-LIBRARlI 

LIBRARY COpy ~Ko.PFRTY OF: 

aJ]. §o ~[jJJ~EAU Of Al1iNES 

UNITED STATES DEPARTMENT OF THE INTER1§fA VI M~NERAL RESOURCE OFF"CE 

Ray Lyman Wilbur, Secretary 

GEOLOGICAL SURVEY 

George Otis Smith, Director 

Professional Paper 155 

LIBRAR~ 

B'IJRfAtf OF MINiS ~18RARV 

SPUI\ANh. ~AStt. 

JUN - 21971 .o~ 

1, "on:".:liiJ 

THE FLORA OF THE DENVER AND AS~OCIATED FORMATI(9NF4Sf ~fTuRH o 1'0 LIBRAI. 

OF COLORADO 

BY 

FRANI{ HALL KNOWLTON 

A POSTHUMOUS WORK 

EDITED BY 

EDWARD W~LBER BERRY 

UNITED STATES 

GOVERNMENT PRINTING OFFICE 

WASHINGTON: 1930 

For sale by the Superintendent of Documents, Washington, D. C. Price (paper cover) 80 cents 
CONTENTS 

Page Editor's preface _____________ . __________ ._ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ III 

Introduction_____________________________________________________________________________________ 1 

Historical review ________________________________ . _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 1 

Area around Denver__________________________________________________________________________ 1 

Dawson arkose _____________________________ - ___________________________________ ._ _ _ _ _ _ _ _ _ _ _ _ _ _ 3 

Middle Park formatioD _______ - ___ - ____ - - ________ --- __________________ -__ - _ __ _ __ ____ __ _ _ __ __ __ _ 5 

Plants in the equivalent of the Arapahoe formation _____________ - _____ :... _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 6 

Th~ flora of the Denver and related formations _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7 

List of species _________ - _____ ._ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 7 

Distribu tion of the flora _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 9 

Ecologic considerations _________________ ,.. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 11 

Physical conditions _____ - _______________ - ___ - - ____ :.. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 13 

Materials studied ______________________ - ___________ - ______________ - _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 13 

Synonyms and changes of name _________________________ . _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 14 

Systematic paleobotany ___________ -. _________ - ____ -. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 15 Index~ ____________________________________________________ ~ _____________________ . _______________ 139 

ILLUSTRATIONS 

Page 

PLATES 1-59. Denver flora ______________________________________________________________ . _ _ _ _ _ _ _ _ _ 137 

L ! i II 
EDITOR'S PREFACE 

The present work was one upon which Doctor I(nowlton had 

labored at intervals over' a long period of years and which was prac-

. tically completed at the time of his death. The editor's task has 

been that of comparing the figures with the types and supplying a 

few gaps in the text. At no point has he deemed it desirable to give 

expression to opinions differing from those of the author, and the 

work as it stands is entirely Doctor Knowlton's. The few comments 

that have seemed advisable have been placed in footnotes signed by 

the editor's initials. 

E. 'V. B. 

III 
THE FLORA OF THE DENVER AND ASSOCIATED FORMATIONS 

OF COLORADO 

By FHANK HALL KNOWLTON 

INTRODUCTION 

In the introduction to my report on the Laramie 

florn of the Denver Basin 1 I stated that that was the 

first of whnt was proposed to be a series of papers 

dealing with the stratigraphy nnd paleobotany of certain Upper Cretaceous and lower Tertiary plant-bearinO' fOl'mntions of the Rocky Mountain region. The 

pl~sent paper is the second of this ~eries and de~ls ' 

with the flora of the Denver formatIOn and certaIn 

identical or closely related formations that occur 

mainly in or adjacent to the Denver Basin of Colorndo. The additional formational units treated are 

the Arapahoe formation, the Dawson arkose, and the 

~fiddle Park formation. 

HISTORICAL REVIEW 

As the history of the recognition and subsequent 

naming of the Arapahoe and Denver formations is 

more or less closely interwoven with that of the Laramie formation, it is necessary to give a brief review of 

certain portions of the Laramie history. This review 

is in the main an adaptation from a historical review 

of opinion concerning the Laramie formation, which 

forms the first part of the paper cited above. 

AREA AROUND DENVER 

The area about the city of Denver, Colo., now well 

known as the Denver Basin, is in a \vay classic ground 

for geology. It has been visited and studied in greater 

or less detail by all the geologists and paleontologists 

,vho have been interested in Rocky 1\10untain geology. 

Included among the pioneers in this study are John 

L. Le Conte, F. V. Hayden, A. C. Peale, A. R. Marvine, 

Leo Lesquereux, C. A. White, L. F. "Yard, S. F. 

Emmons, G. H. Eldridge, and Whitman Cross. The 

coal-bearing rocks lying conformably above the marine 

Fox Hills sandstone were at first believed to be a 

southern extension of the "Lignitic group" of 

the upper Missouri River region and were held by 

Hayden, Lesquereux, White, and others to be of 

Tertiary age. When the Laramie was established by 

IGn 0' as the uppermost member of the conformable 

Cret~ceous series above the Fox Hills this portion of 

1 U. S. Gco1. Survey Prof. Papcr 130, p. 3, 1922. 

the section in the Denver Basin and adjacent areas 

along the Front Range fell within its limits and ~as 

regarded by him and by those who accepted his vie~s 

a~ Cretaceous. This so-called Laramie was for many 

years considered to be a unit, notwithstanding the fact 

that at certain points along the base of the mountains, 

notably at Golden, there is evidence of discontinuity in 

deposition and difference in lithologic composition. 

Thus in the lower or coal-bearing portion of the section some of the beds are nearly vertical, whereas certain other beds, as those in Table Mountain, are nearly 

horizontal. "Yard invoked the' presence of a hypothetical fault to account for this apparent discordance. 

Both lower and upper beds had furnished extensive 

collections of plants, which had been studied mainly 

by Lesquereux, but, as it later appeared, no distinction 

was made in the records concerning them, all being 

called simply Laramie, with the result that when the 

several formations were subsequently proved to be 

distinct the greatest confusion and difficulty resulted 

in attempting to separate these floras. 

As early as the summer of 1881, when Cross 2 began 

I1is studies in the Denver Basin, he "first observed," 

to use his own language, "that the Table Mountain 

strata possessed characteristics pr<?ving them to belong 

to a series distinct from the normal Lar,amie." No 

published announcement of this discovery was made 

at the time, and for a number of succeeding years 

field work was continued in the region, during which 

it was ascertained by George H. Eldridge, who was 

associated with Cross, that there was apparently another distinct ,Tertiary formation between that discovered by Cross and the normal Laramie. A preliminary statement of the nlost significant results of 

this investigation was made by Eldridge and Cross in 

two papers read before the Colorado Scientific Society 

July 2, 1888, and published in its proceedings. In the 

first of these papers Eldridge S named and described 

the Arapahoe formation,4 which, he~ stated, was" the 

2 Cross, Whilman, The Denver Tertiary formation: Am. Jour. ScI., 

3d ser., vol. 37, p. 262, 1889. 

8 Eldridge, G. H., On some stratigraphical and structural features of 

the country about Denver, Colo.: Colorado Sci. Soc. Proc., vol. 3, pp. 

86-118, 1888. 

'This was first named the " Willow Creek beds," but on the ground 

of preoccupation the name was changed in a footnote (p. 97) to 

Arapahoe. 
2 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

formation next succeeding the Laramie in geological 

order, and unconformably resting! on it." It was 

characterized as follows: 

It is composed of a basal member of conglomerate or gritty 

sandstone, according to its distance from the foothills with 

an overlying zone of gray argillaceous or arenaceous ~hales, with an occasional ironstone; when confined between underlying and overlying groups it has a thickness of between 600 

and 1,200 feet. 

In the succeeding article Cross 5 named and described the" D.enver Tertiary formation," which he 

believed to be unconformably overlying the Arapahoe. 

Lithologically it was found to be composed almost entirely of andesitic volcanic' material; its thickness was 

given as 800 to 1.200 feet. 

As already indicated, both the Arapahoe and Denver 

formations were regarded originally as of Tertiary 

age, and both wel~e found to contain fairly abundant 

v~rtebrate remains belonging to turtles, crocodiles, and 

dInosaurs. These remains were studied by O. C. 

Marsh, who pronounced the dinosaurs from'the Denver 

to be "typical Jurassic dinosaurs of both herbivorous 

and carnivorous types." G Later, when the Cer~topsidae, a group of peculiar horned dinosaurs especially 

,abundant in the ~ance formation of Converse County, 

Wyo., was descrIbed, these dinosaurs were found to 

belong to it. The presence of these dinosaurs was considered by the vertebrate paleontologists as' fixing the 

age of the beds containing them as Cretaceous. The 

invertebrates consisted of only a few fresh-water types 

and prove'd of very little value in indicating the age. 

Fossil plants were abundant, especially in the Denver beds; but, as already stated, the large collections 

previously made we~e without specific locality labels, 

and all were then considered to be of Laramie aO'e 

Before they could 'be used in the light of this ne;eI: 

information regarding their stratigraphic relations 

it was necessary to separate them if possible, and it 

was found that this could be done almost invariably 

on the basis of the matrix in which they were preserved. These earlier collections were fortunately deposited in the United States N ationa! Museum, and 

the study of the matrix undertaken by Cross resulted 

in showing that they were distinct from those of the 

underlying Laramie-in fact, out of about 98 species 

in the Laramie and 140 in the Arapahoe and Denver 

only about 15 nominal species were found in common. 

It may be added that subsequent studies of the Arapahoe and Denver plants have shown them to be essentially Tertiary in type. 

15 Cross, Whitman, The Denver Tertiary formation: Colorado Sci. Soc. 

Pl'OC" vol. 3, pp. 110-133, 1888; recast and republished uuder the same 

title in Am. ,Tour. Sci., 3d ser., vol. 37, pp. 261-282, 1889. 

G See Cannon, G. L., On the Tertiary Dinosauria found in Denver 

beds: Colorado Sci. Soc. Proc., vol. 3, p. 143, 1888. 

In 1892 Cross 7 briefly reviewed the relations of the 

Arapahoe and Denver formations of the Denver Basin 

and also enumerated additional Colorado localities in 

the Huerfano Basin, Gunnison County, the Yampa 

River region, the Animas ,River region, Middle Park, 

and elsewhere, in which identical or similar stratigraphic relations had been established. He discussed 

at length the conflicting paleontologic evidence of the' 

age of these deposits, and concluded as follows: 

The writer wishes to advocate the restriction of the term 

"Laramie," in accordance with its original definition, to the 

series of conformable beds succeeding the marine Montana 

Oretaceous, and the grouping of the post-Laramie lake beds 

described, with their demonstrated equivalents, in another 

~eries to which a comprehensive name shall eventually be 

given. ,~, * * The question as to whether the series shall 

be referred to the Cretaceous or to the Eocene can not be 

finally settled until the various conflicting elements of the 

evidence have been adjusted on a ,basis of further and more 

exact information. 

In 1892 Cross 8 published an amplification of the 

previous brief notice, in which it was shown that the 

large area referred to the" Lignite" or Laramie was 

made up of a thick series of mainly andesitic rocks 

resting unconformably on marine Cretaceous, there 

being no beds corresponding to the Laramie of the 

Denver Basin. These beds, called the Middle Park 

beds, were correlated with the Denver beds on account 

of their position, their lithologic similarity, and the 

identity of the contained floras. 

The publication of the monograph on the Denver 

Basin, by Emmons, Cross, and Eldridge,O was delayed until 1896, a delay which in many ways was 

perhaps of advantage, for it permitted the'incorporation and disqussion of many pertinent facts that had 

been made available in the meantime reO'ardinO' the /:) b 

stratigraphic and other relations of, the formations 

under discussion. The Laramie, Arapahoe, and Denver formations of the Denver Basin were , , of course 

described in detail, and the relations between the 

Laramie and overlying rocks especially were made 

plain. The lnagnitude of this time interval was insisted upon as of major importance in Rocky Mountain stratigraphy. The Arapahoe and Denver formations were referred to the Cretaceous out of deference 

to the views of Marsh, at that time vertebrate paleontologist of the Geological Survey. The lithologic and 

stratigraphic relations of these formations and a very 

full discussion of the several lines of paleontologic evid.ence were given by Cross, who prepared this portion 

of the volume. 

7 Cross, Whitman, Post-Laramie deposits of Colorado: Am. Jour. 

ScL, 3d ser., vol. 44, pp. 19-42, 1892. , 

8 Cross, Whitman, The post· Laramie beds of Middle Park, Colo.: 

Colorado Sci. Soc. Proc., vol. 4, PP. 1-27, 1892. 

9 Emmons, S. F., Cross, Whitman, and Eldridge, G. H., Geology of 

the Denver Basin in Colorado: U. S. Geol. Survey Mon. 27, 1896. ' 
HISTORICAL REVIEW 3 

For 10 years after the publication of the Denver 

Busin monograph little active work was prosecuted in 

t.his region, though investigations in adjacent or more 

remote areas were found to have a more or less direct 

bearing on the problems here involved. 

In 1897 Veatch,10 from studies in Carbon County, 

'Vyo., w,as led to question the validity of the current 

application of the term" Laramie." He endeavored to 

prove that Carbon County was intended by Iring to 

be the type locality for the Laramie. In this vicinity 

Veatch discovered the presence in the supposedly continuou,s Laramie section of a profound unconformity, 

which he thought to be similar to that found by Cross 

irJ the Denver Basin, and as the beds studied and described at Carbon by the members of the Iring Survey 

were all above the break, he contended that the name 

should properly be applied only to those beds and not 

to the beds conformable to the Cretaceous section. 

Veatch further .held that the delimitation of the 

Arapahoe and Denver formations by Cross constituted a virtual redefinition of the Laramie. 

Veatch's pa.per was answered by Cross 11 and 

Peale,12, and it is silfficient to say that his view has not 

been a'dopted. Cross especially reviewed the facts relating to the application of the term" Laramie," holding that the separation of the Arapahoe and Denver 

djd not constitute a redefinition, and finally, following 

out a suggestion made by himself in 1892, proposed 

to call these two formations and their equivalents the 

Shoshone group. 

It is proposed to apply the term "Shoshone group" to the 

deposits which unconformably succeed the Laramie and to 

their equivalents, and which are overlain by the Fort Union 

or 'Vasatch beds, when they are present. 

In this paper Cross briefly reviewed the data bearing on the age of these post-LaraIAie beds and concluded as follows: 

In the preceding discussion I have avoided the suggestion 

as to the age of the Shoshone beds, whether Oretaceous 01' 

Eocene, I desire now to urge their reference to the Eocene. 

The Denver beds were originally referred by me to the Eocene, 

but the great weight attached to the Mesozoic affinities of the 

veL'tebrate faunas by paleontologists led to a tentative acquiescence in the assignment of the Arapahoe and Denver 

formations to the Oretaceous in the Denver monograph. 

DAWSON ARKOSE 

Dawson arkose is the name now adopted for the 

lower portion of the "Monument Creek group" of 

10 Veatch, A. C., On the orIgin and definition of the geologic term 

Lammie: Am. JOUl·. ScI., 4th ser., vol. 24, p. 18, 1907; expanded under 

the Sllme title In Jour. Geology, vol. 15, p. 526, 1907. 

U Cross, Whitman, The LaramIe formation and the Shoshone group: 

Washington Acad. ScI. Proc., vol. 11, pp. 27-45, 1900. 

u l'eule, A. C., On the application of the tenD" Laramie" : Am, JOUI', 

ScI.. 4th ser., vol. 28, pp. 45-58, 1909. 

Hayden,13 established in 1869 in an account of a reconnaissance study from. Cheyenne, Wyo., through the 

Denver Basin, Colorado Springs, Canon City, and 

Raton fields. The group was based on "a series of 

variegated. beds of sands and arenaceous clays, nearly 

horizontal, resting on the upturned edges of the older 

rocks, * * * of v,arious colors * * * and of 

v~rious degrees or texture." It occurs along the 

Front Range on the divide between the Platte and. 

Arkansas drainage basins, where it covers an area 

"of about 40 miles in width from east to west and 

50 miles in length north and south." From:' its modern: appe,ar,ance Hayden concluded that it was of 

,- either late Miocene or Pliocene age." 

In a "Report on the vertebrate paleontology of 

Colorado," published in 1873, Cope 14 took occasion to 

speak of the age of the "'Monument Creek group," concluding that it was younger than Eocene and older than 

Pliocene, hence "it may be referred to the Miocene 

until further discoveries enable us to be Inore exact." 

In the following' year (1874) Hayden 16 again reo 

ferred to the "Monument Creek group," and after 

describing 'it at some length alluded to Cope's statement regarding the vertebrate evidence, saying: "As 

to the real age of this group, I am inclined to regard it as Miocene, perhaps llpper Miocene." 

The next publication. that it is necessary to notice 

is the Denver Basin monograph by Emmons, Cross, 

and Eldridge,I6 though their discussion of these rocks 

is brief, as in the series of beds mapped these rocks 

occur only as projecting tongues forming the divide 

between the Platte and Arkansas waters. 

Emmons gave first a brief summary in which, although caliing the beds the" Monument Creek formation," he noted that "two divisions have been disti.nguished, marked by an apparent unconformity and 

period of erosion." The lower division was referred 

tentatively to the Miocene on the basis of the previous 

vertebrate work of Cope; the upper division was 

,thought probably to be referable to the Pliocene. 

Eldridge devoted several pages to the discussion 

of the stratigraphic relations, lithology, and life of 

the "Monument Creek formation," writing in part 

as follows: 

The Monument Oreek formation occurs along the southern 

edge of the Denver field in the steep slopes of a high mesa 

and also stretches from its base prairieward 1n thin sheets. 

la Hayden, F. V., Prellminary field report of the United Stntes geological survey of Colorado and New Mexico, PP. 37-46, 1869; reprint, 

pp. 137-146, 1873. 

14 Cope, E. D., U. S. Geol. and Geog. Survey '.rerr. Ann. Rept. for 

1873, p. 430, 1874. ' 

15 Hayden, F. V., U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 

18;4, pp. 36, 37, 1876. 

10 Emmons, S. F., Cross, Whitman, and Eldridge, G. H., Geology of 

tht) Denver BaSin in Colorado: U. S. Geol. Survey Mon. 27, PP. 38, 39, 

252-254, 1896. 
4 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

The floor of the lake in which the Monument Creek was deposited was more or less irregu1ar from erosion and in one 

part or another consisted of the clays and sandstones of the 

Laramie, Arapahoe, or Denver formations. In the foothill 

region the Monument Creek lies in contact with the Arapahoe; 

between Platte River and Cherry Creek a few hundred feet 

of Denver beds exist, which farther to the east disappear. 

Nortl) and east of Coal Creek, on the eastern edge of the field, 

both Denver and Arapahoe are wanting, and the Monum~nt 

Creek rests directly upon the clays of the Laramie. 

Although the relations between the Denver and 

Arapahoe and the" Monument Creek" were somewhat 

obscure, the interpretation naturally followed that the 

latter was stratigraphically above the former, because 

it was then supposed that the lower division of the 

"Monument Creek" was Miocene in age, whereas the 

Arapahoe and Denver were then referred to the 

Cretaceous. 

In 1902 Lee 11 discussed the "Monument Creek 

group" of Hayden. To the lower portion (of Emmons and others) he applied the name "Monument 

Creek formation," and to the upper division (the 

I' Upper 1\10nument Creek" of Hayden), "the youngest 

formation in the Castle Rock region," he gave the 

name "Castle conglomerate." Lee did not obtain 

paleontologic data bearing on the age of either of 

the divisions of the" Monument Creek ,,- as recognized 

by him, but three years later Darton 18 obtained additional vertebrate evidence regarding the upper division, correlating it with the Chadron formation of 

the White River group-that is, with the Oligocene. 

He did not obtain evidence as to the age of the lower 

division but said: "The presence of the unconformity 

between the upper and lower members suggests that 

the latter may be of \Vasatch or Bridger age." 

The status of the "Monument Creek" problem remained as indicated above until 191~11, when G. B. 

Richardson began a study of this region in preparation 

for a report on the geology of the Castle Rock quadrangle, which is joined by the Denver Basin on the 

north and by the Colorado Springs quadrangle on the 

south. As a result of this study Richardson 19 recognized and confirmed the twofold division of the 

"Monument Creek" and the separation of the two 

divisions by a marked unconformity. To the lower 

division he gave the name Dawson arkose, and to the 

upper division the name Castle Rock conglomerate. 

The Castle Rock conglomerate-that is, the " Upper 

Monument Creek" of Hayden and others; the so-called 

upper division of the" 1\10nument Creek" of Emmons, 

17 Lee, W. T., The areal geology of the Castle Rock region, Colomuo: Am. Geologist, vol. 29, pp. 102, 103, 1902. 

18 Darton, N. H., Age of the Monument Creek formation: Am. Jour. 

SCi., 4th ser., vol. 20, pp. 178-180, 1905. 

10 Richardson, G. B., The Monument Creel, group: Geol. Soc. America 

Bull., vol. 23, pp. 267-276, 1912. 

Eldridge, Darton, and others; and the" Castle" conglomerate of Lee (Lee's name being unavailable on 

account of prior usage)-occurs in the south-central 

part of the Denver Basin, where it crops out in detached areas on the divides between the South Platte 

and Arkansas drainage basins from the vicinity of 

Elbert to Sedalia, a distance of about 30 miles. It is 

a remnant of a once larger formation that has been 

reduced by erosion. Richardson obtained additional 

vertebrate evidence for the correlation of the Castle 

Rock with the Chadron formation, the lower formatiop of. the White River group, of Oligocene age. 

In the present connection of COurse the greater interest 

attaches to the Dawson arkose, the exact age of which 

had been so long in question. It reaches a maximum 

thickness of about 2,000 feet on the west, toward its 

, source in the mountains, and becomes thinner as it 

extends eastward on the plains. The Dawson arkose 

was described at length by Richards~m/o who said: 

This formation is composed of a complex mass of yaricolored andi varitextured arkosic conglomerate, sandstone, 

shale, subordinate carbonaceous deposits, and clay derived 

chiefly from, the rocks of the Front Range and deposited under 

various continental conditions. Sandstones comprise the 

greater part of the Dawson arkose. They are medium to 

coarse textured arkosic grits. ... * * Beds and lenses of 

conglomerate occur throughout the formation. ... * * It also 

includes local bodies of clay. ... • ... In the valley of Cherry 

Creek, at the north end of the quadrangle, there is an exposure 

of sandstone and shale composed of andesitic materia1. 

Con~erning the relation between the Dawson arkose 

and the Arapahoe and Denver formations Richardson 

wrote: 

Recent studies in the Castle Rock quadrangle have led to the 

recognition of previously unsuspected relations between the 

Arapahoe and Denver formations and the Monument Creek 

group of Hayden. ~he upper part of the "Monument Creek 

group" is now known to be Oligocene, and a portion of its 

lower part, the Dawson arkose, is known to be equivalent to 

the Arapahoe and Denver formations. 

The Dawson arkose strikes into and merges into both the 

Arapahoe and Denver formations. Anyone following the Dawson arkose northward along the strike from the vicinity of 

Dawson Butte to the area mapped as Arapahoe in the Denver 

monograph will find it impossible to separate these formations 

even at the type locality of the Arapahoe along the bluffs 

of 1Villow Creek, 3 or 4 miles southeast of the entrance to the 

Platte Canyon, which is only 3 miles west of the Castle Rock 

quadrangle. 

Likewise, the Dawson arkose and Denver formations merge 

into each other. Changes from andesitic to arkosic material 

occurring as interfingering lenses are common near the south 

end of the Denver formation, a few miles north of the Castle 

Hock quadrangle. * • • The marked difference in lithology between the andesitic Denver and the arkosic Dawson 

may be accounted for by the geographic distribution of the 

III Richardson, G. B., U. S. Geol. Survey Geo!. Atlas, Castle Rock 

folio (No. 198), p. 7, 1915. 
HISTOIUCAL REVIEW 5 

l'OC]\:S that supplied the sediments, the arlwse being derived 

mninly fl'om the Pikes Peak granite and associated rocks. 

whel'ens the Denver formation was derived, fiPl)arently simultaneously, from a local source of andesite. 

MIDDLE PARK FORMATION 

For many years it was supposed by the earlier geologists that there was a considerable area of Laramie 

rocks forming the divide between JVliddle and North 

Parks. This was the view held by I-Iayc1en,2l who 

visited Middle Park in 1869 and was the first to describe the beds under consideration, but as his was 

only a hasty reconnnissance trip, mninly on the Colorado (formerly Grand) River near Hot Sulphur 

Springs, the description is necessarily brief. He mentioned especialiy the "Lignitic" beds, which were 

regarded "as of the age of the coal formations of 

the 'Vest-older Tertiary." The beds beneath the 

" Lignitic" I-Iayden called basalt, but they are now 

known to be the volcanic breccia and conglomerate of 

la.ter writers. Fossil plants ,,,ere collected at a number 

of localities. 

The first adequate description of the geology of 

Middle Park was made by l\1arvine:!2 in 1873. The 

geologic map of Colorado issued by the I-Iayden Survey, based largely on l\1nrvine's work, shows the supposed Lara.mie occupying" a large area and extending southeastward from the Continental Divide into 

Middle Park. The area gradually narrov\TS and just 

uefore reaching the Colorado RiYer divides into two 

arms, the eastern of which crosses the river east of 

lIot Sulphur Springs and tenninates at a point some 

4: or 5 miles south of the river. This ann is sommvhat 

spoon-shaped, and as the beds clip inward on all sides 

from the rim of dark volcanic breccia it was desigJUtted the" breccia spoon" by Marvine. The western 

arm ends its continuous exposllre at l\10unt Bross, on 

the north side of the Colorado River, though it continnes intenuptedly for some distance toward l\10unt 

Byers. In describing this area Cross ~3 said: 

In the angle between the arms of "T..Juramie" appear the 

strnta Oof the Cretaceous section, fL'om tile Dakota tOo the Fox 

I-lills, inclusiYc, with n northwest-southeast strike and a nort~leasterly clip of J 5°-25°, which carries them in apparent COI1-

fOI'mitv below the strata of the "breccia spoon," while tIle 

" J..J:Hu·;nie" beds of Mount Bross and the ridge tOo the southeast rest in nearly horizontal position upon Archean and the 

edges of the Dakota find ColorndOo shales. '1'his is the unCOIlfOl'1nity discovered by Marvint'. 

~l Hayden, 11'. V., Preliminary field report of the United States geologIcal survey of Colol'ado and New )Icxico, pp. 80-86, 1869; reprint, 

187:~, pp. 181-186, 193. 

~~ 1\1nr\'ille, A. n., U. S. Geo1. nnd Geog. Sun-cy Terr. Ann. Uept. for 

1873, PI). 154-11:12, 1874. 

~3 C.·oss, Whitman, Thc post-Lnl'aUlic beds ill )I1ddle Park, Colo.: 

COIOl'lIdo ScI. Soc. PI'OC., 1'01. 4, p. 194, 1895. 

80402-30--2 

I 

The doleritic beds next above the Cretaceous were 

stated by l\1arvine to be 800 or 900 feet thick, and 

above them occurs some 5,500 feet of the "Lignitic 

I

i series. " 

" In 1877 C. A. 'Vhite visited Middle Park and 

searched the supposed Laramie for fossils, but without avail, though he refers to a few invertebrates 

procured by l\1arvine in North Park, probably in the 

same formation. Concerning these 'Vhite 24 said: 

Of themselves they are not sufficient to determine the age 

of the strata containing them or their equivalency or otherwise with thOose of the Lfil'flmie group. ... ... ... The Middle 

Park stl'fitn contain no fossils thnt nre certainly identical in 

sDecies with any of the numerous forms found on each side 

of that region fl t the eastern and western bases, reSI)ectively, 

of the Hocky Mountains; and they contain, so fur as known, 

ollly those imperfectly ImOoW11 species [from North Park] 

before referred to that are possibly identical with forms in 

the Fort Union beds of the upper Missouri. 

The status of know ledge concerning the beds in 

l\1iddle Park remained as indicated above for a decade 

or more after 'Vhite's visit, but by that time the 

Arapahoe and Denver formations in. the Denver 

Basin had been demonstrated to be distinct from the 

Laramie. From the early descriptions of the lithology 

and supposed stratigraphic relations of the so-called 

Laramie in :Middle Park it was conjectur~d that 

those beds might show a similarity to the Denver formation for at least a part of the series. In order to test 

this suo'o'ested relation S. F. Emmons, in charge of the 

Colorad~ division of the United States Geological 

Survey, sent George L. Cannon, jr., to l\1idclle Park in 

the summer of 1889, "and his report amply confirmed 

this idea. "Thitman Cross also visited the area in the 

fall of 1890 and ao-ain in 1891, at first devoting his attf>ntion to the lith~logic character of the breccia and of 

the beds above it, as well as to evidences "of unconformity. The early observations were largely of local 

character, but, as Cross said: "The question at issue 

soon expanded to one regarding the geological history 

of .the 'v hole l\1idcUe and North Park areas in the 

period following the conformable series of Cretaceous 

deposits." The results of this study by Cross 25 were 

published in 1892. I-Ie described first the so-called 

" doleritic breccia," which in some places rests on undoubted Fox Hills and in others apparently on Pierre 

shale. This series of beds is composed largely of 

andesitic material; with fine-grained tuff alternating 

irregularly with conglomerate. "There the beds are 

massive bold-faced cliffs are found, but where the beds 

:>1 WlJite, C. A., U. S. Geol. and Geog. Survey Terr. Ann. Rcpt. for 

1877, p. 203, 1870. 

:05 Cross, WlJitman, The post-Laramie beds in Middle Park, Colo.: 

Colorado Sci. Soc. Proc., vol. 4, pp. 192-213, 1895; separate pl'lnt, pp. 

1-27, 1892. 
6 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

are finer grained the cliffs are broken down by cross 

ravines and depressions. Fossil plants occur at a number of horizons within this series. Overlying the 

breccia are the" lignitic " beds, which are separated by 

beds of transition and not by a distinct erosion interval as supposed by Marvine. They are composed of 

tuff, conglomerate, and sandstone and are probably 

over ,5,000 feet thiclc Fossil plants are found at various points but especially toward the base. Cross gave 

a list of the fossil plants then known from the postLaramie I:>eds in Middle Park, prepared by me, and 

tentatively referred these beds to the approximate age 

of the Denver formation. The present study confirms 

this reference. 

PLANTS IN THE EQUIVALENT OF THE ARAPAHOE 

FORMATION 

Before passing to the consideration of the Denver 

formation and its flora it is necessary to dispose of the 

supposed flora of the Arapahoe formation. According to Eldridge, on whom largely devolved the task 

of studying, the Arapahoe formation in the field f01' 

the' Denver Basin monograph, the Arapahoe was 

found'to contain poorly preserved plant remains at a 

number of localities in the Denver reO'ion but no de- • b , 

termInable remains were obtained. In fact, only two 

localities appear. to have been recorded from which 

determinable plants sup'posed to be of Arapahoe age 

had been reported-Sand Creek, east of Denver and 

the vicinity of the old Douglas coal mine, near Sedalia, 

about 20 miles south of Denver. 

The so-called Sand Cl.'e'ek l~cality may be first considet'ed. The following account is taken from my 

report on the flora of the Laramie formation 26 where . ' It was necessary to consider this flora in connection 

with the Laramie flora: ' 

Some 9 or 10. species of plants are recorded as having come 

from the Sand Creek locality, all of which were obtained by 

the earlier workers in this field-that is, probably about 1873. 

Considerable uncertainty has existed as to the exact locality 

whence these plants c'ame, but apparently important light is 

thrown on this point by a chance statement made by A. R. 

Marvine 27 in his paper on "The geology of Middle Park," which, 

on account of its importance, is quoted entire below. 

"East of Denver, in T. 4 S., and probably between Rs. 65 

and 66 W., n shaft has been sunk for some depth in a high 

bank on the south side of Sand Creek but is now abandoned. 

The coal also outcrops on the bank and there appears as of 

very pOOl' quality. Fossil leaves are abundant. 

" It was near here that the first discoveries of coal were made 

in Colorado, and the stream is often known at this point as Coal 

Creek, though called Sand Creek farther down. The latter 

name should be retained to prevent confusion between this 

and the better-known Coal Creek on the west side of the Platte. 

26 Knowlton, F. H., U. S. Geol. S~rvcy Prof. Paper 130 pp. 102-103 

1922. ~ , 

27 U. S. Gcol. and Geog. Survey Terr. Ann. Rept. for 1873, PP. 120, 

121, 1874. 

"About 4 miles to the north, near Box Elder, on the Kansas 

Pacific Railroad, in R. 65 W., T. 3 S., sec. 28 (1), and probably 

at the same horizon as the last, are two shafts which reach 

coal and on which work has been done now and then for some 

years. It is probably in one of these three shafts that the 

following section was made by Mr. E. B. Mally (quoted by 

Lesquereux in Hayden's report for 1872, p. 327), and which 

seems to give an idea of the strata near here. I) * * 

"The work was abandoned on account of the poor quality 

of the coal." 

That the above-indicated plant-bearing horizon is probably 

the important one in the preEent consideration was still further 

indicated by Marvine, who, after mentioning a nu'mber of co~l 

openings in another direction from Denver, said: "The only 

others I have heard of lie from 1.5 to 17 miles east of Denver 

City and near the Box Elder station on the Kansas Pacific 

Railroad." The assumption seems to be justified, therefore, 

that the plants labeled" Sand. Creek" probaply came from this 

area. 

In Lesquereux's "Tertiary flora" the following data are 

recorded concerning six of the species reported to have come 

from Sand Creek: 

Lash'ea (Goniopteris) polypodioides, Sand Creek nV. H. 

Holmes) . 

Gymnogramlllu [Gymnogramme] gardneri. Roof of a coal 

mine, Sand Creek (A. Gardner). 

Equisetum laevigatum. Sand Creek, 8 feet above coal (W. H. 

Holmes) , 

Eriocaulon? porosum. Sand Creek CWo H. Holmes), with 

leaves of Nelumbium. 

Nelumbiulll tenuifolium. Sand Creek, Colo. (A. Gardner). 

Quercus viburnifolia. Sand Creek (A. R. Marvine). 

These species are similarly recorded in the catalogue of fossil 

plants in the United States National Museum, in Lesquereux's 

own handwriting, so it may be presumed that the statemellt~ 

are correct, at least so far as his information went. The 

species obtained by Marvine were doubtless collected on the 

occasion of his visit there in 1873, but no data are available 

as to the date on which the two species accredited to Professor 

Gardner were collected. 

W. H. Holmes assisted Marvine in his geologic work ill 

1873, and the plants recorded as having been collected by him 

were probably obtained at that time, but on consultation with 

Pl'ofessor I-Iolmes I find that he has ilOW no recollection of 

having collected the plants, or, indeed, of havi'ng visited any 

coal mines east of Denver. This injects another element of 

uncertainty into the already sufficiently complicated matter, 

hut we shall probably never come any nearer to a COml)lete 

explanation. To sum up, it appears in its final analysis that 

we do not know the precise locality for a single one of these 

Sand Creek plants, though in all reasonable probability they 

apparently came from the vicinity of the coal openings on 

Coal or Sand Creek about 15 miles east of Denver, but as the 

locality is in doubt, it naturally follows that the horizon is also 

nncertain. It was apparently assumed by Eldridge that they 

came from the Arapahoe, for he stated that along Sand Cre('k 

this formation rests on the Laramie, but according to the map 

showing the areal geology in the Denver Basin monograph the 

Laramie occurs on the north bank of the creek and the Denver 

on the south bank, no Arapahoe being indicated. Cross,2S in 

discussing the Sand Creek locality, said: "The specimens preserved in the National Museum do not satisfactorily indicate 

the horizon fl'om which they came. It seems probable that a 

23 Cross, Whitman, U. S. GeoI. Survey Mon. 27, p. 225, 1896. 
THE FLORA 7 

part of them came from the Arapahoe beds and a part from 

t'lle I.Jaramie." 

Up to this pOint, from the data available, it appeared to me 

that if any of the Sand CL'eek plants came from the Laramie 

they probably all did, and on this basis it was my original 

intention to include them provisionally in the present work 

[l~aramie fiora]. However, important observations made by 

W. T. Lee during the field season of 1915 put an entirely different light 011 the matter and have led me to exclude all the Sand 

CL'eel{ Sl)ecies from both Arapahoe and Laramie and to refer 

them with little 01' no question to the Denver. The evidence is 

ns follows: 

Lee exnmined the log of a deep wen that has been drilled 

11 bout 23h miles northeast of Coal Creel{ (01' Sand Creek). 

l'hls well starts at the surface in sandy coal-bearing beds 

that are lithOlogically and stratigraphically identical with the 

beds in question along Coal Creek and are also the same as 

the coal-bearing beds at Scranton, which from the presence 

of conI nre supposed to be in the upper part of the Laramie. 

1$1 '" I) 

l'11e facts brought out by this well ~:ecord are of far-reaching 

significallce. It shows that the SCL'anton coal, heretofore 

thought to be in the upper part of the Laramie, is about 350 

feet above [the basal conglomerate of the Arapahoe formation. 

This conglomel'llte, which is about 50 feet thick, is characterisl"ic, being made up of the usual assortment of pebbles from 

the older rocl{s]. It also appears that the Scranton coal is 

1ll0L'e than 400 feet above bedS that can with reasonableness 

.ue referred to the Laramie. 

The plants from the vicinity of the old Douglas coal 

mine (later called the Lehigh mine), nearly 4 miles 

west of Sedalia, have' also had a varied history. They 

were collected by Arthur Lakes in 1890 and are recorded as coming from three localities: 

325. Quarry No.2, 3,000 feet east of Douglas coal mine. 

331. Quarry No. l, l,900 feet east of Douglas coal mine. 

332. Douglas coal mine [which is ,in the Laramie formation]. 

These localities were mentioned in the Denver 

Basin monograph 20 and the collections fr~m the localities east of the coal mine were spoken of as ob-' 

tained " in what are apparently Arapahoe beds." The 

larger part of the flora came from quarry No.2, 3,000 

feet east of the Douglas mine. The mine is about 5 

miles south of the southern boundary of the Denver 

quadrangle and 2 miles west of 'the western boundary 

of the Castle Rock quadrangle and is very near the 

t.ype locality of the Arapahoe formation, which was 

originally named" 'V'illow Creek beds" fr01n expoSllres on Willow Creek, 3 or 4 miles southeast of the 

entrance to the Platte Canyon. In the Castle Rock 

folio Richardson maps all the northwest corner of the 

quadrangle around Sedalia as Dawson arkose, and in 

n. sketch map (fig. 5, p. 10) he shows that formation extending to the, narrow band of Laramie in 

which the Lehigh· mine is located. Another sketch 

map (fig. 3, p. 7) shows the areal relations of the 

Dawson to the Arapahoe and Denver in this vicinity. 

~ u. S. (leal. Survey MOil. 27, pp. 225, 469, 1896. 

On page 8 of the text he gives a list of "plants collected by Arthur Lakes from Dawson arkose, NE. 14 

sec. 20, T. 7 S., R. 68 W., 3 nliles west of Sedalia, 

,700 to 900 feet above base [top 30] of the Laramie 

formation." This list is based on Lakes's collections 

from the two localities east of the Douglas mine and 

has since been revised. The rocks from which the 

plants came are in a.n area where the lith~logic distinction between the Arapahoe and the Denver is no 

longer valid for drawing a stratigraphic boundary between these two formations. Nevertheless the horizon 

is so l~w in the Dawson that it is clearly within the 

equivalent of the Arapahoe formation as mapped 

'along the southern boundary of the Denver quadrangle, only 5 miles away, and the species from it are 

designated as Arapahoe species in the general list of 

the flora which follows, though it may reasonably be 

questioned whether they are actually older than the 

plants from Sand Creek above discussed. 

THE FLORA OF THE DENVER AND RELATED 

FORMATIONS 

LIST OF SPECIES 

Before proceeding to a discussion of the flora of 

the Denver and Arapahoe forma.tions, Dawson arkose, 

and Middle Park formation, here spoken of as the 

Denver and related formations, I wish to present the 

following as a complete list of the forms recognized. 

In this list the forms from the Denver formation are 

. indicated by 1, those from the equivalent of the Arapahoe formation in the Dawson arkose by 2, those from 

the Dawson arkose in general by 3, and those from 

the l\1iddle Park formation by 4. 

1 Sclerotites rubellus CCesquereux) Meschinelli. 

3 Sclerotites? cypericola Knowlton, n. sp. 

3 Sclerotites? hesperius Knowlton, n. sp. 

3 Marchantites? coloradensis Knowlton, n. sp. 

1 Hymenophyllulll confusul11 Lesquereux. 

4 Dl'yopteris integra Knowlton, n. sp. 

1,2,4 Dryopteris lakesii (Lesquereux) Knowlton, n. comb. 

2 Dryopteris richardsoniana Knowlton, n. sp. 

1 Dl'yopteris polypodioides? (Ettingshausen) Knowlton. 

2 Dennstaedtia crossiana Knowlton, 11. sp. 

l,2 Woodwardia latiloba Lesquereux. 

4 Woodwardia latiloba sen'ata Knowlton, n. val'. 

1 Pteris pseudopennaefol'mis Lesquereux. 

1,4 Diplazium crossii (Knowlton) Knowlton, n. comb. 

L, 2, 4 Al1antodiopsis erosa (Lesquereux) Knowlton anll 

Maxon. 

1,3 Saccoloma gardneri (I.lesquereux) Knowlton. 

3 Saccoloma sp. 

1 Salpichlaena anceps (Lesquereux) Knowlton. 

3 Anemia mosbyensis. Knowlton, n. sp .. 

4 A.nemia lanceolata Knowlton, n. sp. 

4 Anemia sp. 

30 In n pel'sona1 communication Ml·. Richardson states that the diRtance is above the top of the Laramil', not the base. 
8 .FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

3 

3:4 

1 

1 

1 

4 

1 

1 

1 

1,3 

3 

3 

1 

1 

1 

4 

1 

1 

1 

1 

1 

1 

1 

1 

1 

3 

1 

1,4 

1 

Lygodium coloradense Knowlton, n. sp. 

Equisetum coloradense Knowlton, n. sp. 

Equisetum sp. 

Equisetum sp. 

Selaginella berthoudi Lesquereux. 

Sequoia affinis Lesquereux. 

Ginkgo'! truncata (Lesqucreux) Knowlton, n. comb. 

Arundo? obtusa Lesquereux. 

Carex? bel'thoudi Lesquel'eux. 

Sabalites grayanus Lesquel'eux. 

Sabal? ungeri (Lesquel'eux) Knowlton. 

Geonomites tenuil'achis Lesquereux. 

Geonolllites? gl'alllinifoHus Lesquereux. 

Geonolllites goldianus (Lesquereux) I.esquereux. 

Geonolllites? sp. 

Chamaedorea? coloradensis Knowlton, n. sp. 

Paloreoeloxites plicatus (Lesquereux) Knowlton, n. 

comb. 

Pulmocul'pon palmurum (Lesquereux) Knowlton. 

Pahnocal'pon truncatum Lesquel'eux. 

Pahnocal'pon? subcylindricum Lesquereux. 

Palmocal'pon? corrugatulll Lesquereux. 

Palmocal'pon? linea tum Lesquereux. 

Palmoxylon cannoni SteVE'lls. 

Zingiberites elubius Lesquereux. 

Piper heerii Lesquereux. 

Myrica sp. 

Juglans denveriana Knowlton, n. sp. 

Juglans the'rmalis Lesquel'eux. 

Juglans schimperi Lesquereux. 

1 Juglnl1s l'hamnoides Le'squereux. 

1 Juglans rugosa Lesquereux. 

3 Juglans sp. 

1 Hicoria antiquora (Newberry) Knowlton. 

1 Ptel'ocarya? retusa Lesquereux. 

1 Alnus carpinifolia Lesquereux. 

1 Alnus aural'ia Knowlton anel Cockerell. 

1 Betula schimperi Lesquereux. 

1 Betula gracilis? Ludwig. 

1 Betula fallax Lesquereux. 

3 Castanea intermedia? Lesquereux. 

1, 3, 4 Quercus viburnifolia Lesquereux. 

2 Quercus? seelali.ensis Knowlton, n. sp. 

3 Quercus purelonensis Knowlton, n. sp. 

4 Quercus whitmnni Knowlton, n. sp. 

1 Quercus whitei Lesquereux. 

1 Quercus leonis Knowlton, n. sp. 

1 Quercus celn stl'ifolia r.esquereux. 

I Quercus coloraelensis Lesquereux. 

1 Populus zeilleri (Lesquereux) Knowlton, n. comb. 

I Populus nebrascensis Newberry. 

1 Populus nebrascensis granc1:elentata Lesquereux. 

1 Populus nebrascensis rotundata Lesqu€reux. 

1 Populus nebrascensis .acmidentata Lesquel'eux. 

1 Populus neb'rascensis longifolia Lesquel'eux. 

1 Populus'? sp. 

1 Populus jacksoni Knowlton, n. sp. 

1 Populus tenuinervata Lesquel'eux. 

1 Populus subrotunda Lesquel'eux. 

4 Populus zadelachi Heer? 

1 Populus lacoeana Knowlton, n. sp. 

3 Populus knowltoni Berry, n. sp. 

1 Populus dem"erensis Knowlton, n. sp. 

4 Populus richardsoni Heer? 

1 Ulmus antecedens Lesquereux. 

I Ulmus? quercifoUa Unger? 

1 l!"icus subtruncata I~esquereux. 

2 Ficus? alata Knowlton, n. sp. 

1, 3 Ficus elenveriana Cockerell. 

1, 3 Ficus aguilar Knowltoll. 

1 l!1.CUS be'l'thouc1i Lesquereux. 

1,3 Ficus pseuelopopulus Lesquereux. 

3 Ficus c1awsonensis Knowlton, n. sp. 

3 Ficus sp. 

1 Ficus sp. 

1 Ficus occidentalis ~Lesquereux) Lesquereux. 

3 Ficus ramahensis Knowlton, n. sp. 

1 li'icus? lakesii Knowlton, n. sp. 

1 Ficus coloradensis Cockerell. 

1 Ficus martini Knowlton, n. sp. 

1 Ficus eldrielgi Knowlton, n. sp. 

1,2 Ficus neoplanicostata Knowlton. 

1 Ficus planicostata goldiana Lesquel'eux. 

1 Ficus planicostata problematic'a Knowlton, n. Y~r. 

3 Ficus praetrinervis Knowlton. 

3 Ficus puryearensis Berry? 

3 Ficus sp. 

3 Ficus sp. 

1 Ficus sp. 

1 Artocarpus pungens (Lesquereux) Hollick. 

3 Artocarpus similis Knowlton. 

3 A'rtocarpus? gigantea Knowlton, n. sp. 

1, 3 Platanus rhomboiclea Lesquereux. 

1 Platanus guillelmae Goppel't. 

1 Platanus aceroides latifolia Knowlton. 

1 Platanus raynoldsii Newberry. 

1,4 Platanus raynolc1sii integrifolia Lesquereux. 

1,2 Platanus haydenii Newberry. 

3 Platanus marginata (Lesquereux) Heel'. 

3 Platanus platanoic1es (Lesquereux) Knowlton. 

4 Platanus coloraelensis Knowlton, n. sp. 

3 Platanus sp.? 

1,2,3 Laurus primigenia Unger. 

1?,3 r.aurus sodalis Lesquereux. 

3 Laurus lanceolata ~(nowlton, n. sp. 

4 Per sea brossiana Lesquereux. 

3 Oinnamomum ·sezannense Watelet? 

3 Cinnamomul11 linifolium Knowlton. 

3 Cinnamolllum? sp. 

1 Cinnamol11um sp. 

1 Mngllolia l11agnifolia Knowlton. 

1,2 Nelumbo lakesiana (Lesquereux) Knowlton. 

1 Nelumbo tenuifolia (Lesquereux) Knowlton. 

2 Nelumbo crossii Knowlton, n. sp. 

3 PaleolH~lumbo, nUlcroloba Knowlton, n. gell., 11. st). 

3 Castalia pulcheUa Knowlton, n. sp. 

1, 2, 3, 4 Asimina eocenica Lesquereux, 

1,3 Ohrysobalanus coloraelensis Knowlton, n. sp. 

3 Chrysobalanus? lanceolatus Knowlton, 11. sp. 

1 Crataegus myricoic1es Le~quereux. 

3 Prunus denverensis Knowlton, n. sp. 

3 Phaseolites coloraelensis Knowlton, n. sp. 

3 Sophora richardsoni Knowlton, n. sp. 

3 Sophora puryearensis Berry? 

1,4 Leguminosites? arachioides (Lesquereux) Lesquereux. 

3 Oassia vetustn Knowlton, n. sp. 

3 Cercis coloradensis Knowlton, n. sp. 

1 Celastrus gaudini Lesquereux. 

1 Celastrinites artocal'pidioicles I.esquereux. 

1 Oelastrinites populifolius Knowlton, n. sp. 

1 Negunclo clecurrells Lesquereux. 

2 Acer sp. 
1 Sapindus caudntus Lesquereux. 

3 Sapillc1us berryallus Knowl ton, n. 8p. 

1 Sapindus? obtusifolius Lesquereux? 

1 Ilex'? ovnta Knowlton, n. sp. 

1.,2 Berchemia multinervis. (AI. Bruun) Heel'. 

1 Hhamnus? prnenlnternoides Knowlton, n. 8p. 

1 Hhnmnu8 goldianus Lesquereux. 

1.,3 Hhalllnus cleburni Lesquereux. 

1. Uhnmnus crenulatlls Knowlton nnd Cockerell. 

4 Uhnml1us cannoni Knowlton, n. sp. 

3 Hluunnus salicifolius Lesquereux? 

1 Paliurus coloradensis Lesquereux. 

1., 3 Zi~yphus fibrillosus (Lesquereux) Lesquereux. 

1. 7Jizyplms daphnogenoides Knowlton. 

1 Zi~yphus beckwithii Lesquereux. 

1., 3 Zizyphus distortus' Lesquereux. 

1 Zizyphus lesquereuxii Knowlton. 

1: Zi.zyphus hespel:ius Knowlton, n. sp. 

3 Cissus gros,'Se-dentata Knowlton. 

3 CiS8US hesperia Knowlton, n. sp. 

1,2 Cissus colorndensis Knowlton and Cockerell. 

4 Cissus'l cannoni Knowlton, n. sp. 

1 Cissus cOl'ylifolia Lesquereux. 

1.,3 Cissus lesquereuxii Knowlton. 

1 Cissus obovata Knowlton, n. 8p. 

2,3,4 Ctssus lobato·crenata Lesquel'eux. 

3 Vi tis olriki Heel'. 

1, 3 GI'ewiopsis telluifolia Lesquereux. 

3 Sterculia? heterodonta Kllowlton, n. sp. 

I? SteL'culia saportanea Knowlton. 

1 Sterculia libbeyi Knowlton, n. sp. 

1 Pterospel'luites grulldidentata Lesquereux. 

1, 3, 4 Comus impressa Lesquereux. 

1 Comus lakesii Knowlton, n. sp. 

1 Cornus denverensis Knowlton, n. sp. 

1 Corllus holmesii Lesquel'eux. 

1, 3 Nyssa lnnceolata Lesquereux. 

1 Nyssa denverinlla Knowlton, n. sp. 

1 Nyssa? obovntn Knowlton, n. sp. 

1, 2, 3 Diospyros brnchysepala AI. Bl'Ilun. 

1 Styrax'? lal'mniensis Lesquereux. 

1 ]!'rnxinus eocenica Lesquereux. 

1 lJ'l'axinus sp. 

1.,2,3 Dombeyopsis obtusa Lesquereux. 

3 Dombeyopsis magnifica Knowlton, n. SlJ. 

2 Dombeyopsis? sedaliensis Knowlton, n. sp. 

1,2,3 Viburnum richardsolli Knowlton, n .. sp. 

3 Vibunllun contortum Lesquereux? 

1 Viburnum? hetel'odontum Knowlton, n. sp. 

1,3 Viburnum lakesii Lesquereux. 

3 Viburnum melaenum Knowlton and Cockerell. 

1 VibUL'llum goldianum Lesquereux. 

1 Viburnum solitarium Lesquereux. 

3 Phyllites andromeda Knowlton, n. sp. 

2 Phyllites aristolochioides Knowlton, n. s11. 

1 Phyllites denvel'ensis Knowlton, n. sp. 

3 Phyllites sp. 

3 Phyllites calhunensis Knowlton, n. sp. 

3 Phyllites pellucidus Knowlton, n. sp. 

1. Curpites ovifol'mis Lesquereux. 

3 Carpites qundrivnlvis Knowlton, n. sp. 

3 Carpites templetoni Knowlton, n. sp. 

1 Curpites rostellatus Lesquereux. 

1 Curpites costatus Lesquereux. 

1 CUl'pites coffeneformis Lesquel'eux. 

THE FLORA 

1 Cnrpites lllillUtulus Lesquel'eux. 

1 Carpites myricnrum Lesquereux. 

3 Carpites coryloic1es Knowlton, n. sp. 

1 Eriocaulon? porolSulll Lesquereux. 

1,3 Berrya rncemosn (Knowlton) Knowlton, n. comb. 

DISTRIBUTION OF THE FLORA 

9 

The complete flora as now recognized includes 222 

forms, of which 145 occur in the Denver, 22 in the 

equivalent of the Arapahoe, 95 in the Dawson arkose 

in general, and 24 in the ~1iddle Park formation. Of 

the 222 forms, 69 are regarded as new to science in the 

present paper, and 19 are not named specifically, leaving 134 species previously InlOwn, 70 of which have a 

distribution beyond the limits of the formations here 

considered. 

Naturally much interest attaches to the relation between the flora of the Denver and associated formations and the floras of the underlying Cretaceous 

(Laramie) . Only nine species, three of which are 

more or less in doubt, are known to cross this line, as 

follows: . 

Quercus viburnifolia. 

Ficus denveriana? 

JJ'icus coloradensis. 

Ficus praetrinel'vis. 

Platanus platanoides. 

Nelulllbo telll,lifolia. 

Dombeyol1sis obtusa. 

Rhamnus goldianus', 

Rhamnus salicifolius? 

Of the three species that are questioned two (Ficus 

denve"'iavna and Rhcm711n~lS goldianU'8) are essentially 

Denver species, and their presence in the Laramie 

rests on single poorly preserved examples, and the 

other (Rha17W1/UJS salicifolius) is essentially Laramie 

with a single questionable leaf in the Dawson arkose 

at Templeton Gap, near Colorado Springs. As the 

Laramie flora of the Denver Basin includes 129 forms 

and the Denver and associated formations 222 forms, 

with only 9 in common, it shows that the relationship 

is not marked. 

The closest relationship of the flora under consideratlOll is undoubtedly with that of the Raton formation. 

The following species are held in common: 

Allnntodiol)sis ero:;:.a. 

Pnloreodoxites plicatns. 

Subalites grayunus. 

Sabal? ungeri. 

Geonoll1ites tenuirachis. 

Geonomites goldianus. 

Palll1ocarpon palmnrUlll. 

Juglans schimperi: 

Juglans rugosa. 

Castanea intermedin. 

Ficus denverialla. 

Ficus pseudopopulus. 

Ficus aguilar. 

Ficus occidentalis. 

JNcus neoplanicosta ta. 

Ficus planicostata goldiana. 

Ficus prnetrinervis. 

Artocarpus similis. 

Platanus rholl1boide:1. 

Platanus guillelmllc. 

Platanus aceroides. 

Platanus raynoldsii. 

Platanus platanoides. 

LalU'us socialis. 

Cinllumolllum linifoliull1. 

Magnolia ll1agnifolia. 

Nelull1bo lakesiana. 

Leguminosites arachioides. 

Berchemia multinervis. 

Rhalllnus goldianus. 

Uhalllllus clebul'ni. 

Zizyphus fibl'iUosus. 
10 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Cissus grosse-dentata .. 

Cissus coloradensis. 

Vitis olriki. 

Nyssa lanceolam. 

Viburnum contortum. 

Viburnum lakesii. 

Berrya racemosa. 

This list shows that there are 39 species, or nearly 

60 per cent of the forms having an outside distribution, that are found in the Raton formation. 

There is also a measure of relationship between the 

Denver and related formations and the Wilcox group 

of the Gulf region, as indicated by the following list 

of comlnon species: 

Pte'ris pseudopennaeformis. Ficus occidentalis. 

Sabalites gl'ayanus. Ficus neoplanicostata. 

Juglans rugosa. ]!"icus puryearensis? 

Hicoria antiquora. Artocarpus pungens. 

Ficus denverialla. Leguminosites arachioides. 

Ficus pseudopo,pulus. Diospyros brachysepala. 

The Wilcox flora IS a very large one, comprising 

over 500 species, over 350 of ,,,hich are confined· to 

the 'Vilcox beds. Berry has expressed the opinion 

that the Wilcox is slightly younger than the Raton 

and Denver, but a part at least of the difference in the 

Horas is to be accounted for on the ground that the 

vVilco'x is largely a strand flora, whereas the Raton 

and Denver floras are largely of inland and swamp 

types. 

In southwestern Colorado and adjacent New . Mexico 

the Animas formation shows a very strong relation 

,vith the Denver and associated formations, as shown 

by the following list of common species: 

Allantodiopsis erosa. 

Juglalls schimperi. 

Castanea intermedia. 

Ficus occidentalis. 

Ficus denveriana. 

E'icus subtruncata. 

Artocal'pus pUllgens. 

Platanus aceroides. 

Platanus raynolc1sii. 

Lam'us primigenia. 

Nelumbo lakesiana. 

Magnolia magllifolia. 

Rhamnus cleburlli. 

Rhamnus goldianus. 

Zizyphus fibrillosus. 

Zizyphus daphnogenoides. 

Cornus studeri? 

Berrya racemosa. 

The Animas formation has a flora of 58 species, 38 

of which have been found elsewhere. As shown by the 

list, 18 of these species are found also in the Denver. 

A further comparison discloses the fact that 34 of thesp, 

38 species are found in the Denver, Raton, and 'Vilcox 

floras, which 'indicates that these are approximately of 

the same age. 

Turning now to the north, we may make a compari-

. son with the floras of the Lance and Fort Union 

·formations. The Lance contains the following species 

·common to the Denver and related formations: 

SulJa lites? grayanus. 

'Palmocarpon palmarum. 

. Juglans rugosa. 

:Hicoria nntiquora. 

:Populus nebrascensis. 

:.Populus subl'otunda. 

Quercus viburnifolia. 

Platanus l'homboidea. 

Platanus guillelmae . 

Platanus raYlloldsii. 

Platanus raynoldsii integrifolia. 

Platanus haydenii. 

Platanus marginata. 

Platanus platanoides? 

Leguminosites? arachioides. 

Berchemia multinervis. 

Ficus denveriana. 

The following species are common to the Fort Union 

formation: . 

Juglans rugosa. 

Hicoria alltiquora. 

Populus nebrnscensis. 

Platanus raynoldsii. 

Platanus haydenii. 

Asimina eocenica. 

Leguminosites arachioides. 

Berchemia multinervis. 

Diospyros brachysepala. 

A final comparison can not now be made between 

the Lance flora and that of the Denver and related 

formations, for the reason that the Lance flora is much 

in need of critical revision. The known Lance flora 

numbers about 125 forms, of which over 80 occur also 

in the overlying Fort Union formation. Over 250 

species are listed from the Fort Union formation, and 

it is probable that when the very large unstudied collections are fully described the number of species may 

approximate 500. When the Lance and Fort Union 

floras have been fully worked up the number of species 

common to the Denver flora will probably be considerably augmented. As it stands, nearly 30 per cent of 

the portion of the Denver flora having an outside distribution (8.5 per cent of the total flora) is found in 

the Lance and Fort Union. 

Another flora that can not now he fully compared 

'with the Denve;r and related formations is that of the 

Black Buttes coal group at Black Buttes, Wyo. This 

flora embraces about 60 species and has the following 

species in common with the Denver: 

Diospyros brachysepala. 

Ficus coloradellsis. 

Magnolia magnifolia. 

Platanus platalloides. 

q-rewiopsis eoceniea. 

Juglans rl1amnoides. 

Sabalites grayanus. 

Viburnum eontortum. 

Platanus l'aynolclsii integrifolia. Viburnum meIaenum. 

Platanus margillata. Carpites myricarum. 

Qnercus viburnifolia. Berrya raeemosa. 

In all the comparisons thus far made it is believed 

that the formations are of approximately the same 

age. However, certain of the 70 species found beyond 

the borders of the Denver and related formations 

occur in Tertiary beds at higher horizons. Thus with 

the Green River formation we have the following In 

common: 

Pteris pseudopennaeformis? 

Geonomites goldianus? 

Sapindus obtusifolius . 

Juglans schimperi. 

\Vith the Hanna formation, Carbon: Wyo.: 

Populus subrotunda. 

Ficus coloradensis. 

Platanus raynoldsii. 

Platanus guillelmae. 

Lam'us primigenia. 

Zizyphus lesquereuxii. 

.'Vith the Evanston formation, Evanston, Wyo.: 

Juglans rugosa. 

La urns primigel1ia. 

Leguminosites arachioides. 

Vi tis olriki. 
THE FLORA 11 

ECOLOGIC CONSIDERATIONS . 

The caution that is necessary to be observed in treat-· 

ing the probable ecologic conditions under which a 

flora no,v fossil may have lived was set forth in a 

former report 31 as follows: 

In drnwing conclusions from individual organisms in an inquiry of this kind dependence must, 8f course, be placed on 

oUt· knowledge Qf the present-day requirements of similar 

species, nnd the results must always be subject to possibility 

of errol' from two sources-first, from the incorrect placing 

lJiologically of the orgnnism, and, second, from the fact tbat 

its requirements in past geologic time may not have been the 

Sflllle as these which now dominate tbe life activities of its 

:;;upposed annlogue. However, when all of the elements of a 

Hora nppear to point in tbe same direction the liability to 

serious error is minimized if not eliminated. 

The flora of the Denver and associated formations 

numbers 222 forms. There are three species of fungi, 

aU apparently growing on dead wood or decaying 

leaves and stems, but otherwise these requirements can 

HOt", be determined with certainty. The single wellmarked hcpntic (11/ a'rchantites) is of the type growing on the ground or old logs in moist shaded situations. 

The ferns seem to have been especially abundant, 

there being no less than 18 forms representing 3 

families. Thus, the IIymenophyllaceae are represented by n single species with a thin filmy frond that 

must have required a moist, shaded habitat. ~10st of 

the ferns fall within the Polypodiaceae, and among 

them are some of exceptional biologic interest. The 

most abundant species is Dryoptm'is lakesii, which 

occurs mainly in the Denver beds at Golden, Colo. 

It is so abundant that literally hundreds of specimens 

have been collected. Perhaps the most interesting 

form is Allantodiol)S'tS oJ'osa, which is represented by a 

single finely fruiting specimen and by a considerable 

number of large, perfect fronds. Allantodia, with 

which it is undoubtedly closely related, is a monotypic 

genus now confined to eastern China, ~1alaysia, and 

Polynesia. Another very interesting fern is Saccoloma, with about· eight living species in tropical 

America and the ",Vest Indies and in ~1alaysia and 

Polynesia. The Denver species is clearly referable to 

this genus and is apparently closest to S. elegans I(aulfuss of tropical America. A third interesting form 

is that descr.ibed under the name Salpicl~laena anceps. 

,"!"'he genus SallJichlaena, now by some pteridologists 

merged with Bleclwnt1n, has a single tropical American species. It is a climbing fern, and the Denver 

species was also of similar habit. Still another interesting Denver fOrIn is a species of Diplaziu1n, the 

numerous living species of which are widely spread 

01 Knowlton, F. H., U. S. Geo!. Sl1l'vey Prof. Paper 101, pp. 233-234, 

1017 [1918]. . 

over the tropical portions of both the Old and New 

",Vorlds. In addition to those above mentioned there 

is a small, delicate species of Dennstaedtia and a large, 

robust species of W oodW'a1·dia. The family Schizaea- . 

ceae is represented by three rather inadequately characterized forms of Anemia and a large species of 

LygodiuYln. Although both these genera include many 

tropical or subtropical living species, there are some 

species found in warm temperate regions. Taken altogether, the ferns of the Denver and associated formations would seem to have required a considerable degree of heat for their successful growth, perhaps as 

warm as subtropical. 

The Equisetaceae are represented by three very 

poorly characterized species, and, SeZaqinella by a 

single well-defined form, but they can have very· little 

bearing on the climatic conditions in Denver time. 

Tile conifers are practically a negligible fa~tor in 

this flora, being represented only by a single species 

of Sequoia, and this only by two or three poorly preserved branchlets. The Ginkgoaceae are represented 

by a somewhat questionable fruit. No leaves have 

. been found, and it is quite possible that the fruit has 

been incorrectly placed. 

The monocotyledons are very poorly represented; . 

in fact, with the excepti()n of the palms, only tlll:ee 

forms are recognized-a poorly preserved grass 

(A.1'undo) , a fragmentary Oarew, and a very questionable plant referred to Zingibe1'ites. They are of little 

value in the present connection. 

The palms, however, are a very conspicuous element, 

comprising 14 noniinal forms, 8 of which are based on 

leaves, 5 on fruits, and 1 on a perfectly preserved 

silicified trunk. The leaves are fairly abundant, especially in the Denver beds, and some of them were of 

large size, perhaps ~'eaching a dianleter of 5 feet or 

more., The fruits, except two or three, are not so 

certainly placed. The trunk is very perfect. Although some living palms can withstand a few degrees 

of frost, the vast majority require a temperature well 

above 42°, and Inost of them would be killed at this 

temperature. It seems reasonable to assume, therefore, that the palms of Denver time, being of diverse 

types, could hardly have tolerated a temperature 

cooler than very warm temperate, possibly even subtropical. It is of interest in this connection to recall 

that palms were especially abundant in· the Raton 

formation, where six genera and nine species are 

recognized. Sabal ungeri is probably the nlost abundant and widely distributed species in the Raton formation, occurring at" most of the localities and at many 

of them in great numbers. 

The amentiferous trees were apparently abundant, 

~here being 6 species of J'Mglans, 1 each of Ilico1'ia and 
12 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Pte1'ocarrya, 2 of Alnus, 3 of Betula, 1 of Oastanea, 

8 of Quercus, and 15 species and varieties of Poptblus. 

It is somewhat difficult to interpret the climatic requirements of this assemblage. Although most of the 

genera are most abundant in cool temperate regions 

at the present time, nearly all have representatives 

in the warmer regions of the earth, and considering 

the apparent requirelnents of the ferns and palms it is 

reasonable to assume that the amentiferous trees should 

ha v~ required similar surroundings. 

The Ulmaceae are represented by two species of 

Ul17""us, but they were very rare and none too well 

allocated. 

The figs and breadfruit trees were especially abun- . 

dant during Denver time, 23 forms of FiC1tS and 3 

species of A1'tocarpus being recognized. They undoubtedly argue for a wann climate. 

The sycamores (PlaJtanrus) were also an abundant 

element in this flora, as no less than 10 forms have been 

recognized, though it is possible that too nlany have 

been admitted. Many of them are large-leaved forms, 

and some are poorly preserved; in fact, they give evidence of having been transported for greater distances than many other types. However, if too many 

forms have been recognized, it will be easy enough to 

combine them. 

The Lauraceae did not form a very prominent feature, there being three species of La~(;]'1lS, one of Pe'1'-

sea, and four of Oinna17wlnuln, two of which, however, 

are not named specifically. The laurels and their kin 

are plants requiring a moist, warm habitat. 

The magnolias in the Raton flora were varied and, 

in some forms, extremely abundant, but in the Denver 

flora only a single species has been noted, and this 

was not abundant. 

The Nymphaeaceae were well developed in the Denver and associated beds, there being three genera and 

five species. Of tllese the three species of N elWlnbo are 

especially well marked. Allied to this, but abundantly distinct, is Paleonel1f11nbo, a new type with 

very large, strongly toothed leaves. The lnost interesting is a splendid species of water lily (Oastalia) , 

,vhich is most closely related to a species now living in 

tropical America and the 'Vest Indies. 

Passing over a number of rosaceous and papilionaceous genera (Orataegus, Pnm1JJs, Phaseolites, Sopho1'a, 

Oa8SW" and Oe1'cis) that are without strong climatic 

characters, we come to th~ Celastraceae, with two species of 0 elastrinites and one of 0 elastrus'i the Aceraceae, with a single form each of A(('er and N egundo; 

the Sapindaceae, with three species of Sapind1ls i and 

the Ilicaceae, with a single somewhat doubtful species 

of flew. These are mainly plants of warm regIOns, 

though with representatives in cooler lands. 

The buckthorn family (Rhamnaceae) is strongly de-

,"eloped, having six species each of R ha1nn1t8 and ZizypNuS and one each of Be1'che17~ia and Paliu1"U8. The 

present-day species are mainly of tmnperate.or warnl 

temperate distribution. 

The Vitaceae are represented by eight species of 

OissufS and one of Vitis, the Tiliaceae by a single species. 

of Gre1()riopsis, the ~terculiaceae by three species of 

Ste1'c1JJlia and one of Pterosper11'dtes, and the Cornaceae· 

by four species of 00'r111tS and three of lVys8a. 

If the apparent climatic requirements of the Denver 

and related floras are viewed as a whole, it appears 

from the above review that lnany of the· important 

elements, such as the ferns, palIns, figs, breadfruit 

trees, and laurels, argue for warm, possibly subtropical 

conditions. The other elements almost without exception seenl to call for a t least a wann or wann 

temperate environment; in fact, with minor exceptions 

there is not a type whose living representatives are exclusively temperate. Although a number, such as· 

llly'rica, Magnolia, Oorn'us, and flew, are now dominantly temperate in their distribution, they all have 

forms or closely related genera that extend into the 

Tropics. Therefore it seems safe to conclude that the 

local climate during Denver time was abundantly 

moist and warm temperate or possibly subtropical. 

As the Denver and related floras are compared with 

those of the Raton, 'Vilcox, and Laramie it mny be 

of interest to set forth the interpretation given to the 

apparent clilnatic requirements of these other floras. 

Concerning the Raton flora I concluded 32 as follows: 

The presence of the numerous coal veins as well as the· 

character and luxuriance of the vegetation indicates that 

moisture was abuudant, and the known distrilmtion of the· 

living representatives of the Raton flora make it more than 

probable that the climate was at least warm temperate. 

The following conclusion was drawn as to the c1imate during Laramie time: 33 

From the abundant presence of coal and the apparent requirements of the majority of the plants enumerated, it is. 

beyond question that there must have been abundant moisture. 

It also appears naturally to follow from the presumed requirements of the flora that the climate was warm, at least warm. 

temperate. 

The 'Vilcox flora, is believed by Berry to be slightly 

younger than that of the Raton and Denver floras .. 

It is distinctly a strand flora. Concerning it Berry 34 

,n'ote as follows: 

It may be noted that the 'Wilcox plants, almost without ex-· 

ception, are plants whose modern representatives inhabit the 

warmer parts of the earth. There is not a strictly temperate' 

type in the whole assembly. « '" * It needs but a slight 

acquaintance with the existing Antillean flora or that of the· 

Florida Keys « '" .;. to see at once that the general facies. 

82 Knowlton, F. R., U. S. Geol. Survey Prof. Paper 101, p. 240, 1917 ~ 

~~ Knowlton, F. H., U. S. Geol. Survey Prof. Paper 130. p. 99, 1922. 

M Berry, E. W., U. S. Ceo}. Survey Prof. Paper 91, pp. 136, 137, 1916_ 
THE FLOBA 13 

of the "Wilcox flora is overwhelmingly that of a strand flora, 

some of the elements of which indicate that they grew on 

sandy lJeaches, others in muddy tidal flats, others between 01' 

behind dunes 01' beach ridges, and others in estuary bayous 

or marshes. 

PHYSICAL CONDITIONS 

The probable physical conditions under which the 

Arapahoe, Denver, Dawson, and Middle Park formatlons were deposited may be briefly "considered. They 

. are described by Richardson 35 as follows: 

Uplift of the Jj .... ront Range "region and accompanying erosion, 

afte1' the deposition of the LaramiE' beds, marked the close of 

the Mesozoic and the beginning of the Cenozoic era. A great 

thicklless of Cretaceous and 01de1' rocks was eroded from the 

Jj .... 1·ont Hange region, and a mass of continental deposits accumulated on an uneven surface of the Laramie and older 

formations etlSt of the mountains. The Dawson arkose, derived from the Pikes Peak granite and associated rockS, was 

laid down ullder various continental conditions, chiefly as 

wHsh and fluviatile deposits accompanied by local ponding. 

During the accumulation of the arkose this region may be 

conceived of as a piedmont area having a moist and temperate 

climate. " 

As already mentioned, the lower portion of the 

Dawson arkose appears to interdigitate on the north 

with the Arapahoe and Denver formations. If this 

interpretation is correct, as it is now believed to be, 

t.he Arapahoe and Denver are equivalent in time with 

the lo"\ver portion of the Dawson arkose. Although 

they differ markedly in lithologic composition, the 

physical conditions under which they were deposited 

~ppear to have been the same-that is to say, the uplIft of the Front Range area started extensive erosion. 

and resulting debris was spread out over the adjacent 

plain. 

The streams coming down from the highlands to 

the west became sluggish out on the plains and were 

impounded here and there into ponds and small lakes. 

The material brought down was spread out over the 

plains or deposited in the lakes and ponds. In the 

northern part of the Denver area and in l\1iddle Park 

the material supplied by erosion was largely of volcanic origin, hence the Denver and Middle Park beds 

are composed mainly of andesitic materials; but to 

the south the streams were cutting granitic rocks, and, 

the resulting deposits are mainly arkosic. 

On the highest ridges there were doubtless conifers, 

but they appeal' to have been rare, as only a few 

branchlets of a single species have been detected. On 

the lower slop~s probably grew the oaks, black walnuts, hickory nuts, and birches, with an occasional 

magnolia, chestnut, and maple. Along the streams 

were the alders, sycamores, and certain poplars and 

osiers. In the more open places were the viburnums, 

311 Hichal"(lson, G. B., U. S. Geol. SUI"vey Ceo\. Atlns, Castle Hock 

fOlio (No. 198), p. 12, 1915. 

buckthorns, dogwoods, and hawthorns, with vines 

(V itis) and bittersweet climbing over trees or other 

lower supports. On the low grounds were elms, sycamores, and poplars, and in the swamps was the usual 

tangle to be expected in such situations-the figs~ 

sweet gums, cinnamons, magnificent palms with trunks 

nearly 2 feet in diameter and lea,ves 4 or 5 feet across, 

and soapberries-and underneath were Zizyphu8, Ois8US, PaliU1"U8, and climbing ferns. In shallow water 

and about the margins of ponds were tufts of the tall 

grass Arundo and two apparently large equisetums, 

with probably. smaller grasses and sedges ( 0 (l1'ex) 

near by: On the surface of the ponds and on the slowmoving streams floated the leaves of beautiful ''lhite 

and yellow water lilies. 

In certain parts of the area swamp conditions continued with little variation for long periods, with the 

result t.hat considerable beds of coal were accumulated. 

So far as I have been able to interpret the facts, the 

a vnilable evidence indicates that this coal was not 

formed in any considerable part from transported material but rather was formed in place. 

MATERIALS STUDIED 

The material that has been available for the present 

study is very considerable. This study is based primarily on the early collections Inade by F. V. Hayden 

Hnd other members of the United States GeoloO"ical o " 

Survey of the Territories, mainly in the early seventies, and now preserved in the United States National 

~1useum. Lesquereux and some of his correspondents 

also made collections, and all were identified and described by Lesquereux and the results published from 

time to time in the annual reports of the Hayden survey. In 1878 the results then available were brought 

together by Lesquereux. 3G Up t.o that time the plantbearing beds of this genenll region were supposed to 

be similar in aue and to belonO" to the "LiO"nite '-' 0 0 

I"rertiary," but later, when t.he Laramie formation 

was established, and still later, when the Arapahoe 

and Denver were separated as distinct units. considerable confusion resulted in the collections, as 'they were 

usually without specific locality labels. This is especjally true of the locality" Golden, Colo.," where b~th 

Laramie and Denver are present and plant-bearing. 

Fortunately, however, most of the plants from Golden 

('an b~ distinguished by the matrix, that of the Laramie being usually a white fine-grained sandstone and 

that of the "Denver a yellowish andesitic material. 

In 1883 Lesquereux 37 published a short "Description of species added to the flora of the Laramie 

sa Lesquereux, Leo, The Tertiary flora: U. S. Geol. Survey Terr. 

Uept., vol. 7, 1878. 

37 Lesquereux, Leo, The Cretaceous and Tertiary floras: U. S. Geol. 

Survey Terr. Rept., vol. 8, pp. 121-126, 1883. 
14 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

group," enumerating 11 species, 6 of which were new . 

.J\1:ost of these specimens are now in the museum of 

Princeton UniveI:sity, where I saw and studied them 

some years ago. 

Arthur Lakes, a mining engineer, made a large colkction of Denver plants, mainly from South Table 

Mountain, Golden, that became the property of the 

M:useum of Comparative Zoology, Cambridge, Mass. 

They were worked up by Lesquereux,38 who stated 

that the 873 specimens represented 118 species, 'of 

which 28 were described as new and 32 were recognized for the first time, thus adding 60 species to this 

flora. None of this material has ever been figured, and 

by the courtesy of Dr. Robert T. Jackson, in whose 

custody these plants no,Y are, I have been permitted to 

borrow these types and unfigured specin1ens, and they 

H re here figured for the first time. It has been necessary to revise the no;nenclature somewhat, and a few 

of the forms supposed to be new by Lesquereux have 

been combined with older species or otherwise disposed of. 

In 1890 Lakes was employed by S. F. Emmons to 

make collections of plants from the Arapahoe and 

Denver formations for the United States Geological 

Survey. These collections came mainly from South 

Table Mountain, Qolden, and from the base of Green 

:l\1ountain, in the Denver formation, and included also 

the collections from the equivalent of the Arapahoe in 

the vicinity of the old Douglas coal mine, near Sedalia, 

which are listed as Arapahoe plants. 

Individual specimens or small but interesting and 

valuable collections have been submitted by many persons, among them 'Vhitman Cross, L. F. 'Vard, Junius 

Henderson, Theo. D. A. Cockerell,A. C. Peale, and 

. 'V. T. Lee. These are now maInly in the United States 

National Museum. 

Early collections of plants from the Dawson arkose 

'~vere made in 1896 by T. W. Stanton and F. H. Knowlton in the Colorado Springs area, though they were 

not recognized at the time as distinct from the Laramie. Small collections in the same region were also 

made by M. 1. Goldman and A. C. Peale in 1909, but 

the bulk of the collections from the formation were 

made by G. B. Richardson or by members of the party 

l:nder his direction while they were obtaining data for 

the Castle Rock folio. I also visited the area in 1910 

and in company with Mr. Richardson made collections east of Coloi'ado Springs; at Templeton' Gap, 

near Calhan; on Jimmy Camp Creek; at ~10sby; and 

elsewhere. Small collections were also made in this 

area in 1912 by W. T. Lee. 

The plants available from ~1iddle Park include most 

of the original collection made by F. V. Hayden in 

as Lesquereux, Leo, Harvard CoIl. Mus. Compo Zoology Bull., Yol. 16, 

pp. 43-59, 1888. 

1869, now preserved in the United States National 

Museum, and the la.rger collections made by George L. 

Cannon in 1889, together with a few procured by 

Whitman Cross in 1891 and 1892. The collections 

from Middle Park are not as complete as could be 

wished, though some 24 forms have been recognized. 

The plants are known to be abundant at a number of 

horizons, but it must remain for future n10re exhaustive exploitation to develop knowledge of this flora. 

"Vork on the flora of the Denver formation was 

begun about 1895, and a nmuber of descriptions and 

figures were prepared at that time, but as this was a 

period when views were changing on the distribution 

and stratigraphic relations of the Laramie, with which 

the Denver is more or less intimately related, it was 

thought expedient to lay the manuscript aside for a 

time until views could be more stabilized. Pressure of 

other interests has resulted in this long delay, but the 

'wisdom of the delay is abundantly justified by the increased collections and stratigraphic data now avail-. 

able. The illustrations that were prepared when the 

work was started ,vere drawn by the method then current-that is, pen drawing two-thirds enlarged for 

one-half reduction, which restores them to natural 

size. The present method consists of taking photographs natural size and retouching them to bring out 

the essential characters. As illustrations of th~e two 

kinds can not be mixed on the same plate, it has been 

necessary to arrange the plates in two series. 

SYNONYMS AND CHANGES OF NAME 

In the following list synonyms and former names 

of species here described are given alphabetically, 

with their equivalents in the present ,work. 

Aralia: ptt]1,gens Lesquereux=Artocal'p'ltS p'ltngens. 

Artocarpifti'llm, olmedifoUum? Vnger, Lesquereux (V. S. Geoi. 

and Geog. Survey Terr. Ann. Rept. for 1873, pp. 381, 400, 

1874) =OeZastrinites artocal'pidioides. 

Aspldit~m lakesVi (Lesquereux) Knowlton=Drllopteds lakesU. 

Asplenvum erOStt1n (Lesquereux) Knowlton=AllantodiolJsis 

el'osa. 

Asplenium sltbsimplem (Lesquereux) Knowlton = Allantodiopsis 

erosa. 

Bel'che-mila multinervis AI. Braun, 'Yard (V, S. Geol. Survey 

Sixth Ann. Rept., p. 554, pl. 51, fig. 13, 1887; U. S. Geoi. 

Survey Bull. 37, p. 73, pl. 33, fig. 2, 1887) =Rhal1l'1111tS 

goZdiantts. 

Berchemia par'l)'ifolia Lesquereux=Berchemia, tnultinerms. 

Oarpolithes aracMoides Lesquereux=Legwninositesf arachioides. " 

CarpoZithes palmat'ttl1'll Lesquereux=Pall1WCal'pon palmarum~. 

Ceanoth'lts {ibrillos'lts Lesquereux=Zizllph1tS {ibrUlostts. 

O'iss1ts laevigata Lesquereux=Cissus coZol'a(lensis. 

GiSS1tS pa!rrottiaefolia Lesquereux (Harvard CoIl. Mus. Compo 

Zoology Bu]l., vol. 16, p. 52, 1888) =Cissus obovata. 

Oiss'lts spectabilis Heer, Lesquereux (Harvard CoIl. Mus. Compo 

Zoology Bull., vol. 16, p. 52, 1888) =Oh1'lIsobalan'lls colol'adensis. 
SYSTEMATIC PALEOBOTANY 15 

o iSS'ltS tric1,spidat(t (Heel') IJesquereux= O'iS~1tS lesq1tet'euxU. 

Oornus em!llwnsi "iard=Oornus im,p'I'essa. 

01·(tt(teg'ltS betula..efoZia, Lesquereux (Harvard CoIl. Mus. Compo 

Zoology Bull., vol. 16, p. 56, 1888) =Viburnmn 1'iclulIrdsoni. 

J)aphnogene angl'io(£? Heel', Lesquereux (Tertiary flora, p. 222, 

pI. 37, fig. 9, 1878) =Z'lzyphtts daphnouenoides. 

Dil)/az'i1mt 1/1J1teUeriY Heel', Lesqu('reux (U. S. Geol. and "Geog. 

Survey Terr. ,Ann. Hept. for 1873, p. 3'93, 1874) =D'iplaz'i1l1n O1'oss'i:i. 

J)r!l0lJtoris (l,1·U'I.tfa (Lesquereux) Knowltou=DryolJteris lakes'ii. 

ji'labeU(wla commmnis Lesquereux=SabaUtos Ura,YM/,1tS. 

ji'/o belZOIria eooenica Lesquereux= S£t-b alit es g1'ayanus. 

Plabe11aria? f1'1t.CNfer Lesquereux=Sabalites g1·(tYM1IUS. 

.li'labcll£wia? longirach'is? Unger, Lesquereux (U. S. Geol. aud 

Geog. Survey Terr. Ann Rept. for 1873, p. 396, 1874) =.= 

G-eo'llmnUes tenu'iraohis. 

J;'la,bellaria Z'inlceni Heel', Lesquereux (Tertiary flora, p. 110, 

pl. 9, figs. 6-8, 1878) = Geono'm'ites g01d'ia1ws. 

Oconom,ites 7tllgeri IJesquereux=Sabal? 1tngeri. 

U/IJptost1'ob1tS 'ltngeri? Heel', I.1esquereux (Cretaceous and Terti:H,Y floras, p. 139, pI. 23, figs. 1-5a, 1883) =Sequ,o'ia 

af/inis. 

GoniOl)te1'is lJOlypod'iohloS. Ettingshausen, Lesquereux (U. S. 

Oeol. find Geog. Survey Terr. Ann. Rept. for 1873, p. 394, 

1874) =DI'IJ01)toris 1)o1Y1Jo(l'ioides. 

GY1nllog1'anl1na g(wilncri (I.1esquereux) Lesquereux=Sacooloma, 

ua'l'dner'b. 

La8tl'ea, (G-Ol'l:iOl)toris) ' polypodioides? (Ettingshausen) Lesquereux= D'I'yopteri s l)OlYl)od'ioides. 

'Lulhraea (l1'Ull,fa I,.jesquel'eux=D1'yoptm'is lakes'ii. 

jjauJ'lts bross'iana I.1esquereux=Persea b1·ossiana. 

jjulII'us ob(YVata? Weber, Lesquereux (U. S. Geol. and Geog. 

SUl'vey Terr. Ann. Hept. for 1872, p. 399, 1873) =Lall,r'tlrs 

,pI'1m'igcnia. 

il[a.{/IWlict tenwlnervis I.1esquereux (Tertiary flora, p. 248, pl. 44, 

fig. 2, 1878) =lI[aglloU(t- magnifol'i(t. 

Nclllmbi,ltm lalccsianwn Lesquereux=Nelmnbo lal~cs'iana. 

Nolmnbi1t'ln lal~GsU Lesquereux=Nelu.mbo l£tliics'iana. 

Nel'll1nb'i,lt1l'/, te'n4t'ifolhtmb Lesquereux=Nel'ttmbo telwifoUa. 

Nyssa? raoenwsa Kllowlton=Ber1'ya 1·aoel1wsa. 

Ore'odowi.tes 1)l'ioatus I.1esquereux=Palo/'codoxites pUcatus. 

OiHlloltndn a.f/in'is (Lesquereux) =Salpio1l,laena, anCel)s. 

OSIIl.1t1ul(t nw,jol' Lesquel'eux=A.llantod'iopsis e1'osa. 

l lall1woitcs gold1an1tS Lesquereux=G-eolw1nUes golwian1ts. 

P(tlmoO(l1'lJOn commune Lesquereux=P(tlmocarpon palmar'ltm. 

Pa,1,tnoom'JJOn tnmoatmlb mbnOl' Lesquereux=Ginl~go? trttncata. 

Pan'Mi(t fagifoUa. Ettingshause'll, Lesquereux (Hal'vard ColI. 

Mus. Compo Zoolo~y Bull., vol. 16, p. 52, 1888) =Oelast1'1tS 

Ua1twini. 

Phy.'1ugenJia sp. I,.jesquel'eux (Harva'rd Coll. Mus. Compo 

Zoology Bull.. Yol. 16, p. 43, 1888) =}1]q'ltiset1l1n' sp. 

POPIl,f,US 1tnge'ri Lesquereux, Lesquereux (Harvard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 46, 1888) =Oelastrin!ites 

llU1HlrlifoU,lts. 

p.~·OtOtiOllS zeillol"t I,.jesquereux=Pop1lrl7ts zeilleri (Lesquereux) 

Knowlton. 

Ptcris anceps Lesquereux=Sall)io1l,1acna anoeps. 

Ptcris erosa Lesquel'eux=AllantotZlops';'s erosa. 

Pt,cris uarwll,eri Lesquel'eux=Saocoloma g(l1'dnel"i. 

ptcris l)cnnacfornl'is Heel', Lesquereux (U. S. Geol. and Geog. 

Survey 'ferro Ann. Hept. for 1870, p. 384', 1871)=Ptm'is 

1)SC1Ulol)C'/MWcfo1"l1l i8. 

Plol'is 1>se1td01>ennaeformis Lesquereux (Harvard ColI. Mus. 

Compo Zoology BulL, yol. 16, p. 43, 1888) =SallJichlaena 

anaeps. 

Pte1'is subsi117;plex Lesquereux=A.llantodiopsis er'osa. 

Pte1'is 1tnd1lrlata Lesquereux = A.llantod1·ol)sis el'osa. 

Rhamnus crenat1tS Lesquereux=Rlw.mnus cren7tlat1tS. 

Sabal oampbelli Newberry [part] =Sabal? 1tnge1'i. 

Sa.bal oommunis Lesquereux=Sabalitcs {fl'ayan1ts. 

Sabalf eocen:ica (Lesquereux) Knowlton=Sabalites graya'll1ts. 

Sa.ba,l grandi.foU,a Newberry=Sabal? 1t.nge'l"i. 

Saba1? grayana Lesquereux=Sabalites grayamLS. 

Sa.baZ ~inq1tirenda Knowlton=Sabalites grayanus. 

Sa,balUcs fnwUe1' Lesquereux [part] =Be1'rya 1'aoe11wsa. 

S(~ba.lite8 f1'1totife'r Lesquereux [part] =Sabalites grOllJa1ws. 

Salix media? Heel', Knowlton (Geol. Soc. America Bull., vol. 

8, p. 141, 1897) =L(~1wtLS l)1'imigen,ia.. 

Sclerotium 1'1tbellum Lesquereux=Sclerotites 1"/.t,bel17ts (Lesq uereux) Meschillelli. 

Seq1toia la1l.gsdorfii Bronglliart, Lesquereux (U. S. Geol. and 

Geog. Survey Terr. Ann. Rept. for 1874, p. 310, 1875; , 

Tertiary flora, p. 75, 1878) =Sequoia affi,nis. 

SphelWl)tm'is eooenio£li Ettingshausen, Lesquereux=Dryopteris 

lakes'i'f.. 

Sl)henoptel"is laliiesii Lesquereux=Dryopte1"i,s lalcesi-i. 

Sphenopteris membranaoea Lesquereux=Dryollteris lakes'iI. 

Ste1'OltUa modesta Sapo1'ta, Lesquereux (Cretaceous and Tertiary floras, p. 125, pl. 20, fig. 5, 1883) = SterettUa sapo/,-

tanea. 

V,lbtlrrnulI1t anoeps Lesquereux (Tertiary flora, p. 227, pl. 38, 

fig. 11, 1878) = Platan1ts ?'1wmboi(lea. ' 

Vib1tr1W111J d'iohot011t1tm Lesquereux=Vib7tl'nmn melaenwn. 

Vibw'1Htm mm'gina,tu,m Lesquereux=Platantts 111,arginata. 

V'ib1t1'1W,m maruifnat'ltm Lesquereux (Tertiary flo'ra, p. 223, pl. 

38, fig. 2 [not fig 3], 1878) =Vib'urnu1n conto1'tmn. 

Vib11r1'nwn platanoides Lesquereux=Platanus platanoides. 

Vib1trlwm whymperif Beer, Lesquereux (Tertiary flora, p. 225, 

pI. 38, fig. 7, 1878) =V'ibu1'nllm 1nelaenttm. 

Woowlwrdia latUoba minor Lesquereux=Wooawm'dia la.tUoba. 

Z'ing'iberites? 1Ir1td1tlatus Lesquereux= Z'ingiberites dubi7tS. 

Z1zyphus d'istortus Lesquereux (Tertiary flora, p., 275, pI. 51, 

fig. 7 [not fig. 8], 1878) =Z1zyphtts fibrWos'ltS. 

ZizYl)h1tS fi,brillostLs Lesquereux (Tertiary flora, p. 276, pI. 52, 

fig. 6 [not figs. 1-5], 1878) =Zizyphus £laphnogenoitl,cs. 

Zizyp1l,us hYIJerbo1'eu,s Heel', Lesqueeeux (Haryard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 55, 1888) =ZizYIJh'tts hesZizYl)]WS 1I,yperb01'e'lts? Heel', Lesquereux (Tertiary flora, 

1). 2,76, pl. 51, fig. 15, 1878) =Zizyp7l:ltS lcsq1lr01'e1txi-i. 

SYSTEMATIC PALEOBOTANY 

Phylum THALLOPHYTA 

Class FUNGI 

Sclerotites rubellus (Lesquereux) Meschinelli 

Solcrotites r1tbellus (Lesquereux) l\:1eschinelli, Sylloge fungorum fossile, p. 69, 1892; Fungorum fossilium omnium, 

Icollcgrapllia, p. 101, pI. 27, figs. 12, 13 a-f, 1898. 

Sclerot'tum 1'1tbelZ7tnt Lesquereux, U. S. Geol. and Geog. SUrYey 

Terr. Ann. Rept. for 1872, p. 375, 1873; idem for 1873, 

p. 370, 1874; idem for 1876, p. 496, 1878; Tertiary flora, 

p. 35, pI. 1, figs. 2-2 f, 1878. 

The type'specimen of this species is preserved in the 

United States National Museum (No.2), as are also 

other specimens of the so-called Flabellaria zin7ceni? 

(Nos. 101, 102) upon which it occurs, This is by far 
16 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

the best-marked fungus obtained in these formations 

and is in all probability correctly referred. The perithecia are oval 01' linear, obtuse, 1 millimeter broad, 

2 to 4 millimeters long, and are placed between the 

striae or minute veins of the leaf. 

The species has not been procured since the original 

collection. 

Occurrence: Denver formation, Golden, Colo., on 

leaves of Flabellaria. 

Sclerotites? cypericola Knowlton, n. sp. 

PIa te 1, Figure 7 

In one of the collections from the Dawson arkose, I 

find the single speci~en of a fungus here figured. It 

grew on a finely striated monocotyledonous leaf about 

a centimeter in width, that suggests a large-leaved 

cyperaceous plant or possibly a segment of a palni 

leaf. The fungus is disposed in two Cor three longitudinal rows down the center of the leaf. The pushlles or perithecia, if they should be so called, are 

more or less granular in appearance; some are nearly 

rectangular and others round or even kidney-shaped. 

There is no evidence of an opening in the top of any 

of these perithecia nor of the contents other than a 

black, carbonaceous mass. 

This description is confessedly incomplete and more 

or less unsatisfactory, and therefore the generic reference. is uncertain. The species is placed in Sulerotites because it seems to be like certain forms that have 

been so placed, but this· is little assurance that the 

assignment is absolutely correct. It is presented 

mainly to direct attention to the fact that a plant of 

this character was present at this time. 

Occurrence: Dawson arkose, e9.st bank of Jimmy 

Camp Creek, 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. Knowlton, July, 1910. 

Sclerotites? hesperius Knowlton, n. sp. 

Plate 1, Figure 6 

One of the collections obtained near Ramah, Colo., 

contains a number of specilnens showing an abundance 

of saprophytic fungi. The material is a red, hardbaked shale showing nilmerous fragments of dicotyledonous leaves and a number of stems or decorticated branches that are lnore or less covered with 

black carbonaceous pustules. These are inainlv circular and of all sizes from a pin point up to a millimeter 

or n10re in diameter. The larger ones show distinctly 

a minute osteole in the center. ,\Vhere a pustule or 

" peri theci urn " has been removed there is a circular 

tlepression like the imprint of a small shot, indicating 

apparently where a portion of the substance has been 

removed or absorbed. 

The nature of the host on which this fungus was 

growing can not be detennined with certainty, but it 

appears to be a lnonocotyledon of some kind, perhaps 

a cyperaceous plant or possibly a larger grass. 

The propel' ~ystematic assignment for this fungus 

is difficult to ascertain in the absence of certain essential c!laracters. It is therefore referred provisionally 

to the genus Scle'rotites. It can be compared with 

Sphae?-ites secretani (Heel') ~t[eschinelli,30 from the 

Swiss ~1:iocene and found in Austria on leaves of 

P lwrag171:ites eoningensi8 Heel'. 

This form is certainly very distinct frOIn the last 

described species and indeed n1ay not even be con· 

generic with it. 

Occurrence: Dawson arkose, about 4 miles south of 

Ramah oil well, 40 miles northeast of Colorado 

Springs, Colo. 

:Phylum BRYO:PHYTA 

Class HE:PATICAE 

Order MARCHANTIALES 

Family ldARCHANTIACEAE 

Marchantites? coloradensis Knowlton, n. sp. 

In the material from }\trosby, Colo., is found the single example figured, which appears to belong to the 

IIepaticae. It is a mere fragment that must give only 

a very poor idea of the appearance of the perfect plant 

and ordinarily would hardly be worthy of a description and name: yet, small as it is, it exhibits a number 

of interesting features and can undoubtedly be recognized in future. It js interpreted as representing the 

under surface of the vegetative portion or frond of a. 

hepatic. It is plainly dichotomous, consisting of a' ~ 

main proximad portion about 15 millimeters broad, 

which forks into two narrow distad portions th8,t 

are 9 or 10 millimeters broad and at least 3.5 centimeters long. The most perfect of the two branches is 

rounded at the end, as if it were completB at this 

point, but this is, of course, uncertain, as it comes to 

the very edge of the matrix. The entire stu'face is 

strongly reticulated, the rai~ed portions of the mesh 

being relatively thick and strong. The areas inclosed 

by the raised walls are somewhat irregular, though 

roughly they are quadrangular or oval. Many of 

them are now filled with coaly carbonaceous matter, 

thus proving that the frond had a considerable thick3lI Mescbinelli, Aloysius, Fungorlllll fossiliulll omnium, Iconogl'llpbill, 

p. 37, pl. 13, fig. 25, 1898. 
SYSTEMATIC PALEOBOTANY 17 

]leSS 01' substance. Scattered at irregular distances 

()ver the surface are strong depressions or pits, now 

filled with coaly substance, which are interpreted as 

the points whence rhizoids once arose. 

This specimen is preserved on a fine-grained sandy 

dn,y shale which retains the delicate structure vel'y 

well, and it is to be regretted thn,t so small a portion 

was preserved. 

In some ways the fossil form suggests the living 

.genus ill etzgm'ico, though the frond has no midrib, and 

the cells, if they nrc such, are .larger and much thicker 

\vaUed. 

Occurrence: Dawson arkose, dump of Mosby coal 

mine, ~1:osby, Colo., collected by F. H. l(nowlton and 

-G. B., Uichardson, July, 1910. 

Phylum PTERIDOPHYTA 

Class FILICES 

Order FILICALES 

Family HYMENOPHYLLACEAE 

Hymenophyllum confusum Lesquereux 

Plnte 1, ]j'igure 4 

H1JII/Cl/.oph'llllWIn' OOnflt81t1l'b IJcsquereux, U. S. Geol. and Geog. 

SUI.'\'ey ~l'el'1,·. Ann. Uept. for 1873, p. 39[i, 1874; The Tert'in l'Y Horn, p. [il, pI. 2, figs. 6--6a, 1878. 

Frond tl'iquadripinnate; rachis thin, flexuose; ultiInate pinnae brondly lanceolate; pinnules simple or 

cut into two or three divisions, cuneiform, enlarged 

upward from a decurring base, lobed, the lobes short, 

oblong, obtuse, simple, emarginate or bifid; veins 

strong, dichotomously forked, each branch entering, 

one of the p:innules. 

Th:is species was founded upon a mere fragment 

(U. S. Nat. ~1:us. No. 14) in which the frond is 

crusheil and distorted in a very confusing manner, 

and consequently its complete habit could not be nlade 

. out.. Since that time, however, additional material 

hns been collected at the type locality, and sevel'al of 

these specimens are fairly complete, so that it is now 

possible to ascertain the characters much lnore satisfactorily thnn from the type: The frond is probably 

tripinnate, and the pinnules are seen to be narrower 

than figured by Lesquereux. They nre not as regularly 2-pal.'ted nor turned back as described, but are 

cut into two to several divisions. 

Although it is of course impossible, in the absence 

0:1: fruit, to' be positive as to the correctness of the 

generic reference of any such fragments, there can be 

little doubt concerning these. They are, for example, . 

very much like the living H. f~u.rif01"1ne Swartz, of 

South America, or II. prrot1'o81WJn flooker, of Central 

Anlerica and the vVest Indies. Except fOl; the spinulose-dentate margins, the frond is quite similar to 

H. bivalva Swartz, of New Zealand, and in its branches 

it is suggestive of H. twnbridgen.se Swartz, of northern 

Europe. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo. 

Family. POL YPODIACEAE 

Dryopteris integra Knowlton, n. sp. 

Plate 1, ]j'igul'c 5 

Outline of '\vhole frond not known; apparently coriaceous, bipinnatifid; main rachis rather slender, round, 

not much ridged 01' striate; secondary rachis nearly as 

strong as the other and, of same character; pinnae 

very oblique, narrowly lanceolate, cut nearly or apparently quite to the secondary rachis, the segments 

or pinnules contiguous, oblong, acute at apex, attached 

by their whole base, entire-nlargined; middle vein 01' 

pinnules very strong, secondary nerves obsolete; sori 

large, orbicular or slightly reniform, foul' to six pairs 

on each pinnule, opposite, probably on the back of a 

veIn. 

This fern is quite unlike anything heretofore described from these beds and although obscurely preserved appears to be a reasonably distinct species. 

The specimen is a segment, probably from the lower 

portion of the frond, showing parts of foul' pinnae. It 

is an impression of the under surface of the frond and 

brings out the fruit very clearly, or would if the 

matrix were better fitted to show the details. The sori 

are very large, almost covering the segments, and are 

mostly orbicular, although several are apparently 

slightly kidney~shaped. There is also some induration 

of the impressions of large sporangia disposed in circles. A slight depression in the center of many sori 

seems to 'indicate that the indusiUlll was peitate, but 

t.his can not be' demonstrated with certainty. 

There seems little doubt that this species is correctly referred to D1:Jopteris. It is, for instance, quite 

similar to the living A8pidium 1"addWJnwm flooker (or 

A. quite'nse Christensen) from South America, though 

not quite so deeply cut. It is also strikingly similar' 

to certain fossil Aspidia from Greenland that have 

been described by Heel' under the names A8pidi'Ulln 

fC(YUAUlu17'1J,40 A. 1ney'el"i,41 etc. It differs from A. 1neyel"i 

in being bipinnatifid and in having the pinnules 

pointed rather than obtuse. Froni A. fe(JlJffU]WJn it diff<::rs markedly in size. 

Occurrence: ~1idcUe Park formation, ~1:iddle Park, 

Grand County, Colo., in sandy stratunl just above 

main breccia mass in Potato IEll Gap on the Grand 

40 Heer, Oswa.ld, Flora. fossilis arctica, vol. G, pt. 4, p. 32, pI. 29, 

figs. 5-9, 1882. 

41 Idem, vol. 2, p. 4Gl, pI. 39, figs. 1-3, 18GD. 
18 .FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

[Colorado] River near Hot Sulphur Springs, collected 

by George L. Cannon, 1889. 

Dryopteris Iakesii (Lesquereux) Knowlton, n. comb. 

Plate 1, Figures 1, 2 

La,thraea argu,ta Lesquereux,42 Am. Jour. ScL, 2d ser., vol. 45, 

p. 207, 1868; U. S. Geol. Survey Terr. Third Ann. Rept., 

p. 96, 1869; idem, reprint, p. 196, 1873. 

Sphenopteris eocen,ioa Ettingshausen. Lesquereux, U. S. Geol. 

and Geog. Survey Terr. Ann. Rept. for 1872, p. 376, 

1873; idem for 1873, p. 380, 1874. 

Sphenopteris lakesii Lesquereux, The Tertiary flora, p. 49, pI. 2, 

figs. 1, la, 1878. 

.As])idi1tm la7cesii (Lesquereux) Knowlton, Washington BioI. 

Soc. Proc., vol. 7, p. 154, 1892; U. S. GeoI. Survey Bull. 

, 105, p. 45, .PI. 6, figs. 1-4, 1893. 

Drllopteris arg1tta (Lesquereux) Knowlton, U. S. Geol. Survey 

Bull. 152, p. 91, 1898. 

