The Miocene Seldovia

Poit Flor from the

Kenai Group, Alaska
The Miocene Seldovia 

Point Flora from the 

Kenai Group, Alaska

By JACK A. WOLFE and TOSHIMASA TANAI

GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1980
UNITED STATES DEPARTMENT OF THE INTERIOR 

CECIL D. ANDRUS, Secretary

GEOLOGICAL SURVEY 

H. William Menard, Directorof Congress Cataloging in Publication Data

Wolfe, Jack A. 1936-

The Miocene Seldovia Point flora from the Kenai group, Alaska.

(Geological Survey professional paper ; 1105) 

Bibliography: p. 45-47. 

Includes index.

1. Paleobotany-Miocene. 2. Paleobotany-Alaska-Cook Inlet region.

I. Tanai, Toshimassa, joint author. II. Title. III. Series: United States.

Geological Survey. Professional paper ; 1105. 

QE929.W64 561'.2'097983 79-20550

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

Washington, D.C. 20402

Stock number 024-001-03282-1
CONTENTS

Abstract

Introduction

Geologic occurrence

Age

Floristic composition

Vegetation

Megafossils

Pollen -__ _ _ _______

Page 

1

___________________ 1

3

___________________ 7

q

________________ __ 11

_____ _____________ 11

_ _________ ______ 14

Page

Vegetation—Continued

Distribution ______________________________ — __ 15

Communities _____________________________ —— _-_— _ 16

Floristics _________________________-----------__---------- 16

Paleoclimatology _______________________________--------_ 19

Systematics _________________________________--------_ 24

References cited ________________________________________ 45

Index __________________________-__—— — — — — — — 49

ILLUSTRATIONS

	[Plates follow index]

PLATE 1. Filicales, Coniferales, Magnoliidae.

2. Hamamelididae, Ranunculidae.

3. Hamamelididae.

4. Hamamelididae.

5. Hamamelididae.

6. Hamamelididae.

7. Hamamelididae.

8. Hamamelididae.

9. Hamamelididae.

10. Hamamelididae.

11. Hamamelididae, Dilleniidae.

12. Dilleniidae.

13. Dilleniidae.

14. Dilleniidae, Rosidae.

15. Dilleniidae, Rosidae.

16. Rosidae.

17. Rosidae.

18. Rosidae.

19. Rosidae.

20. Rosidae.

21. Rosidae.

22. Rosidae, Monocotyledones.

23. Rosidae, Monocotyledones.

24. Rosidae.

25. Asteridae, Monocotyledones.

FIGURE 1. Map of Alaska _________________________________________________________________.

2. Map of Cook Inlet region _____________________________________________________ ——————— _.

3. Chart showing changes in usages of stages and ages in the Oligocene through Pliocene series of Alaska .

4. Map of the Seldovia area ____________________________________________________.

5. Diagram showing section of Kenai Group exposed between Seldovia and Barbara Points __—————___.

6. Suggested correlations of the Kenai Group and the coal-bearing group of the Nenana coalfield —____—.

7. Suggested distribution of vegetational types in Alaska during middle Miocene time ______ —— __ — __.

8. Graph showing temperature parameters of modern vegetational types in eastern Asia and in the probable 

	temperature parameters of certain middle Miocene assemblages _____________________.

9. Map of northeastern Asia showing location of some middle Miocene assemblages _____——————.

10. Map of northwestern North America showing location of some middle Miocene assemblages _ ————— __.

Page

_ 2 

_ 3 

_ 4 

_ 5 

_ 6 

_ 9 

_ 15

19

21

.23

in
iv CONTENTS

TABLES

Page

TABLE 1. Average percentages of samples containing pollen of certain broad-leaved genera in Seldovian rocks of the Nenana coalfield,

Alaska Range ________________________________________________________________________ 7

2. Known stratigraphic ranges of Seldovia Point dicotyledonous species in Japan and conterminous United States ______ 9

3. Assumed growth habits of Seldovia Point species __________________________________________——— 12

4. Distribution of most closely related extant species in vegetational types in eastern Asia _______________________— 12

5. Seldovian dicotyledonous species also occurring in Mixed Mesophytic forest in the early and middle Miocene of the Pacific

Northwest _____________________________________________________________________ 13

6. Percentages of pollen types in samples from the Seldovia Point beds __________________— — _________ — _— 14

7. Analysis of the Seldovia Point assemblage in terms of floristic elements ____________________________ ——— __ 17
THE MIOCENE SELDOVIA POINT FLORA FROM THE 

KENAI GROUP, ALASKA

By JACK A. WOLFE and TOSHIMASA TANAI

ABSTRACT

Approximately 60 species of megafossil plants are illustrated and 

are assigned to 45 genera. Most of the species are dicotyledonous, 

although monocotyledons, ginkgo, conifers, and ferns are also represented. Nine species of dicotyledons and one species of 

monocotyledon are described as new. The flora includes the first described representatives of Cyclocarya, Nymphar, Sorbaria, Pueraria, 

Decodon, Kalopanax, Lonicera, and Alisma in the North American 

Tertiary.

The plant-bearing beds are considered to be part of the Kenai 

Group. The West Foreland Formation (late Paleocene) is excluded 

from the Kenai Group, but the following formational units are accepted within the group, in ascending order: Hemlock Conglomerate 

(early Oligocene), Tyonek Formation (early Oligocene through middle Miocene), Beluga Formation (middle and late Miocene), and 

Sterling Formation (late Miocene and Pliocene). Paleobotanical evidence indicates that the Seldovia Point beds are equivalent to part 

of the Tyonek Formation; the geology of these beds indicates that 

they represent deposits that filled a valley to the south of the main 

part of the Kenai basin. Paleobotanical correlations also indicate 

that the Seldovia Point flora is, in provincial terminology, of late 

Seldovian age and further that the upper part of the Seldovian 

Stage is of late early and early middle Miocene age.

Analysis of the Seldovia Point assemblage from both floristic and 

physiognomic standpoints indicates that the assemblage represents 

Mixed Northern Hardwood forest, although palynological data indicate that coniferous forest was close by. The Seldovia Point assemblage has several genera that no longer participate in Mixed 

Northern Hardwood forest and that today are restricted to broadleaved evergreen or Mixed Mesophytic forests. These differences are 

interpreted in light of the history of Mixed Northern Hardwood and 

related forest types during the Neogene.

Analysis of the Seldovia Point assemblage in terms of the derivation of component lineages indicates that of those species whose 

lineages are reasonably well known, about one-third of the species 

are of east Asian origin, one-third of west American (middle 

latitude) origin, and one-third of high-latitude origin. The lack of 

penetration of west American species southward into eastern Asia 

and the lack of penetration of east Asian species southward into 

middle latitudes of western North America are suggested to be the 

result of the loss of genetic plasticity in regard to adaptation to different photoperiodic conditions.

Paleoclimatic conditions inferred from the Seldovia Point assemblage strongly indicate that since the middle Miocene there has 

been a moderate decline in mean annual temperature and a major 

decrease in mean annual range of temperature. Analysis of other 

middle Miocene plant assemblages at middle latitudes in both eastern Asia and western North America indicates that the same basic 

pattern of temperature shifts occurred, although the changes in 

mean annual range of temperature were most pronounced in western North America and particularly Alaska. The primary temperature parameter that was altere 1 was warm-month temperatures, 

which underwent a severe decline; this decline in summer temperatures is probably one of the causes of the initiation of glaciation at 

high latitudes during the late Cenozoic. The Seldovia Point assemblage, when compared to Alaskan assemblages of early and late 

Miocene age, is consistent with the concept of a middle Miocene 

warming, which has been previously documented at middle 

latitudes.

INTRODUCTION

The floras of the Kenai Group of the Cook Inlet region in Alaska have attracted the interest of paleobotanists for over 100 years. Plants collected by or for 

Furuhjelm, at one time governor of what was then 

Russian America, were submitted to the Swiss paleobotanist Oswald Heer; these plants came from two 

localities—one near the village of Ninilchik on the 

east shore of Cook Inlet and the second supposedly 

near English Bay on the southwestern part of the 

Kenai Peninsula (fig. 1). These plants, along with 

those from Kuiu Island in southeastern Alaska, were 

described and illustrated by Heer (1869a) as a part of 

his monumental Flora Fossilis Arctica.

The acquisition of Alaska by the United States in 

1867 led ultimately to investigations of the Cook Inlet 

region by members of the U.S. Geological Survey. 

These investigations included collection and analysis 

of the fossil floras of the Kenai Group (for example, 

Martin and others, 1915), but the only comprehensive 

taxonomic analysis of these floras prior to the 1960's 

was by Hollick (1936). It is notable that these earlier 

collections were small and that none were made by 

paleobotanists. Rollick's fieldwork in Alaska involved 

collecting plants from Cretaceous rocks along the 

Yukon River (Hollick, 1930). Although the late R. W. 

Chaney discussed the floristic significance of the 

Kenai floras in numerous papers (for example, 

Chaney, 1936, 1952, 1967), he never carried out 

paleobotanical field investigations on the Kenai floras, 

which occupied a central position in his concept of an 

"Arcto-Tertiary Geoflora."

The first paleobotanist to undertake collection of the 

Kenai floras was the late R. W. Brown, who made 

small collections from five localities in 1955. Brown, 

however, was nearing retirement and made only tentative, unpublished generic determinations of some of 

the Kenai material. The need for comprehensive field
MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

100 50 0 

I . I

100

I

200MILES 

I

n ' i II

100500 100 200 KILOMETERS

FIGURE 1.—Map of Alaska showing location of Cook Inlet region.

investigations and laboratory studies of not only the 

Kenai floras but also other Alaskan Tertiary floras 

was emphasized by many geologists working in 

Alaska and particularly by D. M. Hopkins. Fieldwork 

was initiated by Wolfe in southeastern Alaska in 1961 

and was extended into the Cook Inlet region in 1962. 

The decision to concentrate on collection and study of 

the floras of the Kenai Group was made because of the 

historical significance of these floras, the discovery of 

petroleum and natural gas in Kenai rocks, and the 

considerable thickness of this unit, which indicated 

the possibility of obtaining numerous assemblages in 

demonstrable stratigraphic succession. Stratigraphically significant species from the Kenai Formation of 

former usage were discussed and illustrated by Wolfe 

(1966), and also incorporating palynological studies by 

Wolfe and E. B. Leopold, these species formed the

basis for a biostratigraphic framework for the Alaskan Neogene (Wolfe and others, 1966). Wolfe (1966) 

also gave a tentative list of the Seldovia Point flora. 

Collections made in 1962 from localities near Seldovia and extending southwestward to Port Graham 

(Heer's "English Bay" material was, in fact, obtained 

from Coal Cove on Port Graham; see also Martin and 

others, 1915) were the largest obtained from the 

Kenai, in terms of both specimens and species. Collectively, the material from USGS palebotanical localities 9856 (Coal Cove), 9857, and 9858 (Seldovia 

Point) is the basis for the Seldovia Point flora. Additional material was collected by Wolfe in 1967 from 

Seldovia Point and from new localities along the north 

shore of Kachemak Bay and elsewhere in the Cook 

Inlet region. Although the Seldovia Point locality was 

not visited, we obtained additional material from the
GEOLOGIC OCCURRENCE

upper part of the Kenai Group in 1973. Added to this 

material is a modest but nevertheless significant collection made by C. E. Allison in 1971 at Seldovia 

Point.

The material now in hand from the Kenai Group 

allows a far better understanding of the floristic, 

climatic, and stratigraphic significance of the various 

Kenai floras. Preliminary statements on the significance of these floras have been published 

elsewhere (Wolfe, 1969a, 1969b, 1972), but this report 

presents the first thorough study of any of the Kenai 

floras, a study requisite for the documentation or modification of previous statements. The Seldovia Point 

flora is particularly significant because it allows an 

understanding of floristic and climatic relationships 

around the North Pacific Basin during the middle 

Miocene; many floras of this time have received detailed attention both in Japan (Tanai, 1972) and in 

the conterminous United States (Wolfe, 1969a).

We gratefully acknowledge the continuing encouragement of D. M. Hopkins in our studies; the geologic 

section of the beds at Seldovia Point is based largely 

on Hopkins' field observations made in 1962. Estella 

B. Leopold has provided us with valuable data based 

on her largely unpublished but extensive studies of 

samples from the Nenana coalfield. For assistance in 

particular taxonomic problems we are grateful to Professor Huzioka (Akita University) and Dr. H. D. MacGinite (University of California, Berkeley). Dr. Carol 

Allison (University of Alaska) allowed us to include 

her university's collections from Seldovia Point in this 

study.

GEOLOGIC OCCURRENCE

The bulk of the Tertiary beds of the Cook Inlet region (fig. 2) have been variously referred to as the 

Kenai Series, Kenai Group, or Kenai Formation. As 

well, various informal lithologic subdivisions of the 

Kenai rocks have been proposed (for example, Kelly, 

1963), but only recently have formal lithologic subdivisions been proposed (Calderwood and Fackler, 

1972).

Three stages have been previously erected for the 

time interval during which the Kenai Group was deposited (Wolfe and others, 1966); in ascending order, 

these are the Seldovian, Homerian, and Clamgulchian. 

The Seldovian Stage was considered to be of 

Oligocene(?) through approximately middle Miocene 

age, the Homerian to be probably of later Miocene 

age, and the Clamgulchian to be largely of Pliocene 

age. The Seldovian stage was, moreover, informally 

subdivided into a lower(?) and upper (?) part based on 

the analysis of certain plant assemblages, some of 

which appeared to be older than those in the type section of the Seldovian Stage at Capps Glacier and

152° 150°

62°

Approximate area of 

outcrops of Kenai

FIGURE 2.—Cook Inlet region, showing approximate extent of outcrops of the Kenai Group.

along Chuitna River and in its reference section of the 

Seldovian Stage at Seldovia Point. Continued work 

(Wahrhaftig and others, 1969) indicated that in the 

Nenana coalfield of the central part of the Alaska 

Range plant assemblages approximately correlative 

with the lower(?) Seldovian assemblages in the Cook 

Inlet region were demonstrably lower stratigraphically than assemblages correlative with those in the 

type and reference sections of the Seldovian Stage; the 

rocks containing these lower(?) Seldovian assemblages 

were consequently excluded from the Seldovian Stage. 

Wolfe (1969b, 1972, 1977) has applied the term Angoonian Stage to these Oligocene assemblages formerly termed lower(?) Seldovian (fig. 3).

We are here accepting the elevation of the Kenai 

Formation to the status of Kenai Group as proposed 

by Calderwood and Fackler (1972), with the exception 

that we exclude their West Foreland Formation from 

the Kenai Group, defined as "a tuffaceous siltstone-
MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Wolfe and others 

(1966)

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FIGURE 3.—Changes in usages of stages and ages in the Oligocene through Pliocene series of Alaska.

claystone unit containing a few conglomeratic 

sandstone and thin coal beds***" (Calderwood and 

Fackler, 1972, p. 741). Such lithology differs markedly 

from the overlying Kenai Group, which typically lacks 

tuffaceous material; indeed, we have observed only 

one bed of volcanic ash (primarily glass) in the Kenai; 

this ash bed occurs in the upper part of the Kenai 

Group south of Clam Gulch. Some coalbeds of the 

upper part of the Kenai Group also contain ash partings (Triplehorn and others, 1977).

The West Foreland Formation, as noted by Calderwood and Fackler (1972), is probably exposed at the 

surface along the west side of Cook Inlet. One of the 

most extensive surface exposures of this unit is south 

of Capps Glacier and includes rocks mapped as "lower 

Kenai" by Barnes (1966). These rocks are highly tuffaceous and are angularly unconformable beneath the 

type section of the Seldovian Stage; elsewhere the 

West Foreland Formation has a consistent angularly 

unconformable relationship to overlying units (Calderwood and Fackler, 1972). Although these rocks at 

Capps Glacier are more conglomeratic than at the

type section of the West Foreland, they are also at the 

margin of the depositional basin and might be expected to be conglomeratic in part. The age of the 

West Foreland Formation is not certainly known. If, 

however, the beds south of Capps Glacier are part of 

this unit, leaf samples from these beds indicate a 

latest Paleocene age for the West Foreland (Wolfe, 

unpub. data). This would indicate that an interval of 

at least 20 million years is represented by the unconformity between the West Foreland Formation and 

the Kenai Group, which is probably no older than 

early Oligocene.

As here recognized, the Hemlock Conglomerate 

(Calderwood and Fackler, 1972) is the basal formation 

of the Kenai Group. Surface exposures of the Hemlock 

Conglomerate are not certainly known, although some 

of the nonvolcanic conglomerate mapped as "lower 

Kenai" by Barnes (1966) may represent this unit. 

Inasmuch as the Hemlock Conglomerate is not separated from the overlying Tyonek Formation by an 

unconformity and is only about 200 to 250 m thick 

(Calderwood and Fackler, 1972), the Hemlock is prob-
GEOLOGIC OCCURRENCE

ably not much older than the Tyonek Formation; the 

Hemlock Conglomerate is considered to be of early 

Oligocene age. Indeed, it is possible that the Hemlock 

is a discontinuous basal conglomerate of the Tyonek 

Formation and that some typical Tyonek beds may be 

isochronous with the Hemlock (for example along 

Harriet Creek).

Calderwood and Fackler (1972) considered that they 

had only redefined Spurr's (1900) "Tyonek beds." Because they designated the type section in a well, however, it is clear that their Tyonek Formation is a new 

concept. Additionally, it was previously noted (Wolfe 

and others, 1966) that fossils from the outcrops of the 

"Tyonek beds" near Tyonek represent the Homerian 

Stage, the type section of which (Wolfe and others, 

1966), as Calderwood and Fackler pointed out, is part 

of the next younger Beluga Formation. Calderwood 

and Fackler (1972) correlated the Kenai Group along 

the upper part of the Chuitna River with the type section of the Tyonek Formation; this correlation places 

the type section of the Seldovian Stage within the 

Tyonek Formation. The Tyonek Formation therefore 

includes rocks of early and middle Miocene age. That 

the Tyonek Formation also includes rocks of early and 

late Oligocene Age (Angoonian) is indicated by Calderwood and Fackler's statement (1972, p. 745) that 

the Chuitna River section is incomplete and evidenced 

by basal coalbearing rocks of the Tyonek Formation 

exposed on Harriet Creek that are of pre-Seldovian 

age. Elsewhere in Alaska (Wahrhaftig and others, 

1969), rocks correlated with the Kenai Group, considered by Wolfe, Hopkins, and Leopold (1966) to be 

"lower(?) Seldovian," have been excluded from the 

Seldovian Stage and are now considered to be of early 

and late Oligocene age. As well, the Tyonek Formation presumably extends into the middle Miocene. 

This is indicated by the fact that the highest exposures of the Tyonek Formation along Chuitna River 

are of Homerian age, and no unconformity between 

the Seldovian and Homerian stages was noted in this 

section. Thus, the Tyonek Formation is here considered to range in age from early Oligocene through 

middle Miocene. Considering that the Tyonek Formation has such a great age range and considering the 

thickness of the unit (over 2,300 m), this unit may be 

capable of further subdivision. The beds at Seldovia 

Point are thus probably equivalent in age to the 

Tyonek Formation.

The Beluga Formation (Calderwood and Fackler, 

1972) is the next youngest unit of the Kenai Group. 

Calderwood and Fackler correlated the Kenai Group 

exposed near Homer, representing the Homerian 

Stage, with the Beluga Formation, and hence this 

part of the Beluga is of middle and late Miocene age. 

However, the lower part of the reference section of the

Clamgulchian Stage along Kachemak Bay includes 

calcareous beds, and Calderwood and Fackler (p. 751) 

indicate that the highest occurrence of calcareous beds 

in the Kenai Group is in the upper half of the Beluga 

Formation. Thus, all the Beluga Formation is here 

considered to be of late middle and late Miocene age. 

We emphasize, however, that we are accepting the 

placement of the Miocene-Pliocene boundary generally accepted by European geologists at about 5-6 

million years.

The youngest unit of the Kenai Group is the Sterling Formation (Calderwood and Fackler, 1972). The 

correlation of this unit with surface outcrops from 

Clam Gulch south beyond Ninilchik (type exposures of 

the Clamgulchian Stage) appears convincing (Calderwood and Fackler, 1972), and thus the Sterling Formation is considered to be of late Miocene and 

Pliocene age.

