _____ -------------------------------- 61,96 

dicksonL----------------------------------- 60,61 

dissecta ___ --------- __ ----------------------- 60 

gigantea ______ ------------------------------- 60 

lessigiana-------------------------38, 60, 61; pl. 25 

liriodendroides ______ ------------ __ -____ ------ 60 

pungens. _ --------- ________ ----------------- 60 

similis _____ ---- __ --------------------------- 61 

artus, Sequoiites--------------------------------- 48 

Asimina _____ __ ------ _____ ---------------------- 73 

eocenica ________ ----------------------------- 67 

vesperalis.--------------------------- 39, 73; pl. 46 

asiminoides, Laurus_____________________________ 66 

asperum, Fiburnum ________ 15, 18, 39, 57, 86, 95; pl. 64 

Aspidiophyllu11!---------------------------------- 81 Aspidium escheri________________________________ 43 

goldianum_ --------------------------------- 42 heeri ____ _________________ ------------------- 43 

meyeri_ __ _______________________ ------- _ ---- 43 

Asplenium alaskamtm__________________________ 43 

eoligniticum ______ ___________ ---------------- 44 

elongatum, Sphenopteris _________ ---_________ 44 

iddingsi __ _____ -___ -------------------------- 43 penhallowL _____________________ -_ _ __ ___ _ _ _ 42 

primero ____ _____ --- ____________ ---- ___ ------ 42 

remotidens _ _ __ ___ _ __ ____ _ ___ _ _ _ ___ ___ ___ ___ _ 41 

asymmetrica, Quercus ___________________________ 38,58 

Atane strata _____________ ------------------~---- 28 

Atanekerdluk strata____________________________ 28 

lower------------------ ___ ------------------ 28 

upper ____ --------------_----------_-------- 28 

atava, Quercus __ -----------------------_-------- 58 

attenuata, Magnolia _____________ --------------__ 67 

auraria, Alnus ______________ -------------------- 79 

auriculata, Ficus ___________________ --_---------- 63 

B 

balsamoides eximia, Populus_____________________ 65 

Barilla Mountains, Texas. ___ ------------------ 27,28 

basilobata, Platanus _____________________________ 51,64 

basiserrata, Betula ______ ------------------------ 57 

Bauhinia. __ ------------- _ ---------------------- 74 wyomingana __________________________ 39, 74;pl.43 

Bear member----------------------------------- 7,17 

Bearpaw for.mation_____________________________ 17 

Bearpaw Mountains, Montana _________________ 12, 18 

Bearpaw shale ____ --------------------------____ 17 

beatricina con[ormis, Betula_____________________ 79 

beckwithii, Zizyphus _____ -------------------- ____ 78 

Belle Fourche River____________________________ 20 

benedini, Celastrophyllum ___ -------------------- 83 

Benitzia minima________________________________ 44 

Berberidaceae __ --------------- _____ ------------ 81 Berchemia multinervis ________________________ 56, 67,77 

Berrya racemosa _____ -------------------------- _ 70 

berryana, Dryophyllum__________________________ 56 

Ficus __________ ----------------------------_ 62 Juglans ____________________________ _________ 38, 56 

Sterculia ______________ ------------------____ 65 

INDEX 

Page 

berryanus, Sapindus __ -------------------------- 76 

berryi, Juglans __ -------------- _ ----------------- 71 

Magnolia_"--------- ____ -------------- 39, 71; pl. 39 

berthoudi, Ficus ___ ------------------------------ 82 

Selaginella __ __ ------------------------- 38, 46; pl. 4 

Betula __ ---------------------------------------- 57, 58 

basiserrata __ -------------------------------- 57 

beatricina conformis_ --- ______ ------------ _ -- 79 

brongniarti__ _________ ----------------------- 79 

claripites ____ -------------------------------- 93 

coryloides __ --------------- _____ ------------- 57 

fallax. -------------------------------------- 79 goppertL __________ --- ____ ------------------ 57 

prisca _____ _ -------- _____ ---------------- ____ 57 

schimperL ---------------------------------- 79 

stevensonL------------------------ 38, 67, 96; pl. 20 vogdesL ___________________________ ---- _ _ _ _ _ _ 57 

Betulaceae _____ ----- _ ----------------------- _ -- 57 

betulaefolium, Vibltrnum________________________ 86 

Betulites __ ______ --------------------------- _ ---- 81 

bicornis, Quercus ___ ------- ______ -----------_____ 62 

Big Dirty, the ______________ ------------------ __ 

Big Horn Basin, Wyoming ___________________ 1; 12,23 

Big Horn Mountains, Wyoming ________________ 13,23 

Bignoniaceae---------------------------- 74, 92; pl. 68 bignonioides, Dipterospermum _______________ 92; pl. 68 

bilinica, Juglans __ --------------- _ -------------- 59 biloba, Ginkgo ___________________________________ 36,47 

Biota borealis ___ ----------------_--------------- 51 Bison Basin, Wyoming _______________________ 8, 14,22 

bivalvi.•, Spriaxis __ -------------- -~---- ---------- 92 

Black Buttes coal groUP------------------------ 22 

Blechnum ____ ---------------------------------- 41 anceps _____________________________ 38, 41, 43; Rl. 6 

blomstrandi, Dennstaedtia_ ---------------------- 42 

Book Cliffs, Utah __ ---------------------------- 26, 27 

borbonia, Per sea ______ ---------------------_____ 67 

boreale, Equisetum __ ------------------------ o__ _ 46 

Rhytisma ____ ---------------------------- __ _ 40 

borealis, Biota _______ ----------------------______ 51 

Cercis ___________ ------ ____ ----------------- 70 

Leguminosites _____ ----------------------- _ -- 70 Magnolia ________________________ 39, 71; pis. 39,66 

Nordenskioldia _______________________ 39, 89; pl. 67 

Nyssa ___ ---------------------------- 39, 85; pl. 61 Paliurus ________________ --------- ____ ------- 70 

Torreya ________ -------------------------- _ __ 48 

Brachyphyllum ___________ ------ _____ ----------- 51 

brachysepala, Diospyros_ ------------------------ 85,89 

Brachysporium sp _______________ ---------------- 40,93 

Brackett Creek, Montana______________________ 17 

Brasenia antiqua________________________________ 51 

breverni, Cupressinoxylon________________________ 93 

brevifolia, Sequoia.---------------------------_-_ 49 brevipedunculata, A mpelopsis _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 79 

Bridger Range__________________________________ 16 

brittoni, Rhamnus_______________________________ 77 

brongniarti, Betula __ ----- ______ ----------------_ 79 brossiana, Laurus _______ -____________ ----- _ _ _ _ _ _ 68 

Persea _______________________________ 39, 67; pl. 34 

browni, Antholithes______________________________ 89 

Nymphaeites __ ------------------ ____ ----- __ _ 52 

bruneri carbonensis, Ampelopsis_________________ 78 

Vitis ___________________ __________ ---------- _ 78 

buckhami, Cinnamomoides______________________ 66 

Bull Mountain coal field _______________________ 14,17 

burgessi, Sequoia _______ ---- ___________ ---_------ 94 

Sequoioxylon __ ____ ----- _____________ --- _ ____ 94 

burlingi, Sphenopteris (Dennstaedtia)____________ 42 

Bursera _______ --------- ____ ------- ___ ------ _____ 76 

c 

Cabomba____ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 68 

gracilis______________________________________ 68 

grand is ___ ------- __ ------------------------- 68 inermis ______________________________ 39, 68: pl. 35 

Calamopsis danaL------------------------------ 53 

calhanensis, Phyllites _______ -----------------____ 85 

California, Elsinore_____________________________ 28 

Riverside County-------------------------- 28 

californica, Cephalotaxus ___ ---------------------- Callitrites __ ____________________________________ _ 

Page 

48 

51 

Calvert, W. R., quoted_________________________ 10 

Calycites hexaphylla _____________________ 39, 88, 89; pl. 4 

polysepala. _____ ----------- __________ 39, 89; pl. 4 

Calyptraphorus ____ --------------- ___ ----------- 8 

campbelli, Amentotaxus __________________ 38, 48; pl. 14 

Podocarpus_ __ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ 48 

Sabal. -- _- -- ________ ----- ___________ ------ _ _ 54 

Sabalites __ ---- _____________________ -- __ . _ _ _ _ 54 

Camptotheca acnminata_________________________ 77 

Canada, Alberta _______________________________ 18,19 

Alberta, Calgary ___ ------------------------ 18 

Cochrane_______________________________ 18 

Red Deer River________________________ 18 

Saskatchewan ______________________________ 18,19 

canadensis, Protophyllum. ----------------------- 61 Canna magnifolia ________________________ 38, 55; pl.15 

sp_ _ ___ _ __ _ _ _ _ _ _ _ _ __ _ __ __ _ _ __ _ _ __ _ _ _ _ _ _ _ _ _ _ _ 55 

Cannaceae_____ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ __ _ _ _ 55 

Cannonball fauna ______________________________ 8,27 

Cannonball member _____________ 5, 8-9,10, 12, 13, 14,18 

Cannonball River_ _____________________ . ________ 9,10 

cannoni, Cissus_________________________________ 78 

Ficus ______ ------------------_______________ 61 

Palmoxylon __ _________________________ . _ _ _ _ _ 93 

Rhamnus _____________________ .______________ 62 

Cannophyllites __ ------------ _____ _ ___ _________ _ _ 55 

magnifolia. _________________________ --- --- _ _ 55 

virlettL ___ ---------------------------------- 55 

Canon City coal field___________________________ 25 

Caprifoliaceae ___ --------- _______________ ---- ___ 81, 86 

Carapa eolignitica ________ ------------ ___ --- __ ___ 67 

Carbon Basin._-------------------------------- 21,25 

carbonensis, Ampelopsis bruneri_________________ 78 

Litsea. _________ -------------- ____ _____ _ ____ 68 

Malapoenna________________________________ 68 

Vitis _______ ______ --------------- ______ ---- __ 78 

Car ex ______________________ -- ___________ ------__ 53 

careyhurstia, Prunus ________________ 39, 73, 74; pl. 42 

carneosus, Phyllites ___ -------------------------- 87 

carolinensis, Androvettia ________ ----------------- 50 

carpinifolia, Alnus .. ---------------------------- 79 

Carpinus.-------------------------------------_ 57 

Carpites ____ ------------------------------------ 36, 37 

cocculoides ---------------------------------- 89 majori _________________________________ _ 

coffeaeformis __ -____________________________ _ 

coryloides. __ ------ _______ ------- ___________ _ cos tat us ____________________________________ _ 

lakesL ____________ -- _______________________ _ 

89 

94 

94 

94 

70 

laurineus ___ ------------------ ___ _ ____ ___ _ __ 94 

tineatus ___ ___ ------------------------------- 94 

minutulus __ -------------------------------- 94 

myricarum ____ ------------------------------ 94 

oviformfs ____ --------------------- _ ---------- 94 

rhomboidalis ____ -~ ____ ---------- _ --------- _ _ 90 

rostellatu.o ____ ---------- _ --- _ --------- _------ 95 

sulcatus _____ ---·------------------- __ ------__ 87 

triangulosus _____ ----------- _____ -- _--------- 95 

utahensis ____ ---"------------ __________ ---- _ 95 

verrucosus ___ ---------------- _ -------------- 52 

Carpolithes cocculoides _____ __ ------------------- _ 89 

spinosus _____________ ----------------------- 39, 89 

Carpolithus (Ginkgoites) selwyn;_________________ 48 hirsntus _________________________________ 91; pl. 67 

Cary a ______ -.- ______________________ ------------ 56, 74 

antiquorum _____________ 38, 55, 56, 57, 95; pis. 17, 18 

veripites _____ ________ ------- _---------------- 93 

viridiflnminipites ___ ------------------------- 93 

Caryojnglan~---- ________ ------------------------ 56 

Cassia concinna ______ _ --- _ ---------- ___________ _ 57 

evanstonensis _______ ------------- __ ---------- 57 

lancifolia _______ _____ --------------------- _ _ _ 67 

puryearensis _ --------- ---------------------- 67 

Castalia leeL __ -------- _ ------------------------ 69 

pulchella ____ ---------------------- _ --------- 69 

Castanaea intermedia____________________________ 57 

Castanea. _________________ ---- ___ -____ ---------- 58 

intermedia ________________________ 38, 58, 95; pl. 22 
Page 

Castanea-Continued 

ungeri. _ --------------------------------- ___ 58 castaneaefolia, Fagus____________________________ 58 

castanopsis, Quercus. ___ ------------------------ 60 

Castle Rock, conglomerate______________________ 24 

castrae, Viburnnm. ----------------------------- 87 

Catalpa .... --••• ----.-----•• ----------------- __ • 82, 92 

crassifolia .. ___ •• _. ------ _. ___ ••• ---- _. _ --•• • 70 

catenulata, Androvettia .. ------------------------ 50 

Fokienia·--------------------------- 38, 50; pl. 11 

Cathedral Bluffs, Colorado_____________________ 26 

caudata, Laurus. ------------------------------- 67 

caudatum, Laurophyllum. ---------------------- 39, 67 caudatus, Sapindus _____________________________ 67,86 

Caulinites ___ ------------------------•. --------- 53 

fecunda. ----------- _ ----------------------- _ 43 

Ceanothus fibrillosus •.•. ---------------------- __ 78 

Cedar Creek, North Dakota____________________ 9 

Cedar Creek anticline, Montana __________ 5, 12, 14, 15 

Cedar Mountain_______________________________ 26 

celastri/olia, Quercus·--------------------------- 76, 79 

Celastrini.tes alatus _____ _____ ------------------- _ 73 

artocarpidioides _ --------------------- _______ 76 

insignis·-------------------------------- 56, 72,85 

populi/olius ... ------------------------------ 76 

celastroides, Grewia _______________ --------------- 72 

Celastrophyllttm benedinL______________________ 83 

Celastrus. -------------------------------------- 59, 73 

alni/olius •• --------------------------------- 72 

curvinervis •• ___ ------- ___ --------------- ____ 72 

I errugineus. __ ------------------------------ 72 gaudinL _________________________________ · __ 76, 79 

grewiopsis __ -------------------------------- 72 

montanensis •• __ ----------- _____ ------- __ ___ 72 

ovatus. ------------------------------- __ ____ 72 

pterospermoides. ---------------------------- 72 serrata •• _. _ ••• _____________ .. _. _ _ _ _ _ _ _ _ _ _ _ _ _ 86 

serrat?ta .•. ____ .•. _. _ ... __ . _ •• ____________ • _ _ 72 

taurinensis. -------------------------------- 65, 72 

wardii. .• ----------------------- ------------ 72 sp. __________ .. ____ ..• _____________ • _ •••• _ _ _ 72 

Celtis lingualis._________________________________ 87 

newberryi__ __________________________ 38, 59; pl. 27 

parvifolia ••. -------------------------------- 59 peracuminata ________________________ 38, 60; pl. 20 

rugosa .. --------------------------------____ 87 

Cenchrus·----------------------------------- 91; pl. 67 Cephalotaxaceae .••. _______ • _. ________ •• _ _ _ _ _ _ _ _ 48 

Cephalota:copsis ___ . _ .. _. __________ • _. _________ • _ 48 

Cephalota:cus ..... ____ . _ .. ____________ • __ • _______ 48, 50 

cali/ornica. ___ ----------------------------- _ 48 

coloradensis. __ ----------- ___ ---------------- 49 Ceratophyllum_____________________ ______ __ _____ 69 

Ceratops beds---------------------------------- 6, 20 

ceratops, Ficus·--------------------------------- 15, 63 

Ceratozamia wrighti.____________________________ 47 

Cercidiphyllaceae. ----------------------------- 70 Cercidiphyllum ____________________ 50, 70, 71, 73, 79, 89 

arcticttm __ 18, 36, 39, 70, 71, 78, 79, 95; pls. 37, 38,52 

ellipticum. __ . ------------------------------ 70, 78 

.faponicum .. --- __ --------------------------- 70, 71 Cercis _ .. _ ••.• ___ • ______ . __ •• _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ 75 

borealis. ___ •••• _________ •• _______ .__________ 70 

coloradensis __________ --------------------- __ 70 

Cercocarpus ________ ------- _____________ ------ ___ 73 

ravenscragensis _____ __ ---------- _____________ 39, 73 

Cerillos coal field ___ ---------------------------- 26 

Ccrnay conglomerate___________________________ 29 

Chaetoptelea. _. ------- ____ -------------- _ _ ___ __ _ 60 

Chamaecyparis .... -------------------------- ____ 50 Chamaedorea ..... __ .. _. ________ • _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ 4 7 

coloradensis. -------------------------------- 47 

danae. _______________ ----------------------- 38, 53 

danai. _________ ---------------------------- _ 53 

Champsosaurus ______ --------------------------- 96 

Chara compressa________________________________ 39 

spp __________ -------------- ___ -------------- 38 

charpenteri, Quercus .•• ______ ------------------_ 59 

Cheyenne River·------------------------------- 20 Chionanthus membranaceus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 71 

'INDEX 

Page 

Chrysobalanus coloradensis. --------------------- 67,85 lanceolatus ___ ____________ • ____ -~ _ _ _ __ _ __ _ _ _ _ 85 

Chuska sandstone______________________________ 27 

Cimarron River, New Mexico__________________ 13 

cinnamomi, Sclerotium__________________________ 40 

Cinnamomoides buckhami_______________________ 66 

cinnamomoides, Zizyphus_______________________ 78 

Cinnamomum ____________________________ 66, 68, 84,96 

affine.----------- __ ------------ ___ ---------- 61, 66 dilleri ... _ •.• ___ •.• _. _ .. ____________ • ___ • __ • _ 67 

dubium_ ••. __ ------------------------------- 66 

ellipsoideum .. ------------------------------ 66 

ellipticum ______ ___ "------------------------- 61 

fici/olium .• ---------- _______ --------- __ ___ __ 61 

formosum ... ---------- __ ------------------.. 66 heeri.. _. _. __ .•.• ___ • ____ . ______ . ___ . _______ • 66 

inaequale .•. --------- ____________ ----------- 66 

lanceolatum ..... ___________ ----------------- 61, 66 

larteti. __ ___ • _. _. _____ . _. ________ .• _. _. _ •. __ • 66 

linifolium ___ -----.------------------- _ ------ 61 

middendorfensis. __ ----- ___ ------------------ 66 

mississippiensis ... -------- _________ --" ___ ___ 63 

newberryi. .•. ----------- __ ------------------ 66 ellipticum _______________ __________ ------ 66 

lanceolatum ___ -------- _______ ----------- 66 

minimum _____ -------------------------- 66 

paucinervum _________ __ ------------------- __ 66 

polymorphum ____________________________ 61, 66,67 

salicoides ___ _________________________ •• _. _ _ _ _ 61 

scheuchzeri__ __ _______________ • _ _ _ _ _ _ _ _ _ _ ____ 66 

sezannense ____________________ 39, 66, 67, 68; pl. 66 

wardi.. _____ ------- _ ----------------- __ ____ _ 66 vera. ________ • ___ .__________________________ 84 

sp ______ ___ ------------ __ --------- __________ 61, 66 

circularis, Isoetites.- ---------------------------- 46 

Cissites. ______ --------------------------- __ ----- 81 

colgatensis. __ ------ _ ------------------------ 81 

lobatus ______ ------ _ ---------------------.-- _ 81 

panduratus ___ ------------- __ · ______ --------- 82 

rocklandensis---·--------------------- 38, 81; pl. 27 

steenstrupL -------------------------------- 82 

CissttB------------------------------------------ 65, 81 

cannoni_ ______ ------------------------------ 78 

coloradensis ... ___ --------------------------- 63 

corylifolia ______ ----------------------------- 79 

grossedentata _____________ ------------------- 65, 79 

hesperia._---------------------------------- 79 

laevigata. _ ---------------------------------· 63, 65 

lesquereuxi__ ________ ------------------------ 78 

lobato-crtnata _____ -------------------------- 78, 79 

marginata. ________ 15, 39, 65, 76, 79, 80; pls. 53, 54,55 

parrotiaefolia ______ . ------------------------. 65 

tricuspidata _______ . ------------------------. 78 

Cladophlebis groenlandica .. --------------------- 45 

cladophleboides, Dryopteris. --------------------- 42 

Claenodon sp ... ------------.------------------- 7 

Claggett formation ... --------------------------. 17 

claripites, Betula .• -------.. ------------ __ ------- 93 

claytoniana, Osmunda___________________________ 45 

cleburni, Dillenites .. ---------------------------- 77 

Rhamnus.--------------------- 39, 62, 77, 80; pl. 42 

clintoni, Ficus planicostata______________________ 62 

Clintonia. _ -------------------- ___ -------------- 52 

oblongifolia. ____ -------------------------... 51 

Coal Bank Creek, Montana____________________ 15 

Coalmont formation____________________________ 25 

Coccolobis .... ____ ... -----------------------.. --- 82 

cocculoides, Carpites _________ -------------- __ ____ 89 

majori, Carpites .. --------------------------- 89 

Carpolithes _______ . -----------------------.. 89 

Cocculus haydenianus.-------------------------- 70 

kaniL ___ -------------------------- __ ---- __ . 88 

laurifolius _ --------------------------------- 88 cockerelli, Ficus ___________ •. ______ -------------- 62 

cojfeaeformis, Carpites.-------------------------- 94 

Colgate member, Fox Hills sandstone__________ 5 

colgatensis, Cissites .. _____ ____ ••• ________ -------. 81 

Thuja. _______ . ______ .•• ______ -------------- 51 

collieri, Acer ------------------------------------ 79 Selaginella ____________________________ 38,46; pl. 4 

109 

Page 

coloradense, Equisetttm .. ___ _ __ _ ___ _ __ _ __ _ __ ___ __ 46 

Hgpnum _____________________________ 38, 41; pl. 5 

Lggodium _________________________ 38, 46, 95; pl. 8 

coloradensis, Aralia.____________________________ 65 

Cephalotaxus. ------------------------------ 49 Cercis ________ --------------- ______ _____ __ __ 70 

Chamaedorea _ _ _ _ __ _ __ __ _ _ _ __ _ _ _ __ __ _ _ _ _ __ _ _ 47 

Chrg,,obalanus ______________ -------------- __ 67,85 

Cissus ........ ________ . __ . __ .. _ _ _ __ _ ___ _ _ _ _ _ 63 

Ficus .. _____________________________________ 77 

Leguminosites ________________________ 39, 74; pl. 68 

Mimosites ___________________________ 39, 75; pl. 43 

Phaseolites .. _________ . _________________ . ___ . 63 

Platanus __ . _______________ . _____ • ___ .. __ ._. 64 

Prunus ______________________________ 39, 75; pl. 42 

Rhus. ______________ ------------------------ 56 Zamia _______________________________ 38, 47; pl. 10 

Colorado, Brighton .... ____ . __ ._ .. _______ ------- 23 

Canon City __________ . ____ .. __ .. __ . ______ ._ 25 

Castle Rock. _______________________________ 12,24 

Colorado Springs. ____________ . __ .. ------_.. 23 

Craig ________________________ .. ____ . ___ ----- 26 

DeBeque _________________________________ 8, 25,26 

Golden ___________________ . _ .. _ ... _. _ 12, 23, 24 

HartseL. __ ._. ___ ... __ ._. ______ . _______ .•... 25 

Jefferson ...... ___ .. ________ ... ____ .... ____ ._ 25 

Meeker __ --------------------------------•. 25,26 

New Castle._------------------------______ 25 Rangely ______________________________ . ___ .. 26 

Rifle_-------------------------------.------ 13, 25 Trinidad._. __ . ___________________________ .. 24 

