Revisions of Ulmus and Zelkova in 

the middle and late Tertiary of 

western North America

GEOLOGICAL SURVEY PROFESSIONAL PAPER 1026
Revisions of Ulmus and Zelkova in 

the middle and late Tertiary of 

western North America

By TOSHIMASA TANAI and JACK A. WOLFE

GEOLOGICAL SURVEY PROFESSIONAL PAPER 1026

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1977
UNITED STATES DEPARTMENT OF THE INTERIOR 

CECIL D. ANDRUS, Secretary

GEOLOGICAL SURVEY 

V. E. McKelvey, Director

Library of Congress Cataloging in Publication Data

Tanai, Toshimasa.

Revisions of Ulmus and Zelkova in the middle and late Tertiary of western North America.

(Geological Survey Professional Paper 1026)

Bibliography: p.

Includes index.

1. Elm, Fossil. 2. Zelkova, Fossil. 3. Paleobotany-Tertiary. 4. Paleobotany North America.

I. Wolfe, Jack A., 1936- joint author. II. Title. III. Series: United States. Geological Survey.

Professional paper 1026. 

QE983.T36 561'.3 77-608067

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

Washington, D.C. 20402

Stock Number 024-001-02991-0
CONTENTS

Page

Abstract ..................................................... .................................................................._ 1

Introduction ................................................................................................................................................... 1

Stratigraphic and areal distribution ...................................................................................................... 2

Systematics ..................................................................................................................... .............................. 3

Doubtful species of Ulmus .......................................................................................................................... 9

Rejected citations of Ulmus ........................................................................................................................ 9

References cited ............................................................................................................................................ 10

Index .............................................................................................................................................................. 13

ILLUSTRATIONS

[Plates follow index]

PLATE 1. Leaves of Ulmus pseudo-americana and U. knowltoni.

2. Leaves of Ulmus knowltoni, U. owyheensis, and U. paucidentata.

3. Leaves of Ulmus owyheensis, U. affinis, and U. speciosa.

4. Leaves of Ulmus chaneyi and Zelkova browni.

TABLE

Page

TABLE 1. Stratigraphic ranges of species of Ulmus and Zelkova in the later Oligocene and Neogene of

conterminous Western United States ...................................................................................................................................................... 2

III

REVISIONS OF ULMUS AND ZELKOVA IN THE MIDDLE AND 

LATE TERTIARY OF WESTERN NORTH AMERICA

By TOSHIMASA TANAI 1 and JACK A. WOLFE

ABSTRACT

Examination of previously described and some undescribed 

leaves of Ulmus and Zelkoua from the later Oligocene, Miocene, 

and Pliocene of western North America indicates that at least 

eight species of Ulmus and two species ofZelkova are represented. 

Three new species are described: Ulmus chaneyi, U. knowltoni, 

and Zelkoua browni.

INTRODUCTION

The elms (Ulmus) and species of the related genus 

Zelkova are among the most conspicuous elements of 

temperate broadleaved deciduous forests in the 

Northern Hemisphere. Judging from the fossil record of both leaves and pollen from the Oligocene and 

Neogene, members of these two genera were also 

important constituents of ancient forests. Although 

foliage of the same basic type also occurs abundantly in many Paleocene and Eocene floras (for example, Brown, 1962; Wolfe, 1968; MacGinitie, 1969), 

the differences between these earlier leaves and 

extant leaves of Ulmus and Zelkova are sufficiently 

great in many specimens to make generic assignments problematic (Wolfe, 1968). Certainly elmlike 

foliage and pollen extend back to the latest Cretaceous (Wolfe, 1973), but we have not undertaken the 

extensive work necessary to unravel the systematics 

of these earlier occurrences.

Indeed, a flowering specimen of the Eocene "Zelkova" nervosa (Newb.) R. W. Br. from the Green 

River Formation clearly indicates that this species is 

unassignable to Zelkova. The specimen, which is a 

young shoot, has clusters of conspicuously pedicellate axillary flowers on both upper and lower parts of 

the shoot, and staminate and bisexual flowers occur 

in the same clusters. In Zelkova, the flowers are 

sessile or shortly pedicellate, and the staminate 

flowers are clustered in the lower axils and the 

bisexual flowers are solitary or few in the upper axils. 

Moreover, the shoot itself is conspicuously straight, 

whereas shoots ofZelkova are conspicuously zig-zag 

(see Tanai and Suzuki, 1963, pi. 18, fig. 5). Specimens

1 Department of Geology and Mineralogy, Hokkaido University, Sapporo, Japan.

of shoots of the early Oligocene "Z. " drymeja (Lesq.) 

R. W. Br. are also straight. We conclude that neither 

"Z." nervosa nor "Z." drymeja can be referred to 

Zelkova and that both species represent some extinct 

(although certainly ulmaceous) genus. Also assignable to the same genus are the leaves referred by 

Chancy (1927) to Ulmus brownellii Lesq.

The major concern of this report will thus be the 

later Oligocene, Miocene, and Pliocene species of 

Ulmus and Zelkova from western North America. In 

the process of determining foliage of these genera 

from the Miocene of Alaska (Wolfe and Tanai, 1977), 

it became apparent that the taxonomy of middle Tertiary species of these genera was poorly understood. 

Many epithets have been proposed; however, some 

authors have erected new species on characteristics 

of little systematic value, and other authors, who 

apparently ignored the descriptions of namebearing 

specimens, have assigned material of widely differing morphology to the same species.

Insofar as we have been able to determine, only 

one species ofZelkova Z. browni is represented in 

the Oligocene and Miocene of conterminous United 

States. An additional species the otherwise Eurasian Z, ungeri Kovats is present in the Miocene of 

Alaska (Wolfe and Tanai, 1977) and is not further 

treated in this report. Brown (1937) was the first to 

recognize the presence of Zelkova in the Tertiary of 

North America, although, as discussed below, we do 

not consider the type of his Z. oregoniana to represent Zelkova. Nevertheless, some of the nontypic 

material referred to Z. oregoniana by Brown (1937) 

does indeed represent Zelkova.

In contrast to Zelkova, Ulmus is represented in the 

middle and late Tertiary of western North America 

by at least eight species. Although nine species are 

discussed in the systematic section of this report, 

we consider one of these (U. moragensis} to be of 

doubtful validity, that is, this species is probably 

synonymous with another previously described 

species. Additionally, the material described as U. 

eolaciniata (MacGinitie, 1937) is considered to be
REVISIONS OF ULMUS AND ZELKOVA IN THE TERTIARY OF NORTH AMERICA

Rosaceae and is thus excluded from the enumeration 

of Ulmus. Because of the past confusion in the 

nomenclature and systematics of Tertiary leaves of 

Ulmus, in western North America, the following key 

is presented.

A KEY FOR THE IDENTIFICATION OF ULMUS

LEAVES FROM THE MIDDLE AND

LATE TERTIARY

A. A prominent tertiary vein forking without reaching the sinus bottom and extending along each 

border of sinus.

1. Principal teeth acutely trigonal.

Subsidiary teeth usually 2 and large, 

having additionally 1 minute tooth 

each .................... U. pseudo-americana

2. Principal teeth bluntly deltoid, rather 

apiculate. Subsidiary tooth absent, or 

sometimes 1 minute 

tooth.................................... U. knowltoni

B. A prominent tertiary vein reaching directly the 

bottom of each principal sinus.

1. Principal teeth bluntly deltoid.

(a) Base rounded, slightly 

asymmetrical.

(i) Subsidiary teeth 2, small, and 

nearly equalsized .................. U. speciosa

(ii) Subsidiary teeth 1 or 2, large, 

and sometimes absent on 

the upper 

margin.......... U. owyheensis

(b) Base asymmetrically cuneate.

(i) Foliage shape oblong-elliptic; 

subsidiary teeth 1 or 2, 

sometimes absent on the 

upper margin. 

................... U. paucidentata

(ii) Foliage shape ovate to orbicular; subsidiary tooth 1 and 

small, or absent ...U. moorei

2. Principal teeth acutely trigonal. 

Subsidiary teeth 1 or 2, small 

........................................................ [/. affinis

3. Principal teeth having subsidiary teeth 

on both upper and lower sides 

.................................................... U, chaneyi

We gratefully acknowledge the assistance of H. D. 

MacGinitie and H. E. Schorn (Museum of Paleontology, University of California, Berkeley) for offering their critical comments on this report. For the 

loan of specimens, we also express our gratitude to H. 

F. Becker, J. A. Doyle, L. J. Hickey, and H. E. Schorn.

STRATIGRAPHIC AND AREAL DISTRIBUTION

The two most geologically long-ranging species of 

Ulmus are U. owyheensis and U. speciosa; both 

species are known to range from the middle Oligocene (Bridge Creek age) through the late Miocene 

(table 1). Somewhat shorter ranging are U. paucidentata and U. pseudo-americana, which range from the 

middle Oligocene through the middle Miocene, and 

U. knowltoni, which ranges from the early through 

the late Miocene. Both U. affinis and U. moorei are 

restricted to late Miocene and younger horizons, and 

thus far U. chaneyi is known from only one middle 

Oligocene locality.