Sphenopteris rnernbrarnaoea Lesquereux, U. S. GeoI. and Geog. 

Survey Terr. Ann. Rept. for 1873, p. 394, 1874; The 

Tertiary flora, p. 50, pI. 2, figs. 2-30" 1878. 

Fronds large, of firm texture, outline uncertain: 

apparently broadly triangular or deltoid, at least tripinnatifid, possibly some fronds fully tripinnate; primary rachis strong, half round, usually with one or 

two faint ridges; pinnae large, apparently broadly 

deltoid, at.an angle of about 45° to the rachis; secondary pinnae long, lanceolate, tap.er-pointed, connected 

along the secondary rachis by a narrow margin, provided with a large number of obliquely set segments 

or ultimate pinnules, which are lineal' or lanceolate 

in shape and have entir~ or more commonly strongly 

and sharply serrate margins; the pinnules are- close 

and united from well below the middle; veins pinnate, 

the divisions either simple or once forked; fruit dots 

large, round, in pairs on opposite sides of the lateral 

veins or in the upper part, in a single row on each 

side of the lnidvein of the pinnules and contiguous to 

it; indusium apparently peltate. 

This fine species is probably the most abundant and 

characteristic plant found in the Denver beds, being 

present in abundance in all collections in fragments 

. of greater or less size. Unfortunately, few large or 

complete specimens have been procured, but the type 

specimen as figured by Lesquereux is one of the largest 

thus far found. It is not possible to obtain a very 

definite estimate as to the full size of the frond, but 

as nearly as can he. made out certain of the larger 

ones must have had a spread of over 20 centimeters, 

and probably a still greater length. Some- of the secondary pinnae, isolated fragments of which are comnl0nly preserved, are fully 12 centimeters long in the 

lower part of the pinnae, but they dimil!ish rapidly 

42 As this is a nomen nudum and never properly described or figured, 

there is Eome uncertainty in associating it with the form under consideration. If authenticated it would be the oldest speCific name 

flpplicd to this species. 

in size toward the upper part. The largest of the secondary pinnae are between 2.5 and 3.5 centimeters in 

width at the base; all the pinnae terminate in a slender 

acuminate point. Curiously enough, out of the many 

hundreds of specinlens of greater or less size that have 

been collected, there appears to have been but one 

found in fruit. . This splendid specinlen, shown in 

Plate 1, Figure 2, appears' fr0111 the label it bears 

to have been collected in 1876 and exhibited at the 

Centennial Exposition at Philadelphia, after which 

it came to the United States National Museum, where 

it has remained in storage until very recently. Its collector is unknown, but there can be no doubt as to its 

authenticity, for the locality-Golden, Colo.-is plainly written on the label, and, moreover, the matrix is 

perfectly characteristic of this well-known locality. 

As may be noted from the figure, there are fragments 

of four secondary· pinnae, all of which are copiously 

fruiting. The lower pinna is a very broad one with 

the pinnules very deeply cut into triangular, acute 

teeth. The nervation of a pinnule consists of a very 

5t.rong midvein and a series of simple or rarely forked 

lateral branches which terl11inate in the teeth. ,Unfortunately, the frond is preserved with the upper SUI'-

:f~ce visible, so that it is difficult to make out the 

exact size and shape of the sori. They show through 

sufficiently well, however, to make it clear that they 

are disp.osed in pairs on each side of the lateral veins, 

or those running to the teeth of the pinnules, and there 

are usually two or exceptionally three pairs to each 

vein. In the upper pinnae as well as in the tips of the 

pinnae, where the pinnules are narrower and the mal'-

gimll teeth reduced, the fruit dots appear in a single 

row on each side of the Inidvein. All of them appear 

to be circular, and from the presence of a minute 

·dot in the center of each it is presumed that the indusium was peltate and that its stalk or support produces this central dot of organic l11atter. This evidence is not strong, however, and should be taken 

with some allowance. 

In 1893, in working over material from Montana 

collected by A. C. Peale, I found several small fragments of a fruiting fern that seemed to be identical 

with the Golden species.43 They came from a locality 

between the middle and north branches of Bear Creek, 

east of the Madison Valley, on the west side of the 

Madison.Range, in' beds supposed to be of Livingston 

age. These specimens represent the extreme distal 

. portions of secondary pinnae, the pinnules with small 

obscure teeth and the fruit dots very small and disposed in a row on each side of the midvein. Although these specimens were small and obscure, I felt 

justified in transferring the Golden species to the 

43 U. S. Geo1. Survey Bull. 105, p. 45, pI. 6, figs. 1-4, 1893. 
SYSTEMATIC PALEOBOTANY 19 

genus .Aspidium~ (now D'ryopte1"is) , and the fortunate 

finding of the above-mentioned fruiting example at 

the type locality proves this reference to have been 

well founded. So far as can be made out this fern 

belongs properly to the genus Dryopteris. The absence of fruit is what caused Lesquereux to place it 

first in the genus Lathraea and later in the impossible 

genus Splwnopte1·is. After a careful examination of 

Lesquereux's type specime~s preserved in the United 

States National Museum, together with more than 100 

additional specimens, collected by Arthur Lakes from 

a number of localities at Golden and Sedalia, Colo., I 

am entirely unable to distinguish more than one species. Extreme forms may be select~d which appeal' 

to be very unlike, but in the large series so many intermediate forms occur that it is impossible to draw any 

line of demarcation between them. Indeed, Lesquereux himself seems to have been in doubt as to 

their specific distinctness, for he says under S. 1nemb1'anaceat' " This form has nearly the same characters 

as the preceding (S. lalcesii) and may be a variety 

of it." That a plant so abundant as this must have 

been should show considerable variation is certainly 

to be expected from what we know of the variations 

existing in living species that have grown under varying conditions of environment. The pinnae and pinnules from different parts of a large compound frond 

vn.ry considerably in certain living species, being narrower in the upper part and quite unlike the lower 

ones. As many of the specimens of the fossil consist 

only of fragments of p~nnae it is impossible to tell 

from what part of the frond they came. 

The most complete specimen that I have seen is the 

specimen partly figured by Lesquereux 45 as the type 

of S. lakesii. The fragment is about 14 centimeters 

long and fully 18 centimeters broad and represents the 

1Iliddle portion of the frond, neither the apex nor t.he 

base being preserved. The primary pinnae appear to 

be broadly deltoid, with the pinnae numerous, close, 

linear, rather abruptly reduced in height in the upper 

portion. The attachment of the primary pinnae to 

the main rachis is not clearly shown, but they were 

evidently obliquely placed. The pinnules are slightly 

connected at the base and are sharply serrate or 

toothed. The nervation, which is incorrectly shown in 

Lesquereu4's figures, is strong, each vein forking once 

or twice or r~rely three times. '. 

As stated above, most of the specimens are fragments of pinn~e of greater or less size, probably from 

various parts of the frond. Some of them, as shown 

in Figure 1, have the pinnae remote but connected at 

base by a narrow decurrent wing. Some of the pin44 Lesqucr('llX, Leo, The Tertiary flora, p. 50, 1878. 

4B Idem, pI. 2, fig. 1. 

'" 

TIules, as also shown in Figure 2, are shorter and looser 

than in the typical forms. The nervation is invariably the same. 

The principal characters upon which Lesquerellx 

separated S. 1nembramacea from S. lakesii were the 

possession of narrower decurrent pinnules and single 

distinct nerves. An examination of the type speeimen/G as well as several others which are evidently 

like it, shows that the pinnae are very long and narrow and the pinnules smaller and Inore scattered than 

in the predominant forIn. It presents, as Lesquereux 

has said, quite a different facies from S. l(J)kesii. The 

statement that the nerves are single and unbranched 

is incorrect, however, as shown not only by the type 

specimen but still more clearly by better-preserved 

specimens from the same locality recently obtained by 

Lakes. These specimens, some of which are typical 

S. 1n.e17vbranacea and others merge into S. lakesii, all . 

have branched nervation precisely as in typical S. 

lakesii. 

The small specimen consisting of a single pinnllle 

which is figured on Plate 2, Figures 3, 3a, of Lesquereux's report on the Tertiary flora was first called by 

him Lath1'1G,ea a1'guta' but in the report mentioned is 

regarded as a form or variety intermediate between S. 

lakesVi and S. rnemb1'aJn(J)cea. The specimen as figured has apparently only one-half preserved, but it is 

fortunately contained in the National Museum collections, and an ,examination showed at once that the 

matrix on the seemingly destroyed half had not been 

removed. By carefully cutting it away the nearly 

complete half was exposed and showed, where fresh 

and unworn, the characteristic outline, toothing, and 

nervation of S. lakesii.. Several other specimens in 

the National Museum recorded by Lesquereux as S. 

'l7Le1nb'l'anacea prove, when carefully examined, to be 

all nearly or quite typical S. lakesii. 

I am therefore compelled to reduce these two species 

to one, which takes the name Dryopte'l'is lalcesii in 

honor of its discoverer. The most abundant form, 

which comes from the middle part of the frond, has 

the pinnae long with numerous contiguous, slightly 

connected pinnules. A less abundant form has shorter 

pinnae less thickly set with pinnules, which probably 

came from the lower portion of the frond. In still 

another form, S. 1nembranacea Lesquereux, the pinnae 

are remote, slender, lanceolate, with the pinnules narrower, plaited in appearance, and the nerves strong. 

Occurrence: Denver formation, Golden, Colo. 

(types), but without specific location. Later collections from Golden and vicinity are as follows: Green 

Mountain, quarry on northwestern base; same local46 Idem, pl. 2, fig. 2. 
20 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

ity but upper sealn or "fern ledge," about 20 feet 

above the first; South Table 1\10untain and about 100 

feet below the lava cap. Dawson arkose, lower part 

(in beds equivalent to Arapahoe formation), Sedalia, 

Colo., about 3,000 feet east of the old Douglas coal 

mine. The specimens thus far enumerated were collected by Arthur Lakes in 1890. 1\t[iddle Park formation, Middle Park, Grand County, Colo., from" doleritic breccia" just above main breccia mass in Potato 

Hill Gap on the Grand [Colorado] River, near Hot 

Sulphur Springs, collected by George L. Cannon, 

1889. 

Dryopteris richardsoniana Knowlton, n. sp. 

Plate 2, Figures 3-5 

Outline of whole frond not known; pinnae linearlanceolate, the rachis extremely strong, prominently 

ridged; pinnules or segments nearly at right angles to 

the rachis, close, parallel, linear, obtuse, cut nearly 

or quite to the rachis; midvein .of pinnules relatively 

strong, straight; lateral veins from 12 to about 18 

pairs" at an angle of 45° or 50°, close, parallel, simple 

or rarely forked near the Inidclle; fruit not known. 

This 'for~n is represented by about a dozen examples. 

As all are merely fragments of pinnae, it is impossible 

to ascertain the size and outline of the whole frond, 

though presumably it ,vas several times compounded. 

The largest fragment of a pinna is about 5 centimeters 

long, and neither base nor apex is shown. The width 

is about 2.25 centimeters. The pinnae are cut in the 

middle and upper parts nearly to the rachis into linear, 

rather obtuse segments. Near the basal portion of 

the pinnae the segments (or pinnules) are farther 

apart and appear to be nearly or quite free. The nervation consists of a relatively strong midvein and 

numerous pairs of simple or very rarely forked veins. 

This fonll was at one time identified provisionally 

with Dryopte'l'is lesqtteretW3ii (Lalf3tre(J) (G01ulopteris) 

goldiana Lesquereux), fr0111 the Denver beds near 

Golden. Although it is of the same type (congeneric) 

and might possibly be shown to be identical with 

that species if a larger series of both were available 

for comparison, as the matter now stands they seem to 

differ in a number of essential particulars. Thus, the 

specimens here described as DT!/Opteris richanison.iana 

are much larger than the Golden species, with the 

pinnae cut deeper to the rachis, the segments or pinnules more nearly at right angles, and the lateral veins 

nearly twice the number present in D. lesqu,eretlXii. 

The rachis is distinctly different in the two species. In 

D. lesquerelllixii it is much more slender, but. in D. 

'l'ichardsoniana it is exceptionally strong and very distinctly ridged. In any event, these two species are very 

o close and may later be shown to be identicaL 

Dryopteris 1"ichmrdsoniana is also very close to Dryopteris weedii I(nowlton,47 from rocks of Fort Union 

E()cene age in the Yellowstone National Park; in fact, 

it is perhaps doubtful if they can be satisfactorily 

separated. The Yellowstone Park form appears to 

have the segments or pinnules narrower and more scattered and the lateral veins at a slightly lower angle, 

but these differences are slight, and if a' series of 

well-preserved specimens were at hand they might 

disappear. As D. wleerJii was described attention was 

called to its undoubted affinity with D. lesqueretlw.ii 

from Golden. I 

I have named this species in honor of George B. 

Richardson, of the United States Geological Survey, 

'w'ho has done so much to elucidate the geology of the 

region whence it came. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Arapahoe formation), 3,000 feet east of 

the old Douglas coal mine, Sedalia, Colo., collected by 

Arthur L~kes, June, 1890. 

Dryopteris polypodioides? (Ettingshausen) Knowlton 

Gonio'ptfJl'is polypodioides Ettinghausen. Lesquereux, U. S. 

Geol. Survey Terr. Ann. Rept. for 1873, p. 394, 1874. 

Last1'ea (GonAopteris) polypoclioi.d6s'l (Ettingshausen) LesqU81'eUX, The Tertiary flora, p. 57, pI. 4, figs. 11, 12, 

1878. 

DJ'lJoptcri-s (Lastrea) 1Jolypodioicles (Ettingshausen) Knowlton, U. S. Geol.' Survey Bull. 152, p. D3, 1898. 

The two examples upon which is predicated the 

presence in this country of the well-known European 

species are preserved in the United States N ationaJ. 

l\1useum (Nos. 24 and 25). As Lesquereux very well 

said in discussing them, "The f.ragnlents figured 

* * * are to'o obscure for positive identification." 

They are preserved on a soft fine-grained sandstone 

which has been so much worn that it is impossible to 

]nake out the margin with certainty in a single place. 

The largest specinlen (Tertiary flora, pI. 4, fig'. 11) 

is about 8 centimeters long and nearly 2.5 centimeters 

wide. There is some evidence to suggest that the 

margin is toothed, but this point can not be verified. 

Concerning the nerva.tion Lesquereux said: "Primary 

veins equidistant, parallel; lateral veins at an acute 

angle of divergence, apparently alternate, simple, and 

cUJ."ved inward." 

As nea.rly as I can make out, the nervation given 

in Lesquereux's figures is almost totally wrong. The 

"primary veins," as he called them, are apparently 

the midveins of pinnules, which probably were free 

at the apices. The" lateral veins" are not simple but 

are thin, at an acute angle of divergence, and at least 

d7 Knowlton, F. H., Fossil flora of the Yellowstone National Park: 

U. S. Geol. Survey Mon. 32, pt. 2, p. 669, pI. 80, fig. 8, 1899. 
SYSTEMATIC PALEOBOTANY 21 

once forked. The whole nervation, as well as can be 

made out, is similar to or practically identical with 

that shown in the fragment from Middle Park shown 

on Plate 8, Figure 7, under the name Ane'l71"ia sp. 

Fr01n the foregoing statenlent it is apparent that 

there is only a remote possibility that these Sand 

Creek specimens can have any relation whatever with 

the European species described by Ettingshausen. 

They should probably be given a new name, but their 

characters are so obscure that it is doubtful if they 

could be so described as to permit their subsequent 

recognition, and under these circumstances it is perhaps as well to leave them as established by Lesquereux. 

I have, however, placed a question niark after the 

species name to indicate its extremely unsatisfactory 

state. 

The locality whence these specinlens came (Sand 

Creek east of Denver) has been involved in much confusion and uncertainty. They were supposed to be of 

either Laramie or Arapahoe age, but the latest information appears to place their horizon in the Denver 

formation. 

Occurrence: Denver formation, Sand Creek, about 

15 miles east of Denver, Colo., said to have been collected by )Villiam H. Holmes in 1873. 

Dennstaedtia crossiana Knowlton, n. sp. 

Plate 2, Figures 7-9 

Frond bipinnate; pinnules alternate, very oblique, 

tl'iangular-Ianceolate, sessile, and decurrent on a 

slightly winged rachis, deeply pinnatifid, the segments oblong, obtuse, pointed upward and one or two 

toothed on tIie upper side, the sinuses rounded; primary vein strong ; secondary veins once forked, one 

branch passing to the apex and the other to the sinus; 

sori seated on a vein at the apex of the lobes and frequently in the sinuses also. 

N otwi thstanding the fact that this fern is in fruit 

it is referred to Dennstaedtia with considerable hesitation, for to distinguish between this genus and Hy-

'111JenopliyllWln or Davallia is next to impossible with 

material so scanty and obscure. Although it is easy 

to distinguish between these genera in a living state, 

where all the organs of fructification can be carefully 

examined, it is exceedingly difficult to decide in regard 

to fossil forms, in which the organs of reproduction 

are more or less obscured by the accidents of preservation. In habit, texture, division, and nervation 

these genera are so similar that it will usually not 

be possible to distinguish them without fruit, and even 

in the fruit much depends upon the character of the 

indusium, which from its delicate nature is not usually preserved. 

The specimens under consideration' have been referred to Dennstaedtia because they are, as nearly as 

can be made out, most like this genus. The sori are 

situated at the margin of the frond and always at 

the apex of a vein, as in the living species. The 

strong, thick primary rachis probably excludes them 

from 11 ym.enopnyllwn, and the character of the sori 

8eems to separate them from Davalli{(). 

This is a very small, delicate fern, the longest pinIlule being hardly over 1 centimeter in length and 

G to 8 millimeters in width. It was probably several times compounded, to judge from the thickness 

of the rachis, but the largest fragment found only 

shows it to be bipinnate. The outline of the whole 

frond can not be determined. The pinnules seem to 

have been easily detached, as four of them were found 

unconnected and not otherwise much mutilated. They 

are elongated, deltoid, or lanceolate, with numerous 

pinnatifid divisions, which are rounded at the apex 

and in the sinuses between them. The nervation consists of a strong central vein and once or rarely twice 

forking lateral veins with the fruit dots borne on them 

at the margin of the frond. The fruit dots are thick, 

globose, and seemingly somewhat cup-shaped, but they 

are not well enough preserved to warrant positive 

. statements regarding the minute details. 

This species is exceedingly rate, for among several 

hundreds of pieces of shale exa.mined only the specimens figured were obtained. But as they are very 

obscure, requiring a glass for their detection, they 

may have been easily overlooked by the collector. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Arapahoe formation), Sedalia, Colo., 

quarry No.2, about 3,000 feet east of the Douglas 

coal mine, in a gulch near the railroad, collected by 

Arthur Lakes, June, 1890. 

Woodwardia latiloba Lesquereux 

Woodwa1'dia lMilobct Lesqtt61'eWlJ, U. S. GeoI. and Geog. Survey Terr. Ann. Rept. for 1873, p. 391, 1874; The Tertiary 

flora, p. 54, p1. 3, figs, 1, la, 1878. 

WOoc~1/)a1'dia Zatiloba var. minor Lesquereux, U. S. Geol. and 

Geog. Survey Terr. Ann. Rept. for 1873, p. 391., 1874; 

The Tertiary flora, p. 54, pI. 4, figs. 9, 9a, 1.878. 

This fine and perfectly distinct species is fairly 

abundant at a number of localities within the Denver 

beds at Golden, Colo., but usua.lly in fragments of very 

much smaller size than the nlagnificent type specimen, 

which is preserved in the United States National 

M:useum (No. 15). The smaller fragments, however, 

retain perfectly their characteristic outline and nervation, thus rendering their identification unquestioned. 

The small fragment described and figured by Lesquereux as variety 1ninor is also preserved in the 

United States National ~1useum (No. 117). In the 
22 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

IJublished accounts of this form it is said to have come 

from Black Buttes, Wyo., but in the Museum catalogue 

it was recorded by Lesquereux himself as being from 

Golden, Colo., and an examination of the specimen 

leads me to believe that this statement is correct. It 

is said to differ from the species in having the" pinnae 

small; rachis narrow; lobes short, obtuse; secondary 

veins less divided." It represents, as Lesquereux has 

said, only the tip of a pinna, so it would naturally be 

smaller and the lobes shorter and more obtuse. It is 

not, however, smaller nor different in outline from 

what must have been the terminal portion of certain 

of the smaller, upper pinnae of the large type specimen 

of W. latiloba. The nervation is of precisely the same 

character as in the species but is, of course, less 

branched to accommodate the smaller size of the pinna. 

I therefore do not hesitate to refer it, locality and all, 

to the species. 

The abundant but rather poorly preserved material 

from Sedalia, Colo., is not to be distinguished from the 

typical material. 

Occurrence: Denver formation, Golden, Colo. 

Dawson arkose, lower part (in beds equivalent to 

Ara12ahoe formation), Sedalia, Colo., 1,900 feet ,east of 

the Douglas coal mine. 

Woodwardia latiloba serrata Knowlton, n. var. 

Plate 2, Figures 1, 2 

Sinlilar to W oodwa:rdia latiloba in all particulars 

except that the margins of the pinnules are finely and 

evenly serrate. 

It is perhaps doubtful if this should be considered 

even subspecifically distinct from the typical form, as 

the only observable difference is in the serrate instead 

of entire margins, though it may be stated that of the 

numerous examples of W. latiloba examined not one 

has shown a tendency to become serrate. 

Occurrence: Middle Park formation, Middle Park, 

Grand County, Colo., from high terrace ridge between 

forks of Kinney Creek, in " breccia spoon " east of Hot 

Sulphur Springs, Gollected by 'Vhitman Cross, October 

ti, 1891. 

Pteris pseudopennaeformis Lesquereux 

Pteris pseu(],ope'nnaeformis Lesquereux, The Tertiary fiOl.'a, p. 

52, pI. 4, figs. 3, 4, 1878. 

Knowlton, U. S. Geol. Survey Bull. 204, p. 22, 1902. 

Hollick, Louisiana Geol. Survey Rept. for 1899, Special 

Rept. 5, p. 279, pI. 32, fig. 1, 1899. 

Berry, U. S. Geol. Survey Prof. Paper 91, p. 168, pI. 9, fig. 

6,1916. 

Pter'is pennaeformis Heer. Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1870, p. 384, 1871; idem 

for 1873, p. 283, 1874. 

?Newberry, U. S. GeoI. Survey Mon. 35, p. 7, pI. 48, fig. 5, 

1898. 

Pinnae large, subcoriaceous, narrow ly lanceolate, 

gradually narrowed to both base and apex; the margin 

entire to about the middle, obtusely dentate at and 

near the apex; midrib stout, more or less flexuose, 

grooved; veins thin, at an angle of 35° or 40°, close, 

straight, mostly simple or rarely once forked. , 

The above description is slightly modified from that 

given by Lesquereux, the changes being based on a reexamination of the type specimen, or rather the specimen figured by Le,squereux in the report on the Tertiary flora (U. S. Nat. Mus. No. 19) as typical. This 

is the only specimen I have had the opportunity of 

examin.ing, as none of the recent collections from 

Golden seem to have included examples. The specimens from Golden in the Museum of Comparative 

Zoology that were identified by Lesquereux as Pteris 

pseudopennaeforrmis are f.ound to belong to Salpichlaena anceps (Lesquereux) Knowlton. 

This species was first characterized by Lesquereux, 

under Heer's name Pte1"is pennaeformis, from speciInens collected by F. V. Hayden on Henrys Fork of 

Green River in Wyoming. It was also later collected 

by Hayden at Golden, Colo., from which came the 

specimen figured in the report on the Tertiary flora, 

Plate 4, Figllre 3, notwithstanding the fact that this 

particular specimen is recorded in the National l\1u-

, seum catalogue as having come from Henrys Forie 

The matrix is unmistakably that from Golden, and 

indeed the specimen is properly so recorded in Hayden's annual report for 1873, page 392, where this 

'specimen was first mentioned. 

This species' appears to be closely related to Pte1'is 

JJennaef01''1nis Heer, as pointed out by Lesquereux, and 

may be identical with it, but in the absence of speci~ 

mens fro111 Greenland and of specimens other than the 

type of P. pseudopennaeformis I have thought best' to 

retain Lesquereux's assignment. It is always possible 

that subsequent collections may contain material that 

will clear the matter up. 

If it has been identified correctly, this species has 

a wide geographic range, having been reported by 

me 48 from the lower part of the Clarno formation at 

Current Creek, Oreg., and by Berry 49 from the 'Vilcox ' 

deposits of Louisiana and Mississippi. It appears to 

be a rare form and thus far is represented by only 

very few poorly preserved specimens, so its characters 

a.re not made out with certainty. Berry has expressed 

the opinion that it is 'more likely to be referable to 

the genus .AsplenVum, and he points out its resemblance 

to .Asplenium) eolignitiewm Berry, a Wilcox species. 

Occurrence: Denver formation, Golden, Colo.; eai'ly 

collections made by F. V. Hayden. Green River 

formation ( n, Henrys Fork of Green River, Wyo. 

Lower part of Clarno formation, Current Creek, Oreg. 

48 Knowlton, F. 'H., U. S. Geol. Survey Buli. 204, p. 22, 1902. 

49 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 168, pI. 9, 

fig. 6, 1916. 
SYSTEMATIC PALEOBOTANY 23 

,\Vilcox formation, Vineyard Bluff, Caddo Parish, and 

N aborton and Mansfield, D.e Soto Parish, La. Wilcox 

group (Grenada formation), Grenada, Miss. 

Diplazium crossii (Knowlton) Knowlton, n. comb'. . 

Plate 2, Figure 6 

A.sl)Zeniu;m (DipZazimn) cTossVi Knowlton, U. S. Geol. Survey 

Bull. 152, p. 44, 1898. . 

Dlplazi'ltm rn'lteZ:~i? Heel'. Lesquereux, U. S. GeoI. and Geog. 

Survey Terr. Ann. Rept. for 1873, p. 393, 1874; The Tertiary flora, p. 55, pI. 4, figs. 10, . lOa, 1878; The Cretaceous 

and Tertiary floras, p. 138, 1883. 

Outline of frond not known; pinnae coriaceous, narrowly lanceolate, gradually tapering upward to a long, 

acuminate apex, and from the lower third tapering 

downward to a wedge-shaped base, cut for one-fourth 

or one-third the width into oblong, rather obtuse segments; midvein of segments exceedingly thin and obscure; veins very thin, some arising from the midvein, 

others from the main rachis, all at an acute angle, 

once or twice forked. 

This species, as indicated in tile above synonymy, was 

doubtfully identified by Lesquereux with Heer's 

DipZaziu(J1v 'I71Iwille'Ji, fr<?nl the brown coal of BornstKdt. 

~I'he specimen on which Lesquereux based this identification is preserved in the United States National 

J\fuseum (No. 18 [by error 19]) and is before me as I 

write. lIe figured only one broken pinna, but as a 

Jnatter of fact there are three pinnae on this piece of 

lnatrix, all lying in the same direction and plane. One 

·of these, although somewhat covered with the matrix, 

is a more perfect specimen than the one figured in that 

it. shows the basal portion nearly complete. A careful 

-examination of Lesquereux's specimen, together with 

about 20 others since collected by Arthur Lakes near 

South Table Mountain, Golden, shows that they differ 

in at least three essential particulars from Diplaziwm 

·muelleri, as described and figured by Heer.50 These 

·differences are as follows: (1) The pinnae are cut int9 

short, entire-margined segments, but in typical D. 

muelle'J'i the pinnae are distinctly doubly serrate. (2) 

'They have no intramarginal nerve as figured by Lesquereux. It is difficult to see how this error of interpretation arose, for although Lesquereux's specimen 

:is rather obscure, there is not the slightest trace of an 

intramarginal nerve such as that described and figured 

for D. 'lnueUf!ri. (3) The veins, although rather close 

together, do not anastomose as described and figured 

by Lesquereux. Even the obscure type specimen fails 

to show a single,place where the veins anastomose, and 

the very well preserved new material proves still more 

conclusively that they do not. The fact that Lesquereux compares the anastomosing to that observed 

in his Gymnog'J'a'l1wna ga'J'l~neri 51 shows cleady that 

150 fleer, Oswald, Braunkohlen Pflanzen yon' Bornstiidt, p. 8, pI. 1, 

figs. 2, 2b. 1869. 

111 I ... esquereux, Leo, The Tertiary flora, p. 58, p1. 4, fig. 2, 1878. 

he was greatly misled concerning it, and it seems 

probable that his description was made entirely from 

the drawing, which, so far as I am able to conclude, 

must have been made for a different plant. The 

veins all arise from the secondary ra'chis or from the 

thin mid-vein and fork once or rarely twice before 

rcaching the margin. 

From this description it becomes evident that these 

specimens can not be considered specifically identical 

with Diplaziwm rnJUelle'J-i Heel', and it is necessary to 

give them a new name. The question of their generic 

reference may be first considered. The absence of 

fructification and of a knowledge of the size, shape, 

and habit of this form makes a generic allocation more 

or less uncertain. It has some resemblance to certain 

li ving species of DiplaeiuVf11J. 

In this connection it may be well to clear up another 

misunderstanding concerning this species. In Hn,yden's annual report for 1873 it is stated that this 

species came from South Table Mountain, Golden, but 

in the report on the Tertiary flora it is accredited to 

Henrys Fork: Wyo. A comparison of the type specimen with those recently obtained by Mr. Lakes shows 

them to be absolutely identical lithologically. There 

can be no doubt that the specimens actually came from 

Golden and not from Henrys Fork. And upon this 

probably hinges a still further error. In Lesquereux's 

report on the Cretaceous and Tertiary floras (p. 138) 

Diplaziu'ln 'lnuelleri is enumerated from the Green 

River formation without mention of exact locality, 

being simply referred to the page of the report on the 

Tertiary flora in which this species is des~ribed and 

recorded from Henrys Fork. If, as there stated, it 

had come from Henrys Fork it would with very little 

doubt be of Green River age, and it seems probable 

that Lesquereux introduced it into his list of species 

from the Green River deposits upon this supposition. 

At least there is no evidence to prove the contrary, as 

thel~e are no specimens extant, so far as known, that 

came without question from the Green River formation of Henrys Fork. It would therefore seem best to 

drop this species from the flora of the Green River, at 

least until further collections prove or disprove the 

correctness of the above surmise. So limited above in 

its distribution DiplaziuYJTh o'rossii is found only in 1 he 

lowest leaf-bearing stratum of the Denver beds. 

Among the material obtained in Middle Park by 

George L. Cannon for S. F. Emmons I find a, single 

specimen with its counterpart that belongs to this 

species. It is rather poorly preserved but does not 

differ in any particular from the Golden specimens. 

Occurrence: Denver formation, Golden, Colo., on a 

bluff of the prairie a quarter of. a mile -south of the 

Reform School and about 500 yards southwest of 

South Table Mountain. 1\1iddle Park formation, Mid-
24 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

dIe Park, Colo., on north bank of the Grand [Colorado] River near Sheriff's coal shaft, on Sheriff Creek, 

collected by George L. Cannon, 1889. 

Genus ALLANTODIOPSIS Knowlton and Maxon 

AllantodAopsis Knowlton and Maxon, U. S. GeoI. Survey Bull, 

696, p. 61, 1919. 

Fronds simply pinnate; venation pinnate, the veins 

free, once or twice forked; sori dorsal, always on the 

upper or distal fork of a vein, short, thick, sausageshaped, with no evidence of the presence of an 

indusium. · 

. Allantodiopsis erosa (Lesquereux) Knowlton and Maxon 

Plates 4, 5 

Allwntodiop'sis el'osa (Lesquereux) Knowlton and Maxon, U. S. 

Geol. Survey BUll. 696, p. 61, 1919. 

Pte1"is erosa Lesquereux, U. S. Geol. and Geog. Survey Terr. 

Ann. Rept. for 1871, SuppI., p. 12, 1872; idem for 1873, 

p. 392, 1874; The Tertiary flora, p. 53, pI. 4, fig. 8, 1878; 

r:rhe Cretaceous and Tertiary floras, p. 121, pI. 19, fig. 

1, 1883. 

Pteris subsimplea; Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1873, p. 392, 1874; The Tertiary 

flora, p. 52, pI. 4, figs~ 5-7, 1878. 

Osmmula major Lesquereux, The Cretaceous and Tertiary 

floras, p. 121, pI. 18, fig. 5, 1883. 

Pter-is ttndttlata Lesquereux, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, p. 43, 1888. 

AspleniUJrn el'osum (Lesquereux) Knowlton, U. S. Geol. Survey 

Bull. 152, p. 45, 1898. 

Asplenitl-l11; sub simp lea; (Lesquereux) Knowlton, U. S. Geol. 

. Survey Bull. 152, p. 45, 1898. . 

Frond large, coriaceous, simply pinnate; pinnules 

simple, alternate or subopposite, apparently sessile, 

oblong-lanceolate, rounded and often very unequalsided at base, long acuminate or taper pointed at apex; 

margin rarely' quite entire, usually undulate, crenulate, or sharply serrate above the middle, sometimes 

~\ntirely serrate; middle nerve strong, round; veins 

free, at an open angle of divergence, distinct, distant, 

very rarely simple, usually forking once near the nlidvein and again at or above the middle; sori dorsal, 

large, sausage-shaped (5.6 millimeters long, 1.5 millimeters broad),. seated on the upper fork of a vein about 

3 or 4 millimeters above the midvein. 

After an examination of the type 'specimens of all 

the so-called species enumerated above, with the ex, 

ception of Osmunda 'mA1Jjor (which, however, is finely. 

figured), together with much additional material representing all the forms obtained a few years ago by 

Arthur Lakes, I ,am absolutely unable to draw any 

sa tisfactory specific lines between them. In size, general outline, and nervation they are identical, merely 

differing slightly in the character of the margin of 

the pinnules.· Thus Pteris subsimplew is described as 

having the margin "entire or nlinutely crenulate"; 

P. erosa has the "borders obtusely dentate' toward 

the point, irregularly crenulate-lacerate in the midcUe" ; P. wnd1.tlata is "regularly deeply undulatecrenate, especially in the upper part"; and OS'm!l1lnda 

'Initjol' is simply described as having the" borders undulate." Ptelris suosi'lnplew and P. wndulat(); were 

rounded on the middle and lower parts of pinnules; P. 

(ff'o;a is 'based on the upper parts of pinnules, and Os~ 

mrumrfa 7lUJ;jol' upon the basal parts of several pinnules 

fortunately preserved so as to indicate the position of 

the rachis. Further, they were all, except O. major', 

established upon fragments which give no indication of 

the nature of the entire frond or plant; consequently P . 

s1.,lbsimplew, P. el'osa, and P. wnuuZ(])ta were thought to 

be entire fronds, while O. majo". was, of course, clearly 

pinnate. It therefore becomes reasonably clear that 

they all represent different parts or slightly different 

forms of a single species, a conclusion to which Lesquereux himself seems to have been almost forced,. 

for in his l~emarks under O. m.ajor 52 he said: "It is, 

therefore, uncertain whether these fragments represent 

two or three species, or whether, perhaps, they may all 

be referable to the same." 

The probability that these four nomlnal species are 

really one and the same thing is further strengthened 

by a knowledge of their geologic and geographic distribution. Thus, Pter'is erosa was described from 

Fishers Peak, in the Raton Mountains, Colo., in beds· 

now referred to the Raton formation, which is cor-· 

related with the Denver formation. This form is' 

found at several places in the Raton Mesa region of 

Colorado and New Mexico, though it is nowhere abundant. The other forms (Pter'is SUOSi171iplew, Pter'isl.tndulata, and OS1n1.l/J~d(f) 171)(f)jor) were all described 

from the andesitic beds of the Denver formation at. 

Golden, Colo. 

From a study of all the material now available it 

becomes apparent that we are dealing with a fern 

that possessed a large, simply pinnate frond with 

alternate or subopposite coriaceous pinnules, which 

are broad, linear, or oblong-lanceolate in outline (10" 

to 16 or 18 centimeters long and 3 to 3.5 centimeters 

broad), with 'an abruptly rounded and unequal-sidecl 

base and an acuminate or ta per-pointed apex. 

It is perhaps not to be wondered at that a fern as· 

large as this should rarely be found preserved entire. 

The conditions for the entombment of a large frond 

might even be favorable, but in exhuming a specimen 

of this size in an easily broken matrix it would be exceedingly difficult to procure a complete fossil, even if 

preserved. It is but natural, therefore, that these 

large isolated pinnules should have been taken for entire fronds, and as' even these were rarely preserved 

52 Lesquereux, Leo, The Cretaceous and Tertiary floras, p. 121, 1883. 
SYSTEMATIC PALEOBOTANY 25 

entire there is perhaps an excuse for regarding the 

basal and' terminal portions as representing different 

species, espe'cially as they have been shown to vary 

from an undulate margin in the lo'wer part to a toothed 

margin in the upper part. Thus in one of the most 

perfect examples thus far obtained the lower portions 

of the pinnules are of exactly the same character, indeed almost a counterpart of the type of 08171JWnda 

majo!)', whereas in the upper portion of at least one 

of the pinnules the margin begins to be slightly 

toothed, showing its transition to the so-called Pte-ris 

e1'osa. This transition from a nearly or quite entire 

outline to a more 01' less pronounced toothed margin 

in the same frond or pinnule is by no means rare 

among living ferns. 

The, small fruiting specimen (pI. 4, fig. 1) obtained 

by Lakes near the Douglas coal lnine, Sedalia, Colo., 

represents, so far as I know, the only example of the 

fruit of these ferns ever obtained. It is well shown 

in the figure and has been described at length. It is 

evidently in very mature condition, for all trace of 

the indusium has disappeared (if, indeed, it was ever 

present), and the sori present a pulverulent appearance under a low power, though no well-defined sporangia can be made olit. Some years ago I was led to 

regard this fruit as indicating the genus Aspleniurn, 

as I then interpreted the sori to be double and contiglIOUS on opposite sides of a vein, but after a reexamination of the specimen I can but decide that conclusive evidence for this assumption is wanting. There 

is no indication of a nliddle parting through a sorus, 

though some trace of it would probably be present if 

it ever existed, and hence the sori appear to be truly 

dorsal. 

This fruiting specimen was submitted to the late 

Prof. D. C. Eaton, of Yale University, a well-known 

pteridologist, who at once suggested its relationship 

to Allantodia brurnoniana Wallich, now known as DipZaziOl)sis javanica (BlunIe) Carl Christensen, a large 

pinnate fern native to the East Indies and Pacific 

islands. In the size and outline of the pinnules and 

in the size and position of the sori this livi~g species 

is almost identical with the fossil form, but the nervation is entirely different. The veins branch at the 

buse as in the fossil and are there only soriferous, but 

they an:astomose into subhexagonal areoles toward the 

margin. 

I have also submitted this material to anotJler student of the ferns, Mr. Willianl R. Maxon, of the 

United States National :.Museum, and have received 

many vaiuable sllggestions from him. He also reH),arked "at once the simIlarity of the fructification to 

that of Diplaziopms javanica, with which, however, 

the fossil plant could not, on account of its free venation, be associated. He further directed my attention 

to the marked similarity of the fossil forms under 

consideration to OoniogTa17Mne frawinea Don (Gy17~­

nog'ra171,'l7W java'11ica of authors), a species of the 

Himalayas, Java, and Ceylon., The agreement here in 

size and nervation is almost perfect, but the fruiting 

is entirely unlike. Instead of a short, sausage-shaped 

sorus confined to the upper fork of a vein,·we have a 

very long, narrow, and usually forked sorus-that is, 

the sori follow both forks of a vein, often for a considerable distance. As 1\1r. Maxon has called to my 

attention, the fossil under consideration appears to 

be excluded from all the present genera of the tribe 

Aspleniae by the dorsal rather than lateral position 

of the sori. A comparison might readily be instituted 

between isolated pinnae of the fossil form and numerous large-fronded living forms, such as certain species 

of Pte1is, Ste1wchlaena, and Acrostichwrn, but all 

would be excluded on the ground of different fruit, 

and, following Mr. Maxon's advice, I have decided to 

institute a new genus for its reception. 

Various relationships among fossil forms have been 

suggested for one or another of the so-called species 

here included under Allantodiopsis erosa, but as they 

were all based on a know ledge of the sterile forms 

only, they can now, in the light of the fruiting specimen, be regarded as of little value. Gardner and 

Ettingshausen, in their monograph on the British 

Eocene flora, stated, in connection with Pteris 

eocenic(J), that "The American Eocene species P. subs-l171J.plew and P. erosa, while possessing the same venation, have much broader pinnae," and later that "Although species of Acrostichwrn, A ngiop teris , Gymrnogrmmxme, etc., possess similar venation, the characters 

furnished by this form [Pteris eO'cenioaJ] so closely 

correspond to what is met with in the recent species of 

Pte'l'is that its reference to this genus can not be 

doubted." If Pteris' eocenioa is really closely allied 

to our form, it should, of course, be considered as 

belonging to this new genus. So also' with several 

other species described under the name Pteri8, as 

P. pa'J'schlugiam.a Unger 53 and P. pennaef01"'171is Heel,.54 

But this is mere conjecture, as size, outline, and 

nerva.tion are often of secondary value in determining relationship in ferns; and it is impossible, in 

the absence of fruit, to determine the genus of many 

Ii ving species. 

. Occurrence: Denver formation, South Table Mountain, Golden, Colo., from early collections by Hayden 

and Lesquereux and later by Arthur Lakes; types of 

Ptmis Und1()lata Lesquereux in Museum of Comparative Zoology, Cambridge, Mass., No. 853. Dawson 

arkose, lower part (in beds equivalent to Arapahoe 

formation), east of Doug:las coal mine, near ~edalia, 

63 UngeJ:,Franz, Chloris Protogaea, pt. 1, p. 122, pI. 36, fig. 6, 1847. 

M Herr, Oswald, Flo'ra tertim'ia Helvetiae, vol. 1, p, 38, 1855. 
26 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Colo., collected by Arthur Lakes. Middle Park formation, Sheriff Creek, north bank of the Grand 

[Colorado] River, Middle Park, Grand County, Colo., 

collected by Geo. L. Cannon. Raton formation, Fishers Peak, Raton Mountains, Colo. 

Saceolomagardneri (Lesquere'ux) Knowlton 

Plate 3, Figures 3-8 

Sa.ccoZorna gardneri (Lesquereux) Knowlton, U. S. GeoI. Survey Bull. 696, p. 560, 1919. 

Ptens gardneri Lesquereux, U. S. Geol. Survey Terr. Ann. 

Rept. for 1873, p. 393, 1874. 

Gym,nogramma. gariJJneri (Lesquereux) Lesquereux, The T¢rtiary flora, p. 58, pI. 4, fig. 2, 1878. 

Outline of whole frond not known, but fron1 its general resemblance to the living SaccolO'm(J; elegans it is 

presumed to have been erect, with the pinnae lanceolate and perfectly entire or slightly undulate; the 

texture is apparently subcoriaceous but thin. The 

sterile pinnae are narrowly lanceolate, acuminate at 

apex, and apparently abruptly rounded at base; the 

margin is slightly undulate. 

No complete specimen has been found, but the most 

perfect one shown is 17 centimeters long and shows 

no evidence of narrowing to either base or apex. It 

nlUst have been 20 to 25 centimeters in length. Midrib very strong, grooved above, finely striate below, 

the veins strong, at an angle of 30° to 40°, usually 

forking just above the midrib and often just above 

the middle, generally anastomosing. near the margin 

though sometimes forking again without anastomosing. The. fertile pinnae are apparently slightly 

smaller than the sterile ones, 2 to 3 centimeters wide, 

with the venatiQn similar, forking near the midrib 

and again at or near the middle, not or rarely anastomosing but running free to the margin. The sori 

a.re small, close, circular, one on each vein or branch 

reaching the margin, slightly submarginal in position, forming a continuous row; indusium obscure, 

perhaps distally attached and folding over the sorus, 

or perhaps the margin of the frond is slightly folded 

over the sorus, thus simulating an indusium. 

The type specimen of what for many years has been 

known under the name Gymrnogramma garilJneri is 

preserved in the United States National Museum (No. 

17). It was found on Sand Creek, some 15 or 18 miles 

east of Denver, Colo., in beds long supposed to be Arap-· 

ahoe in age but now pretty definitely settled as belonging to the Denver formation. This type specimen is 

very poorly preserved and consists simply of two sterile fragments representing the middle portions of pinnules or perhaps whole fronds. They lie side by side 

and less than 2 centimeters apart, and as they are 

without apex or point of attachment there is no evidence to show whether each represents a whole frond or 

whether they may not be piIinules of a very large 

compound frond. The latter interpretation is the one 

that was adopted by Lesquereux, who gave the following description: 

Frond large, simply pinnate; pinnae large, linear, broader 

in the middle, in right angle to the rachis, rounded at the 

base; borders deeply undulate; middle nerve broad, grooved 

in the middle, flattened on the borders; veins in an obtuse' 

angle of divergence, abruptly curved downward at the base, 

or decurring to the rachis, forking once or twice, joined by 

anastomoses and forming by cross branches irregular, . long 

areolae. 

. Saporta, to whom apparently Lesquereux had submitted a' drawing of his so-called Gymnogra1n'l7Ul 

ga'l'dneri, expressed the opinion that it was identical 

with or at least closely related to a fern from the 

Gypse of Aix which he regarded as referable to Oh'l'ysodium" a section of the genus AcrostichU'l7L. Gardner 

and Ettingsha usen 55 also discuss this American fern 

in connection with the description of their 0 hrysodiu'I7~ 

lanzeanU1'n, from the Lower and Middle Bagshot beds 

of England. It is not to be denied that G. ga'l'dne'f'i 

has a degree of resemblance to the living A.f]1'ostichum: 

(0 hrys0 diu171J ) aureU1n Linne, but it differs essentially in its looser, less anastomosing nerves. I was 

at one time inclined to transfer GY'l7~nog'l'a'Jn'l7Ul gardneri to Acrostiohum, but decided to retain it as placed 

by Lesquereux in the hope that sooner or later additional material would come to light that would help 

to settle its status. This new material has fortunately 

now been found. 

In 1910 C. W. Cooke collected for G. B. Richardson 

some excellently preserved ferns and other plants: 

from the basal part of the Dawson arkose near Ramah . ,. Colo. The ferns were represented by the two examples figured, which show the sterile and fertile 

pinnae. The fronds are preserved on a hard baked 

clay shale and exhibit all the details of nervation and 

fructification in a high degree of perfection. The 

fructification as well as the shape and nervation immediately suggested reference to the genus Sacaolo'l7Ul 

of which this was supposed to be the first known fossil 

species. Its relation to Lesquereux's GY'lnnog'l'a111./Ilur; 

gardneri was not then suspected. 

Later .collections from the Dawson arkose on Jimmy 

Camp Creek, east of Colorado Springs, and from. 

Mosby, El Paso County, Colq., were found to include: 

a number of large ferns that were recognized at once 

as belonging to SacCOl0l1W1• ~fost of them are sterile 

fronds, which turn out to be indistingUIshable from' 

Cymnog'l"al1t1na ga'l'dneri. For example, a. segment 

from the base of the large pinna shown could not be 

distinguished from the type of GY1nnogram'l7Ul ga'l'd-· 

til Gardner, J. S., and Ettingshausen, Constantin von, Monograph on. 

the British Eocene flora, vol. 1, Filices, p. 28, Paleont. Soc., 1879. 
SYSTEMATIC PALEOBOTANY 27 

neri, and it is so regarded. The reference of these 

Mosby specimens to SaccolO100 is attested by a number of fruiting examples, one of which is shown in 

Plate 3, Figure 7. This is a mere fragment from the 

middle of. a pinna, about 4 centimeters long and 2.5 

centimeters wide. It has a broad, grooved midvein 

nnd strong forked veins as in G. gardneri, and the 

fruit is apparently more mature and forms an almost 

continuous row. None of the Mosby specimens show 

the basal portion of a pinna, but a number show an 

apparent narrowing toward the apex. One exhibits 

an irregularly erose appearance on one side, but 

whether this is natural or due to injury can not be 

determined. 

The genus SacoolO1na is a small group of davallioid 

polypodiaceous ferns that, with a single exception as 

currently accepted, is confined to tropical America. 

The type of the genus is SaCCOl017W, elegans· I(aulfuss, 

which inhabits tropical America from Guatemala and 

the West Indies southward to Rio Janeiro. Certainly 

congeneric with this, though of quite different form 

and nervation, are two other species-S. im/rayanu17L 

Hooker and S. wircklei Christ, the first occurring in 

the ""Vest Indies and British Guiana and the second 

in Costa Rica. The genus is apparently in need of 

critical revision, but whatever the result of such study 

may be, the fact remains that the type of the genus 

and the two species abovementioned are congeneric, 

and with them undoubtedly belongs the fossil form 

here described. 

SacCOl017W, g(JfJYlineri, as it must now be called, is 

most closely related to S. elegans, with which it agrees 

in the shape and apparent size of the pinnae, in the 

position, size, and appearance of the sori, and, in the 

indusioid character of the m.argin. It differs in nervation from S. ele{}0Jn8, which has the nerves mainly 

simple and only very rarely forked. So far as 

observed the veins in S. elegans never anastomose. 

There is no previous record of Sacco.zo17W, in a fossil 

state, and its presence in eastern Colorado in early 

J~~ocene time is of interest and possible importance in 

its bearing on the climatic conditions that prevailed 

fit that time. 

The sterile fronds of Saccolo'lJw gardneri are very 

similar in size and appearance to Allamtodiop'si8 erOSfJJ 

and might be confused with it unless the nervation 

is studied very closely. In Alla~tod}iop8is the veins 

fork once or twice but never anastomose; in Saocoloma 

the veins fork in a. similar manner but occasionally 

anastomose. If the specimen is poorly preserved, it 

may be difficult to detect the anastomosing of the veins. 

The degree of this anastomosing appears to c;liffer 

somewhat in different specimens-that is, some show 

this condition very plainly and others obscurely. 

Occurrence: Denver formation [type], roof of coal 

mine, Sand Creek, about 15 miles east of Denver, 

Colo., collected by A.Gardner in 1873. Dawson arkose, near Ramah oil well, in the NW. * sec. 33, T. 9 

N., R. 61 W., about 40 miles northeast of Colorado 

Springs, Colo., collected by C. W. Cooke for G. B., 

Richardson, September, 1910; east bank of Jimmy 

Camp Creek, 9 miles east of Colorado Springs, Colo., 

collected by A. C. Peale, 1908; dump of Mosby coal 

mine, Mosby, EI Paso County, Colo., collected by F. H 

Knowlton and G. B. Richardson, July, 1910. 

Saccoloma sp. 

Plate 3, Figures 1, 2 

The fragment of fern here figured is so poorly pres(:rved that under ordinary circumstances its description would hardly be worth while. Yet it belongs to 

a type that has not previously been noted. It is very 

close to or possibly identical with SaJ(JcolO7J!.a r/dff'dneri, just described. It is a sterile frond or pinnule 

that was probably 6 or 8 centimeters long and is nearly 

3 centimeters wide. It differs slightly from S. gardJneri in having the margin obscurely toothed and in 

having the nervation at a less 'acute angle of divergence-both differences that might disappear if we 

had a series of specimens instead of a single one. The 

nerves fork very near the midvein and often fork 

again above the middle. They anastomose not far 

from the margin, as in the sterile frond of S. gardneri. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek, 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., col-' 

lrcted by F. H. Knowlton, July, 1910. 

Salpichlaena anceps (Lesquereux) Knowlton 

Plate 7, Figures 1-3 

Salpichlaena anceps (Lesquereux) Knowlton, U. S. Geol. Survey Bull. 696, p. 574, 1919. 

Ptc1'is anceps Lesquereux, 'u. S. Geol. ,and Geog. Survey Terr. 

Ann. Rept. for 1872, p. 376, 1873. 

Pteris a/fini8 Lesquereux, U. S. GeoI. and Geog. Survey Terr. 

Ann. Rept. for 1873, p. 392, 1874. 

OS11Mtnda atfinis (Lesquereux) Lesquereux, The Tertiary flora, 

p. 60, pI. 4, fig. 1, 1878. 

Knowlton, U. S. Geol. Survey Mon. ii2, pt. 2, p. 673, pI. 80, 

figs. 4, 5, 1899. 

Ptcris pseudopen.naefol"1nis Lesquereux. Lesqu'ereux, Harvard 

ColI. Mus. Compo Zoology Bull., vol. 16, p. 43, 1888. 

Fern apparently of large size, probably climbing, in 

effect bipinnate (primary rachis unknown), secondary 

rachis strong, slightly irregular or zigzag; pinnules 

alternate, closely sessile, narrowly lanceolate, truncate 

or slrghtly heart-shaped at'base, rather obtusely acuminate at apex, margin entire or slightly undulate; midvein very strong, slightly undulate; nervation pinnate, 
28 VLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

free, the veins relatively stro~g, usually at a low angle 

of divergence, rather remote, once or twice forked 

(their apices slightly thickened at the margin ~) ; 

fruit unknown. 

As may be seen from the above synonymy, quite diverse views have been entertained regarding the position of this fern. A mere fragment from Golden, 

Colo., of what was thought by Lesquereux to be this 

plant was first described by him under the name Pteris 

ance:ps.56 In the following year' he obtained more perfect material from the same locality, and, thinking he 

could observe slight differences between this and his 

Pteris anceps, he gave it the name Pteris affinds. 57 

Later, when he came to prepare his report on the 

Tertiary flora, he had received still more perfect examples, and, considering its affinity to be rather with 

Oswunda than, with Pteris, he described and figured 

it as OS17'~unda affinis. On this point he says: 58 

The position of the pinnae (the middle one seemingly terminal), their size and form, and also the characters of the 

nel'vation of this' fern seem to indicate its generic relation 

with Osnmnda rather than with Pte1"is. 

been in the fossil. The pinnules, which are of approximately the same size and shape, are opposite or 

nearly so but, unlike those of the fossil forms, are 

invariably short-stalked. The nervation of the pinIlules in the living species is of the same general 

appearance as in the fossil, but there are differences. 

The veins are nearly at a right angle with the midvein, closer together, and .oftensimple, though usually 

forked once; their apices are slightly thickened and 

form a cartilaginous margin to the pinnule. The 

fruiting in this living species is very peculiar, the 

sori being linear, continuous or nearly so, parallel 

with the midrib and contiguous to it; they are covered 

by a membranous indusium, which unrolls at nlaturity. 

Although no fruit has been found on the fossil 

forms, I do not hesitate to place them· in the genus 

Salpiohlmena, on the ground of the strong resemblances 

above pointed out. With the scanty ma~erial at his 

command Lesquereux was perhaps justified in placing 

them in 08'rn1.llnda, which his best specimen does somevirhat resemble, being the three terminal pinnules of a 

rather small pinna. 

So the matter rested until. the fortunate discovery In the material belonging to the Museum of Comby Lakes, in 1890, of a nUlllber of fairly weH, pre;: parative' Zoology Lesquereux 59 identified two specis(\rved1arge specimens which ,give us,a much better, ' mens as Ptens p'8eudopennaefor17~is Lesquereux. These 

idea of the size and probable habit of this fern than specimens are before me and prove undoubtedly to 

any before obtained. The best of these seems to in- belong to Salpichlaena {JJnceps. The best-preserved 

dicate a large fern, apparently bipinnate, though the example is figured and shows' parts of five pinnae, 

main rachis is not preserved. Fronl the. position of . similar in all particulars to the other figures. 

the pinnae on the matrix it luay be· presumed that Occurrence: Denver formation, South Table Mounthey were alternate on the primary rachis, but this tuin, Golden, Colo., collected by Arthur Lakes. Specipoint is of course uncertain. The secondary rachis men identified as Pte'ris pseudopennaeformis in the 

is very strong and slightly zigzag, as may be observed Museum of Comparative Zoology, Cambridge, Mass., 

in certain living ferns of climbing habit. The posi- No. 852. 

tion, shape, and nervation of the pinnules have been 

indicated in the diagnosis and need not be repeated. 

By the kindness of Mr. William R. Maxon, of the 

United States National Museum, my attention has 

be(3n drawn to the undoubted close similarity between 

the fossil under consideration and the, living Salpichlaena volrubilis J. Smith, or Blech1'llU7n volubile of 

authors, the name by which it was ear1ier known. 

Thi~ fern, the on~y known species of the genus, is 

widely distributed throughout Central and South 

America, being, for instance, especially abundant in 

Brazil, where it is described as climbing to the tops 

of the loftiest trees. It needs but a glance at a suite 

vi specimens to show that Mr. Ma.xon's conjecture 

was well founded, for the living fern has a strong, 

flexuose primary rachis with the odd-pinnate pinnae 

disposed alternately, as they appear possibly to have 

66 U. s .. G~ol. and Geog. Survey Te,rr. Ann. Rept. for 1872, p. 376, 

1873. . 

G7 Idem (or 1873, p. 392, 1874., 

t,R Tbe Tertiary flora, p. 60, 1878. 

Family SCXIZAEACEAE 

Anemia mosbyensis Knowlton, n. ap. 

Plate 8, Figure 9 

The material from the Mosby coal mine at Mosby, 

Colo., contains numerous fragments of an Ane17'~ia that 

it seems best to describe as new l:ather than to refer 

to a named species on what may be insufficient data. 

It is a rather remarkable fact that although Anemia 

appears to be widely distributed, both areally and vertically, it is uncommon to find specimens so perfectly 

preserved as to give a satisfactory idea of the 'yhole 

frond. The present case is an example in :point. 

There are a dozen or more fragments of pinnae but 

only one that gives mucil of an idea of the complete 

frond. This is shown in' the figure. It is t~e apica~ 

portion of the frond, ot' which a length of about 1'0 

centimeters is preserved, b.ut it was undoubtedly lo~ger 

50 Lesquereux, Leo, Harvard CoIl. Mus. Compo Zoology Bull., vol. 16,. 

p. 43, 1888. . 
SYSTEMATIC PALEOBOTANY 29 

than this nnd possibly was forked. The two lower 

pairs of pinnae are alternate, at an acute angle with 

the main raehis, and narrowly linear-Ianceolate. The 

lowest are ,nenrly 8 centimeters long, less than 1 centimeter wide at the base, and slenderly acuminate at the 

apex. They are decurrent at the base and have the 

margins provided with remote, low, sharp teeth. The 

pinnae next a bove the two lower pairs are greatly reduced, being hardly 2 centimeters in length, and are 

not cut to the rachis. The others above are progressively shorter until in the f'.xtreme apical portion they 

ure probably reduced to mere slight indentations 01' 

teeth. The nerva.tion is characteristically that of the 

genus, consisting of a slender secondary rachis or midvein and numerous close, parallel, once or twice forking veins, which arise at a very acute angle. 

The form here described and figured resembles Inore 

or less closely a number of fossil species. For instance, 

it is of the snme type as and at one time was identified 

with Anemia 8'lloC1·eta.cea (Saporta) Gardner and Ettingshausen,6o a very abundant species in the Eocene 

of Bournemouth, England. The European species has 

narrower pinnae, which are abruptly reduced in the 

upper portion of the frond, as in A. '17w8byensis. The 

mnrginal teeth are also different and the veins at a less 

acute nnglc. 

Ane'lnia oooidentalis J{nowlton,61 from the Raton 

fOl'mntion, of the Raton J\tlesa region of Colorado and 

New M:exico, is also of this type but differs in the ab-

~ence of a reduced apical portion-that is, the frond 

is cut to the tip into long, narrow pinnae. The lower 

pinnae are twice or more the size of the lower ones in 

A.. '11wsbye118'lS and have larger, sharper teeth. 

Another species of this type is A ne'l7~ia rre11wnti 

Knowlton ,02 fr0111 the Frontier formation of southwestern 'Yyoming, which is very closely related to 

A. 8ulJ01'etaoea. It is about the size of A. 1nosoyensis 

but is readily distinguished. 

Occurrence: Dawson arkose, dump of Mosby coal 

mine, M:osby, Colo., collected by F. H. J{nowlton and 

G. B. Richardson, 1910. 

Anemia lanceolata. Knowlton, n. sp. 

Plate S, )j"'igure 10 

Fronds apparently coriaceous, evidently of considerable size (15 to 18 centimeters long, 12 or 14 centimeters wide) , probably deltoid in general outline, twice 

pinnatifid; main rachis strong, somewhat ridged; 

pinnae alternate, rather remote, very oblique, stalked 

00 Gtll"dne1', J. S., find. Ettlngshausen, Constantin von, Monograph on 

the British Eocene florn, vol. 1, FiUces, p. 45, pIs. 8, 9, Paleont. Soc., 

1880. 

01 Knowlton, F. H., U. S. GeoI. Survey Prof. Paper 101, p. 285, pI. 

04, fig. 2, 1917 [1918]. 

O!l Knowlton, F. H., U. S. Geol. Survey Prof. Paper 108, p. 84, pI. 31, 

f1~. G; Ill. 3~, fi~s, 1-3, 1917. 

S5,J!)2-:~O--3 

or in the upper portion sessile and decurrent, narrowly 

lanceolate, slenderly acuminate at the apex, wedgeshaped at the base, cut nearly to the secondary rachis 

into numerous opposite or alternate ovate-lanceolate 

oblique pinnules or segments; 1nargin of the large 

lower pinnules usually with a few sharp teeth, other 

pinnules entire; mid vein of pinnules thin, obscure, 

form~d by a very thin vein; lateral veins four or five 

pairs, at a very acute angle, usually only once forked 

in passing to the margin; fruit not known. 

This form is represented by a dozen or more ex9,mples. Although they are not very completely preserved, it can be seen that the frond must have been of 

considerable size. So far as can be made out, the frond 

appears to have been broadly deltoid in outline. 

It is with a good deal of hesitation that I venture to 

describe this form as new to science. So far as American material goes the first form to be det~cted that is 

now known to fall within this group was the GY'l71lno- . 

glJ'a1nma haydenii of Lesquereux,63 which came from 

the south end of the Yellowstone Lake in strata believed to be of Laramie age. Some years later it was 

shown, apparently conclusively, that this species was 

identical with Ane'17~ia 8uboretaoea (Saporta) Gardner and Ettingshausen, from the Eocene of Sezanne, 

Bournemouth, and other European localities. 

Newberry stated that he had specimens of this fern 

collected at Point of Rocks, 'Vyo., Golden and Erie, 

Colo., and Bellingham Bay and Carbonado, Wash. 

I have myself collected specimens at Carbonado, 

"Vash., but they differed so markedly from what I have 

supposed was typical A. pe'rplema Hollick that I was 

forced to give them a new name, and so with the 

specimens in hand. I am able to note so 1nany differences, albeit slight, that it has seemed best to describe 

this fern at least tentatively as a new species. Thus 

from the so-called GY'l1vnogrannma haydenrii it appears 

to differ in having Inuch longer, narrower pinnae with 

more prominent pinnules and fewer but perhaps more 

distinct marginal teeth. The nervation in Lesquereux's species is very much m?re crowded and less 

distinct than in the specimens under consideration. 

From the specimens supposed to be identical with 

Lesquereux's species. but recorded under the name 

Anemia perplema Hollick 64 n1y material differs in the 

more pronounced pillnules with the n1argins simply 

instead of doubly toothed, and the nervation is very 

much sparser .and lighter. 

Occurrence: Middle Park formation, doleritic breccia just above main breccia mass at Potato Hill G'ap, 

on the Grand [Colorado] River near Hot Sulphur 

63 The Tertiary flora, p. 59, pI. 5, figs, ~-3, 1878. 

G4 Hollick, Arthur, in Newbeny, J. S., U. S, Geol. Survey Mon. 

35, pI. 15, figs. I, la, 1898. 
6U FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Springs, ~1iddle Park, Grand County, Colo., collected 

by George L. Cannon, 1889. 

Anemia sp. 

Plate 8, Figure 7 

It is with considerable hesitation that I describe 

this form at all, as it is so fragmentary that its full 

character can not be made out. It has the pinnae 

linenr-Ianceolate and slightly decurrent at the base, 

with the margin perfectly entire or only slightly undulate in parts. The midvein is rather strong, and 

the finer nervation is plainly marked though rather 

slender. The veins arise at an acute angle and fork 

once or occasionally twice near the middle and thence 

pass up for long distances to terminate in the margin. 

This form seems to approach most closelv what has 

been called Anemia elongata (Newberry) Knowlton 65 

. (Anemia peTpleaJa Hollick) ,66 from which it differs 

mainly in the entire nlargins and more spreading finer 

nervation. It is also suggestive of what has been 

called Pten8 p8eudopennaJeformis Lesquereux,67 from 

the Denver formation, which differs in having a bigrooved rachis, dentate margins, and mainly simple 

veIns. 

Occurrence: Middle Park formation, near Hot Sulphur Springs, Middle Park, Grand County, Colo., 

collected by G. L. Cannon, 1889. 

Lygodium coloradense Knowlton, n. sp. 

Plate 8, Figure 8 

Pinnae orbicular in general outline, with a deep, 

rather bro~d sinus at base, deeply palmately cut, with 

about seven oblong-Ianceolate, rather obtuse lobes; 

petiole slender, at least 3 centiineters long; nervation 

as in the living L. pal17wltu1TiJ but proportionately 

rather coarser. 

This species is represented by the fairly . perfect 

specimen figured and by a number of fraO'ments of • 0 

pinllule lobes. It is of the general type of the livinO' 

Lygodiwm pal17~atu17L of the eastern United States bld 

is very mnch larger, the diameter of the pinnule being 

fully 11 centimeters in the fossil forpl' and· only 5 

centimeters in. the living species. The base is deeply 

heart-shaped·, and the lobes oblong-Ianceolate, with the 

upper· ones separated to a point within 1.5 centimeters 

of the top of the petiole. The basal lobes are only 

about 1.5 centime~el~s long, but the central one is more 

than.6 centimeters long. The principal nervation is 

ro:ughly. dichotomous from the top of the petiolethat is, the petiole divides into two equal branches, 

which within a. distance of 2 or 3 millimeters aO'ain o 

fork to supply the large upper lobes, while the basal 

lobes are supplied by a fork from the lower of the 

larger ones. The fine nervation is of exactly the same 

type as in L. palmaturn--that is, the' nerves arise fr0111 

the midnerve at an acute angle and fork usually three 

times before reaching the margin. Fertile fronds 

were not found. 

Of the 25 or more living species of Lygodiu·m, some 

6 or 7 belong to the so-called pal17Lata group, in whi'ch 

the nervation is palmate, or more properly dichotomous, as typified by the little' climbing fern (L. dJiohot01nrWln) Swartz) of eastern North America and L. 

'J'adiatwni Prantl, of tropical America. The genus is 

very widely distributed and is essentially tropical. 

The nominal fossil species of Lygodi1.(jfJ1 also number about 25, but there is more or less doubt about the 

authenticity of certain of the forms from the older 

horizons. As they stand at present in the books, 2 

occur in the" Upper Carboniferous," 1 in the" JuraTrias," 2 in the Upper Cretaceous, 12 in the Eocene 1 

in the Oligocene, and 7 in the l\Iiocene; so far as tIle 

record shows none have been found in either Pliocene 

or Pleistocene. 

The Paleozoic forms appear to be particularly open 

to question. Thus, Lygodiu1n 81nilacifoliul11J (Sternberg) Ettingshausen, fr0111 the" Upper Carboniferous" of Saxony, was first described by Schlotheim in 

1820 as a Phyllites. Two years later SternberO' transferred it to the genus OsmunfkD, in 1825 the same 

author placed it in Neu1'opterw, and . iIi 1865 EttinO'shausen again ~ransferred it; placing it in Lyqodi1.:n. 

It has the pinnate venation of the type of the livinO' 

L. scandens, but i't appears to me to be neuropteroid i~ its affinity and not lygodioid. 

The other supposed Paleozoic species is L. stacltei· 

Stur, from the" Upper Carboniferous" of the central 

Alps. The question of its status was submitted to 

:Mr. David 'Vhite, who writes: " The specimen figured: 

· by I{erner is almost certainly a heteromorphous 

· pinnule of a Neuropteris of the N. 8cheu·chz.eri type." 

The lower Mesozoic species, Lygodiu1n? anti-

, quorul1~ Shirley,68 fr01n the" Trias-Jura" of Queensland; is based on' a very small, supposedly fl~uitinO' 

pjnnule. It is not I descrIbed but· is dIscussed a~ follows: . 

A small fragment, showing a tripartite lobule of a frond 

· surrounded by the characteristic plaited sori of the order: 

: so familiar· ·to Queenslanders -through· the very common L. 

8oanaen8 of our present southern flora and E. ret'iOtftZat1t1n and 

L. jo,ponic·ltm of our northern forests. 

68 ·Shirley, John, Additions to the fossil flora of Queensland: Queensland Geol. Survey BUll. 7, p. 17, fig. 3, 1898. 
SYSTEMATIC PALEOBOTANY 31 

The figure is a mere outline that in itself is hardly 

convincing. 

Of the three supposed Cretaceous forms two are 

American and the other European. Of these, L. 

t'l'ichom,anoides Lesquereux,09 from the Dakota sandstone of I(ansas, is a mere fragment without biologic 

value; in fact, ~esquereux himself stated that although 

he referred it to Lygodium he did not know any species 

of this genus that it resembled. It is far too doubtful 

to be of value in making out the phylogeny of 

Lygodiwn. 

The other American Cretaceous species is L. COrnpactu'In Lesquereux,7o frOID the true Laramie at Marshall, Colo. It is a. single small lobe, supposed to be 

tt lobe of a pinnule of the pabnatu1n type, but is too 

small and obscure to be of convincing valu~. 

LY,qoitiurn O'f'etaoewn Debey and Ettingshaw;en,71 

the European Cretaceous form, is from the Senonian' 

of Aachen, Prussia. The form of the frond, the nervation, and above all the fruit, which is fairly well preserved, agree in general with the L. palmaturn type, 

and hence this species can probably be accepted as the 

oldest known representative of the genus. 

By Tertiary time, however, Lygodiwm had become 

firmly established and widely spread in both hemispheres, as attested by fronds that no one would hesitate to pronounce congeneric with L. p'fil;m;o;twm, as 

well as hyfruit of undoubted identity. As the oldest 

of these (L. gooseleti Fritel, from the" Paleocene" 

of France) is still unfigured, its probable status can 

11 ot be discussed. 

The two forms from the Paris Basin referred by 

Watelet 72 to L'!I!JodJiwm (L. C'rassioostatwm and L. oapillare) appear very doubtful. They are long, lanceolateacuminate fossils with a pinnate and conspicuously 

reticl.Jlate nervation quite unlike the nervation in Lygodium, and they should apparently be excluded from 

this genus. 

In 1872 Saporta 73 described two species (L. pa'f'vifoliu1n and L. ewqwisitwnv) from the Eocene of the 

southeast of France that although without fruit' exhibit the outline and nervation characteristic of Lygodi'l1tm. Later Saporta 74 added two more species to 

this flora (L. tenellwnv and L. dist'f'aotwm) , which confirmed the earlier reference by the discovery of undoubted fruit. 

60 Lcsqucrcux, Leo, The Cretaceous flora, p. 45, pI. 1, fig. 2, 1874. 

70 Lcsqucl"c\Jx, Leo, The '.rertiary flora, p. 64, pI. 5, fig. 9, 1878. 

"Il Debey, M. H., nnd Ettlngshunsen, Constantin von, UrweItl1chen 

Acrobryen des Kreidegeberges von Aacben und Maestrlcbt: K. Akad. 

Wlssensch., Math.-nnturw. CIa sse, vol. 17, p. 18, pI. 2, figs. 18-21; 

pI. 3, fig. 28, 1859. 

7n Watelct, Adolphe, Description des pluntcs fossiles du bassin de 

Parls, p. 49, pI. 13, figs. 2-4, 1866. . 

73 Snporta, Gaston de, :IDtudes sur la v~g~tation du sud-est de In 

I!"rnnce i\. 1't~llo(Jue tertiaire: Annales scI.' nat., 5th ser., vol. 17, p. 87, 

pI. 1, figs. 13, 14, 1.873. 

7' Snporta, Gaston de, Flore fosslIe d'Aix-en-Provence, pp. 23, 24, 

pI. 2, figs. 7, 8, 1888. 

. It is hardly necessary to review the Miocene forms 

that have been referred to Lygodiwn, though mention 

may be .made of the several species described by 

Heer 75 from the Swiss Miocene. These are of the 

type of the living L. ai'f'mnatJum (Burmann) Swartz, 

of tropical Asia and Queensland, with which some 

of them agree closely in form, 'nervation, and fructification. 

We may now return to the consideration_ of the 

Colorado form here described. As already pointed 

out, it agrees most closely with the living L.· pa"lrlwturn, from which it differs in its much larger size, 

longer, narrower lobes, and slightly coarser nervation. 

Among American Tertiary species it approaches nearest Lygodium kaulfussii Heer,76 but it differs from 

that species in the generally smaller size, deeply heartshaped base, dichotomous forking of the main nerves, 

and somewhat more open nervation. . 

Occurrence: Dawson arkose, at Ramah oil well, 

NW. ~ sec. 33, T. 9 S., R. 61 W., 40 miles northeast 

of Colorado Springs, Colo., collected by C. W. Cooke 

for G. B. Richardson, September 7, 1910. 

Order EQUISETALES 

Family EQUISETACEAE 

Equisetum coloradense Knowlton, n. sp. 

Plate 8, Figures 1-6 

Stems of fairly robust size, 12 to 20 millimeters 

wide, usually strongly ribbed and furrowed, the ribs 

9 to· 12 on a side; length of internodes from two or 

three to several times the diameter of the stem; sheath 

short, with about 24 sharp-pointed, closely appressed 

teeth. 

All the known examples belonging· to thi.s form 

ha ve been figured. In the best-preserved stem from 

Middle Park (pI. 8, fig. 4) there appear to be about 

nine strong ribs and furrows, but in the other example (fig. 1) the stem is so poorly preserved that it is 

impossible to make out the furrows. The sheath in 

these specimens, best· shown in Figure 2, is about 5 

millimeters wide and provided with 24 short, sharppointed teeth-that is to say, about 12 teeth can be 

seen on the surface exposed. 

The specimens from Mosby are shown in Figures 

3, 5, and 6. The matrix in which they are preserved 

is a fine-grained sandy clay which has retained the 

characters with fidelity. The largest specimen is 9 

centimeters long and 1.5 centimeters wide and compressed to a thickness of about 3 millimeters. It 

shows only one sheath with its teeth. The ribs take ~ 

75 Heel', Oswald, Flora terUada Helvetiae, vol. 1, pp. 41-43, pI. ·13, 

figs. 1-15, 1855. 

76 cr. Newberry, J. S., The later extinct floras or North America: 

U. S. GeoI. Survey 'Mon. 35, p. 1, pI. 62, figs. 1-4, 1898. 
32 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

their origin at the line of the sheath and ·are double-:-

that is, each tooth separates a wedge-shaped ridge 

which is composed of two appressed ribs at this point. 

These ribs soon sepa.rate, and within a distance of 

2 or 3 centimeters they have spreaod so as to make double the number of ribs that arose at the sheath. The 

interval between the ribs is filled with fine striae. The 

teeth are triangular and about 2 millimeters long; 

they are closely appressed and fill the space between 

the wedge-shaped points of the ribs. It is possible 

that this Mosby specimen is not the same species as 

the specimens from Middle Park, for as well as can 

be made out it exhibits features not present in the 

others; but the general appearance is the same, though 

this may be due to the poor preservation of the Middle 

Park specimens. If material is subsequently found in 

s~flicient quantity to be sure of the characters they can 

easily be separated if not found to be identical. 

The specimen from Mosby, shown in Figure 3, consists of a short segment of a stem about 2 centimeters 

wide, showing the node as in the one first described. 

Lying parallel to and partly on the larger stem is a 

smaller one about 8 millimeters in diameter' that appears to be a branch of the larger one. It does not 

otherwise exhibit any features not present in the specimen first described. 

So far as size goes E quisetu1n colO1~adense seems to 

approach E. haydenrii Lesquereux,71 from the Hanna 

formation of Carbon, Wyo., but as E. haJ!ffknii is 

founded on underground portions of the stem with no 

sheaths showing, it is impossible to make any comparison except in size. 

The closest relative of the form in hand appears to 

be Equisetwn 1'ob'ltst'lt1n Newberry,78 from the" Puget 

Sound group~' of Bellingham Bay, Wash., which: 

indeed, is hardly to be distinguished from it. The 

main points of' difference are the larger size of the 

Washington form and the longer internodes, shorter 

sheaths, and less numerously furrowed stems in the 

Colorado specimens. 

Occurrence: Middle Park fonnation, high terrace 

ridge between forks of IGnney Creek and in "breccia 

spoon," east of Hot Sulphur Springs, Middle Park, 

Grand County, Colo., collected by 'Vhitman Cross, 

October 6, 1891. Da wson arkose, dump of Mosby coal 

mine, Mosby, Colo., collected by F. H. I{nowlton and 

G. B. Richardson, July, 1910. 

Equisetum sp. 

Plate 9, Figure 4 

Among the extensive material from Table ~fountain, 

\ Golden, Colo., I find the specimen here figured, which 

represents a considerable fragment froln the subter77 Lesquereux, Leo, The Tertiary flora, p. 67, pl. 6, figs. 2-4, 1878. 

.s NewbC'l'ry, .T. S., U. S. Geol. Survey Mon. 35, p. 15, pl. 16, figs. 1, 2, 

lS!l8. 

ranean portion of an E q1Jli8etu171l. The stem, the transverse section of which shows as a ring in the center, 

is only about 7 millimeters in diameter, and from it 

radiate some six or seven thickened, constricted tuberous portions, which are not well enough preserved to 

admit of much closer definition. 

The underground portion of E q'uisetWln is so generally without essential characters in all forms of the 

. genus that no attempt has beE:n made either to identify 

this specimen with other so-called species based on 

s}milar fragments or to characterize it in a manner 

worthy of a specific name. It is simply a fragment 

of E qUruetu1n and is presented only to call attention 

to the presence of the genus in these beds. 

Occurrence: Denver formation, South Table Mountain, Gold~n, Co.1o. 

Equisetum sp. 

Physagenia sp. Lesquereux, Haryard ColI. Mus. Compo Zoology 

Bull., vol. 16, p. 43, 1888. 

This form was described by Lesquereux as follows: 

Tubercles attached to filaments diverging in rows from a 

central point, composing the rhizoma of some Equisetacelle. 

Central point exactly round, 2 millimeters in diameter; tubercles oval, 12 millimeters long, 6 millimeters broad in the middle, 

strangled to 2 millimeters at the point of union, and forming a 

chain of which two of the tubercles are seen in close connection. They are deeply, irregularly wrinkled lengthwise. 

It is with some hesitation that this form is admitted. I have examined all t.he supposed types in the 

Museum of Comparative Zoology and could find only 

one specimen (No. 298) that appears to be equisetaceous in ~haracter. This is a specimen showing two or 

three tubercles attached to a central point but so 

poorly preserved that it can not be figured. The re-

. mainder of the so-called types appear to be fragmentary stems or rachIses of ferns, pieces of bark, 

etc., and of course they are without value in the present connection. 

The present form is permitted to stand. as aboye 

desGribed simply to call attention to the fact that 

probably some sort of an equisetaceous plant was living during Denver time, but its size and aerial charac· 

tel'S must remain for future discovery. It may be the 

same as the preceding forln, but the characters are so 

obscure that this can not be ascertained. 

Occurrence: Denver formation, Golden, Colo., collected by Arthur Lakes and deposited in the ~fuseum 

of Comparative Zoology, Cambridge, ~fass. 

Equisetmn perlacvI ga tum Cockerell, 'Vest Am. Scientist, vol. 

6, No. 49, p. 154, 1889. 

Equisetunz. laev·igatunz. Lesquereux, U. S. Geol. and Geog. Survey Terr. AUD. Rept. for. 1873, p. 395, 1874; The Tertiary flora, p. 68, Pl.' 6, fig. 7, 1878. 

One of the type specimens of this supposed species 

came from Sand Creek, Colo., and thus :falls within 
SYSTEMATIC PALEOBOTANY 33 

the scope of the present work, but an examination of 

this specimen shows it to be very obscure; in fact, it 

has more the appearance of having been a piece of 

dicotyledonous bark or the impression of a stem. It 

has the surface wrinkled irregularly rather than striat ely , and the so-called tubercles can hardly be made 

out at all. It is of no value and is excluded from 

systematic enumeration. 

Ordel' LYCOPODIALES 

Family SELAGINELLACEAE 

Selaginella berthoudi Lesquereux 

SelauineUa, be1·thoudi Losquereux, U. S. Goui. and Geog. Survey 

Torr. Ann. Rept. for 1873, p. 395, 1874; idem for 1876, p. 

499, 1878; Tho Tertiary flora, p. 46, pI. 5, figs. 12, 12u, 

1878; Harvard ColI. Mus. Compo Zoology Bull., vol. 16, 

p. 43, 1888. 

Type: United States National Museum, No. 35. 

This fine species appears to be rare, as thus far only 

four specimens have come to light. These include 

the type specimen now in the National Museum, two 

specimens recorded by Lesquereux in the collection of 

the Harvard College Museum of Comparative Zoology, 

and one in the' Hambach collection recently acquired 

by the National Museum. These specimens are all recorded as having come from Golden, Colo., but are 

without precise locality. From the character of the 

matrix, however, it is thought that they. came from 

South Table Mountain. 

This species is described by Lesquereux as havi'np; a 

large diffuse frond which is either creeping or flat~ 

tened on the ground .. It has dichotomous branches 

with two kinds of leaves in four rows; the larger lateralleaves are open, distichous, 3 to 4 millimeters long 

and 1 millimeter broad and imbricated by the lower 

side. They are oblong-lanceolate, acute, sessile, and 

distinctly nei'ved in the middle. The intermediate 

lea ves are small, being scarcely 1 millimeter long 

a,nd 0.5 millimeter broad, oval, obtusely pointed, alid 

marked by a middle nerve. 

This is a perfectly distinct and well-marked species 

and resembles, as Lesquer'eux pointed out, the living 

Selaginella '11w1'tensii of ~Iexico and S. stolonifera of 

the West Indies. 

Since the above account was written two additional 

specimens have come to light. They were collected in 

the Denver formation on Sand Creek near Magnolia, 

just northeast of DenveL'. They are not very well preserved, and the small intermediate leaves are made out 

with difHculty. 

Occurrence: Denver fonnation, Golden, Colo., 

probably South Table ~{ountain; type specimen colIt,('t<~d a.bout 1873 by Capt. Ed. Berthoud, in whose 

honor the species was named; specimens in the Museum of Comparative Zoology collected by Althur 

Lakes in 1883. The lately acquired specimens (U. S. 

Nat. ~{us. No. 33891) were procured from GU'stav 

Hambach, Sanel Creek near Magnolia, northeast of 

Denver, Colo., collected by Francis W. Collins, 1909. 

Phylum SPERMATOPHYTA 

Class GYMNOSPERMAE 

Order CONIFERALES 

Family PINACEAE 

Sequoia atfinis Lesquereux 

Seqtwia, a,ffirl'i8 Lesquereux, U. S. Geo1. and Geog. Surr~y Terr. 

Bull., vol. 1, p. 384, 1875 [1876]; Ann. Rept. fo\' 1874, 

p. 310, 1876; The Tertiary flora, p. 75, pI. 7, flg~, 3·-f), 

1878; The Cretaceous and Tertiary floras, p. 138, 1883, 

Knowlton, U. S. Nnt. Mus. Proc., YOl. 51. p. 248, 1916. 

Scqnoia la'l/us(lor/ii Brongniat·t. Lesquereux, The Te\'tinr~' 

flora, p. 76, 1878. 

GlytJtostl'obu,8 unger'i,? Heel'. Lesquereux, The Cretaceous and 

Tertiary tlOl'US, p. 139, pI. 23, figs. 1-6a, 1883. 

Following is Lesquereux's original description of 

this species: 

Branches long, slender, pinnately branching; leaves ShOI'l', 

oblong, imbricated and obtuse, or longe\', ianceo\ate-acute, erect 

(II' slightly reflexed; branchlets bearing cones open; strobiles 

small, round-oval, obtuse; scales large, rhombOidal, with entire 

borders, a central oval mamilla, and wrinkles passing from it 

to the borders aI1 around; male branches erect, with more acute 

and open leaves, resembling sterile branches of Glyptost1'oblts 

CttrOpaet&8, with small, r~und catkins, covered at the top by, 

imbricated lalleeolate i('aves. 

In his (.u::;\'us;:,ion of t.his species in the report on the 

Tertiary flora (p. 75) Lesquereux stated that the leaves 

are" decurrent, without trace of a middle nerve," but 

(l,n examination of one of the figured types shows that 

the leaves are provided with a clear, fairly prominent 

middle vein. With this exception the: diagnosis appears to have been reasonably well drawn. 

An examination of the history of this species shows 

that it was first described from Middle Park, Colo., 

and was later found at Costello's ranch (Florissant), 

South Park, Colo., and at Elko station, Nev. The 

specimens from the original locality do not seenl to 

be preserved in the United States National Museum, 

nor have I been able to find the specimens said to have 

come from Elko station. Both of the. i::ipecimens figured on Plate 65, Figures 2 and 4, of the report on the 

Tertiary flora are in the National ~{useum (Nos. 87: 

88) and came from Florissant, Colo. 

In the recent material from Middle Park I find a 

number of branchlets that appear to belong to this 

species. They are very slender branchlets with, rather 

acute, one-nerved ~eaves, which correspond closely 

with the slenderer branchlets on the type specimen. 
34 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORAlJO 

The question of the probable relationship of this 

species was comIItented upon by Lesquereux,19 but as 

the material is so scanty it. is perhaps best to defer 

'final judgment until more can be obtained. 

. Occurrence: Middle Park formation (type lost), 

Middle Park, Colo., collected by F. V. Hayden; Mount 

Bross, north side of Grand [Colorado] River oppo8ite Hot Sulphur Springs, Middle. Park, Colo .• collected by G. L. Cannon, 1886. l\1iocene, Florissant, 

Colo. Miocene ( ~), Elko, Nev. 

Order GINKGOALES' 

Family GINKGOACEAE' 

Ginkgo? truncata (Lesquereux) Knowlton, .~. comb. 

Plate 9, Figure 3 

Palrnocarpol'll truMxlltum minor Lesquereux, The Tertiary flora, 

p. 120, pI. 11, figs. 8, 9" 1878. ' 

Oarpite.r; la·u,rineu8 Lesquereux, idem, p. 304, pI. 60, figs. 20, 21. 

Fruits nearly spherical, averaging about 9 to 12 

millimeters in diameter, sometimes slightly flattened; 

testa fairly rigid, hard, thin, obsc~rely bicarinat~, 

seated in a fleshy cup-shaped excavatIOn at the summIt 

of a stout pedu~cle, which is ahout 1 millimeter in 

diameter and 2 centimeters long:. 

Lesquereux based his PalmO'earrpon trun?atum on 

four figured specimens, which he said mIght w.ell 

enough represent two distinct species if based on SIze 

alone. He added: "As their characters are, however, 

the same, and as the difference in size may result fr~m 

the relative position upon racemes of the sa me spe(,I~s 

but of different age, I have considereu Lhe111 as varIeties: var. major and val'. minor." The p~esent discussion is concerned only with the' variety minor. , 

All the type specimens are in the United States 

National Museum. A cQmplication arises concerning 

the specimens supposed to represent the variety :ninor 

in the Museum collection. There are four speCImens, 

, all detached and all bearing the same number (517), 

two of which are preserved in the soft andesitic matrlx 

characteristic of the Denver beds at Golden, Colo., and 

the other two in a hard ironstone matrix wholly unlike 

that at Golden and hence from an unknown locality. 

The figures given in the report on the Tertiary flora 

are so conventionalized that it is impossible to decide 

which of the four specimens served as the originals 

from which the fi<Yures were mad('. This much at least o . 

can be said: The two original speci~ens preserved In 

the andesitic matrix are undoubtedly the same as 

numerous additional examples from Golden collected 

by Arthur Lakes. All the specimens, except one 'of 

the later ones, are free and unattached; in fact, none 

have been observed that give any i,ndication of a point 

~o Lesquercux, Leo, The Tertiary flora, p. 75, 187.8. 

of attachment. The exception is the specimen shown 

ill Plate 9, Figure 3, which is believed to offer important evidence as to its possible affinity .. 

When Lesquereux founded his Pailrnocarpon t~'ttncaJtum, he compared it to the fruits of Sa~al, espeCIally 

Sabal 1newicana Martins. It must be confessed that 

these O'lobular fruits when found isolated have a 

strono' 0 

deO'ree of resemblance to the fruits of several 

gener~ of ~alms, such as Oreodo~a, S(J)bal, and Thn,1UlW. 

In all these, however, the frUIts are ne~rly sessllethat is, the supporting stalk or peduncle IS very short. 

In the specimen here figured the peduncle is fully 2 

centilneters long, and moreover the fruit appears to be 

seated in a cup-shaped expansion at its summit. This 

,vould seem to exclude it from the palms, or at least 

fronl those with which I am familiar, and its affinity 

must be souO'ht in other directions. The requirements b • 

seem to be met in Ginkgo, to which consequently It 

has been referred. 

The flowers of Ginkgo bi~ob(J) Linne, the only living 

species of the genus, are dioecious and are disposed on 

short lateral shoots; The male flowers are arranged 

in the form of a solitary, filiform, axillary catkin. 

The female flowers are placed among the ordinary 

leaves and consist of a petiole with a 2-cleft or rarely 

3-6-cleft apex, each point of which supports an ovule 

instead of a 'blade of the leaf. The ovules are seated 

in a shallow, fleshy cup formed by the dilate~ ap~x of 

the petiole and consist of a bicarinate or trICarln~te 

drupaceous 'seed with a thin outer" flesh" and a tlun, 

hard shell surrounding a fleshy embryo. It frequently 

happens that all but one of the ovules abort, so tl~at 

when mature the ovule appears solitary, though the 

rudiments of the blasted ovules may usually be detected, by the side of or below it. , At ~aturity .the 

ovule is nearly spherical, with a nunute apleal 

projection. 

On account of their exceedingly hard shell and resinous nature, the fruits of Ginkgo SeeID especially well 

fitt~d for preservation in a fossil state, and as a matter 

of fact a considerable number of species have been 

found in association with undoubted leaves of Ginkgo. 

Thus, Ginkgo d'igitata (Brongniart) Heer, from the 

Jurassic of Cape Boheman, Spitzbergen~ has oval 

fruits terminating in a point as in the living species 

and was seemingly covered with a fleshy exocarp. 

Ginkgo sibmicaJ Heer, from Ust-Balei, eastern Sibed:l, 

has small globular fruits borne in .twos at the sununlt 

of a slender peduncle, and G. p11imwrdialis Heel', of the 

Atane beds of Greenland, is described as having oval 

fruits inserted in a pronounced cup-form dilation of 

the oraliferous leaf. 

The fruits under discussion from Golden appear to 

meet the requirements of a. reference to Ginkgo., although no leaves belonging to this type have thus ,far 
SYSTEMATIC PALEOBOTANY 35 

been detected in these beds, and for this reason the 

O'eneric reference has been questioned. Lea,Yes of a b , 

number of species of Ginlego occur more or less abundantly in American rocks ranging in age from J urassic to Tertiary, but so far as I know this is the first 

fruit to be adequately described. As nlay be seen from 

the figure of the best specimen, it is excellently well 

preserved. It shows the fruiting peduncle with the 

cup-shaped ctlvity at the apex, in which is seated the 

small, round, slightly bicarinate fruit. There is no 

visible trace of an aborted ovule, which should normally be' present, nor are there remains of a fleshy 

outer integument. , . 

As already mentioned there are numerous detached 

fruits fonnd in assoeiation with the one just described 

that undoubtedly belong with it. They are uniformly 

spherical or slightly compressed and are obscurely 

bicarinate or tricarinate. One of the best preserved 

of these detached fruits was covered when removed 

from the nlatrix with a thin film of carbonaceous matter, which possibly represented the' fleshy . exterior. 

Anothei' is minutely. and irregularly pitted, suggestinO' that the flesh had dried before the entombment of 

th~ specimen. Still anothe~' specimen has been cracked 

into two nearly equal parts, just as might result when 

a globular fruit with a hard, brittle shell is compressed. . . . 

It seems necessary to refer to Ginkgo? truncata the 

little fruits described by Lesquereux under the name 

Oairpites laurineus.so These fruits; both on one piece 

of matI-ix, are present in the United States National 

:Museum (No. 516): They were said by Lesquereu~ 

to have come from Evanston, 'Vyo., but in the ~1useum 

re.gister they are recorded as coming' from Golden, 

Colo. This . record is undoubtedly correct, for the 

matrix is readily recognizable as, that of the andesitic 

beels nt Golden ~,nd wholly unlike anything known 

from Evanston. In his description of Oarpites laltrine1.l-S Lesquereux wrote as follows: "Berries small, 

nearly round, short-pointed, about 5 millimeters in 

diameter surrounded by a thin shelly epicarp." On , . . comparing the specimens it appears that tIus descrIption must have been drawn by Lesqueteux from the 

figures rather than the specimens. Both fruits are 

broken and not correctly represented by the figures. 

They undoubtedly belong with Ginkgo? tlJ"Uncata. 

Occurrence: Denver formation (types) " Golden,. 

Colo., original collections probably made by Lesqu.eI'ellX about 1875: also collected by Arthur Lakes In 

1890, on South ;.rable ~~[ountain, Golden, at a point 

about 100 feet below the lava cap. 

~Il Lesql1el'Nlx, Leo, The Tertiary flora, p. 304, pI. 60, figs. 20, 21, 

18iK 

Class ANGIOSPERMAE 

Subclass MONOCOTYLEDONES 

Order G RAM IN ALES 

Family GRAMINEAE 

Arundo? obtusa Lesquereux 

An/,ndof obt,lts(/' Lesquereux, U. S. Geol. uud Geog. Survf.:'j' 

Terr. Bull., vol. 1, No.5, 2d ser., p. 385, 1875 [1876]; 

Ann. Rept. for 1874, p. 311, 1876; The Tertiary flora, 

p. 87, pI. 8, figs. 9-9c, 1878. 

So far as can be ascertained, the type specimen: of 

this species has never been' in the collection of the 

United States National Museum, where it should be, 

and its present location is unknown. N 0 addition~l 

examples have beeli obtained, and the status of tlus 

species must be regarded as uncertain. It was described at some length by Lesquereux, but there is no -

evidence other than association on the same rock to 

show that the organisms described as seeds ~nd their 

envelopes reaUy belong to the same plant. 

Occurrence: Denver formation, .South Table :i\lountain, Golden, Colo. 

Family CYPERACEAE 

Carex? berthoudi Lesquereux 

OO!r6(J) bertho'ucU Lesquereux, U. S. Geol., and Geog. Surrey 

Terr. Ann. Rept. for 1872, p. 377, 1873; idem for 1873, 

p. 380: 1874; idem for 1876, p. 500, 1878; The Tertiary 

flora, p. 92, pI. 9, figs. 3, 4, 1878. 

Type: United States Natiomil Museum, No. 98. 

What purport to be the type specimens of this 

species are preserved in the United States National 

Museum (No. 98; three specimens under the same 

number) but after a careful comparison I am not able 

to identi'fy any of them with the figures given by 

Lesquereux. There are a number of fragments of 

sedO'elike or (yrasslike leaves 3 or 4 centimeters lon.(! b b . 

and 3 or 4 millimeters wide, that appear to fit the 

description fairly well, but the so~called seeds 

(perigynia) I am not able to find at all. Th~se have 

either been effaced since the species was establIshed, or 

more probably the wrong specimen was marked as the 

type. This form does not appear to have been contained in any of the' more recent collections from 

Golden. and in fact it is doubtful if it should be continued 'under the name Oa'f'ew. About all that can 

be said concerning it is that it resembles sedgelike 

leaves that are usually described under the name 

Oype·raeites. 

In a table in the annual report of the Hayden sur- . 

vey for 1872 (p. 411), showing the distribution of the 

fossil plants of the North American Tertiary, Lesquereux indicated the presence of Om'ew be'f'thoudi in 
36 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

the Raton :NIountains, but as the column for Golden 

in this table is next to the one for the Raton Mountains it is probable that the cross mark was placed 

there in error. This is the only known mention of its 

coming from the Raton Mountains, and it has not 

been detected in the extensive collections made there 

in recent years. 

, Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Capt. E. Berthoud in 

the early seventies. 

Ordel' ARECALES 

Family ARECACEAE 

Sabalites grayanus Lesquereux 

Plate 9, Figure 5 

Sabal? grayana Lesquereux, Am. Philos. Soc. Trans., vol. 13, 

p. 412, pI. 14, figs. 4-6, 1869. [Not Lesquereux, The 

Tertiary flora, pI. 12, 1878; Knowlton, U. S. GeoI. Survey BUll. 163, pI. 6; fig. 5, 1900.] 

SabaU,tes gm,y.a1WS (Lesquereux) Lesquereux, The Tertiary 

flora, p. 112, 1878. ' 

Berry, U. S. GeoI. Survey Prof. Paper 91, p. 177, pI. 12, 

figs. 1-3; pI. 14, fig. 1, 1916. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 288, 

1917 [1918]. 

Flabellaria eocenica Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1872, p. 391, 1873; idem for 1873, 

p. 380, 1874; idem for 1876, p. 502, 1878; The Tertiary 

flora, p. 111, pI. 13, figs. 1-3, 1878. 

Sabal communis Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 18'14, p. 311, 1876. 

Ji'labcUaria COl1trnunis Lesquereux, U. S. Geo1. and Geog. Survey 

Terr. Bull., vol. 1, No.5, 2d ser., p. 385, 1876. 

Sab(/,l inq16irenda Knowlton, U. S. Geo1. Survey Prof. Paper 

101, p. 288, pI. 56, 1917 [1918]. 

Snbalites tntctite1' Lesquereux, The Tertiary flora, p. 114, pI. 

11, fig. 3 [not fig. 31\ or the three fruits at the base of the 

leaf, which=Be1~rya racmJl.osa], 1878; U. S. Geo1. and 

Geog. Survey Terr. Ann. Rept. for 1876, p. 502, 1878. 

Flabellm'ia? tnwtiter Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1873, p. 396,1874. 

Sabal? eooenica (Lesquereux) Knowlton, U. S. Geo1. Survey 

BUll. 696, p. 557, 1919. 

After a good deal of consideration it has seemed best 

to describe the palm leaves here discussed under the 

name Sabalites grayanus and to refer to that species 

Sabalites eocenwa (Lesquereux), Sabalites fmctiler 

Lesquereux, and Sabal, inqui'renda J{nowlton. In 

order that the position taken may be made clear the 

following account is fairly full. ' 

Sabalite8 graymnu8 was established (as Sabal? 

(j1'ayana) by Lesquereux 81 in 1869, on material from 

the soft white (" Eolignitic") clay of Lafayette 

County, ~1iss., and although the type specimens are 

now apparently lost, the exact locality is thought by 

81 Lesqnerenx, Leo, Am. r'hilos. Soc. Trans., vol. 1.3, p. 41.2, pI. 14, figs, 

4-6, 1.8GD. 

Berry to be in the vicinity of Oxford, Miss. Adrlitional material was procured from the' supposed type 

locality by Berry, as well as from numerous localities 

of similar age in ~iississippi, Arkansas, Tennessee, 

and Texas. As Lesquereux's original description was 

more or less incomplete the new material has enabled 

Berry 82 to give a much more complete description, 

which reads as follows: 

Leaves of large size but mostly fragmentary. Estimated 

diameter in Some of the larger specimens, where nearly half 

the leaf, is preserved, as great as 1.3 meters. Most of the 

leaves are somewhat smaller than this, no doubt because the 

larger the leaves the more fragmentary they would be likely 

to become before fossilization. Petiole long and stout, unarmed, enlarged at the base of the leaf, and tapering into an 

extended and gradually narrowed acumen, which is not visible 

on the upper surface of the leaf, where the petiole is broadly 

rounded and a short and inconspicuous ligule is developed. 

From the manner of preservation and attitude of the rays on 

some of the specimens it is inferred that the acumen was recurved, as it is in the existing Sabal palmetto (Walter) Roemer 

and Schultes. Rays very numerous, about 100 in number, a 

few reduced basal ones on each side free, the remainder 

united for a variable distance above the base. Their dimensions and the relative thickness of tile venation are variable 

features dependent on the size of the leaves. The largest 

specimens seen have thick carinate stout-veined rays, 5 centimeters in maximum width. They increase in size from tbe 

base of the leaf upward, and individually they are narrow 

at their point of attachment, widening medially and becoming 

gradually narrowed into long acuminate tips. Venation characters variable, largely dependent on the size of the leaves 

and the condition of preservation of the epidermis in the fossil 

specimens. In well-preserved material there are four or five 

relatively thin intermediate veins. Between each pair of veins 

there are six to eight fine vein lets which are not visible except 

in well-preserved specimens. 

The above 'description of SabaUte8 grayOlnus may 

be compared with Lesquereux's original description of 

ll'labellaria eocenica"which reads as follows: 

Frond large, rays convex, semicylindrical toward the base, 

flattened in the upper part, diverging from the top of a broad 

rachis, distinctly nerved; primary nerves distant; intermediate 

'veins thin, close, averaging 10 in number; rachis trupcate on 

its upper face, rapidly narrowed to a point on its lower [face]. 

Flabella'ria eocenica was founded by Lesquereux 823. 

in 1872 on specimens from the Black Buttes coal group 

at Black Buttes, Wyo. The type specimens are preserved in the United States National Museum (Nos. 

110, 111, 112) and were well illustrated and described 

by Lesquereux. It was evidently a small or medium 

sized leaf, probably about 1 nleter in diameter. 

Lesquereux stated in the report on the Tertiary 

flora (p. 112) that his Flabellaria eocenic{() is abundant 

in the Denver formation at Golden, 0010., but the 

National ~1useUl~1 does not appear to have speci82 Berry, E. W., U. S. Geol. Snr\'ey Prof. Paper 91., p. 178, 1916. 

S2n Lesquereux, Leo, U. S. Gro!. allfl Geog. SIll'Yey Terr. Ann. Rept. for 

lS72, p. 391., 1.873, 
SYSTEMATIC PALEOBOTANY 37 

BIens of this forlll from Golden, nor has it been noted 

in the collections made by Lakes and others. 

On ngain comparing the types of Sabal inq'll.irenita 

Knowlton 83 wHh the types of Flabellarria eocenica' and 

with the available figures of Sabalites graY(JJJ1/llS I am 

convinced thn t they can not be separated .• :Nly original 

description of Salbal in1q'l.ti1'enda reads as follows: 

]~eaves relntirely smull, probably 1.5 meters OL' less in 

di:lllHlt·Ol', pnlmll te, with 60 to 80 folds, which are much compressed nt the point of attachment and rnpiclly widen out; 

l)eOolo ullUrll\ed, rounded both nbove nnd below, 01' at least 

\lot conspicnously concave nbove, about 2 01' 3 centimeters 

brond; Ullex of potiole rounded on the upper side and prolonged on the under side into a spine or "midrib" 6 or 8 

ccntimetel's ill length. 

On comparing these descriptions or, better, the sevcl'lllfigures, it seems practically impossible to draw 

any satisfactory line of separation, and consequently 

they have been' merged under the older name. 

LesqneL'eux and others identified Sabalites g'l'CfflJaJnUS 

at 11 number of w.idely separated points in Colorado, 

'Yyorning, nnd Vancouver Island, British Columbia, 

Lut, as Berry S'l ha.s well said, 

All t'hese determinations nre open to ycry gl'avo doubt, not 

only on nccount of the inadequacy of the muteriul but also 

been use of H1e n priori hnprobn bility of a single species ranging from the O,'etaceous l\Iontnna group to a horizon well 

nboye the \)use of the lDocene at such widely removed localities, in fhe Que 1H'('a nssocinted with a subtropical coastal 

Hoi'll, which suggest·s the existing flora of the 'Vest Indies and 

lIorthel'H South America which ntlnmced northward in the 

l\1issl~sippi embllyment . region, flnd jn the ",estel'll area associated with n. yery different type of flora. 