The Seldovia Point flora was collected from beds of 

the Kenai Group exposed in a discontinuous series of 

outcrops in the sea cliffs from Barabara Point west 

and south to Coal Cove on Port Graham (fig. 4). These 

beds are typically weakly lithified, although the leaf 

impressions were obtained from limy, indurated 

lenses. These remnants of the Kenai Group rest un15 KILOMETERS

J

FIGURE 4.—Map of Seldovia area. Outcrops of Kenai Group shown 

by stippling; numbers refer to U.S. Geological Survey paleobotanic&l localities.
MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

conformably on altered Mesozoic rocks, which also 

form the headlands that have protected these remnants from total erosion.

The outcrop of the Kenai Group at Coal Cove probably yielded the collection of plants described by Heer 

(1869a), and attributed to a locality on English Bay. 

No Tertiary rocks are exposed on English Bay, and 

the Coal Cove outcrops were mined for coal for Russian vessels. At the east end of this outcrop, the Kenai 

dips gently off the Mesozoic basement and contains 

coarse clasts probably derived from the subjacent 

Mesozoic rocks; that is, the Kenai rocks resemble a 

fossilized talus. Indeed, all outcrops of the Kenai 

Group from the Coal Cove to Barabara Point have 

taluslike appearances near the contacts with the 

Mesozoic rocks. In places, coalified logs or casts of logs 

are abundant in the Kenai. Where observable in 

fiords, the outcrops of the Kenai Group do not extend 

far inland but are apparently continuous along a narrow belt that parallels the present coast (fig. 4).

A reasonable interpretation of the lithology and 

pattern of outcrops of the Kenai Group in this area is 

that the Kenai here represents the remnants of an ancient drainage system. This system drained from the 

southwest into the main part of the Kenai basin, as 

indicated by the greater thickness and areal extent of 

the Kenai Group at Seldovia Point (fig. 5) in comparison to the outcrops near Point Pogibshi and on Coal 

Cove. Although most of the rocks represent fluviatile 

deposits, some of the sediments were deposited in 

slowly moving water; this is indicated not only by the 

presence of some coals but also by the abundance at 

locality 9858 of Potamogeton and Nymphar.

The exact altitude at which the Seldovia Point beds 

accumulated is not known, but it was probably less 

than about 100 m. Although no marine beds are 

known in any part of the Kenai Group, the great 

thickness of this unit (over 8,000 m), the great areal 

extent of the basin (about 110 km wide and 330 km 

long), and the duration of apparently continuous deposition (about 25 m.y.) all indicate deposition near 

sea level for the main part of the basin. The fact that 

the Seldovia Point beds (particularly the plantbearing beds) include fine-grained beds associated 

with coal also indicates that the drainage had a low 

gradient. We thus think that the Seldovia Point beds 

were not deposited at an altitude appreciably higher 

than the remainder of the Kenai Group.

The nearest marine beds of middle Miocene age to 

the Seldovia Point localities are on the northeastern 

part of Kodiak Island, about 170 km to the south (fig. 

1). The marine megafossils from the Kodiak beds indicate shallow water (W. O. Addicott, oral commun., 

Dec. 1973), and this indicates that during the middle 

Miocene the sea did not extend to the north much beCOVERED

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00 - --

- — — —o o 9858

COVERED 

(thickness 

unknown)

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METERS FEET 

35 -r 100

15 -- 50

0 -1-0

FIGURE 5.—Section of Kenai Group 

exposed between Seldovia and 

Barabara Points. Number refers 

to U.S. Geological Survey paleobotanical locality.

yond northeastern Kodiak Island. To the east of Seldovia, the nearest occurrence of middle Miocene 

marine beds is in the Yakataga District, about 500 

km east and slightly north of Seldovia. Some clastic 

rocks of the Kenai Group indicate derivation from the 

area now occupied by the Chugach Mountains, but 

such rocks are younger than the Beluga Formation, 

which is probably of Homerian and early Clamgulchian age. There is thus no evidence that the western 

part of the Chugach Mountains were elevated during 

the middle Miocene.

Most sediments of the Kenai basin deposited during 

the early and middle Miocene were apparently derived from the southern part of the Alaska Range 

(Calderwood and Fackler, 1972, p. 751, 754). Presumably that source area had considerable relief, and 

these mountains—about 150 km from Seldovia—were 

the mountains nearest to Seldovia. The central part of 

the Alaska Range to the north of the Kenai basin was
AGE

apparently a low region during the early and middle 

Miocene, as attested to by the considerable thickness 

of coals in the Suntrana Formation and the extensive 

lacustrine deposits of the Sanctuary Formation 

(Wahrhaftig and others, 1969).

AGE

The Seldovia Point flora has had varied age assignments. Originally dated simply as Miocene by Heer 

(1869a), the flora from the Port Graham locality 

(Heer's English Bay locality) was then—without any 

substantial factual data—assigned to the Eocene 

Epoch (Knowlton, 1894). The Eocene age was accepted 

without question by a number of paleobotanists (for 

example, Hollick, 1936; Chaney, 1936). Reanalysis of 

illustrations of the material from Port Graham led 

Wolfe (in MacNeil and others, 1961) to conclude that 

this flora was no older than late Oligocene; MacGinitie (1962) concurred in this opinion and suggested 

that the flora might be as young as early Miocene. 

Basing their conclusions on new material from both 

Port Graham and Seldovia Point, Wolfe, Hopkins, and 

Leopold (1966) assigned the Seldovia Point flora (including the Port Graham locality) to the Seldovian 

Stage, which was thought to be of late Oligocene(?) 

and earlier Miocene age. Although an attempt was 

made to reassign this flora to the early Oligocene 

(Chaney, 1967), the factual basis for this reassignment proved illusory (Wolfe, 1969b). In the most recent summary of Alaskan Tertiary floras, the Seldovia 

Point assemblage was again assigned to the early 

Miocene (Wolfe, 1972), but it has also been suggested 

that the flora is of middle Miocene age (Tanai, 1973). 

The question of whether the flora is of early Seldovian 

(earliest Miocene) or late Seldovian (late early and 

early middle Miocene) age is particularly significant 

in constructing paleoclimatic models.

The rocks containing the Seldovia Point assemblage 

were designated as a reference section for the Seldovian Stage (Wolfe and others, 1966). This designation 

was made primarily because the flora from this reference section was then (and remains) considerably 

richer than the flora from the type section of the Seldovian Stage at Capps Glacier and along the Chuitna 

River. Considerable documentation for the early and 

middle Miocene age of the Seldovian Stage has been 

presented (Wolfe, 1969b).

The occurrence of the Seldovia Point assemblage in 

an isolated outcrop of the Kenai Group requires that 

the placement of these rocks within subdivisions of 

the Seldovian Stage, however, be based solely on 

paleobotanical evidence. Only two informal subdivisions—lower and upper—of the Seldovian Stage

are recognized at Capps Glacier and along the 

Chuitna River; plant megafossils have been found 

primarily in the lower part of the section (Wolfe and 

others, 1966), but one small flora (loc. 9848 of Wolfe 

and others, 1966) is probably in the upper part of the 

Seldovian Stage, judging from Barnes' (1966) geologic 

map. Several pollen assemblages, have, however, been 

studied from both the upper and lower parts of the 

type section of the Seldovian Stage (Wolfe and others, 

1966).

The upper Seldovian has been equated to the middle 

Miocene, the Homerian to the upper Miocene, and the 

Clamgulchian to the Pliocene (Wolfe, 1969b, 1972). In 

terms of the now widely accepted planktonic foraminiferal chronology, however, the upper Seldovian is 

of late early and early middle Miocene age and the 

Homerian is of late middle and early late Miocene 

age (Wolfe, 1980 ). The Clamgulchian has radiometric 

ages in its lower part that indicate a latest Miocene 

age (Triplehorn and others, 1977); the upper Clamgulchian is assumed to be of Pliocene age.

The rocks of the Nenana coalfield of the central 

Alaska Range (fig. 1) span the entire Seldovian Stage. 

The lowest megafossil and microfossil assemblages in 

this conformable sequence are pre-Seldovian, and the 

highest rocks are of Homerian and Clamgulchian age 

(Wahrhaftig and others, 1969). This sequence has also 

been extensively sampled for pollen. Palynologically, 

some differences are apparent between the pollen assemblages of the lowest part of the Seldovian (the 

upper part of the Healy Creek Formation) and the 

remainder of the stage (Sanctuary Formation and the 

overlying Suntrana Formation).

Upper Seldovian pollen assemblages from the 

Alaska Range appear to have a greater diversity of 

broad-leaved trees than do lower Seldovian assemblages. Ulmus and Liquidambar occur more frequently in upper (table 1) than lower Seldovian assemblages, and Fagus and Tilia, which are present in

TABLE 1.—Average percentages of samples containing pollen of certain broad-leaved genera in the Seldovian rocks of the Nenana 

coalfield, Alaska Range

[Based on unpublished data supplied by E. B. Leopold]

Fagus

Quercus

Liquidambar

Ulmus ... _ ____ __ __.

Tilia ......... . - ... ...

Carva

Juglans __ __ ___.

Pterocarva . .. ..... ..

Ilex

Upper part 

of 

Healy Creek 

Formation 

(12 samples)

0

25

17

„ 0

42

.. 8

. 83

58

Sanctuary 

and 

Suntrana 

Formations 

(28 samples)

29

29

29

89

21

54

18

71

61
8 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

about one-fourth the upper Seldovian samples, are absent in lower Seldovian samples. The sample from 

Seldovia Point has Ulmus, Liquidambar, Tilia, and 

Fagus; the megafossil flora from Seldovia Point also 

contains representatives of all four genera. Of the four 

genera, only Ulmus is yet known from any lower Seldovian megafossil assemblage. These data indicate 

that the Seldovia Point flora is of late Seldovian 

(probably late early and early middle Miocene) age.

Although the megafossil assemblages from the Seldovian rocks of the Nenana coalfield are admittedly 

small, these can also be interpreted to indicate that 

the Seldovia Point flora is of late Seldovian age. The 

flora of the upper part of the Healy Creek Formation 

(lower Seldovian) is known from moderate-size collections that include:

Metasequoia sp.

Populus kenaiana Wolfe

Populus aff. P. eotremuloides Knowlt.

Salix sp.

Carya sp.

Pterocarya nigella (Heer) Wolfe

Alnus cappsi (Holl.) Wolfe

Betula aff. B. thor Knowlt.

Quercus furuhjelmi Heer

Ulmus sp.

Cladrastis aff. C. lutea Michx.

Acer chaneyi Knowlt.

The flora of the overlying Sanctuary Formation was 

thought to be very small (Wahrhaftig and others, 

1969), but this was due to considering locality 7476 to 

be in the Healy Creek Formation. Further discussion 

with Wahrhaftig and reanalysis of Prindle's original 

field notes indicate that this locality is probably in the 

Sanctuary Formation. The flora from this locality includes:

Metasequoia sp.

Pterocarya nigella (Heer) Wolfe

Alnus healyensis Wolfe

Fagus antipofi Heer

Quercus furuhjelmi Heer

Cocculus auriculata (Heer) Wolfe 

To the above list can be added certain elements of the 

overlying Suntrana flora:

Populus kenaiana Wolfe

Cyclocarya ezoana (Tanai et Suz.) Wolfe et Tanai

Alangium milii Wolfe et Tanai

Ulmus sp.

Probably significant is that the known Upper Healy 

Creek flora does not contain certain species that are 

present in both the Seldovia Point flora and in the 

flora of the Sanctuary and Suntrana Formations: Cyclocarya ezoana, Fagus antipofi, Cocculus auriculata, 

and Alangium mikii.

The differences between the microfossil assemblages 

of the upper and those of the lower part of the type 

section of the Seldovian Stage are not as apparent as 

in the Alaska Range section, primarily because fewer 

samples from the type section have been analyzed. In 

the three lowest samples (Iocs. D1946, D1953, D1952 of 

Wolfe and others, 1966), however, Tilia is absent, and 

only one sample contains Liquidambar (and the genus 

is so rare that it did not appear in the pollen counts). 

In contrast, in the four upper Seldovian pollen samples (Iocs. D1720, D1718, D1719, D1949) one sample 

contained rare Tilia, and two samples contained significant amounts of Liquidambar pollen.

The megafossil assemblages from the lower part of 

the type section of the Seldovian Stage are based on 

collections almost as extensive as those from Seldovia 

Point and were obtained from four horizons. The flora, 

however, is depauperate in comparison to that at Seldovia Point. Although some species are common to 

both floras, and in particular Alangium mikii also occurs at Capps Glacier, the Capps Glacier assemblage 

lacks the other three species considered to be significant in correlating the Seldovia Point with the 

Sanctuary-Suntrana flora. The small collection from 

the upper part of the Seldovian section along Beluga 

River (loc. 9848) contains only a few species, but it is 

perhaps significant that this flora has the only other 

known occurrence of Acer ezoanum in Alaska.

Certainly further collecting in both the type section 

of the Seldovian Stage and in the Seldovian rocks of 

the Alaska Range is desirable, but the available 

data—from both megafossils and microfossils— 

indicate that the Seldovia Point flora is probably of 

late Seldovian age. Tentative correlations between the 

Cook Inlet region and Nenana coalfield are given in 

figure 6.

The basis for the early and middle Miocene age of 

the Seldovian Stage has been discussed at some 

length previously (Wolfe, 1969b). The current systematic treatment of the Seldovia Point flora, however, allows a more definitive basis for correlating this 

flora with other plant assemblages at lower latitudes 

on either side of the Pacific Ocean. Most of the gymnospermous species—at least those based on megascopic characters—have such long ranges during the 

Tertiary as to be of little value in biostratigraphy. It 

is only certain dicotyledonous species that are of most 

value in establishing correlations.

Even some of the dicotyledonous species that occur 

outside of Alaska are of less value than others in biostratigraphy. Nymphar ebae, for example, is known 

both at Seldovia Point and in the early Miocene Aniai 

flora of Honshu (Huzioka, 1964), but because so little 

is known of the evolution of Nymphaeaceae during 

the Tertiary, the stratigraphic value of this species is
FLORISTIC COMPOSITION

Ser es

Pliocene

ocene

5

Oligocene

U. 

M

L

U

L

Cook Inlet

UJJMUMUW-M

Sterling Formation

IJP^ 

Beluga Formation

]J)QCP«

Tyonek 

Formation

Hemlock ~z~, 

Conglomerate *C__ ttMirriiTMTr

-

a

D 

0

(5

c 

a>

y

Nonmarine Stage

Clamgulchian

Nenana coalf eld

^/"v^-v_^-x-x-v_/^

Nenana Gravel

Tiinjxu»^rubstake

\y^V^^ Format on

Homerian

Seldovian

Angoonian

Lower

Upper

Lower

Lignite Creek 

Formation

Suntrana Formation

Sanctuary Formation

Healy Creek 

Formation

FIGURE 6.—Suggested correlations of the Kenai Group and the 

coal-bearing group of the Nenana coalfield. Stage nomenclature 

follows Wolfe, Hopkins, and Leopold (1966) and Wolfe (1968, 

1969b).

uncertain. In contrast, the occurrence of Acer ezoanum 

is much more significant in that this species is considered to be a descendant of the Oligocene S. 

kushiroanum (Tanai, 1970, 1972). Similarly, the occurrence in the Seldovia Point flora of species such as 

Platanus bendirei (see p. 29) is also of considerable 

significance. In table 2 are listed the Seldovia Point 

species that occur outside Alaska and the known 

ranges of these species.

Data presented in table 2 indicate that the Seldovia 

Point flora is probably of late early or early middle 

Miocene age. This age is consistent with the occurrence of some Seldovian species elsewhere in Alaska 

in beds independently dated from marine invertebrates (Wolfe, 1969a). We emphasize that, as discussed 

in the section on vegetation, the Seldovia Point assemblage represents a vegetational type cooler than 

that found in the late early to early middle Miocene of 

Japan and the conterminous United States. The occurrence of some species both at Seldovia Point and 

farther to the south is analogous to the present distribution of certain woody species that (1) occur from 

Japan to Alaska and at low altitudes to the conterminous United States (for example Oplopanax horridum (J.E. Sm.) Miq. and Myrica gale L.), (2) occur 

from Alaska to the conterminous United States at low 

altitudes (for example Malus fusca (Raf.) Sarg. and 

Populus trichocarpa Torr. et Gray), and (3) occur from 

Alaska to Japan (for example, Rhododendron 

camtschaticum Pall, and Phyllodoce aleutica (Spreng.) 

Heller). We suggest that the geologic ranges of many 

dicotyledonous species during the Tertiary are 

sufficiently restricted to allow correlations to be made

TABLE 2.—Known stratigraphic ranges of Seldovia Point dicotyledonous species in Japan and conterminous United States

[--.-, Japan; —— U.S.]

Species

Oligocene

Miocene Pliocene

Nymphar ebae ............ ___

Cocculus auriculata ....... _

Liquidambar pachyphylla . 

Platanus bendirei ..___ 

Eucommia montana (cf.)___ 

Ulmus knowltoni ____________

Ulmus owyheensis ......... _ Ulmus speciosa ..............

Zelkova browni ..............

Zelkova ungeri ...............

Alnus cappsi .................

Alnus fairi....................

Alnus healyensis ............

Betula sublutea (cf.) _____

Ostrya oregoniana (cf.) __ 

Fagus antipofi ...............

Carya bendirei ...............

Cyclocarya ezoana ...........

Pterocarya nigella __.___.....

Populus kenaiana ._.____.._.

Tilia subnobilis .............

Cladrastis aniensis (cf.) ... 

Pueraria miothunbergiana 

Hemitrapa borealis .........

Acer glabroides .............

Acer heterodentatum .......

Acer ezoanum ...............

Nyssa knowltoni (cf.) ___ 

Alangium mikii .............

Kalopanax n-suzuki ________

Total .................. 21 29 13

irrespective of the latitudes at which the species are 

found.

FLORISTIC COMPOSITION

In the following list, the classification of the dicotyledons is that of Takhtajan (1969): 

Pterophyta 

Filicinae 

Filicidae 

Aspidiales 

Aspidiaceae 

Dryopteris sp.

Onoclea sensibilis Linnaeus 

Coniferophyta 

Coniferae 

Coniferidae 

Coniferales 

Taxodiaceae

Glyptostrobus europaeus (Brongniart) Heer 

Metasequoia cf. M. glyptostroboides

Hu et Cheng 

Ginkgoidae 

Ginkgoales 

Ginkgoaceae

Ginkgo biloba Linnaeus
10 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Magnoliophyta (Angiospermae) 

Magnoliatae (Dicotyledones) 

Magnoliidae 

Magnolianae 

Nymphaeales 

Nymphaeaceae

Nymphar ebae (Huzioka) Ozaki 

Ranunculidae 

Ranunculanae 

Ranunculales 

Menispermaceae

Cocculus auriculata (Heer) Wolfe 

Hamamelididae 

Hamamelidanae 

Cercidiphyllales 

Cercidiphyllaceae 

Cercidiphyllum alaskanum Wolfe et

Tanai

Hamamelidales 

Hamamelidaceae

Liquidambar pachyphylla Knowlton 

Platanaceae

Platanus bendirei (Lesquereux) Wolfe 

Eucommiales 

Eucommiaceae

Eucommia cf. E. montana R. W. Brown 

Urticales

Ulmaceae 

Celtis sp.

Ulmus knowltoni Tanai et Wolfe 

Ulmus owyheensis H. V. Smith 

Ulmus speciosa Newberry 

Ulmus sp.

Zelkova browni Tanai et Wolfe 

Zelkoua ungeri Kovats 

Fagales 

Fagaceae

Fagus antipofi Heer

Fagus aff. F. crenata Blume

Quercus furuhjelmi Heer

Betulales 

Betulaceae

Alnus cappsi (Hollick) Wolfe 

Alnus fairi (Knowlton) Wolfe 

Alnus healyensis Wolfe 

Betula cf. B. sublutea Tanai et Suzuki 

Carpinus seldoviana Wolfe 

Corylus sp.

Ostrya cf. O. oregoniana Chaney 

Juglandales 

Juglandaceae

Gary a bendirei (Lesquereux) Chaney et 

Axelrod

Cyclocarya ezoana (Tanai et Suzuki)

Wolfe et Tanai

Pterocarya nigella (Heer) Wolfe 

Dilleniidae 

Dillenianae 

Salicales 

Salicaceae

Populus kenaiana Wolfe 

Populus sp. 

Salix cappsensis Wolfe 

Salix hopkinsi Wolfe et Tanai 

Salix picroides (Heer) Wolfe 

Salix seldoviana Wolfe et Tanai 

Malvanae 

Malvales 

Tiliaceae

Tilia subnobilis Huzioka 

Rosidae 

Rosanae 

Saxifragales 

Hydrangeaceae

Hydrangea sp. 