TABLE 1. Stratigraphic ranges of species of Ulmus and Zelkova 

in the late Oligocene and Neogene of conterminous Western 

United States

Oligocene

3 

(Q 

O 

0

35'

3

lower upper

Miocene

a 

o

5'

3

lower upper

+

-f----------+----

Homerian

cene

Clam- gulchian

+ .........+......... + ......... + ........+

 +-+

 +

Epoch

00

r» 

01 s

Ulmus affinis 

U. chaneyi

U. knowltoni 

U. moorei 

U. owyheensis 

U. paucidentata 

U. pseudo-americana 

U. speciosa 

Zelkova browni

Ulmus knowltoni has the same peculiar relation 

between the marginal tertiary veins and sinuses as 

does U. pseudo-americana. Although the ranges of 

the two species overlap during the early to middle 

Miocene, a phylogenetic relation could be suggested. 

An undescribed (because of the lack of sufficiently 

complete material) species from the middle Oligocene (Tsadaka flora) of Alaska is similar to C7. 

knowltoni, except that the teeth are pronouncedly 

attenuated as in U. pseudo-americana; this character indicates that probably the lines of U. pseudoamericana and U. knowltoni had diverged by or 

during the middle Oligocene.

As well as having long stratigraphic ranges, 

Ulmus owyheensis and U. speciosa have broad geographic ranges. During the early to middle Miocene, 

U. owyheensis is known from southeastern Oregon 

north to lowland southern Alaska (Cook Inlet region), and C7. speciosa ranged from northeastern 

Nevada north to Alaska. The somewhat stratigraphically shorter ranging U. knowltoni also had a
SYSTEMATICS

Miocene distribution from Nevada north to Alaska, 

as well as occurring in western Montana.

Zelkova browni also has a long stratigraphic 

range: middle Oligocene through late Miocene. As 

with long-ranging species of Ulmus, Zelkova browni 

was widely distributed during the early to middle 

Miocene; occurrences are known from southwestern 

Nevada north to the Cook Inlet region of Alaska and 

east to western Montana. The combination of long 

stratigraphic ranges with wide geographic distribution is, in fact, to be expected. A wide geographic 

range probably indicates a wide tolerance to different environmental (particularly climatic) factors, 

and the persistence through a long period of time 

would also indicate considerable tolerance to changing environmental factors.

SYSTEMATICS

Foliage of Ulmus is highly similar to that of Zelkova, and such similarity has led to confusion in 

generic assignments. We cannot agree with Brown 

(1937, p. 173) that the areoles of Zelkova are relatively larger than those of Ulmus; from examination of 

ultimate venation of cleared leaves of all extant 

species of Zelkova and 25 species of Ulmus, there is 

complete overlap between the two genera in size of 

the areoles. Although most species of Zelkova are 

singly serrate and most species of Ulmus are doubly 

(or more) serrate, some leaves of Zelkova possess 

secondary serrations and some leaves of Ulmus are 

singly serrate. The only feature that consistently 

separates foliage of the two genera is in the intercostal tertiary venation: in Zelkova, the tertiary 

veins are typically thin, widely spaced, and fork conspicuously midway between the secondary veins, in 

contrast to Ulmus in which the tertiary veins are 

thick, more widely spaced, and at least half the 

tertiary veins are unbranched (that is, they are 

percurrent).

One of the major problems in unraveling the 

nomenclature for the various species (or rather species concepts) discussed in this report is that many 

epithets have been founded on poorly preserved or 

fragmentary (or both) material. In the last decade, 

increasing emphasis has been placed on details of 

fine venation and margin for the characterization of 

leaf species, and such details are in many instances 

lacking on name-bearing specimens. Other species 

concepts have been based on one or two fragmentary 

specimens, and, if no topotypic material is known for 

such species, circumscription of that particular 

taxon is uncertain, that is, even certain gross features and their range of variation are unknown in 

regard to certain specific concepts.

An example of one such problem is the proper 

name to be applied to the widespread species previously called Zelkova oregoniana. Brown (1937) based 

this species on Knowlton's (1902) Myrica oregoniana. Our examination of the type specimen has led 

us to conclude that it, in fact, represents a species of 

Ulmus (renamed below as U. knowltoni), although 

certain other material synonymized to Z. oregoniana 

by Brown (1937) and other workers does represent 

Zelkova. The next available name for this species of 

Zelkova would be fernquisti, based on Knowlton's 

(1926) Ulmus fernquisti. Ulmus fernquisti probably 

represents Zelkova, but Knowlton (1926, p. 39) validly remarks that "finer venation not retained." We 

thus reject the epithet fernquisti because it is based 

on a poorly preserved specimen, and no comparisons 

of the finer venation of the type with other fossil or 

extant species will ever be possible. The next available name would apparently be Brown's (1946) Z. 

hesperia. The only specimen illustrated by Brown 

(1946) is by inference the holotype; this specimen is 

very fragmentary, and moreover, the simple line 

drawing published should not be considered an adequate illustration. In addition, the description offered by Brown (1946) is so vague that it could be 

applied to leaves of many unrelated genera. Our 

collections from the topotypic locality do include 

leaves assignable to Z. "oregoniana," but the fragmentary nature of Brown's holotype combined with 

the inadequacy of both illustration and description 

lead us to reject the epithet hesperia. Further, if one 

of Chaney's (1927) specimens, which were placed in 

Z. hesperia by Brown (1946), was selected as the type, 

this species would become a junior synonym of "Z." 

drymeja; none of the specimens illustrated by 

Chancy (1927) represents valid Zelkova. The next 

available name for Z. "oregoniana" would appear to 

be Axelrod's (1956) nevadensis, based on Z. nevadensis. Fully a third of the holotype is lacking, and the 

paratypes are even more fragmentary. The holotype, 

moreover, is strongly asymmetrical at the base, 

which is atypical for Z. "oregoniana " and could thus 

readily be a source for future error. Indeed, it could be 

argued that a name-bearing specimen that further 

work reveals is probably highly atypical for that 

taxon is a monstrosity and can thus be rejected under 

Article 71 of the International Code of Botanical 

Nomenclature. In any case, we consider that the 

fragmentary nature of the type combined with the 

fact that it is atypical for the definition of the species 

to which Z. "oregoniana" has been previously applied make this specimen an unsatisfactory type. We 

have, therefore, proposed a new species Z. browni 

which is based on several complete (or nearly so) and
REVISIONS OF ULMUS AND ZELKOVA IN THE TERTIARY OF NORTH AMERICA

well-preserved specimens from the Collawash locality. We suggest that, as workers pay greater attention to more detailed aspects of foliar morphology, 

more epithets such as fernquisti, hesperia, and nevadensis will (and should) be relegated to the status of 

nomina dubia.

For the occurrences of each species, we have cited, 

where possible, museum catalog numbers pertaining 

to individual specimens. Specimens that have been 

previously cited by other authors and that are here 

considered to be of doubtful systematic status are not 

cited in this report. The following abbreviations 

have been used: USNM = U.S. National Museum, 

UCMP = University of California Museum of Paleontology (Berkeley), UMMP = University of Michigan 

Museum of Paleontology, NYBG = New York Botanical Garden.

Ulmus affinis Lesquereux

Plate 3B, D, E, G

Ulmus affinis Lesquereux, 1878, Harvard Coll. Mus. Comp.

Zoology Mem., v. 6, p. 16, pi. 4, fig. 4 (part). 

Rhamnus troutdalensis Chaney, 1944b, Carnegie Inst. Washington Pub. 553, p. 348, pi. 64, figs. 2, 3.

Ulmus californica Lesquereux, 1878 [part, nontypic]. Harvard

Coll. Mus. Comp. Zoology Mem., v. 6, p. 15, pi. 4, fig. 2 only.

Condit, 1944a, Carnegie Inst. Washington Pub. 553, p. 46,

pi. 8, fig. 4. 

Condit, 1944b, Carnegie Inst. Washington Pub. 553, p. 77,

pi. 18, fig. 6. 

Axelrod, 1944a, Carnegie Inst. Washington Pub. 553, p. 99,

pi. 22, figs. 4, 5.

Chaney, 1944a, Carnegie Inst. Washington Pub. 553, p. 318. 

Chaney, 1944b, Carnegie Inst. Washington Pub. 553, p. 345,

pi. 63, fig. 4. 

Ulmus cf. tanneri auct. non Chaney, Axelrod, 1944b, Carnegie

Inst. Washington Pub. 553, p. 200, pi. 38, fig. 8. 

Zelkova oregoniana auct. non (Knowlton) Brown. [Chaney],

1944b, Carnegie Inst. Washington Pub. 553, p. 346. 

Ulmus moorei Chaney and Elias, Axelrod, 1964, California 

Univ. Pubs. Geol. Sci., v. 51, p. 119, pi. 12, figs. 14, 15.

Supplementary description. Leaves lanceolateoblong in general outline, gradually narrowed and 

slightly acuminate at apex, asymmetrically rotund 

to cordate at base, 4.5-8 cm long and 2-3.8 cm wide; 

midvein thick, nearly straight; lateral veins distinct, 

13-18 subopposite pairs, leaving midrib at angles of 

40°-50° at the middle and upper part of blade and at 

somewhat wider angles near the base, nearly parallel to each other, ending in larger teeth; intercostal 

tertiary veins forming polygonal meshes that include thrice-branching veinlets; a prominent tertiary 

vein near the margin ending in the bottom of each 

principal sinus, margin doubly serrate with large 

and acute teeth, which are accompanied mostly by a 

small subsidiary tooth or rarely by two subsidiary 

teeth on the lower margin; texture thin; petiole rather

slender, more than 1.5 cm long.