Walsenburg __ ------------------------------ 12,24 

Colorado River ... ------------------------------ 25 

columbi, Zizyphoides. _ --------------- _____ ____ __ 70 

Combretum. ________________ . ________ .. _ ... _.... 64 

commune, Palmocarpon _________________________ 39,89 

communis, Flabellaria. ------------------------- 53 Sabal. _____________________ .. _. __ . _ ........ _ 53 

compactum, Lygodium__________________________ 44 

compositum, Palmocarpon ______________________ 39,89 

compressa, Aclistochara __________________________ 39,40 

Chara _____ ______________________ .. ____ -_. _ _ _ 39 

concinna, Cassia________________________________ 57 

concinmts, Rhamnites.--~----------------------- 87 

conditionalis, Platanus newberryana_ ------------ 79 

co'njormis, Betula beatricina. __ ------------------ 79 

confusum, Hymenophgllum_ -------------- 38, 41; pl. 6 

Coniferoxglon kra?tseL _______ . -- ___ .. ________ . _ _ 93 

conjunctiva, Quercus ______ ---------------------- 59 Conocephalus _______ . __ . ___ . _ _ __ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ 40 

contortum, Viburnum___________________________ 80 

Corbicttla. ----.----- _ -------------------------- _ 8 

Corbula _______ --------------------------------- _ 8 

cordata, Populus __ ------------------------------ 82 

cordatus, Pterospermites .. ---------------- 39, 8$; pl. 41 

cordi folia, Aristolochia _______________ --------- _ _ _ 70 

Magnolia_------------------ ________ -------- 63 

coria ceo, Sterculia------------------_____________ 61 

Cornaceae. __ ---------------------- _____________ 77, 84 

Corn us .• _____ ----------------- ___________ ------ 84 

acuminata .. __ ----- ___ ------- __________ ---- _ 85 

denverensis ____ ___ ------------------. _____ 59, 77, 87 

emmonsi_ __ _____ ---------------------------- 85 fosteri. ___ __ . ______________ • ___ . ___ . _ _ _ __ _ _ _ _ 61 

hyperborea _________________________ 39, 84, 85; pl. 59 

impressa _______ -------- ___________________ -- 77, 85 

lakesii. •. _____ . ___ .. ___ . _. ______________ -- _- 77 

nebrascensis __________________________ 39, 85; pl. 59 

neomexicana .. -----------------------.------ 77 newberryi_ ____________________________ 37, 61, 62,85 

platyphylla. ___ . ____ _____ _ ___ _____ ____ ___ __ _ 84 

rhamnifolia. ____ ---------------------------- 68 

speciosissima ______ _ ------------------- ____ __ 84 

studerL--------------------------------- 63, 67,77 

coronata, Arlistochara. __ ------------- _ _ ___ ____ __ 40 

Corral Bluffs, Colorado_________________________ 24 

corrugatum, Palmocarpum______________________ 95 

corrugatus, Zizyphus __ -------------------. ___ ___ 78 

corrugis, Prunus _________________________ 39, 74; pl. 67 

coryli/olia, Cissus ••• ------ ____ ------------------ 79 
110 INDEX 

Page Page Page 

Cory lites /osterL _________________ ------------- __ 57 dakotensis, Glyptostrobus ________________________ 18,49 Dombeyopsis ___________________ ------ ___ -------- 82 

coryloides, Betula._-----------------_---------__ 57 Prunus ___ ---------------------------------- 56 grandi/olia .• _ ------------------------- _ _ ____ 82 

Carpites. _ ----------------- _ ---------------- 94 Sequoia _________ ---------------------------_ 15 magnifica ___________ --------------- ___ ------ 39, 82 

Momipites __ -------- _____ ------------------- 93 dalmatica, Ficus-------------------------------- 61,66 nebrascensis _________ ------------------------ 78 

Corylopsis sinensis _____ --_-------------------___ 87 Dam mara. ___ --------------------------________ 48 obtusa .• _- ---------------------------------- 63, 81 Corylus .•. _________ ------- __ ------- __________ -- _ 58 acicularis. __ ----------------------------- _ __ 49 sedaliensis. ___ ------------------------ ___ _ __ 82 

americana. __ -----------------------------__ 57 danae, Chamaedorea----------------------------- 38,53 trivialis. _______________________ • ____ ~·- _ _ _ _ _ _ 81 

fossilis. ____ --- _--. --- _------- _____ -•• -_ _ 57 danai, CalamopsiB---------·-·------------------ 53 dombeyopsoides, Phyllites---------- ________ ------ 81 

fosterL ___ _ ---------------------------------- 57 insignis ________________________ 38,57,58,95; pl. 21 Chamaedorea-------- _. -------------- ____ ___ _ 53 

Danforth Hills, Colorado_______________________ 26 

Donacites erdobenyensis. __ ----- ___ -------------- 92 

dorji, ThrinaX-------------------------- 38, 53, 5a:pl.14 

macquarriL ----------------- _ --- _ ---------- 57, 86 Daphnogene kaniL----------------------------- 88 Dragon fauna _____ -------- _______________ ------- 27 

orbiculata ___ ------------------------- ____ ___ 57 daphnogenoides, Zizyphus •• --------------------- 78 Dry Creek, Wyoming_------------------------ 8, 9, 13 rostrata. ___________ • ______ .- _______ • _. __ _ _ _ _ 57 daturae/olia, Credneria.- -------------- 39, 66, pls. 18,32 drymeia, Quercus. __ ._--------------------______ 58 fossilis. ____________ • __ -. _________ • _ _ ___ _ 57 Populus ________ -----_--------------------__ 66 Dryophyllum aquamarum_______________________ 56 

Coryphodon ___ ___ -------------------------- __ ___ 21 Davidia involucrata ________ ---------------------- 87 aquilonium. __ ----------------------------- _ 59 sp. __________________________________ • _ _ _ _ _ _ 22 Dawson arkose.-------------------------------- 23,24 berryana _____ ------------------- _ ----------- 56 

costatus, Carpites ___ --------------- _ ------------ 94 dawsonensis, Ficus ________ ---------------------- 62 moori. _______ _ ------------------------------ 58 

couttsiae, Sequoia. __ -----------_------ ____ ------ 49 dawsoni, Cupressinoxylon_______________________ 93 sub/alcatum ___ __ ----------- _ ---------------- 15, 59 

craspedodroma, Populus_________________________ 78 Peterospermites. ~------- ------------------ _ _ 62 Dryopteris _ ------------------------------------- 42 

crassi/olia, Aristolochia. _ ------------------------ 70 Quercus •• _.----------------------·--------- 62 cladophleboides _______ ---------------------- _ 42 

Catalpa _______________ ---------------------- 70 Deccan traps, India____________________________ 29 integra ___________ ------------------_-------- 42 

Parcmymphaea ______________ 15, 20, 39, 70, 95; pl. 36 decurrens, Negundo. ---------------------------- 79,80 lakesL .. ----------------------------- 38, 42: pl. 8 crassifolium, Alismaphyllites_____________________ 52 delicatula, Anemia •. ----------------------______ 45 lakesii. _____________ • __________ • ________ •• _ _ 42 

crassum, Viburnum_____________________________ 80 DeUoidospora diaphana-------------------------- 93 laramiensis. ___ ----------------------------- 42 Crataegus____ __ _ _ _ _ _ _ __ _ __ _ _ _ _ _ _ __ _ _ _ _ _ __ _ _ _ _ _ _ _ 81 demoresi, Phyllites----------------------- 39, 88; pl. 61 lesquereuxii. _ --------- ____ ------------------ 42 

myricoides ______ ---------------------------- 79 Dennstaedtia americana ___________________ 38, 42; pl. 6 meeteetseana .• ______ • ___________ • _____ 38, 42: pl. 5 

Crazy Mountains ____ --------------- 7, 8, 12, 14, 16, 17 blomstrandi. -------------------------------- 42 polypodioides _______ -----------•• --------·--- 43 

Crazy Woman Creek, Wyoming________________ 21 burlingi, Sphenopteris----------------------- 42 richardsoniana. _ ------- ___ ------ _. -----. ____ 42 

Credneria daturaefolia ________________ 39, 66; pis. 18, 32 crossiana _______ _ ------------------- _ ----- ___ 42 serrata ... ----------------------------- 38, 4S: pl. 8 

crenata, Planer a _____ -------------------------- 60 dentata, Fagus ______________ --__________________ 58 weedi. ___ ----------------------------------- 43 

Woodwardia. _ ----------------------------- 44 Macclintockia _________ _ --------------------- 88 xantholithensis. _ •• -------- __ ---------------- 43 

Crested Butte region.-------------------------_ 26,27 denticulata, Andromeda_________________________ 59 dubia, Quercus ___ ----- ___ ---------------------__ 58 

cretacea, Macclintockia. ________ ----------------- 88 Fraxinus. ____ -------------------------- ____ 57 dubiosa, Osmunda·----------------------------·· 45 crossi, Fagus ____________________ -----------_____ 71 Juglans. ____ ------- _______ --------------- __ 59 dubium, Cinnamomum .. ---------~-------------- 66 Juglans_____________________________________ 59 Pterocarya. _. ___ -- ________________ • _ _ _ _ _ __ _ _ 59 Taxodium _______ _______________ • __________ • _ 50 

Nelumbo____________________________________ 69 dentoni, Viburnum______________________________ 86 dubius, Zingiberites _____________________________ 38,55 

crossiana, Denn.~taedtia__________________________ 42 Denver Basin, Colorado __________ 4, 8, 12, 13, 23, 24,25 duplicata, Ficus _______________________ --- ____ ~__ 63 

;:rossii, Diplazium ______ ________ -_ _ ___ _ _ _ _ _ __ _ _ _ _ 44 Denver formation.------------------- 23, 24, 25, 26, 28 Ficus __________ -- __________ --_______________ 77 denverensis, Cornus _________________________ 58, 77, 87 E 

Cryptomeriteslambii .. -------------------------- 49 Phyllites .. ____ ------------------------------ 79 Eagle coal mine _______________________________ _ 

nordenskioldi, Elatocladus _ _ _ __ __ _ _ _ _ _ _ _ _ _ _ _ _ 49 Populus ________ ---------- ______________ ,___ 79 Eagle formation._--------------------------____ 17 

Cuchara formation_____________________________ 24 Prunus ___ ______________ ---------------- ___ _ 73 Eagle sandstone .• __ ----------------------- _____ 16, 17 

cucharas, Liquidambar _____ -------------------- 65 denveriana, Ficus _______________________ 62, 71, 77, 82 Echinodorus. ___ -------------------------------- 52 

Culmftes __ _________ ------ _____ --- __ ___ __ _ _ _ __ __ _ 53 Juglans _____ ____ ---- ___ ---- ___ -------------- 85 Ectopistes migratorius ____ -------------------- ___ 36 

cuneata, Meliosma______________________________ 61 Nyssa. __ ------ ____________ ----------------- 85 Edmonton formation _________________________ 8, 18,19 

Parrotia. _ _ _ _ __ __ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ 61 Depazea oronlandica. ___ ---------------------- 40 Egypt_ ____ ------------------------------------- 30 

Platanus aceroides _ _ _ _ __ ____ ________ ________ 64 deucalionis, Fagus_____________________________ 58 Elaeodendron polymorphum_____________________ 72 

Trapa ____ ----- _ -- _ _ _ _ _ _ __ _ __ _ _ __ __ _ _ _ _ _ _ _ _ _ 83 deusseni, Dolichites _____ ______ ---------- _ -------- 70 serrulatum ________ -------------------- _ __ _ _ _ 72 

cuneatum, Viburnum___________________________ 62 diaphana, Deltoidospora. ___ -- ------------------ 93 Elatocladus _______ -------------------------- ___ - 49 

Cnnninghamites_____ __ _ ___ ___________ __ ______ ___ 48 dichotomum, Viburnum_________________________ 80 (Cryptomerites) nordenskioldi________________ 49 

elegans_____ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ 48 dicksoni, Artocarpus. ___ ------------------------ 60,61 ( Taxodites) tinajorum. _________ : ____________ 49,50 

recurvatus_____ _ _ __ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ 48 Dicotyledons. ______________ ----- ___ ------------ 38, 55 elegans, Androvettia. _______________ ----- ____ _ _ __ 50 

SP------------------------------------------ 48 dienvalii, Phymatoderma ____ -------------------- 90 Cunninghamites. _. ------------------------- 48 

cnpanioides, Phyllites __ -----____________________ 87 digitata, Aralia _____________ • ____ ---------_______ 64 Elephant Butte Dam___________________________ 26 

Pterospermites_ __ _ _ _ _ _ _ ___ _ __ _ ___ _ _ _ _ _ _ _ _ __ _ 87 Ginkgo ________ ---------------- ______ -----_______ 47 ellipsoideum, Cinnamomum_____________________ 66 

Viburnum ____________ 39, 87, 95; pis. 46, 57, 65 Dilleniaceae .. __________ ------- ______ ____ _ _____ _ 82 elliptica, Kalmia ________________________ 39, 85: pl. 40 

Cupressaceae_--------------____________________ 52 Dillenites. ____ ---------------- ______ ------ ___ ___ 83 Magnolia.--------------- __ ----------- ___ --- 72 

Cupressinoxylon breverni________________________ 93 cleburni__ __ ----- ________________ ----------- _ 77 ellipticum, Cercidiphyllum----------------------- 70,78 

dawsoni_____________ ______ _______ ___ ________ 93 garjieldensis ___________ ------ __________ 39, 82: pl. 57 Cinnamomum. _ ----------- ____ --- __ ------ __ 61 

elongatum _________ ----------- __ ------------ 93 microdentatus_ ---------- ________ ------------ 83 Cinnamomum newberryi____________________ 66 

macrocarpa_ _ __ _____ _______ _ _ _ __ _ __ _________ 93 dilleri, Cinnamomum. __ ------------------------ 67 elongata, Anemia _____________________ 38, 44, 45; pl. 6 

ucranicum_ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 93 Diosypros ____ ___ ~---- __ ------- _ ____ __ _ ___ ___ _ ___ 89 elongatum, Cupressinoxylon ____________________ - 93 

Cupressus _______________________ -.- _ _ _ _ _ _ _ _ _ _ _ _ _ 93 brachysepala. ______________________________ • 85, 89 Sphenopteris (Asplenium)___________________ 44 

curvinervis, Celastrus___ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 72 jicoidea. _______ ----- _ -------- _ -------- ______ 67, 86 Viburnum _______________ ------------_______ 86 

cuspidata, Vitis _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 65, 78 hexaphylla. ________________ ----------- __ ____ 89 emmonsi, Cornua________________________________ 85 

Cycadaceae____ _ _ _ __ _ _ _ ___ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 47 obtusata _____ __ -------- _ --- ______ ----- ____ --~ 67 Engelhardtia ___ _ ------------ _ --------- _ --------- 56 

Cycadopites follicnlaris ___ ---------- ___ _ __ ___ _ ___ 93 wodani. __ ------ __ --- ___ ---- __ ------ __ _ _____ 89 eocenica, Aneimia_______________________________ 44 

Cylichnella____ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ 8 Diplazium crossii_______________________________ 44 Asimina .. _______________________ ----------- 67 

Cyperacites ________________________ ----------- __ 53 Dipterospermum bignonioides _______________ __ 92; pl. 68 Flabellaria .. _____ ------------------------- -~ 54 

tesselatus ____ . ------------------- _ ___ _ ___ _ __ 55 discolor, Rhamnus______________________________ 77 Fraxinus. ------------------ 39, 85, 86, 95; pis. 57,62 

cypericola, Sclerotites _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 40 Disko Island, Greenland_______________________ 28 Osmunda. ______ --------- ___________________ . 45 

Cyperus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 53 disputabilis, Acer __ ----------------------------- 76 

dissecta, Artocarpus. __ -------- ___ --------------- 60 

Phyllites .. __________________ -------------_-- 63 

Sabalites .. ________ --------- ____ -------- _ ---- 54 

D distichum, Taxodium____________________________ 94 Thrinax. ______________ --------------------- 55 

Dakota sandstone.----------------------------- 13 distichum miocenum, Taxodium _________________ 49,50 eolignitica, Carapa. _ ---------------------------- 67 

dakotana, Aralia________________________________ 64 

dakotense, Liquidambar __________________ 39,72; pl. 68 

Podocarpoxylon. _ _ _ __ _ _ _ __ _ _ _ _ _ _ _ __ __ _ _ _ _ _ _ _ 94 

Sequoioxylon ________________ ------------- ___ 94 

Viburnum ________________ .-------------____ 86 

distorta, Populus________________________________ 61 

distortus, Zizyphus________ _____ __ _____ ______ ____ 78 

disturbans, Phyllites _______________________ 39,88: pl. 60 

Dolichites deussenL----------------------------- 70 

Dombeya. __ ------------------------------------ 82 

eoligniticum, Asplenium------------------------ 44 

Equisetaceae. __ --- __ --------------------------- 45 

Equisetum ..• -------------------------------- 45, 46,91 

alexoensis. _____ ----------------------------- 46 

arcticum ___ ------- ___ ---- _______ ---- __ ----.. 46 
Page 

Equisetum-Continued boreale _________________ ___ --------- __ _____ __ 46 

coloradense ____ ------------------------------ 46 

globulosum _______________ _____ -------------- 46 

spp_---------------------------------- 38, 45: pl. 8 

erdobenyensis, Donacites_________________________ 92 

erectum, Vib!trnum_____________________________ 87 

Ericaceac ___ ------------------------ ______ ------ 85 

eridani, Rhamnus_______________________________ 68 

erosa, Allantodiopsis ____________________ 38,41, 43: pl. 5 

escheri, Aspidium _____ -------------------------- 43 

eucalyptifolia, Ficus----------------------------- 66 

Eucommia _______ --------- __ -------------------- 73 

serrata ______________________ 39, 7e, 77, 95; pls. 44,45 

ulmoides .. ____________ --------- _ --------- ___ 73 

Eucommiaccae ___ ------- _____________ --------- _ 72 

Euonymus splendens____________________________ 72 

xantholithensis _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 72 

Euphorbiaceae __ ---------------------------- ___ 89 Euphorbocarpum richardsoni _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 89 

europaea, Thujopsis_____________________________ 51 

europaeus, Glyptostrobus_ ----------------------- 49 

evansii, Rhus _____ -----_------- _____ ------------ 57, 58 

Evanston formation __________________________ 8, 22,27 

evanstonensis, Cassia _____ ----------------------- 57 

Ficns __ ------------------------------------ _ 94 

exaltata, Nephrolepis____________________________ 46 

Exbucklandia ___ --------------- __ ----------- ___ _ 72 

eximia, Populns balsamoides_____________________ 65 

expansa, Hiraea ____ ----------------------------- 52 Hydromystria _________________________ 38,5S: pl.16 

F 

Fagaceae. __ --------------- _____ -------_________ 58 

Fagus ____ ---------------------------------_____ 58 castaneaefolia _____ ------- __ ____ _ __ _____ ___ __ 58 

crossL ------------- _________ ------ _ _____ ___ _ 71 

dentata. _ -------------- _. ------------------- 58 

deucalionis _____ ~-- __ ------------- __ _ ________ 58 

macrophylla ___________ -------------- ____ ___ _ 58 

papyracea __ ---------------- ------- ________ 79 

fallax, Retula____________________________________ 79 

fewnda, Caulinites ___ _______ : _ __ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ 43 

Onoclea___ _ _ _ _ _ _ _ _ _ __ _ __ __ __ _ _ _ _ _ _ __ _ __ __ _ _ _ 43 

Felix coaL_____________________________________ 21 

Ferns __ ---------------------- _______ ----------- 41 

ferretiamtm, Sassafras___________________________ 68 

Ferris coal seam ___________ ·------------------- 19 

Ferris formation ________________________________ 21,25 

ferrngin.ens, Celastms___________________________ 72 

fibrillosus, Ceanothus _ _ ___ ____ _____ ____ _______ __ 78 

Zizyphus ________________ ·-----··------- 39, 78; pl. 50 

ficifoli!tm, Cinnamomnm_ ---------------------- 61 

jicoidea, Diospyros. ____________ ----------------- 67, 86 

Fiws __________ ___ --------------------- ___ 61, 62, 63,66 

a.(finis ____________________ 38, 61, 63, 66; pls. 27, 43 

agltilar _____________________________________ 71,72 

alata __ ------------------------------------- 85 artocartJoides ___________________ 38, 61, 62, 85; pl. 28 

anricnlata. _____________ -------------------- 63 

fJerryana_ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ 62 

berthondL _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 82 

cannoni_ _ ---------------------------------- 61 ceratops __ ----------- _______________________ 15,63 

cockerelli_ _______ __ -------------- _____ __ ___ _ _ 62 

coloradensi-~-------------- __ ___ _ _ _____ __ ___ __ 77 

crossii_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 77 

dalmatica _______ ______ ------------- _________ 61,66 

dawsonensis_ _ ____ ___ ____ ______ _____ ________ 62 

denveriana ___________ _________________ 62, 71, 77,82 

dnplicata. _ _____ ______ _____ __ _ ___ _____ __ ___ _ 63 

encalyptifolia_ ---------------------- _______ 66 

evanstonensis _____ ------------- _______ __ __ __ 94 

haddenL -------------------- __ _ __ __ _ __ __ ____ 61 hispida_ _ _ __ _____ _______ ______ _______ __ _____ 66 

impressa____________________________________ 62 

irregularis_ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 77 

lakesi _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 79 

leeL _ ---------------------- _________________ 62,63 

lig-nitum_ __ __ _ _ _ _ __ __ __ __ ___ _ _ __ __ _ _ _ __ _ _ ___ 83 

INDEX 

Page 

·Ficus-Continued 

limpida_____________________________________ 87 

martini_____________________________________ 63 

minutidens ___________________________ 38,6t; pl. 48 

mississippiensis__ _ _ _ _ _ __ _ _ _ _ __ __ __ _ __ _ __ __ _ 63 

monodon __ --------------------------------- 71 neodalmatica___ _ _ _ _ _ _ __ _ __ _ _ _ _ _ __ _ _ __ _ _ _ _ _ _ _ 62 

neoplanicostata _____________________________ 61, 62 

occidentalis __________ __ -------------------- __ 62, 85 ova lis_______________________________________ 85 

pagosensis ___ --------------------------- ____ 62 

planicostata _________ 38, 61, 6S, 63, 66, 68, 79, 81; pl, 26 

clintoni_ ---------------------------- ____ 62 goldiana ___________ _. ____ ------ ___ __ _ ____ 62 

magnifolia_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 62 

problematica __ ----------------------- __ _ 61 

populoides ______ ---------------------------- 61 

posttrinervis_ ------------------------------ __ 61 

. ~~:::~::~~~:~~t~--:~~~~~~~~~~~~~~~~~~~~~~~-~~~~ 61, :~ preartocarpoides ____ ____ --------------------- 62 problematica_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 62 

(Protojicns) nervosa __________ ------ __ ---~-- _ 94 

pseudopopulus ______ ·--------------------- 61, 62,78 

puryearensis ____ -------- ______ ------------ _ _ 71 

ratonensis _______ ______ ---------------- ____ __ 82 

rhamnoides ____ ------------------------ _ _ ___ 77 

richardsoni__ ___ --------------------- ____ ____ 63 

schimperi_ --------------------------- ___ __ __ 63 

smithsoniana ________________ __ -------------- 67 

speciosissima _______________ __ ------------ ___ 63 

spectabilis ____ __ ----- _- _ ---- __ -- __ _ _ _ _ _ _ _ __ _ _ 77 

squarrosa_ ---------------------------------- 63 

subtruncata. ------------------------- 39, 63; pl. 25 tiliaefolia ____________________________________ 63,65 

trinervis _ -------------------------- _ _ _ _ _ _ _ _ _ _ 61 

uncata .. ------------------------------ 39, 63, 72,77 

viburnifolia. ___ ----------------------------- 86 

wardi_ _________ ---------------------------- _ 63, 78 

wyomingiana ___ ---------------------------- 63 

SP------------------------------------ 61, 63, 71,82 finale, Viburnum_______________________________ 73 

jischeri, Lastrea_ -------------------------------- 43 

Lastrea (Goniopteris)________________________ 42 

Fish Creek, Montana__________________________ 7 

jisheriana, Quercus ___ --------------------------- 75 

Fissidens. ___ ---------------------------------- _ 41 Flabellaria alaskana______________________ __ ___ __ 54 

communis ____ ------------------------------ 53 

eocenica ____ -- _ --------------------------- __ _ 54 

florissantL. -------------------------------- 54 

gronlandica. _______ ------------------------- 53, 54 

johnstrupL ___ ------------------------------ 53, 54 

zinckenL ____ ------------------------------ _ 53 

Flagstaff limestone_____________________________ 27 

florini, Amentotaxus ___ ------------------------- 48 

florissantensis, Geaster _______ -------------- _ _ __ _ 89 

florissanti, Flabellaria___________________________ 54 

Sabalites __ _ -- _____________ ~ _____ -_-- _ _ _ _ _ _ _ _ 54 

Fokienia__ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ __ __ _ _ __ __ ___ __ __ _ 50 

catenulata ________ ___ ----------------- 38, 50; pl. 11 follicularis, Cycadopites _________________________ ~ 93 

fontanella, Liqnidambar _ ------------------------ 72 Fontinalis pristina __________________________ 92; pl. 69 

formosa, Geinitzia ________ --------------- ___ __ ___ 49 

formosum, Cinnamomum ___ -------------------- 66 Fort Peck Dam _______________________________ _ 