Resales 

Rosaceae

Crataegus chamisonii (Heer) Wolfe

et Tanai

Prunus kenaica Wolfe et Tanai 

Prunus aff. P. padus Linnaeus 

Sorbaria hopkinsi (Wolfe) Wolfe

et Tanai 

Fabales

Leguminosae

Cladrastis cf. C. aniensis Huzioka 

Pueraria miothunbergiana Hu

et Chaney 

Myrtanae 

Myrtales 

Lythraceae

Decodon alaskana Wolfe et Tanai 

Trapaceae

Hemitrapa borealiis (Heer) Miki 

Rutanae 

Sapindales 

Aceraceae

Acer ezoanum Oishi et Huzioka 

Acer glabroides R. W. Brown 

Acer grahamensis Knowlton et Cockerell 

Acer heterodentatum (Chaney)

MacGinitie 

Aralianae 

Cornales 

Nyssaceae

Nyssa cf. N. knowltoni Berry 

Alangiaceae 

Alangium mikii Wolfe et Tanai
VEGETATION 11

Araliaceae

Kalopanax n-suzuki Wolfe et Tanai 

Celastranae 

Rhamnales 

Vitidaceae

Vitis seldoviana Wolfe et Tanai 

Oleales 

Oleaceae

Fraxinus kenaica Wolfe et Tanai 

Asteridae 

Lamianae 

Dipsacales 

Caprifoliaceae 

Lonicera sp.

Liliatae (Monocotyledones) 

Alismidae 

Alismanae 

Alismatales 

Alismataceae 

Alisma seldoviana Wolfe et

Tanai 

Najadales 

Potamogetonaceae 

Potamogeton alaskanus Wolfe et

Tanai 

Incertae Sedis

Monocotylophyllum alaskanum

(Heer) Wolfe et Tanai 

Monocotylophyllum spp.

The Seldovia Point flora is composed primarily of 

members of subclass Hamamelididae (23 species), and 

particularly the evolutionarily more advanced families such as Ulmaceae and Betulaceae. Except for 

the specialized aquatic Nymphar, Magnoliidae are absent. Also absent or poorly represented are orders considered to be primitive in the Dilleniidae (for example, 

Theales and Dilleniales) and the Rosidae (Saxifragales). Within the orders of Dilleniidae and Rosidae, 

the Seldovia Point species typically belong to the more 

advanced families in the respective orders, for example Tiliaceae, Trapaceae, Aceraceae, Alangiaceae, 

Araliaceae. Evolutionarily, therefore, the Seldovia 

Point flora is highly advanced.

A comparison with the Eocene Copper Basin flora of 

northern Nevada (Axelrod, 1966) is informative. Although the Copper Basin assemblage grew under 

more equable conditions than did the Seldovia Point 

assemblage (see section on "Paleoclimatology"), both 

assemblages represent cool temperate (microthermal) 

vegetation. In regard to the dicotyledonous flora at the 

specific level, the representation of most subclasses is 

distinctly different in the two assemblages:

Subclass Copper Basin Seldovia Point 

(percent) (percent)

Magnoliidae _. 

Ranunculidae 

Hamamelididae 

Dilleniidae __ 

Rosidae ____. 

Asteridae __

4

11

14

14

57

0

2

2

43

15

36

2

Despite having a number of families in common, the 

Copper Basin flora is less advanced than the Seldovia 

Point flora. One of the most notable differences is in 

the low diversity of Hamamelididae in the Copper 

Basin flora, a diversity that is just as low at the 

generic level as at the specific. Although generic diversification in the Hamamelididae had been largely 

completed during the Eocene Epoch (Wolfe, 1973), it is 

apparent that (1) many of the genera that now include 

microthermal members had not yet invaded microthermal climates in the Eocene and (2) specific diversity within the genera that had entered microthermal climates was low. It is thus evident that 

microthermal forests have undergone major changes 

in floristic composition.

VEGETATION

MEGAFOSSILS

All the species in the Seldovia Point flora that are 

thought to represent trees, shrubs, or vines were 

probably deciduous. Of the vegetational types described by Wang (1961) for eastern Asia, only one 

type—the Mixed Northern Hardwood forest— 

combines the exclusively broad-leaved deciduous 

habit with a diversity similar to that of the Seldovia 

Point assemblage. From the standpoint of foliar 

physiognomy the Seldovia Point assemblage also 

compares well with the Mixed Northern Hardwood 

forest. Only 16 percent of the Seldovia Point species 

have entire-margined leaves, and in Mixed Northern 

Hardwood forest the percentage ranges from 9 to 24 

(Wolfe, 1979a). Of the various species listed'by Wang 

(1961) as occurring in this forest, almost 20 percent 

have palmately lobed leaves in comparison with 16 

percent for the Seldovia Point assemblage.

In floristic composition as well the Seldovia Point 

assemblage resembles Mixed Northern Hardwood 

forest, although there are some notable differences. Of 

the 18 genera listed by Wang (1961, p. 76) as composing the tree stratum of the Mixed Northern Hardwood 

forest in the northeastern provinces of China, 13 are 

represented in the Seldovia Point flora. Other Seldovia Point tree genera (table 3; Cercidiphyllum, 

Pterocarya, Fagus) occur in the Mixed Northern 

Hardwood forest (Fagus zone of many Japanese
12 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

TABLE 3.—Assumed growth habits of Seldovia Point species

Trees

Cercidiphyllum alaskanum 

Liquidambar pachyphylla 

Platanus bendirei 

Eucommia montana 

Ulmus knowltoni 

Ulmus owyheensis 

Ulmus speciosa 

Zelkova browni 

Zelkova ungeri 

Carya bendirei 

Cyclocarya ezoana 

Pterocarya nigella 

Alnus fairi 

Alnus healyensis 

Betula cf. B. sublutea 

Carpinus seldoviana 

Ostrya cf. O. oregoniana 

Fagus antipofi 

Fagus aff. F. crenata 

Quercus furuhjelmi 

Populus kenaiana 

Populus sp. 

Tilia subnobilis 

Prunus kenaica 

Cladrastis cf. aniensis 

Acer ezoanum 

Acer glabroides 

Acer grahamensis 

Acer heterodentatum 

Nyssa cf. N. knowltoni 

Kalopanax n-suzukii

Fraxinus kenaica 

Glyptostrobus europaeus 

Metasequoia cf. M. 

glyptostroboides 

Ginkgo bibloba

Shrubs

Celtis sp. 

Alnus cappsensis 

Corylus sp. 

Salix cappsensis 

Salix hopkinsi 

Salix picroides 

Salix seldoviana 

Hydrangea sp. 

Crataegus chamisonii 

Sorbaria hopkinsi 

Decodon alaskana 

Alangium mikii 

Lonicera sp.

Vines

Cocculus auriculata 

Pueraria miothunbergiana 

Vitis seldoviana

Terrestrial herbs

Dryopteris? sp. 

Onoclea sensibilis 

Monocotylophyllum spp.

Aquatic herbs

Alisma seldoviana 

Potamogeton alaskana 

Nymphar ebae 

Hemitrapa borealis

TABLE 4. — Distribution of most closely related extant species in 

vegetational types in eastern Asia

[Occurrence enclosed in parentheses indicate that the extant 

Asian species may only be distantly related to Seldovia Point 

species]

Broad- Mixed Broad- Mixed 

leaved Meso- leaved Northern 

Ever- phytic Deciduous Hardwood 

green

Carya tonkinensis

Corylus spp

Ostrva spp

Sorbaria hndlevana

Pueraria thunbergiana

Acer henryu

Alangium chmensis

Fraxinus hopeiensis

X

X?

X

XXX 

X X

X

X

---(X) 1X1 (X) (X)

X X

x

X 

XX 

X

X

XXX 

X

X

- ( X IX) (X ) (X)

_ _ ( x (xl (xi ( x )

-. (X (x) (xl (x)

XX X

- (X (XI ( X ) X

- -(X (XI (X) (X)

X

(X) (X) (X) (XI

( X I (XI (X)

y

X

X

xx x

<

X

(X| (X I

XXX 

< X X

X

(xi ( x I (xl (x)

i1 X

( < ) 1 < 1 (X) (X)

.. — -13(101 13(11) 8(10i 12(9)

botanists) in Japan (Kara, 1959). Further, Wang 

(1961, p. 77) notes that the shrubs in the Mixed 

Northern Hardwood forest belong to genera such as 

Corylus, Crataegus, Sorbaria, and Lonicera, and vines 

(including Vitis) are present.

If only floristic associations were relied upon to determine vegetational (and hence climatic) type, the 

Seldovia Point assemblage would be considered as 

most probably representing Mixed Mesophytic forest, 

a conclusion arrived at previously (Wolfe, 1966; Wolfe 

and Leopold, 1967). The majority of the most closely 

related extant species are found in this vegetational 

type (table 4), and several Seldovia Point genera occur 

only in Mixed Mesophytic forest or in broad-leaved 

evergreen forests in eastern Asia: Ginkgo, Metasequoia, Glyptostrobus, Cocculus, Liquidambar, 

Carya, Cyclocarya, Cladrastis, and Nyssa. In contrast, 

of all the Seldovia Point genera, only Sorbaria is presently restricted to cool temperate forests such as the 

Mixed Northern Hardwood. The physiognomic data, 

however, clearly indicate that the Seldovia Point assemblage represents Mixed Northern Hardwood 

forest. The lack of a closer floristic resemblance between the Seldovia Point assemblage and the extant

Mixed Northern Hardwood forest of eastern Asia can 

be explained in terms of the historical development of 

this forest and the related Mixed Mesophytic forest. 

As well, the assumptions commonly made concerning 

present distribution of taxa relative to climate (and 

hence vegetational type) may not be valid.

The inclusion in the Seldovia Point flora of Ginkgo, 

Metasequoia, Glyptostrobus, Cocculus, Liquidambar, 

Platanus, Carya, Cyclocarya, Cladrastis, Nyssa, and 

Alangium indicates a deceptively warm climate if the 

present distributions of these genera are considered. 

The resistance to freezing of extant species of many of 

these genera has, however, been analyzed (Sakai, 

1971, 1972; Sakai and Weiser, 1973). Following are 

listed the results of these investigations in regard to 

some of the Seldovia Point genera (the temperatures 

are the highest of the freezing resistances of bud, leaf, 

or twig):

Ginkgo biloba,-------------------------- -30°C

Glyptostrobus pensilis __________________18°

Metasequoia glyptostroboides _____________ 30°

Platanus occidentalis __________________ —20°

Liquidambar styraciflua _____________ _25°

Liquidambar formosana _______________-17°

Pterocarya rhoifolia ________________^__—30°

Nyssa sylvatica ______________________ — 30°

Sakai's data also illustrate the wide variation of 

freezing resistance from one species to another con-
VEGETATION 13

generic species. Again the highest of the three temperatures are selected:

Acer macrophyllum __________________ -20°C

A. mono _----_________________________-250

A. rubrum _______________________-30°

A. saccharum _______________________ — 40°

Quercus garryana _____________________15°

Q. lyrata ____________________________-20°

Q. mongolica ______________________-30°

Q. macrocarpa ______________________40°

Even within the same species, different geographic 

races may exhibit considerably different tolerances to 

freezing (Sakai and Weiser, 1973). We emphasize that 

the present geographic area occupied by a species may 

have little or no bearing on the paleoecologic significance of that species, its ancestors, or other extinct 

congeneric lineages.

Wang (1961, p. 239-246) has noted that today the 

Mixed Northern Hardwood forest of eastern Asia, although containing many genera in common with the 

Mixed Mesophytic forest, has few species in common 

with that forest. Despite this specific differentiation, 

Wang hypothesized that most species now in Mixed 

Northern Hardwood forest were derived from Mixed 

Mesophytic lineages during the Tertiary and that, 

prior to glaciation, Mixed Northern Hardwood forest 

had more genera and species than now.

That the Mixed Northern Hardwood forest was considerably richer in the past is fully attested to by the 

Seldovia Point flora, as well as by assemblages representing the same vegetational type in eastern Asia. 

Numerous genera are present in these assemblages 

that no longer participate in Mixed Northern 

Hardwood forest. This is valid in regard to the east 

Asian Miocene Mixed Northern Hardwood forest (for 

example, Alangium, Cyclocarya, Liquidambar), which 

is the lineal ancestor of the present Mixed Northern 

Hardwood forest.

Wang's (1961) hypothesis concerning the origin of 

Mixed Northern Harwood lineages is, however, valid 

only in regard to the lineages of the West American 

element (see p. 17) in the Seldovia Point floralineages that typically did not become part of the 

Mixed Northern Hardwood forest in eastern Asia. In 

the Pacific Northwest, several early and middle 

Miocene assemblages represent Mixed Mesophytic 

forest (for example, Latah and equivalents in 

Washington and adjacent parts of British Columbia 

and Idaho, Collawash in Oregon), and these assemblages have several species in common with the 

Seldovia Point assemblage (table 5).

In eastern Asia, however, Mixed Mesophytic forest 

is probably of a more recent origin than Mixed Northern Hardwood forest. The early Miocene Aniai-type

TABLE 5.—Seldovian dicotyledonous species also occurring in Mixed 

Mesophytic forest in the early and middle Miocene of the Pacific 

Northwest

Seldoi'ia Point

Cocculus auriculata 

Liquidambar pachyphylla 

Platanus bendirei 

Ulmus knowltoni 

Ulmus owyheensis 

Ulmus speciosa 

Zelkova browni 

Alnus fairi 

Alnus healyensis 

Ostrya oregoniana 

Carya bendirei 

Pterocarya nigella 

Populus kenaiana 

Acer glabroides 

Acer macrophyllum type 

Acer heterodentatum 

Nyssa knowltoni

Other Seldovian localities

Acer aff. A. pennsylvanicum 

Acer aff. A. saccharinum

floras (Tanai, 1961) all represent broad-leaved deciduous forest without broad-leaved evergreens; that is, 

this type of forest is Mixed Northern Hardwood. This 

is true of assemblages from Kyushu and western Honshu, although admittedly these assemblages are small 

and poorly known. As discussed in the section on 

paleoclimatology, the vegetational zonation during 

the early and middle Miocene excluded Mixed 

Mesophytic forest, but Mixed Northern Hardwood 

forest occupied a broad area north of latitude 44° N. 

The first evidence in eastern Asia of Mixed Mesophytic forest is in the late Miocene. The Mitoku flora 

from western Honshu (Tanai and Onoe, 1961) is 

dominantly broad-leaved deciduous but also contains 

some notophyllous broad-leaved evergreens; this assemblage thus probably represents Mixed Mesophytic 

forest. As noted by Tanai and Onoe (1961, p. 15), some 

of the Mitoku species are also found in the Aniai-type 

assemblages, which would indicate the reverse of 

what Wang (1961) suggested, that is, that s.ome Mixed 

Mesophytic lineages were derived from Mixed Northern Hardwood lineages.

Some of the Mitoku species also occur in Japanese 

middle Miocene assemblages that we interpret as representing Wang's (1961) Deciduous Broad-leaved 

forest. Such a pattern of derivation of some Mixed 

Mesophytic lineages is to be expected as the climate in 

eastern Asia changed from one favorable to the development of this forest to one favorable to the development of Mixed Mesophytic forest (see p. 19). The 

third major source for Mixed Mesophytic lineages has 

been from Wang's Sclerophyllous Broad-leaved Evergreen forest, and these lineages include both broadleaved evergreen and deciduous plants (as well as 

conifers such as Cunninghamia}. Indeed, today the 

Mixed Mesophytic and Sclerophyllous Broad-leaved
14 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Evergreen forests of China share many species.

Assuming the validity of our paleoclimatological 

suggestions (p. 19), most species in the Mixed Northern Hardwood forest of eastern Asia were derived 

primarily from lineages of Deciduous Broad-leaved 

forest, although the ultimate origin of the lineages 

must be in broad-leaved evergreen vegetation; that is, 

in an area such as Japan, the probable sequence during the Oligocene climatic deterioration would have 

been from broad-leaved evergreen forest into Wang's 

(1961) lower oak forest, then into Deciduous Broadleaved forest, and finally into Mixed Northern 

Hardwood forest. At least part of such a sequence has 

been demonstrated in the Oligocene of Hokkaido 

(Tanai, 1970). Lineages that have followed such a pattern are exemplified by Acer kushiroanum-A. ezoanum-A. miyabei (Tanai, 1972). This pattern is 

also indicated by the large number of species common 

to the Yoshioka flora (Decidous Broad-leaved forest) 

and the isochronous Mixed Northern Hardwood assemblages such as the Abura (Tanai and Suzuki, 

1963) and Upper Dui (Fotianova, 1967).

POLLEN

Only three samples from the Seldovia Point 

localities have been analyzed for pollen. The samples 

from localities 9856 and 9858 are poor, and only 67 

and 100 grains, respectively, were counted in these 

two samples. Locality 9857, however, produced an 

abundant and well-preserved pollen assemblage, and 

the percentages (table 6) are based on 300 grains.

The pollen assemblage from locality 9858 at Seldovia Point is dominated by broad-leaved plants. It is 

unknown whether the pollen of Alnus came from trees 

or shrubs or both, but the probable broad-leaved tree 

pollen (Juglandaceae, Betulaceae other than Alnus, 

Fagaceae, Liquidambar} in sample 9858 represents 20 

percent of the total in contrast to only 6 percent pollen 

of Pinaceae.

The samples from 9856 and 9857, however, have a 

very small broad-leaved element. Members of 

Pinaceae account for over half the pollen in both samples. The abundance of Picea and Tsuga is particularly significant and gives strong evidence that the 

forest in this area was dominantly coniferous. Assuming that the assemblages from the three localities are 

approximately isochronous, the data indicate that the 

area around Seldovia was forested primarily by 

broad-leaved trees but that in the area of Port 

Graham the forest was primarily needle leaved, with 

broad-leaved vegetation confined only to the small 

valleys. This, in turn, indicates that in the more 

equable coastal area of southern Alaska, the vegetaTABLE 6.—Percentages of pollen types in samples from the 

Seldovia Point beds

[x, present but not included in counts]

Pollen type

Locality

9857 9856

Aff. Cedrus ___________________________

Picea ____ _ _ _ _ _ _ ____

Tsuga ___ _____ ___ ___ ____ ________ __

Taxodiaceae/Cupressaceae _ ____

Salix _ _

Carya

Juglans ___ ____ ______ ________

Pteroc aryalCyc locar ya

Alnus

Betula _______ _________ ______ ______

OstryalCarpinus __ _____

Fagus

Ulmus/Zelkova

Liquidambar

Ilex _ _ __ _____

Tilia

Ericales

Lonicera

Indet. dicotyledons _____ _________

Typha _ ______ ______ ____ _____

Indet. monocotyledons

Osmunda

?1

________ 4

_______ 1

______ 1

x

x

________ 45

_______ 2

x

_______ 4

________ 14

_______ 1

_______ 12

15

x

x

34

1

1

16

8

x

1

X

1

30

1

3

2

1

x

x

1

X

X

X

33

3

4

22

1

4

1

6

6

3

1

3

x

1

9

x

tion was probably dominantly coniferous.

The lack of Pinaceae in our megafossil collections is 

apparently somewhat contradictory. The largest collections, however, were made at locality 9858, and the 

pollen data indicate that Pinaceae were far distant 

from this locality. In some instances, however, we 

think that megafossil assemblages can yield a highly 

distorted picture of the actual vegetation. One example is the assemblages from the Homerian (late 

Miocene) part of the Kenai Group. Large collections 

made in 1962 from the Homerian contained no 

megafossils of Pinaceae (Wolfe and others, 1966), but 

the pollen assemblages are dominated by Pinaceae 

(Wolfe, 1969b) and were interpreted as indicating the 

presence of a coniferous forest throughout the Cook 

Inlet area during the late Miocene (Wolfe and 

Leopold, 1967). Two new localities, first collected in 

1967, did produce a diverse and abundant pinaceous 

flora, thus substantiating the earlier interpretation. A 

second example is the late Miocene Hidden Lake assemblage from Oregon (Wolfe, 1969b). Although pollen of Picea accounts for almost half the pollen, 

megafossils of Picea represent less than one percent of 

over 2,000 specimens counted; the association of some 

18 species of conifers, however, leaves no reasonable 

doubt that the vegetation was dominantly coniferous 

despite the low representation of conifers in the 

megafossil count. We conclude, therefore, that the pollen data indicate the dominance of conifers in the Port 

Graham area during the early and middle Miocene.
VEGETATION 15

DISTRIBUTION

As discussed previously, the coastal area of southern Alaska was probably occupied by coniferous 

forest. The early and middle Miocene megafossil assemblages of that area are unknown, so a comparison 

with the Seldovia Point flora is impossible. In the 

Wrangell Mountains (fig. 7), however, the late Seldovian assemblage represents coniferous forest, attested 

to by both the dominance of Pinaceae in the pollen 

assemblage and the diversity of this family in the 

megafossil assemblage (Wolfe, 1972). The broadleaved adjuncts to this coniferous forest included 

Populus, Salix, Pterocarya, Alnus, Betula, Fagus, Ulmus, Acer (both macrophyllum and saccharinum 

types), and Lonicera. Presumably the composition of 

the coastal coniferous forest was similar.