Discussion. Despite the previous application of 

the epithet californica to this species of Ulmus, we 

have chosen the epithet affinis over californica for 

the following reasons:

(1) The three type specimens illustrated by Lesquereux (1878) came from different localities; 

the specimens illustrated as plate 4, figure 1 

and plate 6, figure 7a came from the early 

Eocene Chalk Bluffs locality and the specimen illustrated as plate 4, figure 2 came from 

Table Mountain. The Chalk Bluffs specimens clearly represent Chaetoptelea pseudofulva (Lesq.) MacG., whereas only the Table 

Mountain specimen represents valid Ulmus. 

Hence, discordant elements were included in 

Ulmus californica by Lesquereux (1878), and 

thus the name must be rejected "unless it is 

possible to select one of these elements as a 

a satisfactory type." (Article 70 of the International Code of Botanical Nomenclature). 

In this instance, it is possible to select a satisfactory type, but this selection has not previously been made.

(2) Lesquereux (1878, p. 15) describes Ulmus californica as having "borders irregularly denticulate" and secondary veins "more open 

towards the base." In these two features, the 

description clearly applies to the Chalk 

Bluffs Chaetoptelea rather than to the one 

specimen of Ulmus from Table Mountain. 

Thus, designation of the Chalk Bluffs specimen illustrated as plate 4, figure 1 (Lesquereux, 1878) as lectotype is more consistent 

with Lesquereux' intention.

(3) All syntypes of Ulmus affinis are from the 

same locality (Table Mountain), and all represent the same species of Ulmus.

Insofar as we could determine, the type specimens 

of the above-cited taxa are synonymous with Ulmus 

affinis. Leaves described as U. californica from the 

upper Tertiary of California and Oregon (Condit, 

1944b; Chaney, 1944b; Axelrod, 1944a, b, c) are 

particularly valid in specific reference, although 

some of these leaves are more strongly cordate at the 

base and larger than the types of Ulmus affinis.

Ulmus affinis is generally similar to U. speciosa 

Newb. and U. owyheensis Smith, particular to some 

small- and medium-sized leaves. U. affinis is distinguishable in marginal characters from U. speciosa, as well as in the abaxial tertiary veins of the 

basal part of the lamina. Among the extant elms, U. 

americana L. is most similar to U. californica.

Hypotypes: USNM 208512.
SYSTEMATICS

Occurrence.—Table Mountain (USNM 208512; 

UCMP 1917, 2735, 2736, 5735, 5736, 5737), Remington Hill (UCMP 2408, 2409), Sonoma (UCMP 2804), 

and Black Hawk Ranch (UCMP 1743-1749), Calif.; 

Troutdale (UCMP 2601-2604,2616,2617), The Dalles 

(UCMP 2791-2793), and Deschutes, Oreg.; Trapper 

Creek (UCMP 8420-8425), Idaho.

Ulmus chaneyi Tanai and Wolfe, n. sp.

Plate 4B

Description.—Leaves ovate in general outline, 7 

cm wide, to more than 11 cm long; base strongly 

asymmetrical, obliquely cordate; apex known; midvein thick, nearly straight; secondary veins more 

than 14 pairs, subopposite to subalternate, departing 

midrib at angles of 50°-60° one side and 40°-50° 

another side, nearly straight and parallel each other, 

ending in primary teeth; intercostal tertiary veins 

somewhat irregular but convexly percurrent; several 

tertiary veins branching from the secondaries near 

the margin, ending in subsidiary teeth; fourth- and 

fifth-order veins forming polygonal networks that 

enclose fine veinlets that are twice or more branching; margin compoundly serrate, with triangular 

primary teeth on the basal side of which are more 

than four subsidiary teeth, and on the apical side 

typically one or two; petiole strong, 1.5 cm long.

Discussion.—Three incomplete leaves from Twickenham, Oreg., have a characteristic marginal serration that is distinct from other Tertiary elm leaves 

previously described from North America. These 

fossils are closely similar to unlobed leaves of Ulmus 

laciniata Mayer living in northeastern Asia.

Holotype: UCMP 5738

Occurrence.—Twickenham, Oreg.

Ulmus knowltoni Tanai and Wolfe, n. sp.

Plate 1C, F, G; Plate 2A, C, H, I, J

Myrica oregoniana Knowlton, 1902, U.S. Geol. Survey Bull. 204,

p. 33, pi. 3, fig. 4.

Ulmus speciosa auct. non Newberry. LaMotte, 1936 [part], Carnegie Inst. Washington Pub. 455, p. 124. 

Axelrod, 1964, [part], California Univ. Pubs. Geol. Sci., v. 51,

p. 120, pi. 12, fig. 13. 

Graham, 1965, Kent State Univ. Bull. Research ser. 9, p. 97,

pi. 18, fig. 4. 

Ulmus paucidentata H. V. Smith, 1941 [part, nontypic], Am. Mid.

Naturalist, v. 25, p. 512, pi. 13, fig. 4. 

Becker, 1969 [part], Palaeontographica B, v. 127, p. 83, pi. 21,

fig. 7. 

Zelkoua oregoniana auct. non (Knowlton) Brown. Chaney and

Axelrod, 1959 [part], Carnegie Inst. Washington Pub. 617,

p. 31, pi. 31, figs. 5, 8. 

Ulmus newberryi auct. non Knowlton. Graham, 1965, Kent State

Univ. Bull. Research ser. 9, p. 96, pi. 18, fig. 3.

Description.—Leaves highly variable in shape

and size, oblong to lanceolate-oblong in general 

outline, 3-15.5 cm (estimated) long and 1.4-6 cm 

wide; base strongly asymmetrical, obliquely cordate, 

nearly straight on one side and bulged on the other 

side into a typically half-circular shape; apex gradually narrowed, attenuate-acuminate; midrib thick, 

nearly straight; secondary veins opposite to subopposite, 12-20 pairs, diverging at different angles 

on both sides of midrib, nearly straight or slightly 

curving up, rarely forking in the lower part of blade, 

ending in marginal teeth; several prominent 

branches from a basal secondary vein on the bulged 

part diverging outward, ending in marginal teeth; 

tertiary veins in the intercostal area wavy but 

distinct, irregularly percurrent; fourth-order veins 

thin, forming quadrangular or pentagonal meshes 

that include twice- or thrice- branching veinlets; one 

prominent tertiary vein departing from the secondaries near the margin, forking near the bottom of the 

principal sinus, then extending along the margin; 

margin mostly singly serrate with large apiculate 

teeth that are rarely associated with a minute subsidiary tooth on the lower part of blade; texture thin; 

petiole thick, more than 1 cm long.

Discussion.—The above description is primarily 

based on a number of well-preserved leaves from the 

Miocene Collawash locality, Oregon. Leaves of this 

species are usually characterized by the principal 

apiculate teeth, strongly asymmetrical base, and 

prominent tertiary veins forking near each principal 

sinus. The leaves having a simply serrate margin 

could be misidentified as Zelkova in some instances 

but are certainly distinguishable in the marginal serration and venation character of the basal part of the 

lamina.

The specimens of Myrica oregoniana Knowlt. from 

the Mascall flora (USNM 8532 a, b) were transferred 

to Zelkoua by Brown (1937) and are the type specimens of Z. oregoniana. These specimens are closely 

similar to Zelkova leaves having an asymmetrical 

base, which type is sometimes found among the 

abnormal leaves of the extant Z. serrata Makino 

(especially on the young shoots). However, these 

Mascall specimens are quite different in marginal 

serration and tertiary venation of the enlarged part 

of the lamina and are included in U. knowltoni as 

here established. Although it might seem proper to 

use the epithet "oregoniana" for this new species, 

this epithet is already occupied by another elm 

species from the Ashland Eocene (Knowlton, 1900), 

which is of somewhat doubtful generic status (Wolfe, 

1968).

Holotype: USNM 208502.

Paratypes: USNM 208503-208507.
REVISIONS OF ULMUS AND ZELKOVA IN THE TERTIARY OF NORTH AMERICA

Occurrence.—Mascall (UCMP 3095; USNM 8532, 

a, b), Rockville (UMMP 44947, 44950), Collawash, 

Oreg.; Thorn Creek and Trapper Creek (UCMP 8426), 

Idaho '49 Camp, Nevada (UCMP 797,3099); Seldovia 

Point, Alaska; Beaverhead, Mont. (NYBG 601a).

Ulmus moorei Chaney and Elias

Ulmus moorei Chaney and Elias, 1936, Carnegie Inst. Washington Pub. 476, p. 39, pi. 6, figs. 1-5. 

Dorf, 1936, Carnegie Inst. Washington Pub. 476, p. 116,

pi. 2, fig. 4. 

Axelrod, 1956, California Univ. Pubs. Geol. Sci., v. 33, p. 292,

pi. 8, figs. 5, 6, 12. 