Fort Union flora _______ 4, 5, 6, 8, 9, 15, 16, 18, 19, 20, 21,22 

Fort Union formation __ ------------------------ 2, 

5, 7, 11, 13, 14, 15, 16, 17, 18, 19, 20, 21,28 

Fort Union group _________ 3,4,6, 7,8,9, 12, 17, 18,19,26 

fortunatus, Anisonchus__________________________ 27 

Fossil Basin, Wyoming_________________________ 8, 22 

fossil is, Corylus americana _____ ------------------ 57 

Corylus rostrata ---------------------------- 57 

Onoclea sensibilis ______ ____ ----------------- _ 43 

fosteri, Corn us __ ----------------------------____ 61 

Corylities ______ ------------ __ -----·---------- 57 

Corylus ___ ___ :!-:·---~-------- _______ ---------- 57 

[othergilloides, Hamamelites______________________ 72 

111 

Page 

Fowkes formation .• ------------------"--------- 23 Fox Hills sandstone _________ 1, 2, 5, 6, 9, 10, 14, 17, 21,23 

fragile, Acer------------------------------------- 39,75 

fragilis, Acer ------------------------------------ 75 

fraterna, Monimiopsis--------------------------- 86 

Fraxinus ___ ----------------------------------- _ 86 

denticulata __ __ ----------- _ ------------ _- _ __ _ 57 

eocenica _____________________ 39, 85, 86, 95; pls. 57,62 

juglandina _____ ___ ---- __ ------ _- _------- _ _ _ _ 59 

sp ______ ---------------------- ___ _ _ _ ___ __ _ _ _ 79 

Freedom Dome, Montana______________________ 9 

fremontensis, Antholithes------------------------ 89 

fremonti, Anemia.------------------------------ 44,45 fructifer, Sabalites ________________________ --- ____ 70 

Fucus. _ ---------------------------------------- 40 

lignitum. _ ---------------------------------- 40 

FungL _ ---------------------------------------- 38 

furcinervis, Quercus_---------------------------- 58, 59 

G 

Galisteo sandstone _______ ----------------------- 26 gardneri, Saccoloma __________________ 38, 41, 43; pls. 5, 6 

garjieldensis, Dillenites ___________________ 39,8e; pl. 57 

Gashato formation______________________________ 29 

oaudini, Celastrus------------------------------- 76,79 

Celastrus ___ -------------------------------- 76 Populus ________________ --- ___ ---- __ --_--___ 70 

Geaster ___ --- _ ---------------------------------- 89 

jlorissantensis ___ ______ -- ____ ---------- _- _--- 89 

Geinitzia ____ ----------- _ --- _ --- __ --------------- 49 

formosa _______________________ -------------_ 49 

Oelinden, Belgium, Paris Basin_--------------- 28,35 

Geonoma gigantea_______________________________ 53 

Geonomites goldianus____________________________ 53 

graminifolius ____________ ---- _ --------------- 53 

h"ydeniL ___ ---------------- ___ -- _ ---------- 53 

imperialis ____________ ----------------------- 54 

schimperL ---------------------------------- 53 

tenuirachis _______ --------------------------- 53 

ungerL _ ----------------- _________ -------- _- 54 

gigantea, Artocarpus ___ ------------------------- 60 

Geonoma ____ ---------- __ ----- __ ------------ 53 

Sequoia _____ ---------- __ ------- __ ----------- 50, 94 

Gillette coal field_______________________________ 20 

Ginkgo ____________________________________ 36, 47, 48,73 

adiantoides ___________________ 36, 38, 47, 48, 95; pl.lO 

biloba ___ __ ----------------- ____ -- _ ---------- 36, 47 

digitata ___ --------------- ______ ------- _ ---- _ 47 

laramiensis __________________________ -- _---- 15, 4 7 

truncata ______ --------------- _ ------------- _ 48, 90 

Oinkgoaceae ______ ---- ___ ---- ___ --- ___ ---------- 47 

Ginkgoites adiantoides_ -------------------------- 47 

selwyni, Carpolithus________________________ 48 

glabra, Ostrea_ ---------------------------------- 8 

Pterocarya ________________________ 38,57, 95; pl. 18 

glandulifera, Populus___________________________ 70 

Gleichenia ____ --------- __ ----------------------- 44 hesperia ___________________ · ____________ 38, 44; pl. 8 

Oleicheniaceae _______ ----- ___ ----- ___ - --- _-- ---- 44 

Glendive Creek, Montana______________________ 14 

glendivensis, Sapindus___________________________ 76 

globipites, Sparganium__________________________ 93 

globulosum, Equisetum__________________________ 46 

glyptostroboides, Metasequoia ____________________ 50,71 

Glyptostrobus_ ---------------------------- 49, 50, 73,94 

dakotensis _____ _ ------------ ______ ---- ___ ---- 18, 49 

eu.ropaeus__ __ __ __ __ __ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __ _ __ _ _ 49 

nordenskioldi ______________________ 38, 49, 95; pl. 11 

oregonensis ____ ----------------------------- 49 pensilis_ __ _ _ _ _ _ _ __ _ _ _ _ _______ __ ____ __ _ _ ___ _ _ 49 

ungerL __________________________________ -- _ 49 

Golden Valley formation _______________________ 12,20 

goldiana, Ficus planicostata_____________________ 62 

Lastrea _______________________________ 38.4S; pl. 5. 

Lastrea (Goniopteris)________________________ 42 

Rhamnus_--------------------- 39, 77, 87, 95; pl. 48 

goldiamtm, Aspidium___________________________ 42 

Viburnum __________________________________ 39,90 

goldianus, Geonomites___________________________ 53 

Palmacites ____ ___________________________ --- 53 
112 

Page. 

goldianus-Continued 

Rhamnus·---------------------------------- 62,77 

goldmani, Pteris. ------------------------------- 43 

Goniopteris lesquereuxL _ ------------------------ 43 

fischeri, Lastrea. ---------------------------- 42 goldiana, Lastrea____________________________ 42 

knightiana, Lastrea__________________________ 42 

polypodioides, Lastrea _______________________ 37,44 

gopperti, Betula·-------------------------------- 57 

Goshen Hole, Wyoming________________________ 10 

oracilens, Acer ---------------------------------- 65 

gracilis, Aralia---------------------------------- 64 Cabomba ____________________________ ____ ---- 68 

Juniperus __ -------------------------------_ 49 Laevigatosporites. ___ • _ ___ _ __ ___ __ _ _ _ ___ _ _ _ _ _ 93 

oraminifolius, Geonomites_______________________ 53 

Grand Hogback, the.--------------------------- 25,26 

Grand Mesa region_____________________________ 26 

grandifolia, Anemia ... -------------------------- 44 Dombeyopsis_____________________________ _ __ 82 

SabaL __________________ • _ __ __ _ _ _ _ __ _ __ _ _ _ _ _ 54 

Smilax ______________________ ---------------- 70 Sorbus ... _____________________ • _ __ _ __ __ _ _ _ _ _ 60 

Ulmus ......•• ___________________________ .__ 94 

grandifolioluR, Sapindus·------------------------ 76 

grandifolium, Maianthemophyllum______________ 52 

grandifolius, Alismaphyllites ______________ 38, 51!; pl. 15 

grandis, Cabomba. __ ---------------------------- 68 

orandivescipites, Picea _________ _____ ------------- 93 

Grasslike monocotyledons. _____________ 53; pis. 14, 16 

Graveyard Coulee, Montana ___________________ 14,15 

grayana, SabaL ________________ 38, 53, 54, 95; pis. 15, 16 

grayanus, Sabalites ... ------------------------- __ 54 

Great Divide Basin---------------------------- 22 Great Lignite, the ________________________ 1, 3, 4, 5, 8, 9 

Great ·Pine Ridge, Wyoming___________________ 20 

Green River Basin, Wyoming__________________ 12 

Greenland ______________ -------------------- ____ 28, 29 

oreenlandica, Osmunda __________________ 38, 45; pls. 5, 6 

Quercus.---------------- 38, 58, 59, 95; pls. 19, 22,40 

Grewia celastroides. __ ----------------- ____ ___ ___ 72 

obovata---------------------------. ---------- 86 

pealei. ...... -- ________ .. --- ____ . --- ___ ----- _ 72 

Grewiopsis-------------------------------------- 81 

platanifolia .. _ ------------------------------ 65 

populifolia .. -------------------------------. 65 

tenuifolia ... -------------------------------- 79 

viburnifolia .. __ --------------· -------------- 65 

grewiopsis, Celastrus. _ -------------------------- 72 

Populus ____________________ ---------------- 86 

groenlandica, Cladophlebis _______________ -------- 45 

Quercus_---------------------- ______ ------_ 58 

gronlandica, Depazea ____ ___ --------------------- 40 

Flabellaria .. -------------------------------- 53, 54 

Myrsine. __ --------------------------------- 59 

Pteris __ ---------------------------·- _ ___ ___ 45 

grossedentata, Cissu.~. --------------------------- 65, 79 

guillelmae, Platanus._-------------------- 64, 65, 79,80 

guillelmae heeri, Platanus_______________________ 79 

Gunnison Plateau______________________________ 27 

Gymnosperms .. _------------------------------ 38,47 

H 

haddeni, Ficus.-------------------______________ 61 

Hagan coal field________________________________ 26 

Halymenites rn,ajor______________________________ 90 

Hamamelidaceae _______________________________ 70,72 

Hamamelis virginiana __________ ::________________ 72 

Hamamelite.~ fothergilloides. --------------------- 72 inaequalis ____________________________ 39, 7t; pl. 40 

Hanna Basin, Wyoming ___ ------------ __ 11, 12, 21,25 

Hanna formation_______________________________ 21 

hatcheri, Araucaria __________ ------------------ __ 

haydeni, Manicaria. __ ------------------------. _ 

Platanus.---------------- __ -----------_----

haydenianus, Cocculus. ______ -------------------

haydenii, Geonomites ______ • _________ ... ________ _ 

hayeeiana, Pinus ____________________ • __________ _ 

hebridica, Onoclea. _____________________________ _ 

48 

53 

64 

70 

53 

48 

43 

INDEX 

Page. 

Hedera. __ -------------------------------------- 81 parvula ... _____ • -.- __ •• _____ •....• ___ . _ .•... _ 86 

rotundifolia. _. ------------------·----------- 63 

heeri, Aspidium ... ------------------------------ 43 

Cinnamomum .. ---------------------------- 66 

Juglans _. ---------------------------------· 59 

Piper--------------------------------------- . 70 

Platanus_---------------------------------- 79 

guillelmae. _ ---------------------------- 79 

heerii, Osmunda. _ ------------------------------ 45 

heersiensis, Macclintockia. __ -------------------· 88 

Hell Creek formation .• --.--------------------- 5, 

6, 7, 9,10, 11, 14, 15, 16, 17, 18, 19,28 

Hell Creek, Montana. __ ----------------------- 6, 15 

Hemitelites torellL------------------------------ 42 

hendersoni, Zizyphus.--------------------------- 78 

Henefer formation _________ --------------------- 27 

Hepaticae .. -------------- __ -------------------- 40 

herbacea, Phyllites.------------------------------ 61,79 

hesperia, Cissus ... __ ----- __ --------------------- 79 · Gleichenia _____________________________ 38, 44; pl. 8 

Onoclea _______________________________ 38,43; pl. 7 

hesperium, Acrostichum_________________________ 23 

Paleonuphar _________________________ 39,69; pl. 35 

hesperius, Sclerotium.-------------------------- 40 

Zizyphus. ____ -----------------.----------- _ 70 

heterodontt;,, Sterculia ___ . _______ -----------______ 65 

heterophylla, Inga________________________________ 85 

Hewett, D. F., quoted.------------------------ 23 

hexaphylla, Calycites.------------------- 39, 88, 89; pl. 4 

Diospyros ________ --------. _ ------ ______ . ---- 89 

hiatipites, Taxodium. --------------------------- 93 

Hicoria antiquora. _____ . --------------------- _ __ 56 

hilgardiana, Magnolia___________________________ 71 

Hiraea expansa. _ ------------------------------- 52 

hirsuta, Rhamnus.------------------------ 39, 77;pl. 49 

hirsutus, Carpolithus ________________________ 91; pl. 67 

hispida, Ficus ________ ----------_-------------___ 66 

Pterocarya _________________ 38, 57, 95; pls. 18, 19,68 

Hogback Canyon, Wyoming ___________________ 8,22 

Hogback formation_____________________________ 8, 22 

holbiilliana, Rhus .. ------------------------------ 83 

Hornerstown marL.---------------------------- 28 horrid1ts, lsoetites ______________________ 38, 46, 95; pl. 9 

Huerfano formation_--------------------------- 24,25 

Hunter Canyon formation ______________________ 26,27 

Hydrangea _______________________ ---- __ ----- ____ 62, 73 

antica _______________________________ 39, 73; pl. 41 

llydrocharitaceae ____________ -------- _________ -- 51, 52 

H ydromystria. _____ . _.-........... -......... -.. _ 52 

expansa _____________________________ 38, 52,· pl. 16 

Hymenophyllaceae ____ --- _____ .c •.. __ ---------- 41 

H1imenophyllum confusum ______________ 38, 41; pl. 6 

hyperborea, Cornua ____________________ 39, 84, 85; pl. 59 

Pinus. __ ------- __ . _________ ------ ___ -------_ 48 

Hypnum coloradense ______________________ 38, 41; pl. 5 Hyracotherium ________ __________________________ 11, 21 

iddingsi, Asplenium_____________________________ 43 

Ilex artocarpidioides.---------------------------- 39,76 

artocarpoides .. __________ ------- _ __ ___ __ _ _ __ _ 79 

insignia ..•.•. ______ -- ______ ----- _____ ._.---- 72 

longifolia _______ ______________ -- ___ -----.--- _ 59 

microphylla .. __________ ~ _________________ -- _ 94 

microphyllina_______________________________ 94 

ovata_____ _ __ _ _____ _ __ __ _ _ _ ___ _ _ _ __ _ __ _ _ _ _ __ _ 76 

reticulata________ _ __ _ __ _ _ _ __ _ __ ___ _ _ _ ___ __ _ _ _ 59 

triboleti .. ________ --. _. ______ --. _______ . --. __ 59 

Ilicaceae .... ___________ . ______________________ . _ 76 

imperialis, Geonomites. _. ----------------------- 54 

SabaL __________________________ 38, 53, 54; pl. 14 

impresAa, Corn us. __ ---------------------------- 77,85 Ficus .. -------______________________________ 62 

inaequale, Cinnamomum________________________ 66 

inaequalis, Hamamelites. ________________ 39, 7£; pl. 40 

Populus____________________________________ 73 

Protoficus _____ ------------------------------ 59, 72 

Rhamnus ________ -------- __________ --------- 77 

inaequilateralis, Pteris __________ __ --------------- 45 

Page 

India .• ---------------------------------~------_ 29 

indivisum, Acer --------------------------------- 65 inerme, Psilotum________________________________ 68 

inermis, Cabomba _______________________ 39, 68; pl. 35 

Inga heterophylla________________________________ 85 

innominata, Juglans____________________________ 71 

inominata, Vitia._._---------------------------- 82 

inopina, Paleonuphar --------------------------- 69 

inquirenda, Paleoaster --------------------------- 15, 24 

SabaL. ----- ___ -------------- __________ ----- 54 Smilax __________________________________ -·--_ 66 

insigne, Laurophyllum__________________________ 66 

insignia, Celastrinites _________________________ 56, 72,85 

Corylus ________________________ 38, 57, 58, 95; pl. 21 

Ilex .•• _ ------------------------ ____ • --- __ ___ 72 integra, Dryopteris ______________ ;________________ 42 

integrifolia, Platanus raynoldsi. _. --------------- 64,68 

integrifolius, Phoenicites _________________________ 53, 54 

intermedia, Castanaea___________________________ 57 

Castanea ________________________ 38, 58, 95; pl. 22 

interpungens, Sphaeria__________________________ 40 

interrupta, Thuja _________________ 15, 38, 51, 95; pl. 10 

interruptus, Thuites_____________________________ 51 

involucrata, Davidia_____________________________ 87 

Iris. _____ -------- ________ -- ____ ----------------. 52 

sp. ________ ---------- _____ ------------- __ __ _ 53 

Iron Bluff, Montana •.... -.--------------------- 5 

irregularis, Ficus .... ---------------------------- 77 

Isoetaceae. ____________________ ----------------- 46 

Isoetites .. __________________________ ------------ _ 46 

circularis _____ ------------------------------ 46 horridus ________________________ 38, 46, 95; pl. 9 

serratus ... __________________________________ 46 

J 

japonica, Ulmus________________________________ 60 

japonicum, Cercidiphyllum ______________________ 70, 71 

Jenkinsella arctica. __ --------------------------- 70 Jepsen, G. L., quoted. __________ _:______________ 23 

Jimmy Camp Creek, Colorado_________________ 23 

Joes Valley member---------------------------- 27 johnstrupi, Flabellaria ___________________________ 53, 54 

jorgenseni, Phegopteris. __ ----------------------- 44 

Sphenopteris .. _____________ -- __ ------------- 44 

Judith River __ --------------------------------- 4 

Judith.River formation_________________________ 17 

Juglandaceae. ______ . _______ ---- _. ___ .-.---... -- 55 

Juglandicarya _____ _____________________ . --- _ _ _ _ _ 56 

SPP-------------------------------- 38, 56; pl. 19 juglandina, Fraxinus ___ ________ . _______ . _ .. ___ .. 59 

Quercus. ________________ -----------------__ 59 Juglans ______________________________________ 56, 71,75 

acuminata.--------------------------------- 73 

berryana ______ ----------------- _ ------------ 38,56 berryi._ _ _ _ __ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 71 

bilinica. ________________________ ------ _____ _ 59 

crossi. ----------------------------- _ --- _____ 59 denticnlata .. ___ _ _ _ _ _ _ __ _ _ __ _ ___ _ _ _ _ __ _ _ _ _ _ _ _ 59 

denveriana. _______________________ .. -.- _ ---- 85 

heeri. _ _ __ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 59 

innominata_________________________________ 71 

leconteana _____ __________________ -.----.- __ .. - 56 

minutidens ______ ________________ -__ . ----.. -- 75 

nigella .. -------- __ ----------------------- 56, 72, 75 

ru,gosa_ --------------------- _____________ 56, 85,86 

sapindiformis. _ ------------- _ --------------- 71 schimperi_ __________________________________ 56,71 

taurina __________________________ 38, 56, 77; pl. 56 

thermalis _______ ------------- ______ --------- 63, 68 

ungeri. _____________________________ -------- 71 

sp __________________________________________ 59,86 

Juniperus gracilis_______________________________ 49 

K 

Kalmia elliptica. __________________________ 39, 85, pl. 40 

latifolia. ---------- _____ --------------------- 85 kanei, Macclintockia _____________________________ 39,88 

kanii, Cocculus _____ ________ .- _-- _ .. --.---------- 88 

Daphnogene ______ _____________ . --- __ .. -___ -_ 88 

kaulfussi, Lygodium_____________________________ 45 
Page 

Keene dome, North Dakota____________________ 12 

kefersteinii, Alnus_______________________________ 57 

Killpecker Creek, Wyoming .. ~----------------- 22 

Kingsbury conglomera~e------------------------ 13,21 

knightiana, Lastrea (Goniopteris) __ -------------- 42 

knightii, Menispermites__________________________ 70 

Knowlton, F. H., quoted_______________________ 6 

knowltoni, Populus.____________________________ 79 

Koelreuteria. ___ -------------------------------- 73, 76 annosa _______________________________ 39, 76, pl. 59 

Kome strata____________________________________ 28 

krausei, Coniferoxylon___________________________ 93 

krausei, Pityoxylon. ______ ,- ___ __ __ _ _ _ ____ __ __ __ _ 92 

L 

La Barge Creek, Wyoming_____________________ 22 

lacoeana, Populus_______________________________ 78 

laevigata, Ciss1ts.-------------------------------- 63,65 Laevigatosporites gracilis .. ______________________ 93 

ovatus _ _ __ __ ___ __ __ __ ____ __ _ _ __ ____ __ __ __ _ __ 93 

laharpi, Quercus.------------------------------- 59 

lakesi, Carpites __ ------------------------------- 70 

Dryopteris •. -------------------------- 38, 42; pl. 8 Ficus .. ______ ------_________________________ 79 

Viburnum .. ________________________________ 65, 80 

lakesiana, Nelumbo. _ _ _ __ __ __ __ __ __ __ __ _ _ _ _ _ ___ _ 69 

lakesianum, Nelumbium. ----------------------- 69 

lakesii, Cornua__________________________________ 77 

Dryopteris. _ ----- __ __ __ __ _____ __ __ _ _ __ __ _ _ _ _ 42 

Nelumbium. _ ------------------------------ 69 

Virburnum. __ ------ _____________ ---------- _ 78 

lamarensis, Acacia______________________________ 75 

Leguminosites _____ _____ -- ____ _ ___ __ __ __ _ _ _ _ _ 75 

lambii, Cryptomerites. -------------------------- 49 Lance Creek, Wyoming ______________________ 5, 10,20 

Lance flora.____________________________________ 4 

Lance formation________________________________ 5, 

6, 7, 8, 9, 10, 11, 15, 17, 18, 20, 22, 26, 28, 29 

lanceolata, Andromeda__________________________ 67 

Anemia_____________________________________ 44· 

Nyssa.---------------------------------- 85, 86,95 

lanceolatum, Cinnamomum. -------------------- 61,66 

Cinnamomum newberryi.___________________ 66 

Viburnum. ______________ -------____________ 87 

lanceolatum minimum, Cinnamomum___________ 66 

lanceolatus, Chrysobalanus_______________________ 85 

Zizyphus __________ _________________ ------- __ 61 

lancifolia, Cassia.------------------------------- 67 Nectandra. ______________________________ 56, 67, 68 

langeana, Myrica.______________________________ 59 

langsdorji, Sequoia .. ____________________________ 94 

langsdorjii, Sequoia.____________________________ 49 

Sequoiites .. _ _ ____ __ _ _ __ __ ___ _ _ _ _ _ _ __ _ ___ __ _ _ 50 

lapillipites, Sequoia·------------------------~--- 93 laramense, Sequoioxylon. _ _ _ ___ _ ___ __ __ __ _ _ _ _ ____ 94 

Laramie, "Lower"_---------------------------- 21 

Laramie formation.---------------------- 23, 25, 28,29 

Laramie Plains, Wyoming______________________ 4 

Laramie strata.------------------- 4, 6, 13, 16, 24, 25,26 

Laramie, "Upper"_---------------------------- 21 

laramiensis, Dryopteris__________________________ 42 

Ginkgo ______________________________________ 15,47 

larteti, Cinnamom1tm _ _ ___ __ _ _ __ __ _ _ _ _ ___ _ __ _ __ _ 66 

Lastrea fischerL. _ ___ __ _ _ _ _ __ _ _ _ _ __ __ __ __ __ __ ___ 43 

goldiana ______________________________ 38, 42; pl. 5 

(Goniopteris) fischerL _ __ _ _ _ _ _ _ _ __ _ _ _ __ __ __ _ 42 

goldiana _ _ __ _ ___ _ _ _ _ _ __ ___ _ _ _ _ __ __ ____ _ _ 42 

knightiana. _ __ __ __ _ _ _ __ _ _ _ _ _ __ __ _ _ _ _ _ __ _ 42 

polypodioides ___________________________ 37, 44 

latifolia, Kalmia.------------------------------- 85 

Platanus aceroides ______ -------------------- 64, 79 

latifolipites, Typha ___________________ ---------__ 93 

latiloba minor, Woodwardia______________________ 43 

serrata, Woodwardia________________________ 43 

Woodwardia. _ ---------------------------- __ 43 

latipennis, Podozamites__________________________ 47 

latum, Sapoticarpum _______________________ 91; pl. 67 

Lauraceae ______ .. _ ---- _____________________ 66, 68,89 

INDEX 

Page 

laurifolia, Magnolia----------------------------- 67, 71 

Quercus. ___ ----------------------------____ 59 

laurifolius, Cocculus. ___ --------,--------------- 88 

laurineus, Carpites--------------------- --------- 94 

Laurophyllum caudatum ________________________ 39,67 

insigne. ___ --------------------------------- 6tl 

perseanum ___________________________ 39, 67; pl. 34 

Laurus. -------------------------------------- __ 68 

acuminata. __ ------------------"------------ 94 

asiminoides. ------------- __ ---------------- _ 66 

brossiana .• ___ ------------------------------ 68 caudata. _ _ _ __ _ _ _ ___ __ __ _ __ _ _ _ _ _ __ ____ __ _ __ __ 67 

palaeophila. __ ------------------------------ 94 

primigenia.- _ --------------------------- 67, 73, 74 ratonensis ___________________________ ------ __ 67 

resurgens ____ ----------------------------- __ 76 socialis _______________________ 39, 67, 73, 85; pl. 27 

utahensis. ____ ----------------------------- _ 63 Lebo member _____________________ 5, 6, 7, 13, 14, 17,18 

leconteana, Juglans. ______ ---------------------- 56 leei, Castalia. ____ .-__________________ .__________ 69 

Ficus ____ ----------------------------------- 62, 63 

Magnolia._----- __ -------------------------- 71 Nymphaea ________________________ ---------- 39,69 

Phyllites ___________ -----------_------------- 79 

Sabal.----- _ --- __ -- _______ -- _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ 54 