A small assemblage at the northern margin of the 

Kenai basin was collected from beds of the Kenai 

Group. Because these thin beds on Cache Creek include both Seldovian as well as Homerian floras 

(Wolfe and others, 1966), the Seldovian beds there are 

most probably of late Seldovian age. The Cache creek 

assemblage includes:

Metasequoia cf. M. glyptostroboides

Cocculus auriculata

Cercidiphyllum alaskanum

Fagus antipofi

TOO 200 300 400 KILOMETERS

FIGURE 7.—Suggested distribution of vegetational types in Alaska 

during the late early and early middle Miocene. Approximate extent of land shown by heavy solid line.

Quercus furuhjelmi

Alnus cappsi

Populus kenaiana

Salix spp.

Acer aff. A. rubrum

This assemblage is consistent with the occurrence of 

Mixed Northern Hardwood forest both to the south 

(Seldovia Point) and north (Sanctuary and Suntrana) 

of Cache Creek during the late Seldovian.

The Mixed Northern Hardwood forest extended 

north at least to the central Alaska Range. In the 

Sanctuary and Suntrana Formations, the following 

have been found:

Metasequoia sp.

Cocculus auriculata

Ulmus sp.

Alnus sp.

Fagus antipofi

Quercus furhjelmi

Carya bendirei

Cyclocarya ezoana

Pterocarya nigella

Populus kenaiana

Populus spp.

Salix spp.

Alangium mikii

Additional broad-leaved genera from E. B. Leopold's 

(1969, unpub. data) pollen work include Liquidambar, 

Platanus, Juglans, Eucommia?, Betula, Corylus, 

Ostrya/Carpinus, Ericales, Malvaceae, Tilia, 

Rosaceae, Acer, Ilex, Eleagnus, Nyssa, Compositae, 

and Caprifoliaceae (Lonicera type). The SanctuarySuntrana assemblage is similar to the Seldovia Point 

assemblage from the standpoint of both megafossils 

and microfossils. We emphasize, however, that 

warmer elements are fewer in number than in the 

Seldovia Point assemblage, and in all probability the 

Sanctuary-Suntrana climate had a lower mean annual temperature and a higher mean annual range of 

temperature.

The northern limit of Mixed Northern Hardwood 

forest in Alaska during the late Seldovian is not 

known with certainty. A few of the SanctuarySuntrana pollen samples are dominated by Picea, thus 

indicating that the coniferous forest was probably not 

far to the north. This would place the boundary between the two vesetational types at about lat 64° N. 

This placement is consistent with the pollen assemblage from the Noxapaga Formation of Sainsbury 

(1974) on Seward Peninsula (lat 65° N). The two samples examined are dominated by Pinaceae (Picea, 

Pinus, Tsuga, Abies, and Larix/Pseudotsuga, in order 

of decreasing abundance). Except for Betulaceae, none 

of the broad-leaved genera exceeds one percent; this
16 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

element is, however, moderately diverse and includes 

Fagus, Quercus, Carya, PterocaryalCyclocarya, Ulmus, 

Populus, Salix, and Symphoricarpos. Farther north, 

the Nuwok pollen assemblage of middle Miocene 

age (lat 70° N) is almost totally coniferous but has 

minor amounts of PterocaryalCyclocarya and Ulmus.

The coastal coniferous forest during the early 

Miocene was isolated from other coniferous forest. 

To the southeast, the coniferous forest theoretically 

gave way to Mixed Northern Hardwood or Mixed 

Mesophytic forest. To the southwest, coniferous forest 

was clearly bounded by Mixed Northern Hardwood 

forest, which has been recorded from Kamchatka 

(Chelebaeva, 1968). The northern area of coniferous 

forest probably had a great longitudinal extent. 

Neogene pollen assemblages from northeastern 

Siberia (Baranova and others, (1968) are dominated 

by Pinaceae; it should be noted, however, that all 

these assemblages have not been independently dated, 

and their exact placements within the Neogene sequence are not certainly known.

COMMUNITIES

Counts of the leaf impressions in terms of abundance of individual species were not made; such 

counts have not yet been proved reliable in indicating 

the actual abundance of plants in the original vegetation. The most abundant dicotyledonous tree species 

in our collections from the Seldovia Point locality are:

Fagus antipofi

Acer ezoanum

Zelkova spp.

Alnus spp.

Salix spp.

Platanus bendirei

Quercus furuhjelmi

Carpinus seldoviana

The above species may well represent the dominant 

forest species in the vicinity of Seldovia during the 

early and middle Miocene. We thus envisage the main 

forest community to have been an association ofFagusAcer-Zelkova-Quercus-Carpinus-Ulmus. Near Port 

Graham, the forest association was somewhat different; there, Fagus and Ulmus are the most common 

megafossils, Carpinus is absent, and apparently Acer 

ezoanum was replaced by A. grahamensis.

Some of the Seldovia Point species can very clearly 

be relegated to a streamside element. This assignment 

is supported not only by the habitats of extant relatives but also—and perhaps most significantly—by 

the abundance of leaves of these species in association 

on the same slabs. Such species include Salix hopkinsi, S. cappsensis, S. picroides, S. seldoviana, 

Platanus bendirei, Decodon alaskana, and Alnus 

cappsi.

The Seldovia Point flora has a strong aquatic element at locality 9858. Leaves of the extinct Nymphar 

are particularly common, as are seeds and, to a lesser 

degree, leaves of Potamogeton. Nuphar (a genus related to Nymphar) and Potamogeton today typically 

inhabit quiet water, and Nuphar typically lives in 

lakes.

FLORISTICS

Traditionally the species composing a fossil flora 

have been placed into elements, each of which either 

represents the geographic region where the most 

closely related extant species are found (for example, 

East Asian element, East American element) or represents the region where the lineage to which the 

species belongs putatively originated (for example, 

Arcto-Tertiary element, Madro-Tertiary element). 

Both of these approaches to floristic analysis have 

some validity, despite the fact that particular concepts 

such as that of the Arcto-Tertiary element have probably been invalidly applied (Wolfe, 1969b, 1972; 

Tanai, 1971). The areas of origin of the floristic element in the Seldovia Point flora are particularly significant to concepts of floristic history because the 

flora from the Port Graham locality has been one of 

the fundamental bases for the concept of an ArctoTertiary Geoflora, which, in turn, has served as the 

"type" geoflora for the geofloral concept.

We emphasize that the assignment to elements such 

as East Asian or West American by Chaney (for 

example, 1959) and his followers was done primarily 

for paleoclimatic purposes. That is, in west American 

Tertiary floras, a dominant East American-East Asian 

element indicated abundant summer precipitation, 

whereas a dominant West American element indicated little summer precipitation. This usage of "element" has little floristic value and should be abandoned.

To place species such as Liquidambar pachyphylla 

in an "East American" element is, in view of the history of this lineage, unwarranted. There is no direct 

relationship between L. pachyphylla and the extant 

east American L. styraciflua, although the two species 

may belong to lineages that were derived from a 

common stock in the Paleogene. L. pachyphylla is a 

distinctive member of a middle latitude, West American element. Similarly, to the same element belongs 

the type of Cyclocarya found in the Tertiary of the 

Pacific Northwest, a type that is only distantly related 

to the extant Asian C. paliurus. Despite the fact that 

the lineages to which L. pachyphylla and the west 

American Cyclocarya belong are now extinct, these 

lineages have the same floristic significance as the 

Platanus bendirei lineage, which has survived in
FLORISTICS 17

western North America as P. racemosa. Indeed, as 

pointed out previously (Wolfe, 1969b), several 

"Madro-Tertiary" lineages are in fact members of this 

same element.

Prior to the Oligocene deterioration of climate 

(Wolfe and Hopkins, 1967; Wolfe, 1971, 1978), the 

climates of the Paleogene were characterized by high 

equability (that is, a low mean annual range of temperature). Just as temperate broad-leaved deciduous 

forests are today absent in the highly equable Southern Hemisphere temperate regions, it is highly probable that temperate broad-leaved deciduous forests 

were absent in the Northern Hemisphere during the 

Eocene and the Oligocene prior to the deterioration 

(Wolfe, 1978). The Oligocene deterioration was thus 

characterized not only by a dramatic decline in mean 

annual temperature but also by a major increase in 

mean annual range of temperature (Wolfe, 1971). 

Broad-leaved deciduous forests appeared in a geologically short period of time throughout middle and high 

latitudes of the Northern Hemisphere. The lineages 

that composed these forests were derived from two 

major sources (Wolfe, 1972): (1) lineages that were present in broad-leaved evergreen forest prior to the 

deterioration and that were preadapted (or had the 

genetic capability to adapt rapidly) to climates 

characterizing broad-leaved deciduous forest and (2) 

lineages that were present in temperate coniferous 

forests and that moved downslope during the 

Oligocene deterioration. During the later Oligocene 

and Miocene some lineages from broad-leaved evergreen forest also adapted gradually and moved into 

the broad-leaved deciduous forests (Wolfe, 1972).

Terms such as East Asian Element can thus be denned relative to the geographic region in which it is 

thought a given lineage entered broad-leaved deciduous forest for the first time. Species such as Liquidambar pachyphylla and Carya bendirei, although belonging to genera that are no longer native to western 

North America, are consequently considered to belong 

to the West American Element, because the lineages 

to which these species belong were present in western 

North America at middle latitudes prior to the 

Oligocene deterioration.

The significance of the concept of this west American, mid-latitude element is considerable to paleoecological reconstructions. Just as the tolerances of 

Carya in eastern Asia are, at least in part, different 

from the tolerances of Carya in eastern North America, so it can be expected that the tolerances of the 

now extinct west American Carya were also different. 

This, we think, is at least partly the explanation for 

the presence of Liquidambar in the Seldovia Point assemblage; it is also notable that Liquidambar was a 

conspicuous member of fluviatile vegetation in the

late Miocene upland coniferous forest in the Pacific 

Northwest, but today none of the surviving members 

of the genus are associated with Temperate Coniferous forest.

The Seldovia Point flora has three known floristic 

elements: East Asian, West American, and Beringian. 

The Beringian element is composed of those lineages 

that are thought to have entered into the temperate 

forests in Alaska and adjacent high-latitude regions. 

The basis for the assignment to the various floristic 

elements (table 7) is given in the discussions of the 

systematic treatments of the various species.

Assignment to the various floristic elements is well 

documented in instances such as Platanus bendirei 

and less well documented in other instances. It is assumed that if a particular lineage appears in a given 

area immediately following the Oligocene deterioration, that lineage is assigned to that area's element.

TABLE 7.—Analysis of the Seldovia Point assemblage in terms of 

floristic elements

Species East West 

Asian American Beringian Unknown

Dryopteris! sp _____________________

Onoclea sensibihs

Glyptostrobus europaeus ________ x

Metasequoia cf. M. 

glyptostroboides ____________________ x

Ginkgo biloba

Nymphar ebeae

Cocculus auriculata _________________ x

Cercidiphyllum alaskanum _____ ______ x

Liquidambar pachyphylla __ __________ x

Platanus bendirei ____________________ >

Eucommia cf. E. montana _ ________ x?

Celtis sp ____________________

Ulmus knowltoni ________________ _____ x

Ulmus owyheensis ____________________ x

Ulmus speciosa __ ____________ »

Zelkova browni ______________________ x

Zelkova ungeri ___ ______________ x

Fagus antipofi _______________________ x

Fagus aff. F. crenata __ ____________ >

Quercus furuhjelmi ______ ____________ x?

Alnus cappsi ________________________ >

Alnus fairi _________________________ x

Alnus healyengis _____________________ x

Betula cf. B. sublutea __________ x? 

Carpmus seldoviana _________________ *

CoryluK sp ___________________________

Ostrya cf. O. oregoniana ___________ x

Carya bendirei ______________________ x

Cyclocarya ezoana ____________________ x

Pterocarya nigella ____________________ x

Populus kenaiana ____________________ >

Populus sp ______________________

Salix cappsensis ____________________ >

Salix hopkinsi _______________________ *

Sahx picroides _______________________ > 

Salix seldoviana ___________________ /

Tilia subnobilis ______________________ x

Hydrangea sp ______________

Crataegus charnisonn

Prunus kenaica ____________________

Prunus sp _________________________

Sorbaria hopkinsi ___________________ >

Cladrastis cf. C. aniensif, ____________ *

Pueraria miothunbergiana ___________ <

Decodon alaskana ___________________ *

Hemitrapa borealis ___________________ *

Acer ezoanum ________________________ '

Acer grahamensis ____________________ x

Acerheterodentatum _________________ x

Nyssa cf. N knowltoni ___________ *

Alangium miku ______________________ «"

Kalopanax n-suzukii ____________ _____ *

Vitis seldoviana ______________________ *

Fraxinus kenaica __________________

Lonicera sp

Potamogeton alaskana _______________

Monocotylophyllum alaskanum ________

Monocotylophyllum tenuistriatus ______

Total _________________________ 14 16 12 Iff
18 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

For example, Metasequoia is not known in the Alaskan Eocene, but the early Angoonian assemblages 

immediately following the deterioration contain 

abundant Metasequoia, which therefore is assigned to 

the Beringian element; this genus could equally well 

be assigned to the other two elements, but there is no 

need to invoke immigration from another region.

The West American element has suffered far more 

extinction than the remaining two elements. Only 

three West American lineages in the Seldovia Point 

flora still survive: Platanus bendirei (P. racemosa), 

Acer heterodentatum (A. negundo), and A. grahamensis (A. macrophyllum). Of the East Asian element, 

only Cercidiphyllum alaskanum and Decodon alaskana have definitely become extinct. Of the Beringian 

element, possibly many lineages yet survive, particularly in Alnus and Salix.

Today, many boreal species extend southward, particularly in North America, along mountain chains. 

Similarly, during the late Miocene many species are 

known from lowland Alaska to the uplands of the 

Pacific Northwest to the south (Wolfe, 1969b). None of 

these species are, however, members of the East Asian 

element as defined in this report; although not yet 

known in Alaska, Acer scottiae must have had such a 

distribution and would clearly have been East Asian. 

Such data emphasize the difficulty of floristic interchange between midlatitude forests on either side of 

the Pacific during the Neogene, at least in regard to 

woody plants.

The few West American lineages that probably 

migrated into eastern Asia during the Miocene are:

Pterocarya rhoifolia type.—Earliest records of this 

type are in the middle and late Oligocene of Alaska 

(Tsadaka Formation). This type appears in the early 

Miocene of Japan and Oregon and persisted in the 

Pacific Northwest through at least the late Miocene. 

The type of foliage with which the seeds are most frequently associated is that assigned to P. nigella, 

which is thought to be descended from the west American P. pugetensis.

Acer negundo type.—This type extends in North 

America back to at least the early Oligocene (MacGinitie, 1953) and occurs in many younger assemblages. The first record in Alaska is in the Seldovia Point flora, but the occurrence of this group in 

Japan in the early to middle Miocene (Tanai, 1961) 

indicates that the lineage probably migrated through 

Alaska no later than the early Miocene. The lineage 

became extinct in Japan but probably survived on the 

Asian mainland as A. henryi Pax.

Cocculus auriculata. This extinct lineage is 

thought to be descended from an undescribed species 

in the Puget Group and is hence West American. The 

lineage has a Miocene distribution similar to that of

A. negundo type but became extinct in North America 

by the late Miocene.

The only East Asian species that probably migrated 

into middle latitudes of western North America during the Neogene are:

Acer mono type (A. scottiae MacG.).—This type has 

not yet been found in the Alaskan Tertiary sequence, 

but the common occurrence of this type in the middle 

and late Miocene assemblages from the Pacific 

Northwest indicates that the lineage probably passed 

through Alaska during the Seldovian.

Acer miyabei type.—This type has not been previously recorded from the conterminous United States, 

but seeds from Skull Springs, Oreg., (middle Miocene) 

and south-central Idaho (middle or late Miocene) 

probably represent Acer ezoanum. This lineage has a 

long history in eastern Asia (Tanai, 1972).

Excluded from this listing is Acer circinnatum, a 

species closely allied to A. japonicuin, which has an 

Asian record extending back at least into the early 

Miocene (Tanai, 1972). No fossil record of this type is 

known in North America, which is curious in view of 

the current abundance and widespread occurrence of 

A. circinnatum in the forests of the Pacific Northwest. 

We suggest that this is a rare example of longdistance dispersal from Japan to the Pacific Northwest during the Pliocene or Quarternary.

Neither the West American nor the East Asian 

element apparently penetrated significantly southward in Asia or North America, respectively. Few 

species are found in the Neogene of both Japan and 

the Pacific Northwest. We suggest that the northern 

populations of these two elements were unable to 

compete successfully with endemic, more southerly 

populations; that is, while it was apparently easy for 

many species to spread northward, producing populations better adapted to the more rigorous conditions at 

higher latitudes, the reverse process was not easy. 

Possibly these northern populations were genetically 

less plastic than the southern populations of the same 

species. Another explanation is the possibility that the 

midlatitudes of western North America, although 

clearly having more summer precipitation in the 

Neogene than now, had an insufficient amount for 

most East Asian species; the lack of penetration of 

most West American species into Asia, however, 

would remain unexplained. We think that the available data support van Steenis' (1962) suggestion that 

it is improbable that many species could adapt first to 

rigorous conditions and then readapt to conditions 

similar to those of the early member populations of 

the species.

Van Steenis (1962) was particularly referring to the 

problem of day length (particularly the lack of winter 

light) relative to floristic interchange over high-
PALEOCLIMATOLOGY 19

latitude land bridges such as Beringia. We are, however, discussing deciduous species, which are leafless 

during winter, and thus the relative darkness of the 

winter is irrelevant. Temperature is not likely to be 

significant as a barrier to migration, because highaltitude areas at middle latitudes can have the same 

mean annual and mean annual range of temperature 

as low-altitude areas at high latitudes. The only major 

environmental factor that remains is that of day 

length as related to the phenomenon of photoperiodism. Although we have noted probable exceptions, we think that the data in this report suggest 

that generally it is impossible to reverse the trend of 

photoperiodic specialization in woody deciduous 

plants.

PALEOCLIMATOLOGY

Figure 8 shows the distribution of major temperature parameters relative to vegetational types at 

higher middle to high latitudes of eastern Asia (Wolfe,

1979a). The placement of each climatic station in a 

given vegetational type follows the maps of Wang 

(1961), Honda (1928), Kara (1959), and Suslov (1961). 

That the Seldovia Point assemblage is Mixed Northern Hardwood forest has been discussed above, in 

terms of both foliar physiognomy and general floristic 

affinities.

The climatic region now occupied by Mixed Northern Hardwood forest in eastern Asia is broad. The Seldovia Point assemblage is, we think, assignable to the 

warmer and more equable part of that vegetational 

type; that is, the Seldovia Point assemblage probably 

lived at temperatures similar to those of southwestern 

Hokkaido. In this area, some broad-leaved deciduous 

plants of a typically more southerly distribution occur: 

for example, Lindera, Cercidiphyllum, Berchemia, 

Pterocarya, Schisandra, and Cocculus (Hara, 1959). 

Analogous genera in the Seldovia Point assemblage 

are: Cocculus, Cercidiphyllum, Pueraria, Alangium, 

Platanus, and Liquidambar.

(STEPPE)

25°C

20° C

BROAD-LEAVED 

EVERGREEN

BROAD-LEAVED 

Scleropnyllous DECIDUOUS

Broad-leaved

Evergreen

I r r

10 20 30 

MEAN ANNUAL RANGE OF TEMPERATURE, IN DEGREES CENTIGRADE

FIGURE 8. Temperature parameters of modern vegetational types in eastern Asia and the probable temperature parameters of certain early and

middle Miocene assemblages.
20 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

That the Seldovia Point assemblage had a climate 

that at least was along the more equable margin of 

Mixed Northern Hardwood forest is also shown by the 

fact that coniferous forest was not far distant. Although megafossils of Pinaceae have not been collected at any of the Seldovia Point localities, the microfossil assemblages contain up to 34 percent Picea 

pollen, accompanied by a diversity of other pinaceous 

genera (aff. Cedrus, Picea, Pinus, Tsuga). The samples 

that have the numerically and taxonomically most 

abundant representation of Pinaceae are the more 

southerly localities, indicating that probably the 

coastal area was occupied by coniferous forest.