Smiley, 1963, California Univ. Pubs. Geol. Sci., v. 35, no. 3,

p. 218, pi. 8, fig. 6b. 

?Becker, 1972, Palaeontographica B, v. 141, p. 35, pi. 12,

figs. 12, 13. 

Ulmus venustula Brown, 1949, Washington Acad. Sci. Jour., v. 39,

p. 226, figs. 3-5, 9.

Zelkova nevadensis auct. non Axelrod. Smiley, 1963, California 

Univ. Pubs. Geol. Sci., v. 35, p. 219, pi. 8, figs. 2, 6a.

Discussion.—This species is represented mostly by 

small leaves that are ovate to orbicular in shape; the 

base is slightly asymmetrical and broadly cuneate to 

slightly cordate. The margin is mostly simply serrate 

and has large, blunt teeth that may have a minute 

subsidiary tooth. U. moorei closely resembles leaves 

of the extant U. crassifolia Nutt. of eastern United 

States.

As cited above, U. moorei is known in the upper 

Miocene and Pliocene of California, Washington, 

and the High Plains, although the specimens are not 

common everywhere. A single leaf and its counterpart figured as U. moorei from the Metzel Ranch 

locality, southwestern Montana (Becker, 1972), are 

similar in general outline to some of the original 

specimens, but the teeth of Becker's specimen are 

more obtuse. A small leaf figured as Zelkova nevadensis Axelr. from the Ellensberg flora, Washington 

(Smiley, 1963), has serrate teeth on the basal margin, 

and several tertiary veins from the basal secondaries 

end in these teeth; this specimen is included in U. 

moorei.

Occurrence.—Beaver County, Okla. (UCMP 5333, 

5335-5339, 5350-5353); Logan County, Kans.; Aidrich Station, Nev.; Ellensberg, Washington; (?)Beaverhead basin, Mont.; Weiser, Idaho (UCMP 1202, 

1203); Cache Valley, Utah.

Ulmus owyheensis H. V. Smith

Plate 2B, D, E, F; plate 3A

Ulmus owyheensis H. V. Smith, 1939, Michigan Acad. Sci. Papers,

v. 24, p. 113, pi. 6, fig. 4. 

Graham, 1965, Kent State Univ. Bull. Research ser. 9, p. 97,

pi. 18, fig. 1. 

Ulmus speciosa auct. non Newberry. Chaney and Axelrod, 1959,

Carnegie Inst. Washington Pub. 617, p. 174, pi. 32, figs.

1-3 (part). 

Ulmus paucidentata auct. non Smith. Chaney and Axelrod, 1959,

Carnegie Inst. Washington Pub. 617, p. 173 (part). 

Smiley, 1963, California Univ. Pubs. Geol. Sci., v. 35, p. 219,

pi. 8, figs. 4, 5. 

Ulmus plurinervis auct. non Unger. Heer, 1869, Flora Fossilis

Arctica, v. 2, pt. 2, p. 34, pi. 5, fig. 1.

Discussion.—This species was established for a 

single, nearly complete leaf from Sucker Creek of 

Oregon (Smith, 1939a). The species is characterized 

by an ovate shape, slightly asymmetrically rounded 

base, simply serrate teeth, and the lowest secondary 

pair having 3 or 4 abaxial tertiary veins that end in 

small teeth on the basal margin. Several specimens 

collected from the Eagle Creek and Bridge Creek 

(Twickenham) localities add the following characters to this species: principal teeth are rather bluntly 

trigonal and sometimes accompanied by one or two 

small subsidiary teeth, especially below the middle 

of blade; foliar shape is somewhat variable from 

ovate to oval; base is rotund to slightly cordate, 

nearly symmetrical to slightly asymmetrical; a 

prominent tertiary vein near the margin ends in the 

bottom of each principal sinus; intercostal tertiary 

veins irregularly crossed with the secondaries and 

never percurrent. Ulmus owyheensis is not similar to 

any living species of North American elms but seems 

rather to be related to the extant U. pumila L. of east 

Asia.

Several specimens figured as Ulmus speciosa 

Newb. from the Mascall and Stinking Water floras of 

Oregon (Chaney and Axelrod, 1959) are included in 

U. owyheensis by the above-noted characters. 

Although not figured by Chaney and Axelrod, a 

specimen (UCMP 3091) identified as U.paucidentata 

from the Mascall is also referred to U. owyheensis. A 

single incomplete leaf of U. cf. tanneri Chaney from 

the Pliocene Sonoma flora of California (Axelrod, 

1950) was later referred to U. owyheensis in 

marginal serration and basal character.

Hypotypes: USNM 208508-208511; UCMP 5733.

Occurrence.—Sucker Creek (UMMP 20017), Mascall (UCMP 3080, 3082, 3083, 3091), Stinking Water 

(UCMP 3084), Collawash, Fish Creek, Eagle Creek, 

Twickenham,,Oreg.; Seldovia Point, Alaska; Ellensburg, Wash. (UCMP 5346, 5349).

Ulmus paucidentata H. V. Smith

Plate 2G

Ulmus paucidentata H. V. Smith, 1939, Torrey Bot. Club Bull.,

v. 66, p. 478, pi. 13, figs. 2-4, 

Smith, 1941 [part], Am. Midland Naturalist, v. 25, p. 512,

pi. 10, fig. 2; pi. 13, fig. 9. 

Chaney and Axelrod, 1959 [part], Carnegie Inst. Washington

Pub. 617, p. 173, pi. 31, figs. 9-12.
SYSTEMATICS

Seeker, 1969, Palaeontographica B, v. 127, p. 83, pi. 21,

figs. 4-6 (part). 

Ulmus speciosa auct. non Newberry. Newberry, 1898 [part, nontypic], U.S. Geol. Survey Mon. 35, p. 80, pi. 45, figs. 5, 8. 

Dorf, 1936 [part], Carnegie Inst. Washington Pub. 476, p. 116. 

Becker, 1961, Geol. Soc. America Mem. 82, p. 64, fig. 4. 

Axelrod, 1964 [part], California Univ. Pubs. Geol. Sci., v. 51,

p. 120.

Discussion.—Leaves of this species are typically 

small and elliptic to ovate-elliptical; on the margin 

the primary teeth are large and obtusely trigonal, in 

most instances accompanied by one or two small 

subsidiary teeth. The base is typically asymmetrically cuneate or obtuse; narrowly cuneate or obliquely straight on one side and rounded on the other. The 

intercostal tertiary veins are irregularly but prominently percurrent, and a tertiary vein from the 

secondaries near the margin ends in the bottom of 

the principal sinus.

Ulmus paucidentata closely resembles U. owyheensis and C7. knowltoni in marginal characters, 

especially in having large blunt teeth that may 

resemble a simply serrate margin. However, U. paucidentata is distinguishable from these two species 

in the basal shape and character of the tertiary venation character of the basal part, without any intergradation of characters. Considering the foliar 

shape and marginal serration, U. paucidentata may 

be related to the extant U. parvifolia Jacq. living in 

Japan and China, but there are some differences in 

the percurrent tertiary venation.

Hypotype: UCMP 5734.

Occurrence.—Mascall (UCMP 3086, 3088, 3089, 

3090), Bridge Creek (USNM 7064, 201872), Twickenham, Oreg. (UCMP 5734); Thorn Creek (UMMP 

20016), Trapper Creek (UCMP 8427), Weiser, Idaho 

(UCMP 1204); Ruby basin (UMMP 38249), Beaverhead basin (NYBG 602a, 603, 604), Montana.

Ulmus pseudo-americana Lesquereux

Plate 1A, B, D, E, H

Ulmus pseudo-americana Lesquereux, 1883, U.S. Geol. Survey

Terr. Kept., v. 8, p. 249, pi. 54, fig. 10. 

Ulmus speciosa Newberry, 1898 [part, typic], U.S. Geol. Survey

Mon. 35, p. 80, pi. 45, fig. 2.

Knowlton, 1902, U.S. Geol. Survey Bull. 204, p. 53 (part). 

Chancy and Axelrod, 1959 [part], Carnegie Inst. Washington

Pub. 617, p. 174, pi. 32, fig. 5. 

Axelrod, 1964, California Univ. Pubs. Geol. Sci., v. 51, p. 120

(part). 

Ulmus newberryi Knowlton, 1902 [part], U.S. Geol. Survey Bull.

204, p. 54, pi. 9, fig. 4.

Wolfe, 1964, U.S. Geol. Survey Prof. Paper 454-N, p. N22, 

pi. 3, figs. 4, 6.

Discussion.—The larger leaves of elm from the 

middle Tertiary have usually been referred to Ulmus 

newberryi Knowlt., U.pseudo-americanaljesq.,OY U.

speciosa Newb.; most of these specimens have been 

compared to extant U. americana L. by many authors. These three species have, however, been frequently confused, although the criteria for specific 

distinction have been discussed by various authors. 