Vitis ____ ____ --- _ _ _ _ _ _ _ _ _ _ _ __ _ _ ___ _ _ _ _ _ _ _ _ _ _ _ 82 

Leguminosae. ---------------------------------- 74 Leguminosites arachioides___________________ _____ 70 

arachioides minor____________________________ 70 

borealis ___________________ ~-________________ 70 

coloradensis ... ---~------------------ 39, 74; pl. 68 lamarensis. __ _ _ __ _ _ _ _ __ _ _ ___ _ _ _ __ __ _ _ _ _ _ _ _ _ _ 75 

lesquereuxiana. ___ -------------------------- 75 

stagnum ________ ---------------------------- 84 

williamsi. ____ ------ __ ---------------------- 39, 7 4 

Lemna. ____ ------------------------------------ 53 

(Spirodela) scutata__________________________ 52 

Lemnaceae. ------------------------------------ 52 

leonis, Quercus ____ ------------------------------ 76, 86 

lesleyana, Magnolia_____________________________ 71 

lesquereuxi, Apocynophyllum ____________________ 39,86 

Cissus. _ ------------------- __ --------------- 78 

Goniopteris _____ --------------------- _ ______ 43 

lesquereuxiana, Leguminosites___________________ 75 

lesquereuxii, Dryopteris ____________ ___ ----------- 42 

lessigiana, Artocarpus _________________ 38, 60, 61; pl. 25 

lessigii, Myrica _________ ------------------------- 60 

liasinum, Propalmophyllum_____________________ 54 

Libocedrus ___ ____ ------------------------------- 51 

sabiniana. ---------------------------------- 51 Lightning Creek, Wyoming____________________ 20 

lignitica, Marchantia ______________________ 38, 40; pl. 4 

lignitum, Ficus ... ----------------~------------- 83 

Fucus _ ------------------------------------- 40 

Limnobium ______ ______ ------------------------- 52 

limpida, Ficus__________________________________ 87 

limpidum, Viburnum___________________________ 86 

Lindera _______ _________ ------------------------- 67, 68 

obtusata. _ ----------------------- _ 39, 67; pls. 43, 66 

venusta ______ ------------------------------ _ 68 

linearis, Pteris ___ ------------------------------- 41 

lineatum, Palmocarpon. __ ---------------------- 39,90 

lineatus, Carpites. _ ----------------------------- 94 

lingualis, Celtis. _____________ --------------_____ 87 

Planera ______ ______ ----------------------- _ _ 60 

linifolium, Apocynophyllum_____________________ 86 

Cinnamomum_ ----------------------------- 61 Liquidambar _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 51, 72 

cucharas _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 65 

dakotense ____ : _______________________ 39, 72; pl. 68 

Jontanella __________ _________ ------------- __ _ 72 

liriodendroides, Artocarpus_______________________ 60 

Little Big Horn River__________________________ 21 

Little Bitter Creek, Wyoming ___________ _:______ 22 

Little Missouri River-------------------- 8, 9, 10, 12,14 Little Powder River ____________________________ 16,20 

Litsea. __________________________ ------------- _ _ 68 

carbonensis. _______ --------------------- ___ _ 68 

Livingston formation ___________________________ 14, 16 

113 

Page 

lobata, Aralia ______ ------- _______________ ------- 82 

Vitis.-------------------------------- 39, 82; pl. 56 lobato-crenata, Cissus _________ _________ -------- __ 78, 79 

lobatus, Cissites. _____ -------- __ ____ _ _____ ___ ____ 81 

London Basin, England--~--------------------- 29 

longifolia, Araucaria __________________ 38. 48, 49; pl. 13 

Araucarites_ ---------- ___ _ ___ ___ ____ _ __ __ _ __ 48 

flex ___________ ------------__________________ 59 

Sequoia .• ___________________________________ 48 

looziana, Aralia_________________________________ 64 

Louisiana, Caddo Parish________________________ 27 

louisvillensis, Malapoenna_______________________ 61 

Ludlow lignitic member________________________ 6 

Ludlow member _______________________________ 8, 9, 13 

lyallii, Macclintockia____________________________ 88 

lyelli, Quercus. ___ --------- ___ ------- ___ -------_ 59 

Lygodium ______ -- __ ------- __ --------- ____ ------- 45 

co_loradense ________________________ 38, 45, 95; pl. 8 

com pactum._------------------------------- 44 

kaulfussi- ---------------------------------- 45 neuropteroides ______ __ ------------ __ _ _ ___ ____ 45 

pumilum. _ --------------------------------- 45 

M 

Macclintockia ____________ _____ ----- _____ -- _ __ _ _ _ 88 

appendiculata _____ _________ ------- ___ _ ___ __ _ 88 

cretacea __ ------------------ _ --------------- _ 88 

dentata. ___ ----------------- __ ----- ___ ------ 88 

heer1iensis. __ ------------------------------- 88 

kanei- -------------------------------------- 39,88 lyallii. .. _________ -- _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 88 

' trinervis ..... ___ ---------------------------- __ 88 

mackayi, Zizyphoides. -------------------------- 39,78 mocneili, Quercus ________________________ 38, 58; pl. 19 

macquarrii, Alnites ______ ____ -------------- ___ __ _ 57 

Corylus _______________ --------------- _______ 57, 86 

macrocarpa, Cupressinoxylon____________________ 93 

macrodontum, Viburnum________________________ 86 

macroloba, Paleonelumbo _________________ 39, 69; pl. 35 

macrophylla, Fagus.---------------------------- 58 Osmunda __________________________ 38, 45; pis. 6, 8 

macrophyllum, Alisma__________________________ 52 

macrosperma, Acacia____________________________ 75 

magnifica, Dombeyopsis.------------------------ 39,82 magnifolia, Canna _______________________ 38, 55; pl. 15 

Cannophyllites ____ _______ ---------- __ _ _____ _ 55 

Ficus planicostata ____ ________ --------------- 62 

Magnolia _________________________ 39, 71, 72; pl. 39 

Sequoia __________________ ------ ___ ---------- 49 

Magnolia _________________________________ 64, 67, 71,73 

angustifolia. ------------------------"---- 56, 67,68 

attenuata___________________________________ 67 

berryL------------------------------ 39, 71; pl. 39 

borealis-------------------------- 39, 71; pls. 39, 66 cordifolia____________ _ _ _____ __ __ __ ___ __ ____ _ _ 63 

elliptica. ________________ --------- _______ ___ _ 72 

hilgardiana _____ ----------------- _ _ ____ _ _____ 71 

laurifolia _________ __ ----- ____________________ 67, 71 

leei. _______ -------------- _ _ _ __ _____ _ __ _ _ ___ _ 71 

lesleyana ----------------- __________ ------ _ __ 71 

magnifolia ________________________ 39, 71, 72; pl. 39 

regalis.------------------------------ 39, 72; pl. 39 rotundifolia ______________________________ 39, 63,72 

tenuinervis _____ -------------------- _ -------- 71 

Magnoliaceae _____ ------------------- ___ _ __ _ ___ _ 71 

magnum, Viburnum____________________________ 80 

Maianthemophyllum____________________________ 52 

grandifolium ______ ________ -------- ______ ____ 52 

major, Halymenites_____________________________ 90 

majori, Carpites cocwloides______________________ 89 

Malapoenna _______ _______________ ------------- _ 68 

carbonensis. _____ ------------------------ _ __ 68 

louisvillensis. ________ --------------------- __ 61 

praecursoria. ______ ------ _____ ------------ _ _ 68 

sessilijlora. ___ --------- _ --------- __ ------ __ _ 68 

Malva._------------- __ ------------------ __ ----- 89 

Manicaria haydeni______________________________ 53 

Marchantia lignitica _______________________ 38, 40; pl. 4 

pealeL--------~---------------------- 38, 40; pl. 4 polymorpha .••• _______________________ ----- _ 40 
114 

Page 

Marchantia-Continued 

yukonensis__________________________________ 40 

Marchantiaceae. ___ ---------------- _______ ----- 40 

Marchantites ____ ----- _ ------------- ______ ------ _ 40, 41 

pealei_ _____________ __ -------------- __ ____ ___ 40 

wardiL _ ------------------------------------ 40 

marginata, Cissus _____ 15, 39, 65, 76, 79, 80; pls. 53, 54,55 

Platanus _____ ----------- ____ --------------- 79 

marginatum, Viburnum _________________________ 79,81 

marginatum ravenscragensis, Viburnum__________ 80 

marginatus, Rhamnites __________________________ - 87 

marginat~ts apiculatus, Rhamnus __________ .______ 78 

marshallensis, Rhamnus-----·------------------- 77 

Martin, G. C., quoted__________________________ 29 

Martinez formation _____ ------------------------ 28 

martini, Ficus. ___ ---------------------_________ 63 

Maryland, Brightseat_ _____ ~------------------- 28 

Upper Marlboro____________________________ 28 

.Mastixia .• ----.-- ____ . ____ . _ _ _ _ _ _ _ _ __ _ _ _ __ _ _ _ _ _ _ 88 

maxoni, lVoodwardia____________________________ 43 

mclearni, Prunus _______________________________ 39,74 

Medicine Bow formation_______________________ 21 

Medicine Rocks area, Montana.________________ 8 

meedsii, Populus __________________________ ------ 59 

meeki, Zizyphus _. __ ------------- ___ ------------ 70 

meeteetseana, Dryopteris ___________________ 38, 42; pl. 5 

megasperma, Sagittaria ... __ -------------- 38,51; pl. 68 

melaenum, Viburnum___________________________ 80 

Melastomaceae. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 84 

Melastomites montanensis _____________ 39,68,84; pl. 56 

verus. _. ------- _______ ---------------------- 84 

Meliosma cuneata_______________________________ 61 

Melville strata. __ ------------------------------ 17 

membranaceus, Chionanthus_____________________ 71 

Sapindus. ---------.----------------------- _ 76, 87 

Meniscotherium ______ _ ------- ____ ---------- __ __ _ 22 

Menispermites knightiL------------------------ 70 

meriani, Quercus .... ---------------------------- 59 

Mesaverde formation __________________ 16, 22, 25, 26,27 

mesozoica, Robinia. __ -------- ________ ----------- 75 

Metasequoia ______________________________ 49, 50, 51,73 

glyptostroboides ________ --------- _____________ 50, 71 

occidentalis _______________ 18, 38, 49, 71, 94, 95; pl. 12 

mexicanum, Palmocarpon_______________________ 89 

meyeri, Aspidium ... ------------------~--------- 43 

microdentatus, Dillenites________________________ 83 

microphylla, Ilex _______ ------------------------- 94 

Planera ______________________________ 38,60; pl. 24 

microphyllina, flex______________________________ 94 

middendorfensis, Cinnamomum__________________ 66 

Middle Park,· Colorado _________________________ 13,25 

Middle Park formation_________________________ 25 

Midway formation.------------------------- 27, 28,30 

migratorius, Ectopistes _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 36 

Mimosites coloradensis ____________________ 39, 75; pl. 43 

minima, Benitzia __ --------------------------·--- 44 Ulmus _______________ --------------_________ 60 

minimum, Cinnamomum lanceolatum___________ 66 

minor, Leguminosites arachioides. __ ------------- 70 

Pterospermites __________ ------- __ ----------- 87 

Woodwardia latiloba_________________________ 43 

minutidens, Ficus·--------~--------------- 38, 62; pl. 48 

Juglans ________ ------ _____________ --------- _ 75 

Staphylea _________ __________________________ 39, 75 

minutulus, Carpites_____________________________ 94 

minutus, Zizyphus______________________________ 61 

miocena, Pseudotsuga___________________________ 94 

miocenum, Taxodium distichum _________________ 49,50 

mirabilis, Selaginellites_ ------ __ ------------ __ __ _ 92 

Triletes__________________ _____ _ _______ _____ _ 92 

mississippiensis, Cinnamomum_________________ 63 

Ficus ____________ ----------_________________ 63 

Missouri River _____________________ 2, 3, 9, 12, 15, 18,19 

mitella, Aclistochara____ __ _ _ _ __ __ _____ __ _ _ __ _ __ __ 40 

Mizpah coal field_______________________________ 16 

Mnium ______ ___ ------- _____ __ ___________ __ _ ____ 41 

montanense ____________________________ 38, 41; pl. 5 

Momipites coryloides____________________________ 93 

Mongolia_______________________________________ 29 

INDEX 

Page 

Monimiopsis amboraefolia .. _ _ _ ___ __ __ __ _ __ _ _ __ __ 72 

[raterna ____ __ ~- _ __ _ __ ___ _ __ __ _ _ _ _ _ __ __ _ _ __ __ 86 

Monocotyledons ______________________ ---------- 38,50 

monodon, Ficus.-------------------------------- 71 

Populus.___________________________________ 63 

.monstrosa, Selaginella _____________________ 38,46; pl. 4 

monstrosus, Anomalojilicites_____________________ 46 

montana, Sabal.________________________________ 54 

Montana, Absarokee___________________________ 16 

Baker _____________ ---------------------____ 6, 15 Bear Creek _________________________________ 7,16 

Biddle. __ ---------"---------------------___ 16 

Big Sandy_-------------------------------- 18 

Big Timber_------------------------------- 17 

Bridger __ ---------------------------------- 16 

Broadus ___________ ------------------------- 16 

Browning__________________________________ 18 

Circle._------------------------------------ 15, 19 

Coalwood__________________________________ 16 

Ekalaka ___________________________________ 6, 8,15 

Forsyth._--------- ______ ------------------ 2, 9, 14 Fort Peck__________________________________ 18 

Glendive_------------- _______ 5, 6, 7, 9, 11, 14-15, 19 

Harlowton_________________________________ 7 

Hathaway_-------------------------------- 14 Havre._____________________________________ 12 

Jordan.----------------------------------- 6, 9, 12 Livingston ____________________________ 8, 12, 16,17 

McCone County___________________________ 6 

Medicine Lake.____________________________ 7 

Miles City ________________________________ 6, 7,14 

Mizpah __ ----------------------------______ 16 

Powderville._______________________________ 16 

Red Lodge _________________________________ 14, 16 

Reed Point_________________________________ 16 

Roundup __ -------------------------------- 17 

Scobey_____________________________________ 18 

Sidney_____________________________________ 2, 15 

Snowden.__________________________________ 3 

Springdale. __ ------------------------------ 17 

Terry______________________________________ 14 

Van Norman_______________________________ 9 

Wibaux_----------------------------------- 14 

WilsalL------------------------------------ 17 

Wolf Point_-------------------------------- 18 

Montana group __ --------------------------____ 28 

montanense, Mnium ______________________ 38, 41; pl. 5 

Sequoioxylon_ _ __ _ __ _ _ _ __ _ _ _ _ _ __ __ _ _ _ _ __ _ _ __ _ 94 

montanensis, Ammobatrachus_ -----·------------- 96 

A mpelopsis .. _______________________________ 65, 78 

Celastrus __ -~ _ _ __ _ __ _ _ _ __ _ __ _ __ _ _ _ ___ _ __ _ ___ 72 

Melastomites ______________________ 39,68,84; pl. 56 

Moms _______________________________ 39, 64; pl. 20 

montanum, Nelumbium __________________ 39,69; pl. 35 

Viburnum. ________________ ------------- 80 

Monument Creek group________________________ 24 

moori, Dryophyllum_____________________________ 58 

Moraceae. ______ --------- __ _ _____ __ ______ _ ___ __ _ 60 

.i\foriconia. ____________________ -------- __ _ _ ____ _ 51 

Morus ____________ --------------------- _________ 62, 64 

montanensis __________________________ 39,64; pl. 20 

mosbyensis, Anemia_____________________________ 44 

multinervis, Berchemia _______________________ 56, 67,77 

MuscL ___________________ --------- ______ ------- 41 

Myrica acuminata. ___ -------------------------- 59 

langeana___ _ _ ____ __ _______ _ _ ___ ___ ___ ____ __ _ 59 

lessigii _____ ---------------- __ ____ _ __ __ _ ____ _ 60 

torreyi. ______ -------- ______ ----------------- 83 

wilcoxensis __________ __ ----------- ___ -------- 83 

sp.- ~ ------. _ -- ___ . _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 83 

myricarum, Carpites____________________________ 94 

myricoides, Crataegus___________________________ 79 

Myrsine gronlandica____________________________ 59 

Myrtaceae _______________________________ ------- 83, 86 

Myriophyllum torreyi__ ____________________ 39, 83; pl. 50 

N 

Nacimiento group______________________________ 26 

Page 

nebrascensis, Cornus ______________________ 39,85; pl. 59 

Dombeyopsis___________________________ ____ _ 78 

Populus _________________________________ 70, 71,78 

Trochodendroides___ __ __ _ _ _ ___ __ _ __ _ __ _ _ _ _ _ _ _ 78 

Nectandra lanci[olia __________________________ 56, 67,68 

Negundo decurrens------------------------------ 79,80 triloba.- --. _ __ _ _ _ _ _ _ __ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 75 

negundoides, Quercus____________________________ 65 

Nelumbian lakesianum__________________________ 69 

Nelumbites protoluteus__________________________ _69 

striata.-____________________________________ 83 

Nelumbium lakesiL----------------------------- 69 montanum ___________________________ 36,69; pl. 35 

tenuifolium. -------------------------------- 39,69 Nelumbo crossL________________________________ 69 

lakesiana. _. __ ------------ _ --------------- _ _ 69 

protolutea ... -------------------------------- 69 

tenuifolia. _ -- ___ ---------------------------- 69 

neodalmatica, Ficus_____________________________ 62 

neomexicana, Apeibopsis------------------------ 85 

Corn us. __ ---------------------------------- 77 

Quercus~ _______ ---------------------------- 63 neoplanicostata, Ficus ___________________________ 61,62 

Nephrolepis exaUata----------------------------- 46 

neriifolia, Quercus .. ----------------------------- · 86 nervosa, Ficus (Protoficus)_____________________ 94 

Populus. _____ ------------------------------ 65 

Rhus _______ ---------------------_-------_-- 60 

Nesson anticline, North Dakota________________ 12 

Neuropteris angulata____________________________ 83 

neuropteroides, Lygodium ___ -------------------- 45 

New Mexico, Cimarron________________________ 24 Gallup _________________________ ---_________ 27 

Navajo countrY---------------------------- 27 Raton _______________ -------_-----------____ 24 

Ute Park.---------------------------------- 13 

newberrianum, Fiburnum_______________________ 87 

Newberry, J. S., quoted _______________________ _ 

newberryana conditionalis, Platanus __ -___________ 79 

Platanus. ________ ----------- ___ ------------ 79 

newberryi Acer ________________________ 39, 75, 76; pl.46 

Celtis. __________ ------- ______________ 38, 59; pl. 27 

Cinnamomum. ----------------------------- 66 Cornus _______________________________ 37, 61, 62,85 

ellipticum, Cinnamomum___________________ 66 

lanceolatttm, Cinnamomum_________________ 66 

Populus .. ___________________ --------------- 70, 78 

Rha_mnus ... _______________ -------- _______ -- 94 

nielseni, Poacites ... -------- ________ --------- __ _ _ 47 

nigella, Juglans------------------------------ 56, 72,75 

nobilis, Platanus __________________ 18, 39, 64, 65, 95; pl. 29 

nodosa, Ophiomorpha. __ ------------------------- 39,90 

nordenskioldi, Elatocladue ( Cryptomerites) _____ -- 49 

Glyptostrobus ______________________ 38, 49, 95; pl. 11 

Potamogeton .. ______ --- __ --- _ --------------- 52 

Sequoia .. __________ -------------------- ___ -- 49 

Viburnum __________________ ---------------- 86,87 

Nordenskioldia ____ __ ----- _. __ ---------- _ -- __ ---- 89 

borealis------------------------------- 39, 89; pl. 67 

North Dakota, Beulah_________________________ 19 

Bismarck __________ ----------_----------_--- 18 

Bottineau CountY-------------------------- 18 

Buford ... --------------------------------- 3, 7,19 

Dickinson ___ -------- _________ ------________ 19 

Donnybrook. ___________ ------ __ -- ________ -- 7 

llebron .. __ ----------- __________ --- ____ _ _ ___ 19 

Mandan __________________________________ 8, 12,14 

Marmarth __________________________________ 6,15 

Medora. _______ ---------------------------- 20 

Morton County____________________________ 9 

Rolette County____________________________ 18 

Sawyer ____________________ ----------- ____ -- 18 

Sentinel Butte_____________________________ 2 

Sully Springs station_______________________ 19 

Velva. _______________________________ . ___ -- 18 

Washburn _______________ -- ______ ---- __ .---- 18 

Yule _____________________________________ 8,10,12 

North Horn formation__________________________ 27 

North Park, Colorado _______________________ 12, 13,25 

North Park Basin, Colorado____________________ 12 
Page 

· North Park formation-------------------------- 25 

North Platte River .•• -------------------------- 20 

nostratum, Alnt£8 _- ___ ------------------ _______ - 87 

notata, Aralia ____ -----------_--------------_____ 64 

Nugsuak peninsula, Greenland_________________ 28 

Nuphar rttbrodiscum •• -------------------------- 69 

N11mphaea leei. _____________ -------------------- 39, 69 

pulchella __________ ----------- __ ------------- 39, 69 

N11mphaeites angulatus-------------------------- 83 

browni_ _______ --------------- _- --------- __ __ 52 

striatus _____ ----- __ --------------------- ___ - 83 

Nympbeaceae _____ ------------- _______ ------ ___ 68 

Nils sa. __ ---------------------- ____ ---------_ 85, 87, 88 

alata.-------------------------------- 39, 85; pl. 60 

aquatica ____ ------------------------------ __ 85 

borealis------------------------------- 39, 85; pl. 61 

denveriana ____ ---------------------- ___ --~- _ 85 

lanceolata ___ _______ ----------- ___ ----- _ -- 85, 86, 95 

obovata _____ _____ ----------- ____ ---------- ___ 39, 85 

racemosa ______ ------------------------- __ ____ _ 70 

81/lvatica. __________ -- ___ ----------------- ___ 85 

0 

oblonglfolia, Clintonia___________________________ 52 

obovata, Grewia ___ --------------------------- __ _ 86 

N11ssa. ______ ------------------ _ ----------- _ 39, 85 

obtusa, Dombeyopsis ___ ------------------------- 63,81 

obtusata, Diospyros _____ -------- _____ ----------- _ 67 

Llndera. ------------------------- 39, 67; pis. 43,66 

obtusifolius, Sapindus.---------------·----------- 76,85 

occidentale, Artocarpophyllwm___________________ 61 

Tax odium ___ ------------------------------- . 49 ocCidentalis, Anemia____________________________ 44 

Ficus __________ ------------ _________________ 62, 85 

Metasequoia ______________ 18, 38, 49, 71, 87, 95; pl. 12 

Ocotea_ ------- __ --- ____ --------------------- ___ - 68 

oeningensis, Pteris. _ ------------- ____ ----- __ ____ 45 

Ohio Creek conglomerate _______________________ 26,27 

Ojo Alamo sandstone.-------------------------- 26 

Oleaceae __ ·-------------------- __ ------------- __ 85 

olriki, Taxites.---------------------------------- 50,52 Taxodium _____________________ 38,50, 95; pis. 10,11 

llitis _____________________________ 39, 8£; pis. 27,59 

Onoclea ______ • __ ----------------------------- ___ 43, 44 

jecunda ________ _____ -- __ ---- __ -------------- 43 

hebridtca ___________ ------------------------- 43 hesperia _______________________________ 38, 4S; pl. 7 

sensibilia __________________ ------------------ 43, 44 [arctica]___ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ __ _ _ _ _ _ 44 

jossilis _____ ______ ----------------------- 43 

Ophiomorpha. _ --------------------------- ___ __ _ 90 

nodosa ____ ______ ---------------------- ______ 39, 90 

oppositinerve, Virburnum_______________________ 86 

orbicularis, Ulmus._----- __ ----------------_____ 60 

orbiwlata, Corylus_ --· __ ------------- _ ____ __ __ __ 57 

Oredoxites plicatus ___ --------------------------- 54 

oregona, Tilia _____________ -------- __ _ _ ______ _ ___ 57 

oregonensis, Glyptostrobus_______________________ 49 

Oreodaphne _________ ---------------------- __ _ ___ 68 

pagosensis _____ ----------------- ______ ------ 67 ratonensis______ _ _ __ _ ___ _ ___ _ _ __ ___ _ _____ ___ 72 