The temperature regime thus indicated for the Seldovia Point assemblage is 6-7°C mean annual temperature (see fig. 8); if coniferous forest were not far 

distant, then a mean annual range of temperature of 

26-27°C is indicated. These estimates in turn indicate 

that mean annual temperature has declined since the 

middle Miocene by about 3-5°C. As noted previously 

(Wolfe, 1971), however, the Seldovia Point mean annual range of temperature was 9-ll°C greater than 

now; that is, the present climate at Seldovia is considerably milder than the middle Miocene climate. There 

has been almost no change (if anything, perhaps a 

slight increase) in winter temperatures, but there has 

been a drastic change in warm month temperatures; 

the mean temperature of the warmest month has declined by about 7-8°C. We suggest that it is this 

strong decline in summer temperatures during the 

Neogene that was a major factor in the initiation of 

widespread glaciation during the later Cenozoic.

Today areas such as southern Alaska have a considerably lower mean annual range of temperature (that 

is, have a milder climate) than do areas to the southwest. The same situation appears to have prevailed 

during the middle Miocene, judging from paleobotanical data from Sakhalin and particularly Japan. In 

Sakhalin, Fotianova's (1967) analysis of the Upper 

Dui flora (fig. 9) indicated the total absence of members of Pinaceae; that is, the assemblage represents a 

broad-leaved deciduous forest that was geographically 

not close to coniferous forest. That the Upper Dui assemblage represents Mixed Northern Hardwood forest 

is clear from the diversity of lobed leaves (primarily 

Acer); another similarity to the Seldovia Point assemblage is in the abundance and diversity of Ulmaceae. Although Fotianova (1967) does not list 

Fagus as a member of the Upper Dui flora, Heer 

(1978) illustrated many examples of F. antipofi.

Floras from other areas of Sakhalin are generally 

similar to the Upper Dui. The Esutoru, Odasu, Naihoro, and Kashihi floras (Huzioka, 1949) lack Pinaceae 

but have a diversity of Ulmaceae, Betulaceae, 

Aceraceae, and Salicaceae. Among probable trees,

Populus, Alnus, Ulmus, and Acer are particularly 

common; Alangium is also well represented.

Overall the Upper Dui and other floras from Sakhalin do not appear to differ vegetationally from the Seldovia Point assemblage, except for the apparent absence of members of Pinaceae. The warmer elements 

present in the Sakhalin middle Miocene flora— 

Alangium, Leguiminosae—are also represented in the 

Seldovia Point assemblage. What is known of the 

middle Miocene flora of Kamchatka (Chelebaeva, 

1968) supports the suggestion that mean annual temperature during the middle Miocene was almost constant from Sakhalin through Kamchatka and into 

southern Alaska; the same situation prevails today. 

This interpretation of the paleobotanical data would 

also indicate—at least for Kamchatka—a decrease in 

mean annual range of temperature since the middle 

Miocene.

The middle Miocene Yoshioka flora of southwestern 

Hokkaido was interpreted by Tanai (Tanai and 

Suzuki, 1963) to be Mixed Mesophytic forest (Castanea zone of many Japanese botanists). Although 

many of the Yoshioka species occur primarily in this 

vegetational type in both Japan and central China, it 

is significant that broad-leaved evergreen plants— 

typically shrubs but some also scattered trees—occur 

in Mixed Mesophytic forest (Wang, 1961, p. 98, 100). 

In the Yoshioka assemblage, in contrast, the broadleaved evergreen element is represented only by two 

microphyllous species, Quercus elliptica and Camellia 

protojaponica.

In several respects the Yoshioka assemblage resembles what Wang (1961) termed Deciduous Broadleaved forest; this type of vegetation has been greatly 

disturbed, and in fact eliminated, in most areas of the 

Great Plain of North China. In this vegetation, broadleaved deciduous trees and shrubs predominate, and 

broad-leaved evergreens are rare. Notable, however, 

is the occurence in the vegetation of broad-leaved deciduous plants that are typical of more southern vegetational types: Castanea, Gleditschia, Cedrela, Pistacia, for example.

The mean annual temperatures under which this 

Deciduous Broad-leaved forest lives, however, are approximately the same as those under which Mixed 

Mesophytic forest lives, and thus the mean annual 

temperature suggested for the Yoshioka assemblage 

by Tanai (10-14°C; Tanai and Suzuki, 1963) is probably valid. Judging, however, from the diversity of 

lobed leaves in the Yoshioka (most representing Acer), 

Mixed Northern Hardwood forest, in which lobed 

species are common, was not far distant. Indeed, the 

slightly altitudinally higher Abura flora probably represents this vegetational type (Tanai in Tanai and 

Suzuki, 1963), and we thus suggest that the mean an-
PALEOCLIMATOLOGY 21

120° 140° 160° 170° 180° 170° 160° 70°

70°

60°

50°

40°

\ \ \ \ \

A *

Shanwang

/)

KYUSHU \J °L

X

100 200 300 400 500KILOMETERS 

I I I I I

60° 

160°

170°

180°

50°

170°

40° 

160°

150°

120° 30° 130° 140° 

FIGURE 9.—Map of northeastern Asia showing location of some middle Miocene assemblages.
22 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

nual temperature for the Yoshioka assemblage was 

probably 10-12°C.

The Utto flora of northwestern Honshu was interpreted to have lived under a mean annual temperature of 15°C but at the same time to have represented 

broad-leaved deciduous forest (Huzioka, 1963). As 

analyzed by Huzioka, the forest also included broadleaved evergreen trees, and the shrub component included both broad-leaved evergreen and deciduous 

plants. The evergreen element includes members of 

Lauraceae and Fagaceae, which account for almost 

half this element. Comparisons were made with vegetation in southern Honshu, but in that area the 

vegetation—except for secondary vegetation—is dominantly broad-leaved evergreen. Broad-leaved deciduous forests that have a floristic composition highly 

similar to the Utto assemblage do, however, exist in 

mainland China. In the southern parts of Shensi and 

Kansu and adjacent parts of Honan provinces, the 

forest (lower oak forest of Wang, 1961) is primarily 

broad-leaved deciduous, but the trees, as well as the 

shrubs, include both broad-leaved evergreen and deciduous types. Evergreen Lauraceae and Fagaceae are 

particularly notable (Wang, 1961, p. 89-90).

The mean annual temperature suggested for the 

Utto Flora by Huzioka (15°C) is present in the area 

occupied by the lower oak forest, and this would indicate a decline of mean annual temperature of about 

4°C since the middle Miocene. The significant point to 

be made in the above comparison is that in the area 

occupied by this lower oak forest the mean annual 

range of temperature is typically higher than the 

present range at the Utto localities. The change in 

mean annual range of temperature since the middle 

Miocene need not have been great (perhaps only 

2-3°C), but such a change is consistent with the 

change indicated by high-latitude floras.

Fossil assemblages farther south in Japan have 

been uniformly compared to the modern Evergreen 

Sclerophyllous Broad-leaved forest (Tanai, 1961, 

1967a, 1972; Matsuo, 1963; Ishida, 1970). These assemblages typically contain elements indicating 

somewhat warmer mean annual temperatures during 

the middle Miocene than now, but estimates of mean 

annual range of temperature are almost impossible to 

arrive at. It is clearly possible that this vegetational 

type lived under a higher mean annual and mean annual range of temperature than now exists at the fossil localities. Data from these more southerly floras 

neither contradict nor substantiate the changes in 

mean annual range of temperature indicated by the 

more northerly floras. The middle Miocene latitudinal 

vegetational zonation indicated for eastern Asia indicates, however, that there has been a decrease in

mean annual range of temperature since that time.

In western North America at middle latitudes (fig. 

10), most of the known middle Miocene assemblages 

grew in uplands. These uplands, moreover, are east of 

the Sierra-Cascade axis, and interpretations of 

changes in climate since the middle Miocene are 

greatly complicated by tectonic and altitudinal factors. In the Pacific Northwest, comparisons are best 

made to climatic data from the region west of the Cascade Range.

Middle Miocene floras such as the Latah (Knowlton, 

1926; Chaney, 1959), Grand Coulee (Berry, 1931), St. 

Eugene (Hollick, 1927), and Fish Creek (Wolfe, in 

Peck and others, 1964) represent Mixed Mesophytic 

forest. These assemblages are dominantly broadleaved deciduous but contain some notophyllous 

broad-leaved evergreens (Magnolia, Lauraceae, Exbucklandia, Arbutus). These assemblages, however, 

grew at an altitude perhaps over 500-600 m (Wolfe, 

1969a, p. 93). The Latah and similar assemblages 

lived in a mean annual temperature of 10-13°C and a 

mean annual range of temperature of 19-27°C. Comparisons with data for Cedar Lake (a station on the 

west side of the Cascades at an altitude of almost 500 

m) indicate that since the middle Miocene mean annual temperature has declined by at least 1-3°C and 

that mean annual range of temperature has decreased 

by 4-12°C.

Lowland assemblages of middle Miocene age are 

still poorly known in the Pacific Northwest. The undescribed Wishkaw River assemblage from western 

Washington and particularly the undescribed Cape 

Blanco assemblage from southwestern Oregon probably represent Sclerophyllous Broad-leaved Evergreen 

forest. Notophyllous members of evergreen Fagaceae 

and Lauraceae are conspicuous elements in these assemblages. If this vegetation type is represented, then 

there has probably been a moderate (at least 2-3°C) 

decline in mean annual temperature since the middle 

Miocene. In areas lacking major mountains parallel to 

the coast (such as the middle Miocene Pacific Northwest), mean annual range of temperature increases 

only a few degrees a few hundred kilometers from the 

coast. It is improbable that mean annual range of 

temperature in the lowlands during the middle 

Miocene was more than a few degrees less than that 

indicated for upland assemblages such as the Latah 

(19-27°C). If a minimal mean annual range of temperature of 15°C is assumed for the Cape Blanco and 

Wishkaw River climates, then mean annual range of 

temperature in the coastal areas has lessened since 

the middle Miocene; mean annual range is today almost 7°C at Port Orford near Cape Blanco and almost 

12°C near Wishkaw River. On Cape Blanco itself 

mean annual range of temperature is less than 5°C.
PALEOCLIMATOLOGY 23

60° 175° 65° 170° 70° 165" 160°75° 155° 150° 145° 140°135°130'

175'

170° 

55'

165'

160'

50' 

155°

150° 

45 e

145° 

40'

140' 

35°

V/ / / 7 /

W

100 

I

0 100 200 300 400 500 MILES 

I I | I I I

I I I I I I I

100 0 100200300400500 KILOMETERS

125° 

80°

75°

70°

65°

60"

55°

50°

45"

40"

135° 130° 35 0 125" 120° 115° 110"

FIGURE 10.—Northwestern North America showing location of some middle Miocene assemblages.
24 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

The paleobotanical evidence is thus consistent 

around the North Pacific Basin: mean annual temperature has declined since the middle Miocene, although 

the decline has been greatest at high latitudes. Similarly, mean annual range of temperature has declined, and the decline has been greatest at high 

latitudes. Concomitant with the decline in mean annual range of temperature, the decline in mean annual temperature primarily results in a lowering of 

summer temperatures; we emphasize again that such 

a decline in summer temperatures was a major factor 

in the initiation of late Cenozoic glaciation.

Previous analysis of paleoclimates from western 

North America (Wolfe and Hopkins, 1967; Wolfe, 

1971) has indicated that the middle Miocene was 

warmer than either the earliest or late Miocene, and 

paleobotanical analyses based on eastern Asian materials are in basic agreement (Tanai and Huzioka, 

1967; Tanai, 1967a, 1967b, 1971). The Alaskan data 

clearly are also in agreement in comparing the earliest with the late early to early middle Miocene climates.

The vegetation represented by the assemblages 

from the lower Seldovian of the Cook Inlet region was 

also probably Mixed Northern Hardwood forest but 

more depauperate than that of the upper Seldovian. 

The assemblages of probable early Seldovian age in 

the Cook Inlet region are Capps Glacier (Iocs. 9845, 

9846, 9937 of Wolfe and others, 1966) and Houston 

(loc. 9365 of Wolfe and others, 1966). Preliminary determinations of the flora from the localities indicate 

that the following are represented:

Glyptostrobus europaeus

Metasequoia cf. M. glyptostroboides

Keteleeria sp.

Chamaecyparis sp.

Cercidiphyllum alaskanum

Uhnus sp.

Quercus furuhjelmi

Alnus barnesi

Alnus cappsi

Alnus fairi

Alnus healyensis

Carpinus cappsensis

Comptonia naumannii

Myrica sp.

Pterocarya nigella

Populus kenaiana

Populus spp.

Salix cappsensis

Salix picroides

Salix spp.

Hydrangea sp. 

Crataegus sp. 

Sorbaria sp. 

Spiraea weaveri 

Cladrastis aff. C. lutea 

Acer aff. A. pennsylvanicum 

Acer aff. A. saccharinum 

Aesculus cf. A. majus 

Alangium mikii

Notable in these assemblages is the diversity of both 

Betulaceae and Salicaceae, which are also the most 

abundant megafossils. Just as significant is the lesser 

diversity in the lower as compared to the upper Seldovian flora of Fagaceae, Ulmaceae, Juglandaceae, and 

Aceraceae. Genera such as Cocculus, Liquidambar, 

Platanus, and Cyclocarya are absent from the lower 

Seldovian megafossil assemblages.

The most reasonable interpretation of such floristic 

differences is that the middle Miocene warming 

documented at middle latitudes allowed the northward expansion of some species into southern Alaska. 

Because the physiognomy of the early Seldovian assemblages also indicates broad-leaved deciduous 

forest, it appears that mean annual range of temperature also decreased from the early into the middle 

Miocene; this suggestion accords with the trend after 

the middle Miocene.

The Homerian (late middle to early late Miocene) 

assemblages from Alaska, although representing coniferous forest, do not necessarily indicate that mean 

annual temperature declined. From figure 8 it can be 

seen that a change from broad-leaved deciduous to 

coniferous forest can be the result of a decrease in 

mean annual range of temperature alone. The rich 

broad-leaved deciduous tree element that is conspicuous even in coniferous assemblages of Seldovian age 

(for example, Wrangell and Noxapaga assemblages) is 

absent in the early Homerian megafossil assemblages, 

and only sparse amounts of Pterocarya/Cyclocarya and 

Ulmus/Zelkova occur in some microfossil assemblages. 

If, as seems probable, broad-leaved trees are better 

adapted to higher summer heat than are most coniferous trees, and if diversity of broad-leaved trees increases with overall warmth, then the simplest way to 

eliminate broad-leaved trees from Alaska following 

the Seldovian would be to drastically lower summer 

temperatures and overall heat. Such a lowering is accomplished by lowering both mean annual and mean 

annual range of temperature.

SYSTEMATICS

The preliminary list of the Seldovia Point flora pre-
SYSTEMATICS 25

viously published (Wolfe, 1966) differs in several respects from that reported here. In order to clarify any 

differences, the following is presented:

Wolfe, 1966 This report

Equisetum sp _____________ Not recognized; specimen lost?

Dryopteris sp ______________ Dryopteris sp.

Onoclea sensibilis _________ Onoclea sensibilis.

Ginkgo biloba ________________ Ginkgo biloba.

Glyptostrobus europaeus ______ Glyptostrobus europaeus.

Metasequoia glyptostroboides ___ Metasequoia cf. M. glyptostroboides.

Taxodium distichum __________ Not recognized; probably aberrant shoots of Glyptostrobus or 

Metasequoia.

Potamogeton sp. ___—_______ Potamogeton alaskanus.

Smilax sp________________ Alisma seldoviana.

Poacites tenuistriatus ________ Monocotylophyllum spp.

Cyperacites sp________________ Monocotylophyllum sp.

Populus kenaiana __________ Populus kenaiana.

Populus reniformis _________ Populus sp.

Populus sp. aff. P. ciliata ____ Populus kenaiana.

Salix inquirenda _______ Salix picroides.

Salix picroides __________ Salix picroides.

Salix sp_-_-______________ Salix cappsensis.

Carya bendirei ______________ Carya bendirei.

Carya sp. aff. C. sessilis ________ Carya bendirei.

Pterocarya mixta ____________ Pterocarya nigella.

Pterocarya nigella ______________ Pterocarya nigella.

Pterocarya (Cycloptera) sp ____ Cyclocarya ezoana.

Alnus cappsi ____________ Alnus cappsi.

Alnus healyensis ____________ Alnus healyensis.

Alnus fairi __________________ Alnus fairi.

Carpinus seldoviana ____________ Carpinus seldoviana.

Fagus antipofi ________________ Fagus antipofi.

Fagus sp. cf. F. paleocrenata ____ Fagus aff. F. crenata.

Quercus bretzi ______________ Quercus furuhjelmi.

Quercus furuhjelmi _______ Quercus furuhjelmi.

Ulmus longifolia __________ Ulmus owyheensis.

Ulmus newberryi __________ Ulmus knowltoni.

Zelkova oregoniana ___________ Zelkova browni, Z. ungeri.

Nuphar sp __________________ Nymphar ebae.

Cercidiphyllum crenatum ____ Cercidiphyllum alaskanum.

Cocculus auriculata _________ Cocculus auriculata.

Hydrangea sp ______________ Hydrangea sp.

Liquidambar mioformosana ____ Liquidambar pachyphylla.

Platanus bendirei ____________ Platanus bendirei.

Crataegus sp _______________ Crataegus chamisonii.

Prunus sp-_________________ Prunus kenaica.

Spiraea? andersoni _________ Not considered; based only on

Heer's illustration.

Alchornea? sp __________________ Indeterminate leaf.

Mallotus sp _________________ Pueraria miothunbergiana.

Acer ezoanum _______________ Acer ezoanum.

Acer fatisiaefolia __________ Acer ezoanum.

Acer macropterum __________ Acer grahamensis.

Acer sp. aff. A. crataegifolium __ Acer heterodentatum.

Acer sp. cf. A. subpictum _____ Acer ezoanum fruits.

Vitis sp_--_-_____________ Vitis seldoviana.

Tilia sp______________________ Tilia subnobilis.

Nyssa sp. cf. N. knowltoni ___ Nyssa cf. N. knowltoni.

Hemitrapa borealis _________ Hemitrapa borealis.

Kalopanax sp ______________ Kalopanax acerifolius.

Fraxinus sp ____________________ Fraxinus kenaica.

Symphoricarpos sp _________ Decodon alaskana.

Family Aspidiaceae 

Genus Dryopteris Adanson

Dryopteris sp. 

Plate 1, figures 5, 9

Description.—A portion of a pinna 3.5 cm long and 

1.7 cm wide at the lower part, lanceolate; pinnules 

ovate, suboppositely pinnatified by shallow but narrow sinus, nearly entire or having a few teeth on 

margin; midvein of each pinnule leaving pinna axis at 

angles of 40 to 50 degrees, somewhat zigzag, forking 

at the apical part, having 3 or 4 pairs of lateral veins 

that fade out near the margin without any bifurcation.

Discussion.—A single pinna and its counterpart are 

fragmentary and sterile, and the generic assignment 

is uncertain. But these specimens show close resemblance to the pinnae of some extant Dryopteris in 

shape of pinnae and venation, such as D. decipiens O. 

Kuntze and D. tokyoensis (Mats.) C. Chr. living in 

East Asia. Our Alaskan specimens are quite different 

in venation from D. guyotti (Lesq.) MacG. and D. 

idahoensis Knowlt., which were described from the 

middle Tertiary of the conterminous United States.

Occurrence.—9858

Specimen.— USNM 208348A, B.

Genus Onoclea L. 

Onoclea sensibilisis L. 

Plate 1, figures 3, 4

Discussion.—Several fragmentary pinnae from Seldovia Point are undoubtedly referrable to the genus 

Onoclea by their characteristic venation, which is 

quite well preserved. Except for the wavy margin of 

the apical part, they are regularly lobed (each lobe 

having a rounded apex) but are otherwise entire margined. The pinnae axis bears a single series of low and 

long areoles on each side. The midvein of each pinnule 

is thin but distinct and gives off thin secondaries that 

frequently branch, forming a prolonged reticulation 

nearly parallel to or at acute angles to the midvein. 

These characters show that our Alaskan specimens 

are identical with the modern monotypic species, 

Onoclea sensibilis L., distributed in northeastern Asia 

and eastern North America and may represent the 

pinnae of the middle or proximal parts of the frond.

The sterile foliage of Onoclea has frequently been 

recorded from the Upper Cretaceous and Paleogene of 

North America, and Brown (1962) gave a new specific 

name of O. hesperia to them. As stated previously 

(Wolfe, 1966), the original specimens of O. hesperia 

include both finely serrate and entire margins and are 

thus separable from the Seldovian specimens.