For example, LaMotte (1952) placed U. newberryiin 

synonymy under U. speciosa, whereas Knowlton 

(1902) and Chancy and Axelrod (1959) separated 

these two species and included U. pseudo-americana 

in U. speciosa. One of the reasons for such confusion 

is that foliar shape and size are not consistently 

valid criteria for the discrimination of these elm 

species, although U. newberryi is generally more 

slender in shape and more oblique (strongly asymmetric) at the base than U. speciosa. Another reason 

for confusion is that most original specimens from 

the Oligocene or Miocene of Oregon were not always 

correctly illustrated by Newberry (1898) and Knowlton (1902).

From examination of large suites of these larger 

elm leaves, including name-bearing specimens, we 

find that these specimens include divergent 

morphologic characters in marginal serration and 

venation. These leaves typically have a doubly 

serrate margin, as noted by various authors, but are 

separable into three types. The first type has 

somewhat longer primary teeth, between which 

there are typically two large subsidiary teeth, 

accompanied in many instances by a single minute 

tooth (Lesquereux, 1883, pi. 54, fig. 10; Knowlton, 

1902, pi. 9, fig. 4). The second type typically has two 

small teeth (rarely three) of nearly equal size between 

the primary apiculate teeth that the secondary veins 

enter (Newberry, 1898, pi. 45, fig. 2; Knowlton, 1926, 

pi. 18, fig. 4). The third type has blunt dentate 

primary teeth, between which there is typically one 

blunt tooth, accompanied rarely by one minute tooth 

(Newberry, 1898, pi. 45, figs. 5, 8). In leaves of the 

second and third types, one of prominent tertiary 

veins that departs from the secondaries near the 

margin ends in or near the principal sinus; in the 

first type the tertiary vein never reaches the sinus 

but forks and extends along the margin of sinus. 

Generally leaves of the first type are linear-oblong, 

leaves of the second type are lanceolate. Although 

Chancy and Axelrod (1959) relied on foliar shape and 

basal form, these characters may intergrade 

between these three types.

Judging from the original description and the 

above-noted characters, the first morphologic type 

contains the nomenclatorial type of U. pseudoamericana and some of the nomenclatorial types of 

U. newberryi; the second morphologic type contains 

the type specimens of U. speciosa and U. newberryi.
8 REVISIONS OF ULMUS AND ZELKOVA IN THE TERTIARY OF NORTH AMERICA

The third morphologic type is included in the U. 

paucidentata H. V. Smith as discussed above. Thus, 

owing to priority, we reject U. newberryi and we 

retain U. pseudo-americana for leaves having the 

characters of the first morphologic type.

Hypotypes: UCMP 5731, 5732.

Occurrence.—Bridge Creek (UCMP 1758; USNM 

7065, 9217, 9367, 8493a), Twickenham (UCMP 5731, 

5732), Mascall (UCMP 3735), Oreg.; Fingerrock 

(USNM 41942; UCMP 8642, 8643), Nev.

Ulmus speciosa Newberry

Plate 3C, F

Ulmus speciosa Newberry, 1898 [part, typic], U.S. Geol. Survey

Mon. 35, p. 80, pi. 45, figs. 3, 4.

Knowlton, 1902, [part], U.S. Geol Survey Bull. 204, p. 53. 

Chaney, 1920, Walker Mus. Contr., v. 2, p. 171. 

Knowlton, 1926, U.S. Geol. Survey Prof. Paper 140, p. 39,

pi. 18, fig. 4. 

LaMotte, 1936 [part], Carnegie Inst. Washington Pub. 455,

p. 124. 

MacGinitie, 1962, California Univ. Pubs. Geol. Sci., v. 35,

p. 110, pi. 4, figs. 1, 4, 5; pi. 10, fig. 4. 

Becker, 1969, Palaeontographica B, v. 127, p. 84, pi. 21,

figs. 1, 2. 

Ulmus tanneri Chaney, 1920, Walker Mus. Contr., v. 2, p. 172,

pi. 15, figs. 1, 2. 

Ulmus newberryi auct. non Knowlton. Smiley 1963, California

Univ. Pubs. Geol. Sci., v. 35, p. 218, pi. 8, fig. 3.

Discussion.—This species has been discussed in 

detail in connection with Ulmus pseudo-americana 

Lesq. U. speciosa is distinguishable in marginal serration and venation characters from other elm species of the middle Tertiary of North America. The 

name has been misused by various authors for oval 

leaves that have a less asymmetrical base. We here 

designate as the type specimen of U. speciosa USNM 

7067, which is one of the original specimens figured 

from the Bridge Creek locality by Newberry (1898). 

Ulmus speciosa as here circumscribed is generally 

ovate to oblong in outline and is less asymmetrical 

and rounded in the base (as many authors have 

pointed out). Because some leaves of U. pseudoamericana have a less asymmetrical base, these two 

species are not consistently distinguishable only by 

foliar shape and basal form.

On the basis of ovate-oblong and small leaves from 

the Eagle Creek locality, Ulmus tanneri Chan. was 

separated from U. speciosa (Chaney, 1920). Unfortunately, these type specimens of U. tanneri were 

lost after being moved to the University of California 

at Berkeley from Chicago, and we cannot definitely 

determine the status of this species. However, judging from the original illustration and description, U. 

tanneri seems to represent only the ovate-oblong 

leaves of U. speciosa.

Lectotype: USNM 7067.

Hypotype: USNM 208513

Occurrence.—Bridge Creek (USNM 7066, 7067), 

Eagle Creek,' Oreg.; Latah, Ellensberg (UCMP 5340), 

Wash.; Kilgore (UCMP P-746, P-808), Nebr.; Upper 

Cedarville (UCMP 836), Calif.; Beaverhead (NYBG 

605, 844), Mont.; Seldovia Point, Wrangell Mts., 

Healy Creek (USNM 208513), Alaska.

Zelkova browni Tanai and Wolfe, n. sp.

Plate 4A, C-G

Zelkova oregoniana auct. non (Knowlton) Brown. Brown, 1937

[part], U.S. Geol. Survey Prof. Paper 186, p. 173, pi. 51,

figs. 11-15. 

Lakhanpal, 1958, California Univ. Pubs. Geol. Sci., v. 35,

no. 1, p. 29. 

Chaney and Axelrod, 1959 [part], Carnegie Inst. Washington

Pub. 617, p. 174, pi. 31, figs. 6, 7. 

Becker, 1961, Geol. Soc. America Mem. 82, p. 64, pi. 18, figs.

13-18. 

Smiley, 1963, California Univ. Pubs. Geol. Sci., v. 35, p. 219,

pi. 8, fig. 1. 

Wolfe, 1964, U.S. Geol. Survey Prof. Paper 454-N, p. N23, pi. 3,

figs. 2, 3, 5. 

Axelrod, 1964, California Univ. Pubs. Geol. Sci., V. 51, p. 120,

pi. 12, figs. 8, 10-12. 

Graham, 1965, Kent State Univ. Bull. Research ser. 9, p. 97,

pi. 18, fig. 2. 

Becker, 1969, Palaeontographica B, v. 127, p. 84, pi. 21, figs.

14-21.

Becker, 1972, Palaeontographica B, v. 141, p. 36, pi. 6, figs. 1-9. 

Zelkova ungeri auct. non Kovats. Becker, 1969, Palaeontographica B, v. 127, p. 85, pi. 21, figs, lla, b. 

Fagopsis longifolia auct. non (Lesquereux) Hollick. Berry, 1929,

U.S. Geol. Survey Prof. Paper 154, p. 245, pi. 50, fig. 7. 

Quercus mccanni auct. non Berry. Axelrod, 1964, California Univ.

Pubs. Geol. Sci., v. 51, p. 118, pi. 11, figs. 10, 11. 

? Ulmus fernquisti Knowlton, 1926, U.S. Geol. Survey Prof. Paper

140, p. 39, pi. 19, fig. 2. 

?Zelkova nevadensis Axelrod, 1956, California Univ. Pubs. Geol.

Sci. v. 33, p. 292, pi. 8, figs. 3, 8. 

Smiley, 1963, California Univ. Pubs. Geol. Sci., v. 35, no. 3,

p. 219, pi. 8, figs. 2, 6a.

Description.—Leaves variable in shape and size, 

mostly lanceolate-oblong to oblong, rarely linearoblong or ovate, 2.4-10.5 cm long and 1.7-4.9 cm 

wide; base nearly symmetrical to asymmetrical, 

obtuse to rotund, frequently slightly cordate; apex 

gradually narrowed, attenuate-acuminate, some 

slightly tapered; midvein thick, nearly straight; 

secondary veins 9-17 pairs, opposite to subopposite, 

angles of divergence variable, nearly straight or 

gently curving apically, very rarely forking on the 

way, ending in marginal teeth; intercostal tertiary 

veins wavy or forking but crossing to the secondaries; the fourth- and fifth-order veins forming irregular quadrangular or pentagonal areoles, which 

include veinlets that branch twice or more; margin
REJECTED CITATIONS OF ULMUS

mostly singly dentate with large apiculate teeth, and 

sinus usually widely opened; large teeth rarely accompanied by a minute subsidiary tooth, or in some 

instance an additional tooth between two principal 

teeth on the lower half of the margin; texture firm; 

petiole stout, more than 3 mm long.

Discussion.—Axelrod (1956) distinguished Z. 

nevadensis from Z. oregoniana because of size differences, but his small leaves of Zelkova fall well 

within the variation of size and shape displayed by 

leaves of Z. browni from the Collawash locality. 