The correctness of this reasoning is shown-to mention only one instance-by the treatment of specimens from the l\1esnverde formation nt Point of 

Hocks, "TyO., referred by Lesq~leretlx 85 to his Sabalites f/1'aya/n'l.ts. These huye now' been removed under 

the name. Sabal 'J7wntana I{nowlton. 8G 

'rhe type specimens of what Lesquerenx 87 . called 

Savalites f'J'uctife1' are preserved in the United States 

Nutionnl l\~[usetlm (No. 107). The best-preserved 

one-the original of his Figure 3-8hows the distorted 

apex of what might be considered a. pinnate leaf but 

w.ithollt much doubt is .the under surface of n leaf 

such as SalvaUtes ,q'J'aya:n'l.ls. Lesqllereux regarded it 

H,:; It palmate lenf, writing of it as follows: 

Seems to be l'eferable to Sa·bol by the acuminate rachis, 

which \)ear~ 011 \)ot11 sides munerOllS l'nys enlnrgecl and diverg83 Knowlton, 1". n., n. S. Geol. Survey Prof. Paper 101, p. 288, pI. 

GG, 11).17 [l1)18]. 

81 Bcny, E. W., U. S. Ceo!. SUL'yey l'rof. I'npcL' fll, p. 177, 1f116. 

& LCf;quCl'cnx, Leo, ~J;lw Tertiary flora, p. 112, I)\. 12, figs. 1, 2, 

1878. 

60 Knowlton, li'. II., n. S. Ceo\. Survey PL·of. Paper 101, p. 253, pI. 

:-J2, Il!;. a, 1917 [11:118]. 

~7 Le~ql1cl·(,I1X. 1.1'0. The rrel-thl1'~' flor:l. p. ]]4, pI. n, fig'. 8 (in part), 

187R. . 

~tH!I::!·-·;~O--4 

ing upward. The rachis is not very distinct, but as far as 

can be seen it looks narrowed into a point, or with an acumen, along which very numerous rays are attacbed. 

'rhe fruits supposed by Lesquereux to belong to 

Sabalites fructife'Jo are found on the same piece of 

matrix, but there is no evidence whatever to support 

the view that they belong to S. f'J"'uctife'J'. They are 

fairly abundant in the same beds as the palm leaves, 

occurring eit.her as isolated fruits or in compound 

racemes. At present there is no conclusive evidence 

to show what plant produced t.hem. At first they were 

separated as LVyssa? 'J'aee1nosa, but as t.hey can hardly 

belong to LVyssa, they have been described as new under 

the name B errya 'I'aCe1nosa. (See p. 134.) 

Occurrence: Denver formation, Golden, Colo. Dawson arkose, Jimmy Camp Creek, 9 nliles east of Colo· 

rado Sprin~s, Colo. Black Buttes coal group, Black, 

Buttes, ,Yyo. 'Yilcox group, ~1ississippi, Arkansas~ 

and Texas. Lagrange formation, Tennessee, in beds 

of 'Yilcox age. Raton formation, Raton Tunnel, 6 

miles north of Raton, and Dillon Canyon, near BlospLurg, N. ~{ex. 

SabaI? ungeri (Lesquereux) Knowlton 

SabaZ cam'lJbelU Newberry (in part), Boston Jour, Nat. Hist., 

vol. 7; p. 515, 1863. 

S(tbal· C(~m.pbelU Newberry. Lesquereux, Illustrations of Oretaceous and ~rertiary plants, pI. 10, U. S. Geol. anel Geog. 

Survey ~~err., 1878. 

Oeonomlites 1['l1geri Lesquereux, 'rhe 'rertiary flora, p. 118, pl. 

11, fig. 2, 1878. 

Sa·bal g1"(tndifolia Newberry, U. S. Geol. Survey Mon. 35, p. 28, 

pI. 63, fig. 5 [possibly not pI. 25 11(-.1. 64, figs. 2, 2 a], 

1898. 

Sabnlr l.[,l1geri (Lesquereux) Knowlton, U. S. Geol. Survey 

Prof. Paper 101, p. 289, ·Vls. 57, 59, 1917 [1918]. 

Leaves large, probably 2.5 to 3 meters in diameter, 

with 70 to 90 folds; petiole 3 to 4.5 centimeters wide, 

Hat or slightly ridged above but without a keel above 

or below; margins of petiole unarmed, terminating on 

the upper side in nn arch from which the folds radiate, 

on the under side prolonged into a, spine or midrib 15 

to 20 centimeters or more in lengt.h. 

The above description is with some essential modificat.ions that given by Newberry for his S(tbal gTandifolia, of wbich he figured one specimen frOlll the 

Fishers Peak region, in southern Colorado. 'Yhethel' 

or not the other specimens figured and described by 

Newberry under this name are really conspecific is not 

here considered, but there ca.n be no doubt that his 

Fishers Peak specimen is identical with Lesquel.'enx's 

Geono'J71-ites 'I.mgeri, from the same region, and a.s Lesquereux's specific name has priority over the other 

that is the name it must bear. 

The fragment upon which Lesqucreu~ based his 

Geon.O'Jnites 'I.ln'fie'J'i (U. S. Nat. ~fllS. No. 106) came, 
38 FLORA OF THE DENVEH AND ASSOCIATED FORMATIONS OF COLORADO 

as is now.known, from the under side of a large leaf 

"and shows the spikelike prolongation of the petiole. 

He mistook this for the upper· side of the leaf and 

hence concluded that the shape of the perfect leaf 

was flabella to-pinnate, or broadly linear-Ianceolate. 

It was for this reason that he placed it in Geo!,-,01nites, 

aS1n the living genus Geono1TUJJ the leaves are somewhat of this character. 

Notwithstanding the fact that a comparatively large 

number of specimens of what is believed to be a single 

species of palm are now available for study it is extremely difficult to reach definite conclusions regarding 

many of them. On this point Newberry 88 had the 

following to say: 

In the great Humber of tbe remains" o.t= palms found in tbe 

'l'ertiary and Cretaceous rocks of tbe West-trunks, leaves, and 

"fruit-it has been very difficult to define distinqt species, and 

it is probable that many years will elapse before perfect order 

can be brought out of the present confusion. The species 

. under consideration may, however, be identified by the large 

size of its leaf, its plain unkeeled petiole drawn into a long 

acute spine on the under side, the very numerous folds, and 

tbe crowded, subequal nervation. 

'Yith these limitations in Inind, Sabal wngeri is 

thought to be one of the most abundant forms in the 

Raton formation of the Raton Mesa region. It occurs 

a bmidantly from one end of this region to the other, 

and at two localities it has been found "in the top 

of the un~lerlying Vermejo formation, thus apparently 

crossing the line between Cretaceous and Tertiary. 

In the Dawson arkose along Jimmy Camp Creek there 

are great numb~ of remains of palms, but they are so 

fragmentary, OlP'l'ather they break into pieces so small 

when excavated from the bank, that it is difficult to 

identify them with certain·ty. These specimens are 

absolutely indistinguishable from the type of this 

species and also from the large leaves figured in my 

paper on the flora of the Vermejo and ~aton formations.89 

A few words may be said regarding the probable 

affinity and interrelationships of Sabal.~ ungeri. The 

species from outside the Rocky l\fountain area to which 

the present form is most closely related is ,Sabalites 

grayanus (Lesquereux) Lesquel'ellx. This was described originally,90 under the name Sabal, from the 

" Eolignitic" nVilcox group) of l\tlis~issippi, and 

although a mere fragment is distinguished from 

Sabal? wll-geri with difficulty if at aU .. Subsequently 

leaves that have been identified as Sabalites grayanu8 

ha ve been reported from many localities ranging in 

bS Xewberry, .T. 8., The later extinct floras of North Amet'ica: U. S. 

G,'oI. Stuve.r .Mon. ~:5, p. 28, 1898. 

SIl Knowlton. I". H., U. S. GeoI. Survey Prof. PHper .10l, p. 289, 1)ls. 

m, 59, 1917 [Un8] .. 

no Le!';quel'enX, Lf·O. Am. Philos. Soc. T.·ans., vol. 13, }). 412, pI. 14, 

figs. 4-6, 1861:1. 

age from l\fontana to Eocene, but Inost of these are 

fragmentary, and it is extremely doubtful if they have 

been correctly referred to this species. 

The segment of a huge petiole frOIn the Denver fornlation near Golden, Colo., figured by Lesqnereux 91 

:1S Sabalites grayain,u8 is preserved in the United States 

National l\fuseum (No. 109). There is, however, no 

valid reason for referring it to this species. If this 

strongly keeled petiole could really be proved to belong to Sabalites g1'ayanus, the keel would in itself be 

a sufficient character to separate it frOI11 Sabal.fj fwnge1'i, 

in which there is no evidence' that' the peti.ole was 

ei ther grooved or keeled. 

In Iny "Flora of the Vennejo and Raton formations "92 I have made the following stateInent regarding the relationships of Sabal? ~lngeri: 

It undoubtedly approaches most closely to Sa·bal 'inq'lt-i'l'enda 

Knowlton, find unless specimens are of sufficient size and 

perfection to shOW well theiL· character they can not be separated with absolute certainty. Sabal inquiren.do, however, may 

l:e known in general by its small size--not exceeding 1.5 

meters in diameter-and its 40 to 50 instead of 70 to 90 

folds. Sab(l,l 'rugosa is smaller find bas more strongly ribbed 

and plica ted rays, which do not exceed 40 in number .. ' Sa,bal 

leei, also a large-leaved form, may. be distinguished from S. 

u.ngeri by its wedge-shaped base and its great number of very 

closely appressed, obliquely inserted rays. 

Occurrence: Raton formation (types), abundant 

and widely distributed in the Raton Mesa region of 

Colorado and New l\fexico. Dawson arkose, east bank 

. of Jimmy Camp Creek, 0.6 mile above Richfield 

Springs ranch house, 9 Iuiles east of Colorado Springs, 

Colo., collected by F. H. I{nowlton and 'V. T. Lee, 

July, 1910. Vel'lnejo formation, neal' top of formation, Ponil Canyon, N. l\tlex., and near Shumway, Colo. 

Geonomites tenuirachis Lesquereux 

Plate 6; Plate 9, Figures 1, 2, 6 

Geon01ri:ites ten'lti1'achis Lesquereux, The Tertiary fiol':1, p. 117; 

pI. 11, fig. 1, 1878. 

Knowlton, U. S. Geol. Survey Prof. Pfiper 101, p. 291, pI. 62, 

1917 [1918]. 

Flubell(t;ria long'irachis Unger. Lesquereux, U. S. Geol. and 

Geog. Survey Ten. Ann. Rept. for 1873, p. 396, 1874. 

Type: United States Natio'nall\fuseum No. 105. 

This species appears to be rare. The type and only 

example known to Lesquereux came from the Raton 

Mountains of New l\1exico and is a mere fragment 

from near the middle of a leaf of considern.ble size. 

This fragment is so small that the character of the leaf 

can not be made out satisfactorily. It is not improbable that if the complete leaf were known it would be 

01 Lcsquel'enx, Leo, The Tertiary flora, p. 112, pI.. 12, flgs. 1, 2, 1878. 

"2 KnowUoll, F. li., U. S. GeoI. Sur,ey PI·of. Paper 101, p. 290, 1917 

[1918]. 
SYSTEMATIC PALEOBOTANY 39 

found to be referable to some of the other species 

described from beds of this age. In the absence of 

conclusive evidence, however, it seems best to retaill 

this forn1 under the name given by Lesquereux. 

The collections from the Dawson arkose on Jimmy 

Camp Creek contuin a large number of remains of 

pulms, but they are so fragmentary that it is difficult 

to find detern1inable specimens. The fr,agment here 

figured is n, segment fron1 the middle of a leaf of con-

::.ic1erable size. It is not distinguishable' from the type 

specimen of Geon()]n'ites terIJUi1'achis and is referred 

to that species. 

Occurrence: Dawson arkose, about 0.6 mile north 

of Richfield Springs ranch, on east bank of Jimmy 

Cnmp Creek, 9 miles enst of Colorado Springs, Colo., 

collected by 'V. T. Lee and F. H. I(nowlton, July, 1910. 

Raton formation (type), Raton :Mountains, N. Mex., 

ridge ea.st of Yankee mine, near yankee, N. l\1ex. 

Geonomites't graminifolius Lesquereux 

(leonont'itf18 f//'(I,m:iwifoU1t8 Lesquereux, Ha I'vard ColI. Mus. Compo 

Zoology BulL, vol. 16, p. 44. 1888. 

Following is Lesquel'eux's description and discussion 

of this species: 

Bl'okeu Ol' separated rays of :1 palm, varying in width from 

[, to 10 mUlimeters, 1lI1uked in the middle by a I>road nerve, 

the borders thinly rcgulllrly striate by 10 to 15 thin veins 

sea rcely distinct even with a lens. 

l.'heso fragments may belong to leaves of OaZa1nops'is cla1/,oA, 

Lesquereux [now Ohaonaedm'ea M1tad] or to a species much 

liI~e it. One ]eaf in a better state of presel"Vation has the rays 

lIarrowed to n point of connection to a broad, thinly lineate 

1'11('h:s, united three to five together, either connate pf!.rt of 

tlleir length or disconnected to the bnse like the fl'agments 

desct'ibed ubove. l."hough the likeness to Oa,Za,rnops-is da(/'Ulli, 

figUl'pd and described in l.'mns. Am. PhiLos. Soc., vol. 13, 

p. 4,11, pI. XIV, figs. 1-3, is marked, I have nevel' seen the rays 

tli \'ide into nutTOw lnciniae as in this species, generally found 

ill small, nnrrow segments. 

I have studied the type specimens of this supposed 

form in the Harvard College ~~[useum of Comparative 

Zoology, but they are so fragmentary that it is impos-' 

sible to reach any final conclusion. They are simply 

·",hut Lesquereux described them as being-" broken 

01' separated rays of a palm "-and there is no means 

o.f knowing what the complete leaf was like. It should 

probably be merged with some other species or discllrded altogether, but I have finally decided to let it 

stand us established by Lesquereux in the hope that 

subsequent discoveries may resolve its status one way 

ot' the other. 

Occurrence: Denver formation, G;olden, Colo., types 

,in Museum of Comparative Zoology, Cambridge, 

~1nss., collected by Arthur Lakes. 

Geonornites goldianus (Lesquereux) Lesquereux 

Plate 10, Figure 2 

GeullOfnUcs goluictl/ws (Lesquereux) Lesquereux, The Tertiary 

fiora, p. 115, 1)1. I), fig. 9, 1878; U. S. GeDl. and Geog. 

Survey Ten. Ann. Rept. for 1876, p. 502, 1878; Harvard 

Coil. Mus. 'Compo Zoology Bull., vol. 16, p. 44, 1888. 

PaZnwoitcs goliUUII/lt8 Lesquereux, U. S. Geol. and Geog'. Sun'ey 

Terr. Ann. Rept. for 1874, p. 311, 1876. 

fPZabellm·'w. zinkeni Heel'. Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1872, p. 377, 1873; idem 

for 1873, p. 380, 1874; idem for 1876, p. 502, 1878; The 

l.~ertiary fiora, p. 110, pI. 9, figs. 6-8, 1878. 

The type and illustrated specimens as recorded under 

all the above names are in the United States National 

Museum collection§l and are before me. The specimen 

first called Prilnuwites goldianu;8 and later transferred 

to the genus Geonmnites (No. 103) is the best-preserved example. It remained unique for a long time, 

but the collection made by Lakes for the Museum of 

Comparative Zoology, Cambridge, and determined by 

Lesquereux contained 10 specimens. The recent collections include a few exa.mples, and I have also found 

in the National Museum collection another example 

(No. 2059) that seems properly referable to it. 

The type specimen is almost too small for carefill 

description, yet it can probably be l~ecognized in 

future. Following is Lesquereux's description: 

Leaves fiabellate; rays fiat and without carinae; joined in 

an acute angle of divergence and by their whole base to a 

narrow, linear, fiat rachis, with narrow furrows marking the 

line of separation; primary veins generally distinct, with 10 

intermediate veinlets. 

The exact size and shape of this leaf are not known, 

as it is found only in fragments, but it can perhaps 

be recognized by the slender rachis to which the narrow, apparently flat rays are joined by their whole 

bases, the angle being about 20°. Lesquereux's statement that the rays are" without carinae" is not quite 

correct, as may be seen from the specimen. The· 

carinae are undoubtedly present, but the whole leaf 

was apparently thin, and hence the carinae are not 

strongly apparent. 

The new specimen above referred to, shown in 

Plate 10, Figure 2, appears to belong to this sp'ecies. 

It is about 15 centimeters long and about 8 centimeters 

in extreme width and represents the distal extremity 

of the leaf. The rays are united to a point within 

some 10 centimeters of the periphery, where they split 

into long, narrowly acuminate segments, as in many 

living palms of this type. The rays are distinctly 

though not prominently keeled and are separated by 

'7 to 9 or 10 distinct, often very distinct veinlets. It is 

of course impossible to be absolutely certain that this 

specimen is identical with GeonomJites goZdimrw.ts, as 

it does not show the connection of the rays with the 
40 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

rachis, but the general appearance, number of intermediate veinlets, and other features leadl11e to believe 

that it is. 

The specimens identified by Lesqllereux with the 

,European Flabellmria zinkeni Heer (U. S. Nat. ~lus. 

Nos. 101, 102) came from the exact spot that furnished 

the type specimens of Geono1nites goldianus. They 

are mere fragments, the largest of which is less than 

7 centimeters in length, and are entirely without essential characters. Indeed, the European original is 

said by Schenk 03 to be so fragmentary as to be unidentifiable-that is to say, although there can be 

little doubt as to its being a palm, it is not sufficient 

for adequate generic determination. If this is true it 

is still more true for the American examples under 

discussion, and I have referred them as above indicated. They are said by Lesqllereux to be characterized by two to four indeterminate thin veinlets, 

whereas Geonomites goldialf//us is said to have as many 

as ten. A reexamination of all the specinlens proves 

this character to be unreliable, and in the absence of 

others it has seemed best to unite them with G. goldianU8. It is possible that these American specimens 

identified as Flabella;ria zimlceni should be referred to 

Flabellaria sp., but, as pointed out under that form, it 

is too indefinite to be placed with certainty anywhere. 

Occurrence: Denver formation, South 'fable ~lountain, Golden, Colo. Green River formation ( ? ), Barrel Springs, 'Vyo. 

Geonomites? sp. 

Plate 10, Figure 1 

Leaves ev,idently large (rachis and attachment of 

rays unl{nown); rays very strong, approximately 4 

centimeters in width, strongly or Inoderately carinate; 

primary nerves distinct but rather slender, with as 

many as 10 intermediate vein lets. 

This is undoubtedly a very inadequate description, 

and perhaps the Inaterial is so fragmentary that it 

does not Inerit mention, but, as pointed out under 

Flabellwria eocenica and Geonomites goldianru8, LeS'-

quereux has apparently confused the broad rays of the 

present form with one 01' the other of these species, 

and it seems necessary to separate them. Lesquereux 

vms of the opinion that Geonmnites gold/ianus was a 

rather small leaved species, but it is beyond question 

that the present form had very large leaves. Thus, in 

the fragment figured there are 'portions of only three 

rays which show a width of over 8 centimeters, and 

the length, with little decrease in the size of the rays, 

is about 25 centimeters. Other fragments consist of 

single rays many of which show a width of 3.5 or 4 

0:1 Schenk, August, in Zittel, Karl von, Halldbuch del' Palaeontologie, 

Abt. 2, p. 372, 18!)O. 

centimeters and give no indication of narrowing or 

approaching the extremity. They are always distinctly and sometimes strongly carinate, with the pri111ary and finer veinlets as indicated in the diagnosis. 

It is possible that the fragments of palm leu yes 

from these beds referred by Lesquereux to Flabella/'iet 

zinkeni Heel' should be placed with this form, but they 

appear smaller and more grasslike and, moreover, are 

in general so unsatisfactory that it Inakes little difference where they are referred, and I have placed them 

provisionally with Geo'nomiites goldia:nW8. 

At least a part of the material provisionally placed 

under Oyperus ch(JfVannesi Heel' should be referred 

to the present form. 

Occurrence: Denver formation, Golden, Colo., col· 

lected by Arthur Lakes, 1890. 

Chamaedorea? coloradensis Knowlton, n. sp. 

Plate 10, ]j"'igure 3 

Leaves pinnate, evidently of considerable size, 

hardly less than 60' or 70 centimeters in length; rachis 

slender, striate; leaflets remote, narrow, and grasslike, 

probably subopposite, decurrent for some distance 

down the rachis, perhaps reaching the leaflet below; 

leaflets narrowed at base, with about seven or eight 

rather strong parallel nerves, apparently with finer 

striae between the stronger ones, but their number and 

character can not be made out with certainty. 

This form, which is represented by the single fragment figured, is really too fragmentary to admit of 

adequate characterization, but it is of such exceptional 

interest in the present connection that I have ventured 

to describe and figure it. It represents a portion of a 

slender rachis about 6 centimeters in length, which 

hears the basal portions of three leaflets. These lea £lets 

are much na.rrowed, and long decurrent at the base 

hut rapidly become wider above. The length of the 

leaflets is not known, but at the point where they are 

broken they are about 1.5 centimeters in. width. The 

nervation appears to be uniform-that is, there is no 

conspicuously larger midrib nor secondary veins. 

In a specimen as fragmentary as this, with so many 

of the characters obscure or unknown, it is natul'ull)p 

impossible to discuss with satisfaction its possible or 

proba.ble affinities. In many respects this seems to 

approach most closely, a feather palm fr0111 the 

,; Eolignitic" nVilcox group) of !'Iississippi, which 

was described by Lesquereux 04 under the name Oala'll'&-

opsis da'rnai but later transferred by Berry 05 to the 

genus 0 ha'rm(('edo'rea. 

9. Lesquereux, Leo, Am. Philos. Soc. Trans., vol. 13, p. 411, pI. 14, 

figs, 1-3, 1869. 

05 Berry, E. W., U. S. Geol. Survey, Prof. Papel' !)1, p. 179, pI. 12, 

fig. 4; pI. 13, figs. 1-3, 1916. ' 
SYSTEIHATIC PALEOBOTANY 41 

The genus Oala'lnopsi8 wus founded by Heel' no on a 

single species f,'om the Tortonian of Baden, with the 

following generic characterization: 

]j-'l'olldes mngnae, pillnatne, illermes, pillllis gramilleis, llervis 

llrimariis omnino aequnlibus, interstitialibus pluribus, subtilibus; nervis trnnsversnlibus nullis. 

The type species of 0 alal1wp8'i8 (0. b'J'eilana) is seen 

to be a large-leaved f01'111 with nunlerous narrow opposite leaflets wit.hout obvious midrib and not decurrent 

nt the base. 

The Mississippi form was removed from 0 alamopsis 

by Berry on the ground that the leaflets are not opposite n.nd have a decurrent base and a "fairly promiIH:'nt" 111idrib and was referred to the living AUlerican 

genus Olta'Jnaedo'J'ea, the 60 or 1110re species of which 

range from central ~iexico well into South America. 

'rhe speci1nen fronl ~Iiddle Park under discussion 

seems to combine certain of the characters of both 

these genera-that is, it has the decurrent base of 

o llal1wedm'ea but lacks the obvious 1nidvein, like 

Oa.lmnJopsis. More and better material will be necessa,ry to settle its status, but under the circm11stances 

reference to the American genus seems most logical, 

with a question mark. 

Occurrence : ~iiddle Park formation, from south face 

o.E ~iount Bross, ~1iddle Park, Grand County, Colo., 

H bout 1,000 feet above base of mountain, collected by 

George L. Cannon, 1889. 

Genus PALOREODOXITES Knowlton, n. gen. 

Paloreodoxites plicatus (Lesquereux) Knowlton, n. comb. 

Plnte 11, Figures 1-4 

Orcodomites 1)Uoat1t8 Lesquereux, The Cretaceons and Tertiary 

floras, p. 122, pI. 18, figs. 1-4, 1883. 

Knowlton, U. S. Geol. Survey PrOf. Pnper 101, p. 287, 

pI. 63, fig. 1, 1917 [1918]. 

The genus 01'eoitowites was established by Goppert 07 in 1864 nnd is based on a rather obscure seed 

or fruit frol11 the Per1uian of Braunau, Bohemia. The 

type and only species was 01'eodowites ma'rtWinus; it 

has not been subsequently reported in the literature. 

When Lesquereux 98 established his 01'eodowites plrtcat'ltS he did not give a generic diagnosis, nor did he in 

any way indicate that he considered his leaves congeneric with Goppert's Oreodowites, and there is 

every reason to sUPl}Qse that he considered he wns 

founding it anew on the living 01'eoclowa, which, he 

pointed out, it is supposed to resemble. As it is highly 

improbable, not to say inlpossible, that any relation-

('O Heol', Oswald, Flol'n tOl'Uarla Helvctiae, yol. 3, 1). 169, pI. 149, 

1~50. 

97 Goppcrt, H. R., Die fossile l!'loru dCl' permiscllen Formation: 

rnlncontogrnphlcn, vol. 12, .P, 147, pI. 26, fig. 5, 1864, 

~8 Lesquel'cUlt, Leo, The Cretaceous and Tertiary floras, p, 122, pI. 

] S, figs. 1-4, 18i3. 

ship can exist betw'een the Pernlian 01'eodowites of 

Goppert and the Eocene Oreodowites of Lesquereux, 

it seems best to give the latter n new name. I have 

therefore called it Paloreodowites: 

Following is Lesquereux's original description of 

his Oreodowites lJlicatus: 

Leaves acute at both ends, deeply plicate lengthwise in 

numerous rays converging at the base and apex, obscurely 

marked towanl the base by a median nerve [midrib]; rays 

distinctly veined; primat·y nerves distinct, sepfil:ated by three 

or four intermediate ones. 

The type specimens of this very characteristic 

species are preserved in the musemll of Princeton 

University and are apparently the most perfect that 

have been found. The best-preserved example is absolutely perfect except for a slight break at the point 

of attachment. It is about 17 centimeters long and a 

little less than 6 centimeters wide. The Inrgest specinlen is fully 7 centimeters wide and nlust have been 

nearly or quite 20 centimeters long when living. In 

the best-preserved specimen there is little indieation 

of the presence of a midrib, but in the others it is 

very pronounced, extending up for nearly one-third of 

the length. 

Only a very few additional specimens have been 

found at the type locality, and these are even less 

perfect than the least perfect of the type, hence they 

do not throw further light on the habit and relation-

~hi p of this species. . 

The generic name Paloreodowites has been selected 

80 as to preserve its connection with the O'J'eodomtes 

of Lesquereux, rather than the i1nplication of its ancestry of the nlOdern O,,'eodowa; in fact, it is much to 

be doubted if there is any real relationship between 

the living genus and the present fossil form. 

Occurrence: Den vel' formation (types) , Golden, 

Colo., subsequently collected by Arthur Lnkes, L. F. 

Ward, and others on south face of South Table ~Iountain, 100 feet below the lava cap, Golden, Colo. 

Palmocarpon palmarum (Lesquereux) Knowlton 

Ga1'polithes pa7ma1"unt Lesquerp.ux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1871, Suppl., p. 13, 1872; idem 

for 1872, pp. 382, 398, 1873. 

Palmocan;Qn COlIlmUine Lesquereux, The Tertiary flora, p. 

119, pI. 13, figs. 4-7, 1878, 

Palmoc(l1'pon pa,;'marU!I1b (Lesquereux) Knowlton, U. ·S. Geol. 

Survey Bull. 152, p. 158, 1898; Washington Acad. Sci. 

Proc., vol. 11, p. 204, 1909; U. S. Geol. Survey Prof. 

Paper 101, p. 292, 1918. 

The type specimens of this species came f1'0111 the 

Raton formation at Fishers Peak, Raton Mountains, 

Colo., and it has since been found at numerous localities in this formation in southern Colorado and northern New Mexico. It is also abundant in the Denver 

formation at Golden, Colo., and vicinity. 
42 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Notwithstanding the number of these fruits now 

known, very little has been ascertained concerning 

their nlode of growth and probable affini6es. On 

these points Lesquer'eux wrote as follows: 

These fruits, about 3 centimeters in diameter, and nearly 

globular when covered with the outer envelope, are broadly 

oval and a little smallel'-2.5 centimeters long nnd 2 centimeters across-when deprived of their exocarp. This shelly 

covering is thin, straw-colored or yellowish, smooth, and 

easily cl"tlshed; the endocarp appears also thin and, like the 

kernel, soft and easily yielding to compression; therefore few 

of these fruits are preserved in their originnl form. The endocarp is very thinly lined in the length, a character which is 

remarked only with the glass; and. on one side they are 

indistinctly marked by scars resembling the point of a 

chalaza with the raphis and hilum, as seen in the endocarp 

of some seeds. _ * * II< 'l'hey are found very numerous in 

certain localities of limited areas, as if they were derived 

from a comlilon support or a raceme. At least the specimens 

_ of Golden and of Raton are mixed with cylindrical fragments, 

like branches and branchlets or clustered peduncles half destroyed by maceration, and to which these fruits seem to have 

been originally attached; even with pieces of textile filaments, 

like remnants of decolUposed spathes. 

Occurrence: Raton formation (types), Fishers Peak, 

Raton Mountains, Colo., and at many localities in t.his 

formation .in southern Colorado and northern New 

Mexico. Denver formation, Golden, Colo., and numerous localities. Lance formation, 'Veston and Converse 

Counties, Wyo. 

Palmocarpon truncatum Lesquereux 

Palmocalf'])on tnuwatU1n Lesquereux, The Tertiary flora, p. 120, 

pI. 11, figs. 6-9, 1878. 

All the figured types of this species are preserved in 

the United States National Museum (No. 517) and 

have been carefully reexamined. They were fairly 

well -described by Lesquereux, but t~e figures are 

rather poor and more or less misleading. Lesquereux's description reads as follows: " Fruit subglobose, 

_ slightly flattened, truncate on one side, covered with a 

bro,vnish, smooth shaly envelope." On the basis of 

sjze Lesquereux divided this species into two varieties-mmjor and 1nino1'. Concerning them he writes 

as follows: 

The largest specimen of val'. major (fig. 7) meaSlll"eS 17 

millimeters across from side to side and 10 millimeters onl;y 

from the truncate base to the -top; one specimen, which seems 

intermediate between both valieties, is only 14 millimeters broad 

and 8 millimeters high. The variety 1J1,.inor (figs. 8 and 9) is 

representeu by llutle'ts 10 millimeters across the widest part 

and 5 or 6 millimeters in the other direction, generally in a 

better state of preservation, but with the same thin pericar}) 

of the same color. The characters of thE-se small nuts refer 

them to Sabal. 

Occurrence: Denver formation, South Table l\10untain, Golden, Colo., early collections. 

Palmocarpon? subcylindricum Lesquereux 

Palmocarpon su,ocyUnitr-imt1n Lesquereux, _ The Tertiary flora, 

p. ] 21, pI. 11, fig. 12, 1878. 

Following is Lesquereux's description: 

Jj"'ruit oblong or subcylindrienl, truIlcate at one end, split 

at the other in two diverging or slightly recurved, pOinted lobes. 

distantly and obscurely veined near the base. 

The figured t.ypes of this species are in the United 

States National Museum (No. 518), but the figure in 

the report on the Tertiary flora. is not quite accurate. 

There is no evidence to show that the scar at the base 

is as pronounced as the figure would lllake out, nor is 

there indication niat the ribs or costae are so -apparent. 

The base in some of the more recently collected specimens is truncate, but the point of attachment seems to 

have been smaller than figured. 

At first sight it seemed that these fruits could be 

nothing but examples of P(J)ZnlJo(J(JJ'J'pon pauma1"U11h that 

had been split at the apex by pressure, but an examination of the types, as well as a number of additional 

specimens, makes it reasonably certain that they were 

actually bilobed nearly to the middle. The pericarp is, 

as Lesquereux has pointed out, the same as in Palmoc((!rpon paZ/lnapW77V and is found in the same beds with 

that species. -

Occurrence: Den vel' formation, Golden, Colo. 

Palmocarpon? corrugatum Lesquereux 

Palmocm'pon cornt{fatmn Lesquereux, The Tertiary flora, p. 

121, pI. 11, figs. 10, 11, 1878; U. S. Qeol. and Geog. Survey Terr. Ann. Rept. for 1876, p. 503, 1878. 

The larger of the figured types of this species (orjginal of fig. 11) is preserved in the United States N abonal Museum (No. 522). The other does not appear 

to have belonged to the Museum and is now lost. The 

species has not since been found. 

Occurrence: Denver formation, Golden, Colo. 

Palmocarpon? lineatum Lesquereux 

Plate 11, Figure 5 

Pa,lmoca'l'pon Uneat'lt'Il/, Le:"quereux, Harvard Coll. Mus. Compo 

Zoology Bull., vol. 16, p. 44, 1888. 

This species is described by Lesquereux as follows: 

SeedS smnll, oval or oblong, obtuse at apex, subtruncate at 

base, regularly thinly but distinctly striate. The seeds, 4 

millimeters long, 2.5 millimeters in diameter, are very nUllll"rous, apparently derived from racemes. 

I have studied the types of this species and ha \·e 

nothing to add to Lesquereux's description. It is extremely doubtful if they are properly refelTed to the 

palms; but on the other hand no other relationship is 

suggested, and EO they are pennitted to remain as left 

by Lesquereux, but with the genus qnestioned. 
SYSTEMATIC PALEOBOTANY 43 

Occurrence: Denvel' formation, Golden, Colo, 

Types in ~1usenm of Comparative Zoology, Camuridge, :Mass" Nos, 103 and 1616, collected by Arthur 

Lakes. 

Palmoxylon cannoni Stevens 

Palmomylo'lv cnnnonl Stevens, Am. Jour. ScL, 5th sel'., vol. 1, 

p. 442, text figs. 11-16, 1921. 

Described by the author as follows: 

Jj"'ibroynsculnr bundles scnttered, usually about 10 per £qunre 

centhnetel', 0.8 to 1. millimeter in diametet:. Bast region 

rounded in outline, 1iattened where it joins the vascular portion. No marl.:ed difference in size, shape, or arrangement of 

ftbl'OYUSClllur bundles in different parts of the specimen. No 

axillnry scler~nchyma bundles. Jj'undumental tissue comrlOsed of irl'('gular, rnther thin-walled cells, with large intercellulnr spaces, and nl80 beset by groups of thicker-walled 

" stone" cells varying greatly in size, thickness of wall, aud 

number of cells to the group. Stone-cell groups usually measure fL'om 0.1 to 0.3 millimeter in dinmeter. 

- This species is based on a silicified trunk of large 

size and is remarkably well preserved with the exception of the phloem, which had apparently disappeared before silicification. Stevens states that the 

fundamental tissue is rather specialized. 

Like that of P. ccllu,losmn. Knowlton it has lnrge intercellular spaces, but scattered nmong the typical parenchymatous 

cells. usually in groups of 3 to 10, are lUuch thicl~er "stone" 

cells. These stone cells vary considerably in size, in number 

associnted in n single group, and thickness of waUs, while 

sume seem to be pitted. 

Occurrence: Denver formation, "'estern suburbs of 

Denver, Colo., in lower part of formation, collected 

by G. L. Cannon, for whom it is named. Type in 

Yule ~1usellm,-New Haven, Conn. 

Order ZINGIBERALES 

Family ZINGIBERACEAE 

-Zingiberites dubius Lesquereux 

ZingilJel'itcs du"bvlts JJesquereux, The Tertiary flora, p. 95, pI. 

16, fig. 1, 1878; U. S. Geo1. and Geog. Survey Terr. Ann. 

Heot. . for 1876, p. 501. 1878. 

Zinrllberitesf mUlti-latus Lesquereux, U. S. Geol. and Geog. Survey rl'err. Ann. Hept. for 1873, p. 396, 1874. [Homonym, 

Zingibc'I'Ues wnd'ltlat'1t8 Heel', Miocene baltische Jj"'lorn, 

11. 64, 1869.] 

A PHrt of t.he type specimen of this species is in the. 

collection of the United Sta.tes National l\1useum (No. 

136). It is a mere fragment, the nature of which, as 

Lesque'reux saiel, is open to grHve doubt. In his original characterization Lesquereux wrote as follows: 

ll'rng'ments of. hll'ge lcilves, whose ontlines are not presetTe(l, equally ulldulate on the surfnce, marked with oblique, 

distinet, parallel primary veins, 2 millin1eters distant, with 

six or seven very thin intermediate Yeinlets. The surface.is 

covered with fl thick epidermiS, 01' the lenf is subcQriaceous. 

r:rhe surface undulations are formed by deep furrows, which, 

ho\\'eyer, nre more or less distinct and which do not cut the 

connection of the veins. 'l'here is no trace of a rachis to w!lich 

the fragments of an evidently large leaf may have been 

attached. 

A poorly preserved specimen is present in the later 

collections made by Lakes, but it is not large enough 

to throw any additional light on the nature or habit of 

this peculiar plant. Schenk oa has suggested that this 

plant should probably be referred to the genus 11frt_,-,s{()-

phyllu17'&" but in the absence of lnaterial which makes 

this relationshil) plain I have thought it best to retain 

it under the name given it by Lesquereux. 

Occurrence: Denver formation (type) , Golden, 

Colo., collected by Leo Lesquereux, 1873. The recent 

specimen is from the south face of South T'able ~IountHin, Golden, 100 feet below the lava cap, collected by 

Arthur Lakes, 1890. 

Subclass DICOTYLEDONES 

Order PIPERALES 

Family PIPERACEAE 

Piper heerii Lesquereux 

Plate 11, Jj'igure 6 

Piper hcer;;, Lesqlwreux, Harvard ColI. Mus. Compo Zoology 

BulL, yol. 16, p. 44, 1888. 

Described by Lesquereux a::; follows: 

Leaves subcoriaceous, round oi' oval, entire, palmately nerved 

from the base, nerves very curved, the outei' fo1l6\ving the 

borders up to the middle of the leaf, the inner aerodrome. 

The fragment, the half of one leaf cut lengthWise or along 

the median nerve, is, in all its characters, identical with the 

leaf described by Heer 1 as Pipe1' 'U1~tiq'wmn. As the leaf is 

fragmentary, the petiole being absent, I can not well identify 

it with that of the Miocene of Sumatra, though I am unable to 

see any marked difference. The leaf described by Heel' is 

unequilateral and long-petiole(l, find none of these character:-5 

can be see_n from the American fragment. The nerves are distinct; the outer primary follows the borders at a'small distance, 

its branches, nearly at right angles, forming, by anastomosing 

curves, a series of areolae along the borders from the middle 

downward, imd the areas are traversed by nel'villes at right 

angles. 

This leaf has been ,vell described by Lesquereux, 

though, as he says, it is very fragmentary, lacking all 

of one side and a portion of the apex. It does closely 

resemble Pipe,:" antiquwrn Heel', but it is so fragmentary that certain essential features can not be compared satisfact0rily. This leaf also suggests certain 

species of Ficus, such as FiC'ltS pse'ltdopopul1us Lesquereux, which is especially abundant in the Raton 

formation, but it differs essentially in the nervation. 

Occurrence: Denver formation, South Table ~10untain, Golden, Colo., collected by Arthur Lakes. Type 

on Schenl-. August, in :littel, Karl VOII, Handbuch dCl" PalaeOllto](lgie. 

:\ bt. 2, p. 387. 1890. . 

t Heel", Oswald, Beitl'iige ZIIl' fos:,;ilen Flonl von ~lImntnl_ p_ 11, pI. 1, 

fig_ 7, 1880_ 
44 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

jI! .:Museum of Comparative Zoology, Cambridge, 

:Mass., No. 1037. 

Order MYRICALES 

Family MYRICACEAE 

Myrica sp. 

Plate 11, Figure 7 

The specimen here figured is all that was found in 

the interesting collection from reddish baked shale 

near Ramah, Colo., and, although it is rather fragmentary, it is so distinctive that it must be described. 

It is a nledium-sized leaf, clearly lanceolate, and probably acunlinate at both base and a,pex, though both 

are now wanting. It was presumably about 8 or 9 

centimeters long and just 3 centimeters wide. The 

margin is perfectly entire. The nervation is distinctive, consisting of a fairly strong midrib, a very distinct intramarginal vein or connected series of loops 

well inside the margin, and numerous fine irregular 

veins connecting the midrib and the marginal vein. 

SOllle of the stronger veins fork, and all are united 

into a fine mesh or network of. finer nervation. 

.In size, shape, and nervation this species agrees very 

closely with My/rica torreyi Lesquereux,2 but it differs 

in ha ving the nlargin perfectly entire instead of 

sh:ongly serrate. 

Occurrence: Dawson a l;kose , red baked 'shale near 

oil well south of Ramah, Colo., collected by C. )V. 

Cooke for G. B. Richardson, 1910. 

Order JUGLANDALES 

Family JUGLANDACEAE 

'., Juglans denveriana Knowlton, n. sp. 

Plnte 12, Figures 1, 2 ; Plate 13, Fjgures 2-4 

Leaves very large, probably at least 50 centimeters 

long and 30 centimeters broad, coriaceous, pinnate'; 

terminal leaflet lanceolate, long, 'wedge-shaped, ~nd 

equal-sided at the ba~e, rather abruptly narrowed 

a bove to the slenderly acuminate apex; margin slightly 

undulate but otherwise practically entire; nlidrib very 

strong, with about 12 pairs of strong, sub opposite 

secondaries, which emerge at an angle of 20° or 25° 

and curve upward strongly, each joining the one next 

nbove by a long curve just inside the border; nervilles 

numerous, strong, mainly unbroken; finer nervation 

irregularly quadrangular. Lateral leaflets large, the 

upper ones about 15 centimeters long and 5 centimeters 

wide, elliptical-Ianceolate, strongly unequal-sided at 

the base, petioled, margin obscurely serrate, apex 

slenderly acuminate; nervation similar to that of the 

terminal leaflet. Basal leaflets smaller, about 12 centinleters long and 3.5 centimeters wiele; nervation 

similar to that of other leaflets. 

~ Lesql1e1'011X, Leo, The Tertiary flora, p. 129, pI. 16, figs. 3-10, 1878. 

If this fOl'ln has been interpreted correctly, it was 

a very large leaved species. It is represented by a considerable number of well-preserved specimens, a. few 

of which are figured. 'Vhat is presumed to be the 

tel'lninal leaflet is shown in Plate 12, Figure 1. This 

was not less than 19 centimeters long and 6 centimeters 

wide. 'Vhether it was petioled is not known, as this 

portion is Inissing, but it probably was. It is remarkable for its lanceolate shape, slenderly acuminate tip, 

and numel'OUS strollg, camptodrome secondaries. 

There are two or three intermediate secondaries be· 

tween most of the secondaries. 

Two of the lateral leaflets are shown in Plate 12, 

Figure 2. They are not attached to the common petiole, 

but they lie in the same plane, and there is little doubt 

that they are in the same relative positions they held 

when living. Each has a. slender petiole about 1 centimeter long. The upper leaflet is best preserved and 

shows well the unequal-sidedness of the base, but the 

external tip is not preserved. The margin is entire at 

tl~e base, but above it is obscurely serrate. The nervation is of exactly the same type as that in the supposed 

terminal leaflet . 

The leaflet shown in Plate 13, Figure 3, is the counterpart of the upper leaflet in Plate 12, Figure 2. It 

is figured because its preservation is such that the n1arginal teeth are nlore clearly brought out. The manner 

in which the secondaries arch and send fine branches 

t.o the teeth is al~o well shown. 

Smaller leaflets are shown in Plate 13, Figures 2 

and 4. They are presumed to be the lowel' leaflets of 

the large leaf. They agree perfectly in shape and 

nervation with those last described but differ fr0111 

theJn ,in the absence of marginal teeth. In view of 

this difference the question is raised whether all these 

leaflets should be regarded as con specific. On the 

other hand, there are so many points ~f agreement 

among the.l11 that the presence or absence of slight teeth 

is not considerecl to be a sufficient warrant for separating them. The probable correctness of this view 

]s substantiated by the finding of occasional slight 

teeth in leaflets that haye the margin apparently entire, which shows it to be a variable character. 

The probable relationship of this species may now 

be considered. The smaller leaflets, such as those 

shown in Plate 13, Figures 2 and 4, have a very strong 

reselnblance to ce'rtain of the leaflets referred. to J uglan8 'l'Ug'OS((J Lesquereux; 3 in fact, one of these specimens was identified as this species by Lesquereux. 

In general, however, the leaflets of J. 'l'ugostt are 

smaller, 1110re equal-sided at the base, more obtuse at 

the apex, and always-so far as the published figures show-without marginal teeth 01' a trace of a 

petiole. Although the figures given by Lesquereux 

8 Idem, pl. 55, l'Igs. 2, 4. 
SYSTEMATIC PALEOBOTANY 45 

.1It show detached nnd isolated leaflets, it seems probable that S01ne evidence of a petiole should be ob-

~erva.ble if one had been present. The same argument applies to the marginal teeth, but, on the other 

hand, it is to be remembered that all the smaller 

leaflets of J. de'l1/oeriana have entire margins. In 

the leaflets of J. denve'l'iana the nervation is deeply 

impressed, producing the rugose appearance described 

as cha.racteristic of J. 'I'ltgosa, and it will be extremely 

difficult to allocate some specimens-that is, to distingui!:ih the one from the other. However, it is ~elieved that w.ith adequate material they may be dIStinguished without 111uch difficulty. 

,"!,'here are. also some points of agreement between 

the smnUer leaflets of Jugla1'/.s denveriana and a leaflet descr.ibed by Lesquerellx'j, under the name Sa}Jind~ts 

candatus. That leaflet differs, however, in being relatively shorter, much broader, and more abruptly narrowed to the slender tip, in having entire instead of 

obscurely serrate ll)Hrgins, and in minor particulai's 

of nerva.tion. . 

Occurrence: Denver formation, Golden, Colo., probably all from Sonth Table ~10untain, collected by 

Arthur Lakes. 

Juglans thermalis Lesquereux 

PIa te 13, FigUL'e 5 

Jlly7aHs i1bC1'nwZis Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1871, Suppl., p. 17, 1872; idem for 

1873, p. 389, 1874; idem for 1876, p. 517, 1878; The 

'l'crtial'Y florn, p. 287, pI. 56, figs. 3, 4, 1878. 

Types: United States National ~1useum, Nos. 464, 

464a.. 

The lurger of the two type specimens, the original 

. of Plate 56, Figure 3, of the" Tertiary flora," is from 

~1iddle Park, Colo., and is preserved on a hard conO'lomerate characteristic of a ,veIl-known horizon at t-> 

that locality.. The other specimen, the original of 

Lesquereux's Figure 4, is from the andesitic beds at 

Golden Colo. It is much smaller than the Middle 

Park e~ample, and, as Lesquereux pointed out, it diff€TS somewhat in nervation and may belong to a differ-

(lnt species. In the absence of additional material that 

throws light on this point it is left as designated by 

Lesquereux. 

In the material collected by G. L. Cannon in ~1iddle 

Park there is a single specimen that appears to be the 

3ame as the original example front this region. It is a 

little smaller and lacks some of the finer nervation 

but otherwise does JlOt differ essentially. 

Occnrrence: ~1:iddle Park formation (type), Middle 

Park, Grand County, Colo.; Sheriffs Creek, Middle 

~ LrsqllN'PlIx, Leo, The Tet'tiary flora, p. 2134, pi. 48, ftg. 13, 1878. 

Park, collected by G. L. Cannon, 1889. Denver formati on (type), Golden, Colo. 

Juglans schimperi Lesquereux 

Jttglan8 8chi.JII.pori. Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1871, Suppl., p. 8, 1872; The Tertiary flora, p. 287, pI. 56, figs. 5-10, 1878. 

Knowlton, U. S. Geol. Survey :Mon. 32, pt. 2, p. 688, 1899; 

U. S, Geol. Survey Bull. 204,· p. 34, 1902; U. S. GeoI. 

SUl'\'ey Prof. Paper 101, p. 296, pI. 64, fig. 1, 19li [1918]. 

Hollick, Louisiana Geol. Survey, Rept. for 1899, Special 

Hept. G, p. 280, pI. 33, figs. 1, 2, 1899. 

Penh allow, Report on the Tertiary flora of British Columbia, P·. 60, 1908. 

Ben'v The flora of the Wilcox formation: U. S. Geol. 

Su~r~'ey Prof. Paper 91, p. 182, pl. 18, figs. 3-5, pI. 19, 

fig. 4, 1916. 

Juylans 1"ltgosa Lesquereux. Lesquereux, U. S. Nat. Mus. Pl'oc., 

vol. 11, p. 11, 1888. . 

Berry 5 gives the following description . of tl~is 

species as based on the fuller material from the 'VIIcox group: 

J ... eaves ovate-Ianceolate and somewhat inequilateral in outline. Apex gmdually acuminate. Base broadly cuneate or 

rounded, inequilateral. Size variable; length ranges from 

10 to 18 centimeters; maximum width, in middle or lower half 

of the leaf, from 2 to 4.8 centimeters. Margins enti re, slightly 

undulate. Petiolule generally not preserved; in some of the 

specimens from 'Vyoming it ranges from 3 to 7 millimeters in 

length. Midrib stout, usually curved. Secondaries thin, 

numerous, rather evenly spaced, subparallel, about 14 or 15 

subopposite to nlternnte pairs; they branch from the midrib 

at wide angles and curve upward close to the margins in a 

camptodrome manner. Tertiaries mostly percurrent and distinct. Areolation subquadrate. 

Juglans schi'l1Lperi was well described and figured 

oy Lesquereux.G The type specimens came f1'0111 the 

Green River formation at Green River, "Vyo., but in 

the report on the Tertiary flora Lesquereux figured 

also (pI. 56, fig. 9) a single leaflet from the Denver 

formation at Golden, Colo. This leaflet differs slightly 

from the Green River specimens, and it may sometime 

be removed but without further material for cOlnpari- , . son I think: best. to leave it under J uglans scldmlJerz. 

This species was found at a number of localities in 

the Raton formation and by both Hollick and Berry 

in the 'Vilcox group of the Gulf States. It has also 

been reported from the ~1iocene of the Yellowstone 

National Park and with some doubt from the upper 

part of the Clarno fOl'lnation of Oregon. 

As regards the propriety of referring these leaflets 

to the genus Juglans, Berry well says that, although 

the reference is not above question, "no better disposition of theIn has suggested itself." 

Occllrrence: Green River fonnation (types), Green 

River, "TyO. Denver formation (type), Golden, Colo. 

5 BelTY, E. W., U. S. Geo!. Survey Pl·of. Puper m. p. 182, 1916. 

G LCSfjllerCllX, Leo, Th<: ~rel'tial'y flora, p. 287. pI. 513. figs. 5-10, 1878. 
46 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Raton formation, southern Colorado and adjacent 

N ew ~1exico. 

Juglans rhamnoides Lesquereux 

J,ltglams 'rhamno';dc-'S J ... esquel'eux, U. S. Geol. and Geog. Survey 

Terr. ,A.nn. Rept. for 1871, p. 294, 1872; idem for 1872, 

pp. 382, 400, 402, 1873; idem for 1874, p. 307, 1876; idem 

for 1876, p. 517, 1878 ; Bull., vol. 1, p. 370, 1876; The 

Tertiary flora, p. 284, pI. 54, figs. 6-9, 1878; The Cretaceous and Tertiary floras, p. 235, 1883. 

Dawson, Roy. Soc. Canada Trans., vol. 4, sec. 4, p. 31, 

1886. 

Lesquereux, U. S. Nat. :Mus. Proc., vol. 11, p. 32, 1888; 

Harvard CnH. l'Ius. Compo Zoology Bull., vol. 16, p. 56, 

1888. 

Knowlton, U. S. Geol. Survey Bull. 105, p. 58, 1893; U. S. 

Geol. Survey Prof. Paper 101, p. 294, pI. 66, fig. 1, 1917 

[1918]. 

A rather complete history of JUlgla:ns rftmnnoides 

has already been given 7 and need not be repeated here 

except in barest outline. This species appears to have 

been first described in 1871 from fragments said to 

have come from Evanston, Wyo., below the main coal. 

In the following year it was mentioned by Lesquereux 

as occurring at Golden, Colo., and Black Buttes and 

Evanston, 'Yyo., and in 1875 a single leaflet from 

Point of Rocks, 'Yyo., was referred to it. None 6f 

these specimens-at least under this name-:-are now to 

be found in the collections of the United States N ational ~1useum, and it seems likely that they were 

transferred to other forms without a record being 

made of such transfer, as was Lesquereux's occasional 

custom. In the report on the Tertiary flora, which 

was supposed to represent Lesquereux's mature judgment regarding all previously described forms falling 

within its scope, the only two localities mentioned are 

Spring Canyon, Mont., and Point of Rocks, ,Yyo. Of 

the four examples he figured, all of which I have seen, 

three came from Black Buttes and one from Spring 

Canyon. No specinlen was found then in the collections, nor subsequently from Point of Rocks, and 

hence JugZans 'r/w'fJnnoide8 was excluded from my re-. 

port on the flora of the Montana formation. It was 

also reported by Lesquereux as present in Tertiary 

beds at' Cherry Creek, Oreg., but in working over his 

mater'ial I failed to find it, and it was consequently 

excluded fron1 my bulletin on the fossil flora of the 

t.Tohn Day Basin. It was, how'ever, recognized as 

present in my report on the flora. of the Livingston 

formation. 

J1.lrgZan8 lr!ulfJnno·ides.was reported a number of times 

by Lesquereux from Golden, Colo., but these specimens 

have not been recognized in the later studies. Its presence in these beds rests on the' three specimens so 

7 Knowlton. F. If .. PIOI'fI of' the un tOll fOl'mntion: 1.;. ~. Ceol. SUl'\'e.r 

1'1':1'. I'aper ]01, pp. 294-295, 1017 [1!)]S]. 

identified by Lesquereux in the material ~elonging 

to the Harvard College lVluseum· of Comparative Zoology. It was also admitted to the flora of the Ratoll 

forn1at.ion on the basis of a single leaflet collected by 

'iV. T. Lee in Spring Canyon, north of Vermejo Park, 

N. Mex. 

Occurrence : Denver fornlation, Golden, Colo. Black 

Buttes coal group, Black Buttes, ,Yyo. Raton formation, Spring Canyon, Vennejo Park, N. Mex. Living-: 

ston fonnation, "Spring Canyon," ~10nt. [Hodson's 

('oal mine on ~1eadow Creek, 12 miles southeast of 

BozelnanJ. 

Juglans rugosa Les~uereux 

Plate 14, Figure 1 

Jttglall8 1'ltgosa Lesquereux, Am. Jour. ScL, 211 ser., vol. 45, 

p. 206, 1868 [nomen nudum]; U. S. Geol. Survey Terr. 

Ann. Rept. for 1869, p. 96; reprint, 1873, lJ. 196; U. S. 

Geo1. and Geog. Survey Terr. Ann. Rept. for 1871, pp. 

287, 298, 1872; idem, Suppl., pp. 10, 12, 1872; idem for 

1872, pp. 382, 390, 398, 404, 407, 1873; idem for 187Ci, 

p. 517, 1878: The Tertiary flora, p. 286, pI. 54, figs. 5. 

14; pI. 55, figs. 1-9, 1878; U. S. Nat. Mus. Proc., vol. 11, 

pp. 13, 22, 31, 1888. 

Dawson, Roy. Soc. Canada Trans., vol. 3, sec. 4, p. 30, 

1886. 

'Yard, Geol. Soc. America Bull., vol. 1, p. 529, 1890. 

Cross, Colorado Sci. Soc. Proc., vol. 4, p. 211, 1892. 

Knowlton, U. S. Geol. Survey Bull. 105, p. 53, 1893; U. S. 

Geol. Survey Bull. 163, pp. 13, 17, 1900; U. S. GeoI. 

Survey MOll. 32, pt. 2,· pp. 687, 690,779, 1899; in Merriam, 

California Univ. Dept. Geology Bull., vol. 2, p. 289, 1901: 

U. S. Geol. Survey Bull. 204, pp. 34, 90, 102, 103, 1902. 

Stanton and Knowlton, Geol. Soc. America Bun., vol. 8, 

pp. 141, 148, 1897. 

Hollick, Louisiana Geol. Survey Rept. for 1899, Special 

Rept. 5, p. 280, pI. 35, figs. 1, 2, 1900. 

J,ltglans acnminata? AI. Braun. Lesquerf>ux, U. S. GeoI. and 

Geog. Survey Terr. Ann. Rept. for 1871, Suppl., p. 8. 

1872. 

The history of J1.tglanB 'r1.tgo8a~ which is somewhat 

complicated, has recently been given at. length in the 

djscnssion under JugZans praer1.tgosa. ICnowlton. 8 A 

portion of that history is here transcribed. 

Juglans rugosa Lesquereux was nalued in 1868 from 

material obtained fr01n the ~1arshall mine, Marshall, 

Colo., but it was neither described nor figured at that 

time. The only note of explanation then given 9 is 

as follows: "Very nearly related to J. M1.t771/tnata AI. 

Braun, a species extensively distributed in the. European ~1:iocene." This note was copied without 

change by F. V. Hayden in the Third Annual Report 

of the Geological Survey of the Territories, published 

in 1869, page 96, and the species was alluded to in 

several of the later Hayden reports, but it was not. 

8 Knowlton, F. H., Flora of the Laramie fOl'lllntion: (.'. S. Gcol. 

Survey Prof. rape)' 130, pp. 121-]:!2. ] D22. 

o LeRquel'enx, Leo, Am: .Tour. Sei., 2d SP)· .. '"01. 45. p. 206, 1868. 
SYSTEMATIC PALEOBOTANY 47 

nntil 1878 that it was really validated by being prop-

(:.·1'1y described and figured. 10 Now, another complication was introduced with the figuring and describing 

of this species. The original locn,lity of 1\1arshall for 

luglams 'l'u·gosa is nowhere mentioned in the" Tertiary 

flora," and, so far as known, none of the original specilnens nre extunt. The diagnosis and figures in the 

report on the Tertiary fioranre based in the main on 

material from Evanston, ',,"yo., coming from" above 

the conl." Of the 13 figured specimens of Juglan8 

7~(,go8a, 11 are now in the collection of the United 

Stutes N utionn.l 1\1uscum. The missing types are the 

originuls of Lesqllereux's Plate 54, Figure 5, and Plate 

55, Figure 1, both, :it is said, from Golden, Colo., 

though whether from Lurnmie or Denver beds can not 

now be determined. 

From this uccount it appears that, although named 

from materiul of Laramie age at Marshall, Juglans 

?'ugosa as currently accepted is based on· material from 

higher 01' post-Lar~mie horizons. There is thus no 

menns of knowing just what the original Marshall 

leaves were like beyond the statement already Inentioned that they are said to resemble Juglams aawJni-

'nata. It is for these reasons that Juglans ?'ugo'sa is 

excluded fron1 the Laramie flora. l'he Juglans from 

the Laramie formation at 1\1arshall was named Juylans p7'aer'ufJosa, though there is, of course, no cert{tinty that the leaflets on which it is based are of the 

SHIne. type as those to which Lesquereux gave the name 

J~t.qlan8 'l"'Ltgosa. 

In the synonymy given at the head of this discussion I have noted practically all the published references to Ju.glans 'J'ugosa, from which it appears to have 

been reported from many localities. As a large proportion of the specimens upon which this record was 

made are preserved in the United States . National 

1)1 useum, it was of interest to examine them with a 

view of ascertaining if possible the correctness of the 

identifications. The following list shows the illustrations of the types of J'l.lJglans 1'uqosa in the volume 

on the Tertiary florn. and the conesponding numbers 

in the United States Nat.ional 1\1useum catalogue: 

Cntalogue Nt). 

Plnte 54, Fig-me 14 _________________________ ,154 

Plnte GG, Figure L ______________________ :.. __ 851 

Figure 2 __________________________ 45fi 

Figure 3 __________________________ 45fin 

]i'igul'e 4 __________________________ 456 

]l'igul'e ri __________________________ 457 

Figurc H __________________________ 458 

]j'jgme 7---------------0 ---------- 4(11 ]~igurc 8 __________________________ 460 

Figuro 9 __________________________ 459 

Plnj·c fi6, l~igl1re L ________ -: ________________ 46:3 

10 Lt'Sfllll'.'C'IlX. Lf'o, The Tprtin.·y flOln. p. ~R(j, pI. 54, figs. G. 14; VI. 

t,r;, figs. 1-9, 1878. 

Of these specimens Nos. 455, 455a, 45'7, 458, 461, and 

851 are from Evanston, ',,"yo. No. 460 is recorded in 

the ~luseum catalogue as having come from Golden, 

Colo., but attached to the specimen is a small original 

label which gives its locality as Evanston, '¥yo .. and 

as the matrix agrees perfectly with that ·of the other 

Evanston specimens this label is assumed to be correct. No. 455 is from 13lack Buttes, '''yo., and No. 

456 is said to be from Point of Rocks, ',,"yo. ,"!"'he 

matrix of No. 456 appears to be identical with that of 

the specimens from Evanston a,nd unlike that ordinarily found atPoint of Rocks, and on the strength of 

this finding I excluded it from my bulletin on the 

flora of the 1\10ntana formationY However, it is but 

fail' to state that this specimen has an original label 

attached to it, similar to that above mentioned, which 

records it as from Point of Rocks. In spite of this 

evidence I cn.n not escape the feeling· that the label is 

in error, the more so because the abundant recent collections from Point of Rocks did not include specimens referable to J. rugosa. As a further point it may 

be noted that Lesquereux did not give Point of Rocks 

iu his list of localities for this species in tlie report on 

the TertIary flora, this list reading as follows: 

Six. miles above Spring Canyon, near ]j"ort Ellis, :Mont.; 

Evanston, 'Vyo.; very abundant at these locnlities. Plnte 55 

is mostly composed of Evanston specimens. Black Buttes, 

Wyo., and Golden, Colo.; less common. One of the specimens, 

communicated by Dr. A. R. Mnrvine, bears the label" South 

borders of North Park," a locality whose geologicnl station 

is not known to me. 

From this it appears that most of the specimens 

came from Evanston and Spring Canyon, though none 

from Spring Canyon are figured, nor can any of them 

now be found in the Museum collections. However, 

ifl my report on the fossil plants_ of the Bozeman coal 

field I recorded the fact 12 that "a number of good 

specimens were obtained." These came from Hodson's 

coal mine, in beds supposed to be of Livingston age. 

"Ve may now .turn to some of the other localities 

whence luglans ?"ugosa has been reported. In 1886 

Dawson 13 reported this species from the Upper Laramie of Porcupine Creek, Alberta, Canada. He did 

not figure these specimens, nor have I seen them~ but 

from the fact that Dawson at one time identified them 

with J'l.tglans nigella Heel', a species which is really 

quite unlike the present one, it is presumed that this 

determination requires confirmation. 

In 1888 Lesquereux/ 4 in a paper giving the deterlninations of a miscellaneous lot of material that had 

been sent him from the National NIuseum, mentioned 

·11 Knowlton, F. H., U. S. Geol. Survey BUll. HI:). 1900. 

]2 Knowlton, F. B., U. S. Geol. Survey Bull. 105: p. 5:{. 18n:l. 

. 13 Dawson, ,T. 'Y., Roy. Soc. Canada 'l'nllls .. ~'ol. "4: sec. 4. p. 80, 

1880. . 

14 Lesqucreux, Leo, U. S. Nnt. Mns. Proc., vol. 1], pp. 22, 81. 188.';. 
48 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

two specimens fronl Cherry Creek, Crook County, 

Oreg., as referable to Juglans rugosa. I studied t.hese 

specimens when preparing my paper on the fossil 

flora of the John Day Basin 15 and wrote of them as 

follows: " Represented by two fragments that are more 

or less obscure and doubtful." In his paper of 1888 

Lesquereux mentioned a single specimen fr0111 Lassen 

County, Calif., but this also is fragl11entary and not 

well authenticated. 

Hollick 16 reported J'Uglans rugosa fr0111 beds of 

" Eolignitic" (Wilcox) age near Coushatta, La., but the 

species was not accepted by Berry 17 in his paper on 

the flora of the Wilcox formation. In the Yellowstone 

National Park IS I detected this species at a number of 

localities in beds supposed to be of Fort Union age. 

J~(,glans 1"ugosa was found to be fairly abundant in 

the Raton formation of the Raton l\lesa region ot 

Colorado and New Mexico.19 

Occurrence: Denver formation, Sand Creek near 

~lagnolia, northeast of Denver, Colo., collected by 

Francis 'V. Collins, 1909. 

Juglans sp. 

Plate 14, Figure 5; Plate 15, Fjgure 1 

The Dawson arkose at a locality on Jimmy Camp 

Creek has afforded the example here figured. It is a 

segment from near the middle of what appears to be 

an elliptical-Ianceolate leaflet. It must have been 12 

01' 13 centinleters long and is very nearly 6 centimeters 

wide. l\tIargin entire; petiole strong and perfectly 

straight; secondaries numerous, mainly alternate, 

emerging nearly at a right angle, cons~derably curved 

upward, camptodrome; nervilles strong, very oblique 

to the secondaries. 

This is undoubtedly very close to Juglan8 rlJenve'J'-

i((l/Ut, a,bove described, and may be identical with it. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Ci'eek, 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. Knowlton, July, 1910. 

Hicoria antiquora (Newberry) Knowlton 

IIioo1'ia. antiq1t01'a (Newberry) Knowlton, U. S. Geol. Survey 

Bull. 152, p. 117, 1898. 

O((,/,ya. a.ntiq1tOrum Newberry, New York City Lyceum Nat. Hist. 

Annals, vol. 9, p. 72, 1868. 

Lesquereux, The Tertiary flora, p. 289, pI. 57, figs. 1-5; 

pI. 58, fig. 2, 1878; Haryard Co11. Mus. COlllp. Zoology 

Bull., vol. 16, p. 56, 1888. 

Newberry, U. S. Geol. Survey Mon. 35, p. 35, pI. 31, figs. 

1-4, 1898. 

]6 Knowlton, F. fl., U. S. Geol. Survey Bull. 204, p. 34, 1902. 

10 Hollick, Arthur, Fossil plants from Louisiana: Louisiana Geol. 

Survey Rept. for 1899, Special Rept. 5, p. 280, pI. 35, figs. 1, 2, 1899. ' 

17 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, 1916. 

15 Knowlton, Ii'. H., U. S. Geol. Survey Mon. 32, pt. 2, p. 687, 1899. 

19 Knowlton, F. H., Flora of the Raton f'ormation : U. S. Geol. Survey 

Prof. Paper 101, p. 293, pI. 112, fig. 4, 1917 [1918J. 

In the material from the andesitic beds at Golden, 

Colo., identified by Lesquel'eux for the ~1useum of 

COl11parativeZoology, he reports a single example of 

this species. It is a small, rather obscurely preserved 

leaf or leaflet, and its identification is more or less 

doubtful. However, as it seems to differ from any 

other leaves found in the same locality, it. 111ay be permitted to stand until further evidence is available. It 

was not noted in the extensive material fronl this 

locality that. has been passed in review. 

Occurrence: Denver formation, Golden, Colo., speci~ 

men in the l\luseum of Comparative Zoology, Cam~ 

bridge, :Nlass. 

Ptel'ocal'ya? retusa Lesquel'eux 

Plate 14, :Figures 2-4; Plate 15,' li'igurc 5 

Ptel'ocar}}a retwsa Lesquereux, Harvard Co11. Mus. Compo 

Zoology Bull., vol. 16, p. 56, 1888. 

Described as follows by Lesquereux: 

Terminal leaflet large, ovate']anceolate; the lateral [leaflets] 

small, linear-oblong, blunt at apex, rounded-subtruncate at 

base; lateral nerves close, pal'll11el, at a bl'oad angle of di, 

vergence, branching near their ends; surfaces rugose. 'l'he 

leaflets are thic1\:, denticulate or crenulate on the recuned 

borders; the lateral nerves, at an angle of 50°, are not more 

than 5 millimeters distant i-n-the small leaves; deeply marked. 

This species is said by Lesquereux 20 to be "closely 

allied to Pte1'ocarya americaJna Lesquereux, differing 

by the open proximate nen'es." It is also said to show 

relationship with J~()gl(J)ns corrur/at'(J;, Ludwig 21 "by 

its rugose surface and its nervation, * * * differing especially by the leaves being blunt or obtuse, not 

acuminate as in the European species." 

Pterocarrya 0I17U31'icana Lesquereux is said by Lesquereux to have come frOl11 Middle Park, Colo., from 

beds presumed to be in the approxil11ate position of the 

Denver formation, but an exal11ination of the type 

specimen in the Nationall\1useum proves ulllnistakably 

that it came fr0111 the NIiocene lake beds at Florissant, 

Colo. 

Pte1'oca1'ya? 1'etusa is represented by six speciInens, 

four of the best of which are here figured and may be 

regarded as the types. N one of them a re very well 

preserved. The one ta.ken by Lesquereux to be the 

terminal leaflet (pI. 15, fig. 5) is about 7.5 centimeters 

long and nearly 3.5 centimeters wide. It is said by 

Lesquereux to be blunt at the apex, but on reI110val 

of nIore of the 111atrix it was found to be fairly acute. 

The margin is curled under to such an extent that it 

is almost impossible to determine the character of the 

teeth or crenulatio~s, but they appear to be short and 

rather blunt. Another specimen (pI. 14, fig. 4) is fully 

20 LesqueL'ellX, Leo, The TeL·tiary flora, p. 290, pI. 58, fig. 3, 1878. 

21 Ludwig, Rudolph, Palfl.eontogrnphica, vol. 8, p. 178, pI. 70. figs. 11, 

12, 1861. 
SYSTEMATIC PALEOBOTANY 49 

7 centimeters long and about 3 centimeters wide and 

shows the npex to be rather long and slender. The 

base :is destroyed, and the margin is very obscure 

though uppm:ently only undulate. The third specimen (pI. 14, fig. 3) hns the base preserved but Incks 

all of the apical portion; it wa.s probably about 5.5 

centimeters long and 1.75 centimeters wide. The margin is certainly entire ut and near the base, but in the 

upper part it C!1n not be determined. The best-preserved exnmple (pI. 14, fig. 2) shows a tenllinalleaflet 

a.nd a latel'alleaflet in place. The terminal leaflet was 

probably ubout 10.5 or 11 centimeters long and 7 centimeters wiele, and the lateral leaflet must have been 

about 7 centin1eters long and 2.5 centimeters wide. 

The margin in both leaflets is so much turned under 

(whence the specific name 'ret-usa) that it is very difficult to determine whether or not there were teeth, 

though the margin was proba,bly crenulate, as Lesqnereux stutes. The nervation consists of a very 

sh'ong n1idrib and numerous, mainly alternate, irreguludy spaced secondaries that arise at a very low angle, 

curve upwarcl n10re or less, and are apparently craspecloclrome. The nervilles are strong, are lllainly at 

right n.ngles to the seconda.ries, and produce a distinctly 

rugose nppea.rnnce. 

It is doubtful if these lenves are correctly referred 

to the genus Pte1·ocarrya. They have'much l110re the 

appearance of being leaflets of a J'llgZwns; in fact, on 

Lesquereux's or,iginal labe~ with one of the specimens 

it wu,s pluced. in J'l.lJgZans, but later it was referred to 

Pte1'oc(t'}·ya. The resemblance to Pterocarya americana Lesquereux is shown by the lateral leaflet and not 

by the terminnl one, which is, of course, very different. 

On the whole it would probably be better to place this 

form in the genus J'l.lJglans, but as so 111any features are 

obscure it hus been retained as left by its author, 

simply questioning the generic reference. 

Occurrence: Denver formation, South Table ~10untnin, Golden, Colo., collected by Arthur Lukes. Types 

in ~1u~eum of Compnrntiye Zoology, Cambridge, ~1ass., 

Nos. 414, 1558, 1559, anc11561. 

Order FAGALES 

Family BETULACEAE 

Alnus carpinifolia LesQuereux 

Plate 15, lngure 2 

Alm/.s C(irpi,ll'i.foli{b J .. esquereux, Hnryfll'd ColI. Mus. Compo Zoolog~' Bull., vol. 16, p. 45, 1888. 

l. .. enves comparlltively smull, ovate-acute, narrowed and 

nUl'llptly Shol't-decul'ving to the petiole, entire to the middle, 

(lenticulnte above; latel'lll nenes eCluidistallt, the lower pail' 

much brunching, all pllrallel, at fin acute angle of divergence, 

crnspetlodl'ome. ' 

~ehe rt'llltioll 01: this lenf is with that of .·.I1I/o/(.s G1Jcl~J<l'n'ln 

UJlger,22 differing essentinlly by the base, which is rounded and 

,,~ 1;11,1.\'('1'. Fl'nnz, DIe fosslle I!lom yon Kumi, p. 23, p). 3, fig. 10, 1R67. 

abruptly turned down to the petiole. The lower lat('ral nerves 

are joined to the midrib a little above the base of the leaf. 

Occurrence: Denyer formation, South Table ~1ountain, Golden, Colo., collected by Arthur Lakes. Type 

in :Museum of Comparative Zoology, Cambridge, 

Mass., No. 1056. 

Alnus auraria Knowlton and Cockerell 

Plate 15, :t .... igure 6 

Alnus (£1tl'(l1'ict Knowlton and Cockerell, U. S. Geol. Survey 

Bull. 696, p. 63, 1919. 

Al1Mts 1"lt{fOS(t J .. esquereux, Harvard CoHo Mus. Compo Zoology 

Bull., vol. 16, p. 45; 1888. [Homonym, Koch, 1872.] 

Lesquereux's description reads as follows: 

Leaves membranous [membranaceous], elliptic-ovate, narrowed to the base, undulate on the bordel:s, l)innately nerved; 

lateral nerves equidistant, parallel, straight, cruspedodrome, 

with few branches; surface rugose. The leaves nre about of 

the same size ns those of AZw/l,s nost'ratum Unger,23 and the 

nervation is of the same type, but the leaves are narrowed, 

not rounded to the base. 

This species was based on two specimens, the best 

preserved of which is here figured. It lllust. have 

been about 10 centimeters long (the apex is wanting) 

and is a little over 5.5 centimeters broad. It has a 

very strong lllidrib, especially below, and six or seven 

pairs of alternate secondaries. The lowest pair of 

secondaries are very thin and run for only a short 

distance, but the next pair are strong and hn ye several 

branches on the outside. The secondaries and their 

oranches are aU craspedodrome, ending in the margin, 

vlhich is in places slightly undulate-toothed. 

Occurrence: Denver formation, South 'l'able Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in ~1useum of Comparative Zoology, Cambridge, 

~1ass., No. 1057. 

Betula schimperi LesQuereux 

Plate 15, lngures 3, 4 

13et'u.Z(I, sohhlllJCl''i Lesqnercux, Hnl'Yllrd COli. l\lus. Compo Zoology Bull., vol. 16, p. 45, 1888. 

Described by Lesquereux as follows: 

Leaves small, as broad as long, round or subcordate at base; 

ovate, acute, simply or doubly dentate; lateral nerves thick, 

five pairs, the three lower opposite, the lowest more or less 

Qranching outside, all craspedodrome, as wen as the divisions, 

p.ntering the larger teeth directly when simple, and the intermediate short ones by brunches; curved in traversing the 

:ireas at a broad angle of divergence. The leaves meaSUl'e 2 

to 3 centimeters across, both ways, and nre deltoid-acute at the 

apex. The teeth, like the veins, are somewhat distil nt, turned 

outside, sharply pointed, triangular. The base of the median 

nerve is pressed into the stone, and thus the leaves appear 

subcordate; but the base is really rounded and abruptly turned 

down to the pedicel [petiole]. The surface is rugose by the 

deep impression of the nerves and the nervilles, these being 

~3 Unger, Franz, Ch)ol'is l'l'otogncn, pI. 34, fig. 1, 1847. 
50 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

simple or branching in the middle. 'rhe relation of this species 

is with Betula angulata Goppert.2~ 

This species is represented by two specimens, both 

of 'which are figured. It appeal'S to be a good species 

and has been well described by Lesquereux. 

Occurrence: Denver fOl'mation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, l\iass .• 

Nos . .1035, 1036. 

Betula gracilis? Ludwig 

Bet'llla, gracilis Ludwig, Palaeontographica, vol. 8, p. 99, pI. 32, 

figs. 3-6, 1861. 

Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. 

for 1873, pp. 380, 397, 1874; The Tertiary flora, p. 138, 

pI. 17, fig. 20, 1878. 

The only spe<;;imen upon which the presence of this 

European species in American strata depends is in the 

lTnitecl States National Museum (No. 158). It is a 

small, poorly preserved leaf, and its identification is, 

as Lesquereux indicated, open to grave doubt. It does 

resemble Betula somewhat but is too imperfect to be 

of much value. . 

Occurrence: Denver fOl'mation, Golden, Colo. 

Betula fallax Lesquereux 

Plate 15, Figures 7-10; Plate 16, Figures 1, 2 

Betu,la fallaa: Lesquereux, Harvard ColI. Mus. Compo Zoology 

Bull.. vol. 16, p. 45, 1888. 

Lesquel'eux's original description and discussion are 

as follows: . 

Lt'Hves rhomboidal-ovate, cuneate to the base and narrowed 

in the same degree from the middle to the apex, penninerved; 

lower pair of secondaries attached to the midrib above the base 

of the leaves and opposite, the others, five to six pairs, parallel, 

ut Yariable distance, at an acute angle of divergence, somewhat 

curved in passing toward the borders; border distantly dentate 

ill the uppel' part of the leaves, the teeth short and turned 

up, being marked only at the apices of the lateral nerves, 

which are most simple, the lower ones only with few branches. 

The species resembles Betula nigra Linne in the form of th~ 

leaves. The leaves vary from 3.5 to 5.5 centimeters in length; 

from 2.5 to 3.5 centimeters in width in the middle. The angle 

of divergence of the nerves is only 30°, 'and therefore the 

,leaves are only dentate in tile upper part, at the points entered by the secondaries. 

This 'species, according to Lesquereux, is represented 

by no less than 32 specimens in the Harvard College 

~1useum of Comparative Zoology. I have studied all 

this material and have selected six specimens to be fignred, the remainder being largely fragmentary. 

There is so considerable a range in the appearance of 

these types as almost to lead to the conclusion that 

they can not be conspecific. o For example, take the 

question of size. Lesquereux gives the length as 

"4 Giippel't, H. H., TertiUl'e Flora von Schos:snitz, p, 10, pI. 3, fig, 3, 

1855. ' 

being between 3.5 and 5.5 centimeters, but in the 

specimen shown in Plate 16, Figure 1, the length is 

little less than 8 centimeters. The width is said to be 

between 2.5 and 3.5 centimeters, but the width of the 

specimen in Plate 16, Figure 7, is fully 4 centimeters. 

In the marginal toothing there is also considerable 

range. Thus, ill most of the specimens the teeth are 

small and obscure, but in Plate 15, Figure 9, the teeth 

are all relatively large and oaklike. This particular 

specimen closely resenlbles certain of the leaves referred to Orataegu8 betulaefoliu8 Lesquereux, from 

the same horizon, and is also suggestive of small leaves 

of Quercu8 vib~lrnifolia Lesqllerellx. 

Occurrence: Denver fornlation, South Table ~lountatin, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, 

l\Iass., Nos. 1034, 1043, 1044, 1045, 1058, 1620. 

Family FAGACEAE 

Castanea intermedia? Lesquereux 

Ca8tanea 'intermedia, Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Bull., vol. 1, 2d ser., No.5, p. 386, 1876; U. S. 

Geol. and Geog'. Survey Terr. Ann. Rept. for 1874, p. 

313, 1876; The Tertiary flora, p. 164, pI. 21, fig. 7, 1878. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 297, 

pI. 68, fig. 2, 1917 [1918J. 

Th~s species is said to have come from ~1idclle Park; 

Colo., from beds thought to be of Green River age, 

but it apparently has not been since reported fronl 

that locality, and later knowledge of the geology of 

l\1iddle Park makes it seeIn reasonably probable that 

it caIne frOlll beds of Denver age. A single example 

referred to this species was, reported from the Raton 

formation at vVootton, Colo., by me. It is a little 

larger and broader than the type but does not otherwise differ essentially. 

In one of the small collections made near the Purdon 

mine there is a leaf that is most like the speciluen 

figured in the Raton flora, but it differs slightly in 

being proportionately broader and less conspicuously 

toothed, though the margin is not very well preserved. The nervation is the same in both leaves. 

Occurrence': Dawson arkose, a quarter of a mile 

east of Purdon mine, sec. 27, T. 11 S., R. 61 'V~, Colo., 

collected by G. B. Richardson, 1910. 

Quercus viburnifolia Lesquereux 

Plate 16, ngures 3-8; Plate 17, Figure 1 

QuorCtts vib,lt1'111ifoUn Lesquereux, The Tertiary flora, p. 159, 

pI. 20, figs. 11, 12, 1878; U. S. Geoi. and Geog. Survey 

Terr. Ann. Rel1t. for 1876, p. 505, 1878; Haryard CoIl. 

Mus. Com!>. Zoology Bull., vol. 16, p. 46, 1888. 

Ettingshausen, Ro~al Soc. Londou Proc., vol. 30, p. 232, 

1879-80, 

Knowlton, Washington Acad. Sci. Proc., vol. 11, p. 207, 

1909'; U. S. Geol. Survey Prof. Paper 130, p. 127, 1922. 
SYSTEMATIO PALEOBOTANY 51 

Qu.crcn,1I t1'iangula'ri8 GUppel't [part]. Le:-;quereux, U. S. Geol. 

and Geog. Survey Terr. Hept. fol' 1872, p. 377, 1873. 

Quercu8 Mtol1'11(1.ta!t G(jPllert. I ... esquereux, U. S. Geol. and 

Geog. SlIl'vey r.rerr. Uept. for 1873, p. 381, 1874. 

POJ)ltl1t8 ~rena,ta Unger. r ... esquereux, Harvard CoIl. Mus. 

Compo Zoology Bull., vol. 16, p. 48, 1888. 

O'l'(£taoU'lt8 betula.cfoH(t I ... esquel'eux [part], Harvard CoIl. Mus. 

Com!>. Zoology BulL, vol. 16, p. 56, 1888. 

OrataeU1t8 (t:ntiq'na Heel'. Lesquel'eux, HarVlu:c1 CoIl. Mus. 

C0111!>. Zoology Bull., vol. 16, p. 57, 1888. 

Leaves membranaceous, ovate or ovate-oblong, 3.5 

to 9 centimeters long, ::3 to 5.5 centimeters broad, 

broadest neal' or just below the nliddle, whence they 

are abruptly naLTowed to the obtusely wedge-shaped 

base Hnd above to the rather obtuse apex; margin 

entire below, sharply and somewhat irregularly 

toothed l1bove;' petiole sl~nde\r; nervation pinnate, 

consisting of a rather slender, slightly irregular midrib and four or five pairs of slender subopposite or 

alternate secondaries, which end in the marginal 

teeth, lower secondaries often with several outside 

branches, craspedodrome; nerv~lles numerous, strong, 

nbout at right angles to the secondaries. 

One of the figured types of this species-the origiHal of Plate 20, Figure 11, in the report on the Tertiary 

flora-is preserved in the United Sta,tes National MuSl'lIlH (No. 186), together with a number of others 

that wel'e the basis of Lesquereux's discussion of this 

form. This type specimen was found on Sand Creek, 

about 15 miles east of Denver, Colo., in beds long supposed to be of Arapahoe age but now known with reasonable certainty to be in the Denver formation. 

Other specimens not figllred that are in the Musemn 

collection came from the nndesitic beds at Golden, 

Colo., and. the red-baked shale at Black Buttes, Wyo. 

In a hltel' publication Lesquereux 25 recorded the presence of nearly n dozen examples from Golden. It has 

also been reported fr0111 beds of Denver' age on Mount 

13 ross, Middle Pnrk, Colo.; from the Lance formation 

of Converse County, 'Vyo., and somewhat questionably from the true Laramie on Crow 'Creek east of 

Greeley, Colo. 

In the Dawson arkose at ~10sby, Colo., was found 

tlle specilnen with its 'counterpart shown in Plate 16, 

Figure 5. It is intermediate in'size between the two 

figul'ed types as given by Lesquereux." It is about 

6,5 centimeters long and a little over 5 centimeters 

wide. It is proportionately a little shorter than the 

larger of Lesquereux's types but otherwise agrees with 

it :in all essent:ial particulars. 

Another specimen from the vicinity of Calhan is, 

shown in Plate 17, Figure 1. It agrees very closely in 

size, shape, and marginal teeth with the larger of the 

figured types and differs only in having the nerVH,-

~~ LeSqu!'I'(,IIX, Leo, Hnl'\'urcl Coli. Mus. Compo Zoology BulL, vol. 16, 

D. 46. 188tt 

tion somewhat stronger, though of the same disposition. 

Lesquereux:!(1 established O'rataegu8 betulaefolia on 

specimens fronl Golden, Colo., in the Harvard College 

~Iuseum of Comparative Zoology. I have examined 

most of thenl and am unable to draw any line of 

s~pa ration except in size between these and QuerC'tt8 

vibU1"nifoli(() Lesquereux. 27 The two types of Que'reu8 

vib'1llrnifolia are 4 and 9 centimeters long, and the 

l'ancre in size in the leaves of 01'alta'.8g'l()8 betulaefolia o . 

is gi ven as from 3.5 to '7 centimeters. On a single pIece 

of matrix (Mus. Conlp. Zoology No. 1631) there are 

foul' leaves, two of which are types of 01YDtaegu8' bet'ltlaefolia and two were referred by Lesquereux himself 

to Q'l()e'rCU8' vibu1'nifoli(J); all are here figured. Of 

COllrse, close association on the matrix does not necessarily prove identity, but it does suggest that they 

came from the same vicinity and might well enough 

have come from the same tree. 

Lesquereux states that his 01'ataegu8 be,tulaefolia 

is "trinervate from the base," but an examination of 

the specimens shows that the lowest secondaries are 

opposite and arise at a distance above the base of 

the blade, thus producing a sort of f~lse trinervate 

appearance. This character is to be noted in both 

types of Q'UJer(}U)8 vibwl"'nifoUa as well as in the specimens so identified by L~squereux and in many 

others subsequently studied. In the wedge-shaped 

slightly' decurrent base, sharp-toothed margin, and 

craspedodrome secondaries these leaves are aU a ppa 1'-

ently conspecific and are combined under Q'ltJerau8 

viburnifolia. ' 

The specimens identified by Lesquereux 28 as 01oatae-

{JUS antiqua Heel' and now in the ~1useum of Comparative Zoology (Nos. 1563, 1564) are also referred to 

Q'lte1'C1JJS vibwrnito'liaJ.; it has not been thought neces-

, sary to figure any of them. 

In 1888 Lesquereux 29 also referred no less than 17 

specimens from the Denver to Populus C'ren.ata Unger. 

I have now seen these specimens, though the present 

remarks are based on a single leaf (U. S. N at. ~{us. 

No. 51083) so identified by him in the Lacoe collec60n. This is a small leaf about 5 centimeters long 

and 3.5 centimeters wide and has the lnargin above 

the middle provided with few large sharp teeth. The 

secondaries number three or four pairs; the lowest 

pair arise just above the base of the blade and end in 

laro-e o teeth , and each has several branches on the outside. So far as I can determine, this leaf is not essenbally different from the smaller of the two leaves fig20 Lesquereux, Leo, Harvard CoIl. Mus. COIll[). Zoolog'y Bull., vol. 1G. 

p. 56, 1888. 

:l7 Lesquel'eux, Leo, '.rhe Tel·tlal·Y flora, p. 15\), pI. 20, fig. 11, 1878. 

2S I.esquel'eux, Leo, Hnrval'cl CoIl. Mus. Compo Zoolo):y BulL, Yol. 16, 

p. 57. 1888. 

:.'!l Idem. 41. 48. 
52 ]'LORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

ured by Lesquel'eux 30 as the types of Que1.'(}u8 viburnifolia, though it has rather larger teeth and the upper 

secondaries at a slightly Inol'e acute angle. 'Vhether 

all the specimens identified by Lesquereux as Unger's 

Pop~tl~f.,8 ere'JUlia are referable to Q~tereU8 vibtt-rnifolia 

I am not now able to say. 

Occurrence: Denver formation (type), Sand Creek, 

about 15 Iniles east of Denver, Colo., collected by A. R. 

Marvine in 187.3; South Table ~tlountain, Golden, 

Colo., collected by Leo Lesquereux about 1872 and by 

Arthur Lakes in 1885. ~1iddle Park formation, ~tlount 

Bross, Middle Park, Colo., collected by 'Vhitman 

Cross. Dawson arkose, dump of ~tlosby coal mine, 

~losby, EI PasQ County, Colo., collected by F. H. 

I{nowlton and G. B. Richardson, July, 1910; Rice's 

day pit, 1 mile southwest of Calhan, Colo., collected 

by F. H. Knowlton, 1910. Lance formation, gulch 

south of Lightning Creek, Converse County, Wyo., 

collected by T. 'V. Stanton, July, 1896. Black Buttes 

coal group, Black Buttes, ""Vyo. Laramie formation, 

~Canfield ranch, about 25 miles northeast of Greeley, 

Colo., collected by T. 'V. Stanton and F. H. I{nowlton, 

,T une, 1896. ' 

Quercus? sedaliensis Knowlton, n. sp. 

Plate 17, 1!'igure 4 

Leaf large: evidently of fine texture, broadly elliptical, apparently rounded or possibly obtusely wedgeshaped at the base and probably obtusely acuminate 

at the npex j margin undulate-tooth~d throughout, the 

teeth small, low, rounded, and separated by similar 

shallow sinuses; midrib very thick, especially below, 

perfectly straight; secondaries evidently 10 or 12 

pairs, rather slender, alternate, somewhat curving upward, all unbranched and camptodrome. 

This was a, large leaf, probably not less than 15 

centimeters in length and possibly longer, and the 

width in the middle is about 7 centimeters. Unfortunately, it lacks both base and apex, but, so far as can 

be luude out, the shape was broadly elliptical and apparently abruptly rounded to the base; though it may 

have been obtusely wedge-shaped. The most marked 

eharader is the margin, which is provided ~with numerous small, rounded teeth that are separated by 

similar shallow sinuses. 

This leaf is referred with hesitation to the genus 

Qtte'rcu8 solely on the ground of its evident resemblance to what has been called Qtl'ereu8 haidingeri 

Ettingshausen rather than the conviction that it actudly belongs to this genus. Broadly considered its 

facies is hardly that of an oak, yet it seems fair to 

assume that, if Ettingshausen's species is correctly refflTed this must also be correct; in any event it is so 

30 LeSfjlWl"I'UX, Leo, The Tertiary flora, pI. 20, fig. 12, 1878. 

well characterized by its large size and peculiar ma rgin that it can be easily recognized in the future. 

As stated above, it is nearest to Q. haidinge1'i, fr0111 

which it differs in its much larger size, thicker midrib, 

and evidently more rounded base. Additional examples, however, may show that the size and supposed 

difference in shape are "not constant, and it may then 

be referred to Ettingshausen's species. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Al~apahoe formation), 3,000 feet east of 

the Douglas coal mine, Sedalia, Colo. 

Quercus purdonensis Knowlton, n. sp. 

Plate 17, Figure 2 

Leaf small, evidently thick and coriaceous, elliptical, 

rounded and entire below, strongly few-toothed aboye, 

the teeth relatively large, sharp-pointed, apparently 

bristle-tipped, apex acuminate; petiole stout; midrib 

very strong; secondaries four or five pairs, strongly 

alternate, at an angle of about 45°, craspedodrome, 

ending in the Ii1arginal teeth; finer nervation not well 

preserved. 

This little leaf is nearly perfect and is the only one 

noted in the collections. It is 3 centimeters long and 

a little less than 2 centimeters wide. It is remarkable 

for the large, sharp-pointed teeth. 

This species is of the type of a number of species 

living at the present time in the Rocky ~tlountain and 

Pacific coast regions, being not greatly unlike SI11311 

leaves of Que1'Ctt8 engelmanni Greene, Q~ter(}u8 eh1'Ysolapi8 Liebmann, or Querou8 Wi8l1.·zeni A. de Candolle. 

Occurrence: Dawson arkose, a quarter of a 11lile 

east of the Purdon nline, sec. 27, T. 11 S., R. 61 'Y., 

Colo., collected by G. B. Richardson. 

Quercus whitmani Knowlton, n. sp. 

Plate 17, Figure 5 

Leaf relatively large, evidently coriaceous, rather 

broadly lanceolate, wedge-shaped at the base, apparently rather narrowly acuminate at the apex; Inargin 

obscurely toothed, the teeth remote, very low, the 

sinuses very shallow; midrib fairly strong, straight; 

secondaries about 10 pairs, alternate below, subopposite 

nbove, slightly curved upward, occasionally forked, 

the lower ones camptodrome, arch~ng along the nlargin 

Hnd sending nerviJles fr0111 the outside to the Inarginal 

teeth, upper secondaries craspedodrome, entering the 

teeth; nervilles numerous, strong, mainly percurrent, 

approximately at right angles to the secondaries. 

It is" with some hesi tation that this species has been 

descl;ibed as new. It is not a. perfect specimen, lacking both base and apex and a considerable portion of 

one side. So far as can be determined, it nlllst ha '"e 

been about 15 centimeters in length and about 6 centi-
SYSTEMATIC PALEOBOTANY 53 

meters in width. The margin can be made out in 

severnl plnces und shows the teeth to be very low and 

the sinuses correspondingly shallow. One or two of 

the teeth appeur to have been spinescent or possibly 

brisl'le-tipped, and if this appearance has been cor': 

rcctly interpreted it is probable that all the teeth were 

thus terminated. The strong nervation is well shown 

in the figure. 

In some respects the present leaf is similar to what 

is described above as Que'l'cus serlofliensis I\::no~lton, 

but it differs in a number of minor particulars. Thus, 

the teeth are of a different character, the secondaries 

are less numerous, and the finer nervation is much 

more brol{en up. 

Que'l'cus w"l~itm{Jjnri is also in a way suggestive of 

Q~(,erc~ts paucirlentOita Newberry ,81 fronl the upper part 

of the Clarno formation at Bridge Creek, Oreg. The 

Oregon leaf is slightly smaller than the one under consic1el'ntion nnd moreover has the teeth aggregated in 

the upper part of the leaf. 

Occnrrence: Middle Park formation, Middle Park, 

Grnnd County, Colo., from high terrace ridge between 

forks of I(inney Creek, in the "breccia spoon," east 

of I-Iot Sulphur Springs, collected by Whitman Cross, 

October, 1891. 

Quercus whitei Lesquereux 

Plnte 17, li'igm'e 3 j Plat~ 18, Figure 1 

Quc/'ens whUc'i Lesquerenx, Harvard ColI. Mus. Compo Zoology 

BulL, yol. 16, p. 46, 1888. ~ 

J ... efl\·es of medium size, membranous [membranaceous], 

0\"11 to-In IIceola te, cn neiform to the base, short-petioled, reguInrly mOI'e 01' less deeply dentate; secondaries straight, oblique, 

cquidi~tllnt:, simple, plll'allel, each entering one of the teeth, 

which nre gibbous on the back. The secondaries at an angle 

of 40° lire strnight, the upper ones only slightly curved. The 

teeth, ~ltOlt upon some lenYes, as long as 0.5 centimeter upon 

others, fire alwnys distant find bossed on the back. The leaves 

avcrnge 7 centlmeter~ 1011g and 4 c~ntimeters broad in ,the 

mldtlle. 

'1'1118 fine species is closely nllied to Que1'c'u,s etymodrys l\1assalongo,8~ especially with the variety described as casteZZinensis 

by Cnpellini.83 It is nlso allied to QUe1'cus f'lwcinervis Rossmns:sler, differing from both by the subdentate or umbonate 

teel'll. 

Two of the best preserved of the types of this species 

are here figured. The one shown in Plate 18, Figure 

1, was nearly half concealed in the matrix, but on remoying this it is found to be nearly perfect. It is 

Lrondly ovate-Ianceolate, with a very obtusely wedgeshaped base arid an acuminate apex. The margin is 

cntire for about one-fourth the length of the blade and 

thence provided with large obtuse teeth. The midrib 

31 i\cwbcrry, J. S., U. S. GeoI. Survey Mon. 35, pI. 43, fig. 1, 1898. 

r.2 Mnssnlongo, A. B., Synopsis fiora fossilis Senegallicnsis, p. 26, 

lS:;~. 

:-, ('np('lIllll, T. F. G .. l.a fOl'llln?ionc gessosa di Castcllin3, p. 572, pI. 

5. f1;.:'. 1. l~7". 

is relatively strong, with about five pairs of subopposite secondaries. This leaf is about 7.5 centimeters 

long and nearly 4 centimeter~ broad. 

Occurrence: Denver formation, Golden, Colo., collected by Arthur Lakes, 1883, for the Museum of Comparative Zoology, Cambridge, 1\1ass., types Nos. 1072, 

1056,1628. 

Quercus leonis Knowlton, n. SI). 

Plate 18, lI'igures 2, 3. 

Quercus ha-idvngerl Ettingshausen. Lesquereux, Harvard ColI. 

Mus. Compo Zoology Bull., vol. 16, p. 46, 1888. 

Two specimens from Golden were referred by Lesquereux to this species without comment. The best 

preserved of these is shown in Plate 18, Figure 2, and 

is a leaf that must have been about 9 centimeters long 

when perfect, but it now lacks practically all of the 

margin except for the length of about 1 centimeter 

near the middle. The margin here is seen to be distinctly toothed, the teeth low, sharp-pointed, and regular. The nervation consists of a slender straight 

midrib and some 8 or 10 pairs of very slender secondaries that are at irregular distances and much curved 

upward and apparently camptodrome. The finer 

nervation is not preserved. 

In shape and size the leaf here figured resembles the 

smaller of the two leaves figured by Lesquereux U 

under the na,me QUe'l'CUS haidJingeri Ettingshausen, but 

it differs strongly in the secondary nervation. The 

le~f originally figur~d by Ettingshausen from the 

1\1iocene of Austria is narrower than either of the 

leaves figured or identified by Lesquereux and has 

sharp, upward-pointing teeth, whereas the American 

leaves have the margin dosely serrate with the teeth 

outward-pointing. 

There is another complication regarding the leaves 

figured in the report on the Tertiary flora. 'V'hen 

they were first mentioned 85 they were said to have 

come from Green River, Wyo., but subsequently they 

were vaguely thought to have come from the Denver 

a.t Golden, Colo. They are certainly not the same as 

the European type of this species and considering the 

uncertainty of their locality are of little value. 

The leaf from Golden under discussion that was 

identified by Lesquereux as Quercus haidinge1'i Ett.ingshausen is not identical with the European type, 

IJor is it the same as the American leaves discussed 

above. It seems best, therefore, to give it a new nanle, 

especially as it appears to be different from anything 

previously noted. 

Occurrence: Denver formation, Golden, Colo., collected by Arthur Lakes. Type in Museum of ComB4 Lesqucreux, Leo, The Tertiary fiora, pI. 20, fig. 9. 1878. 

311 U. S. GeoI. and Geog. Survey Terr. Buil., yoJ. 1. p, 387, 1875, 

[1876]. 
54 l'LORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

parative Zoology, Cambridge, l\t1ass., No. 1625; one in 

United States Nationall\tluseum, No. 1278. 

Quercus celastrifolia Lesquereux 

Plate 18, Figures 4-7 

QuerOlls oelastrifolia Lesquereux, Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 46, 1888. . 

Qtierous strwl1'11inens? Lesquereux. Lesquereux, op. cit., p. 46. 

Lesquereux's description and reInarks concerning 

this species are as follows: 

Leaves subcoriaceous, oval, equally narrowed at. both ends, 

obtusely pOinted; secondaries yery oblique, distant, parallel 

from the base, curving in passing toward the borders,' ascending along them, nearly simple, passing upward under the teeth 

and joining them by short branches. The. teeth 'are acute, 

turned upward, one at the end of each of the secondaries, which 

are subopposite in five 0'1' six pairs; llervilles thin, at right 

angles to the median nerve, except near the borders, where 

they turn upward like thin tertiary nerves. The leaves are 

5 to 6 centimeters long, 3 to 3.5 centimeters broad; the angle 

of divergence 300 to 350 • 

This species is based on foul~ specimens, all of which 

nre here figured. The best-preserved one is shown in 

Figure 6 and is nearly perfect. It is slightly more 

than 5.5 centimeters long and 3.5 centimeters wide. It 

is entire for the lower third and thence provided with 

few low, apparently spine-tipped teeth separated by 

very shallow sinuses. There are five pairs of nearly 

opposite secondaries at an angle of about 60°, which 

end in marginal teeth. The specimen shown in F~glire 7 is nearly 6 centimeters long and fully 3.5 centiIneters wide. It has the lower secondaries branched 

. but does not otherwise differ frOln the one just described. The smallest specimen, Figure 5, is but little 

more than 4 centimeters long and about 3 'centimeters 

wide; it shows the base well preserved. . 

A single small leaf from Golden that was referred 

by Lesquereux 86 with question to Quercus str(Jfft1inews 

Lesquereux is shown in F.igure 4. He says of it: 

The leaf which I refer to this. species is oval, apparently 

denticulate near the apex, and narrower than those figured 

in the United States Geological Survey of the Territories [Tertiary ft.ora] , VII, Plate 19, Figures 6, 7. It may therefore 

belong to a different species. 

On comparing this leaf with the types of Que1'cus 

ceZastrifolia I am unable to note any essential difference .. It is smaller, being only about 4.5 centimeters 

long and 2.5 centimeters wide, but has the same low, 

sharp teeth and the same kind of nervation. 

The type locality of Querou8 stratrnme'l."8 is in the 

Laramie formation at Golden, Colo., and so far as 

known it has not been found at any other horizon. 

36 Lesquet'ellx, Leo, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, 

p. 46, 1888. 

Occurrence: Denver formation, Golden, Colo., collected by Arthur Lakes about 1883 for the Museum 

of Comparative Zoology, Cambridge, ~1ass.; types, 

Nos. 1071, 1075, 1626. 

Q:uercus coloradensis Lesquereux 

Querous colo1·a.densis Lesquereux, Harvard Coll. Mus. Compo 

Zoology Bull., vol. 16, p. 46. 1~88. 

Leaves subcoriaceous, entire, oblong, obtuse, rounded at tbe 

base, abruptly decurring in joining [to join] the' petiole 

(broken) ; secondaries five to six pairs, at an acute angle of 

divergence, camptodrome, the lower ones branching, aU COIlnected by distant though thin nervilles and running high up 

along the borders, which are parallel in the middle. 

By the size and form of the leaves-5 to 6 centimeters long, 

3 to 4 centimeters broad in the middle, where they are somewhat contracted-the species is related to Quel'Olt8 oread'l.w", 

Saporta.31 

This species is based on two examples supposed to be 

ill the Harvard College Museum of Comparative Zoology, but only one has been located, and this is so 

fragmentary that it was not thought worth figuring. 

Occurrence: Denver fornlation, Golden, Colo., collected by Arthur Lakes in 1883 for the ~t[useU1l1 of 

Comparative Zoology, Cambridge, Mass. 

Order SALICALES 

Family SALICACEAE 

Populus zeiIleri (Lesquereux) Knowlton, n. comb. 

Plate 18, Figure 8; Plate 19, Figures 1-3 

Proto{iO'U8 Zeille1"i Lesquereux, Harvard CoIl. Mus. Compo 

Zoology Bull., yol. 16, p. 50, 1888. 