Hypotypes.—USNM 208349, 208350.

Occurrence.—9858.
26 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Family Taxodiaceae

Genus Glyptostrobus Endl.

Glyptostrobus europaeus (Brongniart) Heer

Plate 1, figure 8

Glyptostrobus europaeus (Brongniart) Heer, 1855, 

Flora Tertiaria Helvetia, v. 1, p. 51, pi. 19; pi. 20, 

fig. 1.

Discussion.—This species is represented only by 

fragmentary foliage shoots, which have slender 

leaves. No fossil cone of Glyptostrobus has been collected from the Seldovia Point localities. This 

Eurasian species was widely distributed during the 

Tertiary, and it is difficult to separate from G. 

oregonensis Brown of the North American Tertiary 

only by foliar characters. Cones collected from other 

Seldovian localities, however, are typical for G. 

europaeus.

Hypotype.—USNM 208351.

Occurrence.—9858.

Genus Metasequoia Miki 

Metasequoia cf. M. glyptostroboides Hu et Cheng

Plate 1, figures 2, 6, 7, 10

Metasequoia glyptostroboides Hu et Cheng, 1948, Fan 

Memorial Inst. Botany Bull., new ser., v. 1, p. 

154.

Discussion.—Numerous short shoots and pistillate 

cones of Metasequoia from the Seldovia Point localities are indistinguishable from homologous parts 

of the extant species. Lacking preservation of epidermal features, however, we hesitate to place the fossil 

material in the extant species. The material should 

not be assigned to M. occidentalis (Newb.) Chan. because the types, as well as other Paleocene shoots of 

Metasequoia, typically have shorter and blunter needles than in middle to late Tertiary specimens of 

Metasequoia.

Specimens.— USNM 208352-208355.

Occurrence. —9856-9858.

Family Ginkgoaceae 

Genus Ginkgo Linnaeus 

Ginkgo biloba Linnaeus

Plate 1, figure 1

Ginkgo biloba Linnaeus, 1771, Linn. Mant., v. 2, p. 

313.

Discussion.—In his extensive review of the geologic 

history of Ginkgo, Tralau (1968) noted that later Tertiary specimens for which epidermal details were 

known did not differ specifically from the extant 

species. We think that there is no reasonable basis for 

excluding later Tertiary material of this genus from 

the extant species.

Hypotype.— USNM 208356.

Occurrence.—9857.

Family Nymphaeaceae

Genus Nymphar Ozaki

Nymphar ebae (Huzioka) Ozaki

Plate 1, figures 11-14

Nymphar ebae (Huzioka) Ozaki, 1978, Yokohama 

Natl. Univ. Sci. Repts., sec. 2, no. 25, p. 17, pi. 1, 

figs. 1, 3-5; text-figs. 4B-D, 5A, B. 

Nuphar ebae Huzioka, 1964, Akita Univ. Mining Coll. 

Jour., ser. A, v. 3, no. 4, p. 82, pi. 11, fig. 6; pi. 12, 

figs. 1-3.

Supplementary description.—Floating leaves ovate 

to oval in shape, 2.1 to 5.2 cm long and 1.3 to 3.2 cm 

wide; apex obtuse to rounded; base auriculate with 

deep sinus; midvein distinct, nearly straight to apex; 

secondary veins slender, 9 to 15 pairs, irregularly 

spaced, leaving the midrib at angles of 50 to 70 degrees, a basal pair more spreading, dichotomizing 3 or 

4 times toward the margin and forming series of loops, 

camptodrome; the intersecondary veins sometimes 

developing but weak; the tertiary veins thin, forming 

irregularly large networks which enclose small polygonal areoles formed by the fourth order veins; margin entire; petiole rather slender, more than 2 cm 

long.

Discussion.—These are the most abundant leaves in 

our collection, and they are identical in shape and venation to Nymphar ebae, which was described from a 

lower Miocene flora of Honshu.

Hypotypes.— USNM 208357-208360.

Occurrence.—9858.

Family Menispermaceae

Genus Cocculus DeCandolle

Coccu/us auriculata (Heer) Wolfe

Plate 2, figure 7

Cocculus auriculata (Heer) Wolfe, 1966, U.S. Geol.

Survey Prof. Paper 398-B, p. B24, pi. 7, fig. 1. 

Hedera auriculata Heer, 1869, Flora Foss. Arctica, v.

2, pt. 2, p. 36, pi. 9, fig. 6. 

Populus heteromorpha Knowlton, 1926, U.S. Geol.

Survey Prof. Paper 140-A, p. 30, pi. 12, figs.

8-10; pi. 13, figs. 1-7; pi. 14, figs. 1-3; pi. 15,

figs. 3-5. 

Berry, 1929, U.S. Geol. Survey Prof. Paper 154, p.

242.

Hoffman, 1932, Jour. Geology, v. 40, p. 735. 

Populus fairii Knowlton, 1926, U.S. Geol. Survey Prof.

Paper 140, p. 30, pi. 15, fig. 2; pi. 16, figs. 1-3. 

Cebatha multiformis Hollick, 1927, New York Bot.

Garden Mem., v. 7, p. 406, pi. 38, figs. 1-6; pi.

39, figs. 1-3. 

Cissampelos dubiosa Hollick, 1927, New York Bot.

Garden Mem., v. 7, p. 408, pi. 37, figs. 4-7; pi.

39, fig. 4. 

Cebatha heteromorpha (Knowlton) Berry, 1931, U.S.

Geol. Survey Prof. Paper 170, p. 37.
SYSTEMATICS 27

Berry, 1934, U.S. Geol. Survey Prof. Paper 185, p.

112.

LaMotte, 1936 [part], Carnegie Inst. Washington 

Pub. 455, p. 126 [unfigured specimen 839 only]. 

Cocculus heteromorpha (Knowlton) Brown, 1946,

Washington Acad. Sci. Jour., v. 36, p. 352. 

Tanai, 1961, Hokkaido Univ. Fac. Sci. Jour., ser. 

IV, p. 324, pi. 21, fig. 7.

Discussion.—This species is one of the most widespread plants in the Miocene of the areas bordering 

the North Pacific, occurring from Honshu north to 

Alaska and south to northern California. In Alaska, 

Cocculus auriculata is known only from the Seldovia 

Point flora and from the Seldovian rocks at Cache 

Creek and in the Nenana coalfield (Wahrhaftig and 

others, 1969).

The specimen assigned by Becker (1969, pi. 26. figs. 

12, 13) to Cocculus heteromorpha is certainly Populus, 

as indicated by the numerous small teeth and the 

closely spaced sinuous tertiary veins that are oriented 

at an acute angle to the secondary veins. In Cocculus 

auriculata (including the junior synonym C. heteromorpha}, the teeth are few and large, and the tertiary 

veins are widely spaced and perpendicular to the secondary veins.

Cocculus auriculata is clearly related to Paleogene 

species of the genus such as C. ezoensis Tanai from the 

Oligocene of Japan. Undescribed material of the same 

group of Cocculus also occurs commonly in the Eocene 

Ishikari floras of Hokkaido. C. ezoensis is, however, 

typically entire margined, although some toothed 

specimens are known (Tanai, 1970), in contrast to C. 

auriculata, which in illustrated specimens and material we have observed always has some teeth. Undescribed material from the lower Ravenian of the Puget 

Group of Washington (loc. 8640 and 9694 of Wolfe, 

1968) has numerous teeth, but the areoles are smaller 

than in C. auriculata and the leaves have small 

mounds of apparently cutinized material. The small 

mounds, which are also present in the Paleocene C. 

flabella (Newb.) Wolfe, indicate a thick leaf and probable evergreen habit (Wolfe, 1966), in contrast to the 

thin leaf and probably deciduous habit of C. auriculata.

Hypotype.—USNM 208369.

Occurrence.— 9856, 9858.

Family Cercidiphyllaceae

Genus Cercidiphyllum Sieb. et Zucc.

Cercidiphyllum alaskanum Wolfe et Tanai, sp. nov.

Plate 2, figures 1, 3, 4; plate 11, figures 3, 5

Cercidiphyllum crenatum auct. non (Unger) Brown. 

Chelebaeva, 1968, Bot. Zhurn., v. 53, p. 744, pi. 4, 

figs. 1, 2; text-fig. 4a-b. 

Description.—Leaves simple; lamina symmetrical,

wide ovate to very wide ovate; apex obtuse to acute; 

base cordate; margin simply to doubly crenate, teeth 

glandular; venation actinodromous; midrib and first 

two lateral pairs of primary veins of moderate size, 

two additional pairs of lateral primary veins weak; 

lateral primary veins markedly curved, irregularly 

brochidodromous, first lateral pair acrodromous; secondary veins irregularly brochidodromous, diverging 

at a widely acute angle, uniformly to abruptly curved, 

branched; intercostal areas between midrib and first 

lateral pair of primary veins braced by widely and irregularly spaced tertiary veins that are approximately perpendicular to primary veins, typically with 

a composite intersecondary vein; composite intersecondary veins typically present in area between first 

and second pairs of lateral primary veins; fourth order 

venation irregularly spaced, tending to be orthogonal; 

fifth order venation forming irregularly polygonal and 

randomly oriented areoles intruded by irregularly 

branching freely ending veinlets.

Discussion.—Typically orbicular leaves of the Cercidiphyllum type were assigned to the European 

species C. crenata (Unger) R. W. Br. (Brown, 1935, 

1939), and most workers have typically followed this 

practice. Comparison of the Seldovia Point material 

with topotypic material from the Swiss Tertiary indicates a marked difference in ultimate venation, particularly in regard to the size of the areoles. Additionally, in the Swiss material the marginal glands are 

incorporated in the foliar tissue, whereas in the Seldovia Point material and the extant C. japonicum 

Siev. et Zucc. the glands are situated abmedial to the 

teeth. The Seldovia Point species also has larger 

areoles than in C. japonicum.

Cercidiphyllum alaskanum also has a characteristic 

pattern in the lateral tertiary venation. The intercostal area between the first (from the midrib) lateral 

primary and the second lateral primary typically is 

braced by an intersecondary vein extending abmedially and giving off tertiary braces that extend to the 

second lateral primary and the most basal secondary 

of the first lateral primary. In Cercidiphyllum 

japonicum and in most other fossil material of Cercidiphyllum that we have examined, an intersecondary is lacking and the bracing is accomplished only 

by tertiary veins. The only other species of Cercidiphyllum that has the type of bracing seen in C. 

alaskanum is C. eojaponicum Endo ex Tanai, which 

was originally described from the early Oligocene 

Fushun beds of Manchuria and later found in approximately isochronous beds on Hokkaido (Tanai, 1970). 

An additional similarity between C. alaskanum and 

C. eojaponicum is the presence of some doubly crenate 

teeth (see, for example, Endo, 1968, pi. 15, figs. 3, 4).
28 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

We have not investigated the areolar size in C. 

eojaponicum, but this species differs from C. alaskanum in that the lateral primary and the secondary 

veins loop well within the margin. The strong 

similarities between these two species, however, indicate that they may be phyletically related.

Other material assigned to Cercidiphyllum crenatum by Brown (1935, 1939) should also be excluded 

from that species. Material from the Eocene beds at 

Republic, Wash., for example, has an areolar size 

comparable to that of the Swiss, material, but the leaf 

base is never deeply cordate (typically it is broadly 

rounded) and the teeth are sharp with glands abmedial to the teeth. We suggest that there is great need 

for a comprehensive examination and analysis of 

other material assigned to C. crenatum.

Holotype.—USNM 208361 A, B.

Paratypes.—USNM 208362, 208363A, B.

Occurrence.— 9856, 9858.

Family Hamamelidaceae

Genus Liquidambar L.

Liquidambar pachyphylla Knowlton

Plate 2, figures 5, 6; plate 3, figures 1-3

Liquidambar pachyphylla Knowlton, 1902, U.S. Geol.

Survey Bull. 204, p. 63, pi. 9, fig. 1. 

Chaney, 1920, Walker Mus. Contr., v. 2, p. 174,

pi. 15, figs. 2, 3. 

Knowlton, 1926, U.S. Geol. Survey Prof. Paper

140, p. 42, pi. 22, fig. 7; pi. 29, fig. 1. 

Brown, 1946, Washington Acad. Sci. Jour., v. 36,

p. 352. 

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

Washington Pub. 617, p. 181, pi. 35, fig. 5 only. 

Liquidambar europaeum patulum Knowlton, 1902,

U.S. Geol. Survey Bull. 204, p. 62, pi. 10, fig. 5. 

Liquidambar acutilobum Chaney, 1920, Walker Mus.

Contr., v. 2, p. 175, pi. 15, fig. 4. 

Arisaema hesperia Knowlton, 1926, U.S. Geol. Survey

Prof. Paper 140, p. 29, pi. 10, fig. 1. 

Liquidambar europaeum auct. non Al. Braun. Heer,

1869, Flora Fossilis Arctica, v. 2, no. 2, p. 25, pi.

2, fig. 7. 

Lesquereux, 1888, U.S. Natl. Mus. Proc., v. 11, p.

14.

Chaney, 1920, Walker Mus. Contr., v. 2, p. 174. 

Liquidambar californicum auct. non Lesquereux.

Chaney, 1920, Walker Mus. Contr., v. 2, p. 174. 

Berry, 1929, U.S. Geol. Survey Prof. Paper 154, p.

250. 

Berry, 1934, U.S. Geol. Survey Prof. Paper 185, p.

113. 

Berry, 1938, Torrey Bot. Club Bull., v. 65, p. 96,

text-fig. 3.

Liquidambar protensum auct. non Unger. Lesquereux, 

1888, U.S. Natl. Mus. Proc., v. 11, p. 13, pi. 8, 

fig. 3.

Knowlton, 1902, U.S. Geol. Survey Bull. 204, p. 

62.

Discussion.—Early collections from the Seldovia 

Point locality contained one poorly preserved specimen of Liquidambar that was three lobed; the specimen was thus assigned to the typically three-lobed L. 

mioformosana Tanai et Onoe. Additional specimens 

obtained in 1967, however, indicated that the Seldovia 

Point Liquidambar varied from three to five lobed, a 

condition that is typical for L. pachyphylla (Chaney 

and Axelrod, 1959, p. 182).

Analysis of available specimens of Liquidambar 

from early and middle Miocene localities in the western United States (Collawash, Latah, Eagle Creek, 

Mascall) indicates that this species is about evenly divided between three- and five-lobed leaves, in contrast 

to late Miocene leaves of Liquidambar (Faraday, Hidden Lake, Blue Mountains, Molalla, Weyerhauser), 

which are almost invariably five lobed (only one 

three-lobed specimen has been found in late Miocene 

beds). Additionally, the typical (early and middle 

Miocene) L. pachyphylla has sharp teeth that have a 

basal side about three times as long as the apical side, 

and the areoles are about 0.6-0.75 mm in diameter. 

The late Miocene material has rounded teeth that 

have a basal side about twice as long or less as the 

apical side and the areoles are about 1.5-2.1 mm in 

diameter. These differences are sufficient to exclude 

the late Miocene material from L. pachyphylla.

The present distribution of Liquidambar makes this 

genus an unlikely member of the Alaskan Neogene 

flora (see p. 17). The genus, however, was apparently 

not common. In most pollen samples from the Capps 

Glacier Seldovian and the Seldovian of the Alaska 

Range the genus is absent, although the genus is 

more persistent in the samples from the Suntrana 

Formation, which is late Seldovian and presumably 

middle Miocene. The high (14 percent) representation 

of Liquidambar in the sample from Seldovia Point is 

the highest we know of in the Alaskan Neogene sequence.

Hypotypes.—USNM 208364-208368.

Occurrence.— 9856, 9858.

Family Platanaceae

Genus Platanus Linnaeus

Platanus bendirei (Lesquereux) Wolfe

Plate 3, figure 4; plate 4, figures 1, 2, 4

Platanus bendirei (Lesquereux) Wolfe, 1964, U.S. 

Geol. Survey Prof. Paper 454-N, p. N24, pi. 4, 

figs. 1, 2, 4.
SYSTEMATICS 29

Acer bendirei Lesquereux, 1888 [part], U.S. Natl. 

Mus. Proc., v. 11, pi. 5, fig. 5; pi. 6, fig. 1; pi. 7, 

fig. 1. 

Acer merriami Knowlton, 1902, U.S. Geol. Survey

Bull. 204, p. 74, pi. 14, fig. 7. 

Platanus youngii Graham, 1963, Am. Jour. Botany, v.

50, p. 925, fig. 12. 

Graham, 1965, Kent State Univ. Bull., Research

Ser. 9, p. 89. 

Magnolia ingelfieldi auct. non Heer. Lesquereux,

1888, U.S. Natl. Mus. Proc., v. 11, p. 13. 

Platanus aceroides auct. non (Goeppert) Heer. Lesquereux, 1888, U.S. Natl. Mus. Proc., v. 11, p. 

19, pi. 5, fig. 7. 

Knowlton, 1902, U.S. Geol. Survey Bull. 204, p.

65. 

Arnold, 1937, Michigan Univ. Mus. Paleontology

Contr., v. 5, p. 88, pi. 3, fig. 1. 

Platanus nobilis auct. non Newberry. Lesquereux,

1888, U.S. Natl. Mus. Proc., v. 11, p. 19. 

Knowlton, 1902, U.S. Geol. Survey Bull. 204, p.

65. 

Platanus raynoldsii auct. non Newberry. Lesquereux,

1888, U.S. Natl. Mus. Proc., v. 11, p. 19. 

Platanus aspera auct. non Newberry. Berry, 1931,

U.S. Geol. Survey Prof. Paper 170, p. 34. 

Platanus dissecta auct. non Lesquereux. Berry, 1929, 

U.S. Geol. Survey Prof. Paper 154, p. 248, pi. 

53, figs. 1, 2; pi. 61. 

Berry, 1931, U.S. Geol. Survey Prof. Paper 170, p.

34. 

Berry, 1934, U.S. Geol. Survey Prof. Paper 185, p.

Ill, pi. 21, fig. 2. 

Arnold, 1937, Michigan Univ. Mus. Paleontology

Contr., v. 5, p. 88, pi. 9, figs. 1-3. 

Brown, 1937.[part], U.S. Geol. Survey Prof. Paper

186, p. 174, pi. 52, fig. 2 only. 

Brown, 1937, Washington Acad. Sci. Jour., v. 27,

p. 515. 

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

Washington Pub. 617, p. 182, pi. 36, fig. 3. 

Graham, 1965 [part], Kent State Univ. Bull., Research Ser. 9, p. 88, pi. 13, fig. 1; pi. 14, figs. 1, 5 

["Sucker Creek" occurrences only]. 

Discussion.—Some workers (for example, Becker, 

1969) have applied the epithet dissecta to any threeto five-lobed leaves of Platanus that also have numerous teeth. Such a morphological concept for P. dissecta 

results in the inclusion in this species of several distinctive (and probably in part unrelated) groupings of 

Platanus. As pointed out previously (Wolfe, 1964, p. 

N24), the late Miocene P. dissecta has leaves that are 

typically five lobed and typically without subsidiary 

teeth between the major teeth, in contrast to the early

to middle Miocene P. bendirei, which has typically 

three-lobed leaves (although five-lobed leaves are also 

present) that have subsidiary teeth. Moreover, in contrast to an undescribed species from the later 

Oligocene of the Pacific Northwest that has several 

subsidiary teeth between any two adjacent major 

teeth, P. bendirei typically has only one or two subsidiary teeth. On this basis, the Seldovia Point material is referrable to P. bendirei.

The reduction in marginal serrations suggested for 

the Platanus bendirei lineage led to the Pliocene P. 

paucidentata Dorf (differing from the next species in 

having larger areoles) and culminated in the extant P. 

racemosa Nutt. Thus far this lineage has been found 

only in North America; the few occurrences of 

Platanus in the Neogene of Japan represent the P. 

aceroides complex.

Hypotypes.—USNM 208370-208373.

Occurrence. —9858.

Family Eucommiaceae

Genus Eucommia Oliv.

Eucommia cf. E. montana Brown

Plate 4, figure 6

Eucommia montana Brown, 1940, Washington Acad. 

Sci. Jour., v. 30, no. 8, p. 349, fig. 3.

Discussion.—This specimen, although incomplete, 

is probably separable from E. montana in size and 

wing shape. Unfortunately this Alaskan material is 

carbonized, and the reticulate venation pattern over 

the seed part, as is characteristic of Eucommia, is not 

visible. In preserved features, our specimen is closely 

similar to the samara of the modern monotypic E. ulmoides Oliver of central China. No fossil leaves referable to Eucommia have been found in the Seldovia 

Point flora.

Specimen.—USNM 208374.

Occurrence.—9858.