Oreopanax sp_ --------------------- ___ ------- ___ 72 

Orites SP---------------------------- _ ----------- 70 

Osmunda affinis-------------------------------- · 41 claytoniana ________ --------- _ ______ __ __ __ __ _ 45 

dubiosa. ----- _____ ------------------------ _ _ 45 

eocenica ___________ ____ -------- _ ------------- 45 

greenlandica _______________________ 38, 45; pis. 5, 6 

heerii. ------------- _____ ----------------- __ _ 45 

macrophylla _____________ ~--------- 38, 45; pis. 6, 8 

regalis.-------------------- _________________ 41, 45 

Osmundaceae ___ --------------------------- ____ 45 

Ostrea glabra ______ ____ --------- ____ _ ___ _____ __ __ 8 

ovalis, Ficus. ___ ----------------- __ ---------____ 85 

ovata, !lex _______ ------------------------________ 76 

ovatus, Celastrus ___ ____ ------ __________ ------- __ 72 

Laevigatosporites _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 93 

ovijormis, Carpites______________________________ 94 

'lNDE.X 

p 

Pacific Creek, Wyoming________________________ 22 

pagei, Aristolochia _____ -------------------- ______ 70 

pagosensis, Ficus ________ ------------------------ 62 

Oreodaphne_ -------------------------------- 67 

Phyllites----------------------------- 39, 88; pl. 18 pulaeophila, Laurus_____________________________ 94 

Paleoaster inquirenda _____ ------------------ ____ 15, 24 

Paleonelumbo macroloba ________________ 39, 69; pl. 35 

Paleonuphar hesperium _________________ 39, 69; pl. 35 

inopina __________ --------------------------- 69 

paliurus, Pterocarva. ___ ------------------------ 57 

Paliur.us ____ ------------------------------------ 77, 78 borealis.: __ . __ -________ ._.--_. __ --.-----______ 70 

zizyphoides ______ -------------------------- __ 78 

sp __ --------- ____ ------- ____ ---------------- 39, 77 Palmaceae ___________________________________ 53, 89,90 

Palmacites goldianus____________________________ 53 

palmata, Vitis ___________ ----------------------- 82 

Palmocarpon. __ -------------------------------- 53 commune ___________________________________ 39,89 

compositum _________________________________ 39,89 

lineatum------------------------------------ 39,90 

mexicanum __ ------------------------------- 89 subcylindricum _______________________ 39, 90; pl. 67 

truncatum_ --------------------------------- 39,90 

Palmocarpum corrugatum_______________________ 95 

Palmoxylon _______ __________ ----- _ -------------- 53 

cannoni --------- ____ ------------------------ 93 Paloreodoxites plicatus ________________ 38, 53, 54; pl. 15 

panduratus, Cissites_____________________________ 82 

Pantolambda SP--------------------------------- 7 

papyracea, Fagus_______________________________ 79 

Paranymphaea __________ __ ------------------ __ __ 82 

crassifolia __________________ 15, 20, 39, 70, 95; pl. 36 

Parataxodium wigginsL.------------------------ 50 Paris Basin, France ____________________________ 28,29 

Parrotia cuneata ______ ------------ ____ --------- _ 61 

parrotiaefolia, Cissus____________________________ 65 

Parthenocissus (Ampelopsis) tricuspidata_________ 79 

quinquejolia __________________ ______ ---- _____ 79, 81 

tertiaria ____________________ -------- ____ __ ___ 82 

ursina _______________________________ 39, 81; pl. 48 

parvifolia, Celtis __________________ --------------- 59 

parvifolius, Rhamnus ___ ------------------------ 94 

parvula, Hedera _____________ -- _ --- _ ------------ 86 

Paskapoo formation_------------------------- 8, 18, 19 Patoot strata___________________________________ 28 

paucidentatum, lliburnum __ -------------------- 87,94 

paucrnervum, Cinnamomum __ ------------------ 66 paulula, Trapa __________________________ 39, 84; pl. 58 

paululum, Adiantum___________________________ 84 

pealea, Phanerophlebites_________________________ 43 

pealei, Grewia___________________________________ 72 

Marchantia ___________________________ 38, 40; pl. 4 

Marchantites _____ _____ ----- _ _ __ _ ___ _ ____ _ _ __ 40 

pellucidens, Phyllites____________________________ 79 

pellucidtts, Phyllites_____________________________ 65 

penhallowi, Asplenium__________________________ 42 

Pterospermites ________ -------- ______ _ __ _ __ __ 62 

Quercus ____ ------- ____________ ------------_ 59 

pennaejormis, Pteris_____________________________ 44 

pensilis, Glyptostrobus~-------------------------- 49 peracuminata, Celtis ______________________ 38, 60; pl. 20 

perjectum, lliburnum___________________________ 86 

perita, Prunus ___________________________ 39, 74; pl. 43 

perplexa, Anemia_______________________________ 44 

perplexum, Viburnum ______________________ ---_ 86 

Persea ______ __ ------- _____________________ ----- _ 67 

borbonia_ _ _ _ _______ __ __ __ _ _____ ___ _ __ ____ _ _ _ 67 

brossiana _____________________________ 39, 67; pl. 34 

perseanum, Laurophyllum ________________ 39,67; pl. 34 

Puerco formation_______________________________ 7 

Phanerophlebites pealeL.----------------------- 43 

Phaseolites coloradensis__________________________ 63 

Phegopteris jorgensen!___________________________ 44 

Phellodendron __ -------------------------------- 76 

Phoenicites integrijolius ___ ---------------------- 53, 54 

Phragmites_ ------------------------------------ 53 alas kana____________________________________ 53 

115 

Page 

Phyllites ___________________________ ---------- ___ 36, 37 

aristolochioides __ ---------- ___ -------- __ _ ___ _ 82 

calhanensis _ ------- _________ ------- _ ------- _ 85 

carneosus_ ----------- _______ ---------------- 87 cupanioides_______________ _ _ _ __ __ _____ _____ _ 87 

demoresi. ---------------------------- 39, 88; pl. 61 

denverensis _____ __ ----------------- _ ----- ____ 79 

disturbans.--------------------------- 39, 88; pl. 60 

dombeyopsoides __ --------------------------- 81 eocenica ________ --------- ____________ ----- ___ 63 

herbacea_ ----------------------------------- 61,79 

leeL __ --------------------------------- _ ____ 79 paoosensis ____________________________ 39, 88; pl. 18 

pellucidens _________________ ----------- ___ ___ 79 

pellucidus __ -------------------------------- 65 populoides. ___ ------------ _________ ----- ___ _ 81 

retusoides _______ ----------------- __ ----- ___ _ 61 

trillioides ________ ------- _ -- __________ -- _ __ __ 78 

trinervis __ ---------------------------------- 81 

venosa ____ ---------- _______ ---------------- _ 87 

sp ____ ------ _____________ ------------ ____ 60, 61, 63 

Phymatoderma dienvalii_________________________ 90 

Picea grandivescipites_ _ _ ___ _ ___ __ ___ __ __ ___ _ _ _ _ _ 93 

Pierre shale ____________________________ 5, 10,17,24,25 

Pinaceac _ -------------------------------------- 48 

Pinus _____ ------------------- ______ -------.____ 48 

hayesiana_"·------------ __ _ ______ _____ _ ___ __ 48 

hyperborea. ____ _ __ ___ ____ __ _ _ _ _ __ __ ___ _____ _ 48 

Pinyon conglomerate.__________________________ 22 

Piper heeri ____ --------------------- ___ _ _____ __ _ _ 70 

Pistia. ___ -------------------------------- _ ----- 52 

Pityoxylon krausei. ----------------------------- 93 sp_ _ __ __ __ __ __ ___ _ ______ _ _ __ __ __ _ _ _ _ _ _ __ _ _ _ _ 94 

Planera ___ ---------- ___________ ----- _ ------- __ _ 60 

crenata __ ----------------------------------- 60 

lingualis ___ ---------- ___ ------ _ _____ __ _ __ ___ 60 

microphylla __________________________ 38,60; pl. 24 

planeroides, Zelkova __________________ 38,60; pis. 22,24 

planicostata clintoni, Ficus---------------------- 62 

Ficus ______________ 38, 61. 6£, 63, 66, 68, 79, 81; pl. 26 

goldiana, Ficus______________________________ 62 

magnifolia, Ficus ___ ------------------------ 62 

problematica, Ficus ___ ---------------------- 61 

Platanaceae. __ --------------------------------- 64, 81 platania, Quercus _______________________________ 62, 65 · 

platanijolia, Grewiopsis _ -------C---------------- 65 

"Vitis _____ -------- ______________ -------- __ --- 80 

platanoides, Platanus ___ ------------------------ 61, 79 Viburnum ___________________ ----------_____ 80 

Platanus ______________________________ 51, 64, 65, 81,93 

aceroides ___ -------- ______ ------ ___ _ ___ _ ___ __ 64 

cuneata_ -------------------------------- 64 latifolia _________________________________ 64, 79 

affinis _______ ---- ________ ------------------. 79 

appendiculata ___ _______ ------.----------- __ . 65 

basilobata .. --------------------------------- 51,64 

coloradensis. --------- ______ ---- ____ --------- 64 

guillelmae.---------------------------- 64, 65, 79,80 

heeri.. _____ . ---------------------------- 79 haydeni. _________________ ------------ __ ____ _ 64 

heeri.. _________________ --------------------- 79 

marginata _____ . ------------------ __ -------- _ 79 

newberryana. ____________ -------- __ --------. 79 

conditionalis. __ ------------------------- 79 nobilis _______________________ 18, 39, 64, 65, 95; pl. 29 

platanoides---------------------------------- 61, 79 

raynoldsi. __ ---------------- ____ --------- ____ 36, 

39, 63, 64, 65, 66, 75, 78, 80; pis. 30, 31, 66 

integrifolia ______________________________ 64,68 

regularis _____ ---- ______ ---- __ . -------------- 65 

rhomboides .. ______________ . _________ --- _ --- _ 79 

sp ___________________________ -------- ___ _ __ _ 79 

Plateau Valley, Colorado_______________________ 26 

Plateau Valley beds____________________________ 8, 26 

Platycarya .. --------- ____________________ ------_ 56 

platyphylla, Cornus_____________________________ 84 

plicatus, Oredoxites______________________________ 54 

Paloreodoxites _____________________ 38, 53, 54; pl. 15 

Poacites _ --------------------------------------- 53 nielseni. ___ ---------- ____________ -------- __ _ 4 7 
116 

Page 

Podocarpoxylon dakotense. ---------------------- 94 Podocarpus _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 49 

campbellL _ --------------------------------- 48 Podozamites latipennis__________________________ 47 

Poison Canyon formation.--------------------- 24,25 

Poison Spider Creek, Wyoming________________ 22 

Polecat Bench__________________________________ 7,14 

Polecat Bench formation ___ -------------------- 7, 23 

polymorpha, Marchantia________________________ 40 

polymorphum, Cinnamomum. _ -------------- 61, 66,67 

Elaeodendron ___ ---------------------------- 72 

Polypodiaceae __________ --------- __ _____ _ __ ___ __ 41 

polypodioides, Dryopteris________________________ 43 

Lastrea (Goniopteris) ________________________ 37,44 

Polyporites sequoiae.---------~------------------ 40 polysepala, Calycites ______________________ 39,89; pl. 4 

Poplar River, Montana ___ --------------------- 18 

populicola, Sclerotium___________________________ 40 

populifolia, Grewiopsis ______ .:.___________________ 65 

populifolius, Celastrinites________________________ 76 

Populites ____________ -------- _____ ----~ ___ __ _ __ _ 81 

populoides, Ficus. __ ---------------------------- 61 Phyllites____________________________________ 81 

Populus ________________________________________ 62, 71 

acerifolia _______________ ~-------------------- 70, 78 

anomala _____________ -------- __ _ __ _ _ _ __ __ _ _ _ 73 

arctica _______ ----- __________________________ 61, 70 

balsamoides eximia. _ ------------------------ 65 cordata______________________________________ 82 

craspedodroma. ----------------------------- 78 daturaefolia _______________ ------- __ ___ ___ __ _ 66 

denverensis____ _ __ ______ __ __ _ __ ___ __ ____ ____ _ 79 

dil!torta ____ -------- __ __ _ _____ __ __ __ _____ _ __ _ 61 

gaudinL _______ __ __ ____ _ ___ __ __ __ _____ _ __ __ _ 70 

glandulifera __ _________________ -- _ __ __ _ ___ __ _ 70 

grewiopsis ____ ------------------------------ 86 inaequalis______ __ __ ____ __ __ _____ _ __ __ __ _ _ __ _ 73 

knowltonL_ __ __ _____ _ __ __ _ _ ____ __ ______ __ __ _ 79 

lacoeana __ ---------------------------------- 78 

meedsiL __ ---------------------------------- 59 monodon____ ___ ____ _ _____ __ _ _____ __ __ _ __ _ __ _ 63 

nebrascensis ______________________________ 70, 71, 78 

nervosa. __ ---------------------------------- 65 newberry;_ __________________________________ 70, 78 

richardsonL __ ------- ________________ ----- _ _ 82 

smilacifolia ____ ----------------------------- _ 70, 78 subrotundata __ ___________________ ------ __ ___ 611 

ungerL _ ------------------------------------ 77 whitei_ _____ ____________ ------ __ __ ______ __ _ __ 78 

wilmattae ____________________ ----- _ __ __ _ _ _ __ 71 

zaddachL _ ---------------------------------- 78 

sp _____ -- __ ------------ __ ------- ___ -~ ------ _ 78 

Porcupine Dome, Montana_____________________ 12 

porcupinianum, Rhamnacianum __ -------------- 94 

posttrinervis, Ficus_----------------------_______ 61 Potamogeton_ __ _ _ __ __ ___ _ _ _ ____ __ _____ __ _ __ _ _ ___ 52 

nordenskioldL __ _ __ ____ __ __ __ _____ _ __ __ ___ __ 51 

Powder River------------------------------- 16, 20,21 

Powder River Basin, Wyoming __________ 11, 12, 21, 23 

Powder River region, Montana_________________ 16 

Powder River Valley, Montana________________ 16 

powelli, Sabal_ ____________________ 38, 53, 54, 55; pl. 16 

Sabalites__ _ __ ___ _ __ _ _ __ _ _ _ __ _ _ _ _ __ _ _ _ _ _ __ _ _ _ 55 

praeangustiloba, Quercus ___ --------------------- 61 

praecursor, Ulmus______________________________ 87 

praecursoria, Malapoenna_______________________ 68 

Tetranthera_ -------------------------------- 68 

praegroenlandica, Quercus_______________________ 60 

praeplanicostata, Ficus__________________________ 62 

praetrinervis, Ficus ______________________________ 61,62 

preartocarpoides, Ficus _______ --------___________ 62 

preauriculata, Salvinia ______ ------- __________ 11, 20,21 

Preissites wardi_ _________________________ 38, 40; pl. 4 

wardiL ---------------------- __ __ __ _____ ___ _ 40 

Price River formation__________________________ 27 

primaevus, Titanoides___________________________ 7,19 

primero, Asplenium_____________________________ 42 

primigenia., Laurus __________________________ 67, 73,74 

prisca, Betula_________________________________ 57 

priscipites, Quercus ________ --------------------- 93 

INDEX 

Page 

pristina, Fontinalis __________________________ 92; pL 69 

Prnnns. ___ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ 74 

problematica, Ficus _________ -------------------- 62 

planicostata _______ --------------------- _ 61 

problematicum, Viburnum______________________ 80 

Propalmophyllum liasinnm______________________ 54 

Protofi.cus inaequalis. _ ------------- _____________ 59, 72 

nervosa, Ficus_______________________________ 94 

protolutea, Nelumbo_____________________________ 69 

protoluteus, Nelnmbites_ _ _ __ __ _ __ _ _ __ __ _ _ _ _ _ __ _ _ 69 

Protophyllum canadensis________________________ 61 

sp_ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ ____ _ ___ _ _ _ _ ___ _ _ __ _ _ __ _ _ 61 

Prunus ______ ___ -------------------------------- 73,74 careyhurstia _______________________ 39, 73, 74; pl. 42 

coloradensis __________________________ 39, 79; pL 42 

cormgis ______________________________ 39, 74; pL 67 

dakotensis___ _ __ _ _ _ __ _ _ _ __ _ _ _ _ _ __ _ _ _ __ _ _ _ _ _ _ _ 56 

denverensis____ _ _ _ _ _ _ _ __ _ _ _ _ _ ___ _ _ __ _ __ _ __ _ _ _ 73 

mclearni_ _________________ __________________ 39,7 4 

perita ________________________________ 39, 74; pl. 43 

pristina___ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ __ _ __ _ __ _ 74 

scotti_ ____ ___________________________________ 59, 74 

pseudoacacia, Robinia___________________________ 75 

pseudopennaeformis, Pteris ______________________ 41,44 

pseudopopulus, Ficus ________________________ 61, 62,78 

Pseudotsuga miocena____________________________ 94 

Psilotu m inerme _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 68 

Pteris goldmani_________________________________ 43 

gronlandica_ ----------------------- _______ _ _ 45 

inaequilateralis ______________ --------- _______ 45 

linearis _ _ _ _ __ _ _ _ _ _ _ _ __ __ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ 41 

oeningensis. ___ ----------------------------- 45 

pennaeformis. ____ -------------------------- 44 peseudopennaeformis ________________________ 41,44 

russelliL. _ __ __ _ ___ __ ___ _ __ _ ___ ___ _ __ _ ___ __ _ 44 

Pterocarya_ _ _ _ _ __ __ ____ __ ______ __ _______ _ __ __ __ _ 73 

denticulata __ ___ ------ ______ ______ ___ _ ______ _ 59 

glabra ______________ ------ _ ------- 38, 57, 95; pL 18 

hispida ___________________ 38, 57, 95; pis. 18, 19, 68 

paliurus __ ---------------------------------- 57 

retusa. _____ -------------------------------- 86 

septentrionale.------------------------------ 59 Pterospermites cordatus ___________________ 39, 8~; pl. 41 

cupanioides _ _ __ __ _ _____ ___ _ __ ____ ___ ___ ____ _ 87 

dawsonL ____ ------------------------------- 62 

minor _____ --------------------------------- 87 pen hall owL_________________________________ 62 

vagans __ _________________ --------- _ ____ __ ___ 49 

whiteL-------------------------------------- 57,87 

pterospermoides, Celastrus ___ -------------------- 72 Pterospermum _______ -- ___ _ __ _ ___ __ ____ ___ _ __ __ _ 82 

Puerco formation_______________________________ 26 

pulchella, Castalia.--_-------------------------- 69 Nymphaea __________________________________ 39,69 

pulchra, Radermachera _______________________ 92; pL 68 

pumilum, Lygodium____________________________ 45 

pungens, Aralia_________________________________ 61 

Artocarpus ____ ----------------------------- 60 

purdonensis, Quercus _______ -------------------- 79 

puryearensis, Cassia. __ ------------------------- 67 Ficus ________________________ ---- ____ ------_ 71 

Sophora ____ -------------------------------- 80 

Q 

quercifolia, Ulmus ____ -------------------------- 80 Quercus _____________________________________ 36, 58,59 

angustiloba____ ___ _ _ _ _ __ __ _ _ __ ___ ___ __ _ _ __ _ _ _ 61 

artocarpites ___ ------------------------------ 59 

asymmetrica __ ------------------------------ 38,58 

atava ________ ------ ________ ------ ______ ---- _ 58 

bicornis. _ __ ___ ___ __ _ _ __ __ ___ _ __ ____ __ __ _ ___ _ 62 

castanopsis. ___ ----------------------------- 60 celastrifolia _____ ______________ -------- __ __ _ __ 76, 79 

charpenterL __ ------------------------------ 59 

conjunctiva__________________________________ 59 

dawsonL ____ _____________________________ -- _ 62 

drymeia___ _ ________ __ _ _ _ ___ __ _____ ___ __ __ __ _ 58 

dubia. _ _ __ _ __ __ ______ __ ____ __ __ ____ __ __ ___ _ _ 58 

fisheriana ______________ -------- ________ ----- 75 

furcinervis. __ ------------------------------- 58, 59 

Page 

Quercus--Continued 

greenlandica ___________ 38, 58, 59, 95; pls. 19, 22, 40 

groenlandica. _ ------------------------------ 58 juglandina ___ ______ -~ __ _ ___ ___ _ _ _ __ _ ___ _ ___ _ 59 

laharpi .. _ ___ _ _ _ _ _ __ _ ___ __ _ _ _ _ __ _ __ ___ ___ ___ _ 59 

laurifolia_____ __ _ _ _ _ __ __ _ _ ____ _ _ _ _ _ _ _ _ _ __ _ __ _ 59 

leonia __ ------- ___________________________ 73, 76, 86 

lyelli_ ___ _____________ ------- _ _ _ _ _ _ _ _ _ _____ __ 59 

macneili. ---------------------------- 38, 58; pl. 19 meriani___ ___ _ _ __ _ _ _ _ __ __ __ __ _ _ ___ _ _ __ _ __ _ _ _ 59 

negundoides____ __ ____ __ _ _ _ _ _ _ _ _ __ __ _ ____ _ _ _ _ 65 

neomexicana_ _ _ ___ __ ___ _ _ _ _____ _ _ __ _ __ ___ _ _ _ 63 

neriifolia_____ __ _ _ _ _ _ _ _ _ _ __ _____ __ _ _ _ _ __ __ _ __ 86 

penhallowi__________________________________ 59 

platania ____________________________________ 62, 65 

praeangustiloba_ _ _ ___ __ _ _ _ __ __ ___ __ __ _ ____ _ _ 61 

praegroenlandica_ _ _ _ _ _ __ _ _ __ _ __ __ __ __ ___ ___ _ 60 

priscipites_____ __ _ _ _ _ _____ __ _ __ __ __ _ _ _ ___ ___ _ 93 

purdonensis______ _ ___ __ ___ __ _ _ _ _ _ __ __ _ _ _ __ _ _ 79 

ratonensis ____ _______________________________ 67, 71 

simplex ___________________________ ---_______ 67 

steenstrupiana ___ --------------------------- 58 straminea _______________________ ------- _ _ _ _ _ 86 

sullyi_ ____________________ 38, 59, 95; pis. 23, 27, 57 

valdensis ____ ____________________ --- ______ --- 56 

viburnifolia _______________________________ -_ 80 

whitei. ___ _______ ----- __________ -- _____ --- _ -- 80 

whitmani. _____________________________ ---- _ 86 

yulensis ______________________________ 38, 59; pl. 19 

quinque folia, Parthenocissus __________ ----------- 79,81 

R 

racemosa, Berrya________________________________ 70 

Nyssa _______________ ----------------------- 70 

Radermachera pulchra _______________________ 92; pl. 68 

Ranch Creek, Montana________________________ 16 

Raton formation _____________________________ 24, 25,28 

Raton Mesa ___ ~-- _____________ --_----- __ ---_--_ 24 

Raton Mesa coal field __________________________ 24,25 

ratonensis, Ficus________________________________ 82 

Laurus __________________________________ --- 67 

Oreodaphne __ _____________________ -___ ------ 72 

Quercus ____________ ------ ___ --------------- 67, 71 

Ravenscrag formation__________________________ 19 

ravenscragen.~is, Cercocarpus _____________________ 39,73 

Fiburrium marginatt£m_ -------------------- 80 

raynoldsi integrifolia, Platanus ____ ----- __ ------- 64, 68 

Platanus_---------------------------------- 36 

39, 63, 64, 65, 66, 75, 78, 80; pis. 30, 31, 66 

Reading beds ___________ ----- ___ ---------------- 29 

rectinervis, Rhamnus _____ .c.--_----------------- 77 

recurvatus, Cunninghamites. _ ---·---------------- 48 

Red Desert, Wyoming_------------------------ 8, 22 

reesidei, Aralia ______________ -____ --------------- 6."i 

regalia, Magnolia _________________________ 39, 72; pl. 39 

Osmunda.-----------------------~---------- 41,45 

regularis, Platanus. ______ -----------------------

reichenbachi, Sequoia._-------------------------- remotidens, Asplenium _________________________ _ 

resurgen,q, Laurus _____ -__ -----------------------

reticulata, Ilex.- ____ -----------------------------

retusa, Pterocarya __ ---- __ --- _ -------------------

retusoides, Phyllites.----------------------------

Rhamnaceae.----------------------------------

65 

49 

41 

76 

59 

86 

61 

77 

Rhamnacinium porcupinianum_- _ -------------- 94 

triseriatim. ___ ------------------------------ 94 

rhamnifolia, Cornus ___ -------------------------- 68 Ulmus ___________________________ 38, 60, 95; pl. 24 

Rhamnites concinnus ____ ------------------------ 87 

marginatus. __ ------------------------------ 87 

rhamnoides, Ficus ___________ -------------------- 77 Rhamnus ____________________________________ 77, 78,94 

brittonL. _____ --------------- _ -------------- 77 

cannonL _____ _______ --- ____ -- ___ ------------ 62 

cleburni_ ______________________ 39, 62, 77, 80; pl. 42 

discolor ______________________ --------------- 77 

eridani_ ___ _ ------ ____ ----------------------- 68 goldiana __________________________ 39, 77, 95; pl. 48 

goldianus. _ ------------------------------ 62, 77, 87 hirsuta _______________________________ 39, 77; pl. 49 
Page 