Following is Lesquereux's original description and 

discussion: 

. Leaves of medium size, coriaceous, rugose on the surface, 

enlarged. and round-cordate at base,. deltoid at the acute apex, 

palmately 3-nerved to 5-nerved from the top of the petiole; 

lateral nerves lDuch branched; borders crenulate. The leaves, 

deeply rugose 'by the impression of strong nervilles, are 6 to 

7 centimeters long, 5 to 5.5 centimeters broad below the middle; 

primary nerves three or more, generally five, the lower at a 

broad angle of diyergence, following the borders, the inner 

ascending in a curve somewhat inclined to the midrib; secondaries two or three pairs at a great distance from the base. 

The borders, mostly destroyed, are seen crenulate at the few 

places where they are preserved.. 

This species was based on five specirp.ens, all of 

which are before me and four of which are here figured. Three of these specimens are uftdoubtedly conspecific, but the other (pI. 19, fig. 1) is less certainly 

so. Lesquereux states that the leaves are" palmately 

3-nerved to 5-nerv.ed from the top of the petiole," but 

81 Saporta, Gaston de, Flora de Cumi: Ecole normale sup~rieure 

Annales sci., 2d ser., vol. 2, p. 337, pI. 2, fig. 11, 1873. 
SYSTEMATIC PALEOBOTANY 55 

a glance a t the foul' specimens above mentioned shows 

that they nre essentially 3-ribbed, with a strong, 

straight midrib and a single pair of nearly as strong 

lateral ribs that curve upward and nearly parallel the 

midri b, each with secondary branches on the outside, 

the lowest arising near the top of the petiole and thus 

producing a sort of false 5-ribbed appearance. In the 

fourth specimen, however (pI. 19, fig. 1), there are five 

distinct ribs, the lowest with three or four secondary 

branches on the outside. The inner pair of ribs do 

llOt pnraUel the midrib, like the others, and their secondary branches, four or five in number, arise some 

djstm;ce above the base. ' 

The' margin in aU these leaves is very obscure1 y 

preserved. It is described by Lesque~'eux as undulate 

or crenate, but in most cases its chara,cter is extremely 

difficult to determine. In one specimen (pI. 19, fig. 

1), which happens to be the anomalous one above 

n1entioned, the margin is provided ,vith scattered low 

teeth which are entered by the secondaries or by nervines from them. l'he only other example in which 

the teeth are pre!Served is that shown in Plate 18, 

Figure 8, and in this only for a shor~ distance just 

a bove the base; the teeth seem to be 1110re rounded 

than in the other one mentioned. 

The question of the generic reference for these 

leaves may be considered. The genus P'roioficus was 

established by Saporta 88 on foul' species .from the 

travertines of Sezanne, the type apparently being 

Protoficus· sez·annensis nVatelet) Saporta (Ficus 

sez((lnne1UJis V,T atelet) . This a large, broadly oblong 

l(~af. with entire margins. It is evidently wedgeshaped at the base and· pinnate, with an extremely 

strong midrib and numerous mainly alternate secondaries which emerge nearly- at a right angle. 

. A. second species, Proto ficus insignis Saporta, is· of 

the same type as Protoficu8 sezaJn/nensis, from which 

it. differs in its more oval sha.pe and upward-curving 

secondaries. 

A· third species, P'rotofic~(;8 crenulata Saporta, is 

grently different and can .hardly be congeneric with 

the others, being ovate with a finely crenulate margin and 3-ribbed from the top of the petiole. It was 

evidently this species that Lesquere~lx had in luind 

when he referred the Golden leaves to P'J'otoficus, but 

these leaves differ so widely from the French species 

. that they can not possibly be congeneric. In p'J'otoflmt8 ()1'enailata the lateral ribs arise at an angle 

of about 45° and pass with very little upward curving to the border, being together with the secondary 

branches distinctly camptodrome, whereas the Golden 

leu-ves have the ribs arising at a more acute angle and 

ru; Snportll, Gllston de, Prodrome d'ulle fiore fossile des tl'avertins 

(1I1<.'I('l1S de S~zIlJlnc: Soc. g~o1. l,'rance Mem., 2d s0r., vol. 7, p. 355 

(OJ), 1868. 

curve abruptly upward to parallel the midrib, ending, 

ns to the secondary and tertiary branches, in the margin. These Golden leaves are of 'course totally unlike 

the type of P'rotoficu8. 

The question next arises as to the generic reference 

of the leaves described by Lesquereux as PTotoficus 

zeille1'i. In size, shape, margin, and particularly in 

the manner in which the lateral ribs curve upward and' 

parallel the midrib, they are at once suggestive of certain Tertiary species of Popul~, such as Populus 

speciosa Ward,89 P. neb'J'ascensis Newberry,40 and P. 

8'milacifolia Newberry,41. all from the Fort Union for- . 

Iuation. In fact, I do not see how they can be sepa~ 

rated from the genus Popul~, nor is it apparent how 

they can be adequately distinguished fron1 the type 

leaves of Pop'l.il/us 'J'U3bTaJ8Censis . . However, as the n1argin is so obscurely preserved in the leaves under con5ideration it may be as well to refer them to Pop~tlus 

but let the species stand under Lesquereux's nan1e. 

Comparison has been made with the genera G'J'ewia, 

Grewiopsis, Fic~, Zizyph~, Srnilaw, etc., and 

although there are certain points of resemblance with 

all these genera the Golden leaves differ in some essential manner. 

Occurrence: Denver formation, South Table ~Ioun- . 

tain, Golden, Colo., collected by Arthur Lakes:' Types 

in l\1useum of Comparative Zoology, Cambridge, 

]\tIass., Nos. 1671, 1672, 1674,1675. 

Populus nebrascensis Newberry 

Populwi nebrascensis Newberry, New York Lyceum Nat. Hist. 

Anuals, vol. 9, p. 62, 1868; Illustrations of Cretaceous 

and Tertiary plants, pI. 12, figs. 4, 5, 1878; U. S. Geol 

Survey Mon. 35,p. 47, pI. 27, figs. 4, 5, 1898. 

Lesquel'eux, .Harvard ColI. Mus. Compo Zoology Bull., vol 

16, p; 47, 1888 . 

PO]JUht8 crenata Uuger. Lesquereux, Harvard ColI. :Mus. 

Oomp. Zoology Bull., vol. 16. D. 48. JRR8. 

. Following is Newberry's original description: 

Leaves long-petioled, 2 to 3 inches long, ovate, pointed, regu· 

larly rounded at base, coarsely and irregularly toothed except 

near the base, where the margins are entire; nervation strong, 

radiating from the base of the leaf; medial nerve straight, 

simple (or supporting very small nerves), except nea r the 

summit, where two. or three larger' branches arise from it; 

lateral nerves, two pairs on each side, springing from a com .. 

mon point of origin; lower pair arched upward, nearly parallel 

with the margin of the leaf, to' which they send off one or 

more simple branches; second pair of laterals diverging from 

these at an angle of 30°, arching upward, and parallel with 

the midrib, terminating in the margin near the summit, each 

giving off about three exterior branches, which curve upward 

and terminate in the dentations of the border. 

30 Ward, L. F., Synopsis of the flora of the Laramie group: U. S. 

Ceol. Survey Sixth Ann. Rept., p. 550, pI. 34, figs. 1-4, 1885. 

40 Newberry. J. S., The later extinct floras of North America: U. S. 

GeoJ. SurveyM'on. 35, p. 47, pI. 27, flgs. 4, 5, 1898. . 

n Idem, pI. 29, fig. 5. 
56 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

This is one of the most abundant and widely distributed Fort Union species and has been well described and figured by Newberry. Curiously enough, 

this species is extremely rare, or rather extremely 

local, in its occurrence in the Denver formation; in 

fact, it has been found at only a single locality, though 

from this place about 300 specimens of it were procured. Although 'excavations were made only a few 

,rods away, not a single specimen of it has been found 

among the many thousands that have been collected. 

This semllS to prove, as Lesquereux has pointed out, 

that the leaves were not transported from any distance 

but, as he says, had "fallen from trees either around 

woody swamps or on the borders of shallow lakes." 

From the fact that only a single very circumscribed 

locality in the Denver Basin has thus far been noted 

for these leaves it seems a reasonable inference that 

they all came from a single tree or at most a small 

clump of trees' and presumably represent a single 

species. There is so much diversity to be noted in 

these leaves, however, that Lesquereux has seen fit to 

divide them up into a number of varieties, some of 

which are indeed so different from what may be called 

the typical form that had they been found at different 

localities and horizons they would probably have been 

designated as full species. 

Lesquel'eux's names and descriptions of these varieties 42 are given below. None of them l].ave previously been figured. 

Populus nebrascensis var. grandidentata Lesquereux 

Plate 19, Figures 4-7; Plate 20, Figures 2, 6, 7 

Leayes broader, rounded, and undulate, toward the base; 

borders cut from the middle ,upward in large deltoid obtuse 

gradually longer teeth. Some of the leaves are subtruncate 

at apex with long irregular teeth; others are rapidly narrowed to an obtuse apex. The nervation is the same in the 

varieties as in the normal form, 3 to 5 palmate from the base, 

with the inner pair of primary nerves curved inward and ascending to near the apexes and the secondary ones at a great 

distance from the base. 

This form was represented by 85 specimens in the 

collection studied by Lesquereux for the Harvard College Musemn of Comparative Zoology. The National 

Museum possesses about a dozen specimens in the 

Lacoe collection, obtained by Arthur Lakes at the 

same locality. Two nearly perfect specimens have 

been figured. The larger one is fully 11 centimeters 

long and a little over 8 centimeters wide. It is 

rounded and nearly truncate at the base and has the 

margin cut above into nuinerous large irregular teeth. 

The nervation is very strongly marked, especially the 

midrib and the inner pair of secondaries or ribs. On 

42 Lesquereux, Leo, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, 

pp. 47, 48, 1888. .' 

one side there are tv"o strong secondaries springing 

from near the top of the petiole, thus making the leaf 

)n effect 6-ribbed. Another example figured is COllsiderably smaller, being only about 7:5 centimeters 

long and 7 centimeters wide; it has fewer but relatively larger teeth and is 5-ribbed from the top of the 

petiole. 

Populus nebrascensis var. rotundata Lesqueretix 

Plate 20, Figures 1, 3, 4, 5, 8 

Much like the preceding, differing by its broader leaves, 

broadly rounded and enlarge~l at the base. 

This was represented by 48 specimens in the, collection studied by Lesquereux and by two specimens, both 

of which are figured, in the National Museum. The 

snla.ller specimen (fig. 1) is 5 centimeters long and 5.5 

centimeters wide and has the petiole preserved for a 

-length of 4.5 centimeters; it has rather few small 

sharper-pointed teeth. The other specimen (fig. 8) 

is about '7 centimeters long Hnd 6.5 centimeters wide. 

n is nearly circular and has rather small sharp teeth. 

Three additional examples from 'the original collection in the Harvard College ~1useum of Compai'ative 

Zoology are also figured. 

Populus nebrascensis var. acutidentata Lesquereux 

Plate 21, Figures 1, 2 

Leaves oval, narrowed at base, lanceolate above, generally 

palmately trinerved, teeth of the borders large, equal, sharply 

pointed. 

This form is represented by 13 specimens in, the 

original collection now in the Harvard College 

Museum of Comparative Zoology but was not noted 

in the National l\1useull1 material. Two of the best 

preserved of the types are figured. Both are large 

leaves, the one shown in Figure 1 being about 8.5 

centimeters long and 7 centimeters wide. It is nearly 

entire for the lower third and thence cut into large, 

equal, sharp teeth. The nervation does not differ from. 

that of the other forms., The other specimen (fig. 2) 

is about 8 centimeters long and a little over 5. centimeters wide; it has the same large, sharp teeth. 

Populus nebrascensis var. longifolia Lesquereux 

Plate 21, Figures 3-5 

Leaves large, Oblong-ovate, rounded at base, 5-nerved; lateral 

primary nerves ascending to abovE' the middle; second curved 

inside, much branching; secondary nerves three or foul' pairs, 

at a great distance from the base, camptodrome or craspedodrome, with their divisions, effaced at the borders, which are 

cut in obtuse large teeth. The leaves are long, lanceolate 

from below the middle, 7.5 to 11 centimeters long, 4 to 7 centimeters broad below the middle; the teeth are large, round, 

equal, marked from near the base; the petiole is long and 

slender. 
SYSTEMATIC l' ALEOBOTAN Y 57 

This form, accordil)g to Lesql1el.'cux, :is represented 

\by 13 specimens in the collections of the Museum of 

Comparative Zoology, three of which are here figured. 

The one shown in Figure 5 is broadly ovate, 7.5 centimeters long and a little over 6 centimeters wide below 

the middle. It is a little prolonged above and has the 

marginal teeth large, regular, and rather obtuse. 

With a base of much the same shape but with the 

upper portion more elongated is the leaf shown in 

Figure 4, which is 8.5 centimeters long and less than 

6 centimeters broad; it has very few teeth. The other 

specimen, shown in Figure 3, is a very narrow leaf 7.5 

cent.jmeters long a~d only about 4 centimeters wide; 

ita 1so has few teeth. 

I..esquel'eux referred a number of leaves, in the collection studied by him for the Harvard College 

Museum of Comparative Zoology, to Populus cre'fWtta 

Unger,68 from the Miocene of Sotzka. On comparing 

the original figures of this species with the leaves from 

Golden it is seen that they agree somewhat as to size 

and outline but differ absolutely in nervation. Populus crena/)a is distinctly pinnate, but the leaves under 

consideration from Golden are distinctly palmate and 

3-ribbed from the top of the petiole. . So far as I can 

determine they are referable to Populus neb'J'ascensis. 

In 1888 Lesquereux 44 established Gissus duplicatoserrata on some seven specimens .from Golden, Colo., 

in the Museum of Comparative Zoology. After chaTacterizing the species he stated: 

'l'11e leaves are referable to Oi.sS1t8, though they have a degree 

of nftlnity with some varieties of P01)'ltZ1lts nebrascensis Newberry. They differ essentially by the primary lateral nerves 

not incUl'ved, much branched, all the divisions craspedodrome, 

the teeth acutc, the substance of the leaves thick. 

I have studied these specimens carefully and am 

at a loss to find any essential character by which 

they can be separated from certain of the forms 

of Poprulu-s nebralScensis. The type specimens of 

Gissus d'ltplicato-se'J"J'ata are all fairly well preserved, 

so there is little difficulty in comparing their characters. Foul' of the best have been figured.. In regard to shape and size they have, it may" be seen, 

the SaIne appearance as Popuil?ts nebrascen8'is-that is, 

they fall easily within the limits of one or another 

of the forn1s of t.his species. The absence of incurving in the primary lateral nerves is not more marked 

than in many leaves of Porntl'ltB neb1'ascensis, nor is 

the secondary branching of these ribs Inore marked. 

Of the four figured types of Oissus dupUcato-ser-

'J'ata, the one shown in Plate 20, Figure 2, is referred 

43 1 Tnge·!·, IJ'l'nnz, Fossllc Flora yon Sotzka, p. 36 (166), pI. 15 (36), 

figEl. 2-5, 1851. 

u I.esf]nel'enx, Leo, lIn l'vn I'd Coli. )IllS. Compo Zoology BulL, vol. 16. 

p. :i:!. 1888. 

to PopUlus nebra~censis grandidentata, and the others 

(pI. 20, figs. 3-5) are referred to Popul!us neb'J'ascensis 

'J'otwndata. . 

At least two of the specimens identified by Lesquereux 45 a~ Gi~sus p(urrottiaefolia Lesquereux, from 

Golden, Colo, are also referred to PopUlus nebrascensis rotundat((;. They are small leaves that do not 

otherwise differ from this form. 

Occurrence: Fort Union formation, mouth of Yellowstone River (type locality). Denver formation, 

South Table Mountain, Golden, Colo., collected by 

Arthur Lakes; a few specimens in the Lacoe collection, now in the United States National 1\1useum, and 

a large number in the 1\luseum of Comparative Zoology, Cambridge, ::l\1ass. The specimens figured are 

all from the Denver formation and are as follows: 

Populus neb'J'ascensis val'. 'J'otundata, Nos. 1219, 1223, 

1226; val'. acutident((;ta, Nos. 1210, 1262; val'. longi-

;olia, Nos. 1239, 1244, 1245. Types of Gissus rlIuplicato-se?'rata Lesquereux= P. nebrascensis rotwndata 

(Nos. 1498, 1499, 1505) and P. neb'l'ascernsis !Jl'andidentata (No. 1683). 

Populus? sp. 

Plate 21, Figure 9 

POIJttl1('S aiJt(JIHHI,ta AI. Bmun. Lesquel'eux, Harvnrd CoIl. 

:Mus. Compo Zoology Bull., vol. 16, p. 48, 1888. 

Leaf very thick and coriaceous, broadly elliptical 

01' broadly ovate, apparently truncate at the base, 

truncate at the apex; margin with about seven large, 

low, rounded teeth or lobes separated by shallow 

sinuses; 5-ribbed from the base, the midrib very 

strong below, much more slender above, with a single 

pair of secondary branches; next (inner) pair of 

ribs very strong~ at an angle of about 500 , straight, 

ending in marginal lobes, each with several slender 

secondary branches; lower (<wter) ribs strong, at 

an angle of about 350 , perfectly straight, craspedodrome; nervilles numerous, light, forming large rectangUlar areas. . 

Lesquereux identified two leaves in the Harvard 

College Museum of Comparative Zoology as Populius 

attenuata AI. Braun, the one here figured and another which is totally different and belongs with what 

he called Gissus duplieato-dentata. The present specimen is fragmentary in that it lacks all of the basal 

portion. It was evidently a very thick leaf, with five 

extremely 8trong, straight ribs apparently from the 

top of the petiole. It appears to have been about 4.5 

centimeters .long and 5.5 centimeters or lnore broad. 

It has only a few broad, low, rounded lobes, each of 

,- Idem, p. 52. 
58 l!'LORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

which is entered by a rib or a secondary branch from 

a rib. 

Populus attemudta AI. Braun, with which this was 

identified by Lesquereux, is a common and widely distributed species in the European Miocene-for instance, in the Swiss Miocene. Heel' 46 has figured 

more than 20 specimens' in his Swiss flora. Although 

there is considerable range in variation, some of them 

somewhat resemble the present specimen in the shape 

of the upper portion, though the marginal teeth are 

small and numerous, but they differ absolutely in 

nervation. P9'pulu:s attenuata is essentially pinnate, 

though in some specimens the lowest pair of secondaries is somewhat strengthened and produces a pseudo 

3-ribbed aspect. The present leaf, however, is distinctly palmately 5-ribbed from the top of the petiole, 

and all the ribs are distinctly craspedodrome, instead 

of being largely camptodrome, as in P. (J)ttenuata. It 

can not.possibly be referred to P. attenuata, and there 

is much doubt as to its even belonging to Populus. It 

suggests certain forms of Hedera, but as it is so fragmentary that its full character can not be made out 

it is permitted to remain with question in Populus. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Type, 

originally under the name PopUlus attenuata; AI. 

Braun, in the Museum of Comparative Zoology, Cambridge, Mass., No. 1286. 

Populus jacksoni Knowlton, n. sp. 

Plate 21, Figures ~; Plate 22, Figure 7 

POP1t~'u8 arctioa Heel'. Lesquereux, Harvard CoIl. Mus. Compo 

Zoology Bull., vol. 16, p. 47, 1888. 

Leaves coriaceous, small, nearly circular or broadly 

ovate, truncate or slightly heart-shaped at the base, 

rounded or pbtusely pointed at the apex; margin from 

nearly entire to crenulate or fairly well toothed, the 

teeth obtuse; petiole long and slender; 5-nerved from 

the base, midrib with two or three secondary branches 

in the extreme upper part, next pair of ribs arching 

inward and reaching the. margin near the apex, usually with several outside branches; outer ribs slender, 

those and all branches craspedodrome; nervi lIes numerous, very strong. 

This species, under the name P OpUlU8 Ulf'ctica Heer, 

is said by Lesquereux to be represented in the Harvard College Museum of Comparative Zoology by 15 

specimens. Of these, four have been selected to show 

the range in size, marginal configuration, and nervation. They ar~ all small leaves from 2 to 4 centimeters in length and about the same in width. The' peti.: 

ole, preserved in only one specimen, was at least 2 

centimeters long and very slender. 

46 HeCi', Oswald, Flora tertiaria Helvetiae., vol. 2, p. 15, pI. 57, figs. 

8-12,1856. 

This species is undoubtedly closely 'related toindeed, possibly identical with-certain leaves that 

have been referred to PopulJus arctioa Heer. For example, Lesquereux 41 ~tated that except that these 

lE:aves 

are generally more distinctly crenulate, nothing in the characters fndicates a diiference from those which I have flgured in 

United States Geological Survey of the Territories [Tertiary 

flora], Plate 23, Figures 1-6, or of those in Heel', Flora fossilis 

arctiea, vO]. 1, pI. 4, flg. 6a. 

Examination of the specimens figured by Lesquereux shows that there is very considerable diversity in 

size, shape, and nervation, and it 'has been necessary 

to separate them into two and possibly three species; 

moreover, it is to be doubted if any should be regarded 

as true Popwl;u;s arctioa. The present species seems to 

ffie to be abundantly distinct from any figured by 

Lesquereux as Populus arctic~; in fact, it is much 

closer to certain leaves of Populus decipiens Lesquereux 48 (now called P. a1'ctica decipiens by Cockerell), 

except that the latter has the leaves always entire and 

more wedge-shaped at the base. 

Populus jacksoni is suggestive of certain very small 

leaves of P opul;us arctiaa, such as that shown in Heer's 

Plate 4; Figure. 6a (Flora fossilis arctica), but from 

the ordinary leaves of this species, such as shown by 

Heer on his Plate 5, there is marked divergence. 

Populus arctica w.as described originally. from the 

Miocene of Greenland and has since been found to be 

very widely distributed throughout the Arctic lands. 

It is especially abundant in the Kenai formation 

of Alaska and has been repo'rted :from the Eocene of 

Canada and the Fort Union and Lance formations of 

Montana and Wyoming. It is probable, however, that 

when all this material is critically studied more than 

one species will be found confused under this name. 

I take pleasure in naming this species in honor of 

Dr. Robert T. Jackson, of Harvard University, who, 

as custodian of the fossil plant collections, has generously placed this Golden material at my disposal. 

Occurrence: Denver formation, South Table Mountain' Golden, Colo., collected by Arthur Lakes. Types 

in the Museum of Comparative Zoology, Cambridge, 

Mass., Nos. 1077, 1083, 1084, 1085, '1441. 

Populus tenuinervata Lesquereux 

Plate 22, Figures 1-4, 6 

Populus tenuinervata Lesquereux, Harvard CoIl. Mus. Compo 

Zoology Bull., vol. 16, p. 48, 1888. 

Lesquereux's original description and remarks concerning this species are given on. page 59. 

47 Lesquereux, Leo, Harvard ColI. Mus. Compo Zoology Bull., 'Vol. 16, 

p. 47, 1888. 

4B Lesquereux, Leo, The Tertiary fiora, p. 179, pI. 23, figs. 7-11, 

1878. 
SYSTEMATIC PALEOBOTANY 59 

Leaves comparatively smull, round 01' o,'ate, broadly cuneate 

01' rounded at ba,se, palmately 5-nerved; inner primary nenes 

curving inward and ascending near the apex; those of the 

outside nlso curvillg and ascending to the middle, all campto- ' 

<'il'ome, thin but distinct; borders irregularly dentate, the 

teeth unequal and pointed; nervilles distinct, obliquely joined 

to the midrib. 

The leaves resemble those of a Pic1ls, being, in their facies, 

like those of IN,()Us crena,ta Unger, which, however, has not 

distinct nervilles. They have a still more marked likeness to 

those of PopuZ'ns lat'ior-t1'a1ts'V(}1'sa Heel', as figured by Ludwig 

in Palaeontographica, vol. 5, pl. 26, fig. 3. The petiole is 

thick. 

According to Lesquereux this form is represented 

in the collections of th.e ~{useum of Comparative 

Zoology by no less than 34 specimens. I have s~en 

and studied these specimens and have selec:ted for 

illustration five examples that seem best to show the 

range in size and general app~arance. They are all 

Heady circular, The smallest one, shown in Fifrure 2 , b , 

is about 2.75 centimeters long and the same in width. 

It is entire for at least the lower third of the marO'in b 

nnd thence provided with few, apparently sharp teeth. 

The next in point of size, sho,vn in Figure 6, is 4.5 

centimeters long and about 4 centimeters bl'oad. It 

has a few scattered teeth below, and larO'er b ones abo,~e , 

all sharp-pointed. A small portion of the petiole pre'-

served is rather thick for the size of the leaf. The 

specimen represented in Figure 4 is of about the same 

s:ize as the last and was selected because the larger 

pail' of ribs diverge with little lncurving in passing to 

the upper margin. The leaf shown in FiO'ure 1 was w • b 

D centImeters or mOl'elong and about 4.5 centimeters 

broad; its petiole is apparently preserved entire .and 

i~ 3.5 centimeters long and very slender. The largest 

leaf observed, shown in Figure 3, is nearly 6 centimeters long and 5 centimeters wide. It is entiremargined below and has n, few relatively very large 

sharp teeth in the upper portion. 

This species is undoubtedly most closely related to 

PO'[YWl'UJ8 neb7'ascensis Newberry; in fact. it is in cer-

. tain respects doubtful if they should be separated. 

For instance, it is almost impossible to separate this 

spec-ies from certain of the smaller leaves referred to 

Populusneb?'(zsoensis rotundata Lesquereux, as may be 

seen on comparing the,two sets of figures. In general, 

h.owever, the present species may b,e known by its small 

S.lze, nearly circular shape, few, equal sharp-pointed 

HHtrginal teeth, and thin, rather delicate nervation. 

In the diagnosis Lesquereux states that the nerves are 

camptodrome, but so far as I can determine they are 

all craspedodrome and end in the marO'inal teeth. 

The petiole is slender in the 'only specim:n in which ,t is fully preserved. ' _. 

Occurrence: Denver formation, Golden, Colo., colkcted by Arthur Lakes. Types in ~{useum of Comparative Zoology, Cambridge, ~{ass.: Nos. 1252, 1256, 

1~66, 1274, 1513. 

PopUlus subrotunda Lesquereux 

Plate 22, l!"igure 9 

'Popul1ls Sttb1'otwllda Lesquereux, Am. Jour. SeL, 2d ser., vol. 

45, p. 205, 1868; U. S. Geol. Survey Terr. Ann. Rept. for 

1869, p. 196, 1869; idem for 1870, p. 380, 1871; idem for 

1876, p. 506, 1878. 

Pop1('hM3 attelHta,ta, AI. Braun. Lesquereux. U. S. Geol. and 

, Geog. Survey Terr. Ann. Rept. for i872, pp. 386, 389, 

392, 1873. 

Pop-ulu:s subl'otu,ndata Lesquereux [apparently a misspelling 

for SUbl'otltnda], The Tertiary flora, p. 173, pI. 24, figs. 

6-8,1878. 

Dawson, Canada Geol. Survey Rept. Progress for 1877-78, 

p. 186B, 1879. 

Two of the figured types of this fine species-the 

originals of Plate 24, Figure? 6 and S, in the" Tertiary 

flora "-from Carbon, Wyo., are preserved in the 

United States National ~{useum (Nos. 220,221). The 

other specimen-the original of Figure 7-said to 

have come from Evanston, ';Vyo., can not be located, 

nor can the specimens reported to have come from 

"Rock Creek on the Laramie Plains." 

As Lesquereux has pointed out, Populus subrotunda 

is apparently closely related to Pop11lAts atte'lllUata AI. 

Braun, especially as figured by Heer 49 from the 

Tertiary of Switz~rland. The American leaves, however, are more nearly round, with a less pointed apex, 

sharper, upturned teeth, and more truncate base; the 

nervation is the same in both. Populus atten,uat(JJ has 

been reported by Lesquereux 50 frOln the Denver for-' 

mation at Golden, Colo., but I have not seen the specimens, two of which are mentioned. " .. 

Among the specimens more recently procured from 

the Denver formation there is a single nluch broken 

example which appears to belong to Popwlus subrotwnda. It is smaller than either of the figured types 

and appears to have only two pairs of secondaries on 

the midrib, in this respect a!!reeinO' with one of Heer's 

fi ~ 0 

gures of P. attenuata. . , 

Among various specimens from the Lance formation of Converse County, Wyo., is a'single very 

perfect example that I am unable to distinguish from 

ccr~~in leav~s of Po'prul1JJ8 suorotwnda. In this diagnOSIS of thIS species Lesquereux stated that "the 

nervation is tripalmate, camptodrome." 'In the specim~n under cons~deration the nervation is' distinctly 

trIpalm~te, but It is not camptodrome, nor for that 

matt~r IS at least one of the type specimens (TertIary 

flora, pI. 24, fig. 6), for although rather" faint the 

43 Heel', Oswald, I!'IOl'fl t~l'tial'ia Helvetiae, vol. 2, p. 15 PI~ 57 'lIgR. 

8-12, 1856. ' , 

50 Lesquereux, Leo, Harvard Coll. Mus. Compo Zoology Bull., vol. 16. 

p. 48. 1888. 
60 li"LORA OF THE DENVER AND ASSOCIATED FOHMATIONS OF COLORADO 

secondary branches may be seen distinctly entering 

the marginal teeth. The Converse County leaf is 

intermediate in si~e between those of Figures 6 and 

1, above mentioned, but the basal secondaries arise at 

a lower point than in Figure 7, though not lower than 

in Figure 6, and the branches from the lower side of 

the ribs clearly enter the marginal teeth. Otherwise 

it is not possible to distinguish between them. 

Occurrence: Hanna formation, Carbon, Wyo. 

(types). Evanston formation, Evanston,. Wyo. 

(type) . Denver formation, northwestern base of 

Green Mountain, Golden, Colo. Lance formation, 

gulch south of Lightning Creek and opposite Box 

Cr.eek, Converse County, Wyo. 

Populus zaddachi Heer? 

Plate 22, Figure 8 

Poplllu .. '1 ,:'addacM Heel', in Zaddach, Phys.-okon. Gesell. 

Konigsberg Schriften, vol. 1, p. 29, pI. 4, 1860 [1861]. 

Heel', Flora fossilis arctica, vol. 1, p. 98, pI. 6, figs. 1-4, 

1868. 

Lesquereux, U. S. Geol. and Geog. Survey T'err. Ann. 

Rept. for 1871, p. 292, 1872; Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 47, 1888. 

In the material collected in Middle Park, Colo., I 

have found a leaf with its counterpart-the one here 

figured-that appears to belong to this species. It 

was at one time identified by Ward as Zizyph!us '/'i1..eekii 

Lesquereux, which is not referred to PopwZus, but it 

seems in both shape and nervation to be more like 

Populus zaddaoh:i. It is about 5 centimeters long and 

3.5 centimeters wide, is rather deeply heart-shaped at 

the base, and has the margin provided with strong, 

rounded teeth. The nervation is strongly marked and 

is well shown in the figure. 

Populus zaitdaohri is essentially a Miocene species, 

though it has been reported from beds somewhat older. 

For instance, in the Hayden annual report for 1871 

Lesquereux reported the probable presence of Populus 

zadx100hi at Evanston, Wyo., "below the coal." It 

was omitted by Lesquereux in his final report on the 

Tertiary flora, however, and has not since been detected with certainty at Evanston. 

Populws zaddachV- is an abundant and characteristic 

~pecies in the auriferous gravel of California 51 and 

has also been reported by Lesquereux 52 from Florissant, Colo. Considering the fact that the specimen 

under discussion is somewhat obscure, its positive 

identification as Popu,zw zaddachi should perhaps be 

questiqned. 

[;l Lp.squereux, Leo, Harvard Co11. Mus. Compo Zoology Mem., vol. 6, 

])1. 8, figs. 1-8, 1878. 

62 Lesquereux, Leo, The Cretaceous and Tertiary floras, p. 158, pI. 31, 

fig. 8, 1883. 

Occurrence: ~1iddle Park formatiol1~ Mount Bross, 

Middle Park, Grant County, Colo., debrts on slope, 

collected by G1eo. L. Cannon. Miocene (auriferous 

gra vel), California. 

Populus lacoeana ](nowlton, n. sp. 

Plate 22, Figure 5 

Leaf coriaceous, very broadly ovate or almost circular, broadly tnlncate and slightly cordate at the 

base, abruptly rounded above to obtusely acuminate 

apex; margin strongly and somewhat irregularly 

toothed, the teeth obtuse and rounded and separated 

by shallow or deep rounded sinuses; petiole very 

strong, of unknown length but 3 centimeters preserved, 

splitting at the very base of the blade into three nearly 

equal ribs; of these ribs the· central or lnidrib is 

straight and terminates in the'tip of the blade; the 

midrib has one or two secondary branches in the extreme upper part; the lateral ribs arise at an angle of 

about 45°, then turn upward, and run nearly parallel 

to the midrib and terminate in large marginal lobes; 

each rib has about six secondary branches on the 

outside, the two lower ones arising together at the 

extreme base, thus simulating a· seIniribbed appearance; the secondary branches are strong, the lower 

ones camptodrome and the upper ones craspedodrome; 

nervilles numerous, strong, often arched but mainly 

unbroken. 

The type specimen-the one here figured-has the 

basal portion showing the petiole and the origin of' 

the primary ribs, as well as one side of the blade, very 

perfectly preserved. It is about 9 or 10 centimeters 

long and 8 centimeters wide. At first sight it seenlS 

to be semiribbed, but it is probably best explained as 

being 3-ribbed, with two secondary branches on the 

lateral ribs arising together and so low down as to 

simulate ribs. The ribs are very strong, the lateral 

ones being perhaps slightly stronger than the midrib. 

They are somewhat irregular. 

PopUlus lacoe{l)'JUZ is closely related to a number of, 

described forms-so closely, in fact, that it is somewhat difficult to draw specific lines. .It is undoubtedly of the same type as a larg~group of species fron1 

the Fort Union formation, some of which are also 

reported from the Denver formation. ,It is especially 

like Populus n.,eb'ra8censis Newberry 63 as regards the 

primary nervation but differs in being much larger 

and much broader and above all in having only a few 

large obtuse teeth instead of numerous small relatively 

sharp teeth. Popul'U/J neo'rasaensis is identified in the 

~3 Newberry, J. S., The later extinct floras of North America: U. S. 

Geol. Survey Mon. 35, p. 47, pl. 27, figs. 4, 5, 1898. 
SYSTEMATIC PALEOBOTANY 61 

Denver Hora : 'Y Lesquereux:~)4 who reports a. large. 

number of specimens in the collection studied for the 

Museum of Comparative Zoology. Lesquereux has 

divided these into a number of vnrieties, among them 

one (g'f'(lIfulriclentata) which may include the form uncler discussion but which is a lnrger leaf with fewer 

und larger teeth than are to be observed in a number 

of specimens in the Lacoe collection identified as this 

Varicl), by Lesquereux. 

Popu7Jus ZacoeO/nCt is also like certain of the larger 

leaves of POP'l.tVtts· (fjJnblyrh;yncM 1Vard,55 a wellknown Fort Union species, but differs in being much 

larger and in having fewer, larger teeth and the 

primary ribs at a n10re acute angle. 

Occurrence: Denver formation, south face of South 

Table Mountain, Golden, Colo., about 100 feet below 

the lava cap, collected by Arthur Lakes, 1890. 

Populus knowltoni Berry, n. Sp.60 

Plate 23, Figures 1-4 

Leaves apparently of thin but firm texture, broadly 

or some narrowly obovate, broadest about the middle, 

whence they narrow below to a wedge-shaped base and 

above to a narrow tip; petiole relatively very strong, 

lq>parently several centimeters long; margin entire for 

about one-fourth the length of the blade above 

the base, thence strongly and somewhat irregularly 

toothed, the teeth obtuse and separated by shallow 

sinuses; nervation palmately 3-ribbed, or in effect 5-

ribbed, from the base of the blade; midrib slightly the 

stronger, straight, with 'usually about three pairs of 

:::tl'ongly alternate secondaries above the middle, the 

lower ones often forked, all craspedodrome; lateral ribs· 

straight, at an angle of about 70°, ending in marginal 

teeth; each with five or six secondary branches. on the 

outside, the lowest arising very near the base of the 

blade and thus producing the 5-ribbed appearance; 

secondaries all craspedodrome; nervilles numerous" 

sf Tong, both broken and percurrent., finer nervation 

very perfectly preserved, irregularly quadrangular. 

There are numerous specimens of this well-marlmd 

species in the collection, and although comparatively 

few were perfect when entombed, they retain .the nervation in the parts present with the greatest degree of 

fidelity. There is considerable range in size and outline, though all have the wedge-shaped base, strongly 

toothed margin, and five ribs from the top of the 

petiole. The larger ones are some 8 centimeters long 

and 5 centimeters or more wide, and the smallest one 

M Lesquereux, Leo, Fossil plants collected at Golden, Colo.: Harvard 

ColI. Mus. Compo Zoology BulI., vol. 16, p. 47, 1888. ' 

t.II Ward, L. 11' .• Flora of the Laramie group: U. S. GeoI. Survey Sixth 

~nD. Rept., p. 550, pI. 35, fig. 6, 1886. 

GO Knowlton inncd to suggest a specific name for this species, and I 

hnve thel'efore nflllled it nftf'I' !lim,-E, W, IJ, 

85492-30--5 

observed (pI. 2·3, fig. 4) is about 5 centimeters long and 

scarcely 2 centimeters wide. The majority, however, 

are perhaps 6 or 7 centimeters long and about 4 centimeters wide. The petiole, which is relatively very 

strong, has not been observed actually attached to the 

blade for a greater length than 1.5 centimeters, but in 

one specimen a petiole which seems to have been broken 

off this particular blade is not less than 5 or 6 centimeters "long. 

The species under consideration has a greater or less 

resemblance to a number of described forms, such as 

PlataJnus rhomb'oide(J) Lesquereux,57 a well-known Denver plant. This differs~ however, in its much longer 

size, broader: less wedge-shaped base, less acute-angled 

primary ribs, and above all relatively larger but sharppointed teeth. They can hardly be even congeneric, 

and as a matter of fact the present generic reference 

for the so-called Platamus rho1rV.boidea is perhaps open 

to some question. 

Occurrence: DawsRn arkose, south of oil well near 

Ramah, Colo., collected by C. 'V. Cooke in 1910. 

Populus denverensis Knowlton, n. sp. 

Plate 23, Figure 5 

Leaf small, 6vidently coriaoeous, elliptical-ovate, 

rounded or almost truncate at the base, apparently 

rather long pointed at the apex; Illargin obscurely 

undulate-toothed above the lower fourth of the blade; 

11lidrib relatively very strong, straight; secondaries 

about five pairs, the two lower pairs arising at practically the same point at the base of the blade and 

together with the midrib producing an obscurely 5-

ribbed appearance; lowest pair of· secondaries slender, 

perhaps best considered as in part tertiary br,anches 

of the second pair, which are far stronger and run 

upward for nearly half the length of the blade, eaGh 

with several branches on the outside that enter the 

obscure teeth; upper secondaries alternate, sometimes 

branched, craspedodrome; nervilles numerous, strong~ 

very much broken. 

The little leaf here figured is slightly ellipticalovate, with a rounded base and an apparently acuminate apex. Its length was about 5 centimeters and it~ 

width 2.5 centimeters. The margin is obscurely 

toothed. 

It is with some hesitation that this leaf is described 

as new to scienr.e. It . is, for instance, very similar 

to what Lesquereux has identified as Populus 1nutabilis ovalis Heer,68 from the Livingston formation of 

Montana, but differs in being more nearly oval and 

in having fewer secondaries. It is even nearer to 

57 Lcsquereux, Leo, The Tertiary flora, p, 186, pI. 26, 11gB. 6, 7, 1878. 

53 Idem, p, 177, pI. 24, fig. '1, 
62 FLORA OF TRE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Populus 1'nutabilis cre1'llUlata Heer,59 from the Swiss 

Miocene, and might almost be identified with th~t 

species, but I recall what Lesquereux once said on thIs 

point: "These varieties [of PopUl'UJS 'fl11I1tabilisJ are 

so numerous and so intimately allied by their characters that it is extremely hazardous to identify single 

leaves with one of the eight subdivisions of this 

species." For these reasons I have ventured to give 

this leaf a new name, although it is acknowledged 

to be close to the. forms above mentioned. 

Occurrence: Denver fornlation, south face of South 

Table Mountain, 100 feet below the lava cap, Golden, 

Colo., collected by Arthur Lakes, 1890. 

Populus richardsoni Heer? 

Plate 23, Figures 6, 8 

POpultt8 1'ichardsoni Reer, Flora f6iSSilis arctica, vol. 1, p. 98, 

pI. 4, figs. 1-5; pI. 6, figs. 7, 8; pI. 15, fig .. Ie, 1868. 

Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. 

Rept. for 1873, p. 411, 1874; The Tertiary fiora, p. 177, 

pI. 22, figs. 10-12, 1878. 

In the material from Middle Park I find' three or 

four leaves of a Populus that seem to belong to Heer's 

Popul'Lt8 richa'IYisoni,60 especially as figured from the 

A tane beds of North Greenland. The best of these 

l~aves is 'shown in Figure 6 and is precisely similar in 

shape and nervation to the figure above referred to, 

thouO'h it is much smaller. It is not, however, smaller 

than b·many leaves. that have been referred to this 

5pecies, notably by Heer himself. The leaf under consideration is seen to be strongly 7-ribbed from the top 

of an extremely thick and evidently flattened petiole. 

'The blade is deeply heart-shaped at the base, and the 

lower pair of ribs is deflected much below a right angle 

to supply these lower. lobes. Although this leaf and 

the others found with it appear to belong to Populus 

'1;iclw1'dsoni, it has been thought best to question the' 

identification, as even the best-preserved leaf lacks the 

upper portion and considerable of the margin. 

Occurrence: Middle Park formation, Mount Bross, 

opposite Hot Sulphur Springs, Middle Park, Grand 

Cqunty, Colo., collected by George L. Cannon, 1889. 

Order URTICALES 

o 

Family ULMACEAE 

Ulmus antecedens Lesqllereux 

Plate 23, Figure 7 

Ulmtt8 antecedens Lesquereux, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, p. 49, 1888. 

This species is described by Lesquereux as follows: 

Leaves small, t;tlickish, oblong-Ianeeolate, acute, subcordate 

and sub equilateral at base, doubly or triply dentate;' teeth 

59 Heer, Oswald, Flora tertiaria Helvetiae, vol. 2, pl. 61, fig. 15. 

1856. . 

60 Heer, Oswald, Flora fossilis arctica, vol. .2, pl. 55, fig. 3b, 1871.. 

short, curved upward; secondaries thick, parallel, strong and 

straight, generally simple, sometimes forking in the middle, 

with thick oblique nervilles. 

The leaf has the same character· as that of Ul;nvu8 cra8sifolia, of Texas. The substance is thick, the size is the same,' 

4 centimeters long, 2 centimeters broad in the middle, the 

widest part; the lateral nerves are 12 or 13 pairs. 

This is a splendid species and has been well described by Lesquereux. As he says, it resembles very 

closely UlmJ/1)s cras8ifolia Nuttall, the so-called cedar 

elm, which ranges frOln Mississippi through southern 

Arkansas into western Texas. 

Occurrence: Denver formation,South Table Mountain, Golden, Colo., collected by Arthur Lakes. A 

single specimen known; type in the Museum of Comparative Zoology, Cambridge, Mass., No. 1692. 

Ulmus? quercifolia Unger? 

Plate 23, Figure 10 

Ulmus quercifolia. Unger, Chloris Protogaea, p. lxxx (nomen 

nudum), 1845; p. 96, pI. 25, fig. 5, 1847. 

Lesquereux, Harvard CoIl. Mus. Compo Zoology Bull., vol. 

16, p. 49, 1888. 

The presence of this species in the Denver flora, 

as determined by Lesquereux, rests on the single 

fragmentary specimen here figured. It is the basal 

part of a leaf that must have been some 8 or 9 

centimeters in length and a little over 5 centimeters 

wide. It is very obtusely wedge-shaped and entire 

below but becomes sharply serrate above. Lesquereux 

says: "The borders are sharply dentate, the lateral 

nerves distant oblique, parallel from the base, passing 

in a curve toward the borders, where they become 

effaced." 

Lesquereux compares this fragment with leaves figured by Unger 61 from the Miocene of Parschlug, 

Styria, and it does closely resemble them, though they 

are considerably smaller. One is nearly entire at the 

base, but the other is sharply serrate nearly to the 

petiole. The base is practically equilateral and the 

margin sharply serrate throughout. 

It is extremely doubtful if the generic reference for 

these leaves is correct; in fact, Unger himself casts 

some doubt on it. They do not have the aspect of elm 

leaves and seem 'much more likely to belong to Quercus. As the leaf from Golden is so broken I have 

hesitated to refer it to another genus and have simply 

questioned both genus and species pending the discovery of more complete material. 

Occurrence: Denver formation, South Table Mountain, Go14en, Colo., collected by Arthur Lakes. ~pe~~.­

men now in the Museum of Comparative Zoology ~ 

Cambridge, Mass., No. 1422. 

61 Unger, Franz, IConogl'aphia plantarum fossilium, p. 115, pl. 43. 

figs. 24, 25, 1852. 
SYSTEMATIC PALEOBOTA.NY 63 

Family MORACEAE 

Ficus sUbtl'uncata Lesquel'eux 

Ficus 8ubt'funoata Lesquereux, The Tertiary flora, p. 205, p1. 30, 

figs. 7-9, 1878. 

INclIs aUl'ioltlata Lesquereux, U. S. GeoI. and Geog. Survey 

Terr.· Ann. Rept. for 1872, p. 379, 18"73; The Tertiary 

flora, p. 206, pI. 30, figs. 4-6, 1878. [Homonym, Lour, 

1834.] 

Ficus tnmcatc('Y Heel'. Lesquel'eux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1873, p. 400, 1874. 

Lea ves variable in size (from 4 to 10 centimeters 

long, 2.5 to 5.5 centimeters wide), entire, ovate-Ianceolute or oblong-obovate, obtuse. or sometimes almost 

acutely pointed at the apex, truncate-cordate or deeply 

cordate, almost aUl'iculate, at the base; basilar veins 

three or four pairs, the upper more remote, all campto-. 

drome. 

This species is most widely known under the name 

Ficu8 o/u/I'iculata Lesquereux, but this name proves to 

be preoccupied by a living species and it becomes necessary to restore the name next in order, Ficus 8ubtrunoata Lesquereux. 

The type specimens on which Fi()U8 au1'iculata Lesquereux was based came from Golden, Colo., and are 

preserved in the United States National Museum (N os~ 

266, 266a, 267). The matrix shows them to have come 

from the andesitic material so characteristic of the 

Denver formation. Lesquereux 62 also reports this 

species from the region north of Grand River, Colo., 

the exact locality being described by Peale 63 as 

follows: 

On Gi'and River, near the mouth of Roan Oreek, there aFe 

bluffs in which about 500 feet of sandstones outcrop. Tl?-ese 

sandstones are probably the upper part of the Laramie group, 

the variegated beds just above representing the Wasatch 

group. In the Grand Hogback range, just east of Oactus Valley, the following fossil plants were obtained, the identifica· 

tiol1S being by Professor Lesquereux. 

Then follows the enumeration of species, among 

them being Ficu"8 auriC'l.IJlata. 

From Peale's description of the stratigraphic relations it appears that the plants mentioned must have 

come :from beds in the approximate position of the 

Denver formation, but the species has not since been 

found in that portion of Colorado, and the exact stratigraphic position must remain in question. 

In the report in which Fi()U8 aWl'icula)ta was named 

Lesquereux recorded it from Spring Canyon, near 

Bozeman, Mont. The specimens so recorded are not 

now to be found in the National Museum collections, 

but the species is present in collections made in the 

same place by A. C. Peale and me during the years 

:1885-1890. They came from the upper part of the 

o:J Lesqucl'eux, Leo, The Tertiary flora, p. 207, 1878. 

08 Peale, A. C., U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 

1876, p. 181, 1878. 

series of rocks comprising the so-called Bozeman coal 

field, in what has been called the Livingston formation, which in my opinion corresponds in position and 

age to the Deriver formation of Colorado. 

The recent collections from Golden, Colo., contain a 

few leaves clearly belonging to this species, but as they 

have only the bases preserved they add nothing to our 

know ledge of the species. 

The specimens originally mentioned by Lesquereux 

under the name Fi<1U8 trwncata)?, later changed to 

Ficu8 8ubtruncata, are fortunately all preserved in the 

United States National M~seum (Nos. 268, 269, 270). 

They came from exactly the same place as those described as· Ficus auriculaJta, and they do not seem to 

. differ from that species sufficiently to warrant their 

separation. Lesquereux described this species as follows: "Leaves comparatively small, entire, truncatecordate at the more or less inequilateral base, ovate or 

oblong, acuminate; bord,ers undulate." Size, it is well 

known, is a very unreliable character. A glance at the 

figures of both species shows that the smallest leaf of 

F. 8ubtT'1Jlnc(J)ta is larger than the smallest leaf of F. 

aU1'ioulata,64 therefore relative size can not be used to 

separate them. The shape is approximately the same, 

ovate or ovate-Ianceolate. F. 8ubtr'1JJncata is said to be 

" truncate-cordate" at the base, as shown in Lesquereux's Plate 30, Figure. 7, but an examination of the 

type specimen of this form (U. S. Nat. Mus. No. 270) 

revealed the fact that the matrix had not been all removed from the base. When this had been done the 

base proved to be of the same shape as that shown in 

Figure 4 of the same plate-that is, distinctly cordate. 

It is also described as "more or less inequilateral," 

presumably as shown in Figure 9 of Plate 30. This 

specimen (U. S. Nat. Mus. No. 268) had not been 

properly excavated from the matrix, and when this 

had been done it showed that the two sides of the leaf 

were approximately equal, thus destroying this supposed character. It was also said to be "undulate" 

on the margins, but on examining the type of Figure 

7, which is shown in the plate as undulate, it is found 

that neither of the margins is preserved, the drawing 

·being wrong. Figure 9 is also· wrong as to the leaf 

being undulate. The leaf does not lie flat on the 

matrix, which causes it to appear undulate. The 

nervation is precisely the same in both of these forms, 

as can be seen by comparing the figures on Plate 30. 

It'therefore seems justifiable to regard them as representing a single species .. 

Occurrence: Denver fornlation (types), South Table 

Mountain, Golden, Colo. Formation uncertain, Grand 

[Colorado] River near the mouth of Roan Creek. 

Colo., collected by A. C. Peale, 1876. Livingston 

64 Lesquereux, Leo, The Tertiary flora, pI. 30, 1878. Cf. figs. 4 and 7. 
64 o FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

formation (?), Hodson's coal mine on Meadow Creek, 

14 miles southeast of Bozeman, Mont., collected by 

A. C. Peale in 1872 and by Peale and !{now lton in 1888. 

Ficus? alata Knowlton, n. sp. 

Plate 23, Figure 9 

Leaf small, very thick, ovaloI' elliptical-oblong, 

rather abruptly rounded to the base and thence narrowly decurrent on the thick, strong petiole, apex 

destroyed; midrib very strong, straight; secondaries 

alternate, thin, at a low angle, camptodrOlne, arching 

and joining well inside the nlargin; nervilles obscure, 

apparently percurrent and at right angles to the secondaries; finer nervation not retained. 

This species is apparently represented by the single· 

example figured, which lacks nearly all of the upper 

portion. So far as can be made out it is oblong or 

elliptical-oblong with perfectly entire lnargins and a 

rounded base, which passes irito a narrow wing down 

the thick petiole. The leaf was probably 3.5 or 4 

centimeters in length and slightly more than 2 centimeters in width. The petiole is about 1 centimeter 

long and gives evidence of having been somewhat 

longer when perfect. The wing to. the petiole appears 

to diminish downward, if the conditions are correctly 

interpreted, and at the base had probably entirely disappeared. This leaf as it i~ preserved exhibits the 

. dorsal surface and is seen to be very thick and probably coriaceous, as the secondaries and nervilles show 

but faintly. 

I am much .in doubt as to the proper generic reference for this little leaf. The contour, coriaceous texture, thick petiole, and midrib suggest many species of 

Ficus, but the presence of a decided wing on the petiole 

can hardly be considered a character of this genus, 

ye~ cer~ain fossil fOrIns having this feature have been 

so placed. Thus Ficus ovalis Lesquereux 65 has approximately the same shape of blade and what appears 

. to be a slight winglike broadening of the petiole; it 

differs in size and somewhat in nervation. In shape, 

size, and nervation it also suggests F. ternuinervis Lesquereux,66 and there is even a slight indication of the 

presence of a wing on the short petiole in this species. 

The species in hand has some resemblance to what I 

have called Oinchonidiwm? turne1'i Knowlton,67 from 

the Esmeralda formation of Nevada. The size and 

winged petiole are somewhat the same, but in the 

Nevada form the base is strongly unequal-sided and 

the nervation much closer and more irregular. The 

genus SapindUs is also suggested, but from this it 

tJIj The Tertiary flora, p. 198, pI. 30, fig. 2, 1878. 

00 The Cretaceous and Tertiary floras, p. 164, pI. 44, flg. 4, 1883. 

07 U. S. Geol. Survey Twenty-first Ann. Rept., pt. 2, p. 218, pI. 30, 

figs. 9-11, 1900. 

appears to be excluded by the long and winged petiole. 

I am therefore constrained to place it under Ficus, 

premising, however, that this reference may not be 

the correct one. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Arapahoe formation), 3,000 feet east of 

the Douglas coal mine, Sedalia, Colo. 

Ficus denveriana Cockerell 

Plate 24, Figures 1-3 

Fious aenveriana Cockerell, Torreya, vol. 10, p. 224, 1910. 

Berry, U. S. Geol. Survey Prof. Paper 91, p. 198, 1916. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 302, pI. 

75, figs. 1, 2, 1917 [1918]. 

Ficus spectabilis Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1872, p. 379, 1873; The Tertiary 

flora, p. 199, pI. 33, figs. 4-6; Harvard CoIl. Mus. Compo 

Zoology Bull., vol. 16, p. 50, 1888. [Homonym, Kunth 

and Bouche, Annales sci. nat., 3d ser., vol. 7, p. 235, 

1847.] 

Populus ungeri Lesquereux, The Tertiary fiora, p. 175, pI. 

24, fig. 5, 1878;· U. S. Geol. and Geog. Survey Terr. 

Ann. Rept. for 1876, p. 507, 1878. 

Populus heUaaum Unger. Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1873, pp. 380, 397, 1874;-

idem for 1876, p. 507, 1878. 

G-i88~ts prirnaeva. Saporta. Lesquereux, Harvard ·Coll." Mus. 

Compo Zoology Bull., vol. 16, p. 51, 1888. 

In one of the collections from the Dawson arkose 

of Jimmy Camp Creek I find the l.arge leaf here 

figured, which apparently belongs to this species. It 

is rather broader and more truncate at the base than 

is usual in this species, but otherwise it does not 

appear to differ essentially. . 

The collection from Mosby, Colo., also includes this 

species, though it is represented by only the upper 

portion of a rather ~mall individual. It has not been 

figured. 

A small collection made by Francis M. Collins for 

Dr. A. C. Peale on Sand Creek near Magnolia, a short 

distance northeast of Denver, includes a very good 

specimen of this species. 

In 1878 Lesquereux 68 described, under the name 

Populus ungeri, a single leaf from the andesitic beds 

at Golden, Colo. This specimen (U. S. Nat. Mus. 

No. 222) is before me and needs but little study to 

show that it has been very incorrectly figured and interpreted by Lesquereux. According to the figure it 

i.~ nearly circular and about equally rounded at base 

and apex, but as a matter of fact neither is represented 

by true margin. By removing the matrix it is found 

that the base is obtusely wedge-shaped and the apex 

prolonged for several centimeters beyond the point 

where it is shown in the figure. That it was prolonged far beyond this point is still further shown 

68 Lesquereux, Leo, The Tertiary fiora. p. 175, p). 24, fig. 5, 1878. 
SYSTEMATIO PALEOBOTANY 

by the fact 0 that the midrib is little if any reduced Ficus aguilar Knowlton 

Plate 24, Figure 4 

65 

in thickness at the point where it enters the supposed 

margin. The margin, shown in the figure as slightly 

undulate on one side, is really practically entire, and 

the nervation is likewise incorrectly figured. The 

secondaries on one side show a number of branches, 

and the finer nervation is well preserved in one or 

two places and is seen to consist of numerous strong, 

orten broken nervilles. 

Popu'z'U)8 unfJ6ri Lesquereux is undoubtedly a medium-sized leaf of Fi0W8 denvm'iana Cockerell, being, 

for example, of the same shape and general appearance as one of Lesquereux's figures of this species,<l!l 

though a little longer. 

In the collection of Golden specimens studied by 

Lesquereux for the Museum of Comparative Zoology 

POP1.l}IUS wngeri is said to be represented by three 

examples, but I have not seen them. 

In the material from Golden elaborated by Lesquereux for the Museum of Comparative Zoology 

there is a single fragmentary specimen that he identified as OisBUS prim(JJeva? Saporta, a species which 

cnme from the travertines of Sezanne .. On comparino- this leaf with Saporta's figure 70 it is seen that 

th~y agree in size and shape but diff.er in marginal 

dentition and in the secondaries, which are camptodrome in the Golden leaf but craspedodrome in the 

French leaf. 

Comparison of the leaf from Golden with figures 

of Fic1.ts denvf31rUtma Cockerell shows that it agrees 

closely with the original figure by Lesquereux, and 

it·, is consequently referred to this sDecies. 

Occurr~nce: Denver formation (types) , Golden, 

Colo., specimen in Museum of Comparative Zoology, 

Cambridge, Mass. (identified by Lesquereux as Oissus 

primaeva.fJ Saporta) , No. 1500; Sand Creek near Magnolia northeast of Denver, Colo., collected by Francis 

~1. Collins. 'Vilcox formation, Arkansas and Louisiana. Lagrange formation, Kentucky, in beds of Wilcox 

nge. Uaton formation, Raton Mesa region of Colorado n.nd New Mexico. Laramie formation, Popes 

Bluff. west of Pikeview, Colo. -[identification questioned]. Dawson arkose, east bank of .Jimmy Camp 

Creek. 1 mile above Richfield Springs ranch house, 9 

miles J east of Colorado Springs, Colo., collected by 

F. H. J{nowiton, July, 1910; dump of Mosby coal 

mine. Mosby, Colo., collected by F. H. J{nowlton and 

G. B. Richardson, 1910; Coal Gulch, about 2 miles 

Hortheast of Ramah, Colo., in sandstone above coal, 

collected by VV. T. Lee, 1914; a quarter of a mile east 

of Purdon mine, sec. 27, T. 11 S., R. 61 W., Colo., 

collected by G. B. Richardson. 

ell Lesqucrcux, Leo, The Tertiary flora, pl. 3H, fig. 4, 1878. 

70 Saportn, Gaston de, Prodrome d'une flore fossile des trn vel'tlns 

nnciens de S~znnne, pI. 11, fig. 2, 1868. 

Ficus ag'uilar Knowlton, U. S. Geol. Survey Prof. Paper 101, 

p. 300, pI. 71, fig. 1, 1917 [1918]. 

This species was described from the Raton formation of southern Colorado and is based on a single 

l:xample which lacks the upper portion .. The present 

specimen permits a knowledge of the apical portion 

of the blade, and the description may be modified to 

read as follows: Leaves large, very thick and coriaceous, oblong,. rather abruptly rounded below to the 

thick petiole and above to an obtusely acuminate apex; 

margin perfectly entire; midrib unusually thick, perfectly straight but becoming t[linner above; secondarIes numerous, about 10 or 15 pairs, at somewhat irregular distances, emerging at a low angle, considerably 

curved upward, ending very near the margin or camptododrome; nervi lIes apparently numerous, unbrokep, 

and at right angles to the secondaries. 

The present specimen is much smaller than the type 

but is nearly perfect. It is about 11 centimeters long 

and nearly 7 centimeters wide. It has the petiole preserved for a short distance only. 

In the collection of Denver plants identified by 

Lesquereux for the Museum of Comparative Zoology 

I find a single leaf under the name P OPIUiU8 un geri 

Lesquereux. It represents only the lower . portion of 

a large oval-lanceolate leaf that must have been about 

12 ceI1timeters long and nearly 7 centimeters wide. 

It is rounded truncate at the base and has. a very 

strong midrib. The secondaries are at a slightly lower 

angle than in the specimen of Fi()Us aguila'}' figured, 

but it does not otherwise differ and has been referred 

to that species; it is not figured. 

Occurrence: Dawson arkose, a quarter of a mile east 

of Purdon mine, sec. 27, T. 11 S., R. 16 W., CoI6., collected by G. B. Richardson. Denver formation, South 

Table Mountain, Golden, Colo., collected by Arthur 

Lakes, specimen (under the name Poplulu8 ungeri 

Lesquereux) in the Museum of Compar~tive Zoology, 

Cambridge, Mass., No. 1087. Raton formation (type), 

Aguilar, Colo. 

Ficus berthoudi Lesquereux 

Plate 25, Figures 1, 2, 6 

li'-iC'l/.8 berthoudi Lesquereux, Harvard ColI. Mus. Compo Zoology 

Bull., vol. 16, p. 49, 1888. 

Leaves thick and coarse, broadly cordate at base, oyatelanceolate, acuminate above, entire, enlarging toward the base 

and rounding to the petiole, descending lower than its top, 

sometimes auriculate, the basilar border in one leaf overlapping 

the top of the petiole; primary nerves deeD and broad; lower 

lateral nerves opposite, the upper alternate, all very deeply 

curving toward the borders and following them in a series of 

'areoles; nervilles deep, close, parallel, cut by branches fit right'. 

angles, forming a square distinct areolation. 
66 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORAD() 

Two of the four specimens referred to this species 

aTe here figured as the types. The more perfect one, 

Figure 2, is a small leaf about 7 centimeters long and 

nearly 5 centimeters broad. The other, Figure 1, is 

much larger, probably about 11 centimeters long and at 

least 8 centimeters qroad. They are, as Lesquereux 

says, rather. thick coarse leaves with strongly impressed nervation. 

, I have also referred to this species a leaf identified 

by Lesquereux 71 as Ficus tiliaefolia (AI. Braun) 

Heel'; it is shown in Figure 6. It is intermediate in 

size between the other specimens but does not seem to 

differ essentially from them. 

FiC'ttS be'l'thoudi is undoubtedly most closely related 

to Fi(JUs subt'l'uncata Lesquereux 72 (which includes 

Fi(JUs auriC11ilata Lesquereux); in fact, it is perhaps 

doubtful if they should be maintained as distinct. It 

is possibly a slightly coarser leaved species, with somewhat larger leaves which are more abruptly narrowed 

above, but these differences are certainly slight. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, 

Mass., Nos. 1089b, 1440, 1443. 

Ficus pseudopopulus Lesquereux 

Plate 25, Figures 3-5; Plate 26, Figures 2-4 

Ficu8 pseudopo'puws Lesquereux, U. S. Geol. oand Geog. Survey Terr. Bull., vol. 1, 2d ser., No.5, p. 387, 1876; U. S. 

Geol. and Geog. Survey Terr. Ann. Rept. for 1874; p. 313, 

1876; The Tertiary fiora, p. 204, pI. 34, figs. la, 2, 1878. 

Berry, U. S. Qeo!. Survey Prof. Paper 91, p. 200, pI. 37, 

figs. 3-5; pI. 113, fig. 3, 1916. 

Knowlton, U. S. Qeol. Survey Prof. Paper 101, p. 304, 

pI. 72, figs. 2-4 j pI. 73, figs. 1, 2; pI. 112, fig. 3, 1917 

[1918J. 

Ficus amLraei Lesquereux, Harvard ColI. Mus. Compo Zoology 

Bull., vol. 16, p. 50, 1888. . 

Ficus paeudopoprul1JJS was described by Lesquereux 

from an unknown locality which was conjectured to 

be in the Green River formation, but the point remains unsettled. The species was subsequently found 

in beds of lower Tertiary age at Evanstqn, Wyo., and 

still later in the Raton formation of the Raton Mesa 

region of Colorado and New Mexico, in the Wilcox 

formation of Louisiana, and in the Lagrange formation of Tennessee, in beds of Wilcox age. It is perhaps most abundant in the Wilcox b~ds. Berry gives 

the following description: 

Palmately veined leaves of medium size, broadly ovate in 

general outline, narrowed and acuminate at the apex, broadly 

rounded or truncate, and more or less decurrent at the base. 

Length about 12 or 13 centimeters. Maximum width, at or 

below the middle, about 6 or 7 centimeters. Texture sub-

• 71 Lesquereux, Leo, Harvard ColI.' Mus. Compo Zoology Bull., VOl. 16, 

p. 49, 1888. 

72 Lesquereux, Leo, The Tertiary'fiora, pI. 30, figs. 4-6, 7-9 [not fig. 

6a, wbIch = Oarpites oviform-is J. . 

coriaceous. Petiole stout, curved, about 1.5 cf'utimeters in 

length. Primaries [ribs] three, curved, the midrib stouter 

than the laterals. Lateral primaries [ribs], one on each side, 

diverging from the midrib just above the top of the petiole 

at angles of about 20°, ascending, and camptodromely joining 

a secondary two-thirds or more of the distance from the base 

to the tip. Secondaries from the upper half ~f the midrib, 

four or five alternate curved pairs, diverghlg from the midrib 

at acute angles, becoming subparallel with the lateral margins 

distad, and arching along them in a camptodrome manner. 

Secondaries from the outer side of the lateral primaries [ribs], 

about seven on each side, thin and camptodrome, the lowest 

on each side longer, stouter, and more ascending than the 

others, diverging at or just above the top of the petiole~ Tertiaries [nervilles] thin, percurrent, at right angles to primaries 

and secondaries. 

The example figured in this paper, although fragmentary,: is undoubtedly identical with leaves of this 

species as figured by Berry fr'om Shandy, Tenn. A 

more perfect specimen from Mosby, Colo., has also 

been figured. It has the petiole preserved for a length· 

of over 2 centimeters and altogether is indistinguishable from leaves in the Raton formation of the Raton 

Mesa region. 73 

The type specimen of Fi(J'Us and'l'aei Lesquereux,74 

the property of the Harvard College Museum of Comparative Zoology, is before me and is shown on Plate 

26, Figure 4. 'It needs but a glance to show that it 

is identical with Fious pseudopopultus Lesquereux. It 

is, for instance, indistinguishable from figures given 

by Berry 16 from the Lagrange formation of Tennessee, 

in beds of Wilcox age, and by I(nowlton 76 from 

the Raton formation of Colorado and New ~Iexico. 

It has the petiole preserved for a length of 2 centimeters. 

Occurrence: Dawson arkose, east bank of Jinllny 

Camp Creek 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., colle.cted by F. H. Knowlton and W. T. Lee, July, 1910; 

same locality but 1 mile above ranch house, collected 

by F. H. Knowlton, July, 1910; dump of Mosby coal 

mine, Mosby, Colo., collected by F. H. Knowlton and 

G. B. Richardson, 1910; Rice's clay bank, 1 mile southwest of Calhan, Colo., collected by F: H. I(nowIton, 

1910. Raton formation, Raton Mesa region of Colorado and N ew ~1exico. Wilcox formation, Louisiana. 

Lagrange formation, Tennessee. Denver formation, 

Sand Creek near Magnolia, northeast of Denver, 

Colo., collected by Francis 1\1. Collins, 1909; South 

l'able Mountain, Golden Colo., collected by Arthur . 

Lakes. Type of FiC1..t8 andiraei Lesquereux, Museum 

of Comparative Zoology, Cambridge, Mass., No. 1677. 

78 See Knowlton, F .. H., U. S. Geol. Survey Prof. Paper 101, p. 304, 

pI. 73, figs. 1, 2, 1917 [1918]. 

14 Lesquereux, Leo, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, 

p. 50, 1888 . 

75 Berry, E .. "V., U. S. Geol. Survey Prof. Paper 91, p. 200, pI. 37, 

figs. 4, 5, 1916. 

70 Knowlton, F. H., U. S. Geol. Survey Prof. Paper 101, p, 304, pI. 72, 

fig. 4; pI. 73, figs. 1, 2, 1917 [1918]. 
SYSTEMATIC PALEOBOTANY 67 

Ficus dawsonensis Knowlton, n. sp. 

PIa te 26, ]j'igul'e 1 

Leaf thick, broadly ovate, rounded and deeply 

heart-shaped at the base, rounded and obtuse· at the 

al:ex; margin absolutely entire; midrib rather slender, 

wIt.h about foul' pairs of thin secondaries, the lowest 

pail' opposite, arising at the top of the petiole and 

giving the leaf a 3-ribbed aspect, each with five or six 

camptodrome branches on th~ outside, upper pairs of 

~econc1aries alternate, subca.mptodrome; nervilles obscure, thin, apparently unbroken. 

This form is represented by a single example with 

its counterpart. It is about 9 centimeters long and 

7 centimeters wide. 

This species is Df the type Df Ficus lee'i KnowltDn,77 

frDm the VermejD fDrmatiDn o.f the Raton Mesa regiDn; in fact, it is distinguished frDm that species with 

difficulty. It appears to differ in being rather brDader 

in prDpDrtiDn to. the length and abDve· all in the absence Df the nervilles, which rire so prDminent in F. 

leei. It is to. be regretted that there are nDt more 

specimens Df F. dawsonen,sis available fDr cDmparisDn, 

fDr then it might be determined whether this is a valid 

species. 

Occurrence: Da WSDn arkQse, l\1Qsby cQal mine, 

lVIQsby, CQIQ., cDllected by F. H. Knowlton and G. B. 

RichardsQn, 1910. 

Ficus sp. 

Plate 27, Figures 1-3 

The material Qbtained near Ramah, Colo., includes 

a number Qf fragmentary l'?aves that appear to belong to. Ficus, but they are hardly well enough preserved to. adlnit Qf specific identificatiQn. They are 

large leaves, prQbably ·not less than 12 or 14 centimeters IQng and 8 centimeters 01' mQre wide. As 

nearly as can be made Qut these leaves were ellipticallanceQlnte 0.1' perhaps slightly Qvate-lanceQlate, with a 

very Qbtusely we~ge-shapec1 Dr rQunded base and apparentlya rather acumina.te apex. The margin is perfectly entire. The nervation consists Qf a very thick 

midrib and numerous pairs of strong secondaries, 

which emerge at an angle Qf about 35° or 40°. 

This fDrm appears to. be mQst clQsely related to. what 

is nQW knQwn as FiC't1J8 wncata Lesquereux and is perhaps identical with it. It is especially suggestive Qf 

certain leaves nQW referred to. FiC'ttS uncata but described and figul:ed by Lesquereux under the name 

Fi(J1J)8 a'J'enacea b1'evipetiolata.78 AbQut the Qnly difference to. be nQted is in the base, which is rather mQre 

wedge-shaped than in Fic'us 'l,(lfLcata!. It would prQb77 Knowlton, li". H., U. S. Geol. Survcy Prof. Paper 101, p. 261, pl. 39'; 

Hgs. I-G; pI. 40, figs. I, 2, 19t7 ·[1918]. 

i8 J.esqnel'cux, Leo, Thc Tertifll:Y flOl'fl, p. 195, pI. 29, figs. 2, 3, 5,' ! 

1878. 

ably be well within the bounds of prQbability to. identify the present form directly with FiC1.tS wncata, but 

in view Qf its fragmentary representatiQn it has seemed 

best to. refrain from a specific d~termination. 

Occurrence: Dawson arkQse, near Qil well 1 mile 

south Qf Ramah, CQIQ., cQllected by C. VV. CODke fQr 

G. B. Richardson, 1910. 

l"icus sp. 

Pious til'iaefoU,(~ Al. Braun. Lesquereux, The Tertiary flora, 

p. 203, pI. 32, fig. 2, 1878. 

In .the repQrt on the Tertiary flora Lesquereux gave 

the figures Qf three fragmentary leaves that he referred 

to the European Fious tiliaefoliaJ AI. Braun. He said 

(p. 204) that he had identified this species" frOln tlle 

lowest stage Qf the lignitic EQcene to' the hiO'hest Ter- • b 

bary measures," the localities enumerated being vVashakie Station, WyO..; Spring CanYQn near FQrt· Ellis, 

l\1Qnt. ; Yellowstone Lake, PQint Qf RQcks, vVYQ.; 

Fishers Peak,· Sand Creek, GQlden, and near CQIQradQ 

Springs, CQIQ.; Black Buttes, WyO..; and the auriferous gravel Qf Cali fQrnia. Lesquereux states that the 

specimens are always mQre 0.1' less brQken and the 

identificatiDn uncertain. 'Vherever the flQra Qf the 

above-mentioned localities has been critically reviewed 

~n recent years, Fim,1J8 tiliaefoli{J). has usually been reJected, and the leaves so. determined have been refened 

to Qther species. . 

One of the specimens figured in the report Qn the 

Tertiary flQra (pI. 32, fig. 2) is frQm GQlden CDIQ. 

It is No. 282 Qf the United States NatiQnal Museum 

collection and is preserved in the andesitic matrix 

.characteristic of the Denver fQrmatiQn. It undeniably resembles certain leaves referred to. the EurQpean 

FiC'ttS tiliaefolia AI. Braun, such as SQme frQm the 

Swiss Miocene as figured by Heer,79 but the Old 'YQrlel 

specimens are all figured as strQngly unequal-sided, 

and the specimen under consideratiQn is so. fragmentary that it is impossib~e to. determine this point. It 

lacks bQth base and apex and all Qf Qne side, and althQugh it may have been unequal-sided there is no. 

positive indicatiQn Qf it. 

This particular specimen is also. Qf abQut the same 

size and shape as the typical leaves of Ficus plmnicostata goldiana Lesquereux,80 except that it is slightly 

mQre wedge-shaped at the base and has a sQmewhat 

. strQnger nervatiQn. Under the circumstances it seems 

best to refer this leaf to. Ficus sp., as it can not be 

positively referred to. F. tiliaefolico Qr to. any Qf the 

varieties of F. pla/nicostata. 

The N atiQnall\1useum contains a specimen (No.. 284) 

that appears to. be the cQunterpart of the leaf figured 

7U Reer, Qswald, Flora tel'tiaria Relvetiae, vol. 2, p. 68, pI. 84, figs. 

1-6, 1856. 

so Lesquereux. I,eo, The Tertiary flora, p. 202, pI. :33, figs. 1-3. 1878. 
68 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

in the report on the Tertiary flora, Plate 32, Figure 1, 

as Ficus tiliaefolia. It is recorded as having come 

from Sand Creek, Colo., but to judge from the matrix 

this can hardly be true. In any event it is a fragment 

lacking both base ~nd apex and has no positive evidence of having been unequal-sided. It is not accepted as Ficus tiliaefolia. 

Occurrence: Denver formation, Golden, Colo., probably collected by Arthur Lakes in the early sev~nties. 

Ficus occidentalis (Lesquereux) Lesquereux 

Plate 26, Figure 5; Plate 27, Figure 4 

Ficus occidental'is (Lesquereux) Lesquereux, The Tertiary 

flora, p. 200, pI. 32, fig. 4, 1878; Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 50, 1888. 

Penhallow, Report on Tertiary plants of British Columbia, 

p. 55, 1908. 

Knowlton, U. S. Geol. SUl'vey Prof. Paper 101, p. 302, pI. 

72, fig. 1, 1917 [1918]. 

Berry, U. S. Geol. Survey Prof. Paper 91, pp. 12, 197, pI. 

28, fig. 3, 1916. 

Dombe.yopsis occidentalis Lesquereux, U. S. Geol. and Geog. 

Survey 'J'err. Ann. Rept. for 1872, p. 380, 1873. 

The type of this species is from the Denver formation at Golden, Colo., and is preserved in the United 

States National Museum (No. 281). It was described. 

by Lesquereux in part as follows: 

Leaves comparatively thick, coriaceous, truncate-cordate at 

the base, narrowed upward into an obtuse acumen, palmately 

triple-nerved; lateral veins equidistant, parallel, camptodrome. 

Ficus occidentali8 app.ears to be rather rare, though 

in his original discussion Lesquereux said, "A number 

of finely preserved specimens of this· species have 

been obtained from the same locality, all, however, 

deprived of the petiole." He also recorded 81 the presence of five specimens in the large collections he identified for the Harvard College Museum of Comparatj ve Zoology. The large collections made by Lakes 

and now for the most part studied for the first time 

contain only a single example that differs from the 

type in being much smaller. 

In the collections from the Raton formation made 

in Dillon Canyon, N. Mex., I noted the presence of 

two leaves of this species.82 The better preserved of 

these was figured and, except for being a little larger 

than the figured type, agrees perfectly with it. 

Ficws occidentaJlis was recorded by Berry 83 from 

the l\1idway(~) formation at Earle, Tex., and the 

Ackerman formation of the Wilcox group at Hurleys, 

Benton County, Miss. The leaves from Mississippi 

51 Lcsquereux, Leo, Harvard CoIl. Mus. Compo Zoology Bull., vol. Hi, 

p. 50, 1888. 

82 Knowlton, F'. H., U. S. Geol. Survey Prof. Paper 101, p. 302, pI. 72, 

fig. 1, 1917 [1918]. 

83 Berry, E. 'V., The lower Eocene flol'fls of southeastern North 

America: U. S. Geol. Survey Prof. Paper 91, pp. 12, 197, pI. 28, fig. 3, 

]016. 

were described as being somewhat smaller than those 

from the Rocky Mountain region, averaging about 8 

c(~ntimeters in length and 6 centimeters in' maximum 

width. 

This species is thought to be most closely related to 

Ficus p'lanicostata Lesquereux, from which it differs 

ill its generally larger size, broadly ovate rather than 

elliptical outline, and thicker nerves. It is also apparently related to Fi(JlJ)S lUJlrrisiana Hollick,s4 a species from the Wilcox formation of Louisiana, which 

is widest in the middle instea d of at the base. The 

venation, Berry states, "though of the same general 

character shows well-marked differences of detail." 

Occurrence: Denver formation (type) , Golden: 

Colo. Raton formation, Dillon' Canyon, near Blossburg, N. Mex. Wilcox group (Ackerman formation), 

Hurleys, Benton County, Miss. Midway ( ~) formation, Earle, Tex .. Eocene, Stanley Park, Vancouver 

Island, British Columbia. 

8 Ficus ramahensis Knowlton, n. sp. 

Plate 27, Figure 5 

Leaf large (about 23 centimeters long and 13 centimeters wide), evidently thick,' broadly ovate, broadest 

near or just below the middle, from which it is 

rounded to the very obtusely wedge-shaped base and 

upward to a relatively long, slender apex; margin 

entire; midrib very strong, especially below, straight; 

secondaries about 15 pairs, mainly 'alternate, strong, 

parallel, at an angle of about 40°, not much curved 

upward, camptodrome, arching very close to the margin; finer nervation not retained. 

This species as repl'esented by the example figured 

is apparently the largest-leaved Ficus found thus far 

in the Denver or related formations. It seems to be 

most closely related to Ficus occidentalis Lesquereux,85 a fairly abundant forn1 in the andesitic 

beds at Golden, but it is much larger and does not 

have the lower secondaries branched, nor are the 

upper secondaries so much curved upward. The 

largest of the type specimens of Flcus occidentalis is 

only ,about 15 centimeters long and 8.5 centilneters 

wide, and the other types. are 'much smaller. Fious 

occidentalis has been identified by Berry in the Wilcox group, but these leaves are smaller than the Rocky 

Mountain specimens. 

Ficus ramahensis is also like Ficus monodon (Lesquereux) Berry,86 especially as figured by Berry from 

the Wilcox group. That species, however, is relatively broader, is more truncate at the base, and has 

84 Hollick, Arthur, Louisiana GeoI. SUl'vey Rept. for 1899, Special 

Rept. 5, p. 281, pI. 46, fig. 2, 1899. 

~5 Lesquel'eux, Leo, The 'l'ertiul'Y flora, p. 200, pI. 32, fig. 4, 1878. 

80 Berry, E. W., U. S. GeoI. Sm vey Prof. Paper 91, p. 201, pI. 32, 

fig. 2; pI. 33, fig. 2, ] ~16. 
SYSTEMATIC PALEOBOTANY 69 

conspicuously undulate margins. It is also like certain of the leaves that have be~n referred to Ficus 

u,ncata Lesquereux, from the Raton formation of 

Colorado and New j)iexico, especially the large leaf 

figured by Lesquereux 87 under the name POPUl1tS 

monodon. 

Occurrence: Dawson arkose, half a mile west of 

Ramah, Colo., collected by 'V. T. Lee, 1915. 

Ficus? lakesii I{nowlton, n. sp. 

Plate 28, lj'igure 1 

Leaf small, nalT01V ly obova te, rounded or almost 

truncate at the apex, obtusely wedge-shaped at the 

base; petiole relatively rather long, slender; margin 

entire or· slightly undulate; nervation strongly 3-

ri bbed from the base of the blade; midrib exceedingly 

strong for the size of the leaf, .nearly straight, with 

one pair of alternate, very strong secondary branches 

in the upper portion; lateral ribs nearly as strong as 

the midrib, passing up to near the apex; there curving inward, each with five or six secondary branches 

on the outside, these camptodrome and arching just 

at the margin; nervil1es numerous, strong, mainly 

broken. . 

The little leaf here figured is rather narrowly obovate, being slightly under 2.5 centimeters in length 

exclusive of the petiole, which is itself nearly 1 centimeter in length. The width at the broadest point, 

somewhat above the middle, is a little less than 1.5 

centimeters. . 

This leaf is referred to the genus Ficus with hesitabon and mainly on the ground of its resemblance to 

certain of the smaller leaves of F. plamicostata Lesqllereux.88 It differs, however, in being obovate instead of ovate or elliptical, with only one pair of seccndaries and somewhat different nervilles, etc. . 

Occurrence: Denver formation, Golden, Colo., south 

fnce of South Table niountain, collected by Arthur 

Lukes, for whom it is nameel. 

Ficus coloradensis Cockerell 

F·iOltS colol'(uZc'nsis Cockerell, Ton'eya, yol. 10, p. 223, 1910. 

Knowlton, U. S. Geol. Survey Prof. Paper 130, p. 134, pI. 

22, fig. 1, 1922. 

Fious i"'J'cgula,ris (Lesquereux) Lesquereux, U. S. Geol. and 

Geog. Survey Terr. Ann. Rept. for 1874, p. 304, 1876; 

U. S. GeoI. and Geog. Suryey Terr. Bull., vol. 1, p. 368, 

1876; The 'l'ertial'Y flora, p. 196, pI. 34, figs. 4-7, 1878; 

Hurvard· CoIl. Mus. Compo Zoology Bull., vol. 16, p. 50, 

1888. 

Ward, U. S. Geol. Survey Sixth Ann. Rept., p. 552, pI. 44, 

fig. 4 [not fig. 5, which=Rh(tnun,1ts gohliantts], 1886; 

U. S. Geol. Survey Bull. 37, p. 38, pI. 20, fig. 4 [not fig. 5, 

which=Rham1H18 gouUan1t8], 1887. 

Knowlton, U. S. GeoI. Suryey Bull. 163, p. 51, 1900. 

bl Lesqnereux, Leo, The Tertiary floro, p. 180, pl. 24, fig. 1, 1878. 

ss Lcsquercux, Leo, 'file TertillJ'Y f1orll, p. 201, pI. 31, figs. 1·-~. 10·-12, 

1878. . 

85492-30--6 

Ul11M.lSf irrcgularis Lesquereux, U. S. Geol. and Geog. Survey 

'l'err. Ann. Rept. for 1872, p. 378, 1873. 

Occurrence: Denver formation, Golden, Colo. 

(types). Laramie formation, Marshall and 114 miles 

south of Golden, Colo. Hanna formation, Carbon, 

Wyo. (doubtful). 

Ficus martini Knowlton, n. sp. 

Plate 40, Figure 5 

Leaf fairly large, ovate, with a broadly roundedtruncate, slightly auriculate base and a pointed tip. 

niargins 'entire and texture apparently coriaceous. 

Length about 15 centimeters; maximum width, below 

the middle, about 9.25 centimeters. Petiole missing. 

Midvein stout, prominent on the under side of the leaf: 

Secondaries stout, about seven somewhat irregularly 

spaced pairs, diverging from the lnidvein at angles of 

about 45°, camptodrome close to the nlargins. Tertiaries not seen: 

This species, of doubtful generic relationship, is 

based upon the single specimen figured .and is named 

for the collector, George C. niartin. 

Occurrence: Denver formation, dump of abandoned 

mine at Scranton, about 22 miles east of Denver, Colo. 

Ficus cldridgi Knowlton, n. sp. 

Plate 59, Figure 4 

Leaf relatively small, ovate, with a slightly cordate 

base and abruptly pointed tip. Margins entire. 

Length about 7 centimeters; maximum width, below 

the middle, about 4.25 centimeters. Petiole missing. 

Midvein, relatively stout, straight, and prominent. 

Secondaries stout, about seven, mostly alternate, camptodrome pairs, diverging from midvein at wide angles 

and all except the basal pair rather straight in their 

courses. Tertiaries obsolete in the coarse matrix. 

This small form, apparently new, is tentatively referred to the genus FiC1.ts, although this relationship 

is somewhat uncertain. It is not unlike FiC"1/'s 8ubtrwncata Lesquereux. It is named for the late George 

H. Eldridge. 

Occurrence: Depver formation, dump of abandoned 

mine at Scranton, about 22 miles east of Denver, Colo~ 

Ficus neopJanicostata Knowlton 

Plute 28, Figures 3, 4, 6, 7; Plate 29, Figures 3, 4 

Pi.o·ll8 neoplMl,icostata, Knowlton, U. S. Geol. Survey Prof. Paper 

101, p. 303, pI. 73, fig. 4; pI. 74, figs. 2, 3; pI. 76, fig. 4, 

1917 [1918]. 

Leaves thick, ovate, broadest at or just below the 

middle, whence they are rounded to the obtusely 

wedge-shaped or nearly truncate base and above to the 

acuminate apex; margin perfectly entire; nervation 

strong, pinnate, the lowest pair of secondaries nearly 
70 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

as strong as the midrib and arising just above the base 

or sometimes nearly at the base," thus producing a 

pseudo 3-ribbed appearance; nlidrib with three or four 

pairs of secondaries, which are slightly curved upward 

and arch just inside the margin; basal pair of secondaries (ribs) with some seven or eight camptodrome 

branches on the outside; nervilles numerous," strong, 

mainly unbroken. 

This species was described originally from the Raton 

formation of southern Colorado" and was based on 

specimens that ranged in length from 4 to 10 centimeters and in width from 3 to 5.5" centimeters. The 

specimens referred to this species from the Denver 

formation average smaller than the original specimens. 

The smallest example is only about 3 centimeters long 

and 2.5 centimeters wide but does not otherwise differ. 

As pointed out in the original' discussion this species 

is undoubtedly of the type of and most closely related 

to Fie'US planieost(J)ta Lesquereux,89 of the Black Buttes 

coal group of Black Buttes, Wyo., regarded by the 

author as of post-Laramie age. It differs from this, 

however, in a number of minor though apparently constant particulars, being ovate-acuminate instead of 

elliptical with a rounded apex, and having the basal 

secondaries or ribs arising slightly above the base of 

the blade instead of at the top of the petiole. The 

finer nervation is practically the same in both forms. 

OcclU'rence: Denver formation, Golden, Colo. Dawson arkose, lower part (in beds equivalent to Arapahoe 

formation), 3,000 feet east of the Douglas mine, Sedalia, Colo. Raton formation, near Walsenburg, Colo. 

"\Vilcox formation, De Soto Parish, La. 

Ficus planicostata goldiana Lesquereux 

Plate 28, Figure 5 

F'ious plarnioostata goldiana Lesquercux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1873, p. 399, 1874; The 

Tertiary flora, p. 202, pI., 33, figs. 1-3, 1878. 

Fious planicostata clint01~i (Lesquereux) Knowlton, U. S. Geo!. 

SUl"Vey Bull. 152, p. 103, 1898; U. S. Geol. Survey Prof. 

Paper 101, p. 303, pI. 76, fig. 3, 1917 [1918]. 

This form is of the type of Ficus plamico8tata Lesquereux but differs by its larger size and above all by 

its thinner texture and lighter nervation. The 

leaves are elliptical or oval, approximating i2 centimeters in length and 6 or 7 centimeters in width, contr.acted above into a short acumen and round-cuneate 

or almost truncate at base. These leaves are conspicuously 3-ribbed from the base of the blade or the very 

top of the petiole, all being of about the same strength. 

The midrib has about three pairs of camptodrome secondary branches in the upper part, and the lateral 

80 Lesquereux, Leo, The Tertiary flora, p. 201, pI. 31, figs. 1-8, 10-12, 

1878. • 

ribs are at an acute angle and have six or eight 

branches on the outside. The nervilles are thin and 

often broken. 

Only one of the three figured types of this fornlthe original of Plate 33, Figure 2, of the report on the 

Tertiary flora-is now to be found in the United States 

National Museum." This specimen and those represented by the other figures are seen to be larger than 

the usual leaves of Ficus pZanicostata, and in this one 

the leaf substance is thinner and the nervation lighter. 

It may be, as Lesquereux once suggested, that this 

form should have full specific rank and might even be 

placed in a different genus. In this connection he 

called attention to its resemblance to Ste'Pculia variabilis Saporta, from the" Paleocene" of Sezanne. 

Some misunderstanding appears to ha ve aris~n in 

connectibn with Lesquereux's reference of his Ficus 

clintoni to F. planicostata goldiana;. Fieus olintoni 

was described 90 in 1872, and the description is on the 

same page as that of F. plarnJioostata. It was described 

as a small species with leaves only 3 to 6 centimeters 

long. In 1878, when Lesquereux published his report 

on the Tertiary flora, he appears to have intended to 

111erge his Fious cl.intoni with Fi01.iJ8 plamicostata: 

goldiana, which has large leaves about 12 centimeters 

long. He stated that four of the small leaves (figs. 7, 

8, 11, 12 of pI. 31), which he described as sinall, undeveloped leaves of F. planicostata and which were 

probably the types or at least typical of F. olintoni, 

may"be referable to this variety, F. p,zanioostata goldia'f/.O;. If this view were accepted it would require" a 

complete redefinition of F. planicostata golcU(J)'JUL, and 

this I am not able to accept. The small leaves above 

mentioned were all found at Black Buttes mixed with 

"the typical leaves of F. pZanicostata, and according 

to present understanding should be "left with the 

typical form. When Lesquereux labeled the specimens now in the National Museum he did not record 

the presence of either F. olintoni or F. planioostata 

goZdiana at Black Buttes, and hence it may be assumed that his final intention was to relegate his Ficu8 

clintoni to FiC'U8 planicostata. 

The above explanation has been nlade especially 

because of the effect on the matter of nomenclature. 

If Ficus clintoni is to be merged with F. planicostata 

goldiana, it would be necessary to replace goldiana by 

tlintoni, on the ground of priority of publication. 

This is the disposition I made of the matter when I 

restored olintoni in the several publications indicated 

in the synonymy at the head of this form. 

Ficus planicostata goldiana is not a very abundant 

form, though several leaves identical with the type 

~pecimens have been found in the recent collections 

90 Lesquereux, Leo, U. S. Geol. and Geog. SUl'vey 'I'err. Ann. Reot. 

for 1872, p. 393, 1873, 
SYSTEMATIC PALEOBOTANY 71 

from Golden. There are several well-marked leaves 

referred to this forln from the Raton formation at 

Trujillo, Colo. 

Occurrence: Denver formation (types) , Golden, 

Colo, Raton formation, Trujillo, Colo. 

Ficus planicostata problematica Knowlton, n. var. 

Plate 28, Figure 2 

Leaf .small and evidently rather thin, ovate-elliptical, very slightly unequal-side-d, rounded about equally 

to both base and apex; nervation palmately 3-ribbed 

from the base, midrib slender, straight, with two pairs 

of alternate, thin secondaries which curve upward, the 

npical pair curving in until they nearly or quite reach 

the extreme apex of the leaf; lateral ribs of about the 

saIne strength as the midrib, slightly curving upward 

and reaching within a short distance of the apex, each 

with six or seven light secondary branches on the outside only, these curving upward and disappearing just 

within the margin; nervilles'numerous, approximately 

at right angles to the midrib; finer nervation obsolete. 

The single example figured is all that has been 

found thus far. It is a small ovate-elliptical leaf, 

about 5.5 Gentimeters in length and 3.3 centimeters 

wide n.ear the middle, and slightly unequal-sided. 

This leaf was at first supposed to be referable to Ficus 

l}lamicostata Lesquereux, with which it is undoubtedly 

most closely related, but it is evidently a much thinner 

leaf, is slightly more elliptical, and has only two pairs 

of secondary branches on the midrib, the upper pair 

curving inward rather more than is usual in the 

former species. As regards apparent texture it is 

nearer F. plamtioos't(JJt(J) goldiana, but it is hardly onefourth the size of leaves o.f that form and moreover 

has 'a different facies. It is quite possible, however, 

thnt additional material may show that this leaf 

should be referred to one or the other of the abovementioned forms, but in the meantime it may be 

referred as above. 

Occurrence: Denver formation, south face of South 

Table ~10untain, 100 feet below the lava cap, Golden, 

Colo., collected by Arthur Lakes, 1890. 

Ficus praetrinel'vis Knowlton 

. Plate 28, Figures 8-10 

Ficus 2))'aetrine'1'vi8 Knowlton, U. S, Geol. Survey Prof. Paper 

101, .pp. 263,304, pI. 41, figs, 1-4 ; pI. 42, fig. 1, 1917 [1918]. 

Following is the original description of this species: 

Leaves of fit'm texture, broadly ovate in shape, more or 

less abruptly rounded below to the decurrent base, and apparently rather obtuse above; margin perfectly entire; 3-

ribbed from the base of the decurrent or wedge-shaped basal 

portion of the blade; central rib (midrib) straight, terminating in the tip of the blade, with two or three pairs of secondaries in the upper portion; lateral ribs nearly or quite 

equal in size to the midrib, equally dividing the distance 

between the midrib and the margin of the blade, curved 

and becoming approximately par~llel to the midrib, thin above 

and apparently lost before reaching the upper third of the 

blade, each with two to four secondary branches on the outside, these being camptodrome and arching well inside the 

margin. 

The type locality for Ficus praetrinervis. is in the 

Vermejo formation of the Canon City field of. Colorado, and it was subsequently determined to be abundant and widely distributed throughout the Raton 

Mesa area of Colorado and New Mexico. The Canon 

City material studied had passed through Lesquereux's hands, this species being identified by him 

as Oinnamomwm affine Lesquereux. There is a considerable . range in size. The larger examples are 

about 10 centimeters long and about 7 centimeters 

wide' , smaller ones not more than 6 or 7 centimeters 

long and 4 centime~rs wide. An average size is 

about 8 by 5 centimeters. The petiole is especially 

well preserved in some of the Canon City material 

and is from a third to almost half as long as the 

blade; the three principal ribs run clear to the 'Very 

base and are compressed and lie side by side in the 

lower part. 

Ficus praetrinervis was also found at severallocalities in the Raton formation of southern Colorado, thus 

proving to be one of the few species crossing the line 

between Cretaceous and Tertiary. 

The collections made in the Dawson arkose on 

Jimmy Camp Creek, east of Colorado Springs, contain 

a dozen or nlore examples that undoubtedly belong to 

Fious praetrinervis and show the same range in size 

as the types from Canon City, or even a greater range. 

These specimens also show more details of the finer 

nervation than the types. ,Four pieces of matrix have 

been figured, depicting six leaves. The smallest leaf 

(fig. 10) was about 4 centimeters long and 2 centimeters wide. Figure 9 is given because it shows the 

upper portion of a leaf of a form which for some 

reason appears to be infrequently preserved. It is 

rather more pointed than the type specimens seem to 

have been. The neryation of these· specimens is well 

. brought out in the figures. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek O.() mile above Richfield Springs ranch 

. house, 9 miles east of Colorado Springs, Colo., collected by F.R. Knowlton and W. T. Lee, July, 1910. 

Raton formation, near Cokedale and Dean, Colo. 

Vermejo formation, abundant in the Raton Mesa re-

. gion of Colorado and New Mexico. 
72 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Ficus puryearensis Berry? 01 

PI~te 29, Figures 1, 2 

Ficus purycm'cnsis Berry, U. S. Geol. Survey Prof. Paper 91, 

p. 205, pI. 27, figs. 4, 5; pI. 28, fig. 5; pI. 30, figs. 4, 5, 

1916. 

This species is described by Berry as follows: 

Leaves of medium size, smooth and coriaceous, oblong-Ianceolate in outline. Apex shortly pointed.. Base broadly rounded, 

slightly cordate, or very broadly pointed. Length ranges from 

9 to 11 c~ntimeters. Maximum width ranges from 3.3 to 5 

centimeters at or somewhat below the middle. There is consideraole variation in the appearance of these leaves, well 

illustrated in the specimens figured. The widest leaf has full, 

regularly curved margins and is short!y and broadly pointed 

distad and still more broadly pointed proximad. From this 

extreme, the leaves vary toward forms that have a rounded, 

almost truncate base and a somewhat extended tip. The extreme form as regards the extended tip has a slightly cordate 

base, with full and rounded lower lateral margins, nearly 

straight sides, and an elongated narrowed tip. Petiole short 

and stout. Midrib stout, prominent on the lower side of the 

leaf. Secondaries relatively thin, 10 to 12 subopposite to alternate pairs, remote and somewhat irregularly spaced, diverging from the midrib at wide angles, 'nearly 90°, nearly straight 

until they reach the marginal region, where they turn abruptly 

upwa rd and form a wide arch to the secondary next above. 

Tertiaries mostly obsolete. Margins entire. 

As may be noted from this description and from the 

several figures given, there is considerable variation 

in the size, shape, and nervation in this species, and 

it seems not impossible that two forms may have been 

confused. The. most abundant and most strongly 

marked form-shown in Berry's Plate 27, Figures 4 

and 5, and Plate 30, Figures 4 and 5-is oblong-ovate 

and has the secondaries very conspicuously at a low 

angle, each joining the one next above by a broad bow 

far inside the margin. The other form-shown in 

Berry's Plate 28, Figure 5-is longer and more sharply 

pointed and has the secondaries, especially the lower 

ones, at a more acute angle and passirrg up nearer the 

margin. 

The leaf here figured is most nearly like that shown 

in Berry's Plate 28, Figure 5, as mentioned above. It 

lacks the base and much of one side, but it has the 

apicat' portion well preserved. It was about 12 centiIneters long and slightly over 4 centimeters wide. It 

has the fairly strong midrib and, numerous alternate 

or subopposite low-angled secondaries, as in Berry's 

specimen, though if anything they seem to pass up for 

a greater distance near the margin before sending 

branches to join the secondary next above. 

Occurrence: Dawson arkose, Rice's clay pit, 1 mile 

southwest of Calhan, Colo., collected by F. H. I{nowlton, 1910. Wilcox group (types), Grenada and Hurleys, Miss. Lagrange formation, Puryear, Tenn., in 

beds of Wilcox age. 

III This deter~ination is extremely doubtful.-E. W. B. 

Ficus sp. 

Plate 29, Figure 5 

The collection from the Dawson arkose on Jimmy 

Camp Creek, 9 miles east of Colorado Springs, Colo., 

contains a large leaf that is here figured. It lacks all 

or nearly all of the margin and hence can not be positively determined. It was probably not less than 18 

centimeters long and 15 centimeters wide and was apparently more or less heart-shaped at the bas~. It is 

5-ribbed, apparently fllom the top of the pet.iole. 

The midrib is slightly the stronger and has a pair of 

opposite secondaries at a great distance above the base. 

The next pair of ribs is at an angle of about 45° and 

has secondary branches on the outside. The lowest 

pair of ribs is approximately at right angles to the 

midrib and probably had branches on the lower side. 

The nervilles are strong and mainly unbroken. 

This leaf resembles a number of described forms, 

but its fragmentary condition makes its full identification a matter of doubt. Thus, it suggests Ficus 

speoio8Wsima Ward,92 from the Mesaverde formation 

a.t Point of Rocks, Wyo., but it is apparently a larger 

leaf wit~ a stronger primary nervation. Without the 

details of the basal configuration I can not compare 

this part with named species. ' 

The leaf under consideration is very much liKe certain large leaves from the true Lara.mie of the Denver 

Basin 93 that have been described a.s Fieus cockerelli, 

but it can not be compared fully on account of its 

fragmentary condition. It also suggests a very large 

leaf of Ficus planicostata latifolia Lesquereux.9.' 

This is so large a leaf that it perhaps merits a fig. 

ure, although it is so fragmentary that its full character can not be made out. But there is probably 

enough of it to permit its identification if found in 

the future. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo.: collected by F. H. I{nowlton and W. T. Lee, July, 1910. 

, Ficus sp. 

PIa te 30, Figure 1 

The specimen here figured lacks so much of being 

perfect that ordinarily it would hardly merit recognition, but it evidently represents so large and striking 

a leaf that I venture to include it. It was apparently 

ovate and could .hardly have been less than 18 centimeters long and 14 centimeters wide. It is truncate 

below, with the base slightly decurrent down the peti02 Ward, L. F., U. S. Geol. Survey Sixth Ann. Rept., p. 552, pI. 65, 

fig. 1, 1886. 

93 Knowlton, F. H., U. S. GeoI. Survey Prof. Paper 180, p. 132, 1922. 

P4 Lesquereux, Leo, The Tertiary flora, p. 202, pI. 31, fig. 9, 1 '378. 
SYSTEMATIC PALEOBOTANY • 73 

ole. It is palmately 5-ribbed; the midrib is considerably stronger than the others and has at least one 

pair of strong opposite secondaries at a considerable 

distance above the base. The inner pair of ribs is at 

an angle of about 50°, straight and with several tertiary branches on the outside. The outer pair of ribs 

emerges at an angle of about 40° and also has a few 

outside branches. The finer nervation is not well 

preserved. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 1 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. !{nowlton, July, 1910. 

Ficus sp. 

fi'icU,8 (£sa1'f,foUa Ettillgshausell. Lesquereux, Harvard CoIl. 

Mus. Compo Zoology Bull., yol. 16, p. 50, 1888. 

This specimen is so fragmentary that ordinarily it 

would not be figured except for the fact that it was 

identified by Lesquereux as Ficus asarifolia Ettingshausen. The American leaves originally referred to 

this species were found in the Mesaverde formation 

at Point of Rocks, Wyo., and are sm~ll leaves not 

exceeding 4 centimeters in width and 3.5 centimeters 

ill length, whereas the present leaf was fully 12 centimeters wide and probably not less than 10 centimeters 

long. It appears to have been broader than long and 

was rounded and obtuse at the apex. The margin 

apears to have- been entire. The nervation is very 

coarse and strongly reticulated but very unlike the 

a l'eolation in the Point of Rocks leaves. 

Occurrence: Denver formation, South Table ~1ountain, Golden, Colo., collected by Arthur Lakes. Specimen identified as FiC'Us {])Sf1YrifoliaJ Ettingshausen in the 

Museum of Comparative Zoology, Cambridge~ Mass., 

No. 1441a. 

Artocarpus }lungens (Lesquereux) Hollicl\: 

Plate 30, Figure 2; Plate 31; Plate 32, Figures 1, 2 . 

Al'tocm'1HtS lHtngcns (Lesquereux) Hollick, Louisiana GeoI. 

Survey Rept. for 1899, Special Rept. 5, p. 281, pI. 38, 

figs. 1, 2, 1899. 

Berry, U. S. Geo1. Survey Prof. Paper 91, p. 195, pI. 25, 

fig. 1; pI. 27, fig. 1; pI. 29, fig. 1, 1910. 

,Aralia pungel/s Lesquereux, The Cretaceous and Tertiary 

floras, p. 123, pI. 19, figs. 3, 4, 1883. 