Chancy and Axelrod (1959) transferred Castanea 

atavia Ung. (Lesquereux, 1883, pi. 52, fig. 2) and 

Quercus horniana Lesq. (Knowlton, 1902, pi. 8, fig. 1) 

to the genus Zelkova, but these two fossils do not 

have the marginal serrations of Z. browni.

Zelkova browni, though highly variable in foliar 

shape and size, typically has large, blunt teeth and is 

easily distinguishable from Z. ungeri Kovats, which 

has been invalidly reported from the Tertiary of 

Western United States, excluding Alaska. Though 

many authors have compared Z. oregoniana with the 

extant Z. serrata Mak. of Japan or Z. sinica Schn. of 

China, Z. browni is rather related to the extant Z. 

carpinifolia Spach. of the Caucasus region in the 

shape of the teeth. Many specimens of Z. browni 

have a minute subsidiary tooth on the basal side of 

the primary teeth, a feature uncommon in extant 

Zelkova.

Holotype: USNM 208514

Paratypes:USNU 208515-208518.

DOUBTFUL SPECIES OF ULMUS

Ulmus moragensis Azelrod

Ulmus moragensis Axelrod, 1944, Carnegie Inst. Washington 

Pub. 553, p. 283, pi. 48, figs. 5, 7-9.

Discussion.—This species, established on material 

from the middle Pliocene of California (Axelrod, 

1944c), is not based on well-preserved specimens; 

they are particularly poorly preserved in marginal 

character. Chaney (1944b, p. 346) emphasized the 

smaller average leaf size for separating this species 

from other Tertiary elms, as did Axelrod (1944c) in 

his original description. As far as we can determine 

from the type specimens of U. moragensis, this 

species seems to be characterized by a strongly 

asymmetrical base and bluntly trigonal teeth that 

have in some instances a small subsidiary tooth; this 

species thus is difficult to distinguish from some 

leaves of U. affinis. Furthermore, owing to the poor 

preservation of the type specimens, U. moragensis 

may be better considered a nomen dubium.

Occurrence.—Alturas, Calif.

REJECTED CITATIONS OF ULMUS

Ulmus borealis auct. non Heer. Hollick, 1936, U.S. 

Geol. Survey Prof. Paper 182, p. 106, pi. 57, figs. 1,2 

= Alnus adumbrata (Holl.) Wolfe (see Wolfe, 1966, 

p. B21).

Ulmus braunii auct. non Heer. Hollick, 1936 [part], 

U.S. Geol. Survey Prof. Paper 182, p. 105, pi. 58, 

figs. 6,12 = Alnus sp. indet. This material probably 

represents A. evidens (Holl.) Wolfe, but it is too 

fragmentary for specific determination. The other 

material assigned to U. braunii by Hollick (1936) is 

of pre-Oligocene age and is thus not further considered.

Ulmus brownellii auct. non Lesquereux. Dorf. 1930, 

Carnegie Inst. Washington Pub. 412, p. 92, pi. 10, 

figs. 1-3, 10 = genus and species indetermined. 

Although Axelrod (1944c, p. 283) assigned these 

leaves to U. moragensis, we consider that the 

preservation is too poor to permit determination.

Ulmus californica Lesquereux, 1878 [part, typic], 

Harvard Coll. Mus. Comp. Zoology Mem., v. 6, p. 

15, pi. 4, fig. 1, pi. 6, fig. 7a = Chaetoptelea pseudofulva (Lesquereux) MacGinitie, 1941, Carnegie 

Inst. Washington Pub. 534, p. 108. Above, we 

designated a lectotype for Ulmus californica that 

is clearly Chaetoptelea. Thus, it would be possible 

for someone at a later date to consider californica 

as a senior synonym of pseudofulva if one epithet 

were not chosen in preference to the other at this 

time.

Ulmus carpinoides auct. non Goeppert. Hollick, 1936, 

U.S. Geol. Survey Prof. Paper 182, p. 105, pi. 51, 

figs. 1, 2 = Alnus evidens (Holl.) Wolfe, 1966, U.S. 

Geol. Survey Prof. Paper 398-B, p. B18.

Ulmus diptera auct. non Steenstrup. Hollick, 1936, 

U.S. Geol. Survey Prof. Paper 182, p. 107, pi. 48, 

fig. 4a = Alnus evidens (Holl.). Wolfe, 1966 U.S. 

Geol. Survey Prof. Paper 398-B, p. B18.

Ulmus eolaciniate MacGinitie, 1937, Carnegie Inst. 

Washington Pub. 465, p. 126, pi. 5, fig. 3 = Rubus 

eolaciniata (MacG.) Tanai and Wolfe, n. comb. The 

sharply acute teeth that are irregualrly spaced and 

the tertiary veins that are conspicuously not perpendicular to the secondary veins exclude this 

species from Ulmus. On the other hand, these 

features are typically found in the rosaceous 

Rubus. The asymmetry of MacGinitie's material is 

interpreted as indicative of a leaflet; this character 

would also fit with an assignment to Rubus.

Ulmus montanensis Becker, 1961, Geol. Soc. America Mem. 82, p. 63, pi. 18, fig. 5 = U. tenuinervis 

Lesquereux, 1883, U.S. Geol. Survey Terr. Kept., 

v. 8, p. 188. Comparison of Becker's specimen with
10 REVISIONS OF ULMUS AND ZELKOVA IN THE TERTIARY OF NORTH AMERICA

a suite of U. tenuinervis from Florissant indicates 

that some leaves of U. tenuinervis have the characters that Becker (1961) thought characterized his 

U. montanensis and distinguished it from U. 

tenuinervis.

Ulmus paucidentata auct. non H. V. Smith. Becker, 

1961, Geol. Soc. America Mem. 82, p. 63, pi. 18, 

fig. 3 = genus and species indetermined. The specimen has glandular teeth and is hence clearly not 

Ulmus. Otherwise, the specimen is poorly preserved, and we doubt if a reliable assignment can 

be made.

Ulmus pseudofulva Lesquereux, 1878, Harvard Coll. 

Mus. Comp. Zoology Mem., v. 6, no. 2, p. 16, pi. 4, 

fig. 3 = Chaetoptelea pseudofulva (Lesquereux) 

MacGinitie, 1941, Carnegie Inst. Washington Pub. 

534, p. 108. LaMotte (1952) transferred MacGinitie's (1941) combination back to Ulmus, and this 

transfer still remains in effect. Although some 

systematists consider the extant Chaetoptelea to 

be a member of Ulmus, other systematists maintain these genera as distinct. Because the foliage 

of Chaetoptelea can be distinguished from that of 

Ulmus, we prefer to recognize the combination 

proposed by MacGinitie (1941).

Ulmus sorbifolia, auct. non Goeppert. Hollick, 1936 

[part], U.S. Geol. Survey Prof. Paper 182, p. 106, 

pi. 57, figs. 4, 5 = Spiraea hopkinsi Wolfe (see 

Wolfe, 1966, p. B24). Although Spiraea hopkinsi is 

being transferred to Sorbaria (Wolfe and Tanai, 

1977), it would confuse nomenclature to introduce 

the new combination in the current report.

Ulmus speciosa auct. non Newberry. Axelrod, 1964 

[part], California Univ. Pubs. Geol. Sci., v. 51, 

p. 120 [unfigured specimen registered as UCMP 

8463] = Betula ashleyi Axelrod, 1964, California 

Univ. Pubs. Geol. Sci, v. 51, p. 116.

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553, p. 167-206, pis. 34-39.

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Survey Prof. Paper 182, 185 p., 122 pis.

Knowlton, F. H., 1900, Fossil plants associated with the lavas of 

the Cascade Range: U.S. Geol. Survey 20th Ann. Rept., p. 37- 

52, pis. 1-6.

 1902, Fossil flora on the John Day Basin, Oregon: U.S. 

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140-A, p. A17-A81, pis. 8-31. 

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LaMotte, R. S., 1936, The Upper Cedarville flora of northwestern

Nevada and adjacent California: Carnegie Inst. Washington

Pub. 455, p. 59-142, pis. 1-14. 

1952, Catalogue of the Cenozoic plants of North America

through 1950: Geol. Soc. America Mem. 51, 381 p. 

Lesquereux, L., 1878, Report on the fossil plants of auriferous 

gravel deposits of the Sierra Nevada: Harvard Coll. Mus. 

Comp. Zoology, v. 6, p. 1-62, pis. 1-10. 

1883, Cretaceous and Tertiary flora of the western Territories: U.S. Geol. Survey Terr., v. 8, 283 p., 55 pis. 

MacGinitie, H. D., 1937, The flora of the Weaverville beds of 

Trinity County, California: Carnegie Inst. Washington Pub. 

465, p. 83-151, pis. 1-15.
REFERENCES CITED 11

—1941, A Middle Eocene flora from the central Sierra Nevada: 

Carnegie Inst. Washington Pub. 534, 178 p., 47 pis.

—1962, The Kilgore flora. A late Miocene flora from northern 

Nebraska: California Univ. Pubs. Geol. Sci., v. 35, p. 67-158, 

pis. 1-16. 

-1969, The Eocene Green River flora of northwestern Colorado and northeastern Utah: California Univ. Pubs. Geol.