Rhamnus-Contlnued 

inaequalis ___ _____________ ---·--- __ ___ _ _ __ _ __ _ 77 

marginatus apiculatus_______________________ 78 

marshallensis ___ ---------------------------- 77 newberryi. _________________________ --------- 94 

parvifolius. _ -------------------------------- 94 

rectinervis ___ ----------------- __ ----------- __ 77 

salicifolia. _ --------------------------------- 77 

Rhoicissus. _ ------------------------------------ 81 

usa mbarensis _____ -------------------------- 81 Rhoiptelea _ _ ___ __ __ _ _ __ ____ __ __ __ __ __ __ _ ____ ___ _ 56 

rhomboidalis, Carpites___________________________ 90 

rhomboides, Platanus _______ --------------------- 79 

Rhus. __ ---------------------------------------- 76 coloradensis _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 56 

evansiL------------------------------------- 57,58 holbOlliana _______________ __________ ------- __ 83 

nervosa. __ ---------------------------------- 60 

unit us _________ ---------_---------__________ 60 

viburnoides ____ ------------------------------ 65, 80 

Rhytisma boreale________________________________ 40 

richardsoni, Euphorbocarpum____________________ 89 

Ficus ___________________________ -----------_ 63 

Populus _____ -----------------------________ 82 

Sophora. __ --------------------------------- 85 

Viburnum ___ ------------___________________ 80 

richardsoniana, Dryopteris____ _ __ __ _ _ __ _ _ _ _ _ _ _ _ _ _ 42 

Rifle Gap, Colorado _________________________ 13, 25,26 

rigida, SabaL ------------------------------- ___ _ 53 

Rio Chico formation____________________________ 29 

Robinia _____________ --------------------________ 75 

mesozoica ___ ___________ ._ _ __ _ _ _ _ _ _ _ _ _ _ __ __ _ _ _ 75 

pseudoacacia. _ ___ __ _ __ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __ __ _ _ _ 75 

wardL -------------------- ------~---- 39, 75; pl. 43 

Hock Creek oil field____________________________ 21 

Rock Springs dome_____________________________ 22 

rocklandensis, Cissites ____________________ 39,81; pl. 27 

Rogers and Lee, quoted________________________ 10 

Roland coaL_--------- __ ------------------_____ 21 

Rosaceae.______________________________________ 73 

Rosebud coal field------------------------------- 16 

rostellatus, Carpites. ______ ---------------------- 95 

rostrata, Corylus. _ -------- __ ---------------- ____ 57 

fossilis, Corylns. ----- _______ -------- _____ ___ 57 

rotundifolia, Hedera _______ _____________ ---~----- 63 

Magnolia ________________________________ 39, 63,72 

rotundifolium, Viburnum_______________________ 80 

Royal coal bed_________________________________ 21 

rnbellnm, Sclerotiltm _________ ___ --------- _______ 40 

rnbra, Vitis _____________ --------- _____ __ ____ ____ 82 

rnbrodiscum, Nuphar ___ ------------------------ 69 

H.uby formation._______________________________ 26 

ru.gosa, Celtis _________ ._---------_______________ 87 

Juglans ______ ____________________________ 56, 85, 86 

Sabal____ __ _ _ _ _ _ _ _ _ __ _ _ _ ___ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 54 

mssellii, Pteris__________________________________ 44 

s Sabal campbellL -------------------------------- 54 communis__________________________________ 53 

grandifolia. __ _ _ _ _ _ _ _ _ _ _ __ __ _ _ _ _ _ _ __ _ _ _ __ __ __ 54 

grayana. ------------------- 38, 53, 54, 95; pls. 15, 16 imperialis _________________________ 38, 53, 54; pl. 14 

inqltirenda ___ _________ ------------ ___ -_______ 54 

leeL ------ __ ___ ______ __ ___ __ ___ ____ ________ _ 54 

montana.___________________________________ 54 

p.o~ellL _______________________ 38,53,54,55; pl. 16 

ngtda ____________ _____ --------------------- _ 53 

rugosa _____ ---------________________________ 54 

wngerL------------- _ -- ___ -_________________ 54, 55 

SP------------------------------------------ 54 Sabalites campbelli______________________________ 54 

eocenica____ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ 54 

j7orissanti___ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 54 

frnctifer ------------------- ________ ---------- 70 

orayanus _______ ----------------------------- 54 

1JOWelli_- _ ---------------------------------- 55 

sabiniana, Libocedrus. __ -------------- _ --------- 53 

Saccoloma gardneri_ _________________ 38, 41, 43; pls. 5, 6 

sp_ -------------------------- ______ __ ___ _ _ __ 43 

sn121 o - 62 - 6 

INDEX 

Page 

Sagitta ria _____ ------- ____ ----------------------- 49, 52 megasperma __________________________ 38, 52; pl. 6~ 

St. Mary River formation ______________________ 18,19 

Salicaceae .. ------------------------------------- 55, 94 

salicifolia, Rhamnus_____________________________ 77 

salicoides, Cinnamomum________________________ 61 

Salix. __ ---------------------------------------- 74 aqltilina ______________________________ 38, 55; pl. 61 

Salpichlaena anceps_____________________________ 41 

Salvinia preauriculata ________________________ 11, 20,21 

Sambucu..~----------------------------------- 91; pl. 67 

Sand Creek, Wyoming_------------------------ 20 

San Jose formation __ --------------------------- 26 

San Juan Basin, Colorado _________________ 7, 12,14, 26 

Sapindaceae _________________ ------- ________ ---- _ 76, 91 

sapindiformis, Juglans __ -~---------------------- 71 

Sap indus ___ --- ______________ ------ __ ----------- 76 

affiniB-------------------------- 39, 76, 85, 95; pl. 47 

alatus.-------------------------------------- 76,85 

angustifolia. __ ------------------------------ 76 

berryanus ____ _______________ ---------------- 76 

caudatus ____________________________________ 67, 86 

glendivensis--------------------------------- 76 

grandifoliolus ___ ---------------------------- 76 

membranaceus __ ---------------------------- 76,87 obtusifolius __________________________________ 76,85 

undulatus _____ __________ ------------------- _ 59 

Sapoticarpum latum _________________________ 91; pl. 67 

Sassafras._------------------------------------- 68,84 

ferretianum ____ ----------------------------- 68 

selwynL --- _ ------------------------------- _ 64 

thermale _________________ 39, 63, 66, 68, 95; pls. 33,34 

Saxifragaceae .. ----------------------------- __ - __ 73 

scheuchzeri, Cinnamomum. _ -------------------- 66 

schimperi, Betula ___ ---------------------------- 79 

Ficus ... __ ------------------------ ____ ------- 63 

Geonomites .. __ ------------------------------ 53 Juglans _____________________________________ 56,71 

Schizaeaceae _____ -------------------------- __ -- _ 44 

Sclerotites cypericola _____ ------------------------ 40 

hesperius ____ ------------------------------- 40 

Sclerotium cinnamomL------------------------- 40 

populicola _______ ---------------------------- 40 

rubellum ___ ____ -------- ____________ -------- _ 40 

Scobey lignite field. __ -------------------------- 18 

Scotland, Ardtun Head, Island of MulL_______ 29 scotti, Prunus ___________________________________ 59, 74 

scripta, Andromeda. ___ ------------------------- 85 

scutata, Lemna (Spriodela)______________________ 52 

Spirodela ___ -------------------------------- 52 

sedaliensis, Dombeyopsis _____________________ -_ _ 82 

Selaginella .... __________ ----------------- __ ------- 46 

berthoudL __ -------------------------- 38, 46; pl. 4 

collierL __ ----------------------------- 38, 46; pl. 4 monstrosa _____________________________ 38, 46; pl. 4 

Selaginellaceae ___________________ -------------- _ 46 

Selaginellites mirabilis___________________________ 92 

selwyni, Carpolithus (Ginkgoites) ---------------- 48 

Sassafras ___ -------------------------------- 64 

sempervirens, Sequoia. __ ------------------------ 50 

Senonian group_________________________________ 28 

sensibilia [arctica], Onoclea_______________________ 44 

Onoclea.------------------------------------ 43,44 fossilis, Onoclea _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 43 

Sentinel Butte, North Dakota__________________ 19 

Sentinel Butte member-------------- 5, 13, 15, 16, 19,21 

Sentinel Butte shale _________________________ 16, 19,20 

septentrionale, Pterocarya __ --------------------- 59 

Sequoia ______________________________________ 49, 50,94 

acuminata .. _ -------------------------------- 48 

aj]ini~ _ ---------~--------------------------- 49 

brevi folia ... ----------- ________ --------------- 49 

burgessi_ __ ----- ________ ------------------ __ _ 94 

couttsiae. _ ---------------------------------- 49 

dakotensis ____ ---------------- __ ------ __ ___ __ 15 

gigantea ____ --------------------------------- 50, 94 

lanosdorji_ --------------------- ______ __ __ __ _ 94 

langsdorfiL _ -------------------------------- 49 

lapillipites. __ ------------------------------- 93 

117 

Page 

Sequoia-Continued longifolia_______ ____ _ _ _ _ _ _____ _ _____ ______ ___ 48 

magnifolia. _ -------------------------------- 49 nordenskioldL ___ ___ __ __ __ _ _____ __ __ _ _____ ___ 49 

reichenbachL____ __ __ __ _ __ _ __ __ ___ _ ____ __ _ _ _ 49 

sempervirens___ __ __ __ ________ ____ __ __ __ __ __ _ 50 

sternbergL ---------------------------------- 49 

sequoiae, Polyporites .. _ -------------------------- 40 

Sequoiites ______________ ________________ -------- _ 49 

art us _______ ------- ______________ ------_____ 48 

langsdorjiL _ -------------------------------- 50 

Sequoioxylon burgessL-------------------------- 94 

dakotense .. _ --------------------------------- 94 laramense ____ -- __ --- _ _ __ ___ ____ _ ___ __ _ ___ ___ 94 

montanense. ----------- __ __ ___ _ __ ____ __ ___ _ _ 94 

serrata, Alnus ____ ------------------------------ 72, 86 Aralia ______________ ----- ________ ------_____ 65 

Celastrus ___ -------------------------------- 86 Dryopteris _____________________________ 38, 42; pl. 8 

Eucommia _______________ 39, 72, 77, 95; pls. 44, 45 

Woodwardia latiloba_________________________ 43 

serratus, Celastm.~- ___ --------------- ___ ----- _ -- 72 

lsoetites ___ ____ ------ _____________ ----- __ _ _ _ _ 46 

sermlatum, Elaeodendron. _ --------------------- 72 

sessilijlora, J.falapoenna ___ ------ __ ------------ _ _ 68 

Tetranthera ___________ __ ------------ _ ----- __ 68, 95 

Sevier River valley_____________________________ 27 

Sezanne, Paris Basin___________________________ 28 

sezannense, Cinnamomum ...... ____ 39, 66, 67, 68, pl. 66 

Sheep Mountain coal field______________________ 14 

Siberia. ___ ----------------- ___ ·_---------------- 29 

silberlingi, Acer -------------------------- 39, 76: pl. 46 

simile, Viburnum_______________________________ 80 

similis, Amelanchier ___ ------------------------- 87 

Artocarpus ___________ -------------------- _ _ 61 

simplex, Quercus·------------------------------- 67 

sinense, Tetracentron ______ ---------- __ --------- _ 70 

sinensis. Corylopsis __ -------- _ ------ _ ------- __ -- 87 

smilacifolia, Popultts.------'--------------------- 70,78 Smilax grandifolia ______________________________ ._ 70 

inquirenda _______ --------------------------- 66 

smithsoniana, Ficus ... --------------------------- 67 

Smoky Butte, Montana __________ -------------- 12 

socialis, Lauru.~------------------- 39, 67, 73, 85; pl. 27 

solitarium, Viburnum ____________________ 39, 90: pl. 67 

Somber beds. __ ----------------------------·---- 1, 6, 9 

Sophora purvearensis. __ ------------------------ 80 

richardsonL _________ -------- ______ ------ _ -- 85 

sorbifolia, Ulmus________________________________ 88 

Sorbus grandifolia_______________________________ 60 

South Dakota, BuffalO------------------------- 15 

South Park, Colorado__________________________ 25 

South Table Mountain, Colorado______________ 24 

Sparganiaceae .. _________ · ______________ ---------- 50 

Sparganium __ ---------------------------------- 50 

antiquum ______ ~------------------ 38, 51, 64; pl. 14 

globipites .. __________ ----------- ___ ---------- 93 

stygium ___ --------------- _ ------------------ 51 valdense _____ _______________________ --------- 51 

speciipites, Alnus. _____ ------ ___ ---------------- 93 

speciosissima, Corn us. __ ------------------------ 84 

Ficus ______ ------_----- _________ -------_---- 63 

Tilia .. ____ ------------- ____________ --------- 82 

speciosum, Viburnum _____________ ·______________ 80 

spectabilis, Ficus .. ------------------------------ 77 Sphaeria annulifera_____________________________ 40 

arctica. _____ ------- ________________ --------- 40 

interpungens _____ _______________ ------------ 40 

Sphagnum antiquasporites_______________________ 93 

Sphenopteris (Asplenium) elongatnm. __ --------- 44 

(Dennstaedtia) burlingL -------------------- 42 

jorgensenL. ______________________ ---------- 44 

spinosus, Carpolithes .. ________________ ---------- 39, 89 

Spiraxis bivalvia __________________ --------------- 92 

Spirodela scutata ___________________ ------------- 52 

Lemna_ -------------------------------- 51 

splendens, Euonymtts __________ -- __ -_ ----------- 72 

Spotted Horse coal field________________________ 21 

Spring Creek, Montana________________________ 16 

squarrosa, Ficus ________________________ -------- 63 
118 

Pag~ 

stagnum, Leguminosites_________________________ 84' 

Stanton, T. W., quoted________________________ 6·: 

stantoni, Vitia. __ -------------------------- __ 15, 80,81 

Staphylea minutidens_ -------------------------- 39,75 

Staphyleaceae ________________ -------------- ____ 75 

statenensis, Androvettia. _ ----------------------- 50 

steenstrupi, Cissites_____________________________ 82 

steenstrupiana, Quercus.------------------------ 58 

Sterculia berryana _______ ------------------------ 65.· coriacea___ _ _ _ _ _ __ __ _ _ __ __ _ _ _ _ _ __ __ _ _ __ __ __ __ 61 . 

heterodonta___ __ __ __ _ _ __ __ _ _ _ _ _ _ _ _ __ _ _ _ __ __ __ 65 · 

Sterculiaceae. ______________________ ---- __ _ _ __ __ 82.' 

sternbergi, Sequoia. _______________________ "_____ 49 

stevensoni, Betula _____________________ 38,57,95; pl. 20 

Stillwater River, Montana_____________________ 16 

straminea, Quercus_---------------------------- 86 striata, Nelumbites. ___ _ _ _ _ _ __ _ _ _ ___ _ _ _ _ __ __ _ _ _ _ _ 83 

striatus, Nymphaeites. _ ------------------------- 83: studeri, Corn us _________ --- __ --- _____________ 63, 67, 77-

stygium, Sparganium. -------------------------- 51 subcylindricum, Palmocarpon _____________ 39,90; pl. 67 

subfalcatum, Dryophyllum. _ -------------------- 15,59 

subrotundata, Populus __ ------------------------ 65 subtruncata, Ficus_ ______________________ 39,63; pl. 25 

sulcatus, Carpites. ------------------------------ 87 

sullyi, Quercus.---------------- 38, 59, 95; pis. 23, 27,57 

supercretacea, Anemia. ___ ------------__________ 44 

Swauk formation ___________________ ------------ 28 

sylvatica, Nyssa _________ ----- ________ --------_-- 85 

sylvestris, Agatha mas____________________________ 4 

symbolicus, Tetraclaenodon ______ ---------------- 8, 17 

T 

Tatman formation.---------------------------- 23 taurina, Juglans ______________________ 38,56, 77; pl. 56 

taurinensis, Aralia._____________________________ 59 

Celastrus ___________________________________ 65, 72 

Taxites olrikL. _ _ _ _ __ __ _ _ _ __ __ _ __ __ _ _ _ _ _ _ _ _ _ _ _ _ _ 50 

valid us_____________________________________ 50 

Taxodiaceae .. __________________________________ 49 

(Taxodites) tinajorum, Elatocladus _______________ 49,50 

Taxodium_ ---------------------------------- 49, 50,94 

distichum _________ ----------- __ --------- ___ _ 94 

distichum miocenum_ ----------------------- 49,50 

d?tbiu.m. ___ --------------------------------- 50 

hiatipites ______ -------------- ___ ------------- 93 

occidentale. ________ ------------------------_ 49 

olriki_ ________________________ 38,50,95; pls.10, 11 

Ta.r:oxylon sp_ ---------------------------------- 94 

Taylor marL.---------------------------------- 27 

Teleodu.~---- ------- ____ -------- __ ___ __ ____ _ _____ 26 

tenuifolia, Grewiopsis _ -------------------------- 79 

Nelumho. __ -------- __ --------- ___ __ _ _______ 69 

tenuifolium, Nelumbium ________________________ 39,69 

tenuinert•is, Magnolia. __ ------------------------ 71 

tenuirachi8, Geonomites. __ ---------------------- 53 tertiaria, Parthenocissus _____ ------ __ _ _ ______ _ ____ 82 

tesselatus, Cyperacites___________________________ 55 

Tetracentron sinense. --------------------------- 70 

Tetraclaedon sp. --------------- _____ ------------ 7 

Tetraclaenodon symboliws_______________________ 8, 17 

Tetranthera praecursoria ____________ ____ --------- 68 

sessilijlora. _ --------------------· _____ ------ 68, 95 

Texas, Fort Davis______________________________ 27 

trans~Pecos region__________________________ 27 

Thanet sands----------------------------------- 29 

thelypteroides, Alsophila ________ ___ ------------- _ 43 

thermale, Sassafras _______ --- 39, 63, 66, 68, 95; pis. 33, 3:1 

thermal is, Juglans. _ ---------------------------- 63,68 

Thrinax. _______ -------------------------- __ ____ 55 

dorfi------------------------------ 38,53,55; pl. 14 

eocenica ___ _______ ------------------- __ _____ _ 55 T huites interruptus_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 5J 

T huja _____ ----------- ___ __ __ _ _____ ____ __ _ _ __ _ _ _ 50 

colgatensis _____ ______ ------- __ __ _____ _ __ ___ __ 51 

interrupta ______________________ 15,38,51,95; pl. 10 

Thujopsis europaea_ ---------------------------- 51 

Tiffany beds.---------------------------------- 26 Tilia_ _ _ _ ___ _ __ __ __ _ _ ___ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ __ __ _ 62 

anti qua.-----------_________________________ 86 

INDEX 

Page 

Tilia-Continued 

oregona. __ ------- _________________ ----- __ _ __ .57 

speciosissima ... _____ . _ ---------------------- 82 

weedi. _________ ------ _____ ----------------- _ 87 tiliaefolia, Ficus _________________________________ 63,65 

tilioides, Viburnum ____________________ 39, 86, 87; pl. 67 

tinajorum, Elatocladus (Taxodites) _______________ 49,50 

Titanoides primaevus____________________________ 7, 19 

Tococa ____ ________ ------- _____________________ . _ 84 

Tohachi shale ..... ------------ ___ ---------______ 27 

Tongue River---------------------------------- 20,21 

Tongue River Basin____________________________ 23 

Tongue River member ______ 5, 6, 10, 13, 14, 16, 18, 19 

torelli, Hemitelites ___ _____ ------ __ ___ _____ __ _____ 42 

Torrejon formation_____________________________ 7, 26 

Torreya _____ ___ -------- ___ -------------- ___ __ _ __ 49 

borealis. ___________________ ----------------_ 48 

torreyi, Myrica ______________ -----------_________ 83 

Myriophyllum _______________________ 39, 88; pl. 50 

Torreyites tyrrelliL _______ --------------- ___ ____ 48 

Trapa ______________________ ------- ___ _ ____ ____ _ 84 

anyulata ___________ 39, 40, 83, 84, 88, 91; pis. 58, 66 cuneata. _ __ _ _ __ ___ ___ __ _____ _ ____ _ _____ _ ____ 83 

microphyllum _____ _____ ------ _________ ------ 83 

paulula ______________________________ 39, 84; pl. 58 

Trapaceae. ____ ---------- __ __ __ __ _ ___ ___ _____ _ __ 83 

trianoulosus, Carpites___________________________ 95 

Triboleti, Ilex. ____ . ___ . _________ ------- _ _ _ _ _ _ _ __ 59 

Tricentes sp _______ ----------- ____ _ _ __ __ _ _ __ __ ___ 7 

Triceratops ___ -------------_---------------- ____ 10, 15 

tricuspidata, Cissus. __ ----------- _______ -------- 78 

Parthenocissus (Ampelopsis)________________ 78 

tricuspidatum, Acer trilobatum. ----------------- 64,65 

Triletes mirabilis________________________________ 92 

trillioides, Phyllite.,______________________________ 78 

triloba, Aralia ________ --------------------------- 75 

Negundo ____ _ -------------------- __ ____ _____ 75 

trilobat?tm tricuspidatum, Acer _ ----------------- 64,65 

trinervi.~, Ficu.~--- ___ --------- _ __ _____ _ _____ __ _ __ 61 

Macclintockia __ _______________ _.______ __ _ ____ 88 

Phyllites .. ________________ -------___________ 81 

Viburnum _________ ------------------------_ 78 

triner1mm, Viburnum antiquum_________________ 87 

Trinidad sandstone_____________________________ 24 

triphylla, Achlys________________________________ 81 

Winchellia---------------------------------- 80,81 

triseriatim, Rhamnacinium______________________ 94 

trivialis, Dombeyopsis. _. ------------------------ 81 

Trochodendroides arctica .. _. _________________ . _ _ _ 70 

nebrascensis ______ ___ --------- _ __ ____ ___ __ _ __ 78 

truncata, Ginkgo __ ------------------------------ 4!1, 90 trnncatum, Palmocarpon ________________________ 39,90 

Tullock Creek coal field ________________________ 10,14 

Tullock member __________ 5, 6, 7, 13, 14, 15, 16, 18, 20 

Turtle Mountains, North Dakota______________ 18 

Tuscher rormation ______________________________ 26,27 

Typha latifolipite.L_____________________________ 93 

tyrellii, Abietite,, _____ ____ ------- __ ____ __ __ _ ___ __ _ 48 

Torreyites _____ ______________ . _ _____ __ _ ___ ___ 48 

u 

ucranicum, Cupressinoxylon_____________________ 93 

Uintacyon. ______ ---------- _ _____ ____ _ _ __ _ ___ __ _ 26 

Ulmaceae _______________ . ______ ------ _________ ._ 59 

ulmoides, Rucommia____________________________ 73 

Ulmus__________________________________________ 60 

antecedens _ _ __ __ ___ __ __ __ _ __ __ ___ _ _ _ _ __ _ ___ _ 60 

orandifolia ______________________ ----- _ _ __ _ _ _ 94 

japonica. _______________ . _ --------. ____ _ ___ _ 60 

minima_____________________________________ 60 

orbicularis ________________________ -----~____ 60 

praecur.,or __ --------- ___ --------------- _____ 87 

querci.folia _____ ------ _____ ------------------- 80 rhamnifolia _______________________ 38, 60, 95; pl. 24 

sorbifolia __ ----------- __ ------- _ _____ _ ____ _ _ _ 88 

wardii_ __ ___ ------- _______ --------------- __ _ 60 sp. _ ~ -------- __ ___ _ _ __ ___ __ _ ___ __ _ ____ _____ _ 88 

uncata, Ficu.s_ ____________________________ 39,63, 72,77 

undulatus, Sapindus. ___________________ ------- _ 59 

Page 

ungeri, Ca11tanea.-- ----------------------------- 5S Geonomites. _______ ------- ___ _ _____ _ ___ ___ __ 54 

Glyptostrobus. __ ___ __ ____ _ ___ _____ __ _____ _ __ 49 

Juglans _____ ----------------------- _ _ ____ _ __ 71 

Populus _____ ------------___________________ 77 

Sabal. --- _ --------- _______ ----------------- _ 54, 55 unit us, Rhus ____ -------_________________________ 60 

ursina, Parthenocissus ___________________ 39, 81; pl. 48 

usambarensis, Rhoicissus________________________ 81 

Utah, Coalville ____ -----------------____________ 27 

Henefer ____ -------------------_____________ 27 

Summit County____________________________ 27 

utahensis, Carpites ____ ___ ---------------- _ ------ 95 Laurus. ------- _ ______ _____ ___ _____ _ ___ _ _ _ __ 63 

v vagans, Pterospermites _______________________ • __ 49 

valdense, Sparganium___________________________ 51 

valdensis, Quercus _____ ------------______________ 56 

validus, Taxites. _____ -------- __ ____ ___ _ __ _ _ ____ _ 50 

venosa, Phyllites. _____________________ ._________ 87 

venusta, Lindera __ ----------- __ ------ ____ _ _ __ _ __ 68 vera, Cinnamomum_ __ ___ __ _ ___ _ _ __ _ _ _ __ ___ _ __ _ _ 84 

veripites, Carya_ _ _ ___ _ ___ ____ _ __ ___ __ _ ___ _ __ _ ___ 93 

Vermejo formation. ____________________________ 24, 25 

Vermillion Creek, Wyoming____________________ 22 

verperalis, Asimina ______________________ 39, 78; pl. 46 

verrucosus, Carpites. ----- _ ----- _ _ __ _ ____ ___ __ _ __ 52 

verus, Melastomites .• _ ------------------------ __ 84 

viburnifolia, Ficus·----------------------------·- 86 Grewiopsis ________ _______ ------ _ ------- _ __ __ 65 