Leaves thick and coriaceous, very large', probably 

reaching a length of 30 centimeters and a width of 

18 or 20 centimeters, oblong in general outline, deeply 

pinnately lobed above and almost if not quite pinnate below; lobes opposite, lower and middle lobes 

directed outward at .an angle of 45° or less, the upper 

ones directed upward; lobes long, oblong-Ianceolate, 

taper-pointed, slightly broadest in the middle and 

lower portion, separated by more or less broad 

rounded sinuses, the sinuses separating the lower 

lobes extending nearly or quite to the midrib; margins entire; midrib very thick and stout, straight, secondaries proportionately thick, of two orders, the 

main ones strong, sub opposite, ascending to the point 

of the lobe and several times branched on either side, 

the others smaller, emerging from the midrib between the large ones, passing up to the sinus, whel~e 

they divide into two branches that arch around the' 

sinus just inside the margin and follow the borders of the lobes in festoons, anastomosing with the 

tertiaries; tertiaries several in each lobe, alternate, 

curved slightly or markedly in passing to the margin, 

along which they often form long festoons just inside the border; ultimate areolation in the main quadrangular, formed by divisions at right angles to the 

tertiaries. 

These were fine large leaves, but leaves of large size 

are rarely preserved complete, and although this species has been known for many years and has been 

found at a number of widely separated localities, we 

are still without a really perfect specimen. The middle and upper parts of the leaf are known from many 

specimens, but the basal portion is still somewhat in 

doubt. The lobes are separated by broad, rounded' 

sinuses, and the extent to which the indentation approaches the midrib increases from the apex downward, until what is perhaps the lowest set of lobes is 

separated from the midrib by only a very narrow 

wing, and still below this there is a pair of apparently 

distinct leaflets. This condition is well brought out 

jn the specimen shown in Plate 31. The upper part 

of this leaf is indistinguishable in shape and nervation from the figures of the type specimens. It may 

also be noted that the third pair of lobes from the 

a pex of the leaf has the sinus on one side cutting 

almost to the Inidrib, but on the other side it is nearly 

a centimeter wide. These lobes are oblong-lanceolate-that is, they are slightly constricted near the 

base. Fully 2 centimeters below the pair of lobes just 

described there is seen a single small, oblong, wedgeshaped and apparently free leaflet, and so far as can 

be made out there is no wing along the midrib connecting it with the pair of lobes p.ext above. An 

exactly similar condition is seen in the specimen 

shown in Plate 32, Figure 2, although it is considerably broken. In this specimen the third pair of lobes 

is about a centimeter below the pair next above, with 

no evidence of a connecting wing along the midrib 

(or rachis). These lobes (or leaflets) are deeply cut 

into on the upper side and expanded on the lower side. 

Two centimeters below is a small petioled leaflet 

(~p.ere was undoubtedly a pair, but the opposite one 

is destroyed) that is in undoubted organic connection. 

with the axis (rachis), which extends fOl' more than 
74 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

2 centimeters below its point of attachment. This is 

probably the real base of the leaf, though there may 

be a still smaller and lower pair of leaflets that is not 

prese"rved. 

Berry reported the presence of Artocarpus pungens 

in Wilcox beds at several localities in Louisiana and 

.A"rkansas, but neither in his figures nor in his discussion of the species is there an indication of free basal 

" leaflets such as just described for the Denver specimens. Berry mentioned the difficulty of getting good 

specimens on account of the friable nature of the matrix in which they are preserved. His best example is 

a splendid specimen from Coushatta, La.95 It represents the upper part of a leaf that was probably 30 

centimeters in length. Two other specimens 96 from 

Benton, Ark., are apparently from near the middle 

portion of the leaves, and although there is no indication of a reduction of the lobes to leaflets, they do show 

the sinus cutting pretty close to the axis. So far as 

can be judged from the specimens figured there is no 

reason to question the correctness of the reference of 

the Wilcox material to A'rtoca-r-p'INJ pungens, bv.t this" 

is confessedly without a knowledge of the configuration of the base of the leaves. 

The species under consideration was first described 

by Lesquereux 97 under the name Aralia pWnge11J8 from 

specimens obtained at Golden, Colo. The types are 

the property of Princeton University and should be 

preserved in the university museum, but at the present 

time only one of them, the original of Figure 3 of 

Lesquereux's plate, can be found." This specimen, by 

the kindness of William Libby, jr., the curator of the 

museum, I was permitted to study. It has been redrawn and is shown on Plate 32, Figure 1. In the 

figure of it as published by Lesquereux the right-hand 

upper lobe is not represented, but an examination 

shows that what is given as true margin is not correct, and there is every reason to suppose that it was 

extended into a lobe similar to the one on the opposite 

side. Its restoration is shown by the dotted lines. 

The other type specimen possessed the two upper lobes, 

a"s do all specimens that are well enough preserved for 

this feature to be made out. 

In referring these leaves to the genus Aralia Lesquereux apparently interpreted them as being the 

upper parts of very large palmately divided leaves 

with the,segments deeply cut into large lobes, such for 

il)stance as may be seen in Aralia dissecta Lesquereux,98 

from the Green River formation. Lesquereux recognized the resemblance of Aralia pwngens to his earlier 

Myrica.@ lessigian</) (no~' Artocarpus lessigiatna), but, 

811 Berry, E. W., The lower Eocene floras of southeastern North 

America: U. S. GeoI. Survey Prof. Paper 91, pl. 25, 1916. 

06 Idem, pl. 27, fig: 1; pI. 29, fig. 1. 

9'7 Lesquereux, Leo, The Cretaceous and Tertiary floras, p. 123, pI. 19, 

figs. 3, 4, 1883. 

08 Idem, pl. 35. 

evidently depending on the drawings rather than a 

study of the specimens, he decided that the nervation 

differed sufficiently to separate them generically. In 

this connection he said: 99 

This species is allied to what has been described in volume 

7 as M[lIr"ica] les8igii, but the nervation differs. In M. Ze8-

sigii the tertiary veins directed toward the sinuses divide 

under them into two branches, passing along on both sides 

and following the borders of the lobes, while in this leaf [his 

Aralia pungens] the tertiary veins do not divide but appear 

merely to pass up on· one side without forking. Though this 

difference may be marked, it is scarcely possible to doubt that 

these fragments represent the same group or the same genus 

of plants, and, as I have remarked it in ·the description of 

"M. lessigii, Saporta and other authors refer plants of this kind 

to the Araliaceae. 

A glance at the figure of Aralia pungens as redrawn 

(pI. 32, fig. 1) shows the nervation about the sinuses to 

be of the same character as that in the so-called Myrica 

lessigiana, and therefore this supposed distinctive 

character fails. 

It is now necessary to consider further the relation 

between Artocarpus pwngens and Artoc(J)1'pus lessigiuiJUt. They are undoubtedly very close, and at one 

time I was inclined to place them together,t but Berry2 

regards them as distinct and recognized both in the 

Wilcox group. He said: 

This species [Artocarpus pungens] differs from Artocarpus 

Zessigiana (Lesquereux) Knowlton, with which it is often confused, by its more orbicular general form and the great elongation and narrowness of the lobes, which are also more ascending and are separated by more open sinuses. 

I have accepted this disposition and in my paper on 

the Laramie flora 8 have kept them distinct, the more 

so because it proved that in the Denver Basin A. lessigiana.is confined to the Laramie and A. pungens to 

the Denver. The peculiar reduced and free lobes or 

leaflets just described as occurring in A. pungens might 

prove an additional and very marked point of difference between the two species if we were able to compare 

this feature in them, but unfortunately no specimen of 

A. lessigiana is known which has the basal portion 

preserved. In the type of A. lessigiaYnoJ. the lowest 

.lobe present is very broad and blunt, and it seems improbable, from the size and configuration of the whole 

leaf, that there could have been any reduced lobes or 

leaflets below that lobe, but this is of course purely 

conjectural. . 

Occurrence: Denver formation, South Table Mountain, Golden, Colo. Wilcox group, Grenaua, Miss., 

Benton, Ark., and Coushatta and N aborton, La. 

99 Idem, p. 123. 

1 Knowlton, F. H., Catalogue of the Cretaceous and Tertiary plants 

of North America: U. S. Geol. Survey BUll. 152, p. 42, 1898. 

2 Berry, E. W., The lower Eocene floras of southeastern North 

America: U. S. GeoI. Survey Prof. Paper 91, p. 195, 1916. 

3 Knowlton, F. H., Laramie flora of the Denver Basin: U. S. Geol. 

Survey Prof. Paper 130, p. 128, 1922. 
SYSTEMATIC PALEOBOTANY 75 

Artocal'pus similis Knowlton 

Plate 32, Figure 4 

ArtoOll1'pus s'im,iZis Knowlton, U. S. GeoI. Survey Prof. Paper 

101, p. 306, pI. 77; pI. 78, figs. 1, 2, 1917 [1918]. 

The type of this species comes from the Raton formation of the Raton ~1esa region of southern Colorado , where it was found at a number of localities. 

It was shown to be most closely related to forms from 

the Denver Basin, which are now believed to represent 

two species. These- are A1>toca1>pus lessigiana (Lesquereux) Knowlton (originally described as Myrica? 

lessigia'fU}) and Artocarpus pungens (Lesquereux) 

}{nowlton (first named Aralia; pUingens). Artocarpus 

Sil1~ilis is, in a way, intermediate between the two Denver Basin forms, having the narrow, acut~-pointed 

Jobes of A. pungens and the ovate, obtuse terminal 

lobe of A. lessigi(]Jna. 

The example here figured is almost exactly the same 

as the type figured on Plate 77 of the Raton Mesa 

report except that it is much smaller. It is only a 

little more than 7 centimeters wide between the points 

of the upper lobes and could hardly hare been much if, 

any over 12 centimeters in length. . 

Occurrence: Dawson arkose, Rice's' clay bank, 1 mIle 

southwest of Calhan, Colo., collected by F. H. Knowlton, Ju(y, 1910. Raton formation, southeastern Colo:. 

rado. 

Artocarpus? gigantea Knowlton, n. SPa 

The collection from Jimmy Camp Creek, east of 

Colorado Springs, Colo., contains the remains of a 

leaf that, although very imperfectly preserved, is so 

bb (TiO'antic and so strikinO' in appearance as to warrant b 

n, name and description. Very little, if any, of the 

margin is retained, yet the portion of th~ leaf pre~ent 

is 30 centimeters long and about 15 centImeters WIde. 

It was probably at least 40 centimeters long, and, to 

judO'e from the thickness of the midrib where it is 

broken at the base and apex, it might well have 

reached a length of 50 centimeters. It appears. to 

have been elliptical or oblong in general outlIne, 

though the absence of the margin makes this Ul~certain. There appears to be a shallow, rounded SI~US 

near the base on the right-hand side, giving indicatlOn 

of the presence of a large lobe, and that there .were 

probably ot1;ler lobes is suggested py the exceedIngly 

~trong secondaries in the middle and upper part of t~e 

blade. The secondaries number at least five or SIX 

pnirs. They are very strong, alternate, and at irregular distances. 

The fragmentary character of this leaf makes a 

knowledge of its generic reference uncertain. Its size 

and apparent shape, the probable lobe i~ the lower I 

portion, and the character of the nervatlOn suggest 

at once the O'enus ..A1'tocarrpus. It is perhaps nearest 

to what Ber~T 4 has identified as Artocarpuslessigiana 

(Lesquereux) I{now lton, from the Wilcox formation 

at Shreveport, La. 5 I am not certain that the vVilcox leaves are conspecific with the type specimens of 

A. lessigiana, which came from the Laramie of the 

Denver Basin, though they may be. The leaves of 

Artocarpus are sometimes identified with difficulty. 

They are such large leaves that it is unusual to find 

them wholly preserved, and moreover they are usually 

very rare, thus making in difficult to procure l,?aterial for comparison. Berry states that the WIlcox 

spec~mens are large leaves having a maximum lengt? 

of 30 centimeters and a maximum width of 20 centImeters. In the figure given by Berry the secondaries 

are seen to be at a low angle with the midrih, whereas 

in the leaf under discussion the secondaries are at 

an angle of about 450 and do not curve do,vnward, so 

far as can be made out. The lower lobe, if it is such, 

in the present species is separated by a deeper sinus 

than in Berry's figured specimen, and there are some 

differences in the details of nervation. But none 

of these differences are great, and it is possible that 

they might be shown to be closer if a sufficient series 

were available for comparison. For this reason I 

have decided to give the leaf under consideration a 

new name and 'trust to the future to settle its status. 

It has some resemblance to A1·tOC(Jff'pus pungens 

(Lesquereux) Hollick, which has just been discus~ed, 

though it is very much larger and of course gIves 

no evidence of the presence of the basal characters 

described in that species. 

The specimen of this form is so fraglnentary that 

I have not been able to figure it. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 0.6 mile north of Richfield Springs ranch 

house and 9 miles east of Colorado Springs, Colo.,' 

collected by F. H. Knowlton and W. T. Lee, July, 1910. 

Order FLATANALES 

Family FLATANACEAE 

Platanus rhomboidea Lesquereux 

Plate 32, Figure 3 

Pla.ta.1'lIl£s rnom'boldeat Lesquereux, U. S. Geol. and Geog. Suryey Terr. Ann. Rept. for 1873, p. 400, 1874; The Tertiary flora, p. 186, pI. 26, figs. 6, 7, 1878; Harvard Co11. 

Mus. Compo Zoology Bull., vol. 16, p. 49', 1888. 

Knowlton, Washington Acad. Sci. Proc., yol. 11, p. 204, 

1909; U. S. Geol. Survey Prof. Paper 101, p. 324, 1917 

[1918]. 

Vi'b~£rnum (M'/,oeps Lesquereux, The Tertiary flora, p. 227, pl. 

38, fig. 11, 1878. 

'Berry, E. W., U. S. GeoI. Survey Prof. Paper 91, p. 194, pI. 26" 

1916. 

II It is undoubtedly identical with the Louisiana material.-E. W. B. 
76 FL·ORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Lea ves small or medium sized (7 to 12 centimeters 

10nO', 5 to 9 centimeters broad), subcoriaceous, rhom- e . . boidal. 3-ribbed and .in some specimens slIghtly trIloba.te; being broadest at about the nliddle, from which 

point they are narrowed and entire to the wedgeshaped base and about equally narrowed above, where 

they are more or less deeply, usually sharply dentate; 

I~lteral nerves (secondaries) at an acute angle of divergence, parallel, the lower pai~ a little the longer 

and entering short acute lobes, each with three or 

four branches on the outside, which also pass to marginal teeth; nervilles nearly at right angles to the 

midrib, mostly percurrent; finer nervation quadrangular. 

The two figured' type specimens of Platanus rh01nboidea are preserved in the United States National 

Museum (Nos .. 245, 246) and are from the characteristic andesitic material of the Denver formation at 

Golden, Colo. They were the only specimens recorded 

in the Museum catalogue that were so identified by 

Lesquereux, but in the large lot of specimens belonging to the J\1useum of Comparative Zoology that were 

studied by Lesquereux in 1888 no less than six additiona,} examples were recorded, and in the large collection fronl Golden procured by Arthur Lakes in 

1890 and studied by me a number of characteristic 

specimens were found. This species has since been 

detected in the Lance formation in Weston County, 

'''yo., and in certain beds southwest of Rawlins, Wyo., 

in beds regard.ed by me as of post-Laramie age. It 

has also been found at a number of localities in the 

Raton formation of Colorado and New Mexico. 

In 1878 Lesquereux described and figured in the 

report on the Tertiary flora (pI. 38, fig. 11). a single 

leaf under the name Viburnum anceps. After a careful exalnination of the type (U. S. Nat. Mus. No. 326) 

I am unable to find any character that will separate it 

Platanus guillelmae Goppert 

Plate 33, Figure 2; Plate 34, Figure 1 

Platan.U8 (Juil elmae Goppert. Lesquel'eux, U. S. Geol. and 

Geog. Survey Terr. Ann. Rept. for 1871, p. 289, 1872; 

The Tertiary flora, p. 183, pI. 25, figs. 1-3, 1878. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 322, 

pI. 93, fig. 1, 1917 [1918]. 

In discussing certain platanoid leaves frOln the 

Raton Mesa region of Colorado and New Mexico, I 

took occasion to say 6 that I found a number of wellpreserved leaves that were indistinguishable from the 

lIsual figures given for Platanu8 guillelmae, such as 

those given by Heel' 7 from the Tertiary of Greenland. 

The same statement applies to certain leaves from the 

DenveI~ formation. The one here figured (pI. 33, fig. 

. 2), for example, is not to be distinguished from the 

figured specimen from the Raton formation, except 

that it is slightly larger. It is about 11 centimeters 

long and 12.5 cent~meters wide, truncate at the base, 

and with margins entire for some distance and thence 

with small sharp teeth. It is distinctly 3-10bed, with 

the lateral lobes small and rather obtuse. The lateral 

. ribs arise some distance above the base of the blade. 

These leaves somewhat resemble small leaves of 

Plat((fnus a'ceroides latifolia I(nowlton,s but they differ 

in being more nearly truncate at the base, in' having 

shorter lateral lobes, and in having the lateral ribs 

arising well above the base instead of at the top of the 

petiole. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Raton 

formation, H.atoll J\1esa region of southern Colorado 

and adjacent New Mexico. H~nna formation, Carbon, Wyo. Fort Union formation, Glendive, Mont. 

Lance formation, near Glendive, Mont., and Dayton 

and Kubey, 'Vyo. 

from Plat(J;nus rho171,boide(J). It is based on the upper Platanus aceroides latifolia Knowlton 

portion of a leaf of the same size as the smaller of the Plate 33, Figure 1 

two types of P. rlv01nboidea. The general shape of the _ . . . .. Platantt8 acel'oides latifolia Knowlton, U. S. Geol. Survey Prof. 

leaves, as well as the nervatIOn, IS preCIsely the same, Paper 101, p. 321, pI. 92; pI. 93, fig. 3; pI. 94, 1917 [1918]. 

the only difference being that in the so-called V ibur- Berry, U. S. Geol. Survey Prof. Paper 91, p. 13, 1916. 

1wmb anceps' the teeth are not quite so prominent nor This variety is similar to' the type but is propol'-

sharp.. . tionately broader and shorter, the margin provided 

Occurrence: Denver formatIOn (types), Golden, with numerous small, rather blunt teeth: It is of the 

Colo.; ~ubsequently c?llected by Arthur Lakes from type of the living Platan~(;s occidentaUs and may be 

South rable MountaIn, Golden. D.awson arkose, a 1 'b db' f d' . 3 'bb d d 3 . . c escn e as emg 0 me Ium SIze, -rI e an - quarter of a mIle east of the Purdon nnne, ~ec. 27, T. lobed, the base truncate, slightly cordate Or rarely 

11 S., R. 61 W., Colo., collected by G. B. RIchardson. more or less subcuneate the lobes broadly trianO'ular 

Raton formation, near Raton and mouth of York Can- , b 

yon, N. Mex., and several localities near Wootton, 6 Knowlton, F. H., U. S. Geo1. Survcy Prof. Paper 101, p. 322, 1917 

Colo., all collected by W. T. Lee. Lance formation, [l~~~~~r, Oswald, Flora fossilis at'ctiea, vol. 7, p. 96, pI. 9;, fig. 6; 

~Teston County, Wyo. Hanna formation, near Raw- pI. 99, fig. I, 1883. 

B Knowlton, F. H., U. S. Geol. SUl'vey Prof. Paper 101, p. 321, pI. 93, lins: WyO. fig. 3, i917 [1918]. 
SYSTEMATIC PALEOBOTANY 77 

\'lith the margins dentate, the m~clclle lobe with two 

to four teeth and the lateral lobes with a few more~ 

the teeth usually strong, in many specimens curved or 

even slightly hooked, and invariably sharp pointed. 

This form is probably the mdst abundant member of 

the genus in the Raton formation of the Raton Mesa region of Colorado and New Mexico. In a single collection obtained near 1Vootton, Colo., there are more 

than n hundred specimens, some of them absolutely 

perfect. It is discussed at some length in my paper 

on t.he flora of the Raton formation. 9 

In the material frOln the Denver formation at 

Golden, Colo., there are a number of specimens that 

clearly belong to this variety. 4 They are smaller than 

lBany from the Raton formation but agree with some 

of the smaller leaves. Thus, the specimen figured 

seems indistinguishable from the one shown on Plate 

94 of the paper above mentioned. 

Platanus aoe1'oides latifolia was found by Berry 10 

in the l\1idway ( ~) formation at Earle, Bexar County, 

Tex. 

Occurrence: Denver formation, South Table Mountajl1~ Golden, Colo. Raton formation, Raton l\1esa 

l'egion of Colorado and New l\1exico. l\1idway( n 

fOl'n"lfition, Earle, Bexar County, Tex. 

Platal\US raynoldsii Newberry 

Pin te 3G, ]j'jgure 1 

I'/atanl/IS ,'ayuoZusi.i Newberry, New York Lyceum Nat. Hist. 

Annals, vol. 9, p. 69, 1868; Illustrations of Cretaceous 

nnd Tel·tiary plants, pI. 18, U. S. Geol. and Geog. Survey 

Terr., 1878 [plates prepared by Newberry; names as-

~igned by Lesquel'eux]; U. S. Geol. Survey Mon. 35, p. 

109, pI. 35, 1898. . . 

]{.Rsquereux, U. S. Geol. anel Geog. Survey 'rei r. Ann. 

Hept. for 1872, pp. 379, 399, 1873; idem for 1873, p. 381, 

1874; idem for 1876, p. 507, 1878; The Tertiary flora, 

p. 185, pI. 27, fig. 2, 1878; Illustrations of Cretaceous and 

Tertial'Y plants, pI. 18, U. S. Geol. and Geog. Survey 

Terr., 1R78 [plates prepared by Newberry; names as-

~igned by Lesquet~eu~]; Harvard CoIl. Mus. Compo 

Zoology Bull.. vol. 16, p. 49, 1888. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 3~4, 

111. 95, fig. 1, 1917 [1918]. 

Tho following is the original description given by 

Newberry: . 

J.JcnYl's of large size, subol'biculaL' 01' rudely triangular in 

outline, more or less rounded below, three-pointed above, often 

decurrent on the petiole; margins at base entire, on the sides 

:llld above coarsely and obtusely doubly serrate, the lobes of 

the upper margin short and broad, less produced than in most 

othC'l' species; nervation strong but open, having the· usual 

cba I'nctcl' of P. oooi!Zentali8 and of the fossil species P. 

(lccroi!Z08. 

o Knowlton, F. II.,' U. S. Gco}' SUl'vey Prof. Papel' 101, p. 321, 1917 

(1918). 

10 R(\l'r~·. E. \\'., U. S. Gcol. Survey Prof. Paper 91, p. 13, 191G. 

The type specimens carne from the Fort lJnion foi'-

mation near the Inouth of the Yellowstone River, wIontana. It was reported as abundant in the collections 

examined by Newberry and was found by 'Vard ll in 

approximately the same locality 20 years later. In 

recent years it has "been collected from a great number 

of localities in l\10ntana and the Dakotas; in fact, it 

proves to be one of the most abundant and widely 

distributed Fort Union species. 

Soon after Platan1.(;s raynoldsii was described from 

the Yellowstone region it was reported by Lesquereux as occurring at Black Buttes, Wyo., and Golden, 

Colo. The specimens from Black Buttes were said to 

have come f1'om the saurian bed, but they are now 

lost, and none have since been obtained. 

Platanus Irayno1ldsii appears to be closely related to 

Plata;nus haydenii, and unless the specimens are fairly 

well preserved it is sometimes very difllcult to distinguish between them. As defined by Newberry they 

could be separated by the characters of the marginal 

dentition, being simply toothed in P. lwydenii and 

doubly serrate in P. raynoldsii, but an examination 

of the figured type specimen of P. 1'aynoldsii/2 now in 

the N ationall\1useum, discloses the fact that it is simply toothed like the other. On removing the matrix 

it was found that what appeared to be smaller teeth 

on the large ones were due entirely to the manner in 

which the matrix was fractured along the margin of 

the leaf. The two species may be distinguished, however, by characters drawn from the shape and nervation of the basal part of· the leaf. In Plat((;l1.1.(,8 1Ylly-

'Tloldsii the base of the blade 'is rounded to a wedgeshaped portion decurrent on the petiole, and the lowest 

pair of lateral ribs arise a considerable distanc~ above 

the base of the blade. In P. ha;yclen,ii the base is 

nearly or quite truncate, with no decurrent portion 

along the petiole, and the large ribs arise wjth the 

midrib at the very base of the blade. This shows why 

in the absence of complete leaves it is so difficult t~ 

distinguish between these species. 

Platan1.lS 1'aY1wldsii is one of the most abundant 

species in the Denver formation at Golden. Colo. So 

far as can be made out, all the speci~ens that pa~sed 

through Lesquereux's hands up to the time of the 

publication of his report on the Tertiary flora (1878) 

he intended to refer to his variety integrifolia, though 

this is by no means certain, as is explained on a subsequent page. In any event, in the list of specimen's 

identified for the :Museum of Comparative Zoology 

the 23 specimens found were recorded under the original specific name only. It is therefore doubtful 

whether it was Lesquereux's intention to abando~l the 

11 Ward, L. F., U. S. Geol. Survey BUll. 37, p. 37, 1887. 

12 Newberry, J. S., Tbe later extinct floras of North Amel'ica: U. S. 

Geol. Survey l\Ion. 35, pI. 35, 1898. 
• 

78 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

so-called varietal form, or whether all th~ specimens 

happened to belong, to the species and not the variety. 

It, is certain that among the recent collections there 

are a number of specimens that belong to the specific 

form as well as some that might be referred to the 

varietal form. It is to be noted, "however, that the 

leaves referred to P. raynoldsii from Golden, Colo., 

are in general much smaller than the specimens from 

the Fort Union. Thus the specimen figured by Lesquereux 13 is absolutely identical with the type as 

figured by Newberry.14 In Newberry's text it is stated 

'that this is one-half the linear size of the largest 

specimen obtained. 

In the material obtained at Carbon, 'V yo., by Ward, 

I find several fragments of a large Platanus leaf, 

among them being one (U. S. Nat. Mus. No. 1080) 

having the decurrent basal portion and a part of the 

petiole fairly well preserved. For some distance the 

margin appears to be perfectly entire, but soon it becomes obscurely toothed, the teeth being low and 

separated by shallow sinuses. The upper part of the 

leaf is not preserved, but so far as can be made out it 

was probably 3-10bed. The basal portion of this leaf 

is undoubtedly like that of typical P. rayrnoldsii, and 

it is so considered. 

Platrunus raynoldsii is also present in the Raton formation of the Raton Mesa region of Colorado and New 

Mexico.11> 

Occurrence: Fort Union formation (types), near 

mouth of Yellowstone River, Mont.; abundant and 

widely distributed in the Fort Union formation. Denver formation, south face of South Table Mountain, 

Golden, Colo. Hanna formation, Carbon, Wyo., collected by Lester F. Ward, 1883. Raton formation, 

Raton Mesa of southern Colorado and adjacent New 

M~exico. 

Platanus raynoldsii integrifolia Lesquereux 

PIa te 34, Figure 2 

Platan'U8 raynoldsii in.tegrifolia Lesquerenx, The Tertiary flora, 

p. 185, pI. 26, figs. 4, 5, 1878. 

There is some confusion and uncertainty regarding 

the status of this form, as it is difficult to make out just 

what Lesquereux intended to do. It was established 

on material said to have come from Golden, Colo., and 

Black Buttes, Wyo., but a study of the matrix of the 

figured types now preserved in the United States N aLional Museum does not bear out its presence at Black 

Buttes. In the report on the Tertiary flora, where the 

variety integrifolia was established, Lesquereux first 

employs the heading Platanus raynoldsii and under it 

18 Lesquereux, Leo, The Tertiary flora, p. 185, pI. 27, fig. 2, 1878. 

14 Newberry, J. S., op. cit., pI. 35. 

16 Knowlton, F. H., U. S. Geo1. Survey Prof. Paper 101, p. 324, pI. 95, 

fig. 1, 1917 [1918]. 

places all the plate references, yet the description follows the introduction of the varietal name. In the discussion of this form given by Lesquereux it is not clear 

whether he intended to call all the specimens integrifolia or whether a part were to be regarded as referable 

to the species. In the explanation of the plates the 

figures on 'Plate 26 are referred to as Plata/m.tS raynoldsii var. i'l'Vtegrifolia, but on Plate 27 the figures are 

all given to the species, and they are all recorded in the 

Museum catalogue under the same designation. 

Lesquereux's description of the variety is as follows: 

Leaves of large size, sub orbicular or obscurely triangular in 

outline, more or less rounded and entire toward the decurrent 

base, dentate, serrate or undulate, even entire, subcoriuceous. 

This description, it will be observed, agrees with the 

characterization of the species given by Newberry, 

except as regards the margin. Four of the five specimens figured by Lesquereux in the report on the Ter'-

tiary flora are now in the National Museum. They 

are all preserved in the andesitic material characteristic of the Denver formation at Golden, Colo., 

although one of them-the original of Plate 26, Fig- , 

ure 4-is said by Lesquereux (p. 186) to have come 

from Black Buttes, Wyo. These leaves are all ilnperfect, and it is difficult if not impossible to determine 

the exact character of the margin~ and I strongly suspect that Lesque~eux's description was prepared 

largely from the drawings and not from the actual 

specimens. For example, t~ke the specimens shown 

on Lesquereux's Plate 26 under the designation Platanus raynoldsii var. integrifolia. The larger leaf is 

Figure 4 (U. S. Nat. Mus. No. 243), which is shown 

as if practically the whole margin were preserved and 

were entire or slightly undulate; but with the exception of a small basal portion below the lowest tertiary 

branch of the basal rib the margin of the, entire lefthand side of tlie leaf is missing, and the margin of 

the right-hand side is indefinite except for perhaps a 

distance of 5 centimeters near the base. Figure 5 of 

this plate is described as perfectly entire. This specimen (No. 244) is very poorly preserved, especially th~, 

margin; in fact, it is doubtful if any margin at all is 

retained. If any of it is true margin it appears to be 

llndulate-toothed rather than perfectly entire. 

On Plate 27 of the report on the Tertiary flora there 

~re three figures referred to Platanus'raynoldsii'without the varietal designation. Two of these specimens 

(figs. 1 and 2) are before me; the better-preserved one 

is that shown in Figure 2 (U. S. Nat. Mus. No. 248). 

In the figure the margin is shown on practically the 

whole of the left-hand side, but the specimen does not 

bear this out, for with the exception of the entiremargined decurrent basal portion of the blad.e· with 

perhaps a length of 4 centimeters adjacent, the whole 

margin of this siae is either absent or so imperfectly 
SYSTEMATIC PALEOBOTANY 79 

preserved that its presence is impossible to determine 

certainly. The other specimens are even more fragmentary and afford no additional information regarding the character of the margin. 

Among the specimens recently collected at Golden 

{Ire a considerable number that fron1 the configuration 

of the base should be referred to Platamws ra:ynolrlsii, 

but unfortunately there is not one that shows the complete outline or gives positive evidence regarding the 

marginal teeth. I think Lesquereux was probably . 

right in believing that these leaves from Golden are 

more nearly entire than the types from the Fort 

Union formation, but they are not so entire-margined 

as the figures would imply. On account of the con-:-

c1itions above set. forth it has been thought best to 

retain the variety integrifoli(() as established by Lesquereux and to leave to the future the determination of 

its exact status. 

In the collections from Converse County, Wyo'J I 

find a single fragmentary leaf that appears to belong 

tv Platoin/us 'ray'rl.,olcJsii infJegrifolia. ' It is of about the 

sume size as the specimens figured by Lesquereux 

being 14 or 15 centimeters long and about 13 centi~ meters wide, and has a portion of a strong petiole preserved. Unfortunately only a very small portion of 

the margin is preserved, but so far as can be made out 

it was not a 3-lobed leaf and has the margin rather 

coarsely undulate-toothed-that is, both teeth and 

sinuses appear to be rounded. It is certainly not 

doubly dentate, as described for the type specimens, 

but seems to agree more with Lesquereux's so-called 

variety integrifoUa. It is possible that if more material were available differences would be observable. 

In the material from Middle Park, Colo., I also find 

n single small, much broken leaf that appears to be 

referable to this variety. Only a portion of the margin near the base of the blade is preserved and this 

is entirely without teeth. ' 

Occurrence: Denver formation [types] South Table 

Mountain, Golden, Colo., collected by Ar~hur Lakes in 

the early seventies and in larger numbers about 1890. 

M~ddle Park formation, Sheriff Creek, north bank of 

Grand [Colorado] River, Middle Park, Colo., colle.cted by George L. Cannon. Lance formation, LightnIng Creek opposite mouth of Box Creek Converse 

County, Wyo., collected by J. B. HatGher.' ' 

Platanus haydenii Newberry 

Plate 36, Figure 1 

Platanu8 haydernii Newberry, New York Lyc. Nat. Hist. Annals, vol. 9, p. 7, 1868; Illustrations of Cretaceous and 

Tertiary plants, pIs. 19, 20, U. S. Geol. and Geog. Survey 

Terr., 1878 [plates prepared by Newberry; names assigned 

by Lesquereux]; U. S. Geol. Survey Mon. 35, p. 103, 

pI,s. 36, 38, pI. 56, fig. 3, 1878. 

Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. 

Rept. for 1871, pp. 287, 289, 290, 1872; idem for 1872, p. 

379, 1873; idem for 1873, p. 381, 1874; idem for 1876, 

p. 507, 1878; Illustrations of Cretaceous and Tertiary 

plants, pIs. 19, 20, U. S. Geol. and Geog. Survey Terr., 

1878 [plates prepared by Newberry; names assigned by 

Lesquereux]; Harvard 'ColI. Mus.) Compo Zoology Bull., 

vol. 16, p. 48, 1888'. 

This species was described by Newb.erry as follows: 

Leaves large, long-petiolecl, when mature three, perhaps 

rarely five lobed; lobes nearly equal, long-pointed, acute; on 

either sicle of the midclle lobe five to eight obtuse teeth; margins of the lateral lobes sinuately toothed to near the base; 

younger leaves ovate, acuminate, coarsely toothecl throughout, 

except near the base, which is slightly decurrent; nervation 

strong, racliate from the base, primary nerves three, which 

are nearly straight and terminate in the three lobes of the 

borcler. From the ~idrib spring seven or eight pairs of lateral nerves above the basilar pair; these diverge at an angle 

of about 35°, are slightly flexed at the base, straight or nearly 

so above, where they are somewhat truncated, their branches 

terminating, in the" marginal teeth. The basilar Jierves diverge 

from the midrib at an angle of about 35° and run nearly 

straight to the extremities of the lateral lobes. They each 

give off on the lower side seven or eight branche::l, of which 

the second or third is straight. These are more or 'less curved 

and branched, the branches terminating in the teeth of the 

margin. Fruit 2-3 lines long, prismatic, clavate. 

This is the original and most complete description 

that has been given of this species, which was first obtained on the banks of the Yellowstone River near its 

mouth. It is said by Newberry to be well represented 

by specimens, though but few of these are present in 

the collections of the United States National Museum. 

Ward, who collected extensively in the same general 

area as the type locality in 1883, did not procure any 

of the large mature leaves, though he did procure a 

number of leaves identical with those referred by Newberry to the young forms of this species. 

I have studied the two leaves figured by Newberry 

as the young, immature leaves 'of P. lianJdenii and am 

inclined to doubt his conclusion. The one shown on 

his Plate 38 is said to be " probably a young or abnormal state of this species, as it occurs with the ordinary 

trilobate form." This leaf is certainly platanoid in 

appe~rance, but it is very different from P. haydenii, 

especIally as regards the nervation at the base of the 

leaf. Without a series connecting this with the normal 

form it seems t.o me to be very doubtful if they should 

be placed together. -

The small leaf shown in Newberry's Plate 56 , FiO"- b 

ure 3, is also platanoid in appearance and may well 

enough haye been the immature leaf of one of the 

large-leaved species, but its association with Platan'UJ8 

hrcydenii seems to me questionable. It is 3-lobed, with 

a large central lobe and smaller lateral lobes, in this 

respect agreeing with P. ha:ydendi. The basal nervation consists of a fairly strong Inidrib and two lighter 
80 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

lateral ribs -which arise at some distance above the 

base of the blade and end in the lateral lobes. In 

this feature this leaf agrees with Platanus 1YlflJnoldsii, 

which has the basal ribs arising well above the base 

of the blade, and differs from P. hGI!Jdenii, in which 

the basal ribs ariseC'at the top of the petiole, with the 

midrib. -

There are sev~ral described and figured Fort Union 

species _ that semll to be identical with the small leaf 

referred by Newberry, as indicated above, to Platanus haydenii. Populus nervosa elongata Newberry,t6 

from the Yellowstone River at or near the type locality 

-for Plata11JU8 lwydemi, is smaller than the leaf of 

Platamus haydenii but does not appear to differ essentially. The form identified by Ward 17 as Plata'nus 

guillel17we Goppert is even more like Platanus hay-

'denii than the Populus just mentioned. Ward described two other species that strongly suggest this 

small leaf referred to Platanus hayd~nii. These are 

Gre1.()iopsis vibwrnifolia 18 and G. populifolia,19 both, 

like the specimen identified as Platanus guillell1we, 

,from the lower Yellowstone River near the locality 

that supplied P. haydenii. When the flora of the 

Fort Union formation comes to be fully worked up 

it may be possible to arrive at a definite conclusion 

regarding the status of the leaves mentioned above, 

but until that time it is perhaps best to leave them as 

fixed by the authors quoted. 

Platar1JU8 haydenii has been reported from a number of localities in Wyoming and Colorado, in beds 

then supposed to be of Laramie age, as well as in certain overlying beds. Thus in the United States National Museum collection I find a si~gle example 

(No. 912) recorded as having come from Evanston, 

'''yo. This appears to be the same that was alluded to 

by. Lesquereux 20 as Platanus nobilis Newberry, as it 

agrees perfectly with the description and moreover 

is the only large Platanus leaf now in the Museum 

collection from this place. This specimen is of much 

importance, for if authenticated it would be an instance of P. IJwbilis found in beds not now recognized 

as Fort Union; but doubt attaches to it, as it was 

referred by Lesquereux in his later work to P. haydenii. An examination of this specimen shows it to 

be a mere fragment from the middle of a large leaf, 

and it is without a trace of margin or any character 

by which it can be positively identified. It is without 

16 Newberry, J. S., The later extinct floras of North America: U. S. 

Geol. Survey Mon. 35, P. 49, pl. 28, fig. 1, 1898. 

17 Ward, L. Ii'. , Types of the Laramie flora: U. S. GeoI. Survey Bull. 

37, p. 37, pI. 20, fig. 1, 1887. 

18 Idem, p. 89, pI. 40, fig. 2. 

10 Idem, p. 90, pI. 40, figs. 3-5. 

20 Lesquereux, Leo, U. S. Geol. and Geog. Survey Terr. Ann. Rept. 

for 1871, p. 295, 1872. 

doubt a piece of a large PlatarIJU8 leaf, but beyond 

this it is absolutely impossible to go with certainty. 

It might have belonged as well to any large-leaved 

species, and this uncertainty is well shown by the fact 

that it was first referred to P. nob'ilis and later to P. 

ltaydenii. 

There is a specimen in the National Museum (No. 

914) from Sand Creek, Colo., that was referred by 

Lesquereux to Platarnus l~{JJlJdenii. It is a fragment 

showing the lower left-hand side of a medium-sized 

leaf. The margin was not exposed at first, but by 

removing some of the matrix it could be made out 

with difficulty. The specimen has large simple teeth 

and is probably correctly named, though more material will be needed for confirming this identification. 

The Museum collection contains a number of speciniens of P. haydenii from the Denver formation at 

Golden, Colo. One of these (No. 242) is the original 

of ,'plate 25, Figure 6, of the report on the Tertiary 

flora. The others (Nos. 843, 909) are broken leaves, 

one of them of larger size, but they do not have the 

margin preserved. 

In the collections from the Denver formation 

studied by Lesquereux 21 for the Museum of Comparative Z~ology of Harvard College no less than 16 

specimens were found. They seem to be the young 

form described above, for the most perfect example 

was compared by Lesquereux with Populus nervosa 

Newberry, the variety of which has already been 

shown to be probably identical with the young leaf 

of PlatantttS haydenii. 

The recent large collections from the Denver formation at Golden, Colo., also include a number of more 

oJ' less perfect leaves that I have referred to P. haydenii, but these lea yes approach P. raynoldsii Newberry so closely in size and nervation as to be separated 

with difficulty. This collection contains a great many 

large leaves of PlataniU8, but they are so fragmentary 

that it is next to impossible to differentiate them. 

According to the catalogue of the National Museum 

there should be specimens of Plat{Jffltus haydenii from 

Bridger Pass, Wyo., the Raton Mountains, N. Mex., 

the vicinity of Colorado Springs, Colo., and elsewhere, 

but-they can not now be found .. 

Occurrence: Fort Union formation (types), YellClwstone River near its mouth, Montana; now known 

to be abundant and widely distributed in the Fort 

Union. Denver formation, Golden and Sand Creek, 

about 12 miles east of Denver, Colo. Lawson arkose, 

lower part (in beds equivalent to Arapahoe formation), Sedalia, Colo. 

21 Lesquereux, Leo, Harvard Coll. Mus. Compo Zoology Bull., vol. 16, 

p. 48, 1888. 
SYSTEMATIC PALEOBOTANY 81 

Platanus marginata (Lesquereux) Heel' 

Plate 36, Figures 2, 3 

"ibul'll11m '/IIa'rgina,t1.tnt Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1872, p. 395, 1873; idem for 

1873, pp. 382, 401, 1874; idem for 1874, p. 306, 1876; 

idem for 1876, p. 510, 1878; U. S. Geol. and Geog·. SurYey Terr. Bull., vol. 1, p. 380, 1875; The Tertiary flora, 

p. 223, pI. 38, figs. 1, 4 [not figs. 2; 3, 5], 1878; Harvard 

ColI. Mus. Compo Zoology Bull., Y01. 16, p. 51, 1888. 

Knowlton, GeoI. Soc. America Bull., vol. 8, p. 145, 1897; 

Jom·. Geology, vol. 19, p. 361, 1911. 

Plata11tt8 m(L1'gilltata (Lesquereux) Heel', Flora fossilis arctica, 

vol. 7, p. 97, pI. 98, figs. 3-5; pI. 99, figs. 2, 3; pI. 101, 

fig. 5, 1883. 

JankO, Engler's bot. Jahrb., yol. 11, p. 454, 1889. 

Cross, Colorado Sci. Soc. Proc., vol. 4, p. 211 [25 of reprint], 1892. 

This species, under the name Vibu'rnum 1narginatw'171J, was first described fro111 abundant material collected in 1872 at Black Buttes, Wyo. Of the great 

number of specimens present Lesquereux was inclined 

to refer a considerable variety to this species, writing 

as follows: 

So different, indeed, fire some of these leaves that but for 

the permanent character of their nervation it would be impossible to consider them as representing the same species. ' 

The small leaves are about 5 centimeters long, half as broad 

in the upper part, or above the middle, tapering downward 

to the petiole. The largest are 12 to 14 centimeters long, 

fully as broad below the middle, abruptly contracted to the 

petiole, rounded upward, or often merely truncate to a short 

pOint entered by the median nerve. 

Lesquereux compared these leaves especially with 

such living species as VibwrnU1n pubescens Pursh, V. 

dentat'U/l11) Linne, and V. lantanoides Michaux. 

In later publications additional material from the 

type locality was mentioned by Lesquereux, and he 

also reported this species as present at Point of Rocks, 

Wyo., and Golden, Colo. All but one of the figured 

types of ViburnU1n 1narginat'l.flln are present in the 

United States N ational ~Iuseum, but there are no 

examples preserved from either of these additional 

localities, nor apparently have any been subsequently 

obtained. . 

In 1883, when Heer studied and described the Tertiary flora of Greenland, he found what he presumed 

to be the sa.me as the V ibu1·11'1.il7n 17UJJrginatwm of Lesquereux, but after fully considering the case he decided that they should be properly referred to the 

genus PlOJtanus, and he consequently called them 

Platanus 1narginata. This view is sustained by 

J ank6,22 who is an authority on the genus Platanus, 

Loth living and fossil. 

I have studied carefully the material, both figured 

and unfigured, in the National Museum, and while I 

2: Jllnl<O, H., Engler's bot. Jllhl'b., ,01. 11, p. 454, 1889. 

am prepared to' admit the correctness of referring a 

portion of them to Platanus, I do not t.hink that all 

belong to a single species or even to the same genus. 

'Vhat I assume to be Platmnus are well shown in Figures 1 and 4 of Plate 38 in the report on the Tertiary 

flora,' and these are evidently the fornls that were 

principally in the minds of Heel' and Janko when they 

transferred Lesquereux's species fro111 V ibuTnwn. 

The type of leaf that I would ascribe to Plalanu8 

1narginata may be described as follows: Leaf of 

medium or small size (12 to 15 centimeters broad and 

about the same in length), broadly ovate or obovate, 

obtusely wedge-shaped at the base, not divided into 

lobes but rather abruptly rounded and entire above 

or with a short acuminate point; Inargin entire below, 

becoming sparsely denticulate above, the teeth small, 

acute; petiole relatively long and strong; primary 

nerves three, strong, the central or midrib straight, 

ending in the tip or apex, with about four pairs of 

strong, subopposite secondaries which usually fork 

unce or twice, the branches passing to the marginal 

teeth; lateral ribs arising at the very base of the blade 

and at the same' angle as the secondaries on the midrib, each with four or five secondary branches on tlie 

lower or outer side, the lowest of which bears numerous camptodrome branches, while the upper branches 

nre once or twice forked and craspedodrome, ending 

in marginal teeth; nervilles numerous, strong, both 

broken and percurrent. 

These leaves from the type locality at Black Buttes 

are usually very perfectly preserved, the matrix being 

a light-yellowish fine-grained sandstone on which the 

ontline and nervation of the leaves stand out in black 

lines. 

The other type of leaf that was included by Lesquereux in his V iburnWln trnarginatu17~ is shown in 

Plate 37, Figure 1, and Plate 38, Figures 2 and 3, of 

the report on the Tertiary flora. It is nearly elliptical 

(II' slightly elliptical-obovate, being about the same at 

both base and apex, with the margin entire below and 

slightly toothed above. The nervation, it seems to me, 

differs essentially in not being strongly 3-ribbed from 

'the base of the blade but having a strong midrib with 

some five or six pairs of rather light secondaries. The 

branches from the lower secondaries are camptodrome 

in the lower part of the blade and craspedodrome in 

the upper part, a feature in which they agree with the 

leaves of Platrunus 7lw,1'gi1wta. In my opinion these 

leaves should be given a new specific name but retained in the genus Viburnwrn. 

Vibu1"nu1n 1nm'ginat1.J(m wa.s reported by Lesquereux 

from Point of Rocks, Wyo., but as no specimens can 

now be found to substantiate this report it is impossible to say whether they belong to what is now referred to Platan118 or to the leaves· it is proposed to 
82 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

retain in Viburnum. This locality must therefore be 

dropped, at least for the present. 

'Platanus ma1'ginata has been identified in Carbon 

County, Wyo., where it occurs in the" Lower Laramie" [Medicine Bow formation], believed by me 

to be the stratigraphic equivalent of the Larami~, of 

the Denver Basin. The specimens are fairly well pre- . 

served and are probably correctly determined. 

This species has also be found in the Lance formation in the Cheyenne Indian Reservation of South 

Dakota. 

A specimen with its counterpart from the Dawson 

arkose. at Mosby, Colo., is here figured. (PI. 36, figs. 

2, 3.) It is unfortunately not very perfect, but enough 

is present to make its identification certain. It appears 

indistinguishable from Figure lin Plate 38 of the 

report on the Tertiary flora, though it is slightly 

smaller. The counterpart shows the perfect petiole, 

which is about 2.5 centimet~rs long and shows the 

enlargement at the end under which the bud. for the 

following year's growth was concealed. This is a wellknown character of Platanus leaves, and this leaf confirms the reference to this genus .. 

. Occurrence: Black Buttes coal group (type$), Black 

Buttes, Wyo. Dawson arkose, dump of Mosby coal 

mine, Mosby, Colo., collected by F. H. Knowlton and 

G. B. Richardson, 1910. Lance formation, Cheyenne 

Indian Reservation, S. Dak. "Lower Laramie" [Medicine Bow formation], Carbon'" County , Wyo. 

Platanus platanoides (Lesquereux) Knowlton 

Plate 36, Figure 5 

Platanus platanoides (Lesquereux) Knowlton, U. S. Geo1. Survey Bull. 152, p. 171, 1898; Jour. Geology, vol. 19, p. 370, 

1911; U. S. GeoI. Survey Prof. Paper 101, p. 323, pI. 95, 

fig. 4, 1917 [1918]. 

Viburnum platanoides Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1874, p. 314, 1876; The Tertiary 

fiora, p. 224, pI. 38, figs. 8, 9, 1878 .. 

The fragmentary specimen from the Dawson arkose 

on Jimmy Camp Creek here figured appears to be 

identical with a specimen from the Raton formation 23. 

figured under this name. It is a little more wedgeshaped at the base than the type specimens but does 

not differ otherwise to a marked degree. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 1 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected 

by F. H. Knowlton, July, 1910. Black Buttes coal 

group, Black Buttes, Wyo. Lance (~ ) formation, 

sec. 20, T. 14 N., R. 10 E., S. Dak. Laramie (~) 

formation, Erie, Colo.' Raton formation, Rouse, Colo. 

28 Knowlton, F. H., U. S. Geol. Survey Prof. Paper 101, p. 323, 

pI. 95, fig. 4, 1917 [1918]. 

Platanus colol'adensis Knowlton, n. sp. 

Plate 37, Figures 1, 2 

Leaves of medium size, evidently coriaceous and 

firm, ovate, lateral lobes slight if any; rather obtusely wedge-shaped at the base and obtusely acuminate at the apex.; margin entire below, undulatetoothed above, the teeth low and separated by very 

shallow sinuses; midrib of moderate strength, 

straight; nervation subpalmate-that is, the lowest 

pair of strong secondaries arise at an angle of about 

450 well above the base of the blade, thence running 

to the margin, though probably not into lobes, each 

provided on the lower or outer side with some five or 

six tertiary branches, which curve slightly upward 

and are apparently camptodrome; below the strong 

pair are one or two pairs of much thinner secondaries, 

one of which is· usually provided with several thin 

camptodrome tertiary branches; above the strong pair 

of secondaries there are six or seven pairs of wellmarked secondaries that run nearly straight to the 

margin, which they enter at the low teeth, the lower 

ones with one or two forks, which also enter the margin; nervilles numerous, strong and deeply impressed, 

usually curved but unbroken. 

This species is represented by several examples, 

of which two 'of the best are figured. They are of 

medium size, the length being about 15' centimeters 

and the width about 10 centimeters; the petiole is not 

preserved. It is probable that the leaves are simply 

ovate, though there may have been slight lateral 

lobes. The lower portion of the blade is perfectly 

entire, but from a point near the middle it becomes 

undulate-toothed. The nervation is strongly marked, 

as may be seen from the figures. 

The most perfect of these 'leaves, shown in Plate 37, 

Figure 1, was found in the collection of the United 

States National Museum under No. 1057, and in the 

catalogue Lesquereux has recorded the fact that it was 

a new species. This specimen was collected in 1875 

by E. A. Barber, who was at that time associated with 

the' Hayden survey. It came from ~Iiddle Park, 

Colo., but is without specific locality. It is, however, 

identical with leaves since obtained by Whitman Cross 

on a high terrace ridge between the forks of Kinney 

Creek, a tributary of the Colorado River east of Hot 

Sulphur Springs, in Middle Park. The matrix is also 

slmilar in the two examples, and it is fair to assume 

that the Barber specimen came from practically the 

same place as the others. 

It is with some hesit~tion that I venture to describe 

this as a new species, yet it does not appear to be very 

closely related to previously named species. In size 

and shape it perhaps approximates most closely what 
SYSTEMATIO PALEOBOTANY 83 

has been called a.young leaf of Platanus haydenii Newberry,24 but the margin and basal nervation are somewhat different. If there were .slight lateral lobes in 

the leaves under consideration, a p'oint which, as above 

stated, it is impossible to demonstrate, this species 

might' come near to several species such as P. heerii 

or P. newbe?"'l'Jja;na, but these may be dismissed without 

further consideration. 

Occurrence: Middle Park formation, on high ter~ 

race diyicl€\ between forks of IGnney Creek, a tributary 

of Grand [Colorado] River on the north, east of Hot. 

Sulphur Springs, ~1iddle Park, Grand CO'unty, Colo., 

collected by vVhitman Cross, October 6, 1891; also 

specimen collected, by E. A. Barber in Middle Park 

but without specific locality .. 

, Platanus sp.? 

PIn te 38, Figure 2 

In the collection from Mosby, Colo., was the speci~ 

men here figured. It is only a fragment of the lower 

side of a leaf with a strong nervation and sharply 

toothed margin. It is a Platanws of the type of P. 

guillel'l1Ute Goppert or P. a<Je1'oides Goppert, as figured, 

for instance, by Lesquereux in the report on the Ter~ 

tiary flora, Plate 25, Figures 1-5. As the origin of 

the strong lateral ribs is not shown in this specimen 

it is not possible to distinguish between these two species, though if anything it appears to approach more 

closely P. gwillelmae. This specimen is, of course: 

of comparatively little value except as indicating the 

presence of this type of leaf in the formation under 

consideration. 

Occurrence: Dawson arkose, dump of ~10sl)y coal 

mine, Mosby, Colo., collected by F. H. I{nowlton and 

G. B. Richardson, 1910. 

Order THYMELEALES 

Family LA URACEAE 

LuUl'us pl'imigenia Unger 

Plnt'c 38, Figures 3, 5, 6 

La1l1"l18 1)1'im .. igcni(£ Unger, Genera et species plantarum fossili UIll, p. 423, 1850. 

Heel', Acad, Nat. Sci. Philadelphia Proc., vol. 10, p. 265, 

1858. 

Lesquereux, U. S, Geol. and Geog. Survey Terr, Ann. Rept. 

for 1872, p. 406, 1873; idem for 1874, p. 385" 1876; idem 

for 1876, p. 510, 1878; The Tertiary fiora, p. 214, pI. 36, 

figs. 5, 6, 8, 1878; Harvard ColI. Mus. Compo Zoology 

Bull., vol. 16, p. 50, 1888; U. S. Nat. Mus. Proc., vol. 11, 

p. 36, 1888. 

Ward, U. S. Geol. Survey 'Sixth Ann. Rept., p. 553, pI. 46, 

figs. 8-10, 1885; U. S. Geol. Survey Bull. 37, p. 47, pI. 23, 

figs. 8-10, 1887. 

24 Newbel"l"Y, J. S., U. S. Geol. Survey Mon. 35, p. 103, pI. 38, 1898. 

Knowlton, U. S. Geo1. Survey MOll. 32, pt. 2, p. 722, pI. 91, 

figs. 4, 5, 1899; Geo1. Soc. America Bull., vol. 8, p. 154, 

1897. 

Hollick, Louisiana Geo1. Survey Rept. for 1899, Special 

Uept. 5, p. 284, pI. 41, figs. 1, 2, 1899. 

Lauru8 obovGltaf Weber. Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1872, p. 399, 1874. 

SaZim media? Heer. Knowlton, Geo1. Soc. America Bull., vol. 

8, p. 141, 1897. 

LauT1J;8 pr'i'l'flfigenia is a cOmllon and widely distributed species in the European Tertiary, being appar~ 

ently most abundant in the upper Eocene, Oligocene, 

and 10weF Miocene. Its presence in North American 

strata' appears to have been first noted by Heel' in 

1858, when he reported Qn a number of drawings of 

fossil plants from the Dakota sandstone of Nebraska, 

snbmitted to him by Meek and Hayden.25 This de~ 

termination was not accepted by either Lesquereux 

or Newberry, for neither mentioned it, though many 

years later Lesquereux procured a specimen from the 

Da~ota sandstone of Pipe Creek, Cloud County, Kans., 

that he at first inclined to regard as a variety of L. 

primigeniaJ but later decided to be more closely allied 

to Fi(YlJ)s.26 

In point of tiine the next mention of Laurus rril1tigenia in North America is by Lesquereux,27 who reported it from Spring Canyon (now Rocky Canyon), 

Mont., but the specimens upon which this identification rests were later, according to a statement in the 

report on the Tertiary flora (p. 214), discarded as 

unworthy of notice. This species was not detected 

when I studied the recent collections from the Boze~ 

-man coal field and may well be dropped from this 

locality. 

In 1874 Lesquereux 28 reported with question the 

presence of LauTUs obovata Weber in . the so~called 

saurian bed at Black Buttes, Wyo., but on the page of 

the report on the Tertiary flora above cited he abandoned this also as being based on too imperfect matel'j al for certain identification. ' 

Apparently the first authentic material referred to 

Laurus pri'l11igenia came from Evanston, Wyo. It 

was described and figured by Lesquereux 29 in 187.8, 

but none of this material, figured or otherwise, can now 

be found in the collections of the United States National Museum, where presumably it should be located. 

Dependence must therefore be placed entirely upon 

Lesquereux's figures. The specimens are said to have 

been found in the same beds with Larurus somalis 

25 Meek, F. B., and Hayden, F. V., Acad. Nat. Sci. Philadelphia Proc., 

vol. 10, pp. 257, 265, 1858. 

~6 U. S. Geol. Survey Mon. 17, p. 85, 1895. 

Zl Lesquereux, Leo, U. S. Geol.. and Geog. Survey Terr. Ann. Rept. 

for 1872, p. 406, 1873. 

28 Idem for 1874, P. 385, 1876. 

l!9 Lesquereux, Leo, The Tertiary flora, p. 214, pI. 36, figs. 5, 6, 8, 

1878. 
84 .FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Lesquel'eux and are certainly very close to it, ,differing, according to Lesquereux, in being "narrowly 

lanceolate, nlore distinctly acuminate, and narrowly 

cuneate," all comparative or relative characters of little weight. Although Laurus pri1nigenia is a very 

polymorphDus species, to judge from the forms that 

have been referred to it by various European students, 

it is hard to reconcile Lesquereux's figures with any 

of them. Thus Figure 6 of his Plate 36 is quite clearly 

a LaU1"lt8, as it has the peculiar arching camptodrome 

secondaries, but Figures 5 and 8 of the same plate may 

not be the same-that is, if they have been. correctly 

drawn. Figure 6 approaches certain of the narrow 

leaves of the European for~ such for instance as those 

figured by. Unger 80 from Sotzka, but the other figures 

do not, and I should incline to exclude them from this 

species. They ~ay be lauraceous, but I doubt very 

much if they belong to Laurus primigenia. 

YVard 81 figured three specimens that he referred to 

L. pri1nigenia. Of these the one from Carbon, Wyo. 

(fig. 8), may be correctly referred on the basis of 

Lesquereux's figures, but the others that came from 

I'oint of Rocks, Wyo., almost certainly belong to 

another species. Indeed, with Ettingshausen,82 I must 

regard Figure 9 at least as more like Laurus ocateaefolia Ettingshausen and quite unlike typical forms of 

L. primigeniaJ. 

In 1888 Lesquereux 88 reported the presence of 

La'UlJ'u.'3 prrimligewidJ in beds of the Denver formation 

nt Golden, Colo., and one of the best preserved is here 

figured. I 'have not been able to detect it among the 

large recent collections from .Golden. In the same 

year Lesquereux 84 also mentioned this species as being 

present. in a small collection from Selma, Cherokee 

County, Tex., but this is extremely doubtful. 

The next in order are two fragmentary examples 

from the Cretaceous of Staten Island more doubtfully 

referred to L. pYf'iJmigenria by Hollick.811 These leaves 

can not possibly belong to this species, and it is even 

doubtful if they belong to the genus Laurus. Nearly 

the same stricture may be passed on the two leaves 

from the Wilcox formation at Coushatta, La., referred 

without question by Hollick 86 to L. primigeniaJ; in 

fact, Berry,37 who has recently revised this flora, has 

relegated the two figures to separate and distinct 

species-that is, Hollick's Figure 1 is referred to 

OreodapAne mississippiensis Berry, and his Figure 2 

to Nectandra pseudocoriacea Berry. 

In my Flora of the Yellowstone National Park 38 

I figured as doubtfully referable to Laurus pri1idgenia 

two rather fragmentary examples from rocks of Fort 

Union age at a point on the Yellowstone River just below the mouth of Elk Creek. This assignment was 

. based largely on their resemblance to the leaf from 

Carbon, Wyo., that was referred by Ward to this 

species, and as that has been accepted it is probable 

that the Yellowstone Park specimens should also be 

accepted as referable to L. primigenia. 

In 1900 89 . I admitted Laurus primigenia as present 

at Point of Rocks, Wyo., but as set forth above, it 

is now believed that the specimen was incorrectly 

identified. 

In the material obtained near the Douglas coal 

mine, Sedalia, Colo., I find the single example here 

figured, which is referred with some question to 

Laurus prilmigenia. It is relatively longer and narrower than the figures given by Lesquereux, but the 

shape and nervation are identical. C 

In 1905 A. C. Veatch, of the United States Geological Survey, made some small collections from the 

Evanston area, and at mine No.7, Almy, he found 

several leaves that are identical with Lesquereux's 

figures of Laurus primigenia. Almy is a small mining camp and, is probably very near, perhaps identical 

with, the locality that afforded Lesquereux's specimens. 

The material from the Mosby coal mine, east of 

Colorado Springs, Colo., includes several leaves that 

agree well enough with the figures of Lauru8 prilmige11lia. 

Occurrence: Dawson arkose, Jimmy Camp Creek, 

near Richfield Springs ranch house, 9 miles east of 

Colorado Springs, Colo., collected by F. H. Knowlton, 

1910. Dawson arkose, lower part (in beds equivalent 

to Arapahoe formation), 3,000 feet east of old Douglas 

coal mine, near Sedalia, Colo., collected by Arthur 

Lakes, 1895. Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes; speci-

----------------------- I mens in Museum of Comparative Zoology, Cambridge, 30 Unger, Franz, Fossile Flora von Sotzka, pI. 19, figs. 1-4, 1851. . 31 War, d L S G S S Mass., No. 1466. Evanston formation, mIne No.7, . F., U. . eol. urvey ixth Ann. Rept., p. 553, pI. 46, 

figs. 8-10, 1886; U. S. Geol. Survey BUll. 37, p. 47, pl. 23, figs. 8-10, near Almy, Evanston, Wyo. Hanna formation, Car1887. 32 cr. Ward, L. F., op. cit. (Bull. 37), p. 48. bon, Wyo. Beds of Fort Union age, Yellowstone a3 Lesquereux, Leo, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, River near mouth of Elk Creek, Yellowstone ~'l.tional 

p. 50, 1888. Park, collected by F. H. I.(nowlton, 1888. 84 Lesquereux, Leo, U. S. Nat. Mus. Proc., vol. 11, p. 36, 1888. 

;<:; Hollkk, Arthur, New York Acad. Sci. Trans., vol. 12, p. 33, pI. 2, 

fig. 20; pI. 3, fig. 3, 1892. 

~6 Hollick, Arthur, Louisiana Geol. Survey Rept. for 1899, SpeCial 

Rept. 5, p. 284, pl. 41, figs. 1. 2. 1899. 

~7 Berry, E. W., U. S.Geol. Survey Prof. Paper 91, pp. 303, 311. 1916. 

as Knowlton, F. H., U. S. Geol. Survey Mon. 32, pt. 2, p. 7::!2. pI. 91, 

figs. 4, 5, 1899. 

89 Knowlton, F. H., Flora of the Montana formation: U. S. Geol. 

Survey BUll. 163. P. 58. 1900. 
SYSTEMATIC PALEOBOTANY 85 

Laurus socialis Lesquereux 

Plate 38, Figme 4 

Laur'u..s socia-lis LeSquerel;x, The Tertiary flora, p. 213, pI. 

36, flgs. 1-4, 7, 1878; Harvard ColI. Mus. Compo Zoology Bull., vol. 16, p. 50, 1888. 

Knowlton, U. S. Geol. Survey BUll. 100, p. 56, 1893; U. S. 

Geol. Survey Prof. Paper 101, p. 317, pI. 91, flg. 0, 1917 

[1918]. 

This species is closely allied to the American specimens identified by Lesquereux as LauTUS pri'lnigenia 

Unger: as Lesquereux stated, and was even regarded 

by him at one time as identical with it. Saporta, 

however, to whom figures had been submitted, considered them distinct, the present species "having 

larger leaves, more obtuse at the base, less distinctly. 

ncumina.te than any of those referred to L. pri'lnigenia." The nervation is of the sam~ character in 

both. 

Most of the type specimens of Larurus socialis are 

preserved in the United States National Museum (Nos. 

306, 307, 308). They were found at the same locality 

and some of them even on the same pieces of matrix 

as those of L. p'l~i'lnigenia, but as no additional examples haye come from that locality we must depend 

upon the figured types. 

Lm.lfI"US socialis was reported by Lesquereux 40 from 

the Denver formation at Golden, Colo., but I have not 

seen these specimens, nor have I been able to detect any 

specimens in the recent collections from thi~ locality. 

LaJU,11U8 sO(Jtarlis was also reported by Lesquereux 41 

from Lassen County, Calif., and this was apparently 

the species largely relied upon to prove the Eocene age 

of the beds whence tJley came. Only three broken 

examples were procured from this locality, all. of 

which are preserved in the United States National 

:Museum. However, they do not seem to me to belong 

to Laruh"l.IS socialis. They are of about the same size 

and sha.pe as leaves of this species, but the nervation 

appears quite different. The finer nervation is well 

preseryed, and although like that of L. socialis, it is 

also as neal' to some of the species of Pe7'sea from California: such as P. dille'l~ Lesquereux.42 One of the 

Lassen County leaves is rounded exactly as in P. dilleri. 

These leaves are also much like Pe7'sea punotulata 

Lesquereux,48 from Corral Hollow, Alameda County, 

Calif. 

Lau?'lts socialis was also reported by Lesquereux 44 

from Ca.mpbell's quarry, Cross Lakes, La. Only one 

leaf wa.s found at this -locality, a small example (U. S. 

Nat. Mus. No. 2501) which lacks both base and apex. 

~o Lc-squereux, Leo, HUl'vurd Coli. Mus. Compo Zoology BulL, VOl. 16, 

p. 00, 1888. 

41 U. S. Nat. Mus. Proc., vol. 11, p. 29, 1888. 

U Idem, P. 27, pI. 13, figs, 2-4. 

43 Idem, p. 26, pI. 14, fig. 1. 

u Id(>m, p. 24. 

This specimen has been studied by Berry,45 who rerers 

it to his j1{ espi~odaph'ne pseudoglauca, a· species that 

has been found to be abundant and widely dis~ributcd 

in beds of· Wilcox age in Louisiana, Mississippi, and 

Kentucky. 

In the Livingston formation near Chestnuts, in the 

Bozeman coal field, Montana, I found 46 a number of 

examples that were referred to Laurus socialis. They 

are fairly well preserved and appear to be correctly 

referred. 

LauTUs socialis was identified 47 in beds of the Raton 

formation at several localities in southern Colorado 

and northern New Mexico. 

Occurrence: Evanston formation [types], Evanston, Wyo~ Denver ( n formation, Golden, Colo. Dawson arkose, Mosby coal mine, Mosby, Colo., collected 

by F~ H. Knowlton and G. B. Richardson, 1910. Livingst.on formation, Chestnuts, near Bozeman, Mont. 

Raton formation, Delagua and near Aguilar, Colo., 

Hnd York Canyon, N. Mex., collected by G. B. Richardson. 

Lam'us lanceolata Knowlton, n. sp. 

Plate 38, Figure 9 

Leaf narrowly lanceolate and slightly falcate,' 

slightly inequilateral, with about equally acuminate 

flpex and base; petiole missing. Midvein stout, 

slightly flexuous, prominent on' the under side of the 

leaf. Secondaries stout, six or seven subopposite to 

r.lternate pairs, prominent, diverging from the midvein at acute angles, sweeping upward in long ascending curves, eventually camptodrome. Tertiaries thin, 

mostly immersed in the coriaceous leaf substance. 

Length between 9 and 10 centimeters; maximum width 

1.4 centimeters. 

This species resembles the Wilcox species N ectandJra 

l)Seudocol"iacea Berry 48 in size and form but is somewhat less acuminate and has fewer and more ascending 

secondaries. 

Occurrence: Dawson arkose; 

Persea brossiana Lesquereux 

Pm'sea b1'ossiana Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1873, p. 407, 1874. 

Law'1f,S brossiana (Lesquereux) Lesquereux, The Tertiary 

flora,. p. 216, pI. 36, fig. 9, 1878. 

The original and so far as known the only specimen 

of this very fine species is preserved in the collections 

of the United States National Museum (No. 309) and 

has 'been well described and figured by Lesquereux. 

46 Berry, E. W., U. S. ,Geol. Survey Prof. Paper 91, p. 306, pI. 80, 

fig. 4, 1916. 

46 Knowlton, F. H., U. S. Geol. Survey Bull. 105, p. 56, 1893. 

47 Knowlton, F. H., U. S. Geo1. Survey Prof. Paper 101, p. 317, pI. 91, 

fig. 5, 1917 [1918]. 

~ Berry, E. W., U. S. Geo1. Survey Prof. Paper 91, p. 311, pI. 88, figs. 

1-3, 1916. . 
86 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

This leaf was first placed in the genus Pe1'sea by 

Lesquereux but was later transferred by him to LaJWl"IJJ8, 

as he regarded it as closely related to his La'Ulf'U8 

utahensis. A comparison of the leaf under consideration and the)iving Persea carolinensis Nees, from the 

southeastern United States, shows them to be so similar that it is almost impossible to separate them. 

This leaf is also closely similar to the living Laurus 

canariensis Weber, as Lesquereux pointed out, and this 

fact, together with the presence of undoubted glands 

in the axils of the secondary nerves, was probably what 

induced him to transfer it to Laurus. But as it was 

first called Per'sea and bears so striking a resemblance 

to the living pe'J'sea carolinensis I have thought best to 

retransfer it. 

The glands that appear so prominently in the type 

illustration are certainly present but hardly seem as 

pronounced as .would be supposed from the figure. It 

is the possession of these glands, together with the fact 

that the leaf is much broader and more obtuse than 

most of the leaves of Persea carolinensis., that separates 

the fossil from the living species. 

Occurrence: Middle Park formation, :Mount Bross, 

Middle Park,Grand County, Colo. 

Cinnamomum sezannense Watelet? 

Plate 38, Figures 7, 8; Plate 59, Figure 2 

Oinnatmomum sozarvnen.s6 Watelet, Description des plantes fossiles du bassin de Paris, p. 175, pI. 50, fig. 2, 1866. 

The specimen with its counterpart that is here 

figured I am not able to distinguish from Oinnamomwm seza'nnens'(j, as described and figured by Watelet 

and others from the Eocene of the Paris Basin. It 

is a narrowly lanceolate leaf 10 or 12 centimeters in 

length and a little less than 4 centimeters in width. 

The petiole is retained for a length of about" 12 millimeters and is not complete. The midrib is strong for 

the size of the blade and has two or three secondary 

branches in the upper third of the leaf. The lateral 

ribs are strong, though lighter than the midrib; they 

arise well above the base and run along well inside the 

margin probably nearly to the apex. There is a single 

series of very light tertiary branches or strong nervilles on the outside of the ribs, these forming a series 

of loops" just inside the margin. The nervilles 'are 

numerous, strong, often broken, and at right angles 

to the midrib. 

This leaf was at first ,thought to be referable to 

OinnoYnw?,)LU1n affine Lesquereux,49. but closer inspection discloses a number of differences. Thus O. affine 

is distinctly ovate or ovate-elliptical, whereas the present leaf is proportionately much narrower or more 

to Lesquereux, Leo, The Tertiary flom, p. 219, pI. 37, figs., 1-5, 7, 

1878, 

nearly lanceolate. The primary nervation, including 

both midrib and lateral ribs, is very much stronger in 

the leaf under consideration than in O. affine, and 

there appear to be fewer secondary branches on the 

midrib. It is possible that this is only a narrow, 

strongly veined leaf of O. affine, but I have not seen 

other similar specimens that could be so interpreted. 

On the other hand, I can not se"e any esse~tial characters by which this leaf can be separated from O. 

sezannense as figured by Watelet, Saporta and Marion, 

and others. In spite of this seeming agreement it is 

with some hesitation that a European species is identified as present in the Rocky Mountain region, and the 

reference has been questioned. 

Occurrence: "Dawson arkose, east bank of Jimmy 

Camp Creek 1 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. Knowlton, July, 1910. 

Cinnamomum linifolium Knowlton 

Plate 59, Figure 3 

Oi'l'llnamomum Umifolium Knowlton, U .. S. Geol. Survey Prof. 

Paper 101, p. 319, pI. 88, figs. 3-7, 1917 [1918]. 

This species was found in the Raton formation of 

the Raton Mesa region of Colorado and New Mexico, 

where it appears to have been rather uncommon. It 

was described as follows: 

Leaves firm in texture, from linear to linear-Ianceolate, long 

and narro~vly wedge-shaped at base, apparently rather obtuse 

at apex; nervation triple-ribbed from the very top of the 

petiole, midrib the stronger, straight; lateral ribs more slender, 

ascending nearly or quite to the apex of the blade. 

The single example here figured from the Dawson 

arkose is referred to Oi11lnarmorruum~ linifoli1.(,1n with a 

considerable degree of certainty. It is a small leaf 

only about 4 centimeters long and 1.5 centimeters 

wide, and although proportionately a little broader 

than in the types it ag~ees in its obtuse apex and in 

the ribs arising from the to,p of the petiole. The firm 

nervation is obscure, as it is in the type specimens, 

though there appear to be one or two secondary 

branches on the outside of the lateral ribs and the 

nervilles seem to be numerous, close, and approxi- ' 

mately at right angles to the midrib. 

Oinnamom;um lirvifoliw:m is 'apparently closely related to Oinn{Wn01num oblo'nr/atwn1J. Berry,50 from, the 

Lagrange formation (in beds of Wilcox age) at Puryear, Tenn. The size and shape are practically identical in the two forms, the most notable difference 

being in the point of origin of the lateral ribs. In 

C. linifoliu1n the lateral ribs arise at the top of the 

petiole, but in O. oblongatwm' the ribs originate a short 

distance above the base of the blade. 

00 Berry, E. W., U. S. Geol. Survey Prof. Paper !)1, p, 297, pI. 79, 

fig~. 1, 2, 1916. 
SYSTEMATIC PALEOBOTANY 87 

The leaf here figured from the Dawson arkose is 

also suggestive of one of the small leaves of Oinna'1no'mwm vevrclJ Berry,51 from the Wilcox group at 

Oxford, Miss. It is not quite so broad, however, and 

differs as regards the' origin of the lateral ribs, as it 

does from O. oblonga1tul1u. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. I(nowlton and W. T. Lee, July, 1910. 

Raton formation, near Starkville, Colo., and Vermejo 

Creek, near Dawson, N. Mex. 

Cinnamomum? sp. 

Plate 36, Figure 4 

The material" from Templeton Gap includes a single 

~pecimen-the one here figured-that appears to belong to OinnGII]W1nU1n, but as 'it lacks both base and 

apex it can not be placed with certainty. It is a small 

leaf about 6 centimeters long and 3.5 centimeters wide. 

It is presumably regularly elliptical, with a wedgeshaped base and an obtuse apex. The midrib is rather 

strong, with two or three pairs of secondaries in the 

upper portion. The lateral ribs are but little lighter 

than the midrib and pass up to at least the upper third 

of the length of the blade. Each has several secondHl'y branches 'on the outside. The manner in which 

the lateral ribs join the midrib can not be ascertained. 

The matrix is so coarse grained that the finer nervation is obscured. This leaf may belong to Oinnamomum affine Lesquereux,52 but it is rather narrower than 

,that species usually is. It also suggests some of the 

narrower leaves of Ficus trine1'vis Knowlton and Ficus 

lJ1'aetrinervis Knowlton, but absence of the base makes 

it impossible to determine this point. It seems more 

likely to belong to Oinnanw1num than to Ficus of the 

type of the above-mentioned species. 

Occurrence: Dawson arkose, Templeton Gap, 4 

miles northeast of Colorado Springs, Colo., collected 

, by F. H. I(nowlton, July, 1910. 

Cinnamomum sp. 

Plate 39, Figure 1 

In the material fronl the Denver beds at Golden, 

Colo., I have found a single leaf that appears to be 

n Oi'f1lnanWl1~wn~. Unfortunately it is fragmentary, 

lacking all of the basal portion and most of the sides, 

but it has more of the apical portion preserved, showing that it was extended into a rather sharp-pointed 

apex. So far as can be made out it is similar to what 

was identified by Lesquereux 58 as Oinna1n01nun~ poly'-

&1 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, pI. 79, fig. 3, 1916, 

5~ LCl!'ql1crcl1x, Lco, The Tertinry flol'n, P. 219. pI. 37, figs. 1-5, 7, 

1878. 

~3 Idem, p. 221, pI. 37, fig. 10. 

1norphu1n AI. 'Braun, later sepa.rated as Oinna1nom'l.tm 

ellipticwm I(nowlton,54 but it is too fragmentary to be 

positively identified. This type specimen of O. elli1Jticwm Was said ,by Lesquereux in the report on the 

Tertiary flora to have come from Golden, Colo., and 

in the catalogue it is recorded from the Bozeman coal 

field, lVlontana. An examination of the specimen, 

which is preserved in the United States N at-ional lVluseum (No. 316), proves clearly that it did not 'come 

from Golden but from the Montana locality. 

The specimen under consideration might bp. l'eferi'ed 

to what has been identified with OinnClJ7n01nU1]~ sezannense? ",Vatelet, or possibly with Oinnamw1]1wln sp.) 

but as it is so broken that its full cha.racter can not be 

made .out, it should be held apart from other forms 

until more nearly perfect material can be procured. 

Occurrence: Denver formation, Golden, Colo., associated with Diplazi'Lt1n c-ro8sii. 

Family MAGNOLIACEAE 

Magnolia magnifolia Knowlton 

Magnolia magnifoUa Knowlton, U. S. Geol. Survey Prof. Paper 

101., p. 311, pI. 84, 1917 [1918). 

JJ:la,gnolia tent(''inervis Lesquereux, The Tertiary flora, p. 249. 

pI. 44, figs. 5, 6, 1878. 

Magnolia 1nagnifolia; as at present understood is 

perhaps the most abundant and widely distributed 

species thus far observed in the Raton formation. It 

is apparently the largest-leaved fossil magnolia yet 

ci.escribed from American rocks, the maximum length 

observed peing approximately 30 centimeters and the 

width 10 or 12 centimeters. It may be known by its 

large size, rather obtuse apex, wedge-shaped base, and 

especially by the irregular spacing of the secondaries 

near the middle of the leaf. 

Magnolia m,agnifolia was established in large part 

to take the place of Magnolia tenuinervi8 of Lesquereux, which is found to be so complicated as regards specific limitation and vertical range that it 

was thought best to abandon the use of the name in 

large part and start afresh. The history of Magnolia te11fUineTVis, both nomenclatorial and distribntional, has been set forth at length in Professional 

Paper ,101 and need not be repeated in full at this 

time. The species was named, though not adequately 

described, by Lesquereux in 1868 from material, said 

. to have been obtained in "lignitic beds near Golden 

City, Colo.," and hence presumably in the true Laramie formation, as the Denver formation has not 

been found coal bearing at this locality. For more 

than 10 years after it was named it reInained a nomen 

nudum, and even where it was described and figured 

it; was based on such a heterogeneous a.ssembla.ge of 

'54 Knon'lton, F. H., U. S. Geol. Survey Bull. 152, p. 68; 1898, 
88 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

forms that are obviously not con specific and may not 

all be congeneric, from unknown or doubtful localities, that it seemed best to abandon the current accept-' 

ance of the" species." The difficulties into which this 

miscellaneous assemblage has led subsequent writers 

are s,hown by the fact that it was supposed to have a 

vertical range from the Mesaverde formation to the 

Denver formation, or even later. But, as the rules 

of nomenclature do not permit the abandonment of 

a name so long as it can not be clearly referred to 

another earlier species, it is proposed hereafter to 

restrict the name Magnolia tenuinervis to Figure 6, 

Plate 45, of the report on the Tertiary flora, for this, so 

far as can be made out, is the figure of the specimen 

from" Golden City, Colo.," upo~ which the" species" 

was based. It is a mere fragment of the upper portion of a lea,f that can not be positively identified with 

anything and may not be a Magnolia. The type specimen itself is lost, and it can not be determined 

whether it belon:gs to the Laramie or the Denver 

formation. 

Although several of the forms figured by Lesquereux 

as Magnolia tenuilnervis are said to have come from 

the Denver formation at Golden, Colo., only one-that 

of Figure 2, Plate 45-is known posItively to have 

come from Golden. This is No. 367 of the National 

Museum collection and is preserved on the characteristic andesitic matrix. It is fragmentary, lacking both 

base and apex and all of the margin on one side. It 

was ,probably about 15 centimeters long and less than 

6 centimeters wide, thus being smaller than most of 

the examples frOln the Raton Mesa region.' Several 

other specimens from Golden obtained in recent years 

are also ,smaller than the average of those from the 

Raton formation, but otherwise they do not differ 

essentially. The originals of 'Figures 3 and 6 of 

Lesquereux's Plate 44 are with little doubt from Black 

Buttes, Wyo., and not from Golden, Colo. 

There is a specimen in the duplicate collection of 

the National Museum (No. 926) from Golden, Colo., 

that was questionably referred by Lesquereux to Magnolia lesle1J(JJlUJ; Lesquereux. It is only a fragment 

from the middle of a leaf that must have approached 

in size the large leaves of M. magnifolia from the 

Raton formation and should probably be referred to 

this ,species. 

Occurrence: Denver formation, andesitic beds at 

Golden, Colo. Raton formation, known from a great 

number of localities in the Raton Mesa region of Colorado and New Mexico; for specific localities see United 

States Geological Survey Professional Paper 101, 

pages 312, 313. Black Buttes coal group, Black 

Buttes, Wyo. 

Order RAN ALES 

Family NYMPHAEACEAE 

Genus NELUMBO 

The leaves of this genus are in general: so characteristic that with adequate material there is little or no 

difficulty in placing them with a great degree of certainty. The genus comprises only two riving species-N elumbo lutea nVilldenow) Persoon, the American lotus or water chinquapin of eastern and southern 

North America from Ontario to Oklahoma, and N el'U77~bo nelumbo (Linne) Karsten, the Indian or sacred 

lotus of Asia and Australasia. The American species 

has leaves from 30 to 60 centimeters (1 to 2 feet) in 

diameter, and the Oriental species has still larger 

leaves, from 60 to 90 centimeters (2 to 3 feet) in diameter. In both species the margins of the leaf are entire and there are 19 to 23 strong, radiatinO' ribs 

which fork some distance below the margin. b 

Although now comparatively restricted in its range, 

the genus N elU1nbo has had an interesting geologic 

history which takes it over a wide area where it is not 

now present. Nearly 30 fossil species have been described, of which about haH are known from North 

America, including Greenland. As certain of these 

species are based on a single specimen, often a mere 

fragment, it is more than probable that the number 

of recognizable forms will be somewhat reduced when 

larger series are available for comparison. 

There is a difference of opinion as to when and 

where this type originated. If Berry is correct in 

:iews recently ~xpressed, it may have had its origin,' 

ln company WIth the oldest known dicotyledons, in 

the Lower Cretaceous (Patapsco formation of the 

Potomac group) of the Atlantic Coastal Plain. In 

1911 Berry 55 established the genus N elU1nbites based 

en the species originally referred by Fontaine to 

M enispermites. Concerning these Berry' wrote as 

follows: 

While the Potomac species have the characteristic peltate 

leaves they are not radially symmetrical as are the later 

species but have the petiole attached nearer to one margin 

giving them an appearance much like that of a number o~ supposed species of Menispermites. The venation is, however, 

nearer that of NeZum'1Jo and its allies, the secondaries being 

prominent on the lower surface, obsolete on the upper surface, 

and forking after the manner of the Nymphaeaceae. 

These leaves are orbicular or somewhat elliptical in 

~hape and 10 or 15 centimeters in diameter. They 

are distinctly peltate, but the pelta is excentric, being 

much nearer one side than the other. When they are 

oriented with the pelta 011 the lower side it appears 

55 Berry, E. W., Maryland Geol. Survey, Lower Cretaceous p. 462 

1911. ' I 
SYSTEMATIC PALEOBOTANY 89 

that there is a slight tendency for the leaves to be 

heart-shaped at the base and slightly pointed at the 

apex. It also appears tha.t there is a more or less 

distinct midrib with from two to four pairs of secondaries, the lower of which are joined by broad arches 

with branches from the next lateral ribs. The margin 

I()f the leaves is often undulate or obscurely crenulate. 

It appears to moe that the characters enumerated 

above tend to make these leaves ancestral to Oal8talia 

l'ather than to Nelu1noo. In Oastalia there is always 

a distinct midrib with a greater or less number of secI()ndary branches' on each side, and the base has the 

margin cut to the point of attachment of the petiole. 

If the indentation already indicated in these leaves 

i~ continued to the point of attachment of the petiole, 

they would undoubtedly be referable to the living 

Oastalia. In order that Nelumoite:8 could be developed 

into N elJumoo it would be necessary to suppress the 

obvious tendency to basal lobation, to move the point 

of petiolar attachment to a central position, and to 

o eliminate the midrib or modify it into an ordinary 

forked' rib. This evolutionary process, while perhaps 

possible, seems a far greater modification than simply 

deepening the basal lobation that is already outlined, 

which would permit its allocation in Oastalia. In 

this connection it is also pertinent to point out that 

this basal lobation may have been more pronounced 

than is now recognized. In discussing N elu'1noites 

virginien8is (Fontaine) Berry, which is the type of 

the genus, Berry 66 stated that the type of the species 

is a fragment of a large leaf about 15 centimeters in 

diameter. 

Tbe preservation is such that its peltate character can not 

be made out witb certainty, tbe specimen baving the appearnnce of baving a deeply auriculate overlapping base, but tbis 

is probably due to maceration or folding, as the 0 additional 

specimens since found whicb resemble it very closely are distinctly peltate. 

Berry also calls attention to the undoubted reselllblance betwen N eZWlnoites virginiensis and certain 

Dakota forms referred to M enispermites, such as M. 

{J1'amd'is Lesquereux 51 and M. cyclophyll'l1ls Lesquereux.fiS It is, indeed, difficult to escape the conviction 

that the Dakota forms are congeneric with the Potomac forms, but even so it shows that there was 

but little evolution between Potomac and Dakota time. 

'The leaves referred to Menisp~tes grandis, for example, are of the same shape and have the incipient 

basal indentation, excentric pelta, and above all the 

distinct midrib that are characters in N elwm,oites 

virginiensis. In other words, it seems to me impos1>0 Bert·y, Eo '",., Maryland Geol. Survey, Lower Cretnceous, p. 463, 

1911. 

~; Lcsql1el'eux, Leo, 'l'he Cretaceous find Tertiary floras, p. 80, pI. 15, 

figs. 1, 2, 1883. 

&3 Idem, p1. 15, fig. 3. 

sible that either N elulJrvoites or the above-mentioned 

species of M enis'Pe1"1nites can be in the direct ancestral 

line of the modern genus N elwrnbo. 

Saporta 59 recorded the presence of two species of 

Nelwmoo (Nelumoiwm hesitf1lnioo1n and N. chotfat~) in 

the supposed Albian of Portugal. This is in approximately the same stratigraphic position as the Patapsco . 

:formation, whence came the forms referred to NeZWl71;-

bites. As neither of Saporta's species is figured or 

adequately described it is impossible to compare them 

critically with the American forms, though from his 

comments it is to be inferred that he regarded them' 

a~ pretty closely related to the living Nelwmoo. He 

did not state that the point of attachment of the ribs, 

of which there are 9 in N. chotfati and 20 in N. hesituniCWln,. is excentric, but the material is fragmentary 

and perhaps this feature could not be ascertained. 

With one of the forms he found portions of the flowers, which were not unlike those of the living species. 

Up to this point we are somewhat in doubt as to the 

value of the evidence pointing to the presence of 

N elumoo or its probable ancestors in the Lower Cretaceous. I have pointed out tp.e reasons why in ll1y 

up inion the two species of N elU17h.oites do not offer 

convincing proof on this point, and the two forms 

from the Albian of Portugal are still too imperfectly 

known to permit a final estimate of their value. However "this may be, when we take into account the strong 

resemblance if not actual identity between the dicotyledonous flora of the Patapsco of Atlantic America and 

the Albian of Portugal it would not be surprising to 

find the four forms at least congeneric. It Inust be 

left to the future to settle these points. 

vVe may now turn to the fossil forms which with' 

little question should be regarded as ancestral or directly referable to the living genus N elurnoo'. So far 

as available knowledge goes the genus with its lllodern 

characteristics was already established in Greenland 

in early Upper Cretaceous time. This form was described by Heer 60 under the name N eZurmOiWln {If}''ctiCWl1~ (Nel1.lRnbo (lIJ"ctica (Heel') Tuzson) from the lower 

Atane beds of Greenland. It is a fragment from the 

central portion of a leaf and shows the pelta with the 

bases of 12 strong radiating ribs. None of the margin 

was preserved. This species is obviously closely related to and perhaps identical with N elu171,oo kempii 

(Hollick) Hollick,ol from the Magothy formation of 

. Long Island. The Atane beds are of practically the 

same age as the Magothy formation of New York and 

on Saporta, Gaston de, Nouveaux details concernant les Nympheninees infracretacees: Compt. Rend., vol. 119, pp. 835-837, 1894. 

00 Heel', Oswald, Flora 'fossills arctica, vol. 6, pt. 2, p. 92, pl. 40, 

fig. 6, 1882. 

61 HOllick, ArtbUl", Additions to tbe paleobotany of tbe Cretaceous 

formation of Long Island, No.2: New York Bot. Garden Bun., vol. 3, 

p. 412, pl. 74, figs. 1, 2; pIs. 75, 76; pl. 77, fig. 1, 1904. 
90 FLO.RA .OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

New Jersey, and there is thus abundant reason for 

the probability of this being identical. Hollick has 

described N. kempii as having leaves approximately 

40 centimeters in diameter.. The only difference that 

can be noted between them is in the number of ribs, 

N. kemp~i having from 15 to 19 and N. atrcticfiJ only 12. 

Under the name N elumoo primaeva Berry 62 described a small form from the ::M:agothy formation at 

Cliffwood, N. J., later found in beds of the same age 

at Round Bay, Md. Available material of this form 

is so fragmentary that it is still imperfectly known. 

So far as can be made out, however, these leaves ranged 

in diameter from· 3 or 4 to about 10 centimeters. 

They are peltate; orbicular or broadly elliptical, and 

with entire margins. The ribs are eight in number, 

"generally st.raight, prominent in the lower surface 

of the leaf, forking dichotomously at variable dista.nces from their origin, and giving off thin, transverse, more or less curved secondaries." 

Berry 63 has recently transferred this species to his 

genus Nelumoites with the statement that it is "probably a descendant of N elumoites virgiwie1lJ8is," which 

has already been discussed. He also expresses the 

opinion that N ellwlnoo kempii should probably be referred to Neltumoites. It does not seem to me that this 

position is well taken, especially as regards N elumoo 

kernpii. This species certainly had a large centrally 

peltate leaf, with numerous, equally strong, eV'enly 

disposed ribs and no particular evidence for selecting 

one that could be interpreted as a midrib. These are 

characters of Nelumooand not of Nelumbites. 

The status of Nelumbo primaeva' is not quite so 

ciear, for the reason that the available nlater~al is so 

scanty. So far as can be seen from the published figures this form also has centrally peltate leaves and no 

midrib such as that seen in the type species of N eliwmbites. Until more conclusive evidence is supplied this 

should be retained in N eZumoo. 

A small but very well characterized species is N elumoo intermedia Knowlton,G4 described from the 

Mesaverde formation at Point of Rocks, Wyo. This 

5pecies has thin leaves 2 to 4 centimeters in diam-:-

eter, with slightly undulate margins and 12 or 13 

rather weak, tortuous, forking ribs. 

Another species that is evidently closely related to 

the last and is of approximately the same· stratigraphic position is N eliwmbo dWUJisoni Hollick 65 (originally named Brasenia antiqua by Dawson but that 

62 Berry, E. W., New York Bot. Garden Bull., vol. 3, p. 75, pl. 43, 

fig. 1, 1903. 

03 Berry, E. W., Maryland Geol. Survey, Upper Cretaceous, p. 84Q, 

pI. 75, fig. 4, 1916. 

64 Knowlton, F. H., The fiora of the Montana formation: U. S. Geo!. 

Survey Bull. 163, p. 53, pI. 13, figs. 3-5, 1900. . 

65 Hollick, Arthur,. A new fossil N elumbo from the Laramie group at 

Florence, Colo.: Torrey B.ot. Club Bull., vol. 21, p. 309, 1894. 

name preoccupied by the earlier B. antiqua of Newberry), from the Belly River formation near Medicine 

Hat, Alberta. This is similar is size and appearance 

to N. intermedia but has 18 strong ribs. Still another 

of these small species is N elumoo lar(Jffniensis Hollick,66 

from the Vermejo formation at Florence, Colo. It 

has leaves 4 centimeters in diameter and with about 

12 relatively strong ribs that branch rather than fork, 

thus differing from N. interrn.edia. 

I recently 67 described under the nalne Nelumoo sp. 

a form from the Montana group (Fruitland formation) in the San Juan Basin, N. Mex. It was based 

on a segment from the center of a leaf that is at least 

12 centimeters in diameter and was probably twice this 

size when living. It is concave and evidently t.hick 

in substance. The ribs are somewhat indistinct, but 

apparently there are 18 or 20. This leaf is of the type 

of the living N elurmoo lute{/). 

Passing still. higher geologically we have N eZumoo 

te11ll1dfoU{/) (Lesquereux) Knowlton (Nelwmoium tenwifolirum Lesquereux 68), fronl the Laramie and Denver 

formations of the Denver Basin of Colorado: This 

species has a rather thin leaf 9 or 10 centimeters in 

diameter and 13 thin ribs. It is very closely related 

to Nelumoo lakesiana (Lesquereux) Knowlton (Nelumoium lakesii Lesquereux 69) and N elumoo crossii 

Knowlton (p. 93), both from the Denver formation. 

The first of these has leaves of about the same 

,size as N. tenuifolia, but the leaf substance appears 

to have been firnler and the 13 or 14 ribs much stronger. 

In N. crossii none of the margin is preserved and there 

are only 7 or.8 ribs. This, it will be recalled, is the 

same number as in LVelumoo primaev{/). The three 

species mentioned in this paragraph are discussed at 

length in the following pages. 

Da wson described but did not figure a small leaf 

from the Canadian Upper Laramie under the name 

Neltumbium saskmtchuense.70 It is said to have only 

seven ribs. The last American species to be mentioned 

is Nelumbium pygmaeum Dawson,71 from beds supposed to be of Miocene age on the Similkameen River, 

British Columbia. This form is ·so poorly. described 

and figured th,at it is difficult to place satisfactorily. 

Dawson says it is about 2 centimet.ers in diameter, 

with the "petiole subcentral " and about 14 thin ribs 

"netting toward the margin." If this is really a 

Nelumoo, the petiole should be approximately centrai, 

in which case it would be at least 4 centimeters in 

diameter and the ribs probably as many as 18. It is 

66 Idem, p. 308, fig. in text. 

67 Knowlton, F. Fl., Flora of the Fruitland and Kirtland formations: 

U. S. Geol. Survey Prof. Paper 98, P. 341, 1916. 

os Lesquereux, Leo, 'l'be Tertiary Hora, p. 253, pI. 46, fig. 3,1878. 

66 Idem, p. 252, pI. 46, figs. 1, 2, 1878. 

70 Dawson, J. W., Roy. Soc. Canada Trans., vol. 5, sec. 4, p. 35, 1887. 

';1 Idem, vol. 8, sec. 4, p. 87, text fig. 22, 1890. 
SYSTEMATIC PALEOBOTANY 91 

obviOl1S that further data are needed concerning this 

form. 

Since the above account was written Berry 72 has 

described a splendi d new species-N elwmbo protolutea-from the upper part of the vVilcox group (Grena,da formation) at l\1eridian, Miss. It is a large species with leaves ranging from 25 to 35 centimeters in 

diameter, and associated with it are fragments of rootstocks showing traces of roots with leaf cushjons and 

rootlet scars. T'here are numerous rootlets in the 

days which show that the plants grew in the exact 

spot where they are now preserved. Associated with 

the other remains are some poorly preserved oblatespheroidal seeds, about 1.3 centimeters in diameter, 

that are considered to be the seeds of this species. It 

rnay be added that the number of ribs in the leaves 

appears to be about 25. Berry considers his N elWlnbo 

p'rotol~ttea to be wholly unlike any Anlerican Tertiary 

species heretofore known. It is, however, obviously 

very closely related to lVelumbiu1n protospeciosum 

Saporta73 from the Aquitanian of southern France; 

in fact, he adds: "If the two occurred at the same 

horizon it would be inlpossible to distinguish them by 

thei t· foliage alone." This French species was considered by Sapol'ta to be descended from Nelumbiu1n 

p1'ovVnciale Saporta,74 of the Aturian of southern 

France, and in turn to have led to the living sacred 

lotus (lVelWlnbo nelu1nbo) of India. 

Paralleling this line of descent in Europe, Berry 

now considers that N elWlnbo lcempii of the Magothy 

formation of Long Island and New Jersey was the 

,; logical ancestor" of N elU1nbo protolutea of the Wilcox, which in turn gave rise to the living N eliumbo 

lu,tea of eastern North America. 

Nelllmbo lakesiana (Lesqllerellx) Knowlton 

Plate 41, Figure 1; Plate 42, Figure 2 

Nu7nmbo lalccsiana (Lesql.lCreux) Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 308, 1917 [1918]. 

'NeZmnb'i.mn Zak6sianu11'I, Lesquereux, U. S. Geol and Geog. Survey Terr. Ann. Rept. for 1873, p. 403, 1874. 

"l\T elmnbi1tm lalcesU Lesquereux, The Tertiary flora, p. 252, pI. 

46, figs. 1, 2, 1878; U. S. Geol. and Geog. Survey Terr. 

Ann. Rept. for 1876, p. 514, 1878; Harvard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 53, 1888. 

The type specimens of this beautiful species are preserved in the United States National Museum (Nos. 

370, 371), together with a number of others more recently found. They were collected about 1875 by 

Arthur Lakes and were named l1y Lesquereux in his 

7~ Berry, E. W., U. S. Geol. Survey Prof. Paper 108, p. 64, pIs. 24-26, 

1017. 

78 Saporta, Gaston de, ReclH'rches sur In v«5getntion du niveau aqui· 

tallien de Munosque, p. 17, pI. 1, figs. 2, 3; pI. 4, figs. 1, 2, 1891. 

74 Supoi·tn, Guston de, Le Nel1t1nbium P1'ovinciale: Soc. geol. France 

Po I. Mem. 5, 1890. 

honor. The original description by Lesquereux is as 

follows: 

Leaves coarse, tllickish, peltate, exactly round, with the 

petiole central; borders turned down; center concave, regular; 

all the ribs (14) equal in thickness, equally diverging from 

the center to the circumference, deeply marked, branching 

near the borders, crossed by thick, flexuous nervilles at right 

angles and disjointed; surface rough. 

Lesquereux added that "this species is represented 

by three specilnens of the same form, two small leaves 

and a much larger one." The two figured specimens 

are the only ones of the original specimens that are 

present in the Museum collection. 

In the living species of N elrwtnbo the ribs are usually 

if not always dichotomously forked some' distance below the margin. In discussing N elwmbo lakesiap,a in 

the report on the Tertiary flora Lesquereux called attention to the fact that the ribs appear to be branched 

rather than forked. 

The ribs are deeply impressed and thick, sparingly branching and, so far as can be seen, only on oue side, and n'ot 

by dichotomous divisions. ... ... ... It is scarcely possible to 

see a difference in the size and in the directions of the nerves 

[ribs], which are more or less tUrned on one side; none of 

them has dichotomous divisions, and thus the leaves of t11is 

species seem far different from those of our time, and also from 

those described from the European. Tertiary. 

A .restudy of the type specimens appears to substantiate the above contention, but both are so fragmentary 

that not all the characters can be made out. The' 

original of Lesquereux's Plate 46, Figure 1, does not 

ha ve any' portion of the margin preserved, and that 

of Figure 2 has only a small portion which shows the 

ribs complete.. Each of these ribs is curved to the 

right and has on the opposite side two or three secondary branches, which are camptodrome and arch just 

inside the margin. This manner of branching is certuinly different fron1 the dichotomous forking that 

is normal in the living and in many fossil species. In 

this character N elwmbo lakesiama approaches the remarkable lobed form described in this report under 

the name Paleonel1.tmbo mac-roloba, except that the 

latter has the ribs straight with secondary branches 

on both sides. In the original of Lesquereux's Figure 

2 the margin is retained for only about 3 centimeters, 

and while there is some slight evidence that it may 0. 

be lobed as in Paleoneltumbo, the evidence is not conclusive. 

Lesquereux stated that the leaves of N elumbo lalcesiana appear to be about 12 centimeters in diameter, 

but an examination of the type of his Figure 1 discloses the fact that a portion of the leaf is folded 

over and is not shown in the figure. Even this does 

not reach the margin, but it permits a measurement 

of over 8 centimeters for the length of the rib, which 
92 FLOR.A OF THE DENVE¥ AND ASSOOIATED FORMATIONS OF OOLORADO 

.' 

would make the whole leaf at least 16 centimeters in 

diameter. It was probably not less than 20 centimeters in diameter. The type (Lesquereux's fig. 

2), in which a portion of the margin is retained, is 

about 12 centimeters in diameter, and this is probably 

the one on which Lesquereux based his statement of 

size. The range in size of the leaves of N elWlnbo 

I a7~esiana is thus shown to be from 12 to 18 or 20 

centimeters. 

The types of N elwmbo ZO)kesianm are preserved in 

the coarse andesitic matrix characteristic of South 

Table Mountain, Golden, Colo. It appears, however, 

to be a rare species, for in the numerous collections 

from Golden and vicinity that have passed through 

my hands in recent years no trace of it has been 

detected. In the collection made by Lakes and now 

the property of the Museum of Comparative Zoology 

Lesqnereux noted the presence of two specimens but 

made no comment on them. 

In the collection made by Lakes near the Douglas 

coal mine, Sedalia, Colo., I find no less than five more 

OJ" less perfect fragments that are referred with 

much certainty to Nelwnbo lake8iarna. They have the 

same thick ribs, which produce, when preserved on 

similar matrix, the rough appearance described by 

Lesquereux for the type specimens. The number of 

ribs is usually 14, though one has but 13, the number 

described as present in N elwmbo ternuiJfoZia. . One 

. specimen shmvs the under surface of a leaf with a 

small portion of the petiole preserved. The petiole 

i& about 3.5 millimeters in diameter, which is about 

'the size of the petiole in the living NeZwmbo lutea. 

The margin is not well enough preserved to show 

with certainty the manner in which the ribs approach 

the border. 

N elttrmbo la)kesWma seems to be rather closely related to N elwmbo airctica (Heel') Tuzson,75 from the 

lower Atane beds of Greenland. The Arctic species 

is based on a small fragment from the center of a 

leaf, but so far as can be made out it appears to agree 

closely with N. lakes'iam(J). It has 13 thick ribs radiatillg from the common center, but all finer nervation 

is obsolete. 

. Occurrence : Denver formation (types), South 

• Table Mountain, Golden, Colo., collected by Arthur 

Lakes, for whom the species is named, about 1875; 

also collected at about the same locality by Lakes in 

1883 and now in the Museum of Comparative Zoology, Cambridge, Mass. Dawson arkose, lower part 

(in beds equivalent to Arapahoe formation), Sedalia, 

Colo., 3,000 feet east of the old Douglas coal mine, colIpcted by Arthur Lakes, 1890. Raton formation, 

j5 Tuzson, J., Magyar tudom. nkad. :Math. tcrmesz. ~rt., vol. 29, No.5, 

p. 827, 1911. 