Family Ulmaceae 

Genus Celtis Linn

Celtis sp. 

Plate 4, figure 7

Discussion.—A single, small leaf is, although lacking the basal part, referred to the genus Celtis partly 

on the basis of the pronounced subprimary veins and 

camptodromous venation. The leaf is similar in general features to C. kansana Chaney et Elias from the 

Miocene and Pliocene of the conterminous United 

States but appears to differ in having apiculate teeth.

Specimen.—USNM 208375.

Occurrence. —9858.
30 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Genus Ulmus Linnaeus 

Ulmus knowltoni Tanai et Wolfe

Plate 5, figures 1, 2

Ulmus knowltoni Tanai and Wolfe, 1977, U.S. Geol. 

Survey Prof. Paper 1026, p. 5, pi. 1C, F, G; pi. 

2A, C, H, I, J [see synonymy and discussion].

Discussion.—Two incomplete specimens are referred to U. knowltoni on the basis of a strongly 

asymmetric cordate base and nearly single blunt 

teeth. This species is somewhat similar to U. 

paucidentata H. V. Smith and U. owyheensis H. V. 

Smith in marginal serrations but differs in shape of 

the base and in the tertiary branches of the basal secondary vein, which develop only on one side of the 

base.

Hypotypes.—USNM 208376, 208377.

Occurrence.—9856.

Ulmus owyheensis H. V. Smith

Plate 4, figure 8; plate 6, figures 7a, 7b

Ulmus owyheensis Smith, 1939, Michigan Acad. Sci.,

Arts and Letters Papers, v. 24, p. 113, pi. 6, fig.

4. 

Tanai and Wolfe, 1977, U.S. Geol. Survey Prof.

Paper 1026 p. 6, pi. 2B, D, E, F; pi. 3A [see

synonymy and discussion]. 

Ulmus plurinervis Unger. Heer, 1869, Kongl.

Svenska Vet.-Akad. Handl., bd. 8, no. 4, p. 34,

pi. 5, fig. 1.

Discussion—Our Alaskan specimens are quite identical in shape, venation, and marginal serrations with 

Ulmus owyheensis H. V. Sm., which was originally 

described from the Miocene Sucker Creek flora. As 

discussed elsewhere (Tanai and Wolfe, 1977), the 

leaves of this species vary in laminar shape from 

ovate to oval and in basal shape from rounded to 

broadly rounded. This species, however, is distinguishable from other Tertiary elms by the following 

characters: single, apiculate teeth having in many instances a minute subsidiary tooth, less unequal and 

slightly cordate base, irregularly percurrent tertiary 

veins, and three or four prominent tertiary branches 

that depart from a basal pair of secondaries and end 

in the teeth. A single specimen figured as U. plurinervia from English Bay (Heer, 1869a) is included in U. 

owyheensis.

Hypotypes.—USNM 208376, 208377. 

Occurrence.— 9856, 9858.

Ulmus speciosa Newberry 

Plate 4, figures 3, 5

Ulmus speciosa Newberry, 1898, U.S. Geol. Survey 

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

2, 5-8).

- Tanai and Wolfe, 1977, U.S. Geol. Survey Prof. 

Paper 1026, p. 8, pi. 3C, F [see synonymy and 

discussion].

Discussion.—Two leaves, although lacking about 

half the base, match well in their venation and marginal characters Ulmus speciosa, which was recently 

reinstituted (Tanai and Wolfe, 1977). The primary 

teeth of our Alaskan specimens typically have two 

subsidiary teeth of nearly equal size, fed by tertiary 

branches from the secondaries. The intercostal tertiary veins are mostly percurrent, crossing to the secondaries, and enclose irregularly polygonal nets 

formed by fourth order veins.

Hypotypes—USNM 208381, 208382.

Occurrence. —9858.

Genus Zelkova Spach 

Zelkova browni Tanai et Wolfe

Plate 5, figures 3, 6, 8a, 8c; plate 6, figure 8

Zelkova browni Tanai and Wolfe, 1977, U.S. Geol.

Survey Prof. Paper 1026, p. 8, pi. 4A, C-G [see

synonymy and discussion]. 

Planera ungeri auct. non Ettingshausen. Heer, 1869,

Kongl. Svens. Vet.-Akad. Handl., bd. 8, no. 4, p.

34, pi. 5, fig. 2.

Discussion.—This species was recently established 

on the basis of well-preserved material from the 

Miocene Collawash flora of Oregon, because the type 

specimen of Z. oregoniana was ascertained to belong 

to Ulmus (Tanai and Wolfe, 1977). Zelkova browni is 

distinguishable from Z. ungeri in marginal characters: larger and rather bluntly deltoid teeth, sometimes with a minute subsidiary tooth. Leaves of this 

species are highly variable in shape; some have a 

slightly cordate base and a minute subsidiary tooth on 

the primary teeth and superficially resemble some 

leaves of Ulmus owyheensis H. V. Smith. Ulmus 

owyheensis, however, differs from Z. browni in the 

serration of basal part of the lamina, the tertiaries 

branching outward from the basal pair of secondary 

veins, and the irregularly percurrent intercostal tertiary veins. Zelkova browni is similar to the modern Z. 

carpinifolia Spach of the Caucasus region in foliar 

shape and marginal serration.

A single specimen figured as Planera ungeri from 

"English Bay" (Heer, 1869a) has the margin of Z. 

browni, although the lower half of the blade is lacking. It is significant in the past distribution of Zelkova 

that all specimens from the conterminous United 

States are of the Z. browni type and that this species 

lived together with the Eurasian species Z. ungeri in 

Alaska during the Miocene. 

Hypotypes.— USNM 208384-208388. 

Occurrence.— 9856, 9858.
SYSTEMATICS 31

Zelkova ungeri Kovats 

Plate 5, figures 4, 7; plate 6, figures 1, 2, 6

Zelkova ungeri Kovats in Unger, 1852, Iconogr., p. 42,

pi. 20, fig. 19. 

Kovats, 1856, Fossile Flora von Erdobenye, p. 27,

pi. 5, figs. 6, 7.

Tanai, 1961, Hokkaido Univ. Fac. Sci. Jour., ser. 

4, v. 10, no. 2, p. 322, pi. 18, figs. 1-4, 6-9, 11.

Discussion.—A number of our Alaskan specimens 

are identical with Zelkova ungeri, which is common in 

the Neogene of Eurasia. These leaves are characterized by apiculate, usually single-serrate teeth with 

somewhat incurved pointing tip. The tertiary veins in 

the intercostal areas are mostly wavy and typically 

appear to form irregularly quadragular or pentagonal meshes due to thick connecting fourth order 

veins. These marginal and venation characters show 

that this species is closely related to the modern Z. 

serrata Mak. from Japan and Z. sinica Schn. from 

mainland China.

As far as we know, Z. ungeri has not been found 

in the Tertiary of the conterminous United States. 

Becker (1969) reported a leaf of Z. ungeri from the 

Beaverhead basin of southern Montana, but his specimens appear to be referable to Z. browni in characters of the serrations.

Hypotypes.—USNM 208389-208393.

Occurrence.— 9856, 9858.

Family Fagaceae 

Genus Fagus Linn. 

Fagus antipofi Heer

Plate 5, figure 8b; plate 6, figures 3, 4; plate 7, figures 1, 5, 6

Fagus antipofi Heer, in Abich, 1858, Acad. Sci. St.

Petersburg Mem., ser. 6, v. 7, p. 572, pi. 8, fig.

2. 

Heer, 1869, Flora Fossilis Arctica, v. 2, pt. 2, p.

30, pi. 5, fig. 4a; pi. 7, figs. 4-8; pi. 8, fig. 1. 

Hollick, 1936, U.S. Geol. Survey Prof. Paper 186,

p. 95, pi. 53, figs. 1, 2. 

Castanea ungeri Heer 1869, Flora Fossilis Arctica, p.

32, pi. 7, figs. 1-3. 

Fagus macrophylla auct. non Unger. Heer, 1869,

Flora Fossilis Arctica, p. 31, pi. 8, fig. 2. 

Discussion.—Leaves referred to Fagus are very 

common in the Seldovia Point localities; these leaves 

are generally elliptical to oblong in shape, typically 

serrate with minute teeth, and have 14 to 26 pairs of 

secondary veins. Although highly variable in gross 

shape and size, these Alaskan leaves match well those 

of Fagus antipofi in venation and marginal characters. 

The specimen figured as F. macrophylla from the "English Bay" locality by Heer (1869a) represents a large 

leaf of F. antipofi. Heer also described three largetoothed leaves as Castanea ungeri from the same locality; these are included in F. antipofi, because we also 

collected several leaves having a similar margin. Our 

material lacks the secondary vein termination near 

the teeth characteristic of Castanea. In measurements 

of many of the Seldovia Point specimens, including 

Heer's material, the foliar characters of the Alaskan 

beech such as leaf indices (length to width ratio) is 

well consistent with measurements of F. antipofi from 

East Asia, as discussed elsewhere (Tanai, 1973).

Hypotypes.— USNM 208394-298399.

Occurrence.— 9856, 9858.

Fagus aff. F. crenata Blume 

Plate 6, figure 5

Discussion.—A single elliptical leaf has, though incomplete, features characteristic of Fagus in venation 

and margin: the midvein is somewhat zigzag in its 

upper portion, the secondaries abruptly arise up along 

the margin, and the margin is sinuately undulate. 

Judging from the restoration, this leaf probably has 

less than 13 pairs of secondary veins. All these 

characters show that this specimen is closely similar 

to leaves of the extant F. crenata Blume in Japan. 

While F. crenata has been known to have appeared 

since the late Pliocene in Japan (Tanai, 1973), this 

Alaskan fossil beech is, though fragmentary, noteworthy for the evolutionary history of beech.

An undescribed leaf from the Oligocene part of the 

Kenai Group (loc. 9884) has a margin of the Fagus 

crenata type. The number of secondary veins is, however, well over 13, and in this feature the leaf is more 

similar to leaves of F. antipofi. We tentatively suggest 

that the lineage to which F. crenata belongs diverged 

from the F. antipofi line at high latitudes and has 

since migrated southward.

Specimen.—USNM 208400.

Occurrence. —9856.

Genus Quercus Linnaeus 

Quercus furuhjelmi Heer

Plate 7, figures 2-4; plate 8, figures 1-3

Quercus furuhjelmi Heer, 1869, Flora Fossilis Arctica,

v. 2, pt. 2, p. 32, pi. 5, fig. 10; pi. 6, figs. 1, 2. 

?Hollick, 1936, U.S. Geol. Survey Prof. Paper 182,

p. 98, pi. 55. 

Quercus pandurata Heer, 1869, Flora Fossilis Arctica,

v. 2, pt. 2, p. 33, pi. 6, fig. 6. 

Populus leucophylla auct. non Unger. Heer, 1869,

Flora Fossilis Arctica, v. 2, pt. 2, p. 26, pi. 2, fig.

6. 

Quercus etymodrys auct. non Unger. Hollick, 1936,

U.S. Geol. Survey Prof. Paper 182, p. 98, pi. 54,

figs. 6, 7.
32 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Quercus pseudocastanea auct. non Goeppert. Heer, 

1869, Flora Fossilis Arctica, v. 2, pt. 2, p. 32, pi. 

6, figs. 3-5.

Hollick, 1936, U.S. Geol. Survey Prof. Paper 182, 

p. 98, pi. 54, figs. 4, 5.

Discussion.—The specimen assigned to this species 

by Hollick (1936, pi. 55) is fragmentary but does have 

several points of similarity to Quercus furuhjelmi. 

Rollick's specimen, however, appears to have considerably more secondary veins than in any specimen of 

Q. furuhjelmi; this apparent difference, as well as the 

fragmentary nature of the specimen, makes specific 

assignment to Q. furuhjelmi questionable.

The numerous large serrations of these leaves allies 

Quercus furuhjelmi to the so-called chestnut oaks. The 

chestnut oaks are, however, an apparently polyphyletic group. The chestnut oaks of eastern Asia (for 

example, Q. crispula Blume) have been placed in the 

subgenus Heterobalanus in contrast to the chestnut 

oaks of eastern North America (for example, Q. 

prinoides Willd.), which are placed in subgenus 

Leucobalanus (Trelease, 1924); the one chestnut oak 

of western North America—Q. sadleriana R. Br.—has 

been allied to Leucobalanus by some authorities and 

to Heterobalanus by others. Examination of cleared 

leaves of almost all species of chestnut oaks does not 

indicate any fundamental distinction between foliage 

of these groups, although some species of Heterobalanus have some aristate teeth, which are absent in 

Leucobalanus. Some specimens of Q. furuhjelmi also 

possess some aristate teeth (pi. 8, fig. 3), which indicates that this species is probably allied to 

Heterobalanus.

Quercus furuhjelmi is similar to the Korean and 

Japanese Q. microcrispula Huz., which has been compared to both Q. crispula Blume (Huzioka, 1954) and 

to Q. mongolica Fisch. var. grosseserrata (Blume) R. et 

W. (Tanai, 1961). In having teeth that are narrowly 

rounded in the basal part of the lamina and that 

grade into acute to acute-aristate teeth near the apex, 

Q. furuhjelmi is distinct from these fossil and extant 

species.

Hypotypes.— USNM 208401-208406.

Occurrence.— 9856, 9858.

Family Betulaceae

Genus Alnus Linnaeus

Alnus cappsi (Hollick) Wolfe

Plate 8, figure 5

Alnus cappsi (Hollick) Wolfe, 1966, U.S. Geol. Survey 

Prof. Paper 398-B, p. B19, pi. 6, figs. 1, 4; pi. 7, 

figs. 2, 6; text-fig. 6 [see synonymy].

Crataegus cappsi Hollick, 1936, U.S. Geol. Survey 

Prof. Paper 182, p. 86, pi. 49, fig. 3.

Discussion.—The close relationship of Alnus cappsi 

to the Alaskan Oligocene A. evidens indicates that 

this lineage is of Beringian origin.

Hypotype.— USNM 208407.

Occurrence.—9858.

Alnus fairi (Knowlton) Wolfe

Plate 8, figure 4; plate 9, figure 2

Alnus fairi (Knowlton) Wolfe, 1966, U.S. Geol. Survey 

Prof. Paper 398-B, p. B17, pi. 7, fig. 3; text-fig. 3 

[see synonymy.] 

Betula fairii Knowlton, 1926, U.S. Geol. Survey Prof.

Paper 140, p. 33, pi. 17, fig. 4.

Discussion.—Although Becker (1973) ignored the 

transfer of the Latah "Betula" fairii to Alnus, the type 

of ultimate venation possessed by both the Alaskan 

and Latah material (see Wolfe, 1966, text-fig. 3) of 

this species does not occur in Betula. 

Hypotypes.—USNM 208408, 208409. 

Occurrence.— 9856 (Heer's material), 9858.

Alnus healyensis Wolfe 

Plate 8, figure 6; plate 9, figure 1

Alnus healyensis Wolfe, 1966, U.S. Geol. Survey Prof.

Paper 398-B, p. B17, pi. 7, fig. 4, text-fig. 4. 

Artocarpidium alaskanum Hollick, 1936 [part], U.S. 

Geol. Survey Prof. Paper 182, p. 108, pi. 59, fig. 5. 

Quercus oregoniana auct. non Knowlton. Hollick, 

1936, U.S. Geol. Survey Prof. Paper 182, p. 103, 

pi. 50, fig. 5.

Alnus relatus auct. non (Knowlton) Brown. Chaney 

and Axelrod, 1959 [part], Carnegie Inst. Washington Pub. 617, p. 159, pi. 22, fig. 6 only. 

Discussion.—Alnus healyensis occurs in both the 

early (Collawash flora) and middle (Latah, Mascall) 

Miocene in the conterminous United States. In 

Alaska, this species occurs in the lower (Capps 

Glacier, Houston, Upper Healy Creek) and upper 

(Seldovia Point, Sanctuary/Suntrana) Seldovian. A 

species having, as in A. healyensis, sharp and basally 

reflexed secondary teeth is represented in the 

Oligocene Bridge Creek flora of Oregon (Klucking, 

1959), and thus A. healyensis is thought to be of West 

American derivation. The Eocene A. kluckingi Wolfe 

also has similar but fewer teeth. 

Hypotypes.— USNM 208410, 208411. 

Occurrence.—9856 (Hollick's material from Coal 

Cove), 9858.

Genus Betula Linnaeus

Betula cf. B. sublutea Tanai et N. Suzuki

Plate 9, figure 8

Betula sublutea Tanai and N. Suzuki, 1963, Tertiary 

floras of Japan, Miocene floras, p. 114, pi. 8, fig. 8; 

pi. 10, figs. 7-9.
SYSTEMATICS 33

Discussion.—A single leaf is referred to Betula on 

the basis of its shape and marginal serration, particularly on the basis of the apically pointing teeth 

and secondary veins. It is nearly identical withfi. sublutea in having rather obtuse teeth and a cordate 

base, though the detailed venation is not visible. Our 

Alaskan leaf is different from the finely and sharply 

serrate B. vera R. W. Br., which was described from 

the Miocene of Washington.

Specimen.—USNM 208413.

Occurrence.—9858.

Genus Carpinus Linnaeus

Carpinus seldoviana Wolfe

Plate 9, figure 7; plate 10, figures 1-3

Carpinus seldoviana Wolfe, 1966, U.S. Geol. Survey 

Prof. Paper 398-B, p. B23, pi. 6, fig. 6; text-fig. 

11D.

Supplementary description.—Involucre obovate in 

shape, asymmetrically cuneate at base, rounded but 

abruptly pointed apex, 19 mm long and 9.6 mm wide; 

palmately three veined, the principal veins leaving 

the base, entering large teeth; secondary veins thin, 

diverging at nearly rights angles or obliquely from 

the principal veins, forming irregular networks with 

thinner tertiary veins; margin compoundly serrate in 

the upper half, sparse, minute teeth, excepting the 

principal three lobes; nutlet not preserved.

Discussion.—This species is characterized by leaves 

having a strongly cordate base and apiculate marginal teeth with aristate tips, thus closely resembling 

the modern Carpinus cordata Blume living in northeastern Asia. The intercostal tertiary veins are nearly 

perpendicular to the secondaries, and typically bifurcate halfway between the secondaries. The fourth and 

fifth order veins form quadrangular or pentagonal 

networks, which are rarely intruded by single 

branched veinlets. These venation characters also resemble well those of the extant species. An incomplete 

involucre is closely similar to those of the extant C. 

cordata in shape and venation and is included in C. 

seldoviana.

C. seldoviana occurs abundantly at the Seldovia 

Point locality and rarely at the Port Graham locality. 

This Alaskan species is probably related to C. subcordata Nathorst, which is common in the Miocene of Japan.

Hypotypes.— USNM 208414-208417.

Occurrence.— 9856, 9858.

Genus Corylus Linnaeus

Corylus sp. 

Plate 9, figure 3

Discussion.—In shape of the teeth, these fragmentary leaves are similar to the extant Corylus chinensis

Franch. The fossils, however, have areoles about half 

the size of the areoles of this extant species; in this 

feature, the fossils are more similar to species such as 

C. rostrata (Ait.) Pursh. These fossils thus represent a 

new species, but we consider the present material to 

be too fragmentary for the basis of a specific concept. 

One of the two specimens on which Heer (1869a, pi. 

10, fig. 2) based his Tilia alaskana represents the 

same species, but that specimen (as well as the second 

fragment that is not Corylus) represents less than half 

a leaf; we, therefore, reject Tilia alaskana Heer 

(1869, p. 36, pi. 10, figs. 2, 3) under Article 70 of the 

International Code of Botanical Nomenclature.

Specimens.—USNM 208418, 208419.

Occurrence.—9856, 9858.

Genus Ostrya Scop.

Ostrya cf. O. oregoniana Chancy

Plate 9, figures 4-6

Ostrya oregoniana Chaney, 1927, Carnegie Inst. 

Washington Pub. 346, p. 106, pi. 9, fig. 12; pi. 10, 

figs. 1, 4.

Discussion.—Our involucre specimens of Ostrya 

vary from elliptical to oval in shape and 7 to 11 in 

number of primary veins; these are nearly identical 

with the involucres of O. oregoniana, which was defined by both leaves and involucres. Unfortunately we 

did not collect any leaves of Ostrya at the Seldovia 

Point localities. These Alaskan specimens are, therefore, only tentatively assigned to O. oregoniana, because of the difficulty of making specific separations 

in Ostrya on the basis of involucres alone.

Specimens.—USNM 208420-208422.

Occurrence.—9858.