Quercus ____________ ----------- __ -----______ 80 

vihurnoides, Rhus ___ ---------------------------- 65,80 

Viburnum ___________________ 7 59, 62, 79, 81, 86, 87, 88,90 

acerifolium. ________ -------------------- ___ _ _ 79 anceps __ ___________ ------ ____ ___ _ __ _ ___ _ ____ 78 

antiquum ____ 18, 36, 39, 65, 67, 80, 86, 87, 95; pis. 63, 64 

trinervum_ ___ ____ ____ ____ _ _ __ ____ _ ___ __ _ 87 

asperum. ___________ 15, 18, 39, 57, 87, 95; pl. 64 

betulaefolium _________ ---------- __ __ _____ _ _ __ 86 

castrae .. ___ ---~ ________________ ---------- _ _ _ 87 

contortum __ _____________________ --------- _ _ _ 80 

crassum. ___ -------------------------------- 80 

cuneatum __ _ -------------------------------- 62 

cupanioides . .' __________ 39, 87, 95, pis. 46, 57, 65 

dakotense. __ -------------------------------- 86 

dentoni. __ ---------------------------------- 86 dichotomum ___ ___ ----~- _____ ------- _ --- _ ___ _ 80 

elongatum. ___ ------------------------------ 86 erectum ___ __________________ --- _ --- _ _ _ _ __ _ _ _ 87 

finale.______________________________________ 73 

goldianum_ --------------------------------- 39,90 

lakesL-------------------------------------- 65, 80 

lakesii. _ ----------------------------------- _ 78 lanceolatum __ __ ---- _ --- _ _ __ ____ __ __ ___ ___ _ _ _ 87 

limpidum ___ ------ _____________ ------ _ _ __ __ _ 86 

macrodontum. _ ----------------------------- 86 magnum _________ ----_______________________ 80 

marginatum. ___ ---------------------------- 79, 81 ravenscragensis .. ___ _ _ ____ ____ __ _ _ __ __ _ __ 80 

melaenum ____ ------------------------------ 80 montanum_____ __ ____ __ __ _ __ ___ __ _ _ __ __ __ _ __ 80 

newberrianum. ___ -------------------------- 87 nordenskioldL ______________________________ 86,87 

oppositinerve __ ------------------------------ 86 paucidentatum ______________________________ 87,94 

perfectum __ ________ --- ________ --- _ --- __ ---- _ 86 

perplexum __ -------------------------------- 86 platanoides ___ __________________ ---- _ ____ _ ___ 80 

problematicum. _ ---------------------------- 80 richardsonL _________________ ---------- __ --- 80 

rotundifolium. _____________ ----------------- 80 

simile. ____________ -______ ------------------- 80 

solitarium---------------------- 39, 90; pl. 67 

speciosum ___ _____________ -------------- _ ---- 80 

tilioides--------------------- 39, 86, 87; pl. 67 

trinervis... _ --------------------------------- 78 

whymperL.-------------------------------- 80,86 

sp_----------------------------------------- 80 
Page 

Vincentown sand_______________________________ 28 

virginiana, Hamamelis.------------------------- 72 

viridijluminipites, Carya_________________________ 93 

virletti, Cannophyllites--------------------------- 55 

Vitaceae------------------------------------- 78, 81,82 

Vitis------------------------------ ________ ------ 62, 81 arctica. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ __ _ __ _ _ _ _ _ _ _ _ _ _ 82 

bruneri. __ ------------------------------ __ __ 78 

carbonensis ____ -------------- __ _____ ____ ___ _ 78 

cuspidata _______ ------------------------- ___ 65, 78 

inominata _____ ____ -------------------------- 82 

leei. -- _ --------- ______ ------ _ ------------ ___ 82 

lobata _________________________________ 39, 8!8: pl. 56 

olriki.- ------ ______ --------------- 39, 8!8: pis. 27, 59 

palmata ____________ ----------- ____ ____ ___ __ 82 

platanifolia ___ -------------------- ---------~ 80 

rubra. ----- ____ --------- ___________ ------ _ __ 82 

stantoni_ ___ ___ ------------------ _________ 15, 80, 81 

xantholithensis __________________ --------- ___ 78 

SP------------------------------------ 39,82: pl. 53 vogde~i, Bet1tla. _ ------- __ ___ __ __ ______ _____ __ __ _ 57 

w Ward, Lester, quoted___________________________ 4, !i 

wardi, Acacia.--------------------______________ 75 

Cinnamomum______________________________ 66 

Ficus ______________ ------------------------- 63, 78 

Preissites ______ ------------- ____ ------- 38, 40: pl. 4 

Robinia ___ --------------------------- 39, 75; pl. 43 

wardiana, Aralia________________________________ 64 

tvardii, Celastrus _____ ____ ------ ___________ ------ 72 

.Marchantites. ·------------------------------ 40 

Preissites ___ -------- _______ --------------- __ 40 

Ulmus __________________ -----_----------____ 60 

Wasatch formation ___ 8, 11, 12, 13, 18, 21, 22, 23, 25, 26, 27 

Wasatch Plateau, Utah_________________________ 27 

weedi, Dryopteris________________________________ 43 

Tilia. ____ ------------- ______ ------------ _ _ _ 87 

Whetstone Falls________________________________ 22 

whitei, Populus--------------------------------- 78 

Pterospermites. __ --------------------------- 58, 87 

Quercus. ________ ------------------_________ 80 

Whitemud formation___________________________ 19 

White River formation_________________________ 25 

whitmani, Quercus______________________________ 86 

whvmperi, Viburnum.-------------------------- 80,86 wigginsi, Parataxodi1tm_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 50 

INDEX 

Page 

Wilcox grOUP----------------------------------- 27 wilcoxensis, Apocynophyllum ____________________ 83,86 

Myrica. __ -------------------------_-------- 83 wilcoxiana, Amygdalus__________________________ 74 

williamsi, Legnminosites ________________________ 39,74 

Williston Basin, North Dakota_________________ 12 

Willow Creek formation________________________ 18 

Willow Creek, Montana________________________ 8,17 

Willwood formation____________________________ 23 

wilmattae, Populu8_____________________________ 71 

Winchellia triphylla __ --------------------------- 80,81 wodani, Diospyros _________________ ----- _ __ _ _ _ __ 89 

Wolf Creek, Montana__________________________ 18 

Woodwardia. ______ -------------------------- 37, 43,44 arctica _____________________________ 38, 43, 44; pl. 7 

crenata. _ _ _ _ _ __ _ _ ___ ____ _ ___ _ __ _ _ _ _ _ __ ___ _ _ _ 44 

latiloba. _ _ _ _ _ _ _ _ _ __ __ _ _ __ __ _ __ ___ _ _ _ ___ __ __ _ 43 

minor----------------------------------- 43 serrafa__ _ __ ___ _ _ _ _ ___ __ _ _____ _ _ __ __ _ _ __ _ 43 

maxoni. __ --------- _ --------------------:.___ 43 Woodwardites arcticns__ _ _ _ _ __ __ __ _ __ __ _ __ __ __ __ _ 43 

Woolwich beds ___________________________ --~--- 29 

mrighti, Ceratozamia __ _ ---------------------·---- 47 

Wyoming, Banner______________________________ 21 

Black Buttes Station _____________________ 4, 13,22 

Boars Tusk_________________________________ 22 

Buffalo_____________________________________ 21 

Carey hurst.._______________________________ 20 

Casper _________ ---------------------------- 22 

Clareton:.___________________________________ 20 

Como______________________________________ 21 

Douglas ________ ----------.------------------ 20 

Evanston _______ ------ ________ ---- ____ ------ 8 

Evanston station___________________________ 22 

Glenrock___________________________________ 20 

GreybulL _________ -----------------------__ 8, 13 

Jackson ___________ -------------------------- 8 

Kaycee ______________ --------------_________ 20 

Kemmerer---------------------------------_ 8 

Knobs Station______________________________ 22 

La Barge___________________________________ 22 

Lander ________________________ ------_______ 22 

Lusk ____ ------ ________________ ------ _ ___ _ _ _ 20 

Moorcroft __ -------------------------------- 20 

Point of Rocks._--------------------------- 22 

Rawlins ____________________________ -------- 22 

Rock Springs_______________________________ 22 

119 

Page 

Wyoming-Continued Sussex _______________________________ ------- 20,21 

Wamsutter_________________________________ 22 

·wvomingana, Bauhinia ___________________ 39, 74, pl. 43 

'wyomingensis, Zamia ____________________ 38, 41; pl. 10 

· .womingiana, Ficus______________________________ 63 

X 

:xantholithensis, A mpelopsis____ __ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ 78 

Dryopteris__ __ _ _ __ __ __ _ _ _ _ _ _ _ _ __ __ _ __ __ _ _ _ _ _ 43 

Euonymus___ _ __ __ _ _ __ _ _ _ _ _ __ _ _ __ _ _ _ _ _ _ _ __ _ _ 72 

Vitis~---- ________________________________ ·_ _ _ 78 

Xanthoxylum ___ --------------- _______ _______ _ __ 76 

y 

, Yellowstone National Park_____________________ 22 

Yellowstone River------- 2, 3, 6, 9, 11, 12, 14, 15, 16, 17, 19 

Yellowstone ValleY----------------------------- 2, 6 

.yukonensis, Marchantia_________________________ 40 

yulensis, Quercus __ ---------------------- 38, 59, pl. 19 

z · zaddachi, Populus_______________________________ 78 

: Zamia------------------------------------------ 47 coloradensis __________________________ 38, 47; pl. 10 

wyomingensis ________________________ 38, 47; pl. 10 

'Zelkova_ ---------------------------------------- 60, 73 , . planeroides _______________________ 38, 60; pls. 22, 24 

iinckeni, Flabellaria_____________________________ 53 

·,q;ingiberaceae ____ ------ __________ -------- __ _ ___ _ 55 · '~ingiberites dubius ______________________________ 38,55 

~zizyphoide.~, Paliurus____________________________ 78 

'Zizyphoides columbL___________________________ 70 

•·· mackay;_ ___ -----·--------------------------- 39, 78 

Zizyphus. __ ------------------------------------ 78 

· beckwithiL----------~---------------------- 78 

cinnamomoides ____ --------- __________ ------- 78 

corrugatu~- _ -------------------------------- 78 

daphnogenoides _______ ________________ --- _ --- 78 

distortus __ ---------------------------------- 78 fibrillosus ____________________________ 39, 78; pl. 110 

hendersonL --------------------------------- 78 

hesperius. ___ ------------------------------- 70 

lanceolatus _____ -------------- ____ ------ __ ___ 61 

meeki. __ --------------------- ____ --------- __ 70 

minutus __ ---------------------------------- 61 
PLATES l-69 

[For locality details corresponding to numbers see list on page 30. All figures on plates 4-69 are natural size unless otherwise indicated) 
PLATE 1 

FIGURE 1. Looking north at the western outcrops of the type locality of the Fort Union 

formation, Snowden, Mont. 1931 (p. 3, 12, 19). 

2. South face of clinker-capped Pretty Butte, 6 miles north of Marmarth, N. Dak. 

Arrow indicates Cretaceous-Paleocene contact. 1931 (p. 1, 11). 

3. Looking east at the unconformable Cretaceous-Paleocene contact indicated by 

arrow at. the base of the white sandstone in the center of the photograph, 

2 miles southeast of Black Buttes station, Wyo. 1938 (pp. 11, 13). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 1 

1 

2 

3 

VIEWS OF CRETACEOUS AND PALEOCENE STRATA 
PLATE 2 

FIGURE 1. Looking west at Cretaceous-Paleocene contact indicated by arrow, 13 miles 

south of Fort Peck dam, Montana. 1948 (p. 1, 11). 

2. Pick rests on oyster bed of the Cannonball member of the Fort Union formation, 

150 feet above the Little Missouri River, left bank, 3 miles southwest of Yule, 

N. Dak. 1936 (p. 8, 12). 

:3. Right bank of Little Missouri River at Phelan ranch, 1 mile southeast of locality 

shown in fig. 2. Below the coal seam in the middle of the bank and river level 

is the lowest known tongue of the Cannonball member of the Fort Union formation. It contains species of Corbula and Corbicula. 1936 (p. 8, 12). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 2 

1 

2 

VIEWS OF CRETACEOUS AND PALEOCENE STRATA 
PLATE 3 

FIGURE 1. Looking west at Cretaceous-Paleocene contact indicated by arrow, west side of 

Powder River 8 miles south of Powderville, Mont. 1946 (pp. 11, 16). 

2. Looking west at Cretaceous-Paleocene contact indicated by arrow, 5 miles 

southeast of Glendive, Mont. 1941 (pp. 6, 11, 14). 

3. Looking north at Cretaceous-Paleocene contact indicated by arrow, at base of 

whitish beds in south face of butte, 10 miles north of Baker, Mont. 1941 

(pp. 11, 15). 

4. Looking northeast at Cretaceous-Paleocene contact indicated by arrow, 4 miles 

south of Rock Springs, Wyo. 1941 (p. 11, 22). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 3 

1 

2 

3 

4 

VIEWS OF CRETACEOUS AND PALEOCENE STRATA 

• 
PLATE 4 

FIGURES 1, 10. Marchantia pealei Knowlton (p. 40). 1, loc. 8517; 10, loc. 4293. 

2, 3, 8. Selaginella collieri Knowlton (p. 46). 2, loc. 2420; 3, 8 (X 2), loc. 7004. 

4, 6. Marchantia lignitica (Ward) Brown, n. comb. (p. 40). Loc. 2420, 6 (X 3). 

5. Calycites polysepala Newberry (p. 89). Exact locality unknown. 

7. Preissites wardi Knowlton (p. 40). Loc. 2420. 

9. Selaginella berthoudi Lesquereux (p. 46). Loc. 8779. 

11. Calycites hexaphylla Lesquereux (p. 88). Loc. 9248. 

12, 13. Selaginella monstrosa (Hollick) Brown, n. comb. (p. 46). 12, loc. 8249; 

13, loc. 4031. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 4 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 5 

FIGURBS 1-6. Oogonia of Chara spp. (p. 39). X 15. 1, loc. 229; 2, :3, loc. 9204; 4-6, 

loc. 9205. 

7. Hypnum coloradense Brown, n. sp. (p. 41). Loc. 5738. 

8. Osmunda greenlandica (Heer) Brown, n. comb. (p. 45). Loc. 317. 

9-11, 18. Lastrea goldiana (Lesquereux) Lesquereux (p. 4 7). 9, 10, loc. 325; 11, loc. 

6105; 18 (X 2) loc. 8258. 

12. Mnium montanense Brown, n. sp. (p. 41). Loc. 8519. 

13-16. Allantodiopsis erosa (Lesquereux) Knowlton and Maxon (p. 41). 13, loc. 

318; 14, loc. 6105; 15, loc. 317; 16 (fertile), loc. 325. 

17. Dryopteris meeteetseana Brown, n. sp. (p. 42). Loc. 4694. 

19. Saccoloma gardneri (Lesquereux) Knowlton (p. 42). Loc. 8897. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 5 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 6 

FIGURES 1, 2, 5-7. Dennstaedtia americana Knowlton (p. 42). 1, loc. 4273; 2, 5, loc. 8222; 

6, loc. 8231; 7, loc. 5029. 

3, 4. Blechmtm anceps (Lesquereux) Brown, n. comb. (p. 41). loc. 8678. 

8. Anemia elongata (Newberry) Knowlton (p. 44). Loc. 6084. 

9. Hymenophyllum confusum Lesquereux (X 3) (p. 41). Loc. 317. 

10, 11. Osmunda macrophylla Penhallow (p. 45). Loc. 8519. 

12, 13. Saccoloma gardneri (Lesquereux) Knowlton (p. 43). 12, loc. 8678; 

13, loc. 4029. 

14, 15. Osmunda greenlandica (Heer) Brown, n. comb. (p. 45). Loc. 8519. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 6 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 7 

FIGURES 1, 4. Onoclea hesperia Brown, n. name (p. 43). 1, exact locality unknown; 4, 

loc. 7004. 

2, 3, 5. Woodwardia arctica (Heer) Brown, n. comb. (p. 43). 2, loc. 5030; 3, 5, loc. 

8519. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 7 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 8 

FIGURES 1-6. Equisetum spp. (p. 45), showing underground tubers (1, ~' 5), and stems 

with sheaths (3, 4, 6). 1, loc. 2417; 2, 5, loc. 2414; 3, loc. 7538; 4, loc. 

8240; 6, loc. 8164. 

7, 8, 13, 14. Osmunda macrophylla Penhallow (p. 45). 7, 14, loc. 9085; 8, loc. 9084; 13, 

loc. 8248. 

9, 10. Lygodium coloradense Knowlton (p. 45). 9 (fertile pinnule), loc. 6105; 10 

(sterile pinnule), loc. 8678. 

11, 12, 16. Gleichenia hesperia Brown, n. sp. (p. 44). 11, loc. 8897; 12 (X 2), 16, loc. 

8519. 

15. Dryopteris lakesi (Lesquereux) Knowlton (p. 42). Loc. 317. 

17. Dryopteris serrata Brown, n. sp. (p. 42). Loc. 8678. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 8 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 9 

FIGURES 1-8. I soetites horrid us (Dawson) Brown (p. 46). 1, corm with sporangia tipped 

by ligules, loc. 8548; 2, detached sporangia, loc. 8881; 3 (X 3), corm with 

circle of sporangia containing megaspores and microspores, loc. 8535; 4, 

corm with rosette of leaves, 5, tip of leaf, 6 (X 15), surface pattern of leaf, 

loc. 2420; 7, corm with leaf scars, loc. 5255; 8, corm with rosette of leaves 

showing rectangular depressions, the external indications of collapsed internal air spaces, loc. 2432. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 9 

2 

6 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 10 

FIGuRE 1. Zamia wyomingensis Brown, n. sp. (p. 47). Loc. 5911. 

2-5, 8. Ginkgo adiantoides (Unger) Heer (p. 47). 2, loc. 8256; 3, loc. 8786; 4 (X 5), 

5, loc. 2416; 8, loc. 9109. 

6, 9, 12, 13. Zamia coloradensis (Knowlton) Brown, n. comb. (p. 4 7). Loc. 8551. 

7, 11, 15. Taxodium olriki (Heer) Brown, n. comb. (p. 50). 7, 11, loc. 2420; 15, loc. 

8549. 

10, 14, 16. Thuja interrupta Newberry (p. 51). 10, loc. 5030; 14, loc. 8897; 16, loc. 7989. 
GEOLOGICAL SURVEY PROFESSIONAL P APER 375 PLATE 10 

MEGASCOPIC PALEOCENE PLANTS 593 12 1 0- 62 - 7 
PLATE 11 

FIGURES 1, 2. Fokienia catenulata (Bell) Brown, n. comb. (p. 50). Loc. 7548. 

3, 7-22. Glyptostrobus nordenskioldi (Heer) Brown, n. comb. (p. 49). 3, loc. 8552; 

7, 13 (seed), loc. 8786; 8, loc. 8203; 9, loc. 9192; 10, 18 (cone scales), loc. 

8224; 11 (seed), 16 (cone scale), loc. 8193; 12 (seed), loc. 8549; 14 (cone 

scale), loc. 8677; 15, 19 (cone scales), loc. 8212; 17 (seed and cone scale), 

loc. 8245; 20, loc. 8249; 21, loc. 8206; 22, exact locality unknown. 

4-6. Taxodium olriki (Heer) Brown, n. comb. (p. 50). 4, loc. 2416; 5, loc. 7548; 

6, loc. 8549. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 11 

10 

~ ~ 15 

' 

14 

12 13 

11 

19 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 12 

FIGURES 1-14. Metasequoia occidentalis (Newberry) Chaney (p. 49). 1, 8 (cones), 10 

(foliage and seeds, X 2), loc. 8521; 2, loc. 4984; 3, loc. 8238; 4, 7, loc. 

8550; 5, loc. 4264; 6, loc. 4582; 9, loc. 8551; 11, loc. 8165, 12 (staminate 

amentS) I }OC. 8212; 13, 141 }OC. 2420. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 12 

6 

9 

10 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 13 

FIGURES 1-7. Araucaria longifolia (Lesquereux) Brown, n. comb. (p. 48). 1-5 (cones 

and foliage), loc. 8519; 6, 7 (stomata, X 10, on lowest left-hand needle of 

fig. 6), loc. ·5886. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 13 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 14 

FIGURE 1. Thrinax dorfi Brown, n. sp. (p. 55). Loc. 8256. 

2, 6. Sabal imperialis Dawson (p. 54). 2 (X 7~), loc. 5886; 6, 8188. 

3. Monocotyledonous leaf (p. 53). Loc. 2417. 

4, 5. Sparganium antiquum (Newberry) Berry (p. 51). 4, loc. 8230; 5, loc. 2416. 

7, 8. Amentotaxus campbelli (Gardner) Florin {p. 48). 7, loc. 9109; 8, loc. 4696. · 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 14 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 15 

FIGURES 1, 4, 6. Alismaphyllites grandifolius (Penhallow) Brown, n. comb. (p. 52). 1, 4, 

loc. 8206; 6, loc. 8227. 

2. Canna? magnifolia Knowlton (p. 55). Loc. 9200. 

3, 7. Paloreodoxites plicat1ts (Lesquereux) Knowlton (p. 54). 3, loc. 8188; 7, 

loc. 317. 

5. Sabal gray ana Lesquereux (p. 54). Loc. 8672. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 15 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 16 

FIGURES 1, 3, 8-11. Hydromystria expansa (Heer) Hantke (p. 52). 1, loc. 4897; 3, loc. 

8212; 8, loc. 2420; 9, 11, loc. 8191; 10, loc. 5512. 

2, 5, 6. Sabal powelli Newberry (p. 55). Loc. 8519. 

4. Sabal grayana Lesquereux (p. 54). Loc. 317. 

7. Monocotyledonous rootstock (p. 53). Loc. 8678. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 16 

7 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 17 

FIGURES 1-7. Carya antiquorum Newberry (p. 55). 1, loc. 541; 2, loc. 8206; 3, 6, loc. 

8224; 4, exact locality unknown; 5, loc. 8523; 7, exact locality unknown. 
GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 376 PLATE 17 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 18 

FIGUREs 1-3,5-9, 13. Pterocarya glabra Brown, n. sp. (p. 57). Loc. 8921. 

4. Carya antiquorum Newberry (p. 55). Loc. 5711. 

10. Phyllites pagosensis Knowlton (p. 88). Loc. 317. 

11, 12. Pterocarya hispida Brown, n. sp. (p. 57). Loc. 8920. 

14. Credneria? daturaefolia Ward (p. 66). Loc. 8920. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 18 

8 9 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 19 

FIGURE 1. Pterocarya hispida Brown, n. sp. (p. 57). Loc. 8910. 

2. Quercus macneili Brown, n. sp. (p. 58). Loc. 8239. 

3, 6, 12. Quercus greenlandica Heer (p. 58). 3, loc. 4010; 6, loc. 4878; 12, loc. 6171. 

4. Quercus yulensis Brown, n. sp. (p. 59). Loc. 8239. 

5, 7-11. Juglandicarya spp. (p. 56). 5, 10, loc. 5917; 7, loc. 8910; 8, loc. 9198; 9, 11, 

loc. 9112. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 19 

7 8 9 10 11 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 20 

FIGURES 1-5,7-9. Betula stevensoni Lesquereux (p. 57). 1, 3 (catkin),loc. 4661; 2, 4,loc. 

8893; 5, loc. 3661; 7 (catkins), loc. 9109; 8, loc. 4984; 9, loc. 9199. 

6. Celtis peracuminata Brown, n. sp. (p. 60). Loc. 7688. 

10. Morus montanensis Brown, n. sp. (p. 64). Loc. 4032. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 20 

10 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 21 

FIGURES 1-11. Corylus insignis Heer (p. 57). 1, 6 (X 3) (nut), 11, loc. 2416; 2, 3, 8, loc. 

9072; 4, loc. 8255; 5, loc. 2414; 7, loc. 9202; 9 (X 2) (catkin), 10, loc. 8519. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 21 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 22 

FIGURE 1. Quercus asymmetrica Trelease (p. 58). 

2. Zelkova planeroides ( 'Vard) Brown, 

unknown. 