Fishers Peak mine, 3 miles southeast of Trinidad, 

Colo., collected by G. B. Richardson. 

Nelumbo tenuifolia (Lesquereux) Knowlton 

PIa te 41, Figure 2 

Nelttmbittm tenttifoZium Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1873, p. 402, 1874; idem 

for 1876, p. 514, 1878; The Tertiary flora, p. 253, 111. 

46, fig. 3, 1878. 

The type specimen of this species came from SaJ?d 

Creek, a few miles east of Denver, Colo., and is preserved in the United States National Museum (No. 

372). It is the only specimen in~ the Museum collection under this name, although Lesquereux, in the 

report on the Tertiary flora (p. 254) Inentioned having two leaves, one of which was" obtained fronl a 

different locality." It is not now possible to determine the source of this unfigured specimen. 

The type specimen' of Ne"Vumbo tenuifoUa is preserved on a soft, friable sandstone and is very indistinct. Lesquereux's figure of it is far from accurate. 

The only part of the true margin retained is a fragment hardly more than 3 centimeters in extent in the 

upp~r left-hand portion of the figure. A careful 

examination fails to show the margin on the lower 

portion, although it is represented in the figure. This 

seems to prove that the leaf was larger than is shown 

in the figure-that is, it was at least 12 centimeters in 

diameter instead of the 8 or 9 centimeters stated by 

. Lesquereux. It has relatively thin ribs as described 

by Lesquereux and altogether appears to have been 

a thin leaf, but otherwise it approaches N eltlllnbo 

lakesiana closely. The number of ribs (13) is 

. probably not of specific value, as some undoubted 

leaves of N. lakesiOJna have only 13. The manner in 

which the ribs turn to one side and have branches 

rather than a distinct fork are also characters common 

to the two forms, as was pointed out by Lesquereux. 

He concluded, however, that" the great difference in 

their substance seems sufficient to authorize a separate 

specification," and it is largely on this basis that they 

a re retained as. distinct. 

The stratigraphic position of the locality on Sand 

Creek which s~pplied the type specimen of Nelu1nbo 

tenuifolia has long been more or less in doubt. It was 

supposed to. be in the Arapahoe formation, but daia 

recently procured place it without 111uch doubt in the 

Denver formation .• The complication regarding the 

stratigraphic position of this locality is fully discussed 

i~ the introductory part of this report (p. 6). 

In a collection made some years ago from the Laramie rocks at Erie, Colo., there was a single example 

that has been referred with considerable certainty to 
SYSTEMATIC PALEOBOTANY 93 

Nelwmbo tenuifolia. 70 It was folded around on both 

sides of a thin piece of ma.trix a.nd appears to have 

been a thin, delicate leaf. Its diameter is about 10 

centilneters. It has the same number of ribs as the 

type specimen, and the ribs tend to be turned to one 

sid~ and to branch rather tha,n fork as in N. lakesiana. 

The finer nervation is obscure. 

Occurrence: Denver formation (type), Sand Creek, 

a bout 15 miles east of Denver, Colo., collected by A. 

Gardne.r about 1873. Laramie formation, Erie, Colo., 

collected by N. L. Britton about 1885. 

Nehunbo Cl'ossii }{nowlton, n. sp. 

Plate 41, Figure 3 

Leaf of thin texture, subcircular, centrally peltate, 

seven or perhaps occasionally eight ribs, the ribs 

rather thin, diverging with approximate regularity, 

forking above; nervi lIes strong, a few percurrent but 

lllainly broken in the middle; finer nervation producing irregular areas. 

It is with some doubt and hesitation that this leaf 

is described as new. It is found associated in the 

same beds with N elwm,bo lakesiana and was at first 

thought to belong to this species, but it has only 7 or 

. possibly 8 ribs, whereas the other always has 13 or 14, 

nnd this is thought to be too great a difference to be 

reconciled with our recognized knowledge of specific 

variation. The number of ribs in the living N. lutea 

is usually 21 but may vary from 19 to 23. So far as 

is known to me no living leaves which showed a reduction to half of the normal number of ribs have been 

found. Therefore it is upon this considerable clifference in the nUlnber of ribs as compared with its 

nearest relative that this species is proposed. 

As shown in the figure, N. c'rossii is founded upon a 

mere fragment of the center of a peltate leaf. At no 

point is the margin preserved, yet from tJle longest 

fragment of a rib retained it appears that the leaf 

must have been at least 8 centimeters in diameter, and, 

as the ribs still appeal' strong at the point where 

broken, it was probably considerably larger. The two 

best-preserved ribs are forked as in typical N elumbo, 

and the intermediate nervation is also similar. It is 

therefore an undoubted N el'lVJnbo, but its specific status 

must remain in more or less doubt pending the discovery of n10re perfect material. 

Dawson 77 has described under the name NelwniJiUffn 

saslcatcl1JUense a small species from the so-called Canadian Upper Laramie of the North Saskatchewan. 

This species was described as "orbicular, pelt ate, with 

seven ribs" and a diameter of 2 centimeters or a little 

70 Knowlton, F. H., The Laramie flora of the Denver Bnsin: U. S. 

Geol. Survey Prof .. Paper 130, p. 141, pI. 26, fig. 7, 1922. 

77 Dawson, J. W., Roy. Soc. Canada Trans., vol. 5, sec. 4, p. 35, 1887. 

85492-30-7 

more. This agrees with iV. 01'ossii, it will be observed, 

in the number of ribs, but the size is so small as apparently to put out of consideration the question of 

specific identity. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Arapahoe formation), 3,000 feet. east of 

the Douglas coal mine, Sedalia, Colo. 

Genus PALEONELUMBO Knowlton, n. gen. 

Similar to the living N eZwrwo but with the leaves 

provided with high, obtuse teeth or lobes, one for 

each rib. 

Paleonelumbo macroloba Knowlton, 11. sp. 

Plate 39, figure 3; Plate 42, Figures 3, ':1 

Leaf very thick and firm, circular, centrally peltate, 

\.!oncave, about 30 centimeters in diameter; marginal 

lobes the same number as the ribs, very obtuse and 

rounded at the tips, about 4 centimeters long and 3 or 

3.5 centimeters wide at the base, separated by deep, 

rounded sinuses; ribs 12 or 13, very thick a.nd strong, 

each terminating in a lobe, with three or four pairs of 

strong secondary branches in the upper portion, these 

arching upward and joining, producing a loop just 

within the margin of the lobes; nervilles numerous, 

especially in the central portion of the leaf, strong, 

unbroken, approximately at right angles to the ribs. 

This large and very striking species is represented 

in the collection by several specimens, three of which 

are here figured. The most perfect of these (pI. 39, 

fig. 3) is an impression of the under side of the leaf 

which brings out the strong ribs and other nervation 

in an admirable manner. The specilnen is cupped 

around the matrix, showing that it was deeply concave above, after the manner of the living lotus (N elumbo nelu1nbo) . The others figured sho'Y the central 

part of leaves with strong, radiating ribs, but no part 

of the margin is retained. 

These leaves belong undoubtedly to the Nymphaen.ceae and are most closely related to the genus N elU1nbo, which comprises only two species,.N el/wmbo 

lutea (Willdenow) Persoon, the American lotus or 

water chinquapin of southeastern and eastern North 

America, and NelWlnbo nelumbo (Linne) Karsten, the 

well-known Indian lotus of Asiatic and Australasian 

countries. Both have entire, more or less concave 

leaves 1 to 3 feet in diameter and more numerous ribs 

which fork into equal or nearly equal branches well 

below the margin.' The fossil fonn differs from the 

Ijving N el'l(;1nbo in the fewer ribs, pronounced marginal 

lobes as many as and entered by the ribs, and secondary 

branches in the upper portion of the ribs instead of 

equal forks. These differences are so marked! as to 

appear to entitle it to generic rank. 

o 
94 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Occurrence: Dawson arkose, about a quarter of a 

mile east of Purdon mine, sec. 27, T. 11 S., R. 61-W., 

Colo., in sandstone about 25 feet above the Purdon 

coal, collected by G. B. Richardson, 1910. 

Castalia pulchella Knowlton, n. sp. 

Plate 42, Figure 1; Plate 57, Figure 3 

Leaf evidently rather thin and membranaceous, apparently nearly circular, fissi-cordate, the lobes rather 

short, rounded, the sinus acute; margin conspicuously 

toothed the teeth low, obtuse, separated by shallow , d' rounded sinuses; ribs, including the midrib, 11, ra IatinO' from a central point or pelta; midrib little if any b • 

stronger than others, with three or four slender paIrs 

of branches; lower ribs turning down into the lobes, 

the others disposed at equal distances, all splitting 

into two equally strong branches which join in a broad 

loop with two or three series of progressively smaller 

loop~ until the margin is reached; nervilles fairly 

strong, mostly unbroken, at right angles between the 

ribs. 

This remarkably fine and distinct species is represented only by the specimen figured and its restoration. The central part of the leaf is preserved, 

showing the radiating ribs or primary veins, as 

Caspary called them, in a high degree of perfection, 

but the margin is retained in only a few places. As 

nearly as can be made out the leaf was nearly c~rcul~r 

or perhaps a little broader than long. The WIdth IS 

approximately 8.5 centimeters, and the preserved 

length only about 7 centimeters, but both base and 

apex are not wholly preserved. The teeth are conspicuously large for so small a leaf of this genus. 

There is some evidence of the presence of smaller 

slight protuberances on the main teeth, but this is not ' 

very conclusive. 

This leaf exhibits distinctively and unmistakably 

the outline and peculiar forking and anastomosing of 

the ribs that are characteristic 'of Oastalia, and there 

can be no doubt about its being correctly referred to 

this genus. Oastal~ (Nymphaea of authors) comprises about 35 living species and is found here and 

there' almost all over the world. As they hybridize 

freely a large number of hybrids ~nd garden varieties 

are known and prized in cultivation. The genus was 

~onographed in 1905 by H. S. Conard,78 of the University of Pennsylvania, and from this splendid pub-' 

lication it is possible to make a number of comparisons 

'With the fossil form under consideration. Thus in 

~ome ways ,this suggests Oastalia rudgearna (Nymph~ea 

rudge(J.lJW, Myer), a species living in tropical AmerIca 

and cert~in West Indian Islands, except that the liv78 Conard, H. S., The water lilies, a monograph of the genus 

~ymphaea.' Carnegie Inst. Washington Pub. 4, 1905'. 

ing leaf is much larger than the fossil. In marginal 

dentition the living Oastalia stellata (Nymp'haea stellata Willdenow), of wide distribution in southern and 

southeastern Asia and neighboring islands, approaches 

the fossil leaf but differs markedly in the number and 

strength of the ribs. Another species with similarly 

toothed, though much larger leaves is Oastalia fia1Jovirens (Nymphaea flavo-virens Lehmann). 

This is the fourth fossil species to be described from 

this country, though neither of the others can be considered, as being so well established as the, present one. 

These are Oastalia stantoni Knowlton,79 from the 

Judith River formation of Montana; Oastalia? duttoniana KnowIton,80 from the Montana group of the 

Laramie Plains, Wyoming; and Oastaiia leei Knowlton , 81 from the Raton formation of southern Colorado. 

These are all very unlike Oastalia pulchella. 

Several species were described by Heer 82 from the 

Tertiary of Greenland, but they are either based on 

seeds or so fragmentary as to be of little comparative 

vaiue in the present connection. Thus his Nymphaea 

arctica is a mere fragment that mayor nlay not have 

been correctly placed. 

A dozen or more European species have been described, most of them being correctly referred to 0 astalia (Nymphaea) , but none approach closely the fossil form here described. 

Although it may perhaps ,seem a wide generalization 

to make on the data available, it seems not unlikely 

that what is here described as Oastalia pulchella may 

be in the direct line from which Oastalia rudgeana 

has been derived. 

Occurrence: Dawson arkose, left bank of Jimmy 

Camp Creek 0.7 mile above Richfield Springs ranch 

house, about 9 miles east, of Colorado Springs, Colo., 

collected by F. R. Knowlton and W. T. Lee, 1910. 

Family ANONACEAE 

Asimina eocenica Lesquereux 

Plate 40, Figure 4; Plate 43, Figures 1, 2, 4, 6 

Asimina eocenica Lesquereux, U. S. GeoI. and Geog. Survey 

Terr. Ann. Rept. for 1872, p. 387, 1873; idem for 1876, 

p. 514, 1878; The Tertiary fiora, p. 251, pI. 43, figs. 5-8, 

1878. 

, Only a part of the type specimens of this fine 

species appear to be preserved in the collections of 

the United States National Museum [No. 362 (" Ter79 Knowlton, F. H., in Stanton, T. W., and Hatcher, J. B., Geology 

and paleontology of the Judith River beds: U. S. GeOI. Survey Bull. 

257, p. 147, pI. 19, fig. 4, 1905. 

80 Knowlton, F. H., Flora of the Montana formation: U. S. Geol. 

Survey Bull. 163, p: 55, pl. 13, fig. 7, 1900. 

81 Knowlton, F. H .. U. S. GeoI. Survey Prof. Paper 101, p. 307, pl. 

79, fig. 2, 1917 [1918]. , 

, 82 Heer, Oswald, Flora fossilis arctic a, vol. 5, pt. 2, p. 44, pI. 13, 

fig. 7, 1878. 
SYSTEMATIC PALEOBOTANY 95 

tiary flora," fig. 7) ; No. 363 (fig. 8)], although there 

are several unfigured examples recorded under the 

same number. No additional specimens appear to 

have been obtained from the type locality. 

In the paleontologic collection of Princeton Univer~ 

sity, kindly placed at my disposal by Prof. William 

Libby, jr., director of the museum,' I find a single 

specimen (No. 2207) from South Table Mountain, 

Golden, Colo., that must be referred to this species. 

'Phis specimen~ figured on Plate 43, Figure 6, lacks 

both base. and apex, yet the size and nervation are 

identical with Lesquereux's Plate 43, Figure 8. The 

camptodrome secondaries with their intermediate secondaries and peculiar branching are clearly. the same. 

This is the only specimen from other than the type 

locality. 

In my bulletin on the flora of the Montana formation (p. 57) I mentioned a single example from a 

small coal mine on the north fork of Dutton Creek, 

near the old stage road, Laramie Plains, Wyo., that 

I doubtfully referred to this species. This leaf has 

the same size, shape, and thick petiole and approximately the same nervation, though the secondaries 

are at a little greater .angle of divergence and perhaps pass up along the margin for a greater distance. 

The differences are slight, yet considering the disparity in horizons it is perhaps doubtful if the identity should be insisted upon, and more material will 

be needed to settle the point. 

Among the recently collected specimens from Mount 

Bross, Middle Park, Colo., I find a single basal portion of a leaf that may well have belonged to this 

species; it is shown in Plate 43, Figure 4. Several 

fragments showing only the basal portions a.nd the 

thick petiole have been found at Sedalia, Colo. 

Still more recently a field party under the direction 

of G. B. Richardson collected in red baked shale near 

Ramah, Colo., a large number of beautifully preserved 

leaves that must be referred to this species. There 

are more than 50 leaves of this species in this collection, 5 of which have been figured. They are very 

uniform in size, 10 or 12 centimeters long and 2.25 to 

3 centimeters wide, with the exceedingly thick petiole 

about 1.5 centimeters long. The apex is prolonged 

and truncate or slightly emarginate at the tip. The 

midrib is very thick and strong in the lower portion 

of the blade but rapidly diminishes in the apical portion. It is strongly ridged and striated in the middle 

and lower portions, and this character extends to the 

thick petiole. 

Occurrence: Hanna formation, Carbon, Wyo. 

(types) . Denver formation, South Table Mountain, 

Golden, Colo., specimen in museum of Princeton University. Middle Park formation, Mount Bross, Middle Park, Grand County, Colo., collected by G. L. 

Cannon, 1889. Dawson arj,{ose, lower part (in beds 

equivalent to Arapahoe formation), Sedalia, Colo., 

3,000 feet east of the old Douglas coal mine, collected 

by Arthur Lakes, 1890. Dawson arkose, south of the 

Ramah oil well, Colorado, in the SE. % sec. 33, T. 9 S., 

R. 61 W., collected by C. W. Cooke for G. B. Richardson, September, 1910. Fort Union formation, Wind 

River Basin, Wyo. Midway ~ formation, Earle, Tex. 

Order ROSALES 

Family ROSACEAE 

Chrysobalanus coloradensis Knowlton, n. sp. 

Plate 40, Figures 2, 3; Plate 43, Figures 3, 5, 7-9; Plate 44, 

Figures 1, 2 

OiS8US speatabilis Heer. Lesquereux, Harvard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 52, 1888. 

Leaves thick and coriaceous, oblong - elliptical, 

abruptly rounded and truncate at the base with a small 

portion slightly decurrent on the petiole, rounded and 

apparently obtusely pointed at the apex; margin perfectly entire throughout; petiole about 1 centimeter 

long, very thick and stout, always striate; nervation 

pinnate, consisting of a very strong midrib, which is 

firmly striate below, and about six pairs of very thin 

alternate, camptodrome secondaries at a low angle; 

nervilles numerous, approximately at right angles to 

the secondaries. 

This species is represented by about a dozen leaves, 

all of which are very well preserved except as regards 

the extreme apex. They are very uniform in size, 

about 8 or 9 centimeters long and 4 centimeters wide. 

The petiole is about 1 centimeter long, very thick and 

provided with a· number of fine striae which extend 

up along the extremely thick midrib. The secondaries 

are very thin and immersed in the thick substance 

of. the leaf. 

There is some doubt about the correctness of this 

generic reference. These leaves have a degree of resemblance to certain leaves that have been referred to 

Fious, such as Ficus pwryearensis Berry 88 from beds 

of Wilcox age in the Lagrange formation at Puryear, 

Tenn. On the whole, however, it seems best to refer 

them to Okry)$O'balamws. This is a small genus of 

anlygdalaceous shrubs or . small trees, with simple, 

leathery leaves and an edible pulpy drupe. Three 

species are recognized in Florida, of which the best 

known is O. iClOOO Linne, the so-called gopher or cocoa 

plum. It occurs in sandy soil in the southern part 

of the Florida peninsula and also in the West Indies 

and tropical America. 0 hrysobald!nluS oblongifolius 

Michaux is found in dry, sandy pine lands from 

Georgia to Florida and Mississippi. It is a low shrub 

28 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 205, pl. 30, 

figs. 4, 5, 1916. 
96 FLORA OF THE DENVER· AND ASSOOIATED -FORMATIONS OF COLORADO 

forming wide patches by the spreading of underground stems. The third species is O. pellocarp'Us 

Meyer, a shrub or small tree of low ground in southern Florida and the West Indies. The genus is also 

represented in tropical west Africa, where it occurs 

along the coasts from Senegambia to the Congo Free 

State, perhaps introduced by the agency of lnan. 

Three fossil species of 0 hryso0 alamJ.us have been 

recorded from this country-O. pollard(J/nxU8' Knowlton,84 from the Esmeralda formation (upper Miocene) 

of Esmeralda County, Nev.; O. eocenica Berry 811 and 

O. inaequalis (Lesquereux) Berry,86 from the Lagrange formation of Puryear, Tenn. 

The present fossil specjes appears to be most like 

the living O. waco, from which it differs in its more 

nearly elliptical shape and more obtuse base. The 

a.pex can not be absolutely determined, but it appears 

to be rather obtuse, though not emarginate. In nervati.on O. colo1'adensis is suggestive of the living O. 00-

longifolius, but it differs in shape. 

Since the above was written I have examined the 

Golden material in the Museum of Comparative Zoology as identified by Lesquereux, and I find that the 

specimen recorded as Oissus speataoilis Heel' 81 is to 

be referred to O.h1'1.Jsooalanus colo1'adensis. It is a 

sHghtly smaller leaf-7 centimeters long and 3.5 centinleters broad-but does no~ otherwise differ essentially. 

.A comparison of this leaf with the figure cited by 

Lesquereux in Heer's flora of Sakhalin 88 shows them 

to be totally unlike. 

Occurrence: Dawson arkose, dump of Mosby coal 

mine, Mosby, El Paso County, Colo., collected by 

It'. H; Knowlton and G. B. Richardson, July, 1910. 

Denver formation, South Table Mountain, Golden, 

Colo., collected by Arthur Lakes, specimen in Museum 

(if Comparative Zoology, Cambridge, Mass., No. 1502. 

Chrysobalanus? lanceolatus Knowlton, n. sp. 

Plate 44, Figure 5 

Leaf small, coriaceous, elliptical-lanceolate, slenderly acuminate at the apex, abruptly rounded and 

slightly unequal-sided at the base; margin perfectly 

entire; petiole strong, at least 1 centimeter long, finely 

striate; midrib relatively strong, finely striate; secondaries about eight pairs, alternate, at an angle of about 

450 , very thin and immersed in the leaf substance, 

somewhat curved upward along the margin; finer 

nervation obsolete. 

84 Knowlton, F. H., U. S. Geoi. Survey Twenty·first Ann. Rept., pt. 2, 

p. 216, pI. 30, fig. 19, 1900. 

811 Berry, E. W., U. S. GeoI. Survey Prof. Paper 91, p. 220, pI. 44, 

figs. 4, 5, 1916. 

so Idem, p. 220, pl. 44, figs. 8-10. 

87 Lesquereux, Leo, Harvard ColI. Mus. Compo ZOology Bull., vol. 16, 

p. 52, 1888. 

sa Beer, Oswald, Miocene Flora des Insel Sachalin, p. 45, pI. 3, fig. 

3b, 1878. 

The example that is here figured as the type of this 

species is absolutely perfect. It is 6 centimeters long, 

including the petiole 1 centimeter long, and is about 

1.8 centimeters wide. It is narrowly elliptical-lanceolate, with a rounded, slightly unequal-sided base and a 

slenderly acuminate apex. With the exception of the 

st.rong midrib the nervation is rather obscure, being 

immersed in the evidently thick substance of the leaf. 

This leaf lies by the side of a stem or something of the 

~nd that was at first supposed to be a rachis, but the 

leaf is perfectly free and there is no scar or point of 

attachment to the axis. 

This species is found in a.ssociation with and is 

undoubtedly congeneric with the preceding species, 

and indeed it may be only a narrow, slenderly acuminate form of that species. . It. has the thick substance, 

immersed secondary nervation, and striate petiole and 

midrib of O. coloradensis but differs in shape, ·especially in the apeX'. This species departs so widely from 

the form of the living leaves that it seemed wise to 

question the generic reference. 

Occurrence: Dawson arkose, dump of Mosby coal 

mine, MosbY1 Colo., collected by F. H. Knowlton and 

G. B. Richardson, 1910. 

Family POMACEAE 

Crataegus myricoides Lesquereux 

Plate 44, Figures 7-10 

01"ataeg'u8 1nyricoide8 Lesquereux, Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 56, 1888. 

Following is Lesquereux's original description and 

discussion: 

Leaves membranous, small, ovate-lanceolate, more or less 

deeply dentate, penni nerved ; lateral·nerves simple or forking, 

oblique and straight to the borders, or curving inside before 

entering the teeth, teeth alternately larger, or large and bitridentate on the back. 

The two leaves representing the species are only 2 or 3 

centimeters long, 1.5 centimeters broad, fragmentary. They 

resemble leaves of Betula, but the irregular nervation is that 

of Myrica,· the two lower pairs of nerves being longer and 

entering the teeth, the upper shorter, curved, and effaced near 

the borders, all at unequal distance, not parallel. They have 

some resemblance to the leaves of Ora-taegu,8 oxycun.fhoides 

Goppert. 

This species is based on two types, both of which 

are here figured. They are small leaves 2.5 to 3 centimeters long and less than 2 centimeters wide and are 

poorly preserved. The one best preserved is shown 

in Figure 9. It is elliptical-Ianceolate, with -a wedgeshaped base and apparently a,n acuminate apex. The 

lnargjn is cut into relatively large, sharp teeth, each, 

as Lesquereux states, bidentate or tridentate on the 

back. The nervation is pjnnate, with a thin midrib 

and five or six pairs of alternate, craspedodrome 

secondaries. 
SYSTEMATIC PALEOBOTANY. 97 

The other specimen (fig. 8) lacks the upper half 

of the blade. It is more obtusely wedge-shaped at the 

base than the other, and the margin is so broken that 

the teeth are obscurely preserved. The nlidrib is 

thicker, but the secondaries are about the same, though 

the lower pair show branching. The finer nervation 

i~ not retained in either specimen. 

The generic reference for thes~ little leaves is perhaps open to question. As Lesquel'eux stated, they 

suggest leaves of Betula, such as Betula glandJulosa 

:Michaux and B. l)'UlmUa; Linne, but there are points of 

difference, and it is perhaps as well to leave thenl at 

least temporarily as left by Lesquereux. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, 

MUS8.: Nos. 1567, 1968. 

Family DRUPACEAE 

Prunus denverensis Knowlton, n. sp. 

Plate 44, Figures 3, 4, 6, 11 

Leaves coriaceous, narrowly ovate-Ianceolate, long 

wedge-shaped at the base, prolonged above into a slenderly acuminate tip; margin almost entire below, then 

finely and evenly serrate, the teeth being small and 

sharp; petiole short, rather stout; midrib fairly strong, 

perfectly straight; secondaries numerous, about 12 or 

15 pairs, alternate, at a low angle of emergence, much 

elU'ved upward, camptodrome, arching near the margin and each joining the one above by a series of bows, 

sending minute nervilles to the teeth; nervilles numerous, strong, irregular, mainly oblique to the secondaries; finer nervation producing a close network of 

1 ather elongated areas. 

I f I am correct in referring all these leaves to a 

si ngle species there is a very considerable range in 

size, but as this appears to be the only noticeable difference they are best considered together for the present. The smallest leaf (fig. 6) is about 4 centimeters 

long and a little over 1 centimeter wide, and the largest ieaf (fig. 11) was probably 10 or 12 centimeters 

long (upper portion missing) and 3.5 centimeters 

wide. The very perfect leaf shown in Figure 4 is 5.5 

centimeters long and 2 centimeters wide. A still 

larger one, shown in Figure 3, is 7.5 centimeters long 

and 2.25 centimeters wide. 

This species suggests a number of living American 

forms, such as P1'unus angustifolia Marsh and more 

particularly P. seIJ'oti11,a Ehrhardt, the common wild 

black cherry of the eastern United States. 

Occurrence: Dawson arkose, oil well near Ramah, 

Colo., collected by C. 'iV. Cooke for G. B. Richardson, 

1910. 

Family PAllILIONACEAE 

Phaseolites coloradensis Knowlton, n. 5p. 

Plate 45, Figure 1 

Leaflets rather thick, asymmetrically ovate, obtusely 

pointed at the apex, obtusely wedge-shaped and inequilateral at the base, the margin perfectly entire; 

petiolule short, very stout; midrib very stout; secondaries five or six pairs, lowest pair stronger, subopposite, arising just above the top of the petiole, at an 

angle of about 45°, joining the next secondaries by a 

broad bow, other secondaries alternate and subopposite, at a low angle, camptodrome; nervilles few, 

strong. 

This species is represented by small leaflets; the best 

one, which is practically perfect, is figured. It is 

lllarkedly unequal-sided. It is 4 centimeters long and 

nearly 2.5 centimeters wide. The petiolule is preserved for a length of 5 lnillimeters. The nervation 

is distinctive, . consisting of the strong midrib and a 

lower pair Qf strong secondaries at a different angle 

from the others, all being camptodrome. 

This species· appears to be certainly congeneric with 

Phaseolites oeningerutis Heer,89. from the Swiss Miocene, though the European species is much larger, and 

the lower secondaries are not so conspicuously larger 

. than the upper ones, nor do they differ in angle of 

origin from the midrib. Heer suggested the resemblance of his species to Ficus tiliaefolia but concluded 

that it is only superficial. 

In shape and general appearance Phaseolites ooloIJ1

adensis has some resembl9nce to Pithecolobiwm eoce-' 

nicumBerry, from beds of Wilcox age in the Lagrange 

formation at Puryear, Tenn., but it is larger and di~­

fers somewhat in nervation. It also agrees very well 

in size and shape with certain of the leaflets referred 

to Sapindus obtusifolius Lesquereux,9o from the Green 

River formation of Green River, Wyo., but differs in 

the nervation. 

Occurrence: Dawson arkose, dump of :Mosby coal 

mine, Mosby, El Paso County, Colo., collected by F. 

H. Knowlton and G. B. Richardson, ·July, 1910. 

Sophora richardsoni Knowlton, n. sp. 

Plate 45, Figure 7 

Leaflets small, rather thick, elliptical, rounded below 

to the short, stout petiolule and in about the same 

degree above to the abruptly acuminate tip; margin 

entire; midrib very thick, straight; secondaries about 

five pairs, thin, at an acute angle, camptodrome; finer 

nervation obsolete. 

so Heel', Oswald, Flora tel'tiaria Helvetiae, vol. 3, p. 103, pI. 133·, fig. 

6, 1859. 

90 Lesquereux, Leo, The Tertiary flora, p. 266, pI. 49, fig. 10, 18i8. 
98 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

The little leaflet figured is almost ~egularly elliptical in outline, being 2.5 centimeters long and 1.3 centimeters wide. The petiolule is at least 3 millimeters 

long. 

This species closely resembles a number of described 

species from the Wilcox group; in fact, it would perhaps be no great violence to identify it with Sophora 

paleolobifoli(J) Berry,91 which is described by Berry as 

elliptical and markedly inequilateral, though the figure given does not exhibit this peculiarity. The size 

and nervation agree very well with the Colorado 

sp'ecies. 

The leaflet under consideration also closely resembles some of the smaller leaflets of Sophora wilcoaJiana Ber,ry,02 and it is also of the same shape and much 

the same nervation as Sop'n;ora pury;e{]Jf'ensis Berry.os 

There are, however, slight differences, and altogether 

it seems best to consider this a new species, at least 

until more material can be procured. 

Occurrence: Dawson arkose, dump of Mosby coal 

mine, Mosby, Colo., collected by F. H. Knowlton and 

G. B. Richardson, 1910. 

Sophora puryearensis Berry 7 

Plate 45, Figure 8 

Sophora pw'yearensis Berry, U. S. Geol~ Survey Prof. Paper 

91, p. 242, pI. 52, fig. 3, 1916. 

The little leaflet here figured appears to belong to 

Sophora puryear'ensis, as described and figured by 

Berry.D4 It is smaller than the Wilcox specimen and 

.has only four pairs of secondaries instead of five, 

but its shape is identical. It is not well preserved, 

especially in the upper portion, but it does not seem 

tc differ essentially, though I have questioned the 

full identification. 

Occurrence: Dawson arkose, red hill 4 miles south 

of Ramah, Colo., collected by G. B. Richardson, 1910. 

Lagrange formation, Puryear, Henry County, Tenn. 

Leguminosites? araehioides (Lesquereux) Lesquereux 

Leg1Jll11linosit8sY arachioides (Lesquereux) Lesquereux, The 

Tertiary fiora, p. 301, pI. 59, figs. 13, 14, 1878; U. S. 

Geol. and Geog. Survey Terr. Ann. Rept. for 1876, p. 

519,1878. 

Ward, U. S. Geol. Survey Sixth Ann. Rept., p. 554, pI. 49. 

fig. 7, 1886; U.· S. Geol. Survey Bull. 37, p. 65, pI. 29. 

fig. 2, 1887. 

Penh allow, Report on the Tertiary plants of British Columbia, p. 61, text fig. 14, 1908. 

Knowlton, Washington Acad. Sci. Proc., vol. 11, pp. 184, 

212, 1909; Jour. Geology, vol. 19, p. 369, 1911; U. S. 

Geol. Survey Prof. Paper 101, p. 326, 1917 [1918]. 

91 Berry, E. W., U. S. GeoI. Survey Prof. Paper 91, p. 243, pI. 52, 

fig. 1,1916. 

92 Idem, p. 241, pI. 47, fig. 10. 

D8 Idem, p. 242, pI. 52, fig. 3. 

94 I regard this as highly improbable,-E. W. B. 

Berry, U. S. Geol. Survey Prof. Paper 91, p. 249, pI. 48, 

fig. 9, 1916. 

Oarpolithes arachioides Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1872, p. 403, 1873. 

This curious organism was described by Lesquereux 

as follows: 

Capsules or siliques alternate and sessile on flexuous, thick, 

woody pedicels; obovate, rounded to a short acumen, mostly 

enlarged on the lower side, bossed under the point as inclosing a round seed, regularly striated with narrow ridges, generally tending in a curve from the borders to the pOint, obscurely and transversely wrinkled. 

These capsules, or pods, are turned upward in the upper 

part of the branches, horizontal or pending in the lower 

part, 2.5 cenimeters long, 1 centimeter broad in the middle, 

fiattened by compression, but generally convex or infiated 

near the point, or from the presence inside Qf a large, round 

seed. 

Lesquereux stated that "the relation of these racemes of fructifications is as yet uncertain," but suggested that they appear to belong to the Leguminosae 

and in some ways suggest the fruits of Araclvis h'ypogaea, the well-known peanut. 

In the years that have come and gone since Lesquereux wrote Legwminosites (JIf'oohioides has been 

found to be very widely distributed, and although its 

exact nature is still a matter of some doubt, it can 

be described with more assurance. Berry,95 who has 

found it at a number of localities in the Wilcox group, 

has revised the description as follows: 

Pogs in compound clusters, arranged alternately in pairs on 

stout flexuous stems, subsessile, of a ligneous conSistency, full 

and evenly rounded, Inllated, about 2.5 centimeters in length 

and about 1 centimeter, wide across the middle, pointed at' 

both ends, mucronate distad, several-seeded, dehiscent. Surface striated; in general there are two series of striae, wrinkles, or corrugations, one set approximately longitudinal and 

the other transverse. These striations are to a certain extent 

the result of compression, since a good many of the pods show 

rounded bases evidently due to deformation. 

As to the affinity of Legwminosites arachioides, Berry 

concludes that it " was either a low straggling plant 

of the sandy beaches, comparable perhaps with the 

modern forms of Baptisiu; or 01'otailoJria, or else it was 

a vine like the modern species of Abrus." It has not 

been found in recognized association with foliage, 

and we must apparently await for future discoveries 

of this or for a closer interpretation of th~ fruits. 

The type specimens of Lefll1Jn1Jinosites (Jff'achioides 

came from Evanston, Wyo., occurring" in a block of 

ironstone taken from the mines," and are all preserved 

in the collections of the United States National Museum (No. 492). Its presence in the Denver Basin 

and adj acent areas depends on some fragmentary yet 

characteristic specimens at Golden and near the base 

of Mount Bross, in Middle Park. They add nothing 

to our previous know ledge of this curious organism. 

oa Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 250, 1916. 
SYSTEMATIO PALEOBOTANY 99 

Occurrence: Evanston formation (types) , Evanston, vVyo. Denver formation, Golden, Colo. Middle 

Park formation, Middle Park, Colo., at base of Mount 

Bross. Raton formation, near Benvynd, Colo. Fort 

Union formation, Clear Creek, near Glendive, Mont. 

Lance formation, Melville, l\1:ont.; Monarch, Wyo.; 

and T. 23 N., R. 20 E., S. DaIr. Wilcox formation, 

Hamilton, Sabine County, Te~., and M'ansfield, De 

Soto Parish, La. 

Family CAESALPINEACEAE 

Cassia vetusta Knowlton, n. sp. 

Plate 45, Figure 2 

Leaflet small, apparently rather thin, lanceolate, 

broadest near the middle, whence it narrows to the 

rather obtusely wedge-shaped base and to the apparently acuminate apex; nervation pinnate, consisting 

of a relatively strong Inidrib and about three pairs of 

secondaries, the lower pair opposite, the others alternate, at an angle of about 60°, all slightly curved upward; finer nervation not retained. 

This little leaflet-the only one observed-is about 

4.5 centimeters long and 1.5 centimeters wide. It 'is 

rather remarkable in that it has only three pairs of 

secondaries, the lowest pair being opposite and the 

others strongly alternate. 

In size and shape this species is similar to Oassia; 

emo:rginataJ. Berry,06 from the Wilcox group of Mississippi, but differs in the secondary nervation. 

Occurrence: Dawson arkose, Templeton Gap, 4 miles 

northeast of Colorado Springs, Colo., collected by F. 

H. I(nowlton, July, 1910. 

Cel'cis colol'adensis Knowlton, n. sp. 

Plate 45, Figure 5 

Leaf membranaceous, broadly ovate or nearly circular, truncate at the base, probably obtusely pointed 

at the apex; margin entire; petiole short, relatively 

stout; nervation palmately 5-ribbed from tpe 

slightly enlarged top of the petiole; a strong basal 

secondary on each side.arises very near the base of the 

outer pair of ribs, thus producing a false 7 -ribbed appe.arance; midrib much the stronger, especially below, 

with at least two pairs of alternate, strong secondary 

branches in the upper part; inner pair of ribs at an 

angle of about 70°, nearly straight, joining the lowest 

pair of secondaries on the miari~ by a broad loop, 

{:Iach of these' ribs with four or five camptodrome secondaries in the upper part; outer pair of ribs at an 

angle of 40° or 50°, a little curved upward, joining 

t.he lowest secondary branch on the inner ribs, earh 

with several secondaries that are approximately at 

00 Berry, E. W., U. S. Geol. Survey Prof. Paper 91,. p. 233, pI. 48, 

fig'. G, 1016. 

right angles to the midrib, camptodronle and forming 

a series of loops near the margin; nervilles numerous, 

strong, mainly broken. . 

The leaf figured was probably about 8 centimeters 

long and a little more than 6 centimeters wide. The 

petiole is preserved for a length of 1.25 centimeters 

but was probably considerably longer. Another very 

fragmentary leaf thought to belong to this species was 

probably 9 or 10 centimeters wide and doubtless as 

much in length. The nervation is so well shown in 

the figure that description is not necessary. 

Oe'}'ois coloTadensis was of about the same size as 

normal leaves of the living OeTcis canadensis ;Linne 

and much resembles it. It differs in the more nearly 

truncate base, proportionately somewhat narrower 

blade, fewer secondaries on the midrib, and fewer 

secondaries on the ribs, but none of these differences 

are of great importance. 

The present species is of about the same size as 

OeTcis wiloowiana Berry,97 from beds of Wilcox age in 

the Lagrange formation at Grand J unction, Tenn. 

It differs, however, in many of the details of nervation, including the . number and angle of divergence 

of the primary ribs, secondary branches on midrib 

and lateral ribs, and character of the finer nervation. '. Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Spri~gs, Colo., collected by F. H. Knowlton and W. T. Lee, July, 1910. 

Order SAPINDALES 

Family CELASTRACEAE 

Celastrus . gaudini Lesquer~ux 

Plate 45, Figures 3, 4, 9 

OeZast'1'U8 gautJ,ini Lesquereux, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, p. 54, 1888. 

Parrotia fagifolia Ettingshausen. Lesquereux, op. cit., p. 52. 

This species is characterized by Lesquereux .as follows: 

Leaves membranous, rugulose on the surface, oval-oblong 01" 

broadly oval, crenate or obtusely serrate, abruptly narrowed 

at base; secondaries alternate, ,about six pairs, much curved 

upward in paSSing to the borders' and effaced; tertiary nerves 

and nervilles obsolete. 

These leaves are 5.5 centimeters long, 4-.5 to 5 centimeters 

broad, apparently obtuse, though the upper part is broken in 

aU the specimens. The species is closely related to OeZast1'u8 

heerii Sismonda.98 

The specimen referred by Lesquereux to PaYrrotia 

fagifolia; Ettingshausen is not to be distinguished from 

97 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 228, pI. 49, 

fig. 1,1916. 

os Sismonda, Eugenio, Materiaux pour servir A la pal~ontologie du 

terrain tertiaire du Pi~mont: Accad. sci. Torino M'em., vol. 22, p. 449, 

pl. 29, fig. 5, 1865. 
100 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

this species and has been so referred. It is shown in 

Figure 9. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, 

Mass., Nos. 1527, 1528; specimen identified as Pa'rrotia 

fagifoUa Ettingshausen, No. 1512. 

Celastrinites artocarpidioides Lesquereux 

Celastl'inites m·tocarpidioides Lesquereux, The Tertiary flora, 

p. 268, pI. 35, fig. 3, 1878. 

A.1'tocarpidue-m olmediaefoZiumf Unger. Lesquereux, U. S. 

Geol. and Geog. Survey Terr. Ann. Rept. for 1873, pp. 

381, 400, 1874; Idem foi'.1876, p. 516, 1878. 

The type specill).en of this species is No. 407 of the 

United States N aUonal Museum collection. It came 

from the andesitic material at Golden, Colo., and 

hence is referred to the Denver formation. So far as 

known no additional specimens have been obtained at 

this locality. As Lesquereux pointed out, the specimen is not well preserved, and its intimate characters 

are made out with difficulty. It is not correctly shown 

in the figure given in the report on the Tertiary flora, 

certain folds in the leaf having been mistaken for 

secondaries. There are six or seven pairs of alternate 

• secondaries, which apparently arch around just inside 

the margin, and not, as Lesquereux said, "curving at 

a distance from the borders, and forming a double 

series of bows by anastomosis of the nervilles." 

This species is apparently closely related to Oelast1'inites alatus I{nowlton,99 from the Laramie at Erie, 

Colo., at least so far as the imperfection of O. a1'toca1'-

pidioide8 permits a comparison. The two species are 

of about the sanle size and shape, but O. a1'toca1'pidioides is without the winged petiole which is so conspicuous a feature in O. alatus. 

Lesquereux first referred this leaf to A1'tocaYrpidiwm 

olmediaefoliWln? Unger, which it much resembles, but 

Inter decided that its affinity was better expressed by a 

reference to Oelast1'inites. 

Occ~llTence: Denver formation, Golden, Colo., collected by Arthur Lakes. 

Celastrillites populifolius Knowlton, n. sp. 

Plate 40, Figure 1; Plate 45, Figure 11 

Popu,l'/ls 'ltngeri Lesquereux. Lesquereux, Havard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 46, 1888. 

Under the name Popu""/lus 'ungeri Lesquereux mentioned and partly described a leaf in the Museum of 

Comparative Zoology from Golden, Colo. He stated 

that the " specimen referred to this species, the lower 

half of a leaf, apparently rOluld, [is] entire on the borders and with the nervation of the species." 

As the specimen was seen by Lesquereux it appeared 

to be a short, round leaf very much like the figure of 

0:1 Knowlton, F. H., U. S. Geol. Survey Prof. Pa.per 130, p. 148, pl. 

25, fi.e:s. -1. 5; pI. 26, fig. 1, 1922. 

Populus ungeri given in the report on the Tel'tiary 

flora, Plate 25, Figure 5, but by removing the matrix 

it is disclosed as something very different. It is a 

medium-sized leaf about 7.5 centimeters long and a 

little over 4 centimeters wide, with the petiole complete, 2.5 centimeters long. It is ovate, abruptly 

rounded to the slightly decurrent base and apparently 

narrowed to a moderately pointed apex. The margin 

is perfectly entire below, but along the sides it is provided with minute, sharp, upward-pointing teeth. The 

midrib is slender and straight, with about nine pairs 

of thin, opposite or subopposite secondaries, which 

arise at an angle of about 40°, much curved upward 

and disappearing just before reaching the margin. 

The nervilles are very fine,apparently mainly simple 

and more or less oblique to the secondaries. 

The generic reference for this leaf is somewhat difficult to settle. In the first place it somewhat resembles 

the lower half of Populus ungeri, but it is much more 

like certain other species of Popultus-for instance, 

PopUlus mut(J;oiZis ovalis Heel', as figured by Lesquereux 1 from the Evanston formation of Evanston, Wyo, 

In size, shape, and nervation it is- much like certain 

lea ves of mutaoilis ovaUs as figured by Heer 2 fronl 

the Swiss Miocene. The presence of small sharp teeth 

on the margin, however, excludes this leaf from the 

genus PopUlus. 

The reference of this leaf to 0 elastrinites seems 

justified. It has, for instance, the same shape and 

nervation as 0 elastriwites a1'tocGJrpidioides Lesquereux,s a Denver species, but differs in the marginal 

teeth. 

Occurrence: Denver Iormation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Type 

ill Museum of Comparative Zoology, Cambridge, 

Mass., No. 1576 (identified by Lesquereux as Populu.s 

ungeii). 

Family ACERACEAE 

Negundo deCUl'l'enS Lesquereux 

Plate 45, Figure 10 

Negtt-ndo decurrens Lesquereux, Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 54, 1888. 

Described as follows by Le~quereux : 

Leaves compound, trifoliolate. Terminal leaflet apparently 

large, trinerved from above the decurring base; lateral nerves 

thick, branching on the lower side; lateral leaflets at a distance from the base of the terminal one, undulate and sparingly dentate on the borders, trinervate abov.e the decurring 

base; se~ondaries thin with few branches; nervilles indistinct, 

oblique to· the median nerve. 

SpeCies related to Negtt-ndo triloba Newberry,' but differing 

by the prolongation of the base of the lateral leaflets along 

1 Lesquel'eux, Leo, The Tertiary flora, ,po 177, p1. 24, fig. 3, 1878. 

2 Heel', Oswald, Flora tertinria Helvetiae, Vol. 2, p. 22, p1. 61, fig. 1, 

1856. 

8 Lesquel'eux, Leo, The ~'ertiary flora, p. 268, p1. 35, fig. 3, 1878. 

4 Newberry, J. S., U. S. Geol. Survey Mon. 35, p. 115, pI. 31, fig. 5, 

1898. 
SYSTEMATIC PALEOBOTANY 101 

the petiole, not cordate. 'l'he nervation is, however, the same, 

nnd the borders also nre marked by few &harp teeth. 

This species j s based on the single specimen figured 

and consists of fragments of two leaflets. All that 

is retained of the terminal leaflet is a part of the base, 

this being wedge-shaped or decurrent to the strong 

rachis. Its whole outline can not be made out, though 

the presence of strong lateral ribs suggests the possibility of lateral lobes. The lateral leaflets are opposite and are 1.5 centimeters below the decurrent base 

of the upper one. 

Although Lesquereux states that this species is related to Neg~o triloba Newberry, from the Fort 

Union formation of North Dakota, the likeness is not 

,'ery marked. In N. trilobo; the terminal leaflet is 

smaller than the lower ones and is hear.t-shaped instead of wedge-shaped at the base. The lateral leaflets 

in N. triloba are truncate or slightly heart-shaped, 

but in N. decurren8 they are slightly wedge-shaped at 

the base. 

Occurrence: Denver formation, South Table ~{ountain, Golden, Colo., collected by Arthur Lakes. Type 

in ~{t1seum of Comparative Zoology, Cambridge, 

Mass., No. 1523. 

Acersp. 

Plate 45, ll'igure 6 

The fragment here figured is the only specimen that 

can with reasonableness be referred to Acer. It is a 

fragment evidently of a basal lobe with a small portion of the smaller central lobe. The lobes were 

toothed, though the teeth were small and only moderately pointed. The nervation is clearly that of Acer. 

There is so little of this leaf present that it is hardly 

worth comparing with described forms, though so far 

as can be determined it seems to agree closely with 

certain of the leaves from the Swiss Miocene referred 

by Heel' 5 to Acer trilobatwmJ prodiuctumj in fact, it 

would be difficult to separate them on the- material, 

available. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Arapahoe form'ation), 3,000 feet east 'of 

the Douglas mine, Sedalia, Colo., collected by Arthur 

Lakes. 

Family SAPINDACEAE 

Sapindus caudatus Lesquereux 

Sapind1t8 ca1.tdat1.t8 Lesquereux, U. S. Geo1. and 'Geog. Survey 

Terr. Ann. Rept. for 1872, 'pp. 380, 397, 1873; idem for 

1873, p. 382, 1874; idem for 1876, p. 515, 1878; The Tertiary flora, p. 264, pI. 48, fig. 6, 1878; Harvard ColI. 

1\:[us. Compo Zoology Bull., vol. 16, p. 54, 1888. 

The type of this species, now preserved in the 

United States National Museum (No. 386), was well 

1\ FIecl', Oswald, Flora tertlnria Helvetiae, vol. 3, p. 47, pI. 114, figs. 

2, 7, etc., 1850. 

85492-30-8 

described and figllred by Lesquereux. It appears to 

be rare, for in addition to the single example on which 

it is based Lesquereux reported only one in the collection from Golden now belonging to the Museuln of 

Comparative Zo010gy, and the large recent collections 

belonging to the United States National ~{useum include only two or three specimens, but one of them is . 

so complete as to render its determination reasonably 

certain. 

Lesquereux 6 identified a single leaf as this species 

(U. S. Nat. Mus. No. 2601) from the 'Vilcox formation of Campbell's quarry, Cross Lake, Caddo Parish, 

La., but Berry,7 who has recently restudied this and 

other similar material, states that it is so fragmentary 

that it can not be identified even generically. 

In a number of particulars Sapindus caudatus 

strongly resembles Jugla'n,s denveriana. Thus it agrees 

in the shape of the base and largely in the type of 

nervation, but it differs in being relatively shorter and 

broader, in having a long, slender apex, and more particul~rly in having the margin perfectly entire instead 

of more or less obscurely toothed. It is possible that 

a larger series of specimens Inight show a. breaking 

down, but. until this is actually demonstrated it seems 

best to keep them distinct. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo. (type); additional specimens from 

same locality, collected by Arthur Lakes, 1890. 

Sapindus berryanus Knowlton .. n. sp. 

Plate 46, Figures 1, 2 

Leaflets of medium size, thickish, linear-lanceolate, 

, abruptly rounded and slightly unequal-sided at the 

base, apex destroyed but apparently acuminate; margin perfectly entire; petiolule short, stout; midrib 

very strong, straight; secondaries numerous, thin, often 

immersed in the substance of the blade, at a very low 

angle of emergence, considerably curved upward; 

finer nervation not retained. 

This species is represented in the collections by several examples, 'wo of the best preserved of which are 

figured. They are 9 or 10 centimeters long and about 

2.5 centimeters wide and are slightly unequal-sided at 

the base. The apex is not preserved in any of the 

specimens, but to judge from the configuration of the 

upper portion of the best-preserv~d specimens it must 

have been fairly acuminate. 

This species is quite distinct from any thus far 

found in the Dawson arkose. It is suggestive of 

Sapindus knowUoni Berry 8 from the 'Vilcox group, 

but that form differs in its smaller size and narrower, 

more acuminate apex. ' 

6 Lesquereux, Leo, U. S. Nat. Mus. Proc., vol. ll, p. 24, 1886. 

7 Berry, E. W., U.S. Geol. Survey Prof. Pnpel' Dlt p. :!:). H))(i. 

S Idem, p. 274, pI. 63, fig. 6. 
102 FLORA OF THE DENVER AND ASSOOIATED FORMATIONS OF OOLORADO 

Occurrence: Dawson arkose, a quarter of a mile 

east of the Purdon mine, sec. 27, T. 11 S., R. 61 W., 

Colo., collected by G. B. Richardson. 

Sapindus? obtusifolius Lesqu6reux? 

Plate 46, Figure 3 

ScWindAtS obtusifoli~ts Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1873, p. 419, 1874; The Tertiary 

flora, p. 266, pI. 49, figs. 8-10, 1878; The Cretaceous and 

Tertiary floras, p. 235, pI. 48, figs. 5-7, 1883. 

There is s~me confusion regarding the type locality of this species. In the original description it 

was stated to have come from the "top of hills, apparently overlying the coal-bearing strata of Rock 

Springs [Wyoming], seen to the east 5 or 6 miles 

distant," where it was associated with numerous fragmentary plants that were la'ter described as llfusopayllwn compUcatu17~. When Sapindu8 obtusifolius 

was taken up in the report on the Tertiary flora the 

exact locality was given as "8 miles southeast of 

Green River Station, Wyo., in connection with three 

beds of coal referable to the Washakie or Carbon 

group." Still later its presence was noted by Lesquereux 9 in the Badlands of North Dakota in beds 

now known to be of Fort Union age. In this connection it may be well enough to call attention to the 

apparent similarity between this species and Sapindus affinis Newberry, a species that is very abundant 

in the Fort Union formation. With the same habit 

and approximately the same size, the main difference 

seems to be in the apex of the leaflets, this being obtuse 

in the present species and long ac'uminate in Sapindus 

afflnis. 

The specimen which is here figured is referred with 

some doubt to Sapindus obtusifoliu8. It is very silnilar in size and shape to Plate' 49, Figure 11, of the 

report on the Tertiary flora, but differs apparently in 

having a more pointed apex than is usual in the species; unfortunately both figure and specimen are without the extreme portion of the tip. The nervation is 

very similar, however. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo. 

Family ILICACEAE 

Hex? ovata Knowlton, n. sp. 

Plate 46, Figure 4 

Leaf coriaceous, ovate, abruptly rounded to the base, 

acuminate at the apex; margin finely serrate, the teeth 

undulate or sharp-pointed; petiole strong; midr~b 

strong, straight; secondaries four or five pairs, alternate, thin, strongly camptodrome and arching far 

9 Lesquereux, Leo, 1.'he Cretaceous and Tertiary floras, p. 235, 48. 

figs. 5-7, 1883. 

'within the border, each joining the one next above and 

sending from the outside of the loop a t4in tertiary 

branch or nerville to a marginal tooth'; nervilles 

strong, often percurrent. 

This little leaf is all that has been thus far detected 

in these beds. Jt is a thick leaf, regularly ovate, with 

rather small undulate or sharp-pointed teeth. The 

length is about 5 centimeters and the width 3 centimeters. The nervation is peculiar in that it consists 

of a rather strong nlidrib and about five pairs of 

strong, ·distinctly alternate secondaries which make 

broad loops far within the border by joining of a 

lower with the one next above it. 

Occurrence: Denver formation, northwestern base 

of Green Mountain, Golden, Colo. 

Order RHAMN ALES 

Family RHAMNACEAE 

Berchemia lUuItinervis (AI. Braun) Heer 

BeTc~~emia parvifoUa Lesguereux, Am. Jour. Sci., 2d ser., vol. 

45, p. 207, 1868; U. S. Geol. Survey Terr. Ann. Rept. for 

1869, p. 96; reprint, 1873, p. 196; idem for 1870, p. 

382, 1871; idem for 1871, Suppl., p .. 15, 1872; idem for 

1873, p. 382, 1874. 

BeTchemia muztinervis (AI. Braun) Heer. Lesquereux, The 

Tertiary flora, p. 277, pI. 52, figs. 9, 10, 1878; U. S. Geol. 

and Geog. Survey Terr. Ann. Rept. for 1876, p. 517, 1878. 

Knowlton, Washington Acad. Sci. Pro c., vol. 11, p. 213, 

1909; U. S. GeoI. Survey Prof. Paper 101, p. 333, pI. 101, 

fig. 5, 1917 [1918]. 

" The sta tus of this species in American strata is not 

very satisfactory. As indicated by the above syn-

(\nymy, Lesquereux first called this form B erc'hemUi 

parvifolia, but when he prepared his account of the 

Tertiary flora he decided that it was not sufficiently 

distinct from the well-known European species Berchemia multinervis. The earlier specimens to which 

the name BercMmia parvifolia was given are said 

to have come from Raton Pass, N. Mex., but these 

specimens are not now in the United States National 

Museum collections. In 1910, however, additional 

material was procured by W. T. Lee from Raton Pass, 

hence in the type locality for Berchemia parvifolia. 

The American form' at least seems to be closely related to Rhamnus fJoldianus, from which it differs, 

according to Lesquereux, in never exhibiting the lower 

pair of secondaries on the outside. This chai'acter 

"\Vard 10 did "not regard as of sufficient weight to be 

treated as generic," and .it undoubtedly is a character 

of slight importance. But, as Lesquereux said, all the 

leaves of Rhamnu8 goldJiwnus, regardless of size, have 

the lower secondaries branched, and as none of the 

l~aves of Bercaemia show this branching it perhaps 

becomes of generic importance. 

10 Ward, L. F., Types of the Laramie flora: U. S. Geol. Survey Bull. 

37, p. 73, 1887. 
SYSTEMATIO PA:{.,EOBOTANY 103 

:Nluch nlore difficult, it appears to lne, is the task of 

discriminating between B e1'chernJia 1wultine1'vis and the 

small leaves of Rhamnus cle 0 wrnti , which are not infrequently associated with it. Rlumnnus cleourni differs from R. goldiamu8 in being usually larger and in 

not ha,ving the' lower pair of secondaries branched. 

This is a sufficient criterion in large leaves, but it is 

almost'impossible to separate small leaves of RhfJJ1n'l'l/US 

cle7)1(',/lni. from B e1'chemia 1nultinevrvis / in fact, they 

can be distinguished only by the shape, Berchemia 

'In'ltlti'Jw'I'vis being nearly elliptical and Rhammus cleo'l.l1'ni more pointed. 

The recent collections from the Denver Basin include this species fronl the south face of South Table 

Mountain, Golden, and from the vicinity of the Douglas coal mine, west of Sedalia. As they do not add 

anything to our knowledge of this species it has not 

been though~ necessary to figure them. 

Occurrence: Denver formation, south face of South' 

Table Mountain, Golden, Colo., collected by Arthur 

Lakes, 1890. Dawson arkose, lower part (in beds 

equivalent to Arapahoe formation), 3,000 feet east 

of the Douglas coal mine, Sedalia, Colo., collected by 

Arthur Lakes, 1890. Raton formation, Raton Pass, 

Raton Mountains, N. Mex., and near Yankee, N. Mex., 

collected by W. T. Lee. Fort Union formation, mouth 

of Yellowstone River, Mont. 

Rhamnus? praealaternoides Knowlton, n. sp. 

Plate 46, Figures 7, 8 

Rhamnus alatel'noidcs Heel'. Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1873, p.405, 1874; idem for 

1876, p. 517, 1878; The Tertiary flora, p. 278, pI. 52, flgs. 

11, 11a, 1878. 

Leaf small, about 14 millimeters long and 6 millimeters wide, subcoriaceous, elliptical, obtusely pointed, 

narowed fr01n near the middle to the wedge-shaped 

base; lnargin obscurely dentate, apparently only above 

the middle; midrib strong for the size of. the leaf; 

secondaries about five pairs, alternate, at somewhat 

irregular distances, emerging at an angle of about 

450 , considerably curved upward, camptodrome, with 

thin branches passing to the obscure teeth; finer nervation not retained. 

This species, it must be confessed, has not a very 

fil'm or satisfactory standing. It is based entirely on 

the single leaf (U. S. Nat. Mus. 432) identified by 

Lesquereux with Rluim'l'l/U8 alaternoides Heer,l1 from 

the Swiss Miocene. Heer has figured three specimens, 

one of which has entire margins and the others are 

obscurely few-toothed. The leaf from Golden resembles the European leaves closely, but it is more 

nearly elliptical and has the secondaries at a slightly 

11 Heer, Oswald, IJilOl'fi tertlarla Helvetlae,. vol. 3, p. 78, pI. 124, figs. 

21-28, 1859. 

more acute angle'. It lacks a good deal of the actual 

margin, and hence it is difficult to compare closely with 

the Swiss leaves. Considering the geographic and 

stratigraphic difference between these American and 

European leaves it seems very doubtful if there can be 

identity between them; hence the American leaf has 

been .given a new name, which, however, preserves its 

supposed resemblance to Heer's species. The generic 

reference has also been questioned, for it is impossible 

to be certain of the affinity of only a single rather 

poorly preserved leaf, especially as the finer nervation 

. is not observable. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Leo Lesquereux, about 

1.875. 

Rhamnus goldianus Lesquereux 

RhamnU8 golcUanrt~8 Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. Rept .... for 1872, p. 381, 1873; idem for 

18~3, p. 405, 1874; idem for 1876, p. 517, 1878; The 

Tertiary flora, p. 281, pI. 53, figs. ~, 1878. 

Be1'cnemia multinervi8 Al Braun. Ward, U. S. Geol. Survey 

Sixth Ann. Rept., p. 554, pI. 51, flg. 13, 1886; U. S. Geol. 

Survey Bull. 37, p. 73, pl. 33, flg. 2, 1887. 

Leaves thick, subcoriaceous, smooth, entire, broadly 

oblong or oval, obtusely pointed or acuminate at the 

apex, subcordate or rounded at the base; petiole short; 

secondaries numerous, 12 to 16 pairs, close, parallel, 

slightly curved, forming a series of simple bows near 

the margin; lowest pair of secondaries branching 

downward in anastomosing with the short, marginal 

veins; nervilles very numerous, both simple and 

forked, usually more or less oblique to the secondaries. 

This species is closely allied to Rhamnus cleowrni 

Lesquereux, from which it differs in its generally 

smaller size, rounded or heart-shaped base, and downward arching' of the tertiary branches of the lowest pair 

of secondaries. The question of size, however, is not 

one of great importance, for the recent collections contain specimens clearly refera.ble to this species which 

are larger than the figured types of R. cleourni. They 

are, however, generally ~maller. The roundecl base 

and branched lowest secondaries become the important 

characterR, and in considering specimens without the 

base it is a question to which form they should be 

referred. 

The type specimens of RhfJJ1n1/JUJ8 goldianus are all 

preserved in the United States National Museum 

(Nos. 441 to 445) and are all in the typical andesitic 

material of the Denver formation at Golden, Colo. 

Accompanying them and having the same number is 

a small example on a hard reddish sandstone that 

seems to have come from a different locality and to 

belong to a different species, possibly Berche'l11l.ia 

m,wltinervis. In the report on the Tertiary flora (p. 

281) Lesquereux reported. this species from Black 
104 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Buttes, Wyo., "where it is rare," but the National 

Museum does not possess and so far as the catalogue 

goes has never possessed a specimen from this locality. 

It is not contained in the large collections procured by 

'Vard at Black Buttes in 1881, nor was it found by 

me on a visit made jri 1896. Lesquereux said also: 

" One specimen, representing the large form, is from 

Carbon, Wyo." As no specimens can be found from 

t.his locality, either among the original or the later collections it seems more than likely that this remark 

was int~nded to apply to Rhamnu8 cleburni, which precedes it on the page and which has larger leaves than 

Lesquereux ever assigned to R. golcliamu8. 

The N at-ional Museum collection contains another 

fragmentary specimen (No. 2375), from the rhyolite 

beds at Silver Cliff, Colo., identified by Lesquereux 12 

as this species, but it represents only the upper half of 

a medium-sized leaf, and as .shown above this is not 

enough to establish positively the identity. of this 

species. 

In workino' up the flora of the Laramie formation I 

found a single fragmentary specimen probably of th is 

species in beds of this age at Marshall, Colo. It seems 

to be identical with some of the smaller leaves referred 

to Rhamnus goldianu8, but as it lacks the base and 

much of the margin its identification has b~en questioned. 

Specimens from the Raton formation of the Raton 

Mesa region of Colorado are referred with but little 

hesitation to Rharnnu8 goldiamu8. 1S 

The specimens figured by Ward 14 under the name 

Bet)'chemia multinervi8 AI. Braun may be briefly considered. Of these; one, the or~ginal of Figure 13, came 

from Golden, Colo., and is undoubtedly referable to 

Rhawnu8 goZdiamus. It is a small leaf, but it has the 

tertiary branches on the lowest pair of secondaries. 

The other specimen (fig. 12) is from the Fort Union 

for~ation in the Bull Mountains, Mont., where it 

was collected by A. C. Peale. This is a small leaf lacking the basal portion and may be correctly placed; in 

any event it may b~ permitted to remain in this species 

for the present. 

Occurrence: Denver formation (types) , Golden, 

Colo., South Table Mountain, collected by Arthur 

Lakes and others; ~Silver Cliff, Colo., collected by 

Whitman Cross. Raton formation, Wootton and near 

Aguilar, Colo., collected by W. T. Lee, 1909. Laramie ( ~) formation, Marshall, Colo., wooded bluff just 

north of station and highest point in the section, collected by F. H. Knowlton, 1908. 

12 I,esquereux, Leo, U. S. Nat. Mus. Proc., vol. 10, p. 44, 1887. 

18 Knowlton, F. H., U. S. Geol. Sur,vey Prof. Paper 101, p. 332, pI. 

101, fig. 4; pI. 112, fig. 5, 1917 [1918]. 

H Ward, L. F., U. S. Geol. Survey Sixth Ann. Rept., p. 554, pI. 51, 

figs. 12, 13, 1886; U. S. Geol. Survey Bull. 37, p. 73, pl. 33, figs. 1, 2, 

1887. 

., 

[Rha,rnn'u,s goldianus latiol' Lesquereux was described briefiy 

in 1873 (U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 

1872, p. 381, 1873) but was not afterward referred to by its 

author. It came from Golden, Colo., but it can not be placed 

without a knowledge of its matrix.] 

Rhamnus cleburni Lesquereux 

Plate 40. Figure 6; Plate 46, Figures' 10, 11 , 

Rhamwlts cleburni LesqueretUE, U. S. Geol. and Geog. SurYeY 

Ten. Ann. Rept. for 1872, pp. 381, 400, 1873; idem for 

1873, p. 382, 1874; idem for 1876, p. 517, 1878; The Tertiary flora, p. 280, pI. 53, figs. 1-3, 1878; Harvard CoIl. 

Mus. Compo Zoology Bull., vol. 16, p. 55, 1888; U. S, Nat 

Mus. Pr,oc., vol. 10, p. 44, 1887; vol. 11, pp. 20, 24, 1888. 

Knowlton, U. S. Geol. Survey Bull. 204, p. 80, 1902; U. S. 

Geel. Survey Prof. Paper 101, p. 332, pI. 113, fig. 3, 1917 

[1918]. 

Berry, U. S. Geol. Survey Prof. Paper 91, p. 283, 1916. 

Oar-nus st'ltde1i Heel'. Lesquereux,The Tertiary flora, p. 244, 

pI. 42, flgs. 4, 5, 1878; Hm'yard ColI. Mus . .comp. Zoology 

Bull., vol. 16, p. 52, 1888. 

Rharnnites knowltoni Berry, U. S. Geol. Survey Prof, Paper 

131, p. 16, 1922 [part]. 

The figured types of this species are preserved in 

the United States National Museum (Nos. 356a: 439, 

440). There are also a large number of ot.her specimens that have been collected in recent years. This 

species is undoubtedly most closely related to Rh(J'7nnU8 gold'iamus, from which it differs in being usu~.ll~ 

larO'er more nearly elliptical, with a tapering pOInt, b , d . I and narrowed to a comparatively long, slen er petlO e. 

The most important difference and one that appears 

to be constant is the absence of branches on the lower 

si de of the basal pair of secondaries. This is really 

the crucial test of the species, for there are leaves of 

Rhamnu8 golilUDnu8 as large as small leaves of R. 

cleb~!trni. It appears plain that the base of th( leaf 

must be present in order that these species may be 

separated with certainty, and the identifications of 

one or the other of them made without knowing this 

character are at least open to question. 

It is also to be noted that very small leaves of fl. 

clebWf'11fi are difficult if not impossible to distinguish 

from large leaves of B e1'che17m(f) 1nultinervis, as pointed 

out under the latter species. 

The question of the generic reference of these two 

species (R. cleburni and R. goldianu8) is still .open. 

According to Lesquereux European paleobot.amstspresumably Saporta and Heer-to whom he had submitted either specimens or drawings inclined to the 

opinion that they showed more affinity with th~ Eu-. 

phorbiaceae or Moraceae, especially with the genus 

FiCfUs and its allies. Take, for example, cert.ain leaves 

of Ficus irre[jU)laris as figured by Lesquereux,15 They 

have the same general shape, much the same cha,ra.cter 

of secondary nervation, and particularly the numerous, 

15 Lesqnereux, Leo, The Tertiary fiora, ~. 196. pI. 34, figs. 4,5,7, ]878. 
SYSTEMATIC PALEOBOTANY 105 

close ne.ryilles. It would seem that all these leaves 

are congeneric-that is: if those above 111entioned 

ha va been correctly referred to Fi()Us those under 

consideration should also be so placed-but so long 

as it is well understood that they occupy· this 

somewhat anomalous position there can be little 

objection. 

Aside from the original locality there are· a number 

of other localities from which Rha1nn'lJ)8 cleburni has 

been reported. Thus a single specimen is present in 

the United States National Museum (No. 2373) from 

the upper I(anab Valley, Utah, as identified by Lesquereux.10 This leaf is much more slender than the 

typical form and it lacks the base. It has the numerous close secondaries of R. oleb~l,rni, and to. judge from 

the general shape is more likely to be· this species than 

R. goldialfw,s, but, as above stated, it can not be positively identified without the base. 

I haye also reported 1i the doubtful presence of R. 

ClebUfl'ni in the lower part of the Clarno formation of 

Cherry Creek, Oreg.; but, as the specin1en is only the 

upper p0L'tion of the blade, it is impossible to make the 

identification positive. 

Among the specimens recently obtained from MidcUe Park, Colo., I find a number of leaves that seem 

to belong to this species, but in the absence of the 

basaL portion the assignment can not he positively 

made. 

RhC1Jmlnus olebul'ni was identified by Lesquereux 18 

from the Wilcox formation of Louisiana, and its presence is confirmed by the later work of Berry,tlJ though 

he considers it to be rather rare. 

Among the collections from the Dawson arkose 

along Jimmy Camp Creek there are a number of fairly 

well preserved leaves that undoubtedly belong to 

Rhamnus oleburnri. 

AfteL' careful consideration I have reached the conelusion that the specimens from Golden, Colo., that 

were referred by Lesquereux 20 to Oorn'l.l-8 8t~tderi? Heer 

should be referred to Rlut1n·nus cleburni. Both of the 

specimens figured by Lesqu~reux as the American types 

of Oornus stucle'l'i are preseryed in the United States 

National Museum (Nos. 355; .356). They are both 

somewhat fragmentary, yet are sufficiently well preserved to show their general character. Both came 

from the andesitic beds at Golden and are like a number of more or less fragmentary specimens obtained 

from tlle same beds in recent years. In his account 

of the Tertiary flora (p.244) Lesquereux gives Evanston, Wyo., us the principal locality for Oornus studeri, 

awl this locality appears to·have been the one at which 

~1 Lcsquereux, Leo, U. S. Nat. Mus. Proe., vol. 10, p. 44, 1887. 

IT Knowlton, F. H., U. S. Geo1. Survey Bull. 204, p. SO, 1902. 

I'! Lesquereux, Leo, U. S. Nat. Mus. Pi·oe., vol. 11, p. 24, 1888. 

1) Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 283, 1916. 

:-I Le~f\u~l't'ux, Leo, Tht' Tertiary flora, 1). 24.4, pI. 42, figs. 4, 5, 1878. 

the species was first detected in American strata,21 

but I have not been able to find any specimens fron1 

this locality in the United States National Museum in 

either the old or the recent collections. 

As the status of 0 ornus studeri Heer as an American 

plant has recently been brought in question by 

Berry,22 it may be of value to' set forth the facts as at 

present understood. Oornus ~tuderi was described by 

Heer 23 from the so-called Swiss Miocene, where apparently it is a not uncommon form. It has since been . 

found at a number of other European ~ocalities. 

The supposed presenc~ of Oornus studeri in North 

America was first made known by Lesquereux 24 in 

1871, when he reported it from Evanston, Wyo. This 

was based on a single large leaf "of which the base 

was destroyed, but whose form and peculiar nervation 

are in accordance with the characters of this species." 

It was said to be similar to Figure 18 of Heer's plate 

above cited. This specimen is not now known to be in 

existence, nor has it been duplicated in subsequent 

collections at this locality. 

The next mention of this species by Lesquereux, except incidental mention in lists, was in 1878 in the 

report on the Tertiary flora. He gave figures of two 

specimens, both of which are preserved in the United' 

States National Museum (Nos. 355, 356), and both 

are on the andesitic material characteristic of the 

Denver formation at Golden, Colo. They are rather 

fragmentary, the larger one especially, which could 

hardly have been less than 16 centimeters in length 

and 8 centimeters in width when growing. This leaf 

is very much larger than any figured by Heel', but 

Lesquereux mentions leaves from Golden that are only 

6.5 centimeters long. The Swiss specimens range 

from about 5.5 centimeters to 12 centimeters in length 

and from 2.25 to nearly 5 centimeters in width. Lesquereux mentions the presence at Golden of numerous 

fragments of this species, and in his report on a collection studied for the Harvard College Museum of 

Comparative Zoology, in 1888, he records two additional specimens. 

vVard 25 identified as Oornus studeri a very perfect 

leaf from the Mesa verde formation at Point of Rocks, 

'Vyo., and in 1900 I admitted this species in the Montana flora and figured another leaf from Point of 

Rocks. 26 A priori it seems improbable that this Euro21. cr. u. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1871, p. 293, 

1872. 

~ Berry, E. W., U. S. Geol. Survey Prof. Paper 131, p. 16, 1922. 

~3 Heel', Oswald, Flora tel'tiaria Helvetiae, vol. 3, p. 27, pI. 105, 6gs. 

18-21, 1859. 

2-1 Lesquereux, Leo, U. S. Geol. and Geog. Survey Terr. Ann. Rept. 

for 1871, SuppI., P. 9, 1872. 

25 Ward, L. F., U. S. Geol. Survey Sixth Ann. Rept, p. 553, pI. 48, fig. 

1, 1886; U. S. Geol. Survey Bull. 37, p. 55, pI. 26, fig. 1, 1887. 

:.'6 Knowlton, F. H., U. S. GeoI. Su\,vey Bull. IG3, p. 68, pI. 15, fig. 3, 

1900. 
106 FLORA OF THE DENVER 4.ND ASSOOIATED FORMATIONS OF OOLORADO 

pean Miocene species should, in America, extend downward into the Montana group, but it is not, easy to 

separate them. The leaf figured by Ward has a much 

thicker petiole and midrib than the specimens f~om 

the Denver beds at Golden, and the specimen figured 

by me is certainly different. It is relatively larger 

and narrower and has 3; different type of finer nervation. The Point of Rocks leaves identified as Oomus 

stuaeri are not accepted as valid at present, pending a 

fuller study of further material. 

As Lesquereux pointed out, and as further emphasized by the question mark placed by him after the 

species, there has always been doubt as to the correctness of identifying Oorn1JJ8 studeri in American rocks. 

They are in'genera'! much larger leaves than the European specimens and differ in having the nerves branch 

near the border. According to Schenk,27 there is also 

some doubt as to the generic reference of certain of the 

leaves usually placed in Oomus, especially O. studeri. 

The American leaves strongly suggest leaves that have 

been placed in Ficus. Thus they may be compared 

with certain ieaves of Ficus denverif11na Cockerell, F. 

irregulwris Lesquereux, F. U1UJa;ta Lesquereux, and F. 

artocaYrpoides Lesquereux. All things considered, it 

'seems best to refer the two leaves from Golden figured 

by Lesquereux 28 as Oomus studeri Heer~' to Rhamnus 

cleburni, as already stated. The smaller of these leaves 

(Lesquereux's,fig. 4) is clearly indistinguishable from 

Rlwmunus cleburni, but the larger specimen (fig. 5) is 

much larger than any previously noted, being about 16 

centimeters long and 8 centimeters wide. 0 

In 1917 I referred the upper portion of a large leaf 

from the Raton forma-tion to Oornus studeri Heel' ~ as 

then currently accepted, some of which it closely resembles.29 However, it is not identical with Rha'l11ll1fU;8 

clebwrni, and if retained in o omus , which it apparently should be, it should r~ceive a new name. 

The status of the American leaves referred to Oor'n'1JJ8 

stiideri has recently been reviewed by Berry,80 who 

concluded that none can be considered conspecific with 

the European leaves, and he then proceeds to give them 

a new name-Rhmmmif}es knowltoni. According to 

the synonymy cited, this proposed new species is made 

to include the original leaves from the Denver formation at Golden, Colo., the several leaves from the 

Mesaverde formation at Point of Rocks, Wyo.-which, 

a~ I have pointed out above, should probably be separated as a new species-and the leaf from the Raton 

formation of southern Colorado. This Raton specimen is probably a Oornus and should also have a new 

~1 Schenk, August, Die fossilen Pfianzenreste, p. 236, 1888. 

2.~ Lesquereux, Leo, The Tertiary fiora, p. 244, pI. 42, figs. 4, 5, 1878. 

20 Knowlton, F. H., U. S. Geo!. Survey Prof. Paper 101, p. 342, pI. 

109, fig. 2, 1917 [1918]. 

30 Berry, E. W., U. S. Geol. Survey Prof. Paper 131, p. 16, pI. 12, 

fig. 7, 1923. 

name. Thus we have left for consideration only the 

specimens first identified as 0 orn'1Ji8 studeri by Holo lick 81 and Berry 82 from the Wilcox formation of 

Louisiana. These southern specimens are in general 

very much larger than the northern ones, sometimes 

reaching a length of 19 centimeters and a width of 9.5 

centimeters, according to Berry. He may be right in 

removing these leaves from Oomu8, but to judge from 

the figures gi ven they are certainly not conspecific 

with the leaves from Golden now referred to Rham~nus 

cleburni. Berry's description of RhOlJ11lJ1:ites knowZton~ reads as follows: 

Leaves large, broadly ovate, many of them slightly inequilateral, the tip somewhat narrowed and acuminate and the 

base rounded or very broadly pointed. Margins entire, in 

some specimens faintly undulate. Texture subcoriaceous. 

Midrib stout and prominent. Secondaries stout, six to nine 

pairs, diverge from the midrib at irregular intervals at angles 

of about 45°, camptodrome. Tertiaries [nervilles] thin. closely 

spaced, percm:rent, typically rhamnaceous. 

These southern leaves are thus se-en to differ from 

those of RhmrilJJwi8 clebu1'ni by their generally larger 

size, broadly ovate shape, rounded, obtuse base, fewer 

secondaries, and apparently slightly different nervilles. 

For the present, therefore, it would seem best to re- ' 

strict the application of Rha'l11lnites knowltoni to the 

leaves from the Wilcox deposits. 

Occurrence: Denver formation (types) , Golden, 

Colo. Dawson arkose, east bank of Jimmy Camp 

Creek 1 mile above Richfield Springs ranch house, 9 

miles east of Colorado Springs, Colo., collected by 

F. H. Knowlton, July, 1910. 

Rhamnus crenatulus Knowlton and Cockerell 

Rham'l1JUs CYrenatulus Knowlton and Cockerell, U. S. Geol. 

Survey Bull. 696, p. 545, 1919. 

Rhamnus - crenatu8 Lesquereux, Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 55, 1888. [Homonym, Siebold 

and Zuccalini, 1843.] 

Following is Lesquereux's original description and 

discussion of this species: 

Leaf large, ovate-Ianceolate, acuminate, rounded or subcordate 

at the base (broken), minutely crenate; lower lateral nerves 

more open, the upper more oblique, all much curved in passing 

toward the borders, and in<!lining to the midrib; the two highest pairs aerodrome; nervilles numerous, close, parallel. 

This fine leaf is 12 centimeters long (base and apex broken), 

6 centimeters broad a little above the base. It has the same 

form, size, and nervation as RhOlYlvnus grosse-serratus Heel' 

(Flora von BornsHidt, p. 20, pI. 4, fig. 10), differing by the 

borders being crenate and the nervilles less distinct. 

This species should be in the Museum of Compara-' 

·tive Zoology, but I was not able to find it, and it must 

stand as left by Lesquereux. 

81 Hollick, Arthur, Louisiana Geol. Survey Rept. for 1899, Special 

Rept. 5, p. 286, pI. 45, fig. 2, 1899. 

82 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 331, PI. 68, 

fig. 3, 1916. 
SYSTEMATIC PALEOBOTANY 107 

Occurrence: Denver formation, Golden, Colo., collected by Arthur Lakes. Type in Museum of Comparative Zoology, Cambridge, Mass. 

Rhamnus cannoni Knowlton, n. sp. 

Plate 46, Figure 5 

Leaf of very firm texture, oblan~eolate, narrowly 

w'edge-shaped from a point apparently above the middle (apex destroyed) ; margin perfectly entire; midrib relatively strong, straight; secondaries numerous, 

probably a dozen or more pairs, strong, alternate, 

emerging at an acute angle, passing upward for relatively long distances, camptodrome or just reaching 

the border; nervilles numerous, strong, approximately 

at right angles to the secondaries, often forked or 

broken. 

Unfortunately the only example found is the one 

here figured, and this lacks considerable of the upper 

part. It is narrowly oblanceolate with a long, narrowed base and entire margin. The length was probably about 6 centimeters and the width about 2.25 

centimeters in the broadest part, which is apparently 

somewhat above' the middle. 

As this is an imperfect leaf it perhaps should not be 

described as new, yet it seems to be fairly well characterized and, moreover, does not agree entirely ·with 

other described forms. It approaches in size and 

shape Rlwnnnus obovatu8 Lesquereux,88 but it differs 

in having fewer secondaries and these at a much more 

acute angle. It may possibly be a very small, unusually narrow leaf of Rha»nnus cleburni, but I do not 

recall having seen one so small. 

Occurrence: MiddI"e Park 'formation, Kinney Creek 

near Hot Sulphur Springs, Middle Park, Grand 

County, Colo., collected by George L. Cannon, of 

Denver, for whom it is named. . 

Rhamnus salicifolius Lesquereux? 

Plate 46, Figure 12 

Rham'l1tU8 8aZioifoZitt8 Lesquereux, Am. Jour. SeL, 2d ser. vol. 

45, p. 206, 1868; U. S. Geol. Survey Terr. Ann. Rept. for 

1869, p. 196, 1869; idem for 1872, p. 400, 1873; U. S. 

Geol. and Geog. Survey Terr. Ann. Rept. for 1873, p. 

382, 1874; idem for 1876, p. 517, 1878; The Tertiary 

fiora, p. 282, pI. 53, figs. 9, 10, 1878. 

Knowlton, U. S. GeoI. Survey BUll. 163, p. 70, 1900; U. S. 

Geol. Survey Prof. Paper 101, p. 271, 1917 [1918J; 

U. S. GeoI. Survey Prof. Paper 130, p. 154, pI. 15, fig. 4; 

pI. 19, fig. 2b, 1922. 

The early history of this species has been given at 

length in Professional Paper 130 and need not be 

repeated here except in the merest outline. The type 

specimen came from the Laramie at Marshall, Colo. 

33 Le~fJUel'cux, Leo, The 'l'ertlary flora, p. 281, pI. 54, fig. 1, 1878. 

This specimen is the original of Figure 9 of Plate 53 

of the report on the Tertiary flora and is No. 446 

of the United States National Museum collection, 

though it is now lost or misplaced. The other' figured 

type (Lesquereux's pI. 53, fig. 10) is said to have come 

from Black Buttes, Wyo., but its exact status is not 

known. It has never been in the National Museum, 

where theoretically it should be, and this species has 

not been again collected at Black Buttes, and consequently its standing is problematical. A study of the 

two figures as given by Lesquereux discloses a number 

of m,inor differences that might be sufficient to separate them. Thus the leaf of Figure 10, said to be 

from Black Buttes, is much narrower, is more acutely 

pointed at the apex, and has fewer secondaries at a 

more acute angle than that shown in Figure 9. 

A specimen of Rhairrlfnws salicifolius was reported 

by Lesquereux from the roof or a coal mine on Sand 

Creek, east of Denver. This is No. 935 of the National 

J\lIuseum collection. It is much larger than the largest 

of the figured specimens, being 11 or 12 centimeters 

long and between 4 and 5 centimeters wide, whereas 

the type is only 9.5 centimeters long and 2 centimeters 

wide. This should probably be separated as a different spec,ies. 

In the recent revision of the Laramie flora a number of specimens of Rha}rnrn/w8 saliciJfolius have been 

found in the Laramie formation of the Denver Basin, 

notably at Cowan station, near Denver, and in a 

cut on the Denver & Salt Lake Railroad near Lyden 

Gulch. This species has also been identified, though 

perhaps not always correctly, at a number of earlier 

horizons, as the;yVermej 0 formation near Rockvale and 

2 miles west of Trinidad, Colo., and the Montana 

group (Mesaverde formati()n)' near Harper station, 

on t;he Laramie Plains, and near the Van Dyke coal, 

Rock Springs, Wyo. 

In the collection from Templeton Gap, near Colorado Springs, occurs the specimen here figured, which 

appears to belong to Rharnnus salicifolius. It is 

rather fragmentary, and the identification is not positive. So far as can be made out it appears to agree 

in size with the larger of the type specimens (Lesquereux's pI. 53, fig. 9), but the nervation is more like 

that of the smaller type figured by Lesquereux. 

Occurrence: Dawson arkose, Templeton Gap, 4 

miles northeast or Colorado Springs, Colo., collected 

by F. H. I(nowlton, 1910. Laramie formation (type), 

Marshall, Colo.; Cowan station, 10 miles south or 

Denver; cut on Denver & Salt Lake Railroad near 

Lyden Gulch, Colo. Vermejo formation, Canon City 

field and near Trinidad, Colo. Montana group (Mesaverde formation), Harper station, Wyo., and near the 

Van Dyke coal, east of Rock Springs, Wyo. 
108 FLORA OF THE DENVER AND ASSOCIATED 'FORMATIONS OF COLORADO 

Paliurus coloradensis Lesquereux 

Pal'iur'U8 aolol'aaoosis Lesquereux, Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 55, 1888. 

Leaf small, obovate, obtuse, denticulate above, triple-nerved 

from above the base, with a single pair of opposite secondary 

veins above the middle of the leaf, parallel to the basilar ones, 

acrodrome and branching outside. 

The top of the leaf is somewhat obHtei'ated; its nervation 

is much like that of Pali'UJru8 O'Vo'iaeu8 (Goppert) Heer.M Its 

size is intermediate betw~en that of Figul'es 58 a,nd 59, but it 

greatly differs by its obovate form and the position of the 

intermediate pair of secondary nerves, as thick as the primaries and parallel to them. 

3' Heer, Oswald, Flora tertinria Helvetiae, vol. 3, p. 76, pI. 121, figs. 

58, 59; pI.' 122, fig. 3, 1859. 
SYSTEMATIC PALEOBOTANY 109 

Zizyphus dallhllogenoides Knowlton 

Plate 48, Figures 5, 6 

Zi:::tIPI"us dapl/lltogfflloides Knowlton, U. S. Geol. Survey Prof. 

Papel' 134, p. 91, 1924. 

Da.I>hnogene a,ngUcaf Heel'. Lesquereux, U. S. Geol. and Geog. 

Survey Terr. Ann. Rept. for 1873, p. 401, 18'74; idem for 

1876, p. 510, 1878; The Tertiary flora, p. 222, pI. 37, 

fig. 9, 1878. 

Zizljph;zts fUwll 08-U8 (Lesquereux) Lesquereux, The Tertiary 

flora, pI. 52, fig. 6 [not figs. 1-5], 1878. 

Leaves of firm texture, ovate-Ianceolate, abruptly 

rounded 01.' very obtusely wedge-shaped at the base, 

long and narrowly acuminate at the apex; margin entire, perhaps slightly undulate; petiole very thick, the 

principal ribs descending into it for some distance below the margin of the blade; nervation peculiar, 

strong, 3-ribbed, the ribs arising well down in the 

petiole, the middle one slightly strongest, straight, 

ending in the ext.reme tip; lateral ribs at a very acute 

angle, arching outward but slightly, then inwa.rd and 

running nearly or quite to the tip; midrib without 

secondary branches, lateral ribs with five or six secondary branches on the outside, the lowest nearly 

basal, all at relatively acute angles, arching and passing high upward, camptodrome; lower secondary with 

a few short, camptodrome tertiary branches on ·the 

outside; nervilles numerous, strong, parallel, unbroken, 

mainly at right angles to the midrib. 

Types: United States National ~{useUln Nos. 317 

(identified as Da;pl~nogene anglica) , 430 (identified as 

Zizyp-hus fibrillos'UJ8), 51066 (with manuscript identification as Zizypl~us 'raincou'rtii var.). 

The species is represented by some half a dozen 

more or less perfect examples from the Denve'r formation at Golden, which together afford a pretty complete knowledge of its size and form. There is considerable range in size, the smallest being the specimen described and figured by Lesqllereux as Daphno~ 

gene anglic(()? Reel' (" Tertiary flora," pI. 37, fig. 9), 

which is 6 centimeters long and only 1.6 centimeters 

wide, and the largest being fully 12 centimeters in 

length and about 6 centimeters in width. The petiole 

is preserved for about 1.5 centimeters and is evidently 

not compl~te. The outline also differs somewhat, ranging from narl'owly ovate-Ianceolate to broadly ovatelanceolate. The base is in no specimen heart-shaped. 

The apex is long and narrowly acuminate. The primary nervation is strongly marked and is peculiar in 

that the three ribs arise well down in the petiole and 

far below the lower margin of the blade. It is really 

only 3-ribbed, but the lowest pair ~f secondaries SOlnetimes arise so low down as to make it appear 5-ribbed. 

The solitary leaf identified by Lesquereux as Daphnogene ooglica? Reel' is present in the United States 

~ ational l\{useum collection (No. 317) and has long 

been a puzzle. Its identification with this European 

species rested on extremely insecure grounds, as D. 

anglica was never figured, Lesquereux making the 

identification on Reer's short description alone. It 

has remained unique in American strata until the 

present time; but although much smaller and narrower than the typical form appears to be, it is so 

absolutely similar in everything except size that I 

have no hesitation in placing them together. 

The question of relationship has to be further con-

&idered. Lesquereux was evidently in doubt as to the 

propriety of referring it to Daphnogene and stated, 

" It. is therefore probable that, as Saporta supposes, 

this leaf may represent a ZizyphulS or a Oeanothus," 

und such seems to me to be the case. The little narrow 

original specimen has seemingly a rel:ationship to 

Zizyphus vulgaris Linne, the common Ii iring form, 

though this differs in having serrate margins and in 

lacking the strong branching on the outside of the 

lateral ribs. But in the light of the additional material now referred to this form it appears beyond 

question to be most closely related to Zizyphus 

fibrillosus Lesquereux; in fact, the line separating 

them is very uncertain. Unfortunately most of the 

leaves of Z. fibrillosus are so fragmentary that it is 

difficult to q.etermine the shape of the base; it appears, 

however, that this is always more or less deeply heartshaped, whereas in the form under discussion the base 

is truncate or obtusely wedge-shaped. The apex is 

also uncertain in many of the examples referred to 

Z. fibriZlosus, but in none does it appear so long or 

narrowly acuminate as in the specimens under conE-ideration. The nervation is certainly very much of 

the same type in both, and it may be that subsequent 

events will show that Z. daphnogenoides is not a wellfounded species, though -in the light of present understanding it seems a far call from the narrow leaf 

figured by Lesquereux in the report on the Tertiary 

flora (pI. 37, fig. 9) to the large, extremely broad 

leaves of typical Z. fibrillos'lt8. 

This is also to be compared with Zizyphus disto1,tus 

Lesquereux, which is really almost too close to warrant 

separation, and if the type and only examples of 

Z. distortus were better preserved or reinforced by 

additional material it is possible that all three would 

be thrown together. 

I have also referred to Zizyp7vus daphnogenoides, 

one of the six specimens figured by Lesquereux in 

his account of the Tertiary flora as Zizyphus fibrillosus-the one shown in his Plate 52, Figure 6. This 

specimen (No. 430, U. S. Nat. Mus. r represents the 

llearly complete apical portion of a long, narrow leaf 

that was probably about 7 centimeters long and not 

much if any over 3 centimeters wide. ,It lnust have 

been narrowed or very obtusely wedge-shaped at the 
110 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

base and is certainly quite unlike the broad truncate 

or heart-shaped leaves normal for Z. fibrillosus. ' 

Among the specimens from the Denver formation 

acquired by the United States National Museum from 

the Lacoe collection is one labeled by Lesquereux as 

an undescribed variety of Zizyphus raincourtii (Sa-

,porta} Saporta. This specimen (No. 51066, U. S. 

N at. Mus.) is a rather fragmentary basal portion of 

a small leaf that app'ears indistinguishable from certain of the leaves now placed in Z. daphnogenoides. 

Saporta's species,37 which is from the Eocene of 

Sezanne, is of about the same size as the leaf under 

consideration but differs in a number of marked particulars. Thus it is 3-ribbed from the top of the 

petiole, but the lateral ribs do not ascend much above 

the middle of the blade, and the middl\,' rib has numerous secondary branches in the upper part. The margin of the leaf is denticulate instead of entire. The 

Lacoe specimen is undoubtedly referable to Zizypnus 

daphnogenoides. 

The Lacoe collection also contains another specimen that must be referred to Zizyphus daphnoge-

"noides. This specimen (No. 51063) was labeled by 

Lesquereux "Smilaw sp.," but it is indistinguishable 

from the ordinary leaves of Z. daphnogenoides, such 

as that figured by Lesquereux in the report on the 

Tertiary flora (pI. 52, fig. 6) as Zizyphus fibrillosus. 

After the above account of the Denver leaves was 

written Zizyphus daphnogenoides was found, in the 

Animas formation, near Pagosa Junction, Colo. This 

identification was based on a very perfect leaf that 

lacks only the extreme tip. It is, narrowly ovatelanceolate, about 6 centimeters long and 2 centimeters 

wide, and has the complete'petiole, 2 centimeters long. 

This leaf is identical in all particulars with the Denver leaf identified by Lesquereux as Daphnogene anglida? Heer, and there can be no doubt" as to their 

being the same. 

As the Animas report (Professional Paper 134) was 

to be published first the description of this species as 

well as most of the discussion was transcribed for that 

report. 

Occurrence: Denver formation, Golden, Colo. The 

leaf figured by Lesquereux as Daphnogene G/ngliea? 

Heer was collected in the seventies by Capt. Ed. 

Berthoud; the specimen removed from Zizyphus 

fibrillosU8 was collected about 1872 by Leo Lesquereux ; 

th~ additional examples :were collected by Arthur. 

Lakes on the south face of South Table Mountain, 100 

feet below the lava cap. Animas formation, Pagosa 

quadrangle, half a mile west of Pagosa Junction, 

Colo., collected by J. B. Reeside, jr., 1921 (lot 7496). 

87 Saporta, Gaston de, Prodrome d'une fiore fossile des travertins 

anciens de S~zanne: Soc. g~ol. France M~m., 2d ser., vol. 7, p. 414, 

text fig. 22, pI. 3'5 (14), figs, 8-10, 1868. 

Zizyphus beckwithii Lesquereux 

Plate 47, Figure 3 

Zizyphu,s becktwithii Lesquereux, The Cretaceous and Tertiary 

floras, p. 125, pI. 19, fig. 5, 1883. 

Type: United States National Museum, No. 1557. 

This species, so far as now known, rests on the 

single example originally described and figured. It 

was characterized by Lesquereux as follows: 

Leaf membranaceous, oval or obovate, rounded at the top, 

narrowed and decurrent to the petiole, palmately trinerved 

from the base j medial nerve narrow, with a single branch 

above the middle, the lateral Curving up at a distance from 

the borders, nearly aerodrome, much branched outside; nerv11les close, distinct, at right angles to the midrib. 

The length is about 4.5 centimeters and the width 

a centimeters. It has a thick petiole a little more 

than 1 centimeter long. 

The basal portion of this leaf is not distinguishable 

from certain leaves of Zizyphus daphnogenoides, as 

already described in this work. The apical por,tion, 

however, is very different. Instead of being prolonged into a slender, acuminate tip it is abruptly 

" rounded and obtuse above. The nervation is of precisely the same type as that in Zizyphus daphnogen~des, even to the single secondary branch on the 

upper portion of the midrib. It may be that the leaf 

of Zizyphus beckwithii is abnormal as regards the 

configuration of th~ upper portion, though there is 

no direct evidence of this condition. As it stands it 

is perfectly distinct from either Zizyphus daphnogenoides or Z. fib rill OSU8, to both of which it is undoubtedly related, and it must be left to the future 

to determine whether it is normal and a good species 

or abnormal and referable to one or the other of the 

obviously related forms. 

Occurrence: Denver formation, Golden, Colo., collected by H. C. Beckwith, in whose honor it was 

"named. 

Zizyphus distortus Lesquereux 

Plate 47, Figure 2; Plate 52, Figure 1 

Zizypkus distortU8 Lesquereux, U. S. Geol. and Geog. Survey 

Terr. Ann. Rept. for 1873, pp. 382, 404, 1874; idem for 

1876, p. 576, 1878; The Tertiary flora, P. 275, pI. 51, flgs. 

8, 9 [not fig. 7, which=Zizypkub' fi,brillo8us], 1878. 

Leaves of small or medium size, membranaceous, 

elliptical or oblong, rounded below to an obtusely 

wedge-shaped or truncate base, apparently obtuse 

above; margin entire; petiole about 1 centimeter 

long, stout; primary nervation palmately 3-ribbed (or 

falsely 5-ribbed) from the top of the petiole, the midrib slightly the stronger, with several pairs of secondary branches in the upper part; outer ribs nearly as 

strong as the midrib, at an angle of 60° or 70° ~ not 
SYSTEMATIO PALEOBOTAWY 111 

much curved, lost in the upper third of the blade, each 

with several secondary branches on the outside, the 

lowest ones arising so near the top of the petiole as to 

give the appearance of being true ribs-hence the 

5-ribbed effect-these secondaries apparently all camptodrome ;.nervilles numerous, strong, mainly unbroken, 

and at right angles to the midrib .. 

Two of the types of Zizyphus distortus-the originals of Lesquereux's Figures 8 and 9-are preserved in 

the United States National l\1useum (Nos. 434, 435). 

The original of Figure 7 is not and, so far as the 

records show, has never been in the Museum collection. 

To judge from the figure alone it appears indistinguishable from leaves of Zizyphus fibril108Us Lesquereux, and it has therefore been transferred to that 

species. 

It thus appears that Zizyph,us distortus was founded 

on two broken specimens which up to the pr.esent time 

uJ.Jpear to be all that have been found. They are in 

effect 5-ribbed, tho)lgh the lower or outer pair of ribs 

are thinner than the others and do not branch. In the 

collection from the Dawson arkose along Jimmy Camp 

Creek I find two examples, both of which are here figured, that seem to belong to this species. They appear 

to be a little larger than the types and add considerably to our knowledge of the species. The best-pres('rved example (pI. 52, fig. 1) must have been 10 

centimeters or more in length and nearly 8 centimeters 

in width. It has the complete petiole, which was' 

about 1 centimeter long.' The other example (pI. 47, 

fig. 2) was only. 8 or 9 centimeters in length and 6 

or 7 centimeters in width. The nervation is well 

brought out in the figures and need not be further. 

described. 

Zizyphus disto'rtus belongs to the group typified by 

Z. fib'J"illosus but differs by its wedge-shaped instead 

uf more or less heart-shaped base, by the lateral ribs 

little or not at all curved upward. 

Occurrence: Denver formation (types), South Table 

Mountain, Golden, Colo.. Dawson arkose, east bank 

of Jimmy Camp Creek 0.6 mile above Richfield 

Springs ranch house, 9 miles east of Colorado Springs, 

Colo., collected by F. H. I(nowlton and W. T. Lee, 

July, 1910. 

Zizyphus lesquereuxii I{nowlton 

Ziz1Jph1tS lesqu.c1·6UXU Knowlton, U. S. Geol. Survey BUll. 152, 

p. 246, 1898. 

Z'izY1J1L1£8 hyperboreu.8? Heer. Lesquereux, U. S. GeoI. and 

Geog. Survey Terr. Ann. Rept. for 1872, p. 389, 1873; 

idem for 1873, p. 387, 1874; idem for 1876, p. 516, 1878; 

The Tertiary flora, p. 276, pI. 51, fig. 15, 1878. 

The single example upon which Lesquereux based 

his origiJ?al identification with Heer's Zizyphus hyperboreus 38 is preserved in the United States National 

Museum (No. 424), but it appears to differ from that 

species in being twice or more times the size .and in 

being clearly 5-nerved instead of 3-nerved. The outer 

nerves in Z. lesquereumii ascend in an even curve nearly 

to the tip of the leaf, branching on the outside. In 

Z. hyperboreus the outer pair of nerves, if they may be 

so called, ascend in a series of bows, or rather the 

principal pair of nerves branch on the outside and 

form by a series of simple bows somewhat the appearance of a nerve. The finer nervation has entirely disappeared from Z. lesquereumii, and consequently it can 

not be compared in this particular with Z. hyperboreuis. 

Occurrence: Denver formation, Golden, Colo. Hanna formation, Carbon, Wyo .. 

Zizyphus hesperius Knowlton, n. sp. 

Plate 49, Figure 2 

Zizyph1ts hyperboreus Heel'. Lesquereux, Harvard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 55, 1888. 

Leaf of rather coarse texture, ovate or ovate-elljptical, abruptly rounded at the base, slenderly acuminate at the apex; margin obscurely preserved but 

apparently remotely undulate-toothed; nervation 

strong, 3-ribbed from the base of the blade, the 

midrib with two pairs of strong secondaries in the 

upper part, lateral ribs as strong as the midrib, arising at an angle of about 50°, much curved inward, 

reaching. the margin just below where the lowest pair 

of secondaries approach the margin, each with numerous thin branches on the outside .which join and make 

a series of loops just inside the margin; nervilles 

strong, at right angles to~the midrib. 

The specimen figured is the onlyone that is referred 

with certainty to this species. It is 8.5 centimeters 

long and about 5 centimeters broad at a point about 

one-third the length of the blade above the base. The 

coarse, heavy nervation is' well brought out in the 

figure. 

The type specimen of this species was identified by 

Lesquereux as ZiZ'!lpaus hype~boreus Heel' on the 

ground of its similarity to a leaf from the Hanna 

formation of Carbon, Wyo., so identified by him 39 

(since renamed ZizYP'hus lesquereumii !(now lton ) . A 

comparison of the two shows that although agreeing in 

size and shape the leaf from Car.bon is distinctly 5-

ribbed, with the inner pair of ribs ascending nearly 

or quite to the tip of the blade, with several outside 

branches, while the outer pair of ribs reach more than 

two-thirds the length of the blade and also have out8!! Heel', Oswald, Flora fossilis al'cticn, vol. 1, p. 123, pI. 49, fig. 2, 

1868; vol. 2, p. 482, pl. 50, fig. 20, 1869. 

3D Lesquereux, Leo, The Tertiary fiora, p. 276, pI. 51, fig. 15, 1878. 

• 
112 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

side branches. The margin also differs, and the whole 

nervation ,is very light, and altogether it is apparent 

that the two can not be conspecific. 

A comparison of this Golden specimen with Heer's 

figures of his ZizypMus hyperboreu8 40 shows a number of essential differences, such as the inner pair of. 

ribs ascending to the tip of the blade. It is also a 

smaller, narrower leaf. 

Occurrence: Denver formation, South Table Moun· 

tain, Golden, Colo., collected by Arthur Lakes. Type 

in Museum of Comparative Zoology, Cambridge, 

Mass" No. 1534. 

rather obtusely pointed at the apex; margin entire at 

the base, thence strongly undulate; petiole short, relatively stout; midrib very strong below, much thinner 

above; secondaries about eigHt pairs, alternate, at 

irregular distances, emerging at an angle of about 450 , 

not much curved upward, craspedodron1e;' nervilles 

numerous, mainly unbroken, at right angles to the 

secondaries. 

The example figured is the only one observed. It 

it about 7.5 centimeters long and a little over 5 centimeters wide. The petiole, which is complete, is 1.3 

centimeters long. 