Sci., v. 83, 140 p., 31 pis. 

Newberry, J. S., 1898, The later extinct floras of North America;

a posthumous work edited by Arthur Hollick: U.S. Geol.

Survey Mon. 35, 295 p., 68 pis.

Smiley, C. J., 1963, The Ellensberg flora of Washington: California Univ. Pubs. Geol. Sci., v. 35, p. 159-276, pis. 1-17. 

Smith, H. V., 1939a, Additions to the fossil flora of Sucker Creek,

Oregon: Michigan Acad. Sci., Arts and Letters Papers, v. 24,

p. 107-120, 7 pis. 

———1939b, A flora of eastern American aspect in the Miocene of

Idaho: Torrey Bot. Club Bull., v. 66, p. 465-481, pis. 10-13.

———1941, A Miocene flora from Thorn Creek, Idaho: Am. Midland Naturalist, v. 25, p. 473-522, 14 pis.

Tanai, Toshimasa, and Suzuki, Nobuo, 1963, Miocene floras of 

southwestern Hokkaido, Japan: Tertiary Floras Japan, Vol. I, 

Miocene Floras, Japan Geol. Survey, p. 9-149, 27 pis.

Wolfe, J. A., 1964, Miocene floras from Fingerrock Wash, southwestern Nevada: U.S. Geol. Survey Prof. Paper 454-N, 36 p., 

12 pis.

———1966, Tertiary plants from the Cook Inlet region, Alaska: 

U.S. Geol. Survey Prof. Paper 398-B, p. B1-B32, 8 pis.

———1968, Paleogene biostratigraphy of nonmarine rocks in 

King County, Washington: U.S. Geol. Survey Prof. Paper 571, 

33 p., 7 pis.

———1973, Fossil forms of Amentiferae: Brittonia, v. 25, p. 334-

355.

Wolfe, J. A., and Tanai, Toshimasa, 1977, The Middle Miocene 

Seldovia Point flora from the Kenai Group, Alaska: U.S. 

Geol. Survey Prof. Paper (in press).

INDEX

[Italic page numbers indicate both major references and descriptions]

Page

A

adumbrata, Alnus .............................................................. 9

affinis, Utmus ....................................2, 4; pi. 3B, D, E, G

Aldrich Station, Nev .......................................................... 6

Alnus adumbrata ................................................................ 9

evidens............................................................................ 9

sp ...................................................................................... 9

Alturas, Calif........................................................................ 9

americana, Ulmus .......................................................... 4, 7

ashleyi, Betula .................................................................... 10

atavia, Costonco.................................................................. 9

B

Beaver County, Okla.......................................................... 6

Beaverhead, Mont .......................................................... 6, 8

Beaverhead basin, Mont .............................................. 6, 7

Betula ashleyi ................................................................... 10

Black Hawk Ranch, Calif ................................................ 5

borealis, Ulmus .................................................................... 9

braunii, Ulmus .................................................................... 9

Bridge Creek, Oreg....................................................._6, 7, 8

Brown, R. W., cited ........................................................ 1, 3

brownellii, Ulmus........ ................................................... 1, 9

browni, Zelkova.................................. 1, 3, 8; pi. 4A, C-G

Cache Valley, Utah ............................................................ 6

californica, Ulmus .......................................................... 4, 9

carpinifolia, Zelkova .......................................................... 9

carpinoides, Ulmus ............................................................ 9

Castanea atavia .................................................................. 9

Chaetoptelea.................................................................... 9, 10

Chalk Bluffs.................................................................. 4

pseudofulva.......... ................................................4, 9, 10

Chalk Bluffs.......................................................................... 4

chaneyi, Ulmus .................................................. 2, 5; pi. 4B

Collawash, Oreg.................................................................. 6

Collawash locality .............................................................. 4

Cook Inlet region, Alaska ................................................ 2

crassifolia, Ulmus .............................................................. 6

D, E

Deschutes, Oreg .................................................................. 5

diptera, Ulmus...................................................................... 9

drymeja, Zelkova ............................................................ 1, 3

Eagle Creek, Oreg .......................................................... 6, 8

Ellensberg, Wash ............................................................ 6, 8

Ellensberg flora, Wash...................................................... 6

Elm, Ashland, Eocene ...................................................... 5

Elm leaves, morphologic characters, divergent........ 7

Elm species, North America, Tertiary ........................ 8

eolaciniata, Rub us .............................................................. 9

Ulmus .............................................................................. 1

eolaciniate, Ulmus.............................................................. 9

evidens, Alnus ...................................................................... 9

F, G

Fagopsis longifolia ............................................................ 8

fernquisti, Ulmus ........................................................3, 4, 8

Fingerrock, Nev .................................................................. 8

Fish Creek, Oreg.................................................................. 6

Forests, deciduous, broadleaved, temperate .............. 1

'49 Camp, Nev...................................................................... 6

Green River Formation .................................................... 1

H, I

Healy Creek, Alaska .......................................................... 8

hesperia, Zelkova ............................................................ 3, 4

hopkinsi, Spiraea .............................................................. 10

horniana, Quercus .............................................................. 9

Introduction .......................................................................... 1

K, L

Page

Kilgore, Nebr........................................................................ 8

Knowlton, F. H., cited........................................................ 3

quoted ..............................................................—........... 3

knowltoni, Ulmus............................2, 5, 7; pis. 1C, F, G,

2A, C, H, I, J

laciniata, Ulmus.................................................................. 5

Latah, Wash.......................................................................... 8

Leaves, fossil ....................................................................... 1

Neogene .....................................................................—.. 1

Oligocene...............,.....................-................"..—........... 1

Lesquereux, L., quoted ...................................................... 4

Logan County, Kans .......................................................... 6

longifolia, Fagopsis............................................................ 8

M, N

mccanni, Quercus................................................................ 8

Mascall, Oreg................................................................ 6, 7, 8

Mascall flora ................................................................... 5, 6

Metzel Ranch, Mont ......................................................... 6

montanensis, Ulmus ................................................... 9, 10

moorei, Ulmus .............................................................2, 4, 6

moragensis, Ulmus ........................................................ 1, 9

Myrica oregoniana.......................................................... 3, 5

oregoniana, Mascall flora ........................................ 5

type specimen ..................................................... 3

nervosa, Zelkova.................................................................. 1

nevadensis, Zelkova ............................................ 4, 6, 8, 9

newberryi, Ulmus........................................................ 5, 7, 8

Nomenclature, problems .................................................. 3

O, P

oregoniana, Myrica ........................................................ 3, 5

Zelkova...................................................... 1, 3, 4, 5, 8, 9

owyheensis, Ulmus ...... 2, 4, 6, 7; pis. 2B, D, E, F, 3A

parvifolia, Ulmus................................................................ 1

paucidentata, Ulmus........................ 2, 5, 6, 8, 10; pi. 2G

plurinervis, Ulmus .............................................................. 6

Pollen ...................................................................................... 1

Neogene .......................................................................... 1

Oligocene........................................................................ 1

pseudo-americana, Ulmus ........2, 7; pi. 1A, B, D, E, H

pseudofulva, Chaetoptelea ....................................4, 9, 10

Ulmus ............................................................................ 10

pumila, Ulmus ...................................................................... 6

Q, R

Quercus horniana................................................................ 9

mccanni......................................................................... 8

References cited.................................................................. 10

Remington Hill, Calif........................................................ 5

Rhamnus troutdalensis .................................................... 4

Rockville, Oreg .................................................................... 6

Rosaceae ................................................................................ 2

Rubus ...................................................................................... 9

eolaciniata...................................................................... 9

Ruby basin, Mont................................................................ 7

Seldovia Point, Alaska.................................................. 6, 8

serrata, Zelkova .................................................................. 5

sinica, Zelkova...................................................................... 9

Sonoma, Calif ...................................................................... 5

Sonoma flora, Calif............................................................ 6

Sorbaria.—............................................................................ 10

sorbifolia, Ulmus .............................................................. 10

Species concepts ................................................................. 3

speciosa, Ulmus ..................2, 4, 5, 6, 7, 8, 10; pi. 3C, F

Spiraea hopkinsi................................................................ 10

Stinking Water, Oreg ........................................................ 6

Stinking Water flora, Oreg .............................................. 6

Page 

Sucker Creek, Oreg ............................................................ 6

Systematics............................................................................ 3

Table Mountain................................................................ 4, 5

tanneri, Ulmus ................................................................ 4, 8

tenuinervis, Ulmus........................................................ 9, 10

The Dalles, Oreg.............................................................. 5

Thorn Creek, Idaho ........................................................ 6, 7

Trapper Creek, Idaho ................................................5, 6, 7

Troutdale, Oreg.................................................................... 5

troutdalensis, Rhamnus.................................................... 4

Tsadaka flora, Alaska ...................................................... 2

Twickenham, Oreg......................................——... 5, 6, 7, 8

u

Ulmus................................................................................ 1, 10

Alaskan genera........................................................... 1

areal distribution ........................................................ 2

areoles............................................................................. 3

doubtful species........—................................................. 9

leaves .........................................—.................................. 3