Exact locality unknown. 

n. comb. (p. 60). Exact locality 

3, 4, 6, 7-10. Castanea intermedia Lesquereux (p. 58). 3, 4, 6, loc. 8928; 7, loc. 541; 8, 

loc. 9203; 9, loc. 5836; 10, loc. 5495. 

5. Quercus greenlandica Heer (p. 58). Loc. 6000. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 22 

4 

5 

MEGASCOPIC PALEOCENE PLANTS 

59312 1 0 - 62 - 8 
PLATE 23 

FIGURES 1-7. Quercus sullyi Newberry (p. 59). 1, loc. 436; 2, exact locality unknown; 

3, loc. 8552; 4, Joe. 8239; 5, loc. 2414; 6, 7, loc. 8886. 
GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 376 PLATE 23 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 24 

FIGURES 1-11, 13, 15, 16. Planera microphylla Newberry (p. 60). 1, 2, loc. 8885; 

3-5, 8, 9, 13, loc. 4264; 6, exact locality unknown; 7, 11, 

loc. 8206; 10, loc 5885; 15, loc~ 8166; 16, exact locality 

unknown. 

12, 14, 19, 20. Zelkova planeroides (Ward) Brown, n. comb. (p. 60). 12, loc. 

4035; 14, 20, loc. 2414; 19, exact locality unknown. 

17, 18, 2i-23. Ulmus rhamnifolia 'Y!tr_d (p. 60). Seeds (17, 18). 17, 

loc. 9207; 18, 23, loc. 8910; 21, loc. 2414; 22, loc. 9109. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 24 

9 

20 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 25 

FIGURES 1-6. Ficus subtruncata Lesquereux (p. 63). 1, 4, loc. 8227; 2, loc. 8551; 3, loc. 

8516; 5, loc. 8901; 6, loc. 317. 

7. Artocarpus lessigiana (Lesquereux) Knowlton (p. 60). Loc. 317. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 25 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 26 

FIGURES 1-8. Ficus planicostata Lesquereux (p. 62). 1, loc. 317; 2, 5, 7, 8, loc. 8774; 3, 

loc. 7 481; 4, loc. 8188; 6, loc. 4876. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 26 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 27 

FIGURES 1, 2, 5. Ficus affinis (Lesquereux) Brown, n. comb. (p. 61). 1, 2, loc. 317; 5, 

loc. 7481. 

3, 4. Celtis newberryi Knowlton and Cockerell (p. 59). 3, exact locality unknown; 4, loc. 9109. 

6, 7. Laurus socialis Lesquereux (p. 67). 6, loc. 4696; 7, loc. 4325. 

8. Cissites rocklandensis Brown, n. sp. (p. 81). Loc. 5679. 

9. Quercus sullY,i Newberry (p. 59). Loc. 8253. 

10. Vitis olriki Heer (p. 82). Loc. 8517. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 27 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 28 

FIGURES 1-7. Ficus artocarpoides Lesquereux (p. 61). 1, exact locality tmkno,yn; 2, 

loc. 8910; 3, loc. 4369; 4, loc. 8261; 5, loc. 2416; 6, loc. 4625; 7, exact locality 

unknown. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 28 

7 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 29 

FIGURES 1, 3-6. Platanus nobilis Newberry (p: 64). 1, 4, loc. 2416; 3, loc. 2414; 5, 6, loc. 

5594. 

2. Leaf of the existing Platanus occidentalis Linnaeus, showing basilar lobes 

(p. 64). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 29 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 30 

FIGURES 1-4. Platanus raynoldsi Newberry (p. 64). 1, exact locality unknown; 2, 3, 

loc. 2420; 4, loc. 8257. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 30 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 31 

FIGURES 1-6. Platanus raynoldsi Newberry (p. 64). 1, 5, 6, Joe. 4035; 2, loc. 4910; 3, 4, 

loc.· 2424. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 31 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 32 

FIGURES 1-5. Credneria? daturaefolia Ward (p. 66). 1, 3, loc. 2416; 2, loc. 7552; 4, loc. 

4975; 5, loc. 8164. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 32 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 33 

FIGURES 1-3, 8. Variant leaves of the existing Sassafras albidum (Nuttall) Nees (p. 68). 

4-7. Sassafras thermale (Lesquereux) Brown, n. comb. (p. 68): 4, loc. 4665; 

· 5, loc. 487 4; 6, loc. 3852; 7, loc. 8551. 
PROFESSIONAL PAPER 375 PLATE 33 

GEOLOGICAL SURVEY 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 34 

FIGURES 1, 2, 4. Sassafras thermale (Lesquereux) Brown, n. comb. (p. 68). 1, loc. 8887: 

2, 4, loc. 3653. 

3, 5. Laurophyllum perseanum Brown, n. sp. (p. 67). 3, loc. 3661; 5, loc. 

2416. 

6-8. Persea brossiana Lesquereux (p. 67). 6, loc. 7496; 7, loc. 4661; 8, loc. 

5612. 
GEOLOGICAL SURVEY 

PROFESSiONAL PAPER 375 PLATE 34 

MEGASCOPIC PALEOCENE PLANTS 

59:H:tl 0- 62 - 9 
PLATE 35 

FIGURE 1. Paleonelumbo macroloba Knowlton (p. 69). Loc. 8673. 

2-4. N elumbium montanum Brown (p. 69). 2, loc. 8786; 3 (fruit), loc. 8262; 4, 

loc. 8227. 

5, 6. Cabomba inermis (Newberry) Hollick (p. 68). 5, loc. 9072; 6, exact locality 

unknown. 

7. Paleonuphar hesperi1tm Brown, n. sp. (p. 69). Loc. 7538. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 35 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 36 

FIGURES 1-5. Paranymphaea crassifolia (Newberry) Berry (p. 70). 1, loc. 2432; 2, loc. 

8553; 3, 4, loc. 8519; 5, exact locality unknown. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 36 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 37 

FIGURES 1-24. Cercidiphyllum arcticum (Heer) Brown (p. 70). Leaves (8-11, 13, 15, 17, 

18, 20, 22); fruits (1, 14, 23, 24); seeds (2-7, 12, 16, 19). 1, 2, loc. 3653; 

3, loc. 8673; 4, 7, 14, loc. 5437; 5, loc. 8227; 6, loc. 8672; 8, 10, 11, 17, loc. 

9334; 9, loc. 8188; 12, 16, loc. 9130; 13, 18, loc. 8567; 15, loc. 8519; 19, 20, 

22-24, loc. 8517; 21, loc. 317. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 37 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 38 

FIGURES 1-17. Cercidiphyllum arcticum (Heer) Brown (p. 70). Leaves (1, 2, 7-14, 16, 17); 

fruits (5, 6, 15); seeds (3, 4). 1, loc. 2416; 2, loc. 8234; 3 (left column in 

3, loc. 8910), others, loc. 9125; 4, 9, 15, loc. 9125; 5, 6, loc. 4256; 7, 8, 10, 14, 

16, loc. 4264; 11, loc. 8167; 12, loc. 8556; 13, loc. 8540; 17, loc. 8774. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 38 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 39 

FIGURE 1. ~Magnolia berryi (Knowlton) Brown, n. comb. (p. 71). Loc. 5679. 

2. Magnolia regalis Heer (p. 72). Loc. 317. 

3. Magnolia borealis Brown, n. name (p. 71). Loc. 8523. 

4. Magnolia magnifolia Knowlton (p. 71).. Loc. 4882. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 39 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 40 

FIGURES 1, 2. Quercus greenlandica Heer (p. 58). Loc. 5526. 

3. Kalmia elliptica Brown, n. sp. (p. 85). Loe. 4661. 

4-6. Hamamelites inaequalis (Newberry) Brown, n. comb. (p. 72). 4, exact locality unknown; 5, loc. 2416; 6, loc. 4977. 

l ~ ,,1 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 40 

2 

6 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 41 

FIGURES 1, 2, 4. Pterospermites cordatus Ward (p. 82). 1, loc. 8678; 2, loc. 7776; 4, loc. 

4625. 

3, 5. Hydrangea antica Brown, n. sp. (p. 73). 3, loc. 337; 5, loc. 4661. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 41 

5 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 42 

FIGuREs 1, 5-7. Rhamnus cleburni Lesquereux (p. 77). 1, 6, loc. 8672; 5, 7, loc. 317. 

2, 8. Prunus careyhurstia Brown, n. sp. (p. 73). 2, loc. 8666; 8, loc. 8552. 

3, 4, 9. Prunus coloradensis Knowlton (p. 73). 3, 4, loc. 5738; 9, loc. 5836. 

, ''I, 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 42 

9 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 43 

FIGURE 1. Lindera obtusata (Ward) Brown, n. comb. (p. 67). Loc. 5720. 

2, 3. Prunus perita Brown, n. sp. (p. 7 4). Loc. 8552. 

4-6. Robinia wardi (Knowlton) Brown, n. comb. (p. 75). 4, 6, loc. 9109; 5, loc. 

9445. 

7, 8. Ficus affinis (Lesquereux) Brown, n. comb. (p. 61). 7, loc. 5796; 8, loc. 4625. 

9. Probably a leaflet of Robinia wardi (Knowlton) Brown (p. 7 5). Loc. 6440. 

10. Bauhinia wyomingana Brown (p. 74). Loc. 4877. 

11. Probably a leguminous leaflet (p. 75). Loc. 8556. 

12, 13. Mimosites coloradensis Knowlton (p. 75).- Loc. 6440. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE43 

8 9 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 44 

FIGURES 1-6. Eucommia berrata (Newberry) Brown, n. comb. (p. 72). 1, loc. 2421; 2, loc. 

5889 ;' 3, loc. 2420; 4, loc. 2423; 5, loc. 4626; 6, loc. 2422. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 44 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 45 

FIGURES 1-7. Eucommia serrata (New'berry) Brown, n. comb. (p. 72). 1, exact locality 

unknown; 2, 5-7, loc. 2420; 3, loc. 8552; 4, loc. 5889. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 45 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 46 

FIGURE 1. Viburmtm cupanioides (Newberry) Brown, n. comb. (p. 87). Loc. 8190. 

2, 3, 6, 8. Acer newberryi Brown, n. name (p. 7 5). 2;- 6, 8, loc. 2420; 3, loc. 8910. 

4. Asimina vesperalis Brown, n. sp. (p. 73). Loc. 877 4. 

5. Acer sp. (p. 76). Loc. 8910. 

7. Acer silberlingi Brown, n. sp. (p. 76). Loc. 8567. 

9, 10. Acer sp. (p. 76). Loc. 9109. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE ~6 

MEGASCOPIC PALEOCENE PLANTS 

593 121 0 62 - 10 
PLATE 47 

FIGURES 1-8. Sapindus affinis Newberry (p. 76). 1, 3, exact locality unknown; 4, 6, loc. 

2416; 2, 5, loc. 8523; 7, 8, loc. 8678. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 47 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 48 

FIGURES 1, 3, 4. Parthenocissus ursina Brown, n. sp. (p. 81). Loe. 8547. 

2, 5-7. Rhamnus goldiana Lesquereux (p. 77). 2, 6, 7, Joe. 8774; 5, Joe. 317. 

8. Ficus minutidens Knowlton (p. 62). Loe. 7 496. 
PROFESSIONAL PAPER 375 PLATE48 

GEOLOGICAL SURVEY 

4 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 49 

FIGURES 1-10. Rhamnus hirsuta Brown, n. sp. (p. 77). Loc. 8519. 
PROFESSIONAL PAPER 375 PLATE 49 

GEOLOGICAL SURVEY 

3 

7 8 9 

6 

MEGASCOPlC PALEOCENE PLANTS 
PLATE 50 

FIGURES 1-4, 7-9. Afyrtophyllum torreyi (Lesquereux) Dorf (p. 83). 1, loc. 9210; 2, 7, loc. 

8551; 3, 4, loc. 8677; 8, loc. 8188; 9, loc. 8881. 

5, 6, 10, 11. Zizyphus fibrillosus (Lesquereux) Lesquereux (p. 78). 5, 6, loc. 8774; 

10, loc. 8776; 11, loc. 317. 
GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 375 PLATE 50 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 51 

FIGURES 1-18. Ampelopsis acerifolia (Newberry) Bro,vn, n. comb. (p. 78). 1, 16, loc. 

8523; 2, 13, loc. 8521; 3, 7, 12, loc. 4974; 4, 11, loc. 8774; 5, 6, 9, 14, loc. 

8922; 8, 15, 18, loc. 2420; 10, loc. 4699; 17, loc. 9130. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 61 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 52 

FIGURES 1-8, 10. Ampelopsis acerifolia (Ne'wberry) Brown, n. comb: (p. 78). 1, loc. 8227; 

2, 3, 6, 10, loc. 8517; 4, loc. 2420; 5, 8, loc. 317; 7, loc. 4699. 

9. Cercidiphyllum arcticum (Heer) Brown (p. 70). Loc. 3981. 

•1 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 52 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 53 

FIGURES 1-4, 6. Cissus marginata (Lesquereux) Brown, n. comb. (p. 79). 1, loc. 317; 2, 3, 

loc. 6431; 4, loc. 9252; 6, loc. 6416. 

5. Seed of Vitis sp. (p. 82). Loc. 9132. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 53 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 54 

FIGURES 1-4. Cissus marginata (Lesquereux) Brown, n. comb. (p. 79). 

loc. 6416; 3, loc. 317. 

1, 4, loc. 7 495; 2, 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 54 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 55 

FIGURES 1-3. Betulites sp. Lesquereux, (pp. 80, 81), from the Dakota sandstone (Upper 

Cretaceous), Ellsworth County, Kans., for comparison with Cissus marginata. 

4, 6, 7. Cissus marginata (Lesquereux) Brown, n. comb. (p. 79). 4, from the 

Mesaverde formation (Upper Cretaceous), south of Gallup, N. ·Mex.; 6, 

from the Hell Creek formation (Upper Cretaceous), east of Glendive, Mont.; 

7 (X ?~),from the Kaiparowitz formation (Upper Cretace·ous) at Parowan 

Gap, Utah. The petiole is perfoliate. 

5. Twig of Cinnamomum sp. (p. 66) with opposite leaves, from the Mesaverde 

formation (Upper Cretaceous) at Black Mesa, south of Kayenta, Ariz. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 55 

4 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 56 

FIGURES 1, 2, 5, 6. 1\felastomites montanensis Brown, n. sp. (p. 84). 1, 2, loc. 8519; 5, 

loc. 7662; 6, loc. 7005. 

3, 7, 10. Vitis lobata (Knowlton) Brown, n. comb. (p.82). Loc. 8519. 

4, 8, 9. Juglans taurina Brown, n. sp. (p. 56). 4, 9, loc. 2423; 8, loc. 4262. 
PROFESSlON AL PAPER 375 PLATE 56 

GEOLOGICAL SURVEY 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 57 

FIGURE 1. Viburnum cupanioides (Newberry) Brown, n. comb. (p. 87). Loc. 5885. 

2, 5. Dillenites garfieldensis Brown, n. sp. (p. 82). Loc. 8774. 

3. Leaf with minute teeth (p. 90). Loc. 877 4. 

4. Probably a small leaflet of Fraxinus eocenica Lesquereux (p. 85). Loc. 8774. 

6, 7. Quercus sullyi Newberry (p. 59). Loc. 8239. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 57 

I I 

I 

J I 

I 

I 

I 

/ • I I 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 58 

FIGURES 1-12. Trapa angulata (Newberry) Brown, n. comb. (p. 83). 1, 2, with purported 

fruits at centers of the rosettes, Ravenscrag formation, Saskatchewan, 

Canada; 3-5, rosette and separate leaves, 7-12, fruits, loc. 2420; 6, rosette 

showing fimbriated submerged foliage, from the Meeteetse formation 

(Upper Cretaceous) in SE% sec. 10, T. 46 N., R. 98 W., Wyoming. 

13. Trapa paulula (Bell) Brown, n. comb. (p. 84). Loc. 7004. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 68 

7 8 9 10 11 

MEGASCOPIC PALEOCENE PLANTS 

593121 0 - 62 - ll 
PLATE 59 

FIGURE 1. Corn us hyperborea Heer (p. 84). I ... oc. 4871. 

2-5, 7, 12. Cornus nebrascensis Schimper (p. 85). 2, 3, 7, loc. 8567; 4, loc. 8195; 5, 

loc .. 4626; 12, loc. 5389. 

6, 11. Ampelopsis acerifolia (Newberry) Brown, n. comb. (p. 78). 6, loc. 8552; 11, 

loc. 2420. 

8, 9. Koelreuteria annosa Brown (p. 76). 8, loc. 9253; 9, loc. 9344. 

10. Vitis olriki Heer ·(p. 82). Loc. 9104 . 

. ~. 

'1 
GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 376 PLATE 59 

9 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 60 

FIGURES 1-4, 6. Nyssa alata (Ward) Brown, n. comb. (p. 85). 1, loc. 8542; 2, 4, 6, loc. 

2416; 3, loc. 8224. 

5, 7-10. Phyllites disturbans Brown, n. sp. (p. 88). 5, loc. 7659; 7, loc. 4262; 

· 8, loc. 6342; 9, 10, loc. 8563. 
GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 375 PLATE 60 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 61 

FIGURES 1-3. Salix aquilina Brown, n. sp. (p. 55). Loc. 9322. 

4. Nyssa borealis Brown, n. sp. (p. 85). Loc. 4981. 

5-8. Phyllites demoresi Brown, n. sp. (p. 88). 5, 6, 8, loc. 4264; 7, loc. 8196. 
GEOLOGLCAL SURVEY PROFESSIONAL PAPER 375 PLATE 61 

8 

7 

6 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 62 

FIGURES 1-7. Fraxinus eocenica Lesquereux (p. 85). 1, loc. 8234; 2, loc. 317; 3-5, loc. 

8774; 6, loc. 6105; 7, loc. 4626. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 62 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 63 

FIGURES 1-8. Viburnum antiquum (Newberry) Hollick (p. 86). 1-7, loc. 2414; 8, loc. 

8166. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 63 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 64 

FIGURES 1-3, 5, 7-11. Viburnum asperum Newberry (p. 87). 1, 3, loc. 2417; 2, loc. 8885; 

5, loc. 8520; 7, 8, loc. 8678; 9, 10, loc. 9056; 11, loc. 607. 

4. Viburnum antiquum (Newberry) Hollick (p. 86). Loc. 9071. 

6. A probable ancestor of Viburnum asperum (p. 87) from the Upper 

Cretaceous, 4 miles east of Reed Point, Mont. 

.• __ 0J,.Jti 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 64 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 65 

FIGURES 1-8. Viburnum cupanioides (Newberry) Brown, n. comb. (p. 87). 1, exact 

locality u·nknown; 2, loc. 6050; 3, 4, loc. 2420; 5, 7, loc. 5889; 6, loc. 4625; 

8, exact locality unknown. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 65 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 66 

FIGURES 1-3, 6, 9. Cinnamomum sezannense Watelet (p. 66). 1, loc. 8426; 2, loc. 4571; 

3, loc. 8777; 6, 9, loc. 4725. 

4. Palmately veined leaf, (p. 90). Loc. 8920. 

5. Serrate leaf, (X 2) (p. 90). Loc. 6215. 

7. Ampelopsis acerifolia (Newberry) Brown, n. comb. (p. 78). Loc. 317. 

8. Small leaf of Platanus raynoldsi Newberry (p. 64). Loc. 9180. 

10. Lindera obtusata (Ward) Brown, n. comb. (p. 67). Loc. 4323. 

11. Magnolia borealis Brown, n. name (p. 71). Loc. 5579. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 66 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 67 

FIGURES 1-4, 8. Prunus corrugis Brown, n. sp. (p. 74). 1, loc. 9492; 2, 3, loc. 

4984; 4, loc. 4050; 8, loc. 541. 

5. A five-loculed fruit (X 2) (p. 91). Loc. 8247. 

6, 7. Maplelike samaras, but probably of sapindaceous affinity (p. 91). 

6, loc. 9201; 7, loc. 9125. 

9, 14'=-16, 20, 22, 23, 27. Seeds with papillose inner surface (p. 91). 9, 10 (X 2), loc. 

8850; 15,22 (X 2) withsmallsnailatleft, 23 (X 3) sameasfig.15, 

27 (X 2), loc. 6215; 16, 20 (X 2), loc. 9236. 

10. Flattened, carbonized fruit with scars (p. 91). Loc. 6382. 

11, 12, 17. Probably fruits with remnants of calyces (p. 91). 11, loc. 4626; 

12, loc. 8566; 17, loc. 8547. 

13, 45. Nordenskioldia borealis Heer (p. 89). 13 (X 3), loc. 8913; 45, 

loc. 8887. 

18, 19, 24, 25, 30, 31. Viburnum tilioides Ward (p. 87). 18, 24, loc. 2416; 19, loc. 2414; 

25, loc. 8554; 30, loc. 8774; 31, loc. 8255. 

21. Spike or catkin of seeds (p. 91). Loc. 8519. 

26. Impression of a seed (X 2) showing pits arranged in longitudinal 

rows (p. 91). Loc. 4618. 

28, 29. Seeds with ridges and scattered glands (p. 91). Loc. 4264. 

32. Heart-shaped fruits or seeds (p. 91). Loc. 8519. 

33, 34. Viburnum solitarium Lesquereux (X 5) (p. 90). Loc. 317. 

35. Elliptic fruit with a fringe of hairs or filaments (p. 91). Loc. 

8522. 

36. Oval fruit with severallocules (X 2) (p. 91). Loc. 8167. 

37. Palmocarpon subcylindricum Lesquereux (p. 90). Loc. 317. 

38. Oval seeds like figs. 33 and 34 but rounded at the top, not squarish 

(p. 91). Loc. 5667. 

39-42, 47. Burlike objects (p. 91). 39-42, loc. 7538; 47, loc. 9109. 

43. Probably the fertile cone of an Equisetum (p. 46, 91). Loc. 6905. 

44. Longitudinal hollow studded with bilobed projections (p. 91). 

Loc. 9130. 

46. Calyx of a flower (X 10) (p. 91). Loc. 8774. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE 67 

44 

43 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 68 

FIGURES 1, 3-6. Leguminosites coloradensis Knowlton (p. 74). 1, 6, loc. 4877; 3, loc. 

5526; 4, loc. 8551; 5, loc. 336. 

2, 7-11. Roots with rootlets (p. 91). 2, loc. 8669; 7, loc. 4468; 8, loc. 750; 9, 

loc. 9533; 10, loc. 8668; 11, loc. 6765. 

12, 13. Pterocarya hispida Brown, n. sp. (p. 57). 12, loc. 8910; 13, loc. 6839. 

14-16. Sagittaria megasperma Brown, n. sp. (p. 52). 14, loc. 4268; 15, 16, 

loc. 9129. 

17-22. Two-winged seeds (p. 92). Loc. 8910. 

23, 24. Liquidambar dakotense Brown, n. sp. (p. 72). Loc. 6384. 

25.- Probably a stone fruit (X 2) (p. 92). Loc. 8913. 

26. Probably the glume of a grass (X 2) (p. 92). Loc. 8913. 

27-29. Pedicelled flowers or fruits (X 2) (p. 92). 27, loc. 9109; 28, loc. 9125; 

29, loc. 5595. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 376 PLATE 68 

10 

9 

8 

17 

23 24 25 26 

12 18 

~ 28 

19 '4P 

27 

11 

20 

13 

21 

t 16 22 29 

MEGASCOPIC PALEOCENE PLANTS 
PLATE 69 

FIGURES 1. Feather of an unknown bird (p. 92, 96). From a locality of clinkered 

shale about 10 miles north of Glendive, Mont. 

2. Dragonfly (p. 96). Loc. 8190. 

3. Wing of a cockroach (p. 96). Loc. 8556. 

4. Wing of a caddisfly (X 2) (p. 96). Loc. 4976. 

5, 6. Views of a Champsosaurus vertebra (p. · 96). From a locality soutp of Watford 

City, N. Dak. 

7, 8. Gar scales (p. 96). Loc. 9342. 

9, 10. Tooth at left in fig. 9, Aphronorus, from Donnybrook, N. Dak. Third tooth 

from left, Tricentes, and all the remaining teeth in figs. 9 and 10, Claenodon, 

from 8 miles west of Medicine Lake, Mont. (p. 96). 

11, 12, 13. Reptilian remains (p. 96). Teeth, 11, 12; claw, 13. Loc. 9342. 

14. Scale of Amia (p. 96). Loc. 8519. 

15-19. Probably fish or reptile coprolites (p. 92). 15, from the Miocene on Salmon 

Creek, Wash.; 16-19, from the Paloecene at loc. 9564, 8 miles south of Rhame, 

N.Dak. ,.__,;.: .. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 375 PLATE ti9 

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7 

' 

8 

15 

14 

MEGASCOPIC PALEOCENE FOSSILS 

U. S. GOVERNMENT PRINTING OFFICE o 1962 0- 593121 
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