Family VITACEAE This species is referred to the genus Oi88u8 on the 

ground of its resemblance to Oissus parrotliaefolia Cissus grosse-uentata Knowlton . Lesquereux,41 supposed to be from the Green RIver 

Plate 48, Figure'2; Plate 51, Figure 10 formation of Wyoming. It agrees with this species 

G-iS8'llS gl'osse-dentata Knowlton, U. S. Geol. Survey Prof. in size and shape and to a less extent in the undulate 

Paper 101, p. 340, pI. 104, fig. 1, 1917 [1918]. lnargin. .The nervation in O. pO!r7'ottiaefolia differs 

With slight modification the original description is from that of O. coloradensis in that the lowest pair 

as follows: of secondaries has several tertiary branches on' the 

Leaf of large size and firm texture, broadly ovate, rather outside. 

abruptly rounded below to a slightly decurrent basal portion, Occurrence: Dawson arkose, Rice's clay pit, 1 mile 

acuminate above; margin entire at base, then strongly few southwest of Calhan, Colo., collected by F. H. Knowltoothed, the teeth large, obtusely acute, separated by shallow 

sinuses; midrib strong below, becoming thin above; second- ton, 1910. 

aries five or six pairs, at an angle of 45° or 50°, the lowest Cissus coloradensis Knowlton and Cockerell 

pair much the stronger, arising well above the decurrent portion of the blade, each with four or five tertiary branches on Plate 47, Figure 6; Plate 50, Figure 2 

the outside which are curved upward and pass to the marginal ' 0'i88US ooloraclensis Knowlton and Cockerell, U. S. Geol. Survey teeth; other secondaries alternate or nearly opposite, terminat- . . I th d l' t Bull. 696, p. 176, 1919. ing in the large teeth; nervll es numerous, ra er e lca e, Gissus laevigata Lesquereux, U.· S. Geol. and Geog. SUl'ves 

irregular. Terr. Ann. Rept, for 1872, p. 380, 1873; idem for 1876, 

The present specimen, although not nearly so per- p. 512, 1878; The Tertiary fiora, p. 238, pI. 40, figs. 12, 

fect as the type specimen, appears to agree with it in 13, 1878. [Homonym, Blume, 1825-26.] 

all essential particulars. The type specimen was Leaves membranaceous, broadly oval, narrowing in 

about 14 centimeters in length, but the present speci- a broad curve to the short petiole; 3-ribbed fron1 the 

men could hardly have been more than 10 or 11 centi- base or slightly above it, the lateral ribs running 

meters; it has the petiole preserved for' more than 2.5 nearly straight to about the middle of the leaf, 

centimeters: The width of the blade is a little over 7 branching on the outside; secondaries alternate or 

centimeters. subopposite, parallel; nervilles prominent, percurrent, 

As pointed out in the original diSCUSSIon of this approximately at right angles to the midrib; areolaspecies it seems ·to be most closely related to Ois8u8 tion quadrangular. 

parrottiaefoUa Lesquereux, fron1 which it differs in its This species, according to Lesquereux, was founded 

greater length, much more pointed apex, and decur- on two specimens, both of which lacked the upper 

rent basal portion and in having its lowest pair of sec- portion. Only one of these type specimens is now 

ondaries arising well above the base instead of at or to be found in the collections of the United States 

near the top of the petiole. National Museum (No. 347). This is the original of 

Occurrence: Da~son arkose, Rice's clay pit, 1 mile Figure 13 of Lesquereux's plate. Both specimens are 

southwest of Calhan, Colo., collected by F. H. Knowl- said to have come from the same place together with 

ton, 1910. Raton formation, Morley, Colo. a number of smaller fragments, but none appear to 

Cissus hesperia Knowlton, n. sp. 

Plate 47, Figure 1 

Leaf of medium size, subcoriaceous, ovate, very obtusely wedge-shaped or nearly truncate at the base, 

40 Heel', Oswald, Flora fossilis arctic:1, vol. 2, pt, 4, p, 482, pI. 50, 

fig. 20, 1869, 

have been since obtained from the type locality. 

The two specimens here figured were found near 

Sedalia, Colo., and agree. in every essential particular 

with the figures of the original specimens. Like the 

originals they both lack the upper portions of the 

41 Lesquereux, Leo, The Tertiary flora, p. 239, pI. 42, fig. 1, 1878. 
SYSTEMATIC PALEOBOTANY 113 

leaf, although one (pI. 47, fig. 6) has fully two-thirds 

of the lamina preserved. Lesquereux thought it probHble that the type specimens were obscurely trilobate 

at. the apex, but there is no indication in the most perfect leaf that this would be the case. Another specimen obtained near Sedalia represents the middle part 

of a leaf but a little higher up than that shown in 

Plate 47, Figure 6, and there is every indication that 

it would be rounded at the apex very much like the 

base. 

Occurrence: Dawson arkose, lower part (in beds 

equivalent to Arapahoe formation), 3,000 feet east 

of the Douglas coal mine, near Sedalia, Colo., collected 

by Arthur Lakes, 1890. Denver formation, G:olden, 

Colo. Raton formation, Abeton, Cola. 

Cissus? cannoni Knowlton, n. sp. 

Plate 47, Figure 5 

Leaf evidently coriaceous, roughly deltoid or deltoidovate, truncate at the base, obtuse at the apex; margin 

irregularly cut into a number of obtuse, often toothed 

lobes; midrib very strong; ~econdaries about four 

pairs, the lower pair as strong as the midrib, much 

arched upward, passing to lobes in about the middle 

of the blade, each with several tertiary branches on 

the outside, these or their branches passing to lobes 

or marginal teeth; upper secondaries simple, ending 

in lobes 01: teeth; nervi lIes few, strong. 

The single leaf figured is all that was observed of 

this species. It is broadly ovate, with an abruptly 

rounded truncate base and an obtuse apex. The margin is provided with about three relatively strong, 

obtuse lobes on each side, in addition to which the 

basal portion and some of the lobes are provided with 

a few obtuse teeth. The strongly marked nervation is 

well shown in the figure. The length of this little leaf 

is about 3.75 centimeters and the width about 2.5 centime tel's; its petiole is not preserved. 

This leaf appears to be most ~losely' related to a 

small leaf referred by Lesquereux to Giss'UJs tricu8pidata I-Ieer,42 the only one of the specimens figured 

llllder this name that can not be found in the collections of the United States National Museum. The locality whence this came is unknown. Our leaf is of 

approximately the same size but is broadly truncate 

instead of obtusely wedge-shaped at the base; the 

secondaries are fewer and the lower pair much curved 

upward. The marginal lobes and teeth are similar in 

both, though rather sharper in Lesquereux's specimen. 

The general arrangement of the secondaries is also 

suggestive of the arrangement in the leaf referred by 

4~ LcsquCI'C'UX, Lco, The TertJD,l'Y flora, p. 240, pI. 41, fig. 6, 1878. 

o 

Lesquereux 43 to V itis ol1'iki, but the size, outline, and 

margin are quite different. It may be that. a larger 

series of specimens would show a further transition 

between the one in hand and Gissus t'ricuspiilata, but 

for the present it seems necessary to keep them apart. 

I have named this supposed new species in honor of 

Mr. George L. Cannon, the collector of much of the 

material from Middle Park. 

Occurrence: Middle Park formation, high terrace 

ridge between forks of IGnney Creek, in, "breccia 

spoon," east of JIot Sulphur Springs, J\1iddle Park, 

Grand County, Colo., collected by 'Vhitman Cross, 

October, 1891. 

Cissus corylifolia Lesquereux 

Plate 47, ll'igure 4; Plate 48, Figure 1; Plate 49, Figure 1 

Gis sus c01'ylifoUa Lesquereux, Harvard ColI. Mus. Compo 

Zoology Bull., vol. 16, p. 52, 1888. 

Lesquereux describes this species as follows: 

Leaves thickish, ovate, blunt at apex, Simply or doubl~' short 

dentate, strongly pinnately nerved; lateral nerves at an acute 

angle of divergence, close, parallel, scarcely curved in passing 

to the borders, the lowest much branched on the under side, 

the upper ones branching near their ends, craspedodrome; 

nervilles at right angles to the nerves, simple or branched in· 

the middle, deeply impressed. 

These finely preserved leaves vary in length from 6 to 9.5 

centimeters and from 4 to 7.5 centimeters in width, being 

broadest a little below the middle. They have a degree of 

likeness to Parrotia pristina Ettingshausen.44 

The three type specimens on which this species was 

based are all nearly perfect and all have been figured. 

The two smaller leaves (pI. 47, fig. 4; pI. 49, fig. 1) 

are undoubtedly conspecific, but the assignment of the 

larger one (pI. 48, fig. 1) is less certain. It is very 

much larger and has the secondaries at a somewhat 

sharper angle, the upper ones being alternate instead 

of opposite and the lower pair with compound branching on the lower side. The margin is entire below and 

seemingly remotely dentate above, though lnost of the 

upper margin is missing. This specimen suggests certain of the leaves that have been referred to Quercus 

viburnifolia Lesquereux, but the marginal cha.racters 

can not be compared. 

The two smaller speciInens are very close to Gis sus 

coloradensis, described above, differing apparently in 

the margin being somewhat doubly toothed and in 

the secondaries being at a more acute angle. 

.... Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, 

Mass., Nos. 1501, 1503, 1504. 

48 Idem, pl. 41, fig. 8. 

44 Ettingshausen, Constantin von, l;']om "o~ Billn, pt. 2, pI. 39, fig. 

23, 1868. . 

• 
114 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Cis sus lesquereuxii Knowlton 

Plate 48, Figures 3 and 4 

Gissus tricu8Pidata (Heer) Schimper, Paleontologie vegetale, 

vol. 3, p. 44, 1874. 

Gi8su8 tricuspidata. (Heer) Schimper. Lesquereux, The Tertiary flora, p. 240, pl. 41, figs. 4-7, 1878; Harvard 

ColI. Mus. Compo Zoology Bull., vol. 16, p. 51, 1888. 

[Homonym, Siebold and Zuccarini, 1846.] 

Gissus ZesquereuaJii, Knowlton, U. S. Geol. Survey Bull. 696, 

p. 177,,1919. 

There are several specimens in a collection obtained 

near Calhan, Colo., that appear to be identical with 

certain leaves from Black Buttes, Wyo., which were 

identified by Lesquereux with Heel"s Oissus tricuspidata. Although all of them are fragmentary, they 

show the basal portion fairly well. The best-preserved specimen is shown in Figure 4. It is a small 

leaf not quite 4: centimeters wide and has the com-' 

plete petiole preserved, 1.5 centimeters long and seemingly strong for the size of the blade. The margin 

of the base for a distance of 1 centimeter from the 

insertion of the petiole is entire, but thence it is 

oharply serrate. The primary nervation consists of 

five ribs, which arise near the top of the petiole. The 

basal pair are much the lighter and have a few tertiary branches on the outside which pass to the teeth. 

The finer nervation is not retained. 

The validity of referring either these Calhan leaves 

or those from Black Buttes to Heer's species may well 

be questioned; in fact, Lesquereux pointed out a number of marked differences in his discussion of the 

Black Buttes specimens. The point that it is desired 

to establish now is the apparent identity between the 

Calhan and Black Buttes leaves, but their reference 

to the European Miocene species may well be'deferred 

until the thorough revision of the Black ,Buttes flora 

is undertaken. 

Since the above statement was written I have had 

opportunity to study the three specimens from Golden 

l'e~erred by Lesquereux 45 to this species. They are of 

about the same size as the leaves from Calhan and 

like them do not have the extreme basal portion serrated. However, two of the type specimens as fig-

\lr~d by L.esquereux 46 show the same tendency, so it 

eVIdently IS not a significant character. One of the 

Golden specimens is represented by Figure 3. 

Occurrence: Dawson arkose, Rice's clay pit, 1 mile 

southwest of Calhan, Colo., collected by F. H. Knowlton, 1910. Denver formation. The examples from 

South Table Mountain, Golden, Colo., identified by 

Lesquereux, are in the Museum of Comparative Zoology, Cambridge, Mass., Nos. 1493, 1493a. 

411 Lesquereux, Leo, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, 

p. 51, 1888. 

46 Lesquereux, Leo, The Tertiary flora, pl. 41, figs. 4, 6, 1878. 

Cissus obovata Knowlton, n. sp. 

Plate 49, Figure 5 

GiSS~t8 pa1'rottiaefoZia Lesquereux. Lesquereux, Harvard CoHo 

Mus. Compo Zoology Bull., vol. 16, p. 52, 1888. , 

Leaf coriaceous, the nervation deeply impr~ssed, 

obovate or elliptical-obovate, broadest near the middle, 

whence it tapers in an almost straight line to the base 

and is rounded to the subacute apex; margin entire to 

a point well above the middle, thence undulatedentate, the teeth remote, rather obtuse; nervation 5-

ribbed from the top of the petiole, the midrib straight, 

with two or three alternate secondary branches in the 

extreme upper part, next pair of ribs as stronO' as the 

midrib, at an acute angle, slightly incurved, r~aching the upper margin, with four or five branches on the 

outside,. some of which are again branched, outer primary rIbs at an angle of about 60°, nearly straight 

each with several secondary branches; branches of th~ outer primaries camptodrome, all other ribs and their 

branches craspedodrome; nervi lIes very strong, usually 

unbroken. The length of this leaf is 6.5 centimeters, 

and the width a little over 5 centimeters. 

This species is based on the single specimen fi O'ured. 

It was identified by Lesquereux as Ois8'U8 pa:rr~ttiae­ folia Lesquereux, but a comparison with the figures of 

this species in his account of the Tertiary flora (pI. 40, 

figs. 15-17) shows distinct differences. Thus, the orio'-

in.al specimens supposed to be from Green River, Wy~., are all distinctly ovate, with a very obtusely wedgeshaped or even slightly heart-shaped base and large 

lmdulate-toothed margin. The present species is obo-

,,'ate, with a wedge-shaped base and a marO'in which is • b 

entIre below and slightly undulate-toothed above, the 

teeth acute rather than obtuse and rounded. The nervation in Oissus pl(JJl'rottiaJefoli~ is essentially 3-ribbed, 

vyith the lateral ribs at an angle of about 50°, little 

curved inward and not reaching the upper portion of 

the blade; none of the secondary branches are forked. 

The whole nervation in O. pa1"f'ottiaefolia is liO'hter 

than in Oissus obovata. b 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Type 

(under the name Oissus parrrottiaefoZia) in the Musemn 

of Comparative Zoology, Cambridge, Mass., No. 1495. 

Cissus lobato-crenata Lesquereux 

Plate 49, Figures 3, 4 

Gissus lobato-crenata Lesquereux, U. S. Geol. and Geog. Survey Terr. Ann. Rept. for 1872, p. 396, 1873; idem for 

1873, pp. 382, 385, 408, 1874; idem for 1876, p. 512, 1878; 

The Tertia'ry flora, p. 240, pI. 41, figs. 1-3, 1878; U. S. 

Nat. Mus. Proc., vol. 11, p. 38, 1888; Harvard ColI. Mus. 

Compo Zoology Bull., vol. 16, p. 52, 1888. 

The three figured types of this species are preserved 

in the United States National Museum (Nos. 349, 354~, 
SYSTEMATIC PALEOBOTANY 115 

354b) and ure before me. They are not particularly 

well ~)reserved but uppear to have been as accurately 

descrIbed and figured as their condition warrants. 

The species was first described from Black Buttes 

Wyo., the original specimen being apparently tha~ shown in Pluto 41, Figure 1, of the report on the Tertiary flora. It was later detected on Mount Bross 

Middle })urk, Colo., and the specimens found ther~ are shown in Figures 2 and 3 of the same plate and are 

preserved on the same piece of matrix. The Black 

Butt.es specimen is the most perfect, having the upper 

portIOll and most of the margin preserved. It is obscurely crenate-lobed; the lobes corh~spond to the 

number of prominent ribs and secondaries, and the 

spacps between the lobes are crenate. The examples 

f1'0111 l\10Ullt Bross do not seem to be lobed but are 

simply crenate, though the margin is not well preserved. The size and general character of the nervation is practically the same in all the type specimens 

and, it is probable that they should be considered 

identical, but more material is needed to settle the 

point. Among the numerous specimens obtained by 

War~ at Black Buttes I do not note any that belong 

to OU8'l.l!t~ looato-01'enata, unless possibly what he has' 

called II edera oruneri 47 should be so referred. This 

specimen is strongly 3-ribbed from the base and has a 

long, strong petiole, but the upper portion of the blade 

is lacking and hence' can not be satisfactorily compared with Lesquereux's species. The nervation of 

H. oruner-i is similar to that of the form under consideration except that the lateral ribs curve slightly 

upward. . 

In the recent . collections from Middle Pa~k I find 

several rather poorly preserved leaves that are undoubtedly similar to the types that came from this 

locality. 

A single fairly well preserved leaf from Sedalia 

Colo., shown in Figure 4, has been referred to thi~ ~pecies. It is of about the same size as the smaller 

of Lesq nereux's types, and although it has not quite 

so broad a base it has the marginal cutting and nervation of this form. It has what the other specimens 

unfortunately lack, namely, the petiole or' a portion 

of it; this is relatively strong. 

Another specjmen from Colorado has also been illusb'ated (fig. 3) for the purpose of showing the comp.lete petiole. This is 4.5 centimeters long and is relatlvely strong. The blade of this leaf is fraO'mentary b , 

but so far as preserved it is not to be distinO'uished I::> 

from one of the type leaves figured by Lesquereux. 

Occurrence: Black Buttes coal group (type), Black 

HuHt's, vVyo. Dawson arkose~ lower part (in beds 

47 Wnl'd, L. F., U. S. Geol. Survey BUll. 34, p. 58, pI. 26, fig. 6, 1887. 

equivalent to Arapahoe formation), 3,000 feet east of 

Douglas coal mine, near Sedalia, Colo., col~ected by 

Arthur Lakes, 1890. Middle Park formation Mount . ' B:os~, MIddle Park, Grand County, Colo. (type), also 

debrIS on slope of Mount Bross, opposite Hot Sulphur 

Springs, collected by G. L. Cannon, 1889. Dawson 

arkose, Red Hill, 4 miles south of Ramah, Colo., collected by G. B. Richardson, 1910. 

Vitis olriki Heer 

Plate 49, ,Figure 6; Plate 50, Figure 4 

Viti8 olri1cl H~er, Flora fossilis arctica, vol. 1, p. 120, pI. 48, 

fig. 1, 1868. 

Lesquereux, U. S. Geol. Survey Terr. Ann. Rept. fOol' 1871, 

Suppl., p. 12, 1872; U. S. Geol. and Geog. Survey Terr 

Ann. Rept. for 1873, p. 385, 1874; idem for 1876, p. 512, 

1878; The Tertiary flora, p. 241, pI. 41, fig. 8, 1878. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 3~8, 

1917 [1918]. 

The presence' of this species in American strata was 

first based on a single example described and figured 

by Lesquereux from Evanston, Wyo., in beds lying 

~, above the coal " and regarded by me as being Tertiary, or in the approximate position of the Denver 

beds. Lesqu'ereux also stated that this species had been 

found in the Raton Mountains of Colorado, but no 

specimen from this region was contained in the older 

collections of the United States National Museum. In 

recent years, however, specimens representing this 

species were found in the Raton formation of the 

Raton region of New Mexico, though they were not 

very well preser.ved and were not figured, as they could 

add nothing to our knowledge of this form. 

In the collections obtained near Ramah there is the 

single example here figured (pI. 50, fig. 4), which is 

certainly the same as the leaf figured by Lesquereux 

in the volume on the Tertiary flora. Although it is 

fragmentary it admirably exhibits the basal portion 

which is deeply heart-shaped with the two lobes clos~ together if not overlapping. The margin of the lower 

part is undulate rather than strongly toothed. The 

nervation is very strong and is precisely like that in 

the Evanston leaf figured by Lesquereux. 

A specimen found near Calhan, Colo., has also been 

figured (pI. 49, fig. 6), and although it is somewhat 

narrower than the figure given by Lesquereux it does 

not differ essentially. 

Occurrence: Dawson arkose, Red Hill, 4 miles south 

of Ramah, Colo., about 100 feet above a coal bed, 

which is equivalent to the Purdon coal or near it , collected by G. B. Richardson, 1910. Raton formation, mesa north of Raton, N. Mex., at top of formation, collected by W. T. Lee. Evanston formation, 

Evanston, Wyo. 
116 FLOB.A OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Order MAL VALES 

Family TILIACEAE 

Grewiopsis tenuifolia Lesquereux 

Plate 50, Figure 3 

Grew'iopsis tenu'ifoUa Lesquereux, The Tertiary flora, p. 258, 

pI. 40, fig. 14, 1878; U. S. Geol. and Geog. Survey Terr, 

Ann. Rept. for 1876, p. 514, 1878; Harvard ColI. Mus. 

Compo ,Zoology Bul1., vol. 16, p. 54, 1888. 

The type specimen of 'this species is fortunately 

preserved in the United States National Museum (No. 

348). It is recorded in the Museum catalogue and 

also in the published accounts as having' come from 

Black Buttes, Wyo., but the matrix shows beyond 

question that it came from the andesitic beds of the 

Denver formation at Golden, Colo. There is also 

another specimen identified as this species in the Museum collection under the same number, and one specimen was reported by Lesquereux among the plants 

from Golden, Colo., examined for the Museum of 

Comparative Zoology of Harvard College. 

The type specimen is fragmentary, consisting of 

only one side of the leaf, the base and apex being 

absent. It is possible that Lesquereux had other specimens that are not now preserved, for he characterized the species as follows: 

Leaves membranaceous, rounded or cordate, taper-pointed, 

subpalmately nerved; borders irregularly dentate; lateral 

nerves mostly craspedodrome or entering the teeth by strong 

nervilles. 

There is a single specimen that undoubtedly belongs 

to this species from Mosby, Colo., and, although it 

lacks much of one side and the extreme apex, it shows 

the base and margin very perfectly. By drawing 

the characterization from the type specimen and the 

~losby specimen, it is possible to give the following 

description: 

Leaf of medium size, membranaceous, broadly ovate, 

t"bruptly truncate at the base, with a small cordate 

portion just above the petiole, and rather acutely 

pointed at the apex; margin entire on the truncate 

basal portion and thence remotely and, somewhat irregularly :few toothed, the teeth low, sharp, or almost 

~piny; petiole about 1.3 centimeters in length, apparently short for the size of the leaf; nervation consisting of a rather slender, straight midrib and three 

or possibly four pairs of craspedodrome secondaries, 

the lowest pair arising some distance above the top 

of the petiole, each with three or four tertiary 

branches on the outside which end in marginal teeth; 

other secondaries, especially ,the lower pair, with tw.o 

or three tertiary branches passing to the teeth; nervilles thin, mostly unbroken, approximately at right 

angles to ~he secondaries or their branches. 

The type specilnen is about 9 centimeters long and 

6 centimeters wide, and the Mosby specimen is probably between 10 and 12 centimeters long and is 8 

centimeters wide. The petiole in the Mosby speeimen 

is a little over 1 centimeter long. 

Occurrence: Denver formation (type) , Golden, 

Colo. (erroneously recorded as coming from Black 

Buttes, Wyo.). Dawson arkose, dump of Mosby coal 

lnine, Mosby, Colo., collected by F. H. Knowlton and 

G. B. Richardson, 1910. Black Buttes coal group, 

Black Buttes, 1Vyo. 

Family STERCULIACEAE 

Sterculia? heteroclonta Knowlton, n. sp. 

Plate 50, Figure 5; Plate 51, Figures 4, 8 

Leaves of medium size, firm in texture, ovate or 

squamose, truncate or slightly decurrent at th~ base; 

margin with four or five large, deltoid, rather obtuse 

teeth or lobes on each side, separated by shallow 

sinuses; nervation pinnate, consisting of a relatively 

very strong midrib and usually three pairs of opposite 

or subopposite secondaries, the lowest pair arising at 

the top of the petiole in the decurrent port.ion of the 

blade and simulating ribs; secondaries at an angle of 

45° to 50°, the lowest pair with two or three branches 

on the lower side which end in the large teeth; middle 

pair of secondaries parallel to the lower pair: craspedodrome; upper secondaries' th,in, camptodrome, 

curving up along the border; short branches :from the 

secondaries pass up to and around the sinuses; finer 

nervation not retained. 

This species is represented by nearly a dozen ex· 

nmples, several o:f which are exceptionaUy well preserved. They are 7 or 8 centimeters long and 5.5 or 6 

centimeters wide. The petiole was not observed. The 

lobes are approximately 1 centimeter long and separated by rather shallow sinuses. The lobes are all 

ra ther obtuse and rounded. 

There are a number of described species to which 

the present :form has more or less resemblance. Among 

these may be mentioned PlataJr//Utl3 1'homooidea I .. esquereux,48 from the Denver formation at Golden, Colo., 

'which is ot about the size of the leaves under consideration but differs in being distinctly wedge-shaped 

at the base and having more numerous as well a.s 

smaller and sharper teeth. They can hardly be eyen 

congeneric. 

There is a remote, perhaps only superficial resemblance between Ste1YfUl~? hete1'ocfonta and wha.t 

Ward 49 called OreibnJ31"ia.~ datWl"iaefolia, :from the Fort 

lTnion' :formation of Montana. This also has more 

numerous teeth, which are of irregular sizes a.nd are 

48 Lesqnereux, Leo, The Tertiary flora, p. 186, pI. 26, figs. 0, ';. 1878. 

(9 Ward. L. F., F. S. Geol. Survey Sixth Ann. Rept., p. 556. pI. 58. 

figs. 1-5, 1886. 
SYSTEMATIO PALEOBOTANY 117 

very sharply pointed. The nervation IS also quite 

different. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 1 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. I(nowlton, July, 1910. 

Sterculia saportauea !{uowltou 

Plate 53, Figure 6 

Stel'C'uUa 8apo1'tanea Knowlton, U. S. Qeo!. Survey BUll. 152, 

p. 224, 1898. 

SfC;"oltZia modesia Saporta,GO Prodrome d'une flore fossile des 

trnvertins anciens de Sezanne: Soc. geol. France Mem., 

3d ser., vol. 8, p. 401, pI. 12, fig. 2, 1868. 

Lesquereux, The Cretaceous and Tertiary floras, p. 125, 

pI. 20, fig. 5, 1883. 

Following is Lesquereux's original description: 

Leaves thick, rounded in the lower part, trilobate at apex; 

median lobe longer, separated from the lateral [lobes] by 

brond sinuses; nervation trifid from the base; lateral nerves 

camptodrome. 

This finely preserved leaf is about 8 centimeters 

long from the base to the apex of the middle lobe and 

6 centilneters broad between the tips of the lateral 

lobes. 'It is enlarged in the middle, a little contracted 

below the lateral lobes, and deltoid to the apex. The 

primary nerves are strong; the lateral nerves are entwined by distinct nervi lIes , and the areolation is in 

loose, irregularly quadrate nleshes. 

This fine species is obviously closely related to Ste1'-

(:ulicD libbeyi, next described, under which the points 

of difference are more fully set forth. The type specimen is the property of Princeton University, but unfortun~tely it could not be found at the time of my 

visit to that institution. It is recorded by Lesquereux 

as having come from Golden, Colo., and there is no 

reason to doubt the statelnent, but without seeing the 

matrix I can n.ot be certain of the horizon from which 

it came. Inasmuch as it is so closely allied with 

SterfYUlia libbey:i, which came from the andesite beds 

at Goldeh, it is assume¢! that Steroulia s((;portanea 

also came from these beds, but obviously this assumption requires confirmation. If this species had been 

subsequently found its horizon would be settled more 

definitely, but so far as known the type specimen 

remains unique. 

Occurrence: Denver formation ( ~), Golden, Colo. 

Type in museum of Princeton ~niversity. 

Sterculia libbeyi Kuowlton, n. sp. 

Plate 53, Figure 2 

Leaf large, broadly ovate, regularly rounded at the 

base, palmately 3-10bed, the lateral lobes short, sharp 

311 As SterO'lllia modesta of Saporta, 1868, Is preoccupied by S. 

tnoaesta Heel', 1853-1855, it was changed to S. 8apo1·tanea, In honor of 

M. Ie Comte Gaston de Saporta, who did so much to make known tl1e 

l)aleobotanic riches of his own country. 

pointed, the central lobe much the larger, long acuminate; nervation tripalmate, the two lateral ribs rather 

slender, slightly curving and entering the points of 

the lateral lobes, the midrib of about the same 

size, straight, entering the apex of the terminal lobe; 

lateral ribs with about seven secondaries on the outside 

only, these camptodrome, each joining the one next 

a bove by a broad loop some distance inside the margin; outside these larger loops is a single series of 

smaller loops just within the margin, producing more 

or less of a line; midrib with about eight or nine pairs 

of alternate, slender camptodrome secondaries having 

the same series of smaller outside loops as the lateral 

ones; nervilles prominent, mostly at right angles to 

the secondaries, a few percurrent, but mostly broken 

in the middle; ultimate nervation thin, quadrangular, 

filling the space between the nervi lIes with a fine 

quadrangular areolation. 

This specimen figured, unfortunately the only one 

found, is a large, beautifully preserved leaf so far as 

nervation goes, although lacking the greater portion of 

one side. It is about 16 centimeters long and 10 centimeters wide in the broadest· portion, which is in the 

lower part of the blade. The base appears to have 

been rounded, probably truncate without being heartshaped. It is broadly ovate, with the small, sharppointed lateral lobes some distance above the middle 

portion. The central portion of the leaf, or terminal 

lobe, a!:? it may be called, is very long and slender in 

comparison with the lateral lobes. The leaf is palmately 3-ribbed, the lateral ribs entering the lateral 

lobes and the middle one the apex. The secondaries 

are all camptodrome, forming broad, even loops with 

a smaller series of loops outside. The nervilles are 

all well preserved and seem to be mainly at right angles 

to the secondaries or to the lateral ribs and lower pair 

of secondaries in the central lobe. The finer nervation 

is also well preserved and produces a fairly regular 

quadrangular areolation between the nervilles. 

This species is undoubtedly closely related to Sterculia saportanea Knowlton as figured by Lesquereux III 

(under the preoccupied name S. modesta Saporta). 

The fine leaf that was the original of Lesquereux's 

figure is the property of Princeton University, as indeed is the present species, but it can not now be 

found, and without seeing it I am unable to determine 

whether it came from Laramie or Denver strata at 

Golden. Ster(JUlia libbeyi differs from S. sap01'tamea 

in being twice as long and nearly twice as broad, with 

the central or terminal lobe twice as long. The nervation is of the same character in both. 

Sterculia libbeyi recalls slightly the larger leaves 

described by Newberry fron1 Cook Inlet, Alaska, as 

61 Lesquereux, Leo, The Cretaceous and Tertiary floras, pI. 20, fig. 5, 

I 1883. 
118 FLORA OF THE DENVER 'AND ASSOCIATED FORMATIONS OF COLORADO 

Ficus.~ alaskana,62 but this species really differs in 

many particulars, such as being heart-shaped at the 

base, with five prominent ribs palmately arranged, as 

well as in nervation. 

The type specimen of Sterculia libbeyi is No. 2339 

of the collection of Princeton University, where it is 

now preserved. This with a small amount of other 

material was kindly placed at my disposal by the 

curator of the museum, Prof. William Libbey, jr., in 

whose honor I take pleasure in naming it. 

Occurrence: Denver formation, Golden, Colo. Type 

in Princeton University. 

Pterospermites grandidentata Lesquereux 

Plate 50, Figure 6; Plate 51,. Figure 3 

Pterosper-mites grandidentata Lesquereux, Harvard ColI. Mus. 

Compo Z·oology Bull., vol. 16, p. 53, 1888. 

This species was described and discussed by Lesquereux as follows: 

Leaves large, sometimes very large,. somewhat like leaves 

of Plata1VU8, palmately sub-5-nerved; t~e outer lateral nerves 

generally thin and shorter, much divided outside; lower secondary nerves opposite, at a dista~ce from the base; borders 

deeply dentate, the teeth acute, turned upward, entered by the 

primary nerves and their branches, while toward the apex 

the secondaries curve in festoons along the borders, joined to 

the teeth by small anastomosing branches; nervilles strong, 

at right angles to the nerves. 

This definition is about t~e same as that given by Saporta ~ 

of Pterospermite.s inaequifoZius. One of the leaves from 

Golden is well preserved and merely differs from those described by Saporta in having the lateral nerves somewhat incurved, not quite straight. The two other specimens represent 

merely the base of two leaves with five primary nerves around 

the point of attachment of the petiole and two smaller ones 

declining downward to the cordate base, as in the leaves of 

Ficus (Dombeyopsis) gra.tnditoZia Unger. 

The two best-preserved specimens are here figured. 

Of these the most perfect, shown in Plate 51, Figure 3, 

is broadly ovate-acuminate, 11.5 centimeters long and 

about 8 centimeters broad. The base is very slightly 

heart-shaped, almost truncate, and entire. Above the 

basal portion the margin is provided with large, sharppointed teeth. The nervation is very strong, being in 

effect 7-nerved instead of 5-nerved as stated by Lesquereux. There is a strong, straight midrib with three 

or four secondary branches above the middle of the 

blade. The next pair of ribs is nearly as strong as 

the midrib. They arise at an angle of about 70° and 

curve upward, reaching within one-fourth the length 

of the blade from the apex. They are provided with 

five or six outside branches that enter the teeth. The 

next pair of ribs are at the same angle as· the branches 

above them but arise at the top of the peti<;>le. There 

52 U. S. Nat. Mus. Proc., vo1. 5, p. 512, 1882 [1883]. 

G3 Saporta, Gaston de, Prodrome d'une flore f'ossile des travertins 

anCiens de S~zanne, vol. 7, p. 402, pI. 35 (12), figs. 3-1), 18fl8. 

is a thin pair of ribs below that supply the basal lobes 

of the blade. These, as well as the ribs next above, 

have a number of outside branches. The nervilles are 

very strong, usually broken. 

This species, as Lesquereux has pointed out, is undoubtedly similar to Pterospermites inaequifolius Saporta, from the travertines of Sezanne. It has the 

large ribs at a less acute angle and little curved upward, and the margin is much less strongly toothed; 

the nervilles are the same in both. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Types 

in Museum of Comparative Zoology, Cambridge, 

Mass., Nos. 1517, 1519. 

Order UMBELLALES 

Family CORNACEAE 

Cornus impressa Lesquereux 

Plate 51, Figures 1, 2 

OornU8 impressa Lesquereux, U. S. GeoI. and Geog. Survey 

Terr. Ann. Rept. for 1873, p. 408, 1874; The Tertiary 

flora, p. 243, pI. 42, fig. 3, 1878. 

Oornus emmonsii Ward, U. S. Geol. Survey Sixth Ann. Rept., 

p. 553, pI. 48, fig. 2 [not fig. 3], 1885; U. S. Geol. Survey 

Bull. 37, p. 55, pI. 26, fig. 2 [not fig. 3], 1887. 

The type speeimen of 001'11lUS impressa, from Mount 

Bross, Middle Park, Colo., is No. 354 of the UnIted 

States National Museum collection. It was character-

~zed as follows: 

Leaves thick, coriaceous, entire, deeply impressed into the 

stone, regularly elliptical, rounded to a very short, scarcely 

marked acumen, rounded also to the base; secondary nel'ves at 

an acute angle (about 40°) of divergence, slightly curving in 

ascending to the borders, regularly camptodrome, simple or 

rarely branched once near the point, and anastomosing in 

festoons along the borders, with strong nerv.illes at right angles 

to the midrib, mostly simple and ~ontinuous. 

The type is about '1 centimeters long and nearly 4.5 

centimeters wide. , o 01"J1lUS impressa was compared by Lesquereux to 

Oomus or'oifera Heer,54 from the Swiss Miocene, 

though he points out a number of minor differences. 

It does not appear to have been a very abundant species, though it has been found rather· widely distributed. One or two broken examples have' been found in 

the andesitic beds at Golden, Colo., and a single wellpreserved specimen .came from the Denver beds in a 

ravine opposite St. Luke's Hospital in North Denver.55 

This specimen was one of the types of Ward's Oornus 

emmwnsii, and although it differs slightly from the 

type of O. impressa seems best referred to that species. 

The 0 ornw e'1nmorl~ii of Ward rests on two specimens, 

&4 Heer, Oswald, Flora tertiaria Helvetlae, vol. 3, p. 27, pI. 105, figs. 

15-17, 1859. . 

55 Wrongly stated by Ward to be from Golden, Colo. 
SYSTEMATIO PALEOBOTANY 119 

the one above mentioned from North Denver and the 

other from the Mesaverde formation at Point of 

Rocks, Wyo. In 1900 GO I referred both these specimens to O. impressa on the basis of the figures given 

by Wal'd, as the specimens were not then available. 

Since then I have studied the specimens and found that 

the Point of Rocks example has been entirely misinterpreted and incorrectly drawn. The margin 

throughout and the distal terminations of the secondaries are exceedingly obscure. Several of the secondaries seem to enter the margin, and in one place it is 

possible to demonstrate with considerable certainty 

that they are camptodrome and arch just within the 

margin; hence it is probable th'at all of them do this. 

The apex is entirely wrong in the figure. For these 

r~asons the name 0 07'nUS emmonSii has been restored 

to the Point of Rocks specimen, and the other from 

North Denver is referred to O. impressa. 

. The example here figured from Mosby, Colo., is 

lacking in considerable degree, but so far as can be 

made out it agrees well enough with O. impressa. 

Occurrence: Middle Park formation (type), Mount 

Bross, Middle Park, Colo. Denver formation, ravine 

opposite St. Luke's Hospital, North Denver, Colo., 

and South Table Mountain, Golden, Colo. Dawson 

arkose, dump of Mosby coal mine, Mosby, Colo., collected by F. H. Knowlton and G. B. Richardson, 1910; 

Rice's clay pit, 1 mile southwest of Calhan, Colo., 

collected by F. H. Irnowlton, 1910. 

Cornlls lakesii Knowlton, n. sp. 

Plate 50, Figure 1 

Leaf coriaceous, ovate-lanceolate, entire, broadest just 

below the middle, tapering upward into a long, siender acuminate apex and downward to a wedge-shaped 

base; nervation pinnate, midrib strong, secondaries 

four pairs, alternate, emerging ·at· an acute angle, 

curving along the margin high above their insertion; 

n~villes numerous, fine, broken, approximately' at 

right angles to the midrib. 

This fine leaf, which is the only one obtained, has 

a length of 6 centiIIl:eters and a width of about 2:5 

centimeters in the broades~ part, just below the middle. The upper portion is slightly oblique-that 

is, turned to one side-after the manner of several 

living species. The petiole, unfortunately, is not preserved. . 

This species seems to have several near relatives 

among living species. It may be compared with O. 

pamiculata L'Heritier, a shrub common on river banks 

in the eastern United States, and more particularly 

with small leaves of O. 8ericea Linne, which grows in 

wet places from Canada and Dakota to Florida. 

BO Knowlton, F. H., Flora of the Montana formation: U. s', GeoI. I 

Survey Bull. 163, p. 68, 1900. 

Among fossil species the present form approaches 

in shape Oornus newbe!"'Yi Hollick,51 from the Fort 

Union beds of the lower Yellowstone in Montana. 

Oornus newberryi differs notably, however, in having 

8 or 10 or more pairs of subopposite secondaries. 1'he 

finer nervation is not shown. 

I have ventured to name this species in honor of the 

Rev. Arthur Lakes, who collected not only this leaf 

but a very large proportion of the material from 

these beds. 

Occurrence: Denver formation, quarry on northern 

base of Green Mountain, Golden, Colo. 

Cornus denverensis Knowlton, n. sp. 

Plate 13, Figure 1 

Leaf regularly oval, abruptly acuminate at the 

apex, evidently regularly narrowed to the base, per-

. fectly entire; midrib strong, straight; secondaries five 

pairs, opposite or subopposite at an acute angle, the 

lower ones slightly arching in, the upper ones much 

arched but not joining the midrib; nervilles numerous, 

percurrent, at right angles to the secondaries. 

'rhis small leaf, which unfortunately lacks the base 

and the greater portion of one side, is the only one 

found. It is perhaps doubtful whether or npt a 

new species should be founded on such scanty material, but there appears to be nothing identical with 

it in the collections. It is a small leaf, only about 

4.5 centimeters long and 2.5 centimeters wide, but it 

shows clearly the characters of Oornus. Among living 

species it resembles some of the small narrow leaves 

of 0O'rnU8 alternifolia Linne, also small leaves of O. 

1na8 Linne, from southern Europe. Among American 

species it is quite like Oornus pubescens val'. californica 

and small leaves of O. sericea Linne. Among fossil 

species its closest relative appears to be Oor'nus newberryi Hollick 58 [0. aouminata Newberry], from the 

Fort Union formation of the lower Yellowstone in 

Montana,.but O. denve'l'ensis differs in its smaller size 

and much less acuminate apex. It is also somewhat 

like Oornu8 'l'hawnifolia O. Weber, as figured by Lesquereux,59 from a locality near Point of Rocks, Wyo. 

Occurrence: Denver formation, Golden, Colo. Type 

specimen the property of Princeton University, No. 

2339 of the Princeton Museum. On same stone with 

Sterculia libbeyi. 

Cornus holmesii Lesquereux 

OornU8 1I,olme8ii Lesquereux, U. S. Geol. and Geog. Survey Terr. 

Ann. Rept. for 1873, p. 402, 1874. 

Following is Lesquereux's description of this form: 

The upper part of an ovate-Ianceolate entire leaf, with 

the secondary veins thin, very distant, alternate, much curved 

G1 Newberry, J. S., U. S. Geol. Survey Mon. 35, p. 124, pI. 28, figs. 

2-4, 1898. 

58 Idem, p. 124, pl. 37, figs. 2--4. 

GO Lesquereux, Leo, The Tertiary fiora, p. 244, pI, 42, fig. 6, 1878. 
120 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

in passing up in an acute angle from the middle nerve [midrib] toward the borders. The point of the leaf is broken. 

Though the specimen is fragmentary, it evidently represents 

a Oornus specifically distinct from other fossil species by the 

great distance of the secondary veins. BY' this character 

only is it distantly related to Oorl1.1us buchi Heel'. 

This species was not figured nor afterward referred 

to by Lesquereux and was probably merged with some 

other form without any synonymic record being made. 

The specimen on which it was based is not now known 

to be in existence. The locality is given as follows: 

"Bituminous shale, Coal Creek, Colo. ; W. H. 

Holmes." This is supposed to be the Coal Creek or 

Sand Creek about 15 miles east of Denver and is presumed to be in the Denver formation. It is mentioned in this place merely to keep the record clear. 

Nyssa lanceolata Lesquereux 

Plate 52, Figure 2 

NY8sa lanceolata Lesquereux, U. S. Geol. find Geog. Survey 

Terr. Ann. Rept. for 1872, p. 407, 1873; idem for 1873, 

p. 385, 1874; idem for 1876, p. 513, 187&; The Tertiary 

flora, p. 245, pI. 35, fig. 5 [not fig. 6], 1878. 

Knowlton, U. S. Geol. Survey Bull. 105, p. 56, 1899; U. S. 

Geol. Survey Prof. Paper 101, p. 343, p1. 108, fig. 1;. pI. 

. 113, fig. 2, 1917 [1918]. 

This species was founded on material from " Spring 

Canyon, Mont." [Meadow Creek, 12 miles southeast 

of Bozeman, Gallatin County], in beds presumed to 

belong to the Livingston formation. It appears to 

have been founded on fragments of leaves which are 

not now preserved in any known collection, and, moreover it does not seeln to have been collected there , . 

subsequently. 

The two additional references to the Hayden reports quoted in the above synonymy are only lists 

which refer to the type locality, so that the renlarks 

in the account of the Tertiary flora constitute the only 

additional utterance by Lesquereux concerning this 

species. Two figures were given at this time, one of 

a leaf and the other of a nnmber of fruits' which Lesquereux said he referred" hypothetically" to this 

species. The original of the leaf figured is No. 303 

in the United States National Museum collection and 

is preserved on the andesitic matrix characteristic of 

the Denver formation at Golden, Colo. Although 

this is not the actual type of the species, it is the first 

a.nd only leaf figured by Lesquereux and hence it 

becomes to all intents and purposes the figured type. 

It is the basis for the identification of the species 

elsewhere. 

As it is obviously impossible to connect the seeds 

with the lear just mentioneu, it has seemed best to 

give them a new name. They may now be called 

Nyssa rlewvel'iana. 

The leaf figureq. by Lesquereux as Nyssa lanceolata 

is very similar to certain of the leaflets he figured 60 

as Juglans rugosa, but it differs by its much lighter 

nervation-that is, it is less rugose. Another species 

figured by Lesquereux,61 Sapindius caudatus, is nluch 

more difficult to distinguish from Nyssa lmnceolata. 

Sapindus caud(J;t'W8, as its name implies, has a long, 

narrowly caudate apex and when complete is very 

easily identified. The leaf ,figured as Nyssa lamceolata 

lacks the apical portion, but it is difficult to note any 

differencve between the portion present and the similar 

portion of the figure of Sapindus cauaatus. 

I have figured two leaves from the Raton formation 

of the Raton Mesa region under the name Nyssa lanceolata.62 One of these (pI. 113, fig. 2) agrees well with 

the type figure in size, shape, and general character 

of the nervation but differs from it in the slightly 

nlOre acute divergence of the lower secondaries. The 

other specimen is slightly nl0re wedge-shaped at the 

base but does not differ markedly from the first. 

The collection from Mosby, Colo., contains the single 

example here figured, which is certainly indistinguishable from Plate 113, Figure 2, of the Raton report 

above cited. 

Occurrence: ,Denver formation (figured type), 

South Table l\1ountain, Golden, Colo., collected by Leo 

Lesquereux about 1873. Raton formation, canyon 

west of Mayne, Colo., and Purgatoire Canyon at mouth 

of Riley Canyon, Colo., collected by W. T. Lee. Dawson arkose, dump of Mosby coal mine, Mosby, Colo., 

collected by F. H. Knowlton and G. B. Richardson 

in 1910. Livingston ( ~) formation, Spring Canyon, 

l\lont. [Meadow Creek, 12 miles southeast of Bozeman, 

Gallatin County]. 

Nyssa dellveriana Knowlton, n. sp. 

Nyssa lanceolata Lesquereux .. Lesquereux, The Tertiary flora, 

p. 245, pI. 35, flg. 6 [not fig. 5], 1878. 

Following is Lesquereux's description of the seeds 

of this species as given under the name Nyssa lanceolat(}): 

Fruit comparatively large, preCisely ovate, round on one side, 

obtusely pointed on the other, deeply, distantly costate, and 

striate. 

In discussing the fruits he continued: 

The fruits, which I 'refer hypothetically to the same species 

[Nyssa lanceolata] for the reason that both kinds of organs 

were found at the same place, lie * * are 15 millimeters 

long and 8 millimeters broad below the middle, marked by 

regular small costae, with the intervening space fiat, and nar· 

rowly lined. For its size the fruit is comparable to some of the 

fossil seeds described by European authors, especially to N. 

60 Lesquereux, Leo, The Tertiary flora, p. 286, pI. 55, 1878. 

61 Idem, p. 264, pI. 48, flg. 6. 

6~ Knowlton, F. H., U. S. GeoI. Survey Prof. Paper 101, p. 343, pI. 

lOS, fig. 1; pI. 113, fig. 2, 1917 [1918]. 
SYSTEMATIC PALEOBOTANY 121 

maminw, Weber, N. ul'ctica Heel', N. ornithobroma Unger, etc. 

'l'he position of the two fl'uits upon the sftme specimen seems to 

indicate them as originally in a cluster. 

The specimen from which the figure of the fruits 

lVas made is preserved in the United States National 

Museum (No. 506). It is recorded in the Museunl 

catalogue as having come from Black Buttes, Wyo., 

but the matrix proves conclusively that it came from 

the andesitic beds of the Denver formation, presumably from Golden, Colo., as this is the locality given by 

Lesquel'eux. The fruits are now very obscure, having 

evidently been much abraded since the figure was 

made. None have since been noted in beds of the 

Denver formation .. 

'rhe National Museum, however, contains a series 

of fine specimens (No. 3579) of this species collected 

by T. W: St~nton from the coal bed at Parker's mine, 

3 miles from White Oaks, N. Mex. They agree jn 

every particular with Lesquereux's figures, although, 

as already stated, the type specimen is very obscure. 

The age. of the coal-bearing rocks at White Oaks is 

st.ill undetermined.68 Additional specimens from the 

White Oaks locality were obtained by C. H. Wegemann, of the United States Geological Survey, in 1912. 

They are now preserved in the National Museum. 

Occurrence: Denver formation, Golden, Colo., collected by Leo Lesquereux, about 1873. Tertiary ( ~) , 

Parker's coal mine, 3 miles from White Oaks, N. Mex., 

collected by T. W. Stanton in 1889 and by C. H. 

Wegemann in 19.12. 

Nyssa? obovata Knowlton, n. sp. 

Plate 54, Figure 1 

Leaf of firm texture, obovate, broadest above the 

middle, whence it is broadly wedge-shaped to the base 

and rather abruptly acute at the apex; margin entire 

for lower two-thirds, coarsely few-toothed above; midrib strong, straight; secondaries eight or nine pairs, 

alternate, at an angle of about 45°, very slightly 

curved upward, camptodrome, becoming craspedochome above and passing to the teeth; nervi lIes obscure, much broken, and forming large irregular areas; 

finer nervation not preserved. 

The example figured is nearly perfect. It is strongly 

obovate with an obtusely wedge-shaped base and obtusely acute apex. The margin above bears a few 

rather large, coarse teeth. The length of the leaf is 

about 9 centimeters and the width a little more than 5 

centimeters. 

It is with some hesitation that this leaf is referred 

to the genus Nyssa, yet it is strongly suggestive of certain others that have been so considered. Among the 

living American species it has much the shape of N. 

II See Wegemann, C. H., U. S. Oeo1. Survey Bull. 541, p. 428, 1914. 

bifim'a Walter, and in the matter of toothing it is 

nearest to N. (f)quatica. Linne. Among fossil forms it 

appears to approach most closely Nyssa? euneata Newberry 64 from the Upper Cretaceous of Orcas Island, 

Washington. 

Occurrence: Denver fonnation, Golden, Colo., south 

face of South Table Mountain, 100 feet below the lava 

cap, collected by Arthur Lakes, 1890. 

Order EBENALES 

Family EBENACEAE 

Diospyros brachysepala AI. Braun 

Plate 51, Figure!? 5, 6, 7, 9; Plate 52, Figure 3 

Diospyro8 braohysepala AI. Braun, Die tertHil' Flora von Oeningen: Neues Jahrb., 1845, p. 170. 

Lesquereux, U. S. Geol. Survey Terr. Ann. Rept. for 1872, 

p. 394, 1873; idem for 1873, p. 401, 1874; idem for 1874, 

p. 306, 1876; idem for 1876, p. 511, 1878; U. S. GeQl. and 

Geog. Survey Terr. Bull., vol. I, p. 367, 1875; The Tertiary flora, p. 232, pI. 40, figs. 7-10, pI. 63, fig. 6, 1878; 

The Cretaceous and Tertiary floras, p. 174, pl. 34, figs. 1, 

2, 1883; U. S. Nat. Mus. Proc., vol. 10, p. 41, 1887. 

Proof that all the leaves in Europe and America 

that have been referred to Diospyros b1YfJehyseptila 

really belong to this species, if obtainable, would show 

not only that it is an exceedi~gly polymorphous species but that it has an extraordinarily wide geographic 

and geologic range. As a matter of fact, it will probably ultimately be shown that perhaps as many as half 

a. dozen species have been confused under this name, 

but until larger series of specimens can be brought 

together it is best to make only such segregation as can 

be pretty well established. 

The type locality of Diospyro8 brachsepaia is the 

Miocene of Oeningen, in Baden.65 The plate devoted 

by Heel' 66 to this species shows a considerable range 

in size and shape. His diagnosis reads as follows: 

Foliis petiolatis, ellipticis, utrinque atte'nuatis, membranacels 

vel subcoriaceis, integerrimis, nervis secundariis aIternantibus, 

l'emotiusculis, subangulo acuto egredientibus, curvatis, ramosis. 

Most of the leaves figured by Heel' are elliptical in 

shape and about equally narrowed to the base and 

apex. Several are distinctly lanceolate, with a rather 

long, sharp-pointed apex and a wedge-shaped base. 

A single leaf is distinctly obovate. The nervation is 

very much the same regardless of shape-that is, it 

61 Newberl'y, J. s., U. S. Geo1. Suryey Mon. 35, p. 125, pI. 17, figs. 4, 5, 

1898. 

6Ii The locality of Oenlngen, long and widely known for its deposits 

of wonderfully preserved plants, insects, fish, etc., Is usually given as 

being in Switzerland. Recently T. D. A. Cockerell, of the University 

of Colorado, visited the region and determined that the celebrated 

fossil beds are located near the town of Wangen, wbich is on the 

north bank of the Rhine in tbe German Province of Baden. See 

Scientific Monthly, vol. 1, pp. 287-291, 1915. 

66 Heer, Oswald, Flora tertiarln Helvetlne, ,01. 3, p. 11, pI. 102, figs. 

1-14, 1859. 
122 FLORA OF THE DENVER AND ASSOCIATEb FORMATIONS OF COLORADO 

consists of a fairly thick midrib and 8 to 12 pairs of 

thin, usually alternate, camptodrome secondaries, 

usually at a low angle of origin. The leaves range 

from 4 to 9 centimeters in length and 1.5 to 3.5 centimeters in width. 

As already indicated, this species has been widely 

identified in European rocks which range in age practically throughout the Tertiary. 

Diospyros brachysepala was first noted in American 

rocks by Lesquereux,67 who identified it at Black 

Buttes, ,Wyo. The following year Lesquereux 68 reported this species from Sand Creek, Colo., in beds 

then supposed to be Miocene but now known to belong 

to the Denver formation. Re apparently had only a 

single leaf, which he described as follows: 

Leaves broadly oval or slightly obovate, narrowed in a 

curve to the base, entire, rather membranaceous, but not thick; 

secondary nerves alternate, curving to and along the borders, 

mostly simple or with few branches, deflected downward in 

reaching the middle nerve. 

Lesquereux especially compared this leaf with Figure ,6 of Reer's plate for size and shape and Figure 2 

for nervation. 

In 1874 Lesquereux identified Diospyro8 braahyljepala at Point of Rocks, Wyo., comparing the leaves 

from this locality with Figure 2 of Reer's plate, which 

is a long, narrow, sharp-pointed leaf. The remarks 

concerning these Point of Rocks leaves were republished without change in 1876.69 

In 1878 Lesquereux 70 figured for the first time the 

v~rious specimens he had referred to Diospyros brachysepala. They were said to have come from Golden, 

Colo., and Black Buttes and Point of Rocks, Wyo. Of 

the five figures given, only three appear to be represented by specimens in the Unit.ed States National Museum collections. These are Figures 7 (No. 339), 8 (No. ' 

338), and 10 (No. 337) of his Plate 40. FigUre 7 is said 

to have come from Black Buttes, Figure 8 from Point 

of Rocks, and Figure 10 from Golden. None of these 

leaves are very well preserved; the best one is that of 

Figure 7, Which in size and shape is very similar to 

Figure 8 of Reer's plate,71 but the secondaries are at a 

slightly more acute angle. The specimen shown in 

Lesquereux's Figure 8, from Point of Rocks, is only the 

tip of a very small leaf that mayor may not be referable to Diospyros brachysepal(J). The Point of Rocks 

material is mentioned again below. Figure 10 of Lesquereux's plate shows a very fragmentary specimen, 

67 Lesquereux, Leo, U. S. Geol. and Geog. ~urvey Terr. Ann. Rept. 

for 1872, p. 394, 1873. 

88 Idem for 1873, p. 401, 1874. 

69 Lesquereux, Leo" On some new species of fossil plants from the 

Llgnitic formations: U. S. Geol. and Geog. Survey Terr. Bull., vol. 1, 

No.5, 2d ser., p. 367, 1876. 

70 Lesquereux, Leo, The Tertiary flora, p. 232, pl. 40, figs. 7-10; PI. 

63, fig. 6, 1878. 

11 Heer, Oswald, Flora tertlal'ia Helvetiae, vol. 3, p. 11, pI. 102, 1859. 

lacking both base and apex and much of one side. Its 

reference to this species can hardly be considered 

firmly established. In 1883 Lesquereux 72 admitted 

Diospyros brachysepaZa as present in the Miocene 

Florissant lake beds of Colorado. Re figured two fine 

leaves that are now preserved in the museum of 

Pri,nceton University. They are elliptical-obovate, 

and, as Lesquereux said, are" more positively identified with the European species" than those figured in 

the report on the Tertiary flora. 

In passing it may be noted that the leaf described 

by Cockerell 78 under the name Diospyros princetonia 

does not seem to me to be distinguishable from the 

lea ves from Florissant figured by Lesquereux as 

Diospyros brachY8epala, though Cockerell says: " Both 

apex and base are more pointed than in normal D. 

oracNy8epala."· . 

:fwo leaves from the Fort Union formation of the 

lower Yellowstone in Montana were referred by 

Ward 74 to Dio8pyros braahY8epala. They both have 

a distinctly decurrent base and I suspect should be 

referred to Sapindu8 ala-tus Ward [ S. glendJivensis 

Knowlton] ,75 though otherwise they agree pretty 

closely with Reer's figures of D. br(iohY8epa7AJJ. 

In 1900 I reviewed the flora of the Montana group 76 

and referred a single leaf to Dio8pyro8 bracNy8epala. 

It is a narrowly lanceolate, long acuminate leaf very 

unlike most of the lea ves usually referred to this 

species, though not greatly unlike the narrow sharppointed leaves so referred by Heer. On again studying this specimen I am convinced that I was in error 

in referring it to Diospyro8 bracNysepala/ in fact, 

even its reference to Dio8pyros is considered questionable. 

The presence of Dio8pyros brachysepala in the Wilcox group of the Southern States has been reported 

by Berry.77 Of the three leaves figured by him two 

(figs. 3 and 6) appear to be referred to this species 

with a considerable degree of certainty, but the other 

one (fig. 4) seems to me to belong with Diospyro8 wilcowiana Berry, as figured on the same plate. Although 

it is slightly smaller than one of the figures of D. wil-

(Joanana (fig. 1) it has the same markedly decurrent, 

base and the same nervation. 

To recapitulate concerning the presence of Diospyro8 

brachysepala AI. Braun in 'American rocks: The speci72 Lesquereux, Leo, The Cretaceous and Tertiary floras, p. 174, pl. 84, 

figs. 1, 2, 1883. 

73 Cockerell, T. D. A., Am. Mus. Nat. Hist. Bull., vol. 24, p. 105, pI. . 

10, flg. 36, 1908. 

74. Ward, L. F., U. S. Geol. Survey Sixth Ann. Rept., p. 556, pI. 60, 

figs. 4, 5, 1886; U. S. Geol. Survey Bull. 37, p. 104, pI. 49, figs. 1, 2. 

1887. 

76 Ward, L. F., op. cit. (Sixth Ann. Rept.), pI. 50, figs. 9, 10. 

76 Knowlton, F. H., U. S. Geol. Survey Bull. 163, p. 74, pI. 18, flg. 3, 

1900. . 

T1 Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 333, pl. 101, 

figs. 3, 4; pl. 107, fig. 6, 1916. 
SYSTEMATIC PALEOBOTANY 123 

mens identified by Lesquereux from Black Buttes, 

'Vyo., and Golden, Colo., should probably be accepted, 

but the fragment from Point of Rocks, Wyo., is too 

small and doubtful to be of convincing value. The 

two leaves from the Miocene at Florissant, Colo., 

identified by Lesquereux are apparently correct gen-

~l'ically but are open to question specifically. The 

leaves from the Fort Union formation of the lower 

Yellowstone in l\10ntana, as determined by Ward, 

should probably be referred to Sapindus alatus Ward 

(S. glcndiAJensis Knowlton). The si~gle leaf identified by me in the Mesaverde formation at Point of 

Rocks, Wyo., is not Diospyros brachyscpala and should 

be given a new name. Of the three leaves from the 

Wilcox group figured 5y Berry, two may be accepted 

as Diosp1jros brachyscpala and the other referred to 

D. wilcowiana Berry. 

We may now consider the leaves that the present 

study has brought to light in the Denver Basin and 

adjacent areas. Several leaves have been found in 

the Denver formation at Golden that may be referred 

to Diospyros brachysepala as currently accepted. One 

of these is shown in Plate 51, Figure 6. It is ovateelliptical, about 5.5 centimeters long and nearly 3 

centimeters wide. The midrib is fairly strong, but 

the secondaries are thin and at a low angle. 

The leaf shown in Plate 51, Figure 5, .is from the 

Dawson arkose near Mosby, Colo. It is practically 

perfect, including the petiole, which is 1 centimeter 

long. It is elliptical, with a sharp-pointed apex. Its 

length is 6 centimeters and its width 2.5 centimeters. 

Leaves that are referred to Diospyros braehysepala 

have also been found in the lower part of the Dawson 

ar kose . near Sedalia, Colo.; they ha ve not been 

figured. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Dawson arkose, dump of l\10sby coal mine, Mosby, Colo., 

collected by F. H. Knowlton and G. B. Richardson, 

1910; also in lower part of Dawson arkose near Douglas coal mine, Sedalia, Colo., in beds equivalent to 

Arapahoe formation, collected by Arthur Lakes, 1890. 

Family STYRACACEAE 

Styrax? laramiensis Lesquereux 

Plate 52, Figure 7 

StyraaJ Zamtniens6 Lesquereux, Harvard ColI. Mus. Compo Zoology Bull., vol. 16, p. 51, 1888. 

Leaf subcoriaceous, smooth on the surface, elliptical, apparently about equally rounded to both base 

and apex (apex absent), abruptly slightly decurrent 

at base; petiole evidently strong; margin perfectly 

entire; nervation pinnate but 3-ribbed in eff€ct, midrib very strong, straight, with four or five pairs of 

secondaries, the lowest pair nearly as strong as the 

midrib, arising some distance above the base of the 

blade, at an angle of about 45°, slightly curved upward, camptodrome, each with four or five tertiary 

branches on the outside, all camptodrome and joining 

along the border, upper secondaries alternate, camptodrome; nervilles numerous, thin, mainly broken. 

This species was based on a single specimen by Lesquereux-the one here figured. It is a small ovaloI' 

elliptical leaf that must have been about 5.5 centinleters long and 3.5 centimeters wide. Lesquereux 

apparently did not note that it is slightly decurrent 

on the stout petiole or that the lowest pair of secondaries are nearly as strong as the midrib and arise 

above the base of the blade, thus producing a pseudopalmate aspect. 

There is some question as to the proper generic reference of this leaf. Lesquereux states that' it is related to the European Styrt1w offiainailis Linne, but it 

does not greatly resemble the American species, espe· 

cially as they usually have more or less toothed or 

lobed leaves and this is perfectly entire. In some 

respects it strongly resembles certain of the leaves referred to FiC'UJ8 pZa11ficostataJ Lesquereux,78 but this 

never has the blade decurrent, only rarely are the lowest pair of secondaries inserted above the top of the 

petiole, and the nervilles are not of the same character 

as in StyraaJ'I laramien$is. 

There is also some resemblance between this leaf and 

certain leaves referred to O'teodaphne, as, for instance, 

small leaves of O. obt'W8ifolia Berry,79 from the Wilcox 

deposits of Mississippi and Louisiana. This species 

is often slightly decurrent at the base, and the lowest 

secondaries always oarise well above the base of the 

blade, but they never branch on the outside like the 

"ribs" in Styrax? laramiensis. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by. Arthur Lakes. Typ~ 

in Museum of Comparative Zoology, Cambridge: 

Mass., No. 1488. 

Order OLEALES 

Family OLEACEAE 

Fraxinus eocenica LesquerEmx 

PIa te 52, Figures ~ 

FraaJinu8 eocenica Lesquereux, The Tertiary fiora, p. 229, 1878; 

The Cretaceous and Tertiary fioras, p. 123, pI. 20, figs. 

1-3 [figures here r~produced], 1883. 

The type specimens of this fine species are the property of Princeton University and are preserved in the 

Princeton Museum of Natural Histor:y. It appears 

78 Lesquereux, Leo, The Tertiary flora, p. 201, pl. 31, figs. 1-8, 10--12, 

1878. 

7D Berry, E. W., U. S. Geol. Survey Prof. Paper 91, p. 301, pI. 83, 

fig. 3, 1916. 
124 FLORA OF THE 'DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

to be a rare species, for so far as I can learn no others 

have been found. It was not reported by Lesquereux 

. in the large collection from Golden in the Museum of 

Comparative Zoology of Harvard University, nor does 

it appear to be present in the still larger collections in 

the United States National Museum. 

This species was described and discussed by Lesquereux as follows: 

Leaves of large size, subcol'iaceous, distantly obtusely dentate or merely undulate on the borders, broadly lanceolate, 

rounded in narrowing to the inequilateral base; nervation 

subcamptodrome. 

The largest of these leaflets is about 15 centimeters long 

and 5 centimeters broad below the middle, where it is the 

widest; the other leaflets are somewhat smaller but exactly of 

the same form. The secondary nerves, on an open angle of 

divergence, nearly parallel and eqUidistant, generally curve 

in passing toward the borders, where they form, close to them, 

a series of'simple bows, from which emerge the nervilles which 

enter the blunt distant teeth. '" '" * The nervilles are distinct, irregular in direction, much divided; the areolation, as 

far as it is discernible, is in irregular polygonal meshes. The 

base vf the leaf is cuneate on one side, descending lower and 

rounded on the other, or rounded on both sides, which then 

are nearly equal; the upper part is gradually tapered upward 

and then contracted to a slightly obtuse acumen, as in the 

leaves of FraaJinus pubescens Lamarck and F. americana Linne, 

to' which this fossil species is related. 

In some respects F'J'awinus eocenic(J) suggests certain 

leaflets of Juglarns denver'iam.a as described above, 

especially in the shape of the base, but it differs in 

being relatively shorter and broader and more particularly in the margin, which is undulate or strongly 

toothed in F. eocenic(t and entire or obscurely serrate 

in J. denve1'iama . . Also, the secondaries in F. e'ocenica 

emerge at a lower angle and are somewhat less curved 

upward than in J. denveriana. 

Occurrence: Denver formation, Golden, Colo., collected by Arthur Lakes; about 1875. 

Fraxinus sp. 

Plate 58, Figure 7 

FraaJinus aenticulata, Heel'. Lesquereux, Harvard CoIl. Mus. 

CO'mp. Zoology Bull., vol. 16, p. 51, 1888. 

In the collection of Denver plants identified, by Lesquereux for the Muselun of Comparative Zoology he 

listed with question a single specimen under the name 

PTa-minus (~enticulat(t Heer. He wrote of it as tollows: 

A. fragment only, the upper part of a small O'blO'ng-O'val leaf, 

with the borders slightly denticulate, the lateral nerves mostly 

craspedodrome, entering the teeth. • • • The identity is 

not positively ascertained. 

This is a mere fragment of the upper portion of a 

leaf that must'have been about 5 centimeters long and 

2.5 centimeters in width. So far as can be made out it 

resembles FrarIJinU8 denti(}1J)lata Heer ,80 from the Mio80 Heer, Oswald, Flora fossills arctica,' vol. 1, p. 118, pI. 16, fig. 4, 

1868. 

cene of Greenland, but it is so fragmentary t.hat the 

identification can not be made positive. The specimen 

has been figured here in the hope that it will be sufficient to call attention to the presence of a type of 

foliage not otherwise represented. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo., collected by Arthur Lakes. Type 

in Museum of Comparative Zoology, Ca,mbridge~ 

Mass, No. 1487. 

Order TUBIFLORAE 

Family BIGNONIACEAE 

Dombeyopsis obtusa Lesquereux 

D011'tbeyo1J8'is obtt,sa Lesquereux, U. S. GeO'l. Survey Terr. Ann. 

Rept. for 1872, p. 375, 1873; i4ilem for 1873, p. 382, 1874; 

idem for 1876, p. 514, 1878; The Tertiary flora, p. 255, 

pI. 47, flgs. 4,5, 1878. 

Knowlton, U. S. Geol. Survey Prof. Paper 130, p. 162, pI. 

13, fig. 4; pI. 20, fig. 11; pI. 27, figs. 1-4, 1922. 

This species was named and described from specil11ens obtained about 1871 from a locality known as 

Gehrung's coal mine, about 15 Iniles southeast of Colorado Springs, Colo., in the Laramie formation. These 

specimens can not now be found. They are not in the 

lTnited States National Museum, unless there has been 

an error in recording them, nor has the species been 

noted in material recently obtained at or near the 

original loc'itlity. 

The two specimens figured as types of this species 

are preserved in the United States National Museum 

(Nos. 380, 381), and, if correctly labeled, both came 

from the true Laramie at Golden, Colo. In, any event 

they are preserved on the hard, white sandstone matrix characteristic of the Laramie, whether they came 

from Golden or not. The further occurrence of this 

species in the true Laramie is set forth at some length 

in my report on the Laramie flora 81 and need not be 

1 epeated here. 

D01nbeyopsw obtu8a is one of the few species kllo~n 

to cross the line between the Laramie and Denver or 

Dawson arkose. A very fine leaf of this species was 

found in the andesitic beds of the Denver formation 

at Golden. It was preserved on the same piece of 

lnatrix as the type of Ficus occidentalis Lesquereux 

(see p. 68), which is No. 281 of the National Museum 

collection, but as the specimen was somewhat irregular an attendant who attempted to reduce its size 

entirely removed this leaf of D01nbeyopsis obfiu,$a.. 

This is much to be regretted, for it proved to be one 

of the most perfectly preserved examples known. It 

was nearly circular in outline, being a.bout 9.5 centimeters long and 10 centimeters wide. 

Among the collections made by Lakes at the old 

Douglas coal mine, near Sedalia, Colo., there a,re three 

81 Knowlton, F. H., U. S. Geol. Survey Prof. Paper 130, p. 162, 1922. 
SYSTEMATIC PALEOBOTANY 125 

or four somewhat fragmentary specimens that appear 

to belong with D01nbeyopsis obtusa. They represent 

only basal portions of the leaves, but so far as I can 

see they are not to' be distinguished from this species. 

Occurrence: Laramie formation (type), Gehrung's 

coal mine, southeast of Colorado Springs, Colo.; figured types, Golden, Colo.; also found at Coal Creek, 

Marshall, Hoyt's coal mine, 1 mile south of Golden, 

and Crow Creek, 25 miles northeast of Greeley, Colo. 

Denver formation, South Table Mountain, Golden, 

Colo. Dawson arkose, Pulpit Rock, near Colorado 

Springs, Colo.; also lower part of Dawson arkose (in 

beds equivalent to Arapahoe formation), 3,000 feet 

cast of Douglas coal mine, Sedalia, Colo. 

Dombeyopsis magnifica Knowlton, 'n. sp. 

Plnte 54, Figure 4; Plate 55, Figure 4; Plate 57, Figure 1; 

Plate 58, Figure 8 

DombeyolJsis gl'((,JuUfoli(t Unger'l Lesquereux, U. S. Geol. and 

Geog. Survey 'rerr. Ann. Rept. for 1873, pp. 382, 404, 

1874; idem for 1876, p. 514, 1878; Tbe Tertiary fbm, 

p. 255, pI. 47, fig. 6, 1878. 

Leaves large and thick, roughly deltoid, from truncute and slightly heart-shaped to deeply heartshaped at the base, where the margin is entire, other 

portions of the margin cut into four or five very large, 

deltoid, rather obtuse lobes on a side, the apical lobe 

sjmilar; the lowest lobe is turned downward, the 

others outward or upward; petiole exceedingly large 

and strong, split at the base of the blade into seven 

OJ' ni:Je strong radiating ribs, all except lowest ones 

ending iI). lobes; midrib slightly the strongest, with 

three or four pairs of secondaries, one or two of which 

end in marginal lobes; next pair of secondaries at an 

angle of about 45°, craspedodrome, with one or more 

branches entering lobes; next pair slightly smaller, a 

little above a right angle in insertion; in 7 -ribbed 

leaves the lower ribs to enter the lowest lobe have several camptodrome secondaries on the lower side, the 

first one arising with the large ribs and thus producing the 7 -ribbed appearance; in 9-ribbed leaves 

the lowest pair of ribs fall below a right angle, and 

here the first secondaries arise with the ribs, thus producing this 9-ribbed eff.ect; nerviUes numerous, strong, 

not often broken, approximately at right angles to the 

ribs or their secondaries. 

This splendid species is represented by half a dozen 

or more examples, two of them being exceptionally 

perfect. One of these, shown in Plate 57, Figure 1, 

is about 10 centimeters long and was not far fr01n 

16 centimeters wide. It has four lobes on a side, 1 

to 3 centimeters long and separated by broad, shallow sinuses. The petiole is not preserved in this 

F.pecimen. The other nearly perfect example is shown 

86492-30--9 

in Plate 58, Figure 8. It is about 11 centimeters long 

and 14 centimeters wide and has the petiole preserved 

for a length of nearly 8 centimeters. It also has four 

prominent lobes on each side, but they are longer 

and more sharply pointed, especially the terminal 

lobe. 

The ot.her specimens have the base well preserved, 

but otherwise they all lack most if not all of the margin. They are much larger than those first mentioned, 

one shown in Plate 54, Figure 4, for instance, being 

at least 16 centimeters long and when perfect probably not far from 20 centimeters wide. This specimen has the petiole preserved for a length of about 3.5 

centimeters, and in the one shown in Plate 57, Figure 

1. the petiole is preserved for over 5 centimeters. This 

specimen was clearly more than 20 centimeters wide, 

but its length can only be conjectured. It is safe to 

,say that these leaves were from 15 to nearly or quite 

25 centimeters wide and from 10 to 16 or 18 centimeters long. 

This species is undoubtedly congeneric with Dom-. 

beyopsis t'1'ivialis Lesquereux,82 from the Laramie formation at Golden, Colo. The Laramie species differs 

in being more nearly square in general outline, much 

smaller, and only 5-ribbed. It is only 7 centimeters 

long and less than 8 centimeters wide and has only 

one or two large lobes on each side of the blade. It is 

also more deeply heart-shaped at the base than D. 

magnifica. 

In 1873 Lesquereux referred with question a single 

fragmentary leaf from the andesitic beds of South 

Table Mountain, Golden, Colo., to the European species D01nbey'opsis gramdilolia Unger. This was figured and further described by Lesquereux in 1878,83 

but no additional examples appear to have been found 

at Golden. This specimen-No. 38~ of the United 

States National Museum collection-is a mere fragment from the base of what ,vas apparently a large 

.leaf. It sho,YS a portion of a very thick petiole, somewhat expanded at the apex, where it gives rise to six 

strong, nearly equal, radiating ribs. The base of the 

blade appears to be heart-shaped, but this point is obSClu·e. It seems probable that there were at least seven 

ribs when this leaf was pe'rfect, for" six is an unusual 

number. On comparison this Golden fragment is 

practically indistinguishable from some of the leaves 

here described as D01nbeyop8is 7nagnifica, especially 

the one shown in Plate 57, Figure 1. It has the same 

thick petiole, strong, radiating ribs, and coarse nervilles. The size, as nearly as I can judge, must have 

been about the same in both. Dombeyopsis grandifolia Unger is a handsome species from the Miocene of 

B2Lesquereux, Leo, The Tertiary tIora, p. 255, pI. 47, fig. 3, 1878. 

53 Idem, p. 255, pI. 47, fig. 6. 
126 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Styl'ia and Bohemia. Unger 84 devoted two plates to 

this species, but, as Schimper long ago pointed out, 

,all but one of 'Unger's figures probably represent the 

polymorphous FiouJs' tiliaJefoliaJ. The exception is 

Figure 1, Plate '27, which is the one most. like what I 

have here called D. 1nagnfific(J). It is so fragmentary 

that no part of the margin except the base is to be 

seen. The size of the blade, number and disposition 

of the ribs, nervilles, and petiole are like those features 

in D. 17WJgnifio(J), but it is impossible to compare the 

configuration of the margin in the two forms. 

Occurrence: Dawson arkose, a quarter of a mile 

east of the Purdon mine, sec. 27, T. 11 S, R. 61 W., 

Colo., collected by G. B. Richardson. 

Dombeyopsis? sedaliensis Knowlton, n. sp. 

Plate 56, Figure 5; Plate 59, Figure 1 

Leaves large, apparently rather thin and membranaceous, nearly circular or possibly very broadly 

ovate, markedly heart-shaped at the base (apex unknown) ; margin entire; petiole very strong ~nd long; 

5-ribbed from the extreme base of the blade, the ribs 

of about equal strength; midrib with two or three 

pairs of strong, alternate secondaries in the middle 

and upper portions of the blade; next pair of ribs 

arising at an angle of about 45°, considerably curving 

upward, each with three or four secondary b~'anches 

on the lower or outer side; lowest pair of ribs arising 

nearly at right angles to the midrib, soon curving 

slightly or sometimes considerably upward, each with 

several camptodrome secondary branches on the lower 

side which supply the rounded basal lobes of the blade ~ 

nervilles numerous, mainly unbroken, mainly at right 

angles to the ribs or to the secondary branches. 

This species is represented apparently by several 

specimens, but unfortunately they are so fragmentary 

that they are made out with difficulty. It is a largeleaved species, being some 10 or 12 centimeters in 

width and probably a little longer. The petiole is 

very strong, being fully 3 millimeters thick in some 

specimens where it merges into the Llade. The length 

of the petiole is at least 6 or 7 centimeters. 

It is with some hesitation that these leaves are referred to the genus D01noeyopsis, but they are found 

jn intimate association with other species of this genus 

which they closely resemble and are therefore tentatively referred to it. At first it was supposed,that 

they belonged to D01noeyopsis ootusa, wit.h which they 

.are associated, but they differ in having five instead of 

three ribs; otherwise the differences are not great. 

Occurrence : Dawson arkose, lower part (in beds 

-equiv~lent to Arapahoe formation), 3,000 feet east of 

the Douglas coal mine, Sedalia, Colo. ' 

RI TInge)', Frauz, Fossile Flora von Sotzlw: K. Aknd. Wiss. Wien 

Denkschr., vol. 2, p. 175 (45), pIs. 47 (26),48 (27),1850. 

Order R UBIALES 

Family CAPRIFOLIACEAE 

Viburnum richardsoni Knowlton, n. sp. 

Plate 52, Figure 8; Plate 53, Figures 1, 3, 4, 5; Plate 54, Fig. 

ures 2,3 

Gt'ataegu8 betu,zaetoUa Lesquereux, Harvard 0011. Mus. Oomp. 

Zoology Bull., vol. 16, p. 56, 1888 [part]. 

Leaves evidently very firn1, broadly ovate to nearlv 

circular, generally more or less obtusely wedge-shape~l 

at. the base, rounded and rather obtuse at the apex; 

margin regularly and finely toothed throughout except at the very base, the teeth deltoid in shape; petiole 

long, relatively strong; nervation strong, deeply impressed, consisting of a markedly zigzag Inidrib and 

four to six pairs of strong, alternate, straight secondaries arising at an angle of 60° or 70°, the lowest 

pai~' arising at the very base of the blade, each with five 

or SIX strong branches on the lower side, some of which 

are fork~d, all entering the marginal teeth; upper 

secondarIes close, parallel, with one or two branches 

which enter the teeth; nervilles numerous, strong, at 

right angles to the secondaries or their branches. 

This species is the dominant form at Palmer's ranch 

and is represented by more than 25 more or less 

perfect specimens. As shown by t.he examples selected 

for illustration there is considerable range in size and 

a minor range in shape. The smallest specimen (pI. 

52, fig. 8) is 4 centimeters long and a little less than 3 

centimeters wide. The longest specimen is 8.5 centirneters long and was probably not less than 6 or 7 

(;entimeters wide; this particular specimen has the 

petiole preserved for a length of 2 centimeters. A 

still larger specimen not figured is fully 10 centin1eters 

long. The range in the shape of the base is 'also well 

shown in the five examples figured. Thus the specimen 

shown in Plate [>,2, Figure 8, is n1arkedly wedgeshaped, that shown in Plate 53, Figure 3, is n1uch less 

wedge-shaped, and that shown in Plate 54: Figure 3, 

is so much rounded that the whole leaf is nearly circular. This last-ment.ioned specimen is the n10st perfectly preserved of the lot. It is 5 centilneters lono-

(exclusive of the petiole, which is 2 centimeters long)' 

and about 4.5 ce.ntimeters wide. The rather obtuse 

apex is brought out in four of the examples figured. 

As may be seen from the. excellent figures this species 

iF; extremely well marked. It is clearly distinct from 

any species heretofore described from the Denver formation, though some of the forms approach some of 

the named fOrIns rather closely. Thus Plate 53, Figllre 3, suggests what has been identified by Lesquereux 85 as Viou1"nWln dichoto1nWln, from Black 

Buttes, Wyo. This diffei's in having fewer, smaller 

rnarginal teeth and fewer, more irregular secondaries. 

85 Lesquereux, Leo, The 'l'ertinl'Y flora, p. 225, pI. 38, fig. G, 1878. 
SYSTEMATIC PALEOBOTANY 127 

The perfect specimen seen in Plate 54, Figure 3, is 

very similar to an undescribed species from the Livingston formation of l\10ntalUt: at least in general 

appearance. The Livingston species, however, has a 

more truncate base, minut.e marginal teeth, and 

s1ightly different nervation. 

Since the above was written I have studied the types 

of O'ra.taegus bet'ulaefolria Lesquereux in the l\1useum 

of Comparative Zoology, a,nd I have found that one 

of tlH'se is undoubtedly ide.ntical with TT ib'U1"nWln 

1·icl~a1·d80111t. ,"!"'his specimen is practically complete, 

even to the major part of the petiole. It is shown in 

Plate 5B, Figure 4. It is 4 centimeters long and 3 

centimeters wiele and agrees very well with the figure 

of TTiburm.t1n ')'tohm'dsonrt given on Plate 52, Figure 8. 

Occurrence: Dawson arkose, Palmer's ranch, Templeton Gap, 4 m:ileH lIorthenst of Colorado Springs, 

Colo., about 40 fpet above base of section, collected by 

F. JI. I(nowltoll, ,July, 1910; also lower part of Dawson 

arkose On beds equi.valcnt to Arapahoe formation), 

;-3,000 feet east of Douglas eoal mine, Sedalia, Colo. 

Denver formation, South Table l\10untain, Golden, 

Colo., collected by Arthur Lakes; specimen in l\/fuseum 

of Comparative Zoology, Cambridge, l\1nss. (No. 1576)) 

one of the types of 01'ataegut~ betulaefolia Lesquereux. 

Viburnum contortum Lesquereux? 

Plate 55, Figure 1 

Yibu.nt1tnt oontort-/(:II1. Lesquereux, U. S. Geol. Survey Terr. 

Ann. Rept. for 1872, p. 396, 1873. 

Knowlton, U. S. Geol. Survey Prof. Paper 101, p. 346, pI. 

108, 1ig. 3, 1017 [1918]. 

Vibnrnum nw.l'uinM,lt'ln I.esqueL·eux, The Tertiary flora, p. 223, 

pI. 38, fig. 2· [not flg. 3], 1878. 

In rny report on the flora of the Raton formation 811 

I gave the following account of the present disposition of this species: 

V';'bm'IMt1li contort'llm was described originally from Black 

Buttes, "\Vyo., where it wus found in association with such 

wull-lmown forms as lNmts 1)Z(l!nicostata, and Fic/f,.I! 1 a,t-;"o1A·a. 

At tho tilne it was founded I.esquereux stated tlwt it might 

perhaps be considered as merely a variety of Vibw'lllliU '11/(/.1'-

uinat'U1n [PIManlltS 1IUH'g'i1uftfa], and in "The 'I'<:,rtial'Y flora" 

lie so considered it und met'ged the name of V. contortmn 

with 1)w1'uinat'lt11t anel placed the two specimens figured (the 

types of oontOl't1tm) undet· that name. This disposition by 

f .. esquel'cux bas been accepted until the present time, though 

all who have compared the figures giYen undE'r Vibnl'rltUm nta1'-

uvnatum could hardly have failed to note the marked difference 

between them, particularly in shape, the olle being broadly 

obovate wHh a truncate 01' rounded Hl)ex and the other with 

an acuminate apex. If ]enyes of t~e cont01·tmlL type were 

only found in direct association with the ma1·u'ilw.fwn type, 

it might perhaps be permissible to allow them to remain 

together, but where ll'aves that are obviously of the type of 

oonfO'l-tmn are found at widely c}istant points where they 

86 Know) ton, F. H., U. S. Geo)' Survey Prof. Paper 101, p. 346, 1917 

(U)1S). 

are not associated with the 11W1'uinatwn type, it seems best to 

regard them as specifically distinct. 

A specimen that is almost exactly similar to Lesquereux's Figure 2 was found in the Raton formation· 

and is figured in the report above quoted. The only 

lloticeable difference is the possession of fewer, 

slightly larger teeth. 

The leaf figured here under the name V. contort'tt1n 

is rather fragmentary, lacking all of the apical portion. The base including the petiole is fairly complete, and the nervation is strongly preserved. The 

preservation of the Inargin is only sufficient to show 

. two or three of the lower sharp teeth. Although this 

leaf so far as it is preserved agrees perfectly with 

Lesquereux's figure, as well as with the Raton specimen, so much of the upper portion is wanting that it 

. 8eems best to qnestion the identification. 

Occurrence: Dawson arkose, Rice's clay bank~ 1 

mile southwest of Calhan, Colo., collected by F. H. 

I(nowlton~ .July, 1910. Raton formation, Rockland 

mine, Cuehal'as Canyon, Colo. Black Buttes conI 

group, Black Buttes, ',,"yo. 

Viburnum? heterodontum I(nowltoll, n. sp. 

Plate 55, Figure 2 

Leaf firm or coriaceous, approximately circulur, 

truncate 01' very slightly heart-shaped at the base but 

equally rounded in other portions; margin conspicuously but rather finely serrate, the teeth of two sizes, 

a series of slightly larger ones which are entered by 

the principal branches of the nervation and a smaller, 

more j l'l'egular series of intermediate teeth; nervation 

strong, palmately 5-l'ibbed from the base of the blade, 

the centml or midrib perfectly straight and bearing 

in the upper part one or two pairs of alternate, forked, 

craspedoc1rome secondaries; lowest pair of ribs at an 

angle of about 40°, passing with a slight upward 

curve to a marginal tooth, and bearing on the outside 

only six or seven strong secondary branches, which 

nro mostly simple and enter the marginal teeth; other 

pail' of ribs diyiding about equally the space between 

the midrib hnd lower pair of ribs, very strong, nluch 

curved upward, each with three or four very strong 

secondary branches on the outside, each of which forks 

and passes to the larger marginal teeth; nervilles 

I-I.umerous, strong, mainly percurrent and at right 

angles to the primary branches of the nervation, between which they pass. . 

The single example here figured is apparently all 

of the species thus far brought to light, and even this 

is not quite perfect, as it lacks the major portion of 

one side. This leaf appears to have been of very firm 

texture, nearly circular, with the margin bearing numerous rather small teeth of two distinct series-a 
130 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

southern Colorado; with which it appears to agree in 

'shape, but it has apparently a camptodrome instead of 

a somewhat craspedodrome secondary nervation. In 

general appearance and the finer nervation it is quite 

similar to the Raton species. 

Occurrence: Dawsoll arkose, lower part (in beds 

equivalent to Arapahoe formation), 3,000 feet east 0 f 

the' Douglas coal mine, Sedalia, Colo., collected by 

Arthur Lakes. 

Phyllites dcnverensis Knowlton, n. sp. 

Plate 59, :E'igure 5 

The material preserved on a red baked shale from 

the vicinity of Ramah, Colo., contains a number of 

fragments of a large palmately ribbed leaf that 'can 

not be made out with distinctness. The best preserved 

of these specimens is shown in the figure. As 

nearly as can be discerned it was an ovate leaf with a 

broad, shallow heart-shaped base and sharply serrate 

margin. The most conspicuous feature is the extremely 

large petiole, which is about 8 centimeters long and 

nearly 2 millimeters thiclc The petiole splits at the 

lower border of the blade into three strong branches, 

of which the midrib is the strongest. The midrib has 

a single pail' of alternate secondaries on the preserved 

portion. The lateral ribs arise at an angle of about 

45° and have several strong, somewhat irregular secondary branches on the outside, the lower pair with 

tertiary branches on the lower side. The' finer nervation consists of numerous strong. irregular nervilles. 

The width of this leaf was at least 10 centimeters. It 

has about 5 centimeters of the length preserved, but . 

there is no means of knowing its total length. 

So little of this leaf is present that its probable 

affinity is diffi_ult to determine. In some respects' it 

suggests the genus Tilia, but it could hardly be referred to this genus, and it must await adequate material for its allocation. The obvious size, long, thick 

petiole, three ribs, and serrate margin will probably 

permit its recognition if it should be 'subsequently 

found. 

Occurrence: Denver formation, red, baked shale 

near Ramah oil well, in the SE. 1;4 sec. 33, T. 9 S., R. 

61 'V., Colo., collected by C. W. Cooke for G. B. 

Richardson, 1910. 

Phyllites sp. 

Plate 56, ,Figure 3 

The little leaf here figured is the only one of its 

kind noted in the collections. It" is so fragmentary 

that ordinarily a desq'iptipn :would hardly be worth 

while, but the portion preserved is so well marked 

that it may be included. It was apparently ovate, 

probably 'about 3 centimeters long and 2 .centimeters 

wide, slightly heart-shaped at the base, and with entire. margins. The petiole is short-about 7 lnillimeters-and comparatively very stout. The midrib 

is extremely thick, and the lowest pair of secondaries 

are of the nature of ribs. They arise at the very top 

of the petiole and have three or. four camptodrome 

tertiary branches on the outside and an equal number 

on the upper side, ,yhich join the next secondary above. 

The number of secondaries is hot known, though there 

were at least three pairs, all opposite and at an angle 

of 45° or 50°. The finel: nervation is not retained. 

Occurrence: Dawson arkose, duml)' of Mosby coal 

mine, Mosby, Colo., collected by F. H. Knowlton and 

G. B. Richardson, 1910. 

Phyllites calhanensis Knowlton, n. sp. 

Plate 56, Figure 1 

The collection obtained near Calhan, Colo., includes 

cmong lImny other leaves a specilnen with its counterpart that is here figured. It is evidently a leaflet, 

. being unequal-sided and slightly arcuate. It· is 

Janceolate, about equally narrowed to both base and 

apex, and entire margined. The length is 4 centimeters and the width 1.75 centimeters. The petiolule 

is hardly more than 3 millinleters long. The nervation, with the exception of the midrib, is very obscure. 

The midrib is strong, and there appear to be several 

pairs of light, much curved secondaries. The seconda:ries are obscured by the thick substance of the leaflet. 

In the absence of adequate information concerning 

the nervation it is ilnpossible to place the leaflet satisfactorilv. It is not unlike some species of Sapind'l.ts, 

such as~ .8apind'tls affinis Newberry,S9 from the Fort 

Union formation, but it is too obscure for certainty. 

It also resembles certain leguminous leaves and leaflets, 

bnt again th~ obscurity of the nervatioH precludes positive identification. Under the circumstances it seems 

best placed under Phyllites. 

Occurrence: Dawson arkose, Rice's clay bank, 1 mile 

southwest of Calhan, Colo., collected by F. H. I{nowlton, July, 1910. 

Phyllites pellucidus Knowlton, n. sp. 

Plate 39, Figure 2; Plate 56, Figure 4 

In the material collected near Calhan, Colo., there 

are a number of very fragmentary leaves that seem to 

merit description. Two of the largest and best preserved ·of the fragments have been figured. These 

leaves were thin in texture. The outline is difficult 

to make o.ut beyond the fact that the apex was prolonged into. a rather narrow acuminate point. The 

margin is toothed and the teeth are.low, sharp-pointed, 

and separated by very shallow sinuses. The width 

au Newberry, J. S.; The Intel' extinct floras of North. America: U. $. 

Geol: Survey Mon. 35, p. 116, pI. 30, fig. 1; pI. 40, fig. 2, 1898. 
SYSTEMATIC PALEOBOTANY 131 

was at least 9 centimeters and the length could hardly 

have beon less than 12 or 15 centimeters. The nervation conRists of an apparently straight midrib and numerom; pairs of opposite or subopposite secondaries at 

an angle of abotlt 45°; the secondaries are craspedodrome. The nervilles are relatively strong and are' 

both percnrl.'ent and broken. • 

] t is diilicu Lt with fragments of the size available to 

suggest affinities for these leaves. 

Occnrrence : Dawson arkose, H.ice's clay bank, 1 mile 

southwest of Calhan, Colo., collected by F. H. I(nowlton, 1910. 

Carpites oviformis Lesquereux 

OU,fl)lics ovifonni,s Lesqucl'cUX, The TeL"tinl'Y florn, 1). 302, pI.' 

30, fig. Oa, 1878. 

The type specimen of this species is in the United 

States National J\1useum (No. 266) and is preserved 

on the same piece of matrix with a small leaf of Ficus 

aU'J"ic'l.IJlata. It has suffered to a considerable extent 

in the, yen.rs that hay;e intervened since it was described) and at present is so worn that little can be 

made out concerning it. It was described by Lesquereux as follows: 

)j~ruit exactly ovoid, 10 millimeters long, 6 millimeters broad 

in the middle. This ~ll1all nut is apparently a hard drupe, 

a8 it is not flattened by compl'Cssoll. Its surface is neither 

striate 1101.' lincate but somewhat rough. 

I have not found this species in the large collections 

made by Lukes and others, but in the material acquired 

by the National b1useum with the collections donated 

by H.. D. Lacoe there is a single specimen (No. 51027) 

that was 'referred to 0 a17tJYites o1Jif01"11mS by Lesquereux, ' 

who identified and labeled these collections. It is a 

smooth, ovoid fruit about 12 millimeters long and 8 

millimeters broad and appears to be slightly narrowed 

at one end. Lesquerel.lx cOlllpared this fruit to that of 

P1~I(jj1fltS scottvi I-Ieel', from Greenland, but it is unwise 

t.o attmnpt a definite allocation or interpretation of 

possible affinity. , 

Occurrence: Denver formation, Golden, Colo., collected by Leo Lesquereux, about 1875. Additional 

specimen from Lacoe collection. 

Carpites quadrivalvis I{nowlton, n. sp. 

Plate 56, Figure 2 

Fruit small, narrowly conical, ligneous, apparently 

4-valved, the sutures deeply furrowed, the backs of the 

yalves rough, with strong transverse lines or ridges . 

. This little fruit appears to be a 4-va1ved capsule, as 

it shows one valve complete and portions of two 

, others, one on each side of the perfect one, which probably means that there is a similn.r one on the concealed 

side. It is narrowly conical in shape, about 11 millimeters long, obtuse or trunca,te at the base and acute 

at the al;e·x. The perfect valve is about 2 mil1im~ters 

wide in the middle. The back of this valve and the 

sides of the lateral valves are thick and raised above 

the suture and are crossed by deeply impressed fine 

lines which' are often broken. 

The affinity of this fruit has not been recognized. 

Occurrence: Dawson arkose, Palmer's road, Templeton Gap, 4 miles northeast of ,Colorado Springs, 

Colo., sandy bed 6 feet thick, about 90 01' 100 feet above 

base of section, collected by F. H. I(nowlton, July, 

1910. 

Carpites templetoni Knowlton, n. sp. 

Plate 58, Figure 6 

In one of the small collections from the Dawson 

arkose at, "rempleto~ Gap, above Colorado Springs. I 

find two small fruits that do not appear to be deseribed. The best-preserved specimen-the one here 

figured-is circular, about 6 millimeters in diameter, 

and compressed to a thickness of about 1 millimeter. 

The llnfigured specimen is only about 5 millimeters 

in cliameter but does not otherwise differ. The figured 

specimen has a number of rather strong ridges, but 

their direction and character are uncertain. The surface otherwise is apparently roughened 01' possibly 

striate, but this is also uncertain. The point of attachment is not shown, hence it is impossible to orient the 

specimen with any certainty of this being its correct 

position. 

These little fruits have somewhat the appearance of 

certain thin-fleshed "berries" of certain palms, but 

obviously this is hardly more than conjecture. 

Occurrence: Dawson arkose, Palmer's road, Templeton Gap; 4 miles northeast of Colorado Springs, Colo., 

sandy bed 6 feet thick, about 90 01' 100 feet above 

base of section, collected by F. H. I(nowlton, July, 

1910. 

Carpites rostellatus Lesquereux 

O(tl'1Jites 1'ostellahts Lesquel'eux, The TC1'tial'Y flora, p. 303, 

pI. 60, figs. 11, 12, 1878. 

The figured types ?f this species are preserved in 

the United States National,l\1useum (No. 503). They 

are recorded by Lesquereux as coming from Golden, 

Colo., a reference that is sustained by the J\{useum 

record as well as by the matrix in which they ~'ire 

preserve<:l. In the same box and under the same nUlllbel' is another un figured 'specimen, preserved in a very 

hard, nearly black matrix unlike anything known 

from Golden, It is close to but possibly not identical 

,yith the other specimens of OaYrpites 1'o'8tellaJtus. It 

is probably from older rocks, possibly from the Laramie of Coal Creek, Boulder County, Colo., but this is 

not known positively. 

oAt first it was thought that O(JJ)'pites 1'ostellalms 

should be referred to Ginkgo? tnm,cata,"and it is still 
132 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

possible that this should be done, but they are much 

more distinctly and promInently ribbed than any" so 

referred. Nothing exactly like this species has been 

noted in the recent collections. 

Occurrence: Denver formation, Golden, Colo., original collections probably made by Leo Lesquereux 

~bout 1875. 

Carpite's costatus Lesquereux 

Oarpites CQstatus Lesquereux, The Tertiary fiora, p. 303, pI. 

60, fig. 5, 1878; U. S. GeoI. and Geog. Survey Terr. Ann. 

Rept. for 1876, p. 520, 1878. 

The type of this species is No. 497 of the United 

States National Museum collection and appears to be 

the only example obtained. It represents the base of a 

fruit with the scar of attachment. Its affinity has not 

been made out. 

Occurrence: Denver formation, South Table Moun- . 

tain, Golden, Colo. 

Carpites coffeaeformis Lesquereux 

OarpUes coffeaetormis Lesquereux, The Tertiary fiora, p. 303, 

pI. 60, figs. 6, 7; U. S. GeoI. and Geog. Survey Terr. 

Ann. Rept. for 1876, p .. 520, 1878. 

The types of this species are Nos. 495 and 502 of 

the United States National Museum collection. They 

represent the only examples thus far discovered and 

have been as well described and figured by Lesquereux 

as the specimens warrant. Their affinity has not been 

made out. 

Occurrence: Denver formation, Golden, Colo. 

Carpites minutulus Lesquereux 

Oarpites minutulus Lesquereux, The Tertiary flora, p. 305, 

pI. 60, fig. 25, 1878; U. S. Geol. and Geog. Survey Terr. 

Ann. Rept. for 1876, p. 520, 1878. 

This specimen, which represents, so far as known, 

the only one ever obtained, is not to be found in the 

collection of the United States National Museum, 

where it should be. There is no record of its having 

been sent here, and it is undoubtedly lost. It was 

described by Lesquereux as follows: 

Seeds very small, 3 millimeters long, only half as broad, inflated and rounded at one end, graduall:r narrowed to a short 

acumen, smooth. * * * It is mixed with fragments of 

stems and branches which appear to belong to some. conifer. 

Occurrence: Denver formation, South Table Mountain, Golden, Colo. 

Carpites myricarum Lesquereux 

Oarpites myrica1''ltm Lesquereux, The ·Tertiary flora, p. 303, 

pI. 60, figs. 8-1l! 1878. 

'rhis species was recorded by Lesquereux as having 

come from Black Buttes, Wyo., but at least one of the 

type specimeIls (the original of Lesquereux's pI. 60, 

fig. 9, equivalent to No. 499, U. S. Nat. Mus.) certainly 

came from the andesitic beds at Golden, Colo. Only 

one additional type specimen (the original of pI. 60, 

fig. 10) has ever been the property of the United 

States National Museum, so far as the records show. 

Thjs probably came from Black Buttes, as recorded 

by Lesquereux. 

o arpites rmyricarwm is obviously close to 0 arpites 

laurineU8 Lesquereux, which is now referred to 

Ginkgo? truncata. Lesquereux interpreted Oarpites 

mtyrica:rwm· as being seeds, which he says resemble 

large seeds of M y1'ica, but 0 arpites Zau1'ineus was 

thought to be a berry. Be this as it may, it is thought 

best to keep O. 1nyricaru1Z~ as a distinct species, at least 

until additional examples can be procured. 

Occurrence: Denver formation, Golden, Colo., Black 

Buttes coal group, Black Buttes, Wyo. 

Carpites coryloides Knowlton, n. sp. 

Plate 58, Figures 3-5 

One of the collections from the Dawson arkose on 

Jimmy Camp Creek, east of Colorado Springs, contains several specimens of a fruit that seems worthy of 

description. It was evidently a hard nut inclosed in 

a thick fibrous covering or "husk," somewhat after 

the manner of the common hazelnut and its inclosing 

covering. The nut itself is ovoid, about 22 millimeters long and 12 or 14 lnillimeters broad. It is 

somewhat angled: though doubtless it has been more 

or less distorted and compressed by fossilization. The 

outer covering was about 30 millimeters long. 

The most instructive specimen with its counterpart 

is that shown in Figures 3 and 4. In Figure 3 the 

outer covering is shown~ It is emarginate at the base, 

as if it fitted around an axis of some kind. The depression in which the nut rested is well shown. The 

counterpart shown in Figure 4 shows the nut lying in 

the opposite half of the outer covering. The nut is 

considerably flattened. In Figure 5 the nut has not 

been so much disto~ted, though the outer covering is 

not well preserved. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek opposite Richfield Springs ranch house, 9 

miles east of Colorado Springs, Colo~, collected by 

A. C. Peale, September 18, 1908. 

Eriocaulon? porosum Lesquereux 

Erioc(l.ulon'f poros1.tm Lesquereux, U. S. GeoI. and Geog. Survey 

'l'err. Ann. Rept. for 1873, p. 396, 1874; The Tertiary 

flora, p. 106, pI. 16, figs. 2, 2a, 1878. . 

Lesquereux's original description reads as follows: 

Lea "es basilar, rosulate, spreading, entire, lillear-Ianceolate, 

broader at the middle, gradually tapering upward t') a slightly 

obtuse point and downward to a very short petiole; media n 
SYSTEMATIC PALEOBOTANY 133 

nerve broad, concave; lateral veins two, nearly parallel, With 

apl)arent rumifications toward the borders, forming round 

polygonal slllall areolae. 

The leaves are thick, of a spongious texture apparently; the 

meshes along the borders are not distinct and may be formed 

by contraction of the epidermis. I do not find any species to 

which this form may be compared, except the leaves of some 

large rosulate llJrio(J(('1tlon·. The specimen is cut through by 

rootlets as thick as the leaves are broad. o 

. Thus fnr the type specimen, which is before me 

as I write (D. S. Nat. Mus: No. 137), is the only one 

that has been found. The type specimen is preservedo 

on a piece of soft, fine-grained clay and has been fairly 

well described and figured by Lesquereux. It is preserved on and partly vHapped around the end of the 

piece of matrix, and the" leaves," therefore, do not 

all lie in the same plane. Apparently, however, as 

Lesquereux supposed, they were disposed in a circle, 

possibly around a stem. The nervation in these 

" leaves " is uncertain. There is clearly a strong central ncrve. or midrib and apparently about two thinner 

ribs on each side, but the remainder is obscure and 

uncertain. The" leaf" was evidently a very thick 

und leathery one, and the surface has been more 01' 

less wrinkled, in addition to which the;e are numerous 

obvious joint checks, so that any actual nervation, if 

present, is difficult to make out. It seems probable~ 

however, that some of the meshing or areolation must 

belong to the nervation. 

'In subsequent years a number of o:rganisms ,were 

fonnd, mainly in older rocks, that were at first thought 

to be at least congeneric with the E'Jioc(JfUlon.f! PO'J'08U'ln 

of Lesquereux. They were mostly isolated" leaves" 

or segments, though occasionally several were found 

that showed these segments pretty closely associated. 