Miocene species............................................................ 1

Oligocene species ........................................................ 1

Pliocene species............................................................ 1

Table Mountain............................................................ 4

western North American species ......................... 1

species, Bridge Creek age ........................................ 2

stratigraphic range .................................................... 2

affinis ....................................... .....2, 4; pi. 3B, D, E, G

restricted range.................................................... 2

syntypes.................................................................. 4

Table Mountain.................................................... 4

americana.................................................................. 4, 7

borealis........................................................................... 9

braunii ............................................................................ 9

brownellii.................................................................... 1, 9

californica.................................................................. 4, 9

Chalk Bluffs locality .......................................... 4

California .......................................................... 4

Oregon ................................................................ 4

Tertiary .............................................................. 4

lectotype.................................................................. 9

synonym ................................................................ 9

Table Mountain...........................—...................... 4

carpinoides .................................................................... 9

chaneyi.......................................................... 2, 5; pi. 4B

leaves ...................................................................... 5

restricted range .....——..................................... 2

Twickenham, Oreg..............—........................... 5

crassifolia ...................................................................... 6

diptera ............................................................................ 9

eolaciniata...................................................................... 1

eolaciniate...................................................................... 9

fernquisti................................................................ 3, 4, 8

*nou)/tonz—..............................2, 5, 7; pis. 1C, F, G,

2A, C, H, I, J

Collawash locality, Oreg .................................. 5

geographic range ..................................———. 2

leaves .............................................—...............— 5

phylogenetic relationship ................................ 2

stratigraphic range ...——...........—............... 2

laciniata, northeastern Asia...—........——............. 5

leaves, identification key..................——................. 2

Tertiary, late ........................................................ 2

middle..................——...........——.—................. 2

nomenclature —————.......—........................ 2

systematics........................................................ 2

western North America ................................ 2

montanensis............................................................ 9, 10

moorei.................—.........................................—..... 2, 4, 6

leaf.......——.............................___.......___.. 6

Metzel Ranch, Mont .............——.............—— 6

restricted range .................................—............. 2

13
14 INDEX

Page 

Ulmus—Continued

moragensis ................................................................ 1, 9

newberryi..-...-.......................................................5, 7, 8

owyheensis .............. 2, 4, 6, 7; pis. 2B, D, E, F, 3A

Bridge Creek locality.......................................... 6

Eagle Creek, Oreg .............................................. 6

geographic range ................................................ 2

leaf............................................................................ 6

stratigraphic range ............................................ 2

Sucker Creek, Oreg ............................................ 6

parvifolia, China ........................................................ 7

Japan ...................................................................... 7

paucidentata................................2, 5, 6, 8, 10; pi. 2G

Mascall flora ........................................................ 6

stratigraphic range ............................................ 2

plurinervis...................................................................... 6

pseudo-americana ................2, 7; pi. 1A, B, D, E, H

phylogenetic relationship ................................ 2

stratigraphic range ............................................ 2

pseudofulva .................................................................. 10

pumila, east Asia ........................................................ 6

sorbifolia ...................................................................... 10

speciosa ..........................2, 4, 5, 6, 7, #, 10; pi. 3C, F

Bridge Creek locality.......................................... 8

geographic range ................................................ 2

Page 

Ulmus speciosa—Continued

stratigraphic range ............................................ 2

type specimen ...................................................... 8

tanneri ........................................................................ 4, 8

leaf............................................................................ 6

Sonoma flora ........................................................ 6

tenuinervis .............................................................. 9, 10

Florissant ............................................................ 10

uenustula ........................................................................ 6

ungeri, Zelkova ............................................................1, 8, 9

Upper Cedarville, Calif .................................................... 8

V, W

uenustula, Ulmus ................................................................ 6

Weiser, Idaho.................................................................... 6, 7

Wrangell Mountains, Alaska .......................................... 8

Zelkova.................................................................................... 1

Alaskan genera............................................................ 1

areoles.............................................................................. 3

Page 

Zelkov a—Continued

flowers ............................................................................ 1

leaves ......................................................3, 5, 9

Miocene species............................................................ 1

Oligocene species ........................................................ 1

Pliocene species............................................................ 1

browni............................................ 1, 3, 8; pi. 4A, C-G

Collawash locality .......................................... 4, 9

geographic range ................................................ 3

stratigraphic range ............................................ 3

carpinifolia, Caucasus region ................................ 9

drymeja ...................................................................... 1, 3

hesperia ...................................................................... 3, 4

Brown's specimen................................................ 3

holotype.................................................................. 3

nervosa............................................................................ 1

nevadensis ...................................................... 4, 6, 8, 9

Ellensberg flora, Wash ...................................... 6

holotype.................................................................. 3

oregoniana ..............................................1, 3, 4, 5, 8, 9

type specimens .................................................... 5

serrata ............................................................................ 5

Japan ...................................................................... 9

sinica, China ................................................................ 9

ungeri...................................................................... 1, 8, 9
PLATES 1-4

Contact photographs of the plates in this report are available, at cost, from U.S. Geological Survey 

Library, Federal Center, Denver, Colorado 80225.
PLATE 1

FIGURES A. Ulmuspseudo-americana Lesquereux. Bridge Creek, Wheeler County, Oreg.: UCMP1758 (original

specimen by Lesquereux, 1883: pi. 54, fig. 10). 

B, H. Ulmus pseudo-americana Lesquereux. Twickenham, Wheeler County, Oreg.: hypotypes UCMP

5731, 5732. 

C. Ulmus knowltoni Tanai and Wolfe, n. sp. Collawash, Clackamas County, Oreg.: paratype

USNM 208503. 

D. Ulmus pseudo-americana Lesquereux. Bridge Creek, Wheeler County, Oreg.: USNM 8493a

(Knowlton, 1902: pi. 9, fig. 2).

E. Ulmus pseudo-americana Lesquereux. Bridge Creek, Wheeler County, Oreg.: USNM 9367. 

F, G. Ulmus knowltoni Tanai and Wolfe, n. sp. White Hill, Belshaw Ranch, Grant County, Oreg.: 

USNM 8532a, b (Knowlton, 1902: pi. 3, fig. 4). These specimens were originally described as 

Myrica oregoniana.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1026 PLATE 1

LEAVES OF ULMUS PSEUDO-AMERICANA AND U. KNOWLTON1
PLATE 2

FIGURES A. Ulmus knowltoni Tanai and Wolfe, n. sp. Collawash, Clackamas County, Oreg.: holotype USNM

208502.

B. Ulmus owyheensis H. V. Smith. Twickenham, Wheeler County, Oreg.: hypotype UCMP 5733. 

C, H, I. Ulmus knowltoni Tanai and Wolfe, n. sp. Collawash, Clackamas County, Oreg.: paratypes

USNM 208504-208506. 

D, E, F. Ulmus owyheensis H. V. Smith. Collawash, Clackamas County, Oreg.: hypotypes USNM 208508-

208510.

G. Ulmus paucidentata H. V. Smith. Twickenham, Wheeler County, Oreg.: hypotype UCMP 5734. 

J. Ulmus knowltoni Tanai and Wolfe, n. sp. Collawash, Clackamas County, Oreg.: paratype USNM 

208507. Showing margin and fine venation (about X 5.8).
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1026 PLATE 2 

•*.

LEAVES OF ULMUS KNOWLTONI, U. OWYHEENSIS, AND C7. PAUCIDENTATA
PLATE 3

FIGURES A. Ulmus owyheensis H. V. Smith. Collawash, Clackamas County, Oreg.: hypotype USNM 208511. 

B. Ulmus affinis Lesquereux. Table Mountain, Tuolumne County, Calif. (USGS loc. 6902): hypotype

USNM 208512. 

C. Ulmus speciosa Newberry. Healy Creek, west of Suntrana, Alaska (USGS loc. 9925): hypotype

USNM 208513. 

D, E. Ulmus affinis Lesquereux. Table Mountain, Tuolumne County, Calif.: UCMP 5735, 5736. These

are unfigured specimens assigned to U. californica by Lesquereux (1878). 

F. Ulmus speciosa Newberry. Bridge Creek, Wheeler County, Oreg.: USNM 7067 (original specimen

by Newberry, 1898: PL 45, fig. 2). 

G. Ulmus affinis Lesquereux. Table Mountain, Tuolumne County, Calif.: UCMP 5737 (original

specimen by Lesquereux, 1878: pi. 4, fig. 4).
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1026 PLATE 3

D

LEAVES OF ULMUS OWYHEENSIS, U. AFFINIS,AND U. SPECIOSA
PLATE 4

FIGURES A, D. Zelkova browni Tanai and Wolfe, n. sp. Collawash, Clackamas County, Oreg. a, holotype USNM 

208514. X 1. d, Enlargement of holotype specimen showing margin and fine venation. Approximately X 4.

B. Ulmus chaneyi Tanai and Wolfe, n. sp. Twickenham, Wheeler County, Oreg. Holotype UCMP

no. 5738. X 1.

C, E-G. Zelkova browni Tanai and Wolfe, n. sp. Collawash, Clackamas County, Oreg. Paratypes USNM 

208515-208518. X 1.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1026 PLATE 4

LEAVES OF ULMUS CHANEYI AND ZELKO VA BROWNI

; oPO: 1977-792- 204