Family Juglandaceae

Genus Carya Nuttall

Carya bendirei (Lesquereux) Chaney et Axelrod

Plate 10, figure 7; plate 11, figures 1, 2

Carya bendirei (Lesquereux) Chaney and Axelrod,

1959 [part], Carnegie Inst. Washington Pub.

617, p. 155, pi. 19, figs. 1-5 [not many items

synonymized.] 

Wolfe, 1964, U.S. Geol. Survey Prof. Paper

454-N, p. N20, pi. 1, fig. 7. 

Graham, 1965, Kent State Univ. Bull., Research

Ser., 9, p. 81, pi. 12, fig. 8. 

Rhus bendirei Lesquereux, 1888, U.S. Natl. Mus.

Proc., v. 11, p. 15, pi. 9, fig. 2. 

Salix engelhardti Lesquereux, 1888, U.S. Natl. Mus.

Proc., v. 11, p. 17, pi. 8, fig. 2. 

Hicoria dentata Chaney, 1920, Walker Mus. Contr., v.

2, p. 163, pi. 8, fig. 1.
34 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Hicoria orientalis Chaney, 1920, Walker Mus. Contr.,

v. 2, p. 163, pi. 8, fig. 2. 

Hicoria pecanoides Chaney, 1920, Walker Mus.

Contr., v. 2, p. 164, pi. 8, fig. 3. 

Carya egregia auct. non Juglans egregia Lesquereux.

LaMotte, 1936, p. 116, pi. 4, figs. 5, 6; pi. 6, figs.

1,2. 

Brown, 1937, U.S. Geol. Survey Prof. Paper 186,

p. 169, pi. 57, fig. 4.

Beck, 1938, Mineralogist, v. 6, no. 8, p. 22, textfig. 3. 

Smith, 1939, Michigan Acad. Sci. Papers, v. 24, p.

Ill, pl.7, fig. 1. 

Smith, 1941, Am. Midland Naturalist, v. 25, p.

500, pi. 5, fig. 3. 

Juglans egregia auct. non Lesquereux. Berry, 1931,

U.S. Geol. Survey Prof. Paper 170, p. 35, pi. 11,

fig. 3. 

Ptelea miocenica auct. non Berry. LaMotte, 1936

[part], Carnegie Inst. Washington Pub. 455, p.

133, pi. 11, fig. 4 only. 

Arbutus matthesii auct. non Chaney. LaMotte, 1936

[part], Carnegie Inst. Washington Pub. 455, p.

140, pi. 14, fig. 3 only.

Discussion—Many of the items synonymized under 

Carya bendirei are rejected by us. For example, the 

types of Prunus? merriami Knowlton (1902, p. 67) 

represent Cyclocarya. Additionally, Carya simulata 

(Knowlt.) R. W. Brown (1937, p. 169) is considered to 

represent a valid species distinct from C. bendirei. In 

C. simulata, (1) the secondary veins fork conspicuously to give a ladderlike appearance to the secondary 

external veins and (2) the apical and basal sides of the 

teeth are equal in length.

The Seldovia Point leaflets are assigned to Carya on 

the basis of (1) finely serrate margin with uniformly 

spaced and sized teeth, (2) mixed craspedodromous 

and camptodromous secondary venation, (3) central 

entry of the teeth by secondary or external secondary 

veins, (4) fourth rank venation, and (5) typically 

quadrangular areoles formed by fifth order veins and 

either lacking veinlets or intruded by simple veinlets. 

Carya is also known from the Neogene of eastern Asia 

(Hu and Chaney, 1940; Tanai and Suzuki, 1963), but 

the Asian C. miocathayensis Hu et Chan. is consistently camptodromous, has about 25 pairs of secondary veins, and typically has a narrowly rounded 

base. In contrast, the Seldovian material—as in material of C. bendirei from the conterminous United 

States—has some camptodromous and some craspedodromous secondary veins, has fewer (about 

12-18) pairs of secondary veins, and has some leaflets 

with a broadly rounded base. No extant species of 

Carya closely resembles C. bendirei. In members of

section Carya, which is exclusively North American, 

the teeth have a pronouncedly elongated basal side 

and tend to be pointed apically. In members of section 

Apocarya, which is disjunct between North America 

(Mexico and eastern United States) and east Asia 

(Vietnam and China), the teeth have apical and basal 

sides of almost equal length and many of the teeth are 

pointed abmedially. On this basis, C. bendirei is referrable to Apocarya.

Hypotypes.—USNM 208423, 208424.

Occurrence.— 9856, 9858.

Genus Cyclocarya Iljinskaja

Cyclocarya ezoana (Tanai et N. Suzuki) Wolfe et Tanai, comb.

nov.

Plate 10, figures 4-6, 8, 9

Pterocarya ezoana Tanai and N. Suzuki, 1963, Tertiary Floras Japan, v. 1, p. 110, pi. 6, figs. 2-5, 

8, 9, 11; pi. 19, fig. 1; pi. 21, fig. 10. 

Ishida, 1970 [part], Kyoto Univ. Fac. Sci. Mem., 

Ser. Geology and Mineralogy, v. 37, no. 1, p. 70, 

pi. 5, fig. 2 only.

Discussion.—In establishing this species, Tanai and 

Suzuki (1963, p. 110) noted the close resemblance to 

leaflets of the extant Pterocarya paliurus Skan. This 

extant species, however, is now generally accorded 

monotypic status as Cyclocarya paliurus (Skan.) Ilj., 

and we consequently transfer the fossil material to 

Cyclocarya.

As noted by Tanai and Suzuki (1963, p. 110), Cyclocarya ezoana is readily distinguished from Pterocarya asymmetrosa in that the secondary veins bend 

sharply towards the apex about two-thirds of the distance from the midrib to the margin and in having 

sharper and more widely spaced teeth. The same 

criteria also distinguish C. ezoana from P. nigella. 

Additionally, the areoles of Cyclocarya (including C. 

ezoana) are about one-half the size of the areoles in 

Pterocarya.

Leaflets similar to the extant Cyclocarya also occur 

in the Neogene of the conterminous United States 

(Wolfe, 1959), but the material from the Pacific 

Northwest is specifically distinct from the Alaskan 

and Japanese material. In particular, the teeth of the 

material from the Pacific Northwest are broader, and 

the areoles are large as in Pterocarya, although in 

secondary venation and overall shape of the teeth the 

material is most similar to Cyclocarya. We suggest 

that the species from the Pacific Northwest represents 

an extinct lineage of Cyclocarya; the leaflets in the 

Collawash flora are associated with a fruit of Cyclocarya.

Hypotypes.—USNM 208425-208429.

Occurrence.— 9856, 9858.
SYSTEMATICS 35

Genus Pterocarya Kunth 

Pterocarya nigella (Heer) Wolfe

Plate 11, figures 4, 6

Pterocarya nigella (Heer) Wolfe, 1966, U.S. Geol. Survey Prof. Paper 398-B, p. B15, pi. 3, fig. 3. 

Juglans nigella Heer, 1869, Flora Fossilis Arctica, v.

2, pt. 2, p. 38, pi. 9, figs. 2-4.

Juglans oregoniana Lesquereux, 1878, Harvard Coll. 

Mus. Comp. Zoology, Mem., v. 6, no. 2, p. 35, pi. 

9, fig. 10.

Salix varians auct. non Goeppert. Heer, 1869 [part], 

Flora Fossilis Arctica, v. 2, pt. 2, p. 27, pi. 2, 

fig. 8 only.

Pterocarya mixta auct. non (Knowlton) Brown.

Chaney and Axelrod, 1959, Carnegie Inst.

Washington Pub. 617, p. 157, pi. 21. figs. 1, 2.

Axelrod, 1964, California Univ. Pubs. Geol. Sci.,

v. 51, p. 115, pi. 10, figs. 8, 9.

Discussion.—As noted previously (Wolfe, 1966, p. 

B15), these leaflets have features that ally them to the 

extant members of section Platyptera. In particular, 

leaflets of P. rhoifolia Sieb. et Zucc. are similar to the 

fossils. The Japanese Neogene P. asymmetrosa Kon'no 

is also closely similar to P. rhoifolia (Tanai and 

Suzuki, 1963, p. 110), and probably some material 

now referred to P. asymmetrosa is conspecific with P. 

nigella. Other material referred to P. asymmetrosa 

may, however, be referrable to other species, and we 

consequently defer a formal synonymy until such time 

as the Japanese Neogene material of this species is 

thoroughly reviewed. P. nigella was probably one of 

the most geographically wide ranging species, occurring from Oregon to Alaska and to Honshu. 

Hypotypes.— USNM 208430, 208431. 

Occurrence.—9856 (Heer's material), 9858.

Pterocarya sp.

Plate 11, figure 7

Description.—Fruit representing nut with two 

wings, incomplete in preservation, more than 17 mm 

wide and 8 mm high; wings linear-oblong in shape, 

rather thin in texture; alate veins thin, more than 10 

in each wing, leaving the nut part, then branching 

once or twice toward the apical part of wing; nut 

ovoid, 6 mm high and 3.5 mm wide, pointed at apex; 

axis of nut parallel to the wing plane.

Discussion.—A two-winged nut, though poorly preserved, is certainly referable to Pterocarya; this 

specimen is closely similar to the extant P. fraxinifolia 

Skan. of the Middle East by its shape and the direction 

of nut axis. This fossil fruit may represent P. nigella, 

but leaflets of that species are more similar to P. 

rhoifolia than to P. fraxinifolia.

Specimen.—USNM 208432.

Occurrence.—9856.

Family Salicaceae 

Genus Populus Linnaeus

Populus kenaiana Wolfe 

Plate 11, figure 9; plate 12, figure 1

Populus kenaiana Wolfe, 1966, U.S. Geol. Survey Prof.

Paper, 398-B, p. B12, pi. 3, fig. 1. 

Vitis crenata Heer, 1869, Flora Fossilis Arctica, v. 2,

pt. 2, p. 36, pi. 8, fig. 6. 

Populus lindgreni Knowlton, U.S. Geol. Survey 18th

Ann. Kept., pt. 3, p. 725, pi. 100, fig. 3. 

Chaney and Axelrod, 1959, Carnegie Inst. 

Washington Pub. 617, p. 151, pi. 17, figs. 1-3. 

Vitis heeriana Knowlton and Cockerell, 1919, U.S.

Geol. Survey Bull. 696, p. 648. 

Discussion.—Although a member of the aspen 

group of Populus, relationships to extant species appear distant. This or a closely related species occurs in 

the middle and late Oligocene Alaskan flora; the 

youngest occurrence of the species in both Alaska and 

the conterminous United States is late Miocene. 

Hypotypes.— USNM 42264B, 208433. 

Occurrence.—9856 (Heer's material), 9858.

Populus sp.

Plate 11, figure 8

Discussion.—This fragmentary specimen has lateral primary veins that extend more admedially than 

apically (as is typical in Populus kenaiana). Although 

representing a separate species, the material is inadequate for specific determination.

Specimen.—USNM 208434.

Occurrence.—9858.

Genus Salix Linnaeus

Four morphologic types can be differentiated in the 

Seldovia Point material of Salix. The previously described S. picroides is characterized by the small, apically pointing teeth that may be appressed to the 

lamina; the tertiary venation is irregular but tends to 

be percurrent, and the lamina tends to be narrowly 

oblanceolate. In shape, S. cappsensis is similar to S. 

picroides, but the former species has large, widely 

spaced, and basally reflexed teeth and the tertiary venation is more uniformly percurrent. Salix seldoviana 

has an elliptical shape, but most characteristic are the 

small, closely spaced, and irregularly shaped teeth. 

Salix hopkinsi is almost entire margined, has a narrowly elliptic shape, and has closely spaced tertiary 

veins.

None of the Seldovia Point species of Salix are 

known outside of Alaska; the material listed as S. inquirenda Knowlt. by Wolfe (1966) is now considered to 

represent S. picroides. Salix seldoviana is similar to
36 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

some leaves attributed to S. varians Goepp., but 

leaves of the varians type from the German Miocene 

do not have a margin as finely serrate as S. seldoviana. The relationships of the Seldovia Point 

species of Salix to extant species are not clear. Salix 

cappsensis is thought to be in the same lineage that 

gave rise to the extant Alaskan S. richardsonii Hook., 

and S. picroides has some similarities to the extant S. 

pseudomonticola Ball (Wolfe, 1966); these suggested 

relationships, as well as the relationships of S. seldoviana and S. hopkinsi, require further study, particularly considering the large amount of as yet specifically undetermined material of Salix in the collections from the Homerian and Clamgulchian. This 

material should provide much information on the relationships of the Seldovia Point willows to extant 

species.

Salix cappsensis Wolfe 

Plate 12, figure 8; plate 13, figures 1, 2, 4; plate 14, figures 3, 4

Salix cappsensis Wolfe, 1966, U.S. Geol. Survey Prof. 

Paper 398-B, p. B12, pi. 4, fig. 6.

Discussion.—A closely related but undescribed 

species occurs in Oligocene beds at Kukak Bay; this 

older species has fewer teeth than Salix cappsensis. As 

noted previously (Wolfe, 1966), S. cappsensis appears 

to be ancestral to the Homerian S. chuitensis Wolfe 

and, in turn, to the extant S. richardsonii Hook.

Hypotypes.— USNM 208435-208439.

Occurrence. —9858.

Salix hopkinsi Wolfe et Tanai, sp. nov.

Plate 12, figure 4; plate 13, figures 5, 6

Description.—Leaves simple, venation pinnate; 

shape narrowly elliptic, apex acute, base acuminate; 

length 4.5 to 7.0 cm, width 2.0 to 2.8 cm; about seven 

pairs of secondary veins, irregularly spaced, departing 

at an angle of 60 to 80 degrees, some slightly decurrent, bending sharply towards apex, eucamptodromous, approaching margin closely; intersecondaries 

numerous, gradational with tertiary veins; tertiary 

veins irregularly spaced, oblique to secondary veins, 

branching, sinuous; fourth order venation forming a 

highly irregular reticulum of thinning veins and apparently forming areoles; margin entire except for a 

few irregularly spaced small teeth; petiole incomplete, 

at least 0.5 cm long.

Discussion.—Leaves of Salix from the Homerian 

appear similar to Salix hopkinsi; whether these 

younger specimens are conspecific with S. hopkinsi 

requires further study.

We take pleasure in naming this species for David 

M. Hopkins, who has aided this study in numerous 

ways.

Holotype.—USNM 208440. 

Paratype.—USNM 208441. 

Occurrence. —9858.

Salix picroides (Heer) Wolfe

Plate 12, figures 5-7; plate 13, figures 3, 7

Salix picroides (Heer) Wolfe, 1966, U.S. Geol. Survey

Prof. Paper 398-B p. B14.

Juglans picroides Heer, 1869, Flora Fossilis Arctica, 

v. 2, pt. 2, p. 39, pi. 9, fig. 5.

Discussion.—Salix picroides is also represented in 

collections from the lower Seldovian at Capps Glacier 

and Houston. In the Nenana coalfield, S. picroides 

also occurs in both the lower and upper Seldovian.

Hypotypes.— USNM 208442-208445.

Occurrence.— 9856 (Heer's material), 9858.

Salix seldoviana Wolfe et Tanai, sp. nov.

Plate 12, figures 2, 3; plate 14, figure 2

Description.—Leaves simple, pinnately veined; 

shape elliptical, apex not known, base cuneate; length 

8.5 to more than 10 cm, width 5 to 6 cm; about nine 

pairs of secondary veins, irregularly spaced, departing 

at an angle of 50 to 70 degrees, eucamptodromous, approaching margin closely; a few short intersecondary 

veins; tertiary veins oblique to secondary veins, tending to be oriented perpendicular to midrib in outer 

part of intercostal area, percurrent to branching, 

somewhat sinuous; fourth order venation forming a 

reticulation of irregular shape and size; fifth order and 

ultimate venation indistinct; margin finely serrate, 

basal side of tooth convex, about twice as long as apical side, which is slightly convex to straight; teeth entered centrally by branches of secondary veins; petiole 

incomplete, at least 0.5 cm long.

Discussion.—The finely serrate margin combined 

with the few secondary veins and few short intersecondary veins separate this species from others found 

at Seldovia Point. Salix seldoviana does not closely resemble any other known Alaskan Tertiary Salix.

Holotype.—USNM 208446.

Paratype.—USNM 208447.

Occurrence. —9858.

Family Tiliaceae

Genus Tilia Linnaeus

Tilia subnobilis Huzioka

Plate 15, figure 1

Tilia subnobilis Huzioka, 1943, Hokkaido Univ. Fac. 

Sci. Jour., ser. 4, v. 7, no. 1, p. 125, pi. 22, figs. 

2,3.

Tanai, 1961, Hokkaido Univ. Fac. Sci. Jour., v. 

11, no. 2, p. 370, pi. 29, fig. 1.
SYSTEMATICS 37

Discussion.—A single leaf is referred to the genus 

Tilia on the basis of its venation and marginal serrations with glandular teeth. The tertiary veins are 

nearly straight or slightly convex and percurrent; the 

fourth order veins are thin, forming quadrangular or 

pentagonal meshes that are penetrated by once- or 

twice-branched veinlets. This Alaskan specimen is referred to T. subnobilis in the oval shape, deeply cordate base and venation, although the marginal teeth 

are smaller in the Alaskan leaf. T. subnobilis has 

been reported from a few localities in the lower and 

middle Miocene of Hokkaido and Korea. Our specimen 

shows a slight resemblance in general outline to T. 

oregona LaMotte from the Bridge Creek flora of Oregon (LaMotte, 1935) but differs in venation characters.

Hypotype.— USNM 208448.

Occurrence.—9856.

Family Hydrangeaceae 

Genus Hydrangea Linnaeus

Hydrangea sp. 

Plate 14, figure 1

Discussion.—A single leaf and its counterpart, although fragmentary, show characters of the genus 

Hydrangea in having finely and closely serrate margin and the secondary veins loop well within the margin. Superficially these specimens resemble Prunus elliptica but differ in the sharply rising secondary veins 

and marginal looping. This Alaskan specimen is 

closely similar in venation to an incomplete leaf described as Ilex insignis Heer from the "English Bay" 

(Heer, 1869a) locality but differs in the marginal 

teeth, if his specimen was accurately illustrated.

Specimen.—USNM 208449A, B.

Occurrence.—9858.

Family Rosaceae

Genus Crataegus Linnaeus

Crataegus chamissoni (Heer) Wolfe et Tanai, comb. nov.

Plate 14, figures 5, 6

Quercus chamissoni Heer, 1869, Kongl. Svenska 

Vet.-Akad. Handl., v. 8, p. 33, pi. 6, figs. 7, 8.

Discussion.—The type specimens of Quercus 

chamissoni have a cuneate base, irregularly 

duplicate-dentate teeth, and irregularly spaced secondary veins that enter the teeth that are mixed 

craspedodromous-camptodromous. The camptodromous secondaries join with the next secondary above or 

with the intersecondaries. These characters show that 

Heer's specimens are not leaves of Quercus but are 

those of Crataegus.

Two incomplete specimens in our collections are referred to Crataegus chamissoni; one specimen is

ovate-lanceolate in shape, quite similar to one of the 

type specimens (Heer, 1869a), pi. 6, fig. 8), while 

another is obliquely ovate, having well-preserved fine 

venation. In venation and marginal characters C. 

chamissoni is similar to the extant C. douglasii Lindl. 

of western North America, which includes ovatelanceolate to oval leaves.

Hypotypes.—USNM 208450, 208451.

Occurrence.—9856 (Heer's material), 9858.

Genus Prunus Linnaeus 

Prunus kenaica Wolfe et Tanai, sp. nov.

Plate 15, figures 2, 5

Description.—Leaves elliptic to elliptic-oblong in 

shape, 10.7 to 17 cm long (estimated) and 4.2 to 5.3 cm 

wide; base gradually narrowed, broadly cuneate, apex 

unknown; midrib stout, nearly straight or slightly arcuate; secondary veins somewhat slender but distinct, 

more than 11 pairs, subopposite, somewhat irregularly spaced, diverging from the mid vein at angles of 

40 to 50 degrees in the lower half of blade, at wider 

angles in the upper, nearly straight and then turning 

up near the margin, camptodromous, looping distinctly with the next secondary above; tertiaries 

branching outward from the secondaries also looping 

well within the margin; fourth order veins branching 

from the looping tertiaries entering teeth or irregularly looping; intercostal tertiaries forming large, irregularly polygonal networks that enclose small 

fourth order nets; veinlets once or twice branching; 

margin finely serrate with inward-pointed, acute 

teeth; texture rather thin; petiole missing.

Discussion.—Three leaves and their counterparts 

are referred to the genus Prunus on the basis of their 

marginal loops and serrations. The fourth and fifth 

order veinlets show a close similarity to those of many 

modern cherry leaves. In