It was not, however, until some very perfect examples 

were found in the Vermejo formation in the Canon 

. City field of Colorado that the distinctness from Lesquereux's species became clear. These were described 

under the name Pala730aster inquirenda I(nowlton,9o 

with the frilnk admission that their affinity was unrecognized. They ~onsist of 8 to 12, usually about 

{) narrow, erect "leaves" or members 3.5 to about 

4.5 centimeters long and 6 to 10 millimeters wide in 

the nliddle. They are slightly narrowed to the sessile 

base, where they are in contact, though evidently perfectly free from each other. Above they are narrowed 

to a very slender acuminate point, usually somewhat 

incurved. The segments are thick and leathery, if 

not indeed woody, and are traversed dorsally by a 

deep median furrow. These specimens, preserved' 

in rather coarse sandstone, show very little trace of 

~tructure other than the median rib. 

00 Knowlton, F. H., U. S. Geol. Survey Prof. Paper 101, p. 278, pI. 

69, figs, 5, 6, 1917 [1918]. 

It seems probable that these organisms were terminal for there is some evidence of the presence of a , . dcar or point of attachment at the base, but no aXIS 

lin which they might have stood has ever been noted. 

They are certainly not leaves whorled around a stem, 

for had thev been some trace of the stem should , '" , . have been detected in some of the numerous speCImens. It appears much more likely that they were 

L!apsular in nature, for, if the now spreading segments were brought together, they would apparently 

make a tightly closed" capsule." The incurved tips 

of: the segments lend support to this view, though no 

evidence of seeds or other interior structure has been 

observed. 

. Among the several lots of specimens from the Raton 

Mesa reo-ion there is one found near Walsenburg that b , 

c:onsists of a number of detached leaves or segments. 

Only three or four leaves are preserved on any piece 

of matrix, but these seem to arise from a central point, 

as in the' other examples referred to Palaeoaster inqui'Joenda and as in the type of Eriocaulon? poro8urn. 

The leaves in these Walsenburg specimens are rather 

longer and narrower than common, and they appear, 

if anything, to have been thicker, as shown by the 

layer of carbonaceous matter remaining in favored 

places. The midrib is very deeply impressed--channeled, in fact--and at right angles to it there are 

minute parallel lines connecting with the margin. 

These specimens approach most closely Eriocaulon? 

}){)7'08Um, and it is possible that they should be united. 

However, the specimens involved are so few in numLeI' and their nature C so obscure that it seems best to 

hold them apart until more conclusive data are forthcoming. It is, of course, practically certain that Lesquereux's species has no connection whatever with 

the genus Eriocaulon, but, on the other hand, it is not 

at all clear that there is any closer affinity. 

The specimen from Black Buttes, Wyo., that was 

referred by Lesquereux 91 to Eucal/yptu8 haeringiana? 

Ettingshausen is probably also congeneric with Eriocaulon.f! pOT08um and possibly also with the leaves 

above mentioned from Walsenburg. In any event it 

'is fairly clear that this specimen has nothing to do 

with Eucalyptus. 

Occurrence: Denver formation, Sand Creek about 

12 miles east of Denver, Colo., supposed to have been 

collected by W. H. Holmes. 

Genus BERRYA Knowlton, n. gen; 

Generic characters the same as the specific char8.cters. 

91 Lesquereux, Leo, The Tertiary flora, P. 296, pI. 59, fig. 10, 1878. 
134 FLORA OF THE DENVER AND ASSOCIATED FORMATIONS OF COLORADO 

Berrya racemosa (Knowlton) Knowlton, n. comb. 

Plate 41, Figures 4, 5 

Nyssa? racemosa Kn:owlton, U. S. Geol. Survey Bull. 152, p. 153, 

1898. 

Salbal~te8 fruotif(}1' Lesquereux [part], The Tertiary flora, 

p. 114, pI. 11, figs. 3 (fruits at base of figure), 3a (fruit· 

enlarged), 1878. 

Fruits in a compound raceme with a very thick, 

strong rachis; individual fruits 10 to 16 millimeters 

h,ng, 4 to 6 millimeters in diameter, erect, shortpediceled, oblong, acute at both ends or sometimes 

slightly obtuse at apex, provided with probably about 

12 to 15 faint longitudinal striae or ribs and very 

numerous, delicate transverse striae. 

The type locality for this striking fossil is in the 

Denver beds at Golden, Colo. It is represented by· 

two large, nearly perfect compound racemes, by a 

number of parts of racemes, and by a very consicler-

" able number of detached fruits. The larger and more 

perfect specimen was at least. 8 centimeters long, with 

the rachis fully 2 millimeters-thick. It is branched 

near the base into two nearly equal portions, along 

which the fruits are dispersed alternately, each branch 

bearing six or seven fruits. 

The other of the more perfect specimens found had 

at least three branches-a lateral o!1e, the base with 

five fruits; one on the opposite side and a little higher 

up with three fruits; and the basal portion of another 

branch still higher. Each fruit has a short, thic.k 

pedicel, from which it appears- to have been detached 

at maturity by an abscission at the base of the fruit. 

Each fruit shows a number of longitudinal lines or 

ribs, the exact number of which can not be positively 

determined, though there appeal' to be about 12 or H). 

The transverse striae are very fine and delicate and 

may possibly have been produced by the wrinkling of 

a fleshy exoeatp .. These fruits appear to have been 

drupes with probably a thin, fleshy exocarp ,vhich in 

drying may have caused some of the corrugations observed, though it seems unlikely that this could have 

produced all of the very regular fine lines. 

These fruits were first detected by Lesquereux. who 

found them in close association with a small pinnate 

palm leaf on a small piece of matrix. On account of 

this association Lesquereux concluded that they represented the fruit of this palm. The original specimen 

upon which this determination rests is preserved in 

the United States National Museum under the name 

S(J)b(J)Zites frwoti/m'. It needs but a glance at this specimen, or indeed at the well-executed figure of it, to 

show that there is no organic connection between them, 

nor is such union implied by Lesquereux. On this 

point he said: "The fruits are probably referable to 

the same species, as they are not only seen at the base 

of the leaf but more numerous still embedded in the 

stone." The specimen consists of the upper part of a 

raceme with three fruits. 

It was with milch hesitation that these fruits were 

at one time referred to the genus Nyssa. It had not 

then been found possible to locate a living genus to 

which they could be referred with reasonable certainty, and they ,yere placed tentatively in NY'8sa 

mainly on account of slight resemblance to certain 

fossil fruits that have been described under this 

cgeneric name. 

The fossil species to which the fruits fr0111 Golden 

show the most resemblance is perhaps NyssaJ arctica 

IIeer,!l2 from the Eocene of Cape Lyle, Spitzbergen. 

It differs in being much larger (17 to 21 millimeters 

long and 8 to 10 millimeters wide) and in having more 

numerous and more prominent longitudinal ribs. The 

transverse striae are of about the same character in 

both. There is no evidence that Nyssa; aJrcticaJ had the 

fruits arranged in a raceme, although several are figured as having two fruits sessile on the top of a short 

peduncle, as in the modern Nyssa. 

Some of the nlore elongated fruits of N yssidiu1n 

el~l]utni Heer,D3 .from Cape Staratschin, Spitzbergen, 

are also similar to the Golden fruits, but differ mainly 

in being smaller and apparently without transverse 

striae. 

A number of other fruits from Cape Staratschin are 

described by Heel' under the name Nyssidiw'/1v, and although they all resemble in a general way the indiyidual fruits of Be·rrya, their manner of growth is not 

a t all simila r. 

Since this species was first made known it has been 

found at a number of localities in beds of approximately the same age. Thus it occurs in the Dawson 

arkose near Ramah, Colo., in the Raton formation 

Ileal' Raton, N. ~iex., in the Anilllas. formation on the 

northern rim of the San Juan Basin, Colo., and in the 

Black Buttes coal group at Black Buttes, 'Vyo. 

I ta ke pleasure in dedicating this genus to Prof. 

Edward 'Vilber Berry, professor of geology and paleontology in the Johns Hopkins University, Baltimore, 

Md. 

Occurrence: Denver. formation (type), Golden, 

Colo., probably collected by Lesquereux, about 1872; 

additional material, including the two figured specimens, collected by Arthur Lakes on the south face of 

South Table MOllntain, 100 feet below the lava cap. 

pawson arkose, Coal Gulch, about 2 miles northwest 

of Ramah, Colo., collected by VV. T. Lee, 1914. Black 

Buttes coal group, Black Buttes, Wyo., collected by 

F. H. Knowlton ~nd A. C. Peale in 1909. Raton f.o~~-

02 Heer, Oswald, Flora fossilis arctiea, vol. 4, p. 80, pI. 19, figs. 

1-10, 1877. 

03 Idem, vol. 2, p. 62, p1. 15, figs. 1-7, 1870. 
SYSTEMATIC PALEOBOTANY 135 

mation, Yankee mine, near Raton, N. ~1ex., collected 

by 'V. T. Lee. Animas formation, Ignacio quadrangle, 

Colo., pebbles lIear top of beds, collected by J. H. 

Gardner, 1909. 

Decorticated roots? 

Plate 58, Figures 1, 2 

The collections from the Dawson. arkose along 

.Ji III Illy Camp Creek eontain a large number of fragmentary specimens that appeaJ.· to be decorticated 

roots or rootstocks, but their exact affinity has not been 

recognized. The largest portion of the main axis that 

is preserved is about 1 centimeter in diameter. It is 

sligHtly and irregularly ridged and also is finely 

striate. It gives off the" roots " ~t alternate intervals, 

these emerging at an angle of about 45° and of unknown length but at least 10 centimeters. The" roots" 

or rootstocks are more strongly ribbed or striate than 

the main axis and show a regular series of scars whence 

apparently finer roots have emerged. These scars, if 

this is what they are, are strong, fairly regularly disposed, and probably though not certainly spiral. 

I have at present no knowledge as to the probable 

affinity of this plant. 

Occurrence: Dawson arkose, east bank of Jimmy 

Camp Creek 0.6 mile above Richfield Springs ranch 

house, 9 miles east of Colorado Springs, Colo., collected by F. H. I(nowlton and ""V. T. Lee, July, 1910. 

PLATES 

SG4!)!;!-i:lO--lO 137 
NOTE.-The museum numbers are given in parentbeses-U. S. N. M., UniteclStates National Museum; M. C. Z., Harvard 

College Museum of Comparative Zoology, Cambridge, Mass.; Pr., Princeton University Museum, Princeton, N. J 

138 
U. S. GEOLOGICAL SVRVEY PROFESSIONAL PAPER 155 PLATE 1 

1 

') JI # / 

~rj 

~ l.r-, 2 

./ 

3 

7 

DENVER FLORA 

I, 2. Dryopteris takesii (Lesquereux) Knowlion, n. comb. (2, U.S.N .M . 37750), Den ver formation . 3. "'1archantites' coloradensis Knowlton, 11. sp. (U.S.N .M . 37791), Dawson arkose. 4. IIymenophyltum con/mum Lesquereux (U .S.N. 1. 37782), Denver formation. 5. Dryopteris integra Knowlion, n. sp., Middle Park formation. 

G. Sclerotites1 hesperiu8 Knowlion, n. sp., Dawson arkose. 

7. Seterolites' cypericola Knowlion, n. sp. (U.S.N.M. 37834), Dawson arkose. 
U. S. GEOLOGICAL SURVEY 

1 

2 

7 8 

PROFESSION AL PAPER 155 PLA TE 2 

3 4 

5 

9 6 

I , '.1 , II 

.. ( )u 

,,,", 

, I 

I 

, 

I 

I 

I 

I,.W. 

, I 

'_ ...... I 

I 

I 

I 

, I I, 

~!.-l , I 

\ : 

'" 

17 

J.; I 

/ 

DENVER FLORA 

t,2. Woodwardia iatiloba serrata Knowlton, n. var. (1, U.S.N.M. 37846), l\'1iddle 

Park formation. (1 X 2; 2 X 4.) 

3-5. Dryopteris richardsollialla Knowlton, n. sp. (U.S.N.M. 37751), Dawson 

arkose. . 

6. Dipiazium crossii (Knowlton) Knowlton, X 2 (U.S.N.M. 37752), Denver 

formation. 

7-9. Dennstaedtia cTossiana Knowlton, n. sp., Dawson arkose, lower part (in beds=Arapahoe formation). (8, single pinnule X 2.) . 
U. S. GEOLOG ICAL SURVEY PROFESSIONAL PAPER 155 PLATE 3 

1 

4 

2 

5 

3 

6 7 8 

DENVER FLORA 

1,2. Sacc%ma sp. Knowlton (U.S.N.l\J. 37831), Dawson arkose. 3-8. Saccoloma gardner; (Lesquereux) Knowlton (3,5, 8, U.S.N.~1. 37832; 4, M.C.Z. 853), Dawson arkose. (7, fruiting specimen.) 
U. S. GEOLOGICAL SURVEY 

·1 

2 

DENVER FIJORA 

Allalltodiopsis erosa (Lesqllerellx) Knowlton ancI l\'faxon (U.S.N.M. 37712-37714). (I, fruiting specimen. All X 2.) 

PROFESSION AL PAPER 155 PLATE 4 

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..... ~----------------....... ."" / \ "~ ~ ':;:?" 

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3 
U. S. GEOLOGICAL SURVEY 

Allantodiopsis erosa (Lesquereux . ) Knowlton 

PROFESSION AL I 'APER 155 

O

DENVE RA (1 and 5 after R FL . S 1\.M. 37715). 

and Maxon (1, U . . S N.M. 3618; 5, U .. Lesquereux.) 

PLATE 5 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 6 

DENVEH FLOHA 

Geonomites tenutrachis LesQuereux (u.S.N.M. 3nn), Dawson arkose. 
U . S. G EOLOGICAL 

2 

SURVEY PROFESSION AL 

3 

Salpichlaena DENVER FLORA anceps (Lesquereux) K . nowlton (1, M .C.Z. 852 ), Denver format' IOn. 

PAPER 155 PLATE 7 

' ) 

i , , 

1 

) , -,-.. , ( 

, ' .. .1 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 8 

3 

9 10 4 

DENVER }'LORA 

1-6. Equisetum colora dense Knowlton, ll. sp. (U .S.N.M. 37753). 1,2,4 X 2, Middlo 

Park formation; 3. 5. 6 natural size, Dawson arkose. i. Anemia sp. Knowlton, Middle Park formation. 

8. Lygodium coloradense Knowlton, n. sp., Dawson arkose. 

9. Anemia mosbyensis Knowlton, n. sp. (U.S.N.M. 37719), Dawson arkose. 10. Anemia lanceolata Knowlton, ll. sp., Middle Park formation. 
U. S. GEOLOGICAL SURVEY PROFESSION AL PAPER 155 PLATE 9 

3 

2 

1 

4 

6 

DENVEH l?LORA 

1,2, G. Geonomites tenuirachis Lesquereux (1, U.S K.~I. 37ii8; 2, U.S.N.l\I. 3ii77), Dawson arkGse . 3. Ginkgo? truncata (Lesquereu x) Knowlton, n. comb., X 2 (U.S.N.M. 3i(80) , Dellyer formation. 

4. Equisetum sp. Knowlton (U.S.N.~I. 37i54), DeD\'er formation. 5. Sabatites grayanus (Lesqucreux) Lesquereux (U.S.N.M. 37830), Den· ver formatioll. 
U. S. GEOLOGICAL SURVEY 

PROFESSION AL PAPER 155 PLATE 10 

2. I. mation. Geonomites a,,,,,.mu,,, goldianus spc Knowlton (Lesquereux) (U.S.N.M. Lesquereux 37779), Denv" (U.S.N.M. '''mation. 2059), Denver for- •• formation. Ch.m",d."., ''''''ad .. ,;, Knowlton, n. sp. (U.S.N.M. 37727), MMd'. P"", 

DENVER FLORA 
U. S. GEOLOGICAL SURVEY 

1 

6 

5 

DENVER FLORA 

~. Paloreodoxites plicatus (Lesquerem) Knowlton, n. comb., Denver formation. S. PalmocarponY lineatum Lesquereux (M.C.Z. 1616), Denver formation . 

PROFESSIONAL PAPER 155 PLATE 11 

7 

6. Piper heeri Lesquereux (M.C.7-. 1037), Denver formation. 7. Myrica sp. Knowlton (U.S.N.M. 37792), Dawson arkose. 
U. S. GEOLOGICAL SURVEY PHOFESSION AL P APEH 155 PLATE 12 

I 

, 

DE VER FLORA 

Juglan. denveriana Knowlton, n. sp. (U.S.N.i\I. 2062, 2063), Dem-er formation. 
U.· S. GEOLOGICAL SURVEY PROFESSIONAL P APER 155 PLATE 13 

1 

4 

DE VER FLORA 

t. Comus denverensis Knowlton, n. sp., X 2 (Pr . 2339), Denver formation. 

2-4. Juglan" denl>eriana Knowlton. n. sp. (U.S.N.M. 2061 ... 37785, 3626), Denver formation. 

D. Juglam thermalis Lesquereux (U.S.N.M. 37787), Middle .t'ark formatiou. 

5 
ru. S . GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 14 

DENVER FLORA 

I. Jugian8 rugosa Lesquereux (U.S.N.M. 37786), Dem'er formation. 

2-4. Pterocarya? retusa Lesquereux (M.C.Z. 1558, 1559, 1561), Denver formation. 5. Jugians sp. Knowlton (U.S.N.M. 37784), Dawson arkose. 

3 

4 
U. S. GEOLOGTCAL SUHVEY PHOFESSIONAL PAPEH 155 PLATE 15 

3 

7 

5 

8 9 10 

DENVER FLOHA 

1. Juglans sp. Knowlton (U.S.N.I\L 37784), Dawson arkose. (Photograph 

made before specimen was broken as shown on pI. 14, fig. 5.) 2. Alnus carpiniJolia Lesquereux (l\LC.Z. 1056), Denver formation. 

3,4. Betula schimper; Lesquereux CU .C.Z. 1035, 1036), Denver formation, 

85,j!)2-30--11 

5. Pterocarya? retusa Lesquereux (lVI.C.Z. 914), Denver formation. 

S. A lnus auraria Knowlton and Cockerell (lVI .C.Z. 1057), Denver formation . 7- 10_ Bet'ula Jallax Lesquereux (lVI.C.Z. 1034, 1045, 1058, 1620), Denver formation. 
U . 8. GEOLOGICAL SURVEY PROFESSIO)lAL PAPER l.?5 PLATE 16 

3 

2 

1 

4 5 

6 

8 

DENVER FLORA 

1,2. Betula Jaltax Lesquereux (M.C.Z. 1043, 1044), Denver formation. 

3- S. Quercus viburniJolia Lesql1ereux (3, 4, 6-S, M.C.Z. 1572, 157S, 1585, 1631; 5, U.S. ".M. 37825), Dem"er formation. 
U . S. GEOLOGICAL SURVEY 

, 

I 

I 

I 

I 

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,....... ) \, ... , - ' 

\ 

1 

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_.) 

--

2 

'-. ~; 

, 

, 

, 

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, 

, 

I 

I 

, 

, 

, 

, 

I 

I 

, 

PROFESSIONAL PAPER 155 I' LATE 17 

3 

~- ;( /"., / '-., .... ,--._,/ .'/ ":-~-"­ '--.., -,\ 

, . . ... .---' 

": 

4 

, ' 

... _.....," 

5 

DENVER FLORA 

I. Quercu8 viburnijolia Lesq uereux (U.S.N.M. 3i824), Dem'el' formation. 2. Quercus purdon ens is Knowlton, n. sp. (U.S.N.M. 37822), 1)a\\'50n arkose. 3. Q'uercu8 whitei Lesquereux (M.C.Z. 1056), Denver formation. 

4." Quercu8Y 8edalien8is Knowlton, n. sp., X 1~" (U .S.N.M. 37823), D awson ark ose, lower part (in beds=A rapahoe formation ). 5. Quercus w hilmani Knowlton, n. sp. (U.S.N.)I. 37826), Middle Pnrk formation. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 18 

4 

6 

'\ 

. 

' \ 

.,' .. ~-, . ~. 

. , 

, 

, 

, 

/- ~""i 

l -_i ",-,/--"- .----- -" 

8 

DENVER FLORA 

1. Quercus whitei Lesquereux (M.C.Z. 1073), Denver formation. 

2,3. Quercusteonis Knowlton, n. sp. (2, U.S.N .M. 37838; 3, M .C.Z. 1625), Denver formation , 4- 7. Quercus cetastriJolia Lesquereux (M.C.Z. 1071, 1075, 1076, 1626), Denver formation. 

8. Populus zeilleri (Lesquereux) Knowlton, n. comb. (M.C.Z. 1672), Denver formation. 

5 

7 
U. S. GEOLOGICAL SURVEY PROFESSION AL PAPER 155 PLATE 1U 

1 

8 

5 

6 

7 

DENVER FLORA 

1- 3. Populus zeilleri (Lesquereux) Knowlton, n. comb. (M.C.Z. 1671, 1674, 1675), Dem'er formation. ' -7. Populus nebrasemsis var. grandide71tata Lesquereux (4-6, M.C.Z. 1123, 1219, 1226; 7, U .S.N.M. 37817), Denver formation. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 20 

6 

7 8 

D E YER F LORA 

I, ~, 5, 8. Populus nebraseensis var. rotundata Lesq uereux (3-5, l\L C.Z. 1441,1498,1505), Denver formation. 2,6,1. Populus "ebraseemis var. grandidentata Lesquereux (2, 6, ~l.C.Z. 1499, 1683; 7, U.S. .M. 37817), Denver formation. 
U. S . GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 21 

7 

2 

1 

5 

9 

DE VER FLORA 

1, 2. Populus nebrascensis yar. acutidentata LesquereLLx (lILC.Z. 1202, 1210), Dem-er formation. 3- 5. Popl!lus nebrascensis var. longi!olia Lesquereux (M.C.Z. 1239, 1244, 1245), Dem'er formation. 6- 8. Pop/du .• jacksoni Knowlton, n. sp. (M.C.Z.I077, 1083, 1084), Dem'er formaLion. 9. Populus? sp. Knowlton (M.C.Z. 1286), Denver formation. 

3 

6 

8 
U . S . . GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 22 

5 

j 

) 

\ 

....... :;:::--7 ... 

7 9 

8 

DENVER FLORA 

1- 4.6. Populus lenuinervala Lesquereux (M.C.Z. 1252, 1256, 1266, 12i4, 1513), Den· ver formation . 5. Populus lacoeana Knowlton, n. sp. (U.S.N.M. 37816), Denver formation. 7. Populus jacksoni Knowlton, n. sp. (M.C.Z. 1085), Denver formation. 8. Populus zaddachi lIeer? (U.S.N.M. 37820), Middle Park formation. 9. Populus subrolunda Lesquereux (U.S.N.M. 37819), Denver formation. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 23 

3 

1 2 

5 

7 

6 

9 

8 10 

DENVER FLORA 

1-4. Populus knowltoni Berry, n. sp. (U.S.N.M. 37815), Dawson arkose. 5. Populus den"erens;s Knowlton, n. sp. (U.S.N.M. 37814), Denver formation. 6,8. Populus richardsoni Heer? (U.S.N.M. 37818), Middle Park formation. 7. Ulmus antecedens Lesquereux (M.C,Z. 1692), Denver formation . 

9. Ficus? alala Knowlton, n. sp. (U.S.N.M. 37760), Dawson arkose, lower part (in beds=Arapaboe formation). 10. Ulmus? querci{olia Unger? (M.C.Z. 1628), Denver formation. 
U. S." GEOLOGICAL SURVEY 

4 

PROFESSIOK AL PAPER 155 PLATE 24 

i'·~/ .'. --- ~ 

3 

DENVER FLORA 

1- 3. Ficus denveriana Cockerell (I, 2, U.S.N.l\T. 37762; 3, U.S.N.lVI. 3621) Dem"er formation. 4. Ficus aguilar Knowlton (U.S.N.M. 37759), Dawson arkose. 
U. S . GEOLOGICAL SURYEY PROFESSIONAL PAPER 155 PLATE 25 

1 2 

6 

DENVER F LORA 

1, 2,6. Ficus berthoud; LesQuereux (M.C.Z. 1089a, b, 1440, 1442), Denver formation . 3- 5. Ficus pseudopopulus LesQuereux (4, U.S.N.M. 377i2; 5, U.S.N.M. 3n,3), Dawson arkose. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 26 

3 

1 

DENVER FLORA 

I. Ficus dawsonensis Knowlton, Il. sp. (U.S.N.M. 3ii61), Dawson arkose. 2-4. Ficus puudopopulus LesQuereux (2,3, U.S.N.M. 37770, 37771; 4, M.C.Z. 16ii), Dawson arkose. 5. Ficus occidentalis (Lesquereux) Lesquereux (U.S.N.l\L 37768), Denver formation. 

5 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 27 

DENVER FLOHA 

1-3. Ficus sp. Knowlton (U.S.N.M. 37758), Dawson arkose. 

4. Ficus occidentalis (Lesquereux) Lesquereux, Denver formation. 

5. Ficus ramahensis Knowlton, n. sp. (U.S.N.M. 37776), Dawson arkose. 
U. S . GEOLOGICAL SURVEY PROFESSIONAL PAPEU 155 PLATE 2S 

7 

3 

2 

5 

6 

4 

9 

8 10 

DENVER FLOllA 

1. Ficus? lakesii Knowlton, n. sp. (U.S.N.M. 37763). Denver form ation. 2. Ficus l)lanicostata var. problematica Knowlton, n. var. (U.i'.N.i\.J". 37767), De",'er form ation. ~,~, .6,7. Ficus neoplanicostata Knowlton (3, U.S.N.M. 37764; 4, U.S.N.M. 3633; 6, U .S.N.M 37765), Dem'er formation . • • FlCu~ l)lamcostata var. gotdlOna LesQuereux (U .S.N.M. 37769), Denver formation. 8-10. FIcus 7JraetnnerVls Knowlton, n. sp. (U.S.N.i\I. 37774), Dawson arkose. 
U. S. GEOLOGICAL SUHVEY 

2 

3 

DENVEll FLOl1A 

1,2. Ficus puryearensis Berry? (U.S.N.M. 3i775) , Dawson ark ose. 3, 4. Ficus ne01)/anicostata Knowlton (U.S.N.i\I. 37766), Den\'er formation . • . Ficus sp. Knowlton (U.S.K.i\I. 3ii56), Dawson arkose. 

pnOFEI-'SlONAL llAPEH 1.1 5 rLATE 29 

4 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 30 

1 

DENVER F LORA 

1. Ficus sp. Knowlton (U.S.N.M. 3i757), Dawson arkose. . Artocarpus pung'''8 (Lesquereux) Hollick. (After Lesqnereux.) 
u. S. GI~OLOGlCAf, SURVEY 

:::;;:;4!):.!-aO--12 

" \.. ,/ 

~~~" 

DENVEH FLOHA 

I 'j! 

,:"/ 

, 

PROFESSIONAL P·:\.PEH 155 PLATE 31 

Artocarpus Imngens (Lesquereux) Hollick (U.S.N.].-!:. :;:O(j9), Del1\"er formation. 
U. S. GEOLOGICAL SURVEY 

,. 

< 

\~-~V :..,,- ---~---

1 

I 

I 

--.-' 

I 

, 

I 

I 

I 

PROFESSIONAL PAPER 155 PLATE 32 

2 

4 

DENVER FLORA 

1. Artocarpus punge",s (Lesq L1 ereL1x) IToJlick, refigured type (Pr.). (After L es· 

qucreux.) 2. Artocarpus punge",s (Lesquereux) IToJlick (U.S.N.M. 206i), Dem'er formation. 

3. Platanus rhomboidea LesQuereux (U.S.N.M. 3i812), Dawson arkose. . Artocarpus similis Knowlton (U.S.N.Iv!. 3nn), Dawson arkose. 
U. S GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 33 

2 

1 

DENVEH FL OHA 

1. Platanus aceroides "al'. lati/olia Knowlton (U.S.N.M. 37805), Denver formation. 2. Platanus yuillelmae Goppert (U.S.N.M. 37807), Denver formation. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL 'PAPER 155 PLATE 34 

2 

1 

DENVEH FLORA 

1. Plalanu8 yuillelmae GOlJpert (U.S.N.:'L 3622), Denver [ormation. 2. Plalanu8 raynold8ii var. inlegri!olia LesQ uereux (U .S.N.:' L 37811) Dc",'er [ormation. 
l:. S. GEOLOGICAL SURVEY PHOFESSlONAL PAPEH 155 PLATE 35 

DENVER FLORA 

P latanus ravnotdsii l\ewberry (U.S.N .lI!. 3i8JO), l)em"er [ormation. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 36 

DENVEH FLORA 

1. Plalanus haydenii Newberry (1\[.('.Z. 1371), Denver formation. 

2,3. Platanus marginata (Lesquereux) lI eer (U.S.N.M. 37808), D awson arkose. 4. Cinnamomum? s p. Knowlton (U.S.N.M. 37732), Dawson ark ose. 5. Platanus platanoides (Lesquereux) Knowlton (V.S.N .M. 37809), Dawson arkose. 
U. fl. GEOLOGICAL SURVEY 

PROFESSIONAJ, PAPER 155 PLATE 37 

2 

1 

DENVER FLORA 

P/ata1lllS c%radensis Knowlton, n. sp. (U.S.N .M. 37806), Middle Park formation. 
u. S. GEOLOGlC ~L SURVEY' PROFESSIONAL PAPER 155 PLATE 38 

.J 

6 7 8 9 

DENVER FLORA 

1. Platartus c%radensis K nowlion, n. sp. (Drawing of sl:ecimen shown on pI. 37, fig. 2.) 

2. Platal1 us sp. ? Knowlion (U.S.N.M. 3781 3), Dawson arkose. 

3,5, fl. LaltTUs pTimigenia Unger (3, M .C.Z. H66; 5, 6, U.S.N.M. 37789), Dawson arkose. 

4. L(LUTUS socia lis Lesquerellx (U.S.N.M. 3i/90), Dawson arkose. 7, 8. Gil1l1am 07nUm selal1l1C11Be Waielei? (U.S.N.i\f. 37735), Dawson ark ose. 9. LauTus lal1ceolata KnowltOIl. Il. sp. (U.S.K. M. 37788), Dawsoll arkose. 
U . S. GEOLOGICAL SURVEY PIWFESSION AI. PAPER 1.,5 PLATE 3n 

1 

2 

3 DENVER FLORA 

1. Cinnamomum sp. Knowlton (U.S.N. Col. 37733), Dem'er formation. 

2. Phyllites peUucidus Kno\\'lton, n. sp. (U.S.K.lIL 37801), Dawson arkose. 3. Paleonelumbo macroloba Knowlton, n. sp. (U.S.K.lIl. 37796), Dawson arkose. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 40 

1 2 

3 

4 

6 

5 

DENVER :FLORA 

1. Celastrinites populiJotius Knowlton, n. sp. (M.C.Z. 1422), Denver formation. 

2,3. Chrysobalanus coloradensis Knowlton, n. sp. (U.S.N.M. 37729, 37730) . 4. Asimina eocenica Lesquereux (U.S.N.M. 37718), Denver formation. 

0. Ficus martini Knowlton, ll. sp., Denver formation. 6. Rhamnus cleburni Lesquereux (U.S.N.M. 37828), showing character oflarger leaves from tbe Denver formation formerly referred to Comus studeri neer. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 41 

4 

:2 

3 5 

DENVER FLORA 

1. Nelumba lakesiana Lesquereux. (After Lesquereux.) 2. Nelumba tenuiJalia (Lesquereux) Knowlton. (A fler Lesquereux.) 3. Netumba crassii Knowlton, n . sp. (U.S.N.M. 3i(93), Dawson arkose, lower part (m beds=Arapaboe formation). 

4,5. Berryu racemasa (Knowlton) Knowlton, n . comb. (5, U.S.N.M. 37720), Dawson arkose. 
U. S. GEOLOGICAL SURVEY PROFESSIOXAL PAPER 155 PLATE 42 

4 

DE VEH FLORA 

I. Castalia pulchella Knowlton, n. sp. (restoration), Dawson arkose. 2. Nelumbo takesiarUL Lesqllerellx. (After Lesquereux.) 

a, 4. Pa/eone/umbo macroloba Knowlton, n. sp. (U .. N.M. 3ii96), Dawson arkose. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPE R 155 PLATE 43 

4 

, - , 

1 

5 

7 8 9 

DENVER FLORA 

1, 2,4,6. A8imina eocenica Lesquereux (I, 2, U.S N.~\r. 37f04; 4, U.S.N.M. 37717; 6, PI'. 2207). 4, Midd le Park formation; 1, 2, 6, Dawson arkose; 6, X 2. 3, 5,7- 9. ChrY8obaian'U8 co[oraden8i8 Knowlton, n. sp. (3,5,8,9, 1'.S.N.l\I. 37728; 7, M .C.Z. 1502). 
U. S. GEOLOGICAL SURVEY 

8 

PROFESSIONAL PAPER 155 P L ATE 44 

5 

9 

10 

DENVER FLORA 

1, 2. Chrysobalanus coloradensis Knowlton, n. sp. (U.S.N.1\T. 37728). 

3,4, G, 11. Prunus denvere"sis Knowlton, n. sp. (U.S.N.M. 37821), Dawson arkose . 5. Chrysobalanus? lanceolatus Knowlton, n. sp. (U.S.N.1\ 1. 37731), Dawson arkose. 7- 10. Grataegus 71Iyricoides Lesquereux (8- 10, 1\1. C.Z. 1072, 1566, 1567), Den ver formation. 

3 

7 

11 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 45 

1 

2 4 

3 

7 

6 8 

5 

9 10 11 

DE VER FLORA 

1. Phaseolites coloraliensis KnowlLon, n. sp. (U.S.N.1\f. 3i798), Dawson arkose. 

2. Cassia vetusta Knowlton, n. sp. (U.S.N.~1. 37724), Dawson arkose. 

3,4,9. Celastrus gaudini Lesquereux (3, n, M.C.Z. 1528, 1512), Dem'er formation. 

5. Cercis coloradensis Knowlton, n. sp. (U.S.N.1\L 37726), Dawson arkose. 

6. Acer sp. Knowlton (U.S.N.1\l. 377J6), Dawson arkose, lower part (in 

beds=Arapahoe formation). 

7. Sophora richardsoni KnowlLon, n. sp. (U.S.N.1\I. 3i836), Dawson arkose. 

8. Sophora puryearensis Berry'! (U.S.N.1\I. 37835), Dawson arkose. 

10. Negundo decurrens Lesquereux (M.C.Z. 1523), Denver formation. 

11. Cetastrinites poputiJolius KnowlLoJ], 11. sp. (1\.I.C.Z. 15i6), Denver formation. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER Ij5 PLATE 4G 

4 

12 

DENVER FLORA 

1,2. Sapindus berryanus Knowlton, n. 51'. (U.S.N.M. 37833), Dawson arkose. 3. Sapindus' obtusifolius Lesquereux, Denver formation. 4. flex? ovata K nowlton, n. sp. (U.S.N .M. 37783), Denver formation . 5. Rhamnus cannoni Knowlton, n. sp. (U.S.N.M. 37827), Middle Park formation. ~, 9. Zizyphus fibrillosus (Lesq uereux) Lesquereux (U .S.N.M. 37849), Dem'er 

formation. 

7, 8. Rhamnus?praealaternoides Knowlton, n. sp. (U.S.N.M. 432), Denver formation. 

(After Lesquereux.) 

10,11. Rhamn'us cleburni Lesquereux (U.S.N.M. 37828), Dawson arkose. 12. Rhamnus salicifolius Lesquereux? (U.S.N.M. 37829), Dawson ar kose. 
• U. S. GEOLOGICAL SURVEY 

PROFESSI ONAL PAPER 155 PLATE 47 

3 

'. '--\ 

'\ '. 

1 

4 

5 6 

DENVER FLOHA 

1. Gissus hesperia Knowlton, n. sp. (U.S.K. M. 3i(40), Dawson arkose. 2. ZlZyphus dis/or/us LesQuereux (U S.N.M. 37848), Dawson arkose. 3. Zizyphus becku'Uhii LesQuereux, Denver formation. (After Lesquereux.) 4. Gissus coryli/olia LesQuereux (11.C.Z. 1501), Denver formation. 

5. Gissus? cannoni Knowlton, n . sp. (U.S.N.M . 37737), Middle Park formation. 6. Gissus coioradensis Knowlton and Cockerell, X 2 (U.S.N.M. 37736), Dawson arkose, lower part (in beds=Arapaboe formation). 

S549:!- :.!O--1;; 
U. S . GEOLOGICAL SURVEY 

3 

4 

PROFESSION AL PAPER 155 PLATE 48 

1 

2 

5 

DENVER FLORA 

1. Gissus corylifolia Lesquereux (M.C.Z. 1503), Denver formation. 2. Gissus grosse-dentata Knowlton (U.S.N.M. 37738), Dawson arkose. 

3, i. GiSBUS lesquereuxii Knowlton (3, M.C.Z. 1493; 4, U.S.N.M. 37741), Denver formation. 5,6. Zizyphus daphnogenoides Knowlton (U.S.N .M. 37847) , Denver formation. 

6 
U. S. GEOLOGICAL SURVEY PROFESSIO" AL PAPER 155 PLATE 49 

5 

DENVEH FLOHA 

1. Gissus coryli/olia Lesquereux (M.O.Z. 1504), Denver formation. 2. Zizyphus hespeTius Knowlton, n. sp. (M.O.Z. 1534), Denver formation. 

3,4. Gissus lobato,cTenata Lesquereux (U.S.N.M. 37742, 37743), Dawson arkose, lower part (in beds= Arapahoe formation) 5 . GiS8US obovata Knowlton, n. sp., Denver formation. G. Vitis OITiki Heel' (U.S.N.M. 37844), Dawson arkose. 
U. S. GEOLOGICAL SURVEY • 

5 

PROFESSIONAL PAPER 155 PLATE 50 

3 

6 

DENVEH FLOHA 

I. Corn us lakesii KnowlLon, n. sp. (U.S.K.l\L 3i746), Denver formation. 

2. Cissus colora dens is Knowlton a nd Cockerell, X 2 (U.S.N.M. 37736), Dawson arkose, lower part (in beds=Arapahoe formation). 

3. Greu'iopsis tenuifolia Lesquereux ('G.S.N.M. 37i81), Dawson arkose. 

t. Vitis olriki lIeer (U.S.N.M. 37845), D~wson arkose. 

5. Sterculia? heterodonta Knowlton, n. sp. (U.S.N.M. 37837), Dawso n arkose. 6. Pterosllerrnites grandidentata Lesquereux (l\I.C.Z. 1519), Denver formation. 
U. S. GEOLOGICAL SURVEY PROFESSIOKAL PAPER 155 PLATE 51 

2 

1 

{} 

5 

4 

8 9 10 

DE TVER FLORA 

1, 2. Cornus impressa Lesq uereux (U.S.N.M. 37744, 37745), Dawson arkose. 3. Pterosper1nites grandidentata Lesquereux (M.C.Z. 1517), Denver rormation. 

t, 8. Sterculia' heterodonta Knowlton, n. sp. (U.S.N.M. 37837), Dawson arkose. 

5, G, 7, 9. Diospyros brachysepata AI. Braun (5, U.S.N.M. 37748; 6, 7,9, U.S.N.M . 

37749),5, Dawson arkose; 6,7,9, X , Denver rormation. 

10. Gissus grosse·dentata Knowlton (U.S.N.M. 37739), Dawson arkose. 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 52 

3 

1 

4 

5 

6 7 8 

• DENVER FLORA 

1. Zilyphus distortus Lesquereux (U.S.N.M. 37848), Dawson arkose. 

2. Nyssa Lanceolata Lesquereux (U.S.N.M. 37794), Dawson arkose. 

3. Diospyros brachysepaLa AI. Braun (U.S.N.M. 51055), Denver formation. 

4- 6. Fraxinu8 eocenica Lesquereux, Denver formation. (After Lesquereux.) 7. Styrax? laramiensis Lesquereux (M.C.Z. 1488), Denver formation. 

8. Viburnum richardsoni Knowlton. n. sp. (U.S.N.M. 37843), Dawson arkose. 
U. S. GEOLOGICAL SURVEY 

PROFESSIONAL PAPER 155 PLATE 53 

3 

2 

DENVER FLORA 

6 

J, 3, 1, 5. Viburnum richardsoni Knowlton, n. sp. (J, 3, 5, U .S.N.M. 37843; 4, :\I.C.Z. 1576), Dawson arkose. 2. Sterculia libbeyi Knowlton, n. sp., X IH (Pr.), Denver formation. 

G. Sterculia saportanea Knowlton, Denver? formation. (After Lesquereux.) 

4 
U . S. GEOLOGICAL SURVEY 

1 

PROFESSIONAL PAPER 155 PLATE 54 

2 

4 

DENVER FLORA 

I. Nvssa' obooata Knowlton, n. sp. (U.S.N.M. 37795), Denver formation. 

2,3. Viburnum richardsoni Knowlton, n. sp. (U.S.N.M. 37843), Dawson arkose 

4. Dombeyopsis magnifica Knowlton,n. sp. (U.S.N.M. 37755), Dawson arkose. 

3 
U . S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 55 

3 

1 

2 

5 

4 

7 

DENVER FLORA 

I. Viburnum contortum Lesquereux? (U .S.N.M. 37839), Dawson arkose. 2. Viburnum' heterodontum Knowlton, ll. sp. (U.S.N.M. 37840), Denver formation. a. VIburnum lakesii Lesquereux (U.S.N.M. 3784 1), Denver formation. 4. Dombeyopsis magnifica Knowlton, n. sp. (U.S.N.M. 37755), Dawson arkose. 

5. Viburnum me/aenum Knowlton and Cockerell (U.S.N .M. 37842), Dawson arkose. 6. Phyllites aristolochioides Knowlton, n. sp. (U.S.N.M. 37799), Dawson arkose, 

lower part (in beds=Arapahoe formation). 7. Phyllites andromeda Knowlton, n. sp. (U.S.N.M. 37797), Dawson arkose. 
(7. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 56 

2 

3 

,', /--'-' ' ... .. 

" " '-_.'1..... 

'M'" ........ "\'1'. " , \J \ _ .... -..... 

--.-

-". 

\ 

'\ '. I ._' 

! 

; -.......... ..-.. ] 

, 

" , 

... ... -"' .. 

5 

( 

/ 

, 

'\ 

'-

4 

I 

I , 

, 

, 

".I i 

, 

, 

,/ 

I 

, 

\ 

\ 

, 

, 

I 

I 

) 

\ 

I 

, 

, 

DE VER FLORA 

1. Phyllites calhanensis Knowlton, n. sp. (U.S.N.lH. 378(0), Dawson arkose. 

'2. Carpi/es quadrivalvis Knowlton, n. sp. (U.S.N.M. 37i22), Dawson arkose. 3. Phyllites sp. Knowlton (U.S.N.M. 37802), Dawson arkose. 

,. Phyllites pellucidus Knowlton, n. sp. (U.S.N.M. 37801), Dawso n arkose. 5. Dombeyopsis? sedaliwsis Knowlton, n. sp., X 2, Dawson arkose, lower part (in 

beds= Arapahoe formation). 
U . S . GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLA'l'E 51 

'.. /--- ."-~--.-. ' .. 

''-'-'---'~ '\"" 

\\/'\""'''':.-.-.: -. ~\.-' ' . . '''. fk' : .. '. \ '. .. ---'.. . -". -~---\ '-.., ./ .--;-' ..... , 

\ ; '-. i ./ ". \/ } , ;' 

"',! ,\ 

" 

3 

1 

DENVER FLORA 

1. Dombeyopsis magni,~ca Knowlton, n, sp, (U,S.N.1L 37755), Dawson arkose, 

2. Phyllites aristolochioides Knowlton, n, sp" X 2, Dawson arkose, lower part (in beds=Arapahoc formation), 

3. Castalia pulchella Knowlton, n, sp, (U,S.N,M. 37i25) . Dawson arkose, 
U. S. GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 58 

3 

4 

1 

6 

7 

DENVER FLORA 

1,2. Decorticated roots (U.S.N.M. 37747), Dawson arkose. 

3-5. Carpites coryloides Knowlton, n . sp. (U.S.N.M. 37723), Dawson arkose. 

6. Carpites templetoni Knowlton, n. sp. (U.S.N.M. 37721), Dawson arkose. 

7. Fraxinus sp. Knowlton (M.C.Z. 1487), Denver formation. 

8. Dombeyopsis magnifica Knowlton, n. sp. (U.S.N.M. 3ii55), Dawson arkose. 
U. S . GEOLOGICAL SURVEY PROFESSIONAL PAPER 155 PLATE 59 

-~- .. ---~ 

2 

1 

3 

DENVEH FLORA 

1. Dombeyopsisf sedaliensis Knowlton, n. sp., Dawson arkose, lower part (in 

beds=Arapaboe formation). 2. C'ir>namomum sezannense Watelet? (U.S.K.l\I. 37735), Dawson arkose. 

3. C'innamomum lini/olium Knowlton (U.S.N.i\I. 37734), Dawson arkose. 

4. Ficus eldridgi Knowlton, n. sp., Denver formation, Scranton, Colo. 

S. Phyllites denverensis Knowlton, n. sp. (U.S.N.M. 37803), Denver formation. 
INDEX 

Names in italics are synonyms; figures in black face refer to. descriptions] 

A Pago 

Acor sp. Knowlton, pI. 45, fig. G ___ .___________ 101 

Acor triloblltum productulll HeOl· ______________ ------------------------------ 10l· 

AcrostichulIl (Chrysodium) auroum Linn6 ____ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 2G 

Allantodia bmnoninna Wallich________________ 25 

AllnntQdiopsis Knowlton and Maxon __________ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 24 

AUcl7ltoriiopsis crosCl (Losquoroux) :Knowlton and l'd:axon, pI:;. 4, 5 __ 9,10,11,24-26,27 

Alnus aurada Knowlton and Cookoroll, pI. 15, fig. G_________________________ 49 

Alnus cnrpinifoIin 1,osquoroux, pI. 15, fig. 2__________________________________ 49 

Alnus cyclndum Ungor________________________ 49 Al.nus rugosCl 1,osquorOllx ______________________ ------------------------------ 49' 

Andromoda polifolin 1,inn6____ ------------------------------ 129 

Anomit\ sp. pI. 8, fig. 7.. _______ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 30 

Anomia olongntn (Nowborry) Knowltou_____________________________________ 30 

Anomin fromon(;i Knowlton 29 

A nomin Inncoolntn Knowlto;l~ -I;i.-8: fi~'- iii~~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ ~ ~ ~ ~ 29· 

Anomitl mosbyonsis Knowlton, pI. 8, fig. 9 ___________________________________ 28-29 

Anomill occidontlllis Knowlton______ __________ __ __ __________ __ ___ _______ __ __ 29 

Anolllill porploxn Hollick_______________________ _ 29 

Anomin suborotncoa (Snportn) Gardnor and Etti~~~h~~S-~~~~~~~~~~~~~~~~~~~~ 29 Al'llchis hypognon_ _ _ _ ___ __ __ ________ ______ ________ __ ____________ ________ __ __ 98 

A rnlil\ dissoctll Losquorollx- _ _ __ ___ ____ ________ __ __________ ______________ ____ 74 

Ara/iCl1mngcns Losquoroux ____________________________________________ 14,73-74,75 

Arnpnhoo formntion, doscription oL _____________________________________ 1-3 4 5 G 

110ra of. _________________________________________________________________ : ' '6 

Artocclr1Jidiu,711 ol.mec!;ac/oU·l£11I? Ung~r _______________________________________ 14,100 

Artoonl'J)Us'! gignntoa Knowlton_ ____ ________ __ __ ______________ __ __________ __ 75 

Artocnrpus lossigillnn (Losqnoroux) Knowltou _______________________________ 74,75 

ArtocllrplIs pllI.lgons (Losquoroux) Hollick, pI. 30, fig. 2; pI. 31; pI. 32, figs. 1, 2_ 73-74,75 

Artocnrpus similis .Knowlton, pI. 32, fig. 4 ___________________________________ 9 75 

Arundo'! obtusa Losquoroux ______________________________ ~__________________ '35 

Asimina ooconicn Lcsquoroux, pI. 40, fig. 4; pI. 43, figs. 1, 2,4, G __________ 10,94-95 

Aspidium fccundUlIl :IIoor ______ .____________________________________________ 17 

AS1Jic!;u,'1IIla/rcsii (Lcsquoroux) Knowlton ____________________________________ 14,18 

Aspidiulllllloyori Hoor_____________________ _ 17 

Aspidium quitollso Christonson - -------------------------------

Aspldillm rnddillllUlll Hookor __ -_--~~~~~~~~ ~~~~~~~--- ------------------ -- -----

Asplonium (Dipln.zium) crossii Knowlton ------~---------------------

17 

Aspionilllll ooligniticu III Berry ___________ -_ ~ ~~ ~ ~ ~~ ~ ~~ ~~ ~~ ~~~ ~ ~ ~~~ ~ ~ ~ ~~ ~~ ~ ~ ~ ~ ~ 

17 

23 

22 

AS1J/cniu:m cro.vn'1ll (Lcsquoroux) Knowlton _____________________ ~__________ 14,24 

AS1J/clliu,'III snbsi'llllJ/ex (1,osquoroux) Knowlton ____________________________ ~~ 14,24 

D 

13crchc'IIIia 'IIl'll/tincrvis (AI. Brnun) Hoor ____________________________ 9,10 14,102-103 

/3crchc'lllia lJarvi/o[iu LOSquoreux ________________________________________ ~ ___ 14,102 

Borrya .Knowlton, n. gen____________________________________________________ 133 

Borryn rnCOIllOSf\ (Knowlton) Knowlton, pI. 41, figs. 4,5 ___________________ 134-135 

Botuln Ilngulnta Gopport; __ · ____________________________________ -'_______ _____ _ 50 

Bot.ulll fnllnx Losqlloroux, pI. 15, figs. 7-10; pI. 16, figs. 1, 2 _____ .______________ 50 

Botula glnndulosa Michnux __________________ · 97 

g~i~ ~]f:if:;~~]:,~;,~~,~;~~~,: I~~: ~;~ :~~.: ~~ ~~.~ ~;;; ~ ~;;;;;;:;;;;;;;;;;;:;:;;: j 

BlochllUIll volubilc___________________________ - 28 

~;::~:~:;:~ ~:~:::~::::~ ~~~~:~~:\~~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ ~~ ~ ~ ~ ~~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ :~ C 

~~~l~~B~Jl::,;:{fi~~:f:~~.:~ ~ ~ ~ ~ ~: ~ ~"~ ~~ ~ ~ ~ ~: ~: ~ ~ ~ ~:::~ ~ ~. ~ ~: ~: ~ ~ ~ ~~::::. ::j Cnl'p~tos cofl'onofol'lnis LOSqllOI'OllX________________ _ __ ____________ ________ __ _ _ 132 

Cnl'p~tos ooryloidos Knowlton, pI. 58, figs. 3-5_______________________________ 132 

g:;~:i~: ';~:~:i:;Ei:~~:~~~::::::::::::::::::::::::::::::::::::::::::~" ::: 

Cnl'j.lltos mynclll'llln Losqllorollx ___________________________________________ 10,132 

Cnl'pitos oviforlllis Losquoroux_ _ __ __ __________ __ __ __ __________ __ __ _____ ___ __ 131 

Cnl'p~tos quadrivnlvis Knowlton, pI. 5G, fig. 2________________________________ 131 

Cnrpltcs rostollotus Losquorollx ___________________________________________ 131-132 

. Page 

Carpites templetoni Knowlton, pI. 58, fig, 6_________________________________ 131 

Carpol~thes arachioides Lesqllereux ___________________________________________ 14,98 

Carpollthes pulrnarum Lesquereux- ______________________ ~ ___________________ 14,41 

Carya antiquomm Newberry ___________________________________ ~ __ __________ 48 

Cassia emarginata Derry _ _ _ _______ __ ___________ _ ____________ __ __ _ _______ ____ 99 

Cassia vetusta Knowlton, pI. 45, fig. 2_______________________________________ 99 

Castalia? duttoniana Knowlton_________________ __ 94 

Castnlia flavo-virens _____________________________ ~ __ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ ~~ 94 

Castalia leei Knowlton______________________________________________________ . 94 

Castalia pulchella Knowlton, pI. 42, fig. I; pI. 57, fig. 3_______________________ 901, 

Castalia rudgeana _________________________________________________ . ___ __ ____ 94 

Castalia stantoni Knowlton _________________________ . _________ -_____ --- __ ---- 94 

Castalia stellata___ _______ _________ ______________ ____ __ ____ __ ______ __________ 94 . 

Castanea intermedia? Lesquereux ________________________________________ 9,10,50 

Ceanothus fibrillosus Lesquereux ____________________________________________ 14,108 

Celastrinites alatus Knowlton ______________________________________________ -- 100 

Celastrinites artocarpidioides Lesqueroux_________________ ___ ____ ________ ____ 100 

Celastrinites populifolius Knowltou, pI. 40, fig. 1; pI. 45, fig.H_______________ 100 

Celllstrus gaudini J~esquereux, pI. 45, figs. 3, 4, 9 ____________________________ 99-100 

Cerais oanadonsis Linnc_______ __ ____________ _____ ___ __ __________ ____ ________ 99 

Cerais ooloradensis Knowltou, pI. 45, fig. 5 __________ ._________________________ 99 

Cerais wilcoxiana Derry _________________________ .___ ___________ ____ __ ________ 99 

Chamaedorea? ooloradensis Knowlton, pI. 10, fig. 3 __________________________ olO-U 

Chamaedorea danai. ________________________________________________ ._ ------- 39 

Chrysobalanus coloradensis Knowlton, pI. 40, figs. 2, 3; pI. 43, figs. 3, 5, 7-9; pI. 44, figs. 1, 2 ___________________________________________________ 95-96 

Chrysobalanus eocenica Berry ___________________________________________ ---- 9G 

Chrysobalanus icaoo Linnc __________________________________________________ 95~96 

Chrysobalanus inaequalis (Lesquereux) Berry_______________________________ 96 

Chrysobalanus? lanceolatus Knowlton, pI. 44, fig. 5 ________________ "-_________ 9G 

Chrysobalanus oblongifolius Michau·x- ______________________________________ 95-96 

Chrysobalanus pellocarpus Meyer _________________________________ ·__________ 96 

Chrysobalanus pollardanus Knowlton _____________________________________ -- 96 

Cinchonidium? tumori Knowlton G4 

Cinnamomum sp. Knowlton, pI. 3~:fi~~-4;-~1~39~-fi~~~--~~~~~~~~~~~~~~~~~~~~~~ 87 

Cinnamomum affine Lesquereux _________________________________________ 71,86,87 

Cinuamomum ellipticum Knowlton __________________ " ______________ ._______ 87 

Cinnamomum linifolium Knowlton, pI. 59, fig. 3 ____________________________ 86-87 

Cinn::nnomum oblongatum Derry __________ " ________________________________ 8G-87 

Cinnamomum polymorphum AI. Braun_____________________________________ 87 

Ciunmnomum sezannense Watelet?, pI. 38, flgs. 7, 8; pI. 59, fig. 2 _____________ 86.87 

Cinnamomum vera Derry _________________________________________________ -- 87 

Cissus? oannoni Knowlton, pI. 47, fig. 5_____________________________________ 113 

Cissus coloradeusis Knowlton au'd Cockerell, pI. 47, fig. G; pI. 50, fig .. 2 __ 10,112-113 

Cissus corylifolia Lesqucreux, pI. 47, fig. 4; pI. 48, fig. 1; pI. 49, fig. L _______ 113-1U 

Cissus duplicato-serrata Lesquereux ____________________________________ ----- 57 

Cisslls grosso-dentata Knowlton, pI. 48, fig. 2; pI. 51, fig. 10 _________________ 10,112 

Cissus hesperia Knowlton, pI. 47, fig. L_____________________________________ 112 

Cissuslacvigata Lesquereux- ________________________________________________ 14,112 

Cissus lesquereuxii Knowlton, pI. 48, figs. 3, 4_______________________________ lU 

Cissus lobato-crenata Lesquereux, pI. 49, figs. 3, 4 __________________________ 114-115 

Cissus obovata Knowlton, pI. 49, fig. 5______________________________________ 11.1 

Cissus parrottiae/olia Lesqllereux- __ . __________________________________ 14,57,112, Hoi, 

Cissus prirnaeva Saporta _____________________________________________________ G4-G5 

Cissus speCtabilis Heel' _________________________________ ~ __________________ 14,95-96 

Cissus triwspiriata (Heer) Lesqueroux ___________________________ .------ 15,113,114 

Coniogramme fraxinea Don____ _ ____________________________________________ 25 

Cope, E. D., cited___________________________________________________________ 3 

Comus acuminata Newberry _____ _____ ______ ____ __________ _____ ___ __________ 111} 

Comus altemifolia Liune____________ ______ __ __ ________ ___ __ ___ ______ ____ ____ 119 

Comus buohi Heel' _________________________ -r- __________________________ ---- 120 

Comus deuverensis Knowlton, pI. 13, fig. L_________________________________ 119 

Comus emrnonsi Ward _________________________________________________ 15, lI8-UI} 

Comus holmesii Lesquereux _______________________________________ ~ _______ 119-120 

Comus lakesii Knowlton, pI. 50, fig. L______________________________________ 119 

Comus impressa Lesquereux, pI. 51, figs. I, 2 ______________________________ 118-119 

Corllus mas Linn6 ______________________________________________________ ---- 119 

Comus newberryi Holliok _________________________________________________ -- 119 

Comus orbifera Heer _____________________________________________ ~ __ __ ______ 118 

Comus paniculata L'H6ritier. _____________________________________________ ~_ 119 

Comus pubescens val'. oalifomica___________________________________________ 119 

Comus rhamnifolia O. Weber_______________________________________________ Ill} 

139· 
140 INDEX 

.Page Page 

Cornus sericea Linne__________________ __ _____ _____ ______________ _ _____ __ __ _ _ 119 Ficus planicostata latiColia Lesquereux ________________________________ ~_____ 72 

Cornu8 studeri? HeeL __________________________________________________ ]0,104-106 Ficus planicostata problematica Knowlton, pI. 28, fig. 2_____________________ ·71 

Crataegu8 antiqua Heel'. _ ____ ____ __ ______ ____________ ________ __ ___ _____ __ _ _ __ 5~ Ficus·praetrinervis Knowlton, pI. 28, figs. 8-10 ______________________________ 71.87 

Crataegu8 betulae/olil! 1,esquereux _________________________________ ]5,50,51,126-:-127 Ficus pseudopopulus 1,esquereux, pi. 25, figs. 3-5; pI. 26, figs. 2-4 ___________ 9,43,66 

Crataegus myricoides1,esquereux, pI. 44, figs. 7-10 ___________ .. _______________ 96-97 Ficus puryearensis Berry? _____________ c· __________________________________ 10,72.95 

Crataegus oxycanthoides Goppert_ ______________ _____ __ _ ___ __ _ __ __ ____ __ ____ 96 Ficus ramahensis Knowlton, pI. 27, fig. 5 ____________________________________ 68-69 

. Credneria? daturiaeColia Ward _________________________________ " _ __ ___ ______ 116 Ficus sezannensis \VateleL______________ ___ _ __ ____ ____ __ __ __ ____ ___ _ ________ 55 

Cross, ·Whitman, cited _______________ " ____________________________________ 1-3,5,6 Ficus speciosissima Ward_ __________________________________________________ 72 

D 

Daphnogene angelica? HeeL ____________ .. ______________________________ ]5,109-110 

Darton, N. H., cited _____________________ .__________ __ ______ _____ ___ ______ __ 4 

Dawson arkose,' description oC______________________________________________ 3-5 

Decorticated roots? pI. 58, figs. 1,2_ _________________________________________ 135 

Dennstaedtia crossiana Knowlton, pI. 2, figs. 7-9____________________________ 21 

Denver Basin, description _____________________________________ "___ _ _ _____ _ __ 1-3 

monograph, publication oL __ ~_ __ __ ______ __ ________ __ ____ __ ___ __ _ __ __ ____ 2 

. Denver Cormation, description oC __________________________________ ·_________ 1-3 

flora of. _ _____ ___ __ ______ __ ______ __ ________________ ______ ______ __ __ ___ ___ ·7-9 

Diospyros brachysepala AI. Braun, pI. 51, figs. 5, 6, 7, 9; pI. 52, fig 3 _______ 121-123 

Diospyros princetonia CockereIL____ ______ __ ______ __ _____ ___ __ __ ______ ____ __ 122 

Diospyros wilcoxiana Berry _________________________________ . ______________ 122-123 

Diplazium crossii (Knowlton) Knuwlton, pI. 2, fig. 6 ________________________ 23-24 

Diplaziopsis javanica (Blume) Carl Christensen_____________________________ 25 

Ficus spectabilis Lesquereux. _ ___ ____ ________ __ __ __ ____ ________ __________ __ __ 64 

Ficus subtruncata Lesquereux _____ ~ ______________________________________ 10,63.69 

Ficus tenuinervis Lesquereux. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 64 

Ficus tiliaefolia (AI. Braun) HeeL __ . __ . ______________________________ 66,67-68.97 

Ficus trinervis Knowlton____________ ___ _ _________ _ ____ ___ _ __ _ ___ __ __ ____ __ __ 87 

Ficus truncata? HeeL __________________________________________________ ~_ __ 63 

Ficus uncata Lesquereux ________________________________________________ 67,69.100 

Flabellaria communis Lesquereux ____________________________________________ 15,3() 

Flabellaria eocenica Lesquereux ________________________________________ ]5,36-37.40 

Flabellaria? fructi/er Lesquereux. ____________________________________________ 15,3() 

Flabellaria? longirachis'? UngeL ______________________________________________ 15,38 

Flabellaria zinkeni Heer ___________________________________________________ 15,39-40 

Fraxinus sp. pI. 58, fig. 7_____________________________________________________ 124 

Fraxinus denticulata HeeL ________ ._ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 12 .. 

Fraxinus eocenica 1,esquereux. pI. 52, figs. 4-6 _____________________________ 123-12" 

Diplazium muelleri? HeeL _______________________________________________ ]5,23-24 G Dombeyopsis grandi/olia Unger? ____________________________________ ~ __ ___ _ __ 125 

Dombeyopsis magnifica Knowlton, pI. 54, fig. 4; pI. 55, fig. 4; pI. 57, fig. ]; 

pI. 58, fig 8 ____________________________________________________ 125-126 

Dombeyopsis obtusa Lesquereux ____________________________________ 9,124-125.126 

Dom beyopsis occidentalis 1,esquereux. _ _ _ _ _ _ __ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 68 

Dombeyopsis'! seclaliensis Knowlton, pI. 56, fig. 5; pI. 59, fig. L______________ 126 

Dryopteris arguta (1,esquereux) Knowlton ___________________________________ 15,18 

Dryopteris integra Knowlton, pI. ], fig. 5____________________________________ 17 

Dryopteris lakesii (1,esquereux) KilOwlton, pl.], figs.], 2 ________________ ll,18-20 

Dryopteris lesquereuxii Knowlton_______ __ ______ __ _____ _ __ __ ___ ___ __ __ _ _ __ __ 20 

Dryopteris polypodioides? (Ettingshausen) Knowlton ________________ . ______ 20-21 

Dryopteris (Lastrea) polypodioides (Ettingshausen) Knowlton ________________ 20-21 

Dryopteris richardsoniana Knowlton, pI. 2, figs. 3-5_________________________ 20 

Dryopteris weedii Knowlton_ _ _________ _____________ __ ______ _____ __ _ ____ _ _ __ 20 

Geonomites sp., pI. 10, fig. L________________________________________________ 40 

Geonomites goldianus (Lesquereux) Lesquereux, pI. 10, fig. 2 __ "~ ________ 9,10,39-10' 

Geonomites? graminfolius Lesquereux.______ __ ________ ____ __ ____ ___ _ __ __ ___ _ 39 

Geonomites tenuirachis Lesquereux, pI. 6; pI. 9, figs. 1, 2, 6 __________________ 38-39 

Geonomites ungeri Lesquereux ____________________________________________ 15,37-38 

Gingko biloba Linne ___________ .____________________________________________ 34 

Ginkgo digitata (Brongniart) Heer. ____________ · ______ ~______________________ 34 

Ginkgo primordialis HeeL _ _________________________________________________ 3J 

Ginkgo siberica Heer _______________________________________________________ : 3 .. 

Ginkgo? truncata (1,esquereux) Knowlton, pI. 9, fig. 3 _______________ 34-35,131,132 

Glyptostrobus ungeri'! Heer_______________________________________________ ]5,33 

Goniopteris polypodioides Ettingshausen ______ . ___________________ ~_ ___ ___ _ _ __ 15 

Grewiopsis eocenica_ _ _______ ______________ _ _____ ____ _______ _ ___ ___ ____ ___ ___ 10 

Grewiopsis populaColia Ward ____________________________ :___________________ 80 

E GrewiopsistenuiColia Lesquereux, pI. 50, fig. 3_______________________________ 116 

Ecologic considerations ___________ . _____ ~_ ___ ___ _ _____ __ __ ___ _ __ __ ____ __ ___ ___ ] 1 Grewiopsis viburniColia Ward_ ____________ ____ _ _ ____ ____ ______ _ _ __ _____ ___ _ __ 80 

Eldridge, G. H., cited ________________________ : ______________________________ 1-2,3 Gymnogramma gardneri (Lesquereux) LeSC!llCreUx- ________________________ 6.15,26 

.Emmons, S. F., cited _______________________________________________________ ],2,3 Gymnogramma haydenii Lesquereux _________________________________ c______ 29 

Equisetum sp _____________________________ .__ _ __________ _ ________ _____ ___ ____ 32 Gymnogramma javanica_ _ _ ____ _________ ____ __ ___ ___ __ __ _ _ ____ ___ ___ __ ______ 25 

Equisetum sp. pI. 9; fig. 4___________________________________________________ 32 

Equisetum coloradense Knowlton, pI. 8, figs. ]-6~ ___________________________ 31-32 H 

Equisetum haydenii 1,esquereux. _ __________________________________________ 32 Hayden, F. V., cited ________________________________________________________ 1,3,5 

Equisetum laevigatum 1,esquereux ________________________________________ 6,32-33 Hedera bruneri Ward ______________ ~________________________________________ lIS 

Equisetum perlaevigatum CockereIL _________________________________________ 32-33 Hicoria antiquora (Newberry) KnowltoiL __________________ ~ ________________ 10,48 

Equisetum robustum Newberry_______________________________________ ______ 32 Holmes, \V. H., cited________________________________________________________ 6 

Eriocaulon? porosulll 1,esquereux ________________________________________ 6,132-133 Hymenophyllum bivalva Swartz _______ -- -7---- ___ __ __ _________ _ __ __ __ __ _ _ __ 17 

Eucalyptus hae.ringiana? Ettingshausen ________________________ .__ _ __ ___ _ _ _ _ ]33 Hymenophyllum confusum Lesquereux, pI. 1, fig. 4_________________________ 17 

F 

Hymenophyllum fuciCorme Swartz_ _________________________________________ 17 

Hymenophyllum protrosulll Hooker ______ ____ __ _ ___ _ __ _ _ __ ____ __ _ __ _ __ _ _ __ __ 17 

Ficus sp. pI. 27, figs. h3; pI. 29, fig. 5; pI. 30, fig. ] ________________________ 67.72.73 

Ficus aguilar. Knowlton, pI. 24, fig. 4 _________ . ______________________________ 9,65 

Ficus? alaskan a Newberry ____________ ~_____________________________________ 118 Ilex? ovata Knowlton, pI. 46, fig. 4___________________________________________ 102 

Ficus? alata Knowlton, pI. 23, fig. 9_________________________________________ 64 J Ficus andrai 1,esquereux___________________ _ ___ __ __ __ ___ ___ __ _ ____ _ __ _____ _ __ 66 

Ficus arenacea brevipetiolata Lesquereux____________________________________ 67 

Ficus artocarpoides 1,esquerux. _______________________________________ : _ __ __ ]06 

Ficus asari/olia Ettingshausen _________ c_ c _________________ .__ __ _ _ _ _ __ _ __ ___ __ i3 

Ficus auriculata Lesquereux________________ _ _ ___ __ _ _ ____ __ __ _ __ _ _ _ __ _ __ _ ____ 63 

Ficus berthoudi 1,esquereux, pI. 25, figs. 1, 2, 6 _______________________________ 65-66 

Juglans sp., pI. 14, fig. 5; pI. 15, fig. L________________________________________ 48 

Jug/a1l8 acnminata? AI. Braun _______________________________________________ 46-4i 

Juglans corrugata Ludwig_____ ____ ______ ______ __ __ __ _ _ __ ___ _ __ _ _ ___ _ __ ___ _ __ 48 

Juglans denveriana Knowlton, pI. 12, figs. 1, 2; 1)1. 13, figs. 2-4 ____ .... -45.48, 101, 124 Juglans nigella HeeL____ ____ ______ ______ ____ __ ____ __ __ _______ _ _ _ __ __ __ ______ 47 

Juglans praerugosa Knowlton _______________________________________________ 46-47 Ficus clintoni 1,esquereux ________________ ._ ______ __ __ ___ _ _ _ _ _ __ __ ___ _ __ __ __ _ _ 70 

Ficus cockerelli Knowlton___ __ ______ ______________ __ __ _ __ __ _ __ __ __ __ __ __ _ __ _ 72 

Ficus coloradensis CockereIL ______________________________________________ 9,10,69 

Juglans rhamnoides 1,esquereux_ --- _______ -c- --- - - -- - - -- - - -- __ - ______ - -_ - ___ . 10,46 

Juglans rugosa Lesquereux, pI. 14, fig. L _________________________ 9,10,45, -16-48. 120 . 

Juglans schimperi Lesquereux _____________________________________________ 9,10,45 

Ficus crenata Unger. ____________________________________________ • __ __ _ __ _ ___ 59 

Ficus clawsonensis Knowlton, pI. 26, fig. L__________________________________ 67 

Juglans thermalis 1,esquereux, pI. 13: fig. 5 ______________________ . _ ____ _ __ __ __ 45 

Ficus denveriana Cockerell, pI. 24, figs. 1-3 ______________________________ 64-65.106 L 

Ficus eldridgi Knowlton, pI. 59, fig. 4________________________________________ 69 Laramie Cormation, description oC _________________________________________ _ 

Ficus harrisiana Hollick ______________________________ " _________________ "_ _ _ _ 68 flora of. ______________________________________________________ ._ __ _ _ ____ __ 9 

Ficus irregularis (1,esquereux) 1,esquereux. _________________ ~----------- 6\),104; lOt} Lastrea (Goniopteris) polypodioirles'! (Ettingshallsen) Lesquereux ___________ 6,15,20 

Ficus lakesii Knowlton, pI. 28, fig. L________________________________________ 69 Lathraea argutu Lesquereux _______________________________________________ 15,18,19 

Ficus latiColia (1,esquereux) Kno\dton______________________________________ ]27 Laurus brossiana (Lesquereux) 1,esquereux. ______________________________ 15,85-86 

Ficus leei KnO\dton____ _ ___ _ ______ ______ ______ ____ ___ _ __ __ __ _ __ __ _ ___ _ __ __ __ 67 Lallrus lanceolata Knowlton, pI. 38, fig. 9____________________________________ 85 

Ficus martini Knowlton, pI. 40, fig_ 5________________________________________ 69 LrL'UT1ls obovata'! Weber ______________________________________________________ 15,83 

Ficus monodon (Lesquereux) Berry ___ ~ _____________________________________ 68-69 Laurus ocateaeColia Ettingshausen_________ _____________ _ ___ _ __ __ __ __ ___ _ ____ 84 

Ficus neoplanicostata Knowlton, 1;1. 28, figs. 3, 4, 6, 7;'p1. 29, figs. 3, 4 ___ 9, 10,69-70 1,aurus primigenia Unger, pI. 38, figs. 3, 5, 6 ___________________________ 10,83-8-1.85 

Ficus occidental is (Lesquereux) Lesquereux, pI. 26, fig. 5; pI. 27, fig. 4 _____ 9,10,68 Laur-us socialis Lesquereux, pI. 38, fig. 4 ________________ · _____________________ 9,85 

Ficus ovalis 1,esquereux___ __ __ __ __ ____ __ __________ __ __ _ _____ __ __ ______ __ _ ___ 6f Laurus utahensis Lesquereux________ __ ____ ____________ _____ _ ____ ____ __ _ _ ____ 86 

Ficus planicostata Lesquereux ______________________________ 67,68,69,70,71,123,127 1,e Conte, J. 1,., cited ______________________________________________________ _ 

Ficus planicostata clintoni (Lesquereux) Knowlton___________________________ 70 Lee, \Y. '1'., cited ___________________________________________________________ _ 

Ficus ·planicostata goldiana Lesquereux ________________________________ 9,67,.70-71 Leguminosites? arachioides (Lesquereux) 1,esquereux ___________________ 9,10,98-99 
INDEX 141 

Page 

Losquoroux, Loo, cltod _______________________________________ --------------_ ],6 

Llquldombar'l cuchoras KnowltOIl _________________________________________ ]29-130 

LygodlulU'lnntlquorulU Shlrloy ___________________ ~__ _____ __ ___ _ ____________ 30 

LygodllllU cnplllnro Wntolot__________________ _____ ____ ______ ____ ____ ________ 31 

Lygodllllll clrcillotulll (Bllrmnnn) Swnrtz___________________________________ 31 

Lygodllllll colorndonso Knowlton pI. 8, fig. 8 ____________ .' __ -" _________________ 30-31 

LygodllllU cOlllpnctllm Losqlloroux __________________________________ '_ _ __ __ _ _ 31 

[,ygodllllll crnsslcostatllm Wntolot. ___________________ -'__________________ ____ 31 

Lygodiulll crotncoum Doboy nnd Ettlngshnllsoll_____________________________ 31 

Lygodlulll dlchotOlllum Swnrtz______________ ____ __ ___ ____ __ _______ __ ____ ___ _ 30 

Lygodillm dlstrnctulll_______________________________ _ ___ __ ________ __________ 31 

Lygocliulll oxqulsltulll Snport.n_ _ _ _____________ __ _ _____ ______________________ 31 

Lygocliulll goosoloti :Frl tol. _ _ __ _ _ _ _ _ _ _ _ __ __ _ _ _ _ _ _ ____ __ __ _ _ _ _ ___ _ _ _ __ _ _ _ _ _ _ __ 31 

LygoellulIl jrlpolllculIl _________________________________________ ~_______ _ _____ 30 

LygodluUl koulrussli Iloor_________ __ ___ _ __ ______________ __ __ ____ _ __ ___ ______ 31 

I~ygodllllll pairnatlllll ________________________________________________________ 30-31 

Lygodlurn pllrvifulillm Snportn______________________________________________ 31 

Lygocllulll rnclintlllll PnllltL.________ ______ ______________ __ ____ ________ __ ____ 30 

Lygocliulll.roticullltulI1- _ ______________________ _ _______ _____ _ __________ ______ 30 

Lygodilllll scundons_ ____________ ______________ ____________ __________________ 30 

Lygodiunl smilnciroliulll (Stornborg) Ettillgshnuson_________________________ 30 

I.,ygoellmll stllchoi Stur ____ -_______________ -_______________ : _____________ .,_ _ _ 30 

Lygocliulll tonolllllll Snporta ___________________________________________ .. ____ ' 31 

LygodlulIl trichomnnoldcs Lcsquorcux ______________________________ ,._ ______ 31 

:IV!: 

1\1 ngllolin losloynnn Losquoroux __________ -- -- -- -------- -- -- -_ _ _ _ _ ___ __ _______ 88 

Mllgnolill IIlllgnirolill Knowlton ________________________ , _________________ g, 10,87-88 

,Ma(l'lloliu tenll.illervi8 Losquoroux __________________________________________ 15,87.-88 

Mnrchautitos'l colol'lldollsis Knowlton, pI. I, flg. :1_, __________________________ 16-17 

M'arvino, A. n., citod _________________________ .______________________________ ],5 

Mntorlnls stud lod ____________________ - ----- _ __ __ __ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ 13 

1\1 oulspormitos cyclophyllus l,osquoroux ____ - --_ ------------ __ _ __ __ __ __ _ _ _ _ _ _ 89 

1\1'olllsporlllitos grnlldis Losquoroux_ ------- --___________ - _____________ ,_ __ _ _ _ _ 89 

Mospllc:idaphuo psolldoglnucn Borry _ ------ - __ '_______________________________ 85 

l\-Jlddlo ]>1\I'k rorlllotion, doscrlption ot. _____________ " ______________________ _ 

M:OIlUlllout Crook group, ronallloe1.__________________________________________ 3-4 

Musophyllulll complicatuIll Lcsqllorollx_____________________________________ '102 

1Vlyrlca sp., pI. 11, flg. 7_____________________________________________________ 4t 

1\1:yrlca? losslglnna Losquoroux ______________________________________________ 74,75 

Myrica losslgii Losquoroux_ _ ____________ ____ ______________ ________________ __ 74 

Ivlyrlca torroyi Losquoroux __________ -- -------- ______ - __ -_ _ _ __ _ _ __ _ _ _ _ ___ __ __ 44 

N 

Noct.nlldrn psoudocorioCOl\ Borry ____________________________________________ 84,85 

Nogundo docllrrcns Losquoroux, pI. '15, fig. 10 ___________________________ ' ___ 100-101 

NOglllldo trilobll Nowboq'y ________________________________________________ 100-101 

Nellllllbitos virgillionsis (Fontnillo) Borry ___________________________________ 89-90 

Nolll!ll biulll nrtlculll Boor ____________ - __ --- -'_ _ __ __ _ _ _ _ __ _ __ _ _ __ _ _ _ __ _ _ _ _ __ __ 89 

Nolumbillm cholYnti Saporta________________________________________________ 89 

Nolulllbiuul hosltl\IlICllm Saporta _______ ----- __ -- ________ --- ___ ~_______ ______ 89 

Ne/1£'IlIbiu711 lClkcsi(I7IWIII .Losquoroux _____ --- -- ________________________________ ]5,91 

Netl£7I!bill,'11I1ukesii Losquorollx ____________________________________________ 15,90,91 

Nolulllbiulll prot.ospoclosunl Sl\portll_ _ __ ______________________ __ ____ __ __ ____ 91 

NolulUiJlulll provlnclalo Saportll_____ ____________ ______________________ ______ 91 

Kolumblulll pyglllOOUIll Dawson____________________________________________ 90 

Nolulllbium snslmtchuonso Dawsoll _________________________________________ 90,93 

Nelu'lllbil£'1II tenni/olil£'/II Losqllorollx ______ ---- ___________________________ 6,15,90,92 

Nolumbo nrcticll (Hoor) 'fuzsou _____________________________________________ 89,92 

Nolumbo crossli Knowlton, pI. 41, fig. :3 _____________________________________ 90,93 

Nolulllbo dnwsoni Hollick _________________ - _________________ __ __ ____ __ __ ____ 90 

N clum bo intcl'lllocilo Knowltol.l ________ - -- -- _____________ -_ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ 90 

Nolurnbo kompli (Hollick) Hollick _______________ .' __________________________ 89-90 

Nolumbo lalwslalla (Losqllorollx) Knowlton, pI. 41, fig. 1; pI. 42, flg. 2 ___ !JO, 91-92,93 

Noillmbo 1Ill'UluioLlSIs Hollick________________________________________________ 90 

Nolllmbo lutoa (Willtlonow) Porsoon _______________________________ 88, !JO, 91. 92, 93 

Nolurnbo nolulllbo (Linno) Karston ______________________________________ 88,91,93 

Nolumbo prilllaovl\ Borry __ _ ________________________________________________ gO 

Nolurnbo protolutoa Bcrry _ _ _ _ _ ____________ ___ ____________ _ ____ __ _ _ __ _ __ __ __ 91 

Nolumbo tonllirolia (Losqlloroux) ,Knowltoll, pI. 41, flg_ 2 ________________ !J, 90, 92-93 

Nouroptoris sChouchzori. _ _ _ _________ __ __ __ _ _______ ______ ____ _ _ _ _ ___ __ _ ______ 30 

Nymphaoa lIavo-vlrons LohmulIlI_ _ _ _ _______________________________________ 94 

Nymphaoa J'lIc1gonnl\ :Myor ____________ ______ ______ ______ _ _ ____ __ ___ _ __ _ _____ 94 

Nymphooa stollntll Willclonow _______________________________________________ ' 94 

Nyssa aquatica I,inlll)_ _ _ ___ _____ ___ _ _________ __ _ _ ____ _____ __ __ ______ __ __ __ __ 121 

NYSSll nrctica Hoor' __________________________ ' ______________________________ 121,134 

Nyssa billol'U Wultor_ _ _ _ ______ __________ ______ __ _ ___ _ __ __ _ _ __ ____ __ _ _ _______ 121 

Nyssa? cunoatll Nowborry _ ____________ __ ____ _ ___ __ __ ____ __ _ _ __ __ __ __________ 121 

Nyssa donvorinna Kllowlton _______________________________________________ 120-121 

NYSSllluJlcoolatu Losquoroux, pI. 52, flg. 2 ___________________________________ 10,120 

Nyssn maxinlll Wobor ____________________________________________ ~__________ 121 

Nyssa? obovntll Knowlton, pI. 50\, IIg. 1._____________________________________ 121 

Nyssa ornithobl'oma Unger____ ___ _ _____________________ _________ ______ __ ____ 121 

1\'1188(1'1 raccmOS(1 ,Kllowlton ________________ , ________________________________ 15,134 

Nyssidium oklllfini Hoer______________________ ______________ ___________ ______ 134 

Ho;W:!-i:\O--14 

o Pu~ 

Oreoduphne mississippiensis Berry _______________________ . ________ _ __ _____ __ 84 

Oreodaphne obtusifolia Berry _______________________________________ : __ ______ 123 

Oreodoxites martionlls ,oopper.t. ________________________________ . __ ______ ____ 41 

Oreodoxite8 pliCCltus Lesqllerellx ______________________________________________ 15,41 

Osmllnda amnis (Lesquereux) Lesquereux ________________________________ 15,27-28 

Osmunda major Lesquerellx _______________________________________________ 1.5,2,(,-25 

P 

Palaeoaster inquirenda _________________________________________ ~__ __ ______ __ 133 

Paloonelum bo KnowltOll, n. gen _____________________________________________ 93-94 

Paleonelumoo macroloba Knowlton, pI. 39, fig. 3; pI. 42, figs. 3, 4 ____________ 91,93 

Paillirus coloradensis Lesquereux______ __ _____ _ _____________ _________________ 108 

Paluirus ovoideus (Goppert) HceL ______________________________________ ~___ 108 

PCllmacites goldiClnlls Lesqllereux ____________________________________ .. _____ 15,39-40 

Palmocarpon commune Lesquereux _______________________________________ 15,41-42 

Polmocarpon? corrugatum Lesquereux_ _ ____________________ ________________ 42 

Palmocarpon? subcylindricum Lesquereux ________________ :_____________ ____ 42 

Palmocarpon? lineatum Lesquereux, pI. 11, fig. 5 ____________________________ 42-43' 

Polmocarpon palmarum (Lesquereux) Knowlton ____________________ ~ __ 9,10,41-42 

Palmocarpon·truncatum Lesquereux_ _ _____________ ______ _______ _ _ _ _ __ __ ____ 42 

Palmocarpoll truncatulU major Lesquereux ______________________ ,___________ 42 

PCllmocarpon truncatum minor LesQuereux _______________________________ ]5,34,42 

Palmoxylon cannoni Stevens__________ ____________ _ ___________ ___ _ __________ 43 

Paloreodoxites Knowlton, n. gen____________________________________________ 41 

Pnloreodoxitos plicatus (Losquerellx) Knowlton, pI. 11, figs. 1-4 ______________ 9,41 

Parrotia ragirolia Ettingshallsen_ - ___________________________________________ 15,99 

Parrotia pristina Ettingshallsen_ - __________ _____ __ __ _______ _ __ _ _ __ _ ___ ____ __ 113 

Peale, A. C., cited ___________________________ -- --____ ____ _ ____ _ __ __ ______ ____ 1,3 

Persea brossiann LesquereulC. _ -- -___________ -- ______________________________ 85-86 

Persoa carolinensis Nees ____________ ---- -- ---- ----__ __ ____ __ _ __ ____ ___ _______ S6 

Persen dilleri Lesquereux __________ ------ -- -------_____ __ __ _ _ _______ _ __ ____ __ 85 

Persen punctulata LesQuereux _____________ ---- -- -_ ____ ________ _ _ ______ ____ __ 85 

Pharagmites ooningensis HeeL _______________________________ .. _____ __ _ ___ __ J(j 

Phaseolites coloradensis Knowlton, pI. 45, fig. 1. __ :_________________________ !Ii 

Phaseolites oeningensis Ileer - -------------------- ________________________ ~__ . 9i 

Phyllites sp. pI. 56, fig. 3 ___ ~ __________________________ . _________________ . ____ 130 

Phyllites andromeda Knowlton, pI. 55,fig. 7________________________________ 129 

Phyllitos aristolochioides Knowlton, pi. 55, fig. 6; pI. 57, fig. :? ______________ 129-130 

Phyllites calhanensis Knowlton, pI. 56, fig, L_______________________________ 130 

Phyllites denverensis Knowlton, pI. 59, fig. 5________________________________ 130 

Phyllites pellucidus Knowlton, pI. 39, fig. 2; pI. 56, fig. 4 __________________ 130-131 

Phl1sageniCl sp _________________________ ----------- ___________________________ 15,32 

Physical conditions _______________ -________ --________________ ___________ _ __ __ 13 

Piper antiquum Heel' __ _ ____________________________________________________ 43 

Piper heerii Lesquereux, pI. 11, fig. 6_________________________________________ 43 

Pithecolobium eoconicum Derry - - _. _________________________ ._______________ 97 

Platanus sp., pI. 38, fig. 2____________________________________________________ 83 

Pilltanus aceroides Goppert. ___________________________________ ' __________ 9,10,83 

Platanus aceroides latifolia Knowlton, pI. 33, fig. 1. ______ ~ ___________________ 76-77 

Platanus coloradensis Knowlton, pI. 37, figs. 1, 2 __________ ~ __________________ 82-S3 

Platanus guillelmae Goppert, pI. 33, fig. 2; pI. 34, fig. L __________________ 76,80.83 

Platanus hayclenii Newberry, pI. 36, fig. L ____________________________ 7i,79-80,8:l 

Platanus heeri Newberry --------------- ----------_ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 83 

Plat!lnUS marginata (Lesquereux) Heer, pI. 36, figs. 2, 3 __________ , ________ 10,81-82 

Platanus newberryana Heel' _____ -- __ -- __ _ _ __ ___ _ _ ___ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ 8:~ 

Platanus nobilis Newberry ----- ------------------ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ SO 

Platanus occidentalis _______________ .' ____________ -______________ __ _____ ______ 71l ' 

Platonus platanoicies (V'lsquereux) Knowlton, j)1. 36, fig. 5 ________________ 9,10.82 

Platanus rnynoldsii Newberry, pI. 35, fig. L ____________________________ 9,10.77-78 

Platanus raynoldsii integrifolia Lesquereux, pI. 34, fig. 2 ______ . ________ 10,7i. 78-79 

Platanns rhomboidea Lesquereux, pI. 32, fig. 3 ___________________ !J, 10, (il. 75-76,'116 

Populus sp., pI. 21, fig. 9 _________________________ • ____________________________ 57-58 , 

Populus amblyrhynca Ward_ --_____________________________________________ 61 

Populus arctica Hoer _ _ _ _________________________________________ __ __ __ ______ 5S 

Populus arctica decipions CockoreIL________________________________________ 58 

Populu8 attenuata AI. Braun, pI. 21, fig. 9 _______________________ ' _________ 57-58,59 

Populus crenatCl Unger ____________________________ ~ ___________________ 51-52,5.1,57 

Populus denverensis Knowlton, pI. 23, fig. 5 _________________________________ 61-62 

Populus decipiens Lesquereux_________________________ __ __ _ __ _ __ __ __ __ _ _ __ __ 58 

Populus heliaclum UngeL __________________________________________ . ________ 64 

Populus jacksoni Knowlton, pI. 21, figs. 6-8; pI. 22, flg. 7 __________________ .__ 58 

Populus knowltoni Berry, pI. 23, figs. 1-4 ________ ~""_________________________ 61 

Populus lacoeana Knowlton, pI. 22, fig. 5 ____________________________________ 60-61 

Populus latior-transversa TIeer. _ _ __ ___ ________ ___ __ _ __ __ _____ ___ _ _ _ __ _ __ _ ___ 5!J 

Poriulus Inonodon Lesquereux ________________________________ . ___ '___________ 69 

Populus mutabilis crenulata Ileer - _________ - __ ' _____________ : ___________ . _ _ _ _ 62 

Populus mutabilis ovalis Heer ___________________________________________ 61-62,.100 

Populus nobrascensis Newberry _______________________________________ 10,55-57, fi9 

Populus nebrascensis acuticlelltata Lesquereux, pI. 21, figs. 1.2______________ 56 

Populus nebrascellsis grandicientata Losquoreux, pI. 19, figs. 4-7; pI. 20, figs. 2,6,7 ___________ : ______________________________________________ _ 

Populus nebrascensis longifolia Lesqucrcux, pI. 21, figs. 3-5 _________________ _ 

Populus nebrascensis rotundntn LcsCluerollx, pI. 20, figs. 1; 3,4,5, S _________ _ POPl!lus nervosa Newberry _______________________________________________ . _. 

Populus nervosa elongata I'\cwberr~' .. ________ . _ . __________ .. _________ . _____ _ 

56 

56-57 

56,59 

80 

SO 
142 INDEX 

Page 

Populus richardsoni Heer?, pI. 23, figs. 6,8__________________________________ 62 

Populus smilacifolia Newberry _______________________ _______________ ________ 55 

Populus speciosa Ward ______________________________________________ ~_______ 55 

Populus subrotunda Lesquereux, pI. 22, fig. 9 _____________________________ 10,59-60 

Populus subrotundata Lesquereux_ __ _______________________________________ 59 

Populus tenuinervata Lesquereux, pI. 22, figs. 1-4, 6 _________________________ 58-59 

Populus ungeri Lesquereux __________________________________________ '_ 15,64-65,100 

Populus zaddachLHeer, pI. 22, fig. 8_________________________________________ 60 

Populus zeilleri (Lesquereux) Knowlton, pI. 18, fig. 8; pI. 19, figs. 1-3 ________ 54-55 

Protoficus crenulata Saporta _____________________ ,_ _ _ _ _ _ __ _ _ _ _ _ _ __ _ __ _ _ _ _ _ _ _ __ 55 

Protoficus insignis Saporta __ ' ____________________ ,---- ______ __ ________________ 55 

Protoficus sezannensis (Watelet) Saporta____________________________________ 55 

ProtoficlLS zeilleri LesquereuL _________ " _______ ~ _____ ~ ____________________ 15,54-55 

Prunus angustifolia Marsh ______________________ ~___________________________ 97 

Prunus denverensis Knowlton, pI. 44, figs. 3, 4, 6; 1L________________________ 97 

Prunus scottii HeeL _ _ __________________________________ __ ________________ __ 131 

Prunus serotina EhrhardL_ _ __ __ _ _ _ _ __ _ _ _ __ _ __ _ __ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ 97 

Pteris ajJinis Lesquereux _____________________________________________________ 27-28 

Pleris anceps LesquereuL ________________________________________________ 15,27-28 

Pteris eocenica ___________________________________________________________ , ,__ 25 

Pteris erosa Lesquereux ___________________________________________________ 15,2<1-25 

Pteris gardneri Lesquereux ___________________________________________________ 15,26 

pteris parschlugiana Unger---- ____ ____________ __ ________ ____ ________________ 25 

Pteris pennae/orrnis HeeL ________________________________________________ 15,22,25 

pteris pseudopennae/ormis LesquereuL _____________________________ 10,15,22,27,28 

Pteris subsimplex Lesquereux _____________________________________________ 15,24-25 

Pteris undulata Lesquereux- ______________________________________________ 15,2<1':'25 

Pterocarya americana LesquereuL __________________________________________ 48-49 

Pterocarya? retusa Lesquereux, pI. 14. figs. 2-4; pI. 1.5, fig .. 5 __________________ 48-49 

Pterospermites grandidentata Lesquereux, pI. 50, fig. 6; pI. 51, fig. 3_________ 118 

Pterospermites inaequifolius Saporta ______ c_ ________ ________________ ______ __ 118 

Q 

Querc'us attenuata GopperL__________________________________________________ 51 

Quercus celastrifolia Lesquereux, pI. 18, figs. 4-7_____________________________ 54 

Quercus chrysolepis Liebmann ____________________________ ,__________________ 52 

Quercus coloradensis LesquereuL ______________________________________ ~____ 54 

Quercus engelmanni Gree~e_________________________________________________ 52 

Quercus etymodrys Massalongo--------------------------------------------_ 53 

Quercus etymodrys castellinensis CapellinL_________________________________ 53 

Quercus furcinervis RossmassleL ____________________________ ~_ _ _ _ _ _ _ _ _ __ ___ _ .53 

Quercus haidingeri Ettingshausen---------------- ____________________________ 52,53 

Quercus leon is Knowlton, pI. 18, figs. 2, 3 _______ ,_____________________________ 53 

Quercus paucidentata Newberry ________________________________ '____________ 53 

Quercus purdonensis Knowlton, pI. 17, fig. 2 ________________ '________________ 52 

Quercus? sedaliensis Knowlton, pI. 17, fig. 4 _________________________________ 52,53 

Quercus stramineus? Lesquereux-_ _ _ _ _ _ __ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ __ _ _ _ _ 54 

Q'uercus triang'u{aris GopperL _ _ ______ _______ _ __ ____ _ ________ ______________ __ 51 

Quercus viburnifolia Lesquereux, pI. 16, figs. 3-8; pI. 17, fig, L ___________ 50-52,113 

Quercus whitmani Knowlton, pI. 17, fig. 5 ___________________________________ 52-53 

Quercus whitei Lesquereux, pI. 17, fig. 3; pI. 18, fig. L_______________________ 53 

Quercus wislizeni A. de Candolle ______________________________ ~_____________ 52 

H 

Haton formation flora ________ ~____ __________________________________________ 9-10 

Rhamnites knowltoni Berry _______________________________________________ 105-106 

,Rhamnus alaternoides HeeL_________________________________________________ 103 

Hhamnus cannoni Knowlton, pI. 46, fig .. 5___________________________________ 107 

Hhamnus cleburni Lesquereux, pI. 40, fig. 6; pI. 46, fig~. 10, 11_ 9,10,103,104-106,107 

Hhamnus crenatulus Knowlton and CockerelL ____________________________ 106-107 

Rhamnus crenatus Lesquereux ______________________________________________ 15,106 

Hhamnus goldianus Lesquereux ___ ~ ______________________________ 9,10,102,103-104 

Rhamnus goldianus latior Lesquereux-______________________________________ 104 

Rhamnus grosse-serratus HeeL __ '___ ______ ____ ____ __ __ _____ ___ ________ ______ 106 

Rhamnus obovatus Lesquereux- _ ___________________________________________ 107 

Rhamnus praealaternoides Knowlton, pI. 46, figs. 7,8________________________ 103 

Rhamnus salicifolius Lesquereux'?, pI. 46, fig. 12______________________________ 107 

Richardson, G: B., cited ________________________________________________ ~ ___ 4-5, i 

S Sabal campbelli Newberry ___________________________________________________ 15,37 

Sabal communis LesquereuL __ " _____________________________________ -_______ 15, 36 

Sabal? eocenica (Lesquereux) Knowlton _____________________________________ 15,36 

Sabal grandi/olia Newberry _______________________________ ~ _________________ 15,37 

Sabat? grayana Lesquereux __________________________________________________ 15,36 

Sabal inquirenda Knowltol1 ___________________ · _____________ , _______________ 15,36-37 

Sabal mexicana Martins ____________ ' __________________________ ._ ________ __ ____ 34 

Sabal montana Knowlton _____ --c --- ___ - -- -- -- ______ --- -____ __ ______ _____ ___ 37 

Sabal? 'IIngeri (Lesquerellx) Knowltoll ____ . ____________________________ 9,11,37-38 

Sa ba lites /ructi/er Lesqllereux- ______________________________ ~ ______ 15,36-37,134 

Sabalites grayanus (Lesqllereux) Lesquereux, pI. 9, fig .. 5 _______________ 9,10,36-37 

Saccoloma sp., pl. 3, figs. 1, 2________________________________________________ 27 

Saccaloma elegans Kaulfuss __ c ____ -- _________________________________ • ______ 11,27 

Saccoloma gardneri (Lesquereux) Knowlton, pI. 3, figs. 3.:.8 ___________________ 26-27 

Saccoloma imrayanum HookeL_____________________________________________ 27 

Page 

Saccoloma wircklei Christ __________________________________ ~ _________ .,______ 27 

Salix media? HeeL __________________________________________________________ 15,83 

Salix plicata Knowlton______________________________________________________ 129 

Salpichlaena anceps (Lesquereux) Knowlton, pI. 7, figs. 1-3 ___________ 11,22,27-28 

Salpichlaena volubilis J. Smith______________________________________________ 28 

Sapindus affinus Newberry ____ _ _ __ _ _ _ _ _ _ _ _ __ _ _ __ _ _ _ _ __ _ _ __ __ _ _ __ _ _ _ _ _ _ _ _ _ ___ 102 

Sapindus alatus Ward _____________________________________________________ 122-123 

Sap indus berryanus Knowlton, pl. 46, figs. 1,2 ____________________________ 101-102 

Sapindus caudatus Lesquereux- _________ ~ ______________________________ 45,101,120 

Sap indus glendiyensis Knowlton ___________________________________ '_ _______ _ 122 

Sapindus knowltoni Berry ________________________ :_~___________ ____________ 101 

Sapindus? obtusifolius Lesquereux?, pI. 46, fig. 3 _________________________ 10,97,102 

Sclerotites? cypericola Knowlton, pl. 1, fig. 7________________________________ 16 

Sclerotites? hesperius Knowlton, pI. 1, fig. 6_________________________________ 16 

Sclerotites rubellus (Lesquereux) lVIeschinellL _________________________ " _____ 15-16 

Sclerotium rubellum Lesquereux- _ _ ____________ ________ __ ____ ________________ 15 

Selaginella berthoudi Lesquereux- ________________________ ,___ ____ ___________ 33 

Selaginella mertensiL_______________________________________________________ 33 

Selaginella stolonifera_ _ _ __ __ __ __ _ _ __ __ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ ___ _ _ _ _ __ __ _ _ ____ 33 

Sequoia affinis LesquereuL ____________________________________________ ~ ____ 33-34 

~~~~o~;al~:i:~~~~:f!;:;ge~~~~-t---~~ ~~ ~ ~ ~~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~ ~ ~~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~ ~~~ 15,~: Sophora puryearensis Berry?, pI. 45, fig. 8___________________________________ 98 

Sophora richardsoni Knowlton, pI. 45, fig. 7. _________________________________ 97-98 

Sophora wilcoxiana Berry _ _ __ ________________ ________________ ____ __ ____ _____ 98 

Sphaerites secretani (Heer)lVIeschinellL_______ ______ ______ __________________ 16 

Sphenopteris eocenica Ettingshausen-------------------------c------"--------_ 15,18 

Sphenopteris lakesii Lesquereux ___________________________________________ 1.5, 18-19 

Sphenopteris tnembranacea Lesquereux ____________________________________ ] 5, 18-19 

Sterculia? heterodonta Knowlton, pI. 50, fig. 5; pI. 51, figs. 4, 8 _____________ 116-117 

Sterculia libbeyi Knowlton, pI. 53, fig. 2 ___________________________________ 117-118 

Sterculia modesta Saporta, Lesquereux." _______ ~ ____________________________ 15, Hi 

Sterculia saportanea Knowlton, pI. 53, fig. 6_________________________________ 117 

Sterculia variabilis Saporta_______________________________________ ______ _____ 70 

Styrax? laramiensis Lesquereux, pI. 52, fig. 7 _______________________________ '__ 123 

Styrax ofIicinalis Linnc__ _ _ _ _ __ _ _ _ _ _ _ _ ___ _ _ __ __ _ _ __ _ _ __ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ ] 23 

U 

Ulmus antecedens Lesquereux, pI. 23, fig. 7 ____ ,______________________________ 62 

Ulmus crassifolia NuttalL_ _ _____ _ _ _ _ __ _ _ __ _ _ _ _ _ _ _ _ __ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ 62 

Ulmus? irregular is Lesquereux ______ ~ __ _ ___ ___ _ __ ___ _ __ _ _ __ _ _ __ _ _ __ _ _ __ _ _ __ __ 69 

Ulmus? quercifolia Unger?, pI. 23, fig. 10_____________________________________ 62 

V 

Veatch, A. C., cited ____________________________________________ ' ____________ _ 

Viburnum acerifolium Linne _______________________ '_ __ _ _ _ __ _ _ __ _ _ _ _ _ _ _ _ _ _ _ __ 128 

Viburnum anceps Lesquereux ___________________________________________ ~_ 15,75-76 

Viburnum con tor tum Lesquereux'!, pI. 55, fig. L____________________________ 127 

Viburnum dentatum Linne_ _ _ ____________________ ________________________ __ 81 

\' iburnum dichotomum Lesquereux- ____________________________ : ____ 15,126, ]28-129 

! Viburnum ellipticum Hooker ____________________________________________ :___ 128 

Viburnum goldianum Lesquereux___________________________________________ 129 

Viburnum? heterodontum Knowlton, pI. 55, fig. 2 _________________________ 127-128 

Viburnum lakesii LesquerCllx, pI. 55, fig. 3 __________________________________ 10,128 

Viburnum latanoides Michaux______________________________________________ 81 

Viburnum marginatum ~esquereux.------------------------~------------ 15,81,127 

Viburnum melaenum Knowlton anel Cockerell, pI. 55, fig. 5 ____________ 10,128-129 

i Viburnum platanoides Lesquereux. ______________________________________ .1.5,82,128 

Viburnum pubescens PurslL __ _ _ __ __ _ _ __ _ _ _ _ __ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ 81 

Viburnum richardsoni Knowlton, pI. 52, fig. 8; pI. 53, figs. ], 3, 4, 5; pI. 54, 

figs. 2, 3 _______________________________________________________ 126-127 

Viburnum solitarium LesquereuL _ _______ __________________________________ 129 

Viburnum whymperi? Heel'. _________________________________________ ~ __ 15, 128-129 

Vitis olrika Heer, pI. 49, fig. 6; pI. 50, fig. 4 ______________________________ 10,113,115 

W 

'Yard, L. F., cited __________________________________________________________ _ 

White, C. A., cited __ ~'_______________________________________________________ ],5 

Woodwardia latiloba Lesquereux ____________________________________________ 21-22 

Woodwardia latiloba minor Lesquereux ____________________________________ ]5,21-22 

Woodwardia latiloba serrata Knowlton, pI. 2, figs. 1, 2_______________________ 22 

Z Zingiberites elubius Lesquereux ____ ~_____ ______ ____ ____ __ __ __ __ __ __ __ ______ __ 43 

Zingiberites? undulatus Lesquereux. _________________________________________ 15,4:1 

Zizyphus beckwithii Lesquereux, pI. 47, fig. 3________________________________ 110 

Zizyphus daphnogenoides Knowlton, pI. 48, figs. 5, 6 ______________________ 109-110 

Zizyphus distortus Lesquereux, pI. 4i, fig. 2; pI. 52, fig. L _______ 15,108,109,110-111 

Zizyphus fibrillosus (Lesquereux) Lesquereux, pI. 46, figs. (i, 9_ 9,15,108,109-110,111 

Zizyphus hesperius Knowlton, pI. 49, fig. 2 ________________________________ 111-112 

Zizyphus hyperboreu8? HeeL ____________________________________________ 15,111-112 

Zizyphus lesquereuxii Knowlton_ _ _ _________________________________________ 111 

Zizyphus meeki Lesquereux ___ -_______ -___ __________ ____ ____ ____ ____________ 60 

Zizyphus raincourtii (Saporta) Saporta ____________________________________ ]09-110 

Zizyphus vulgaris Linne _________________________________ ~_ ____ __ ____ ______ __ 109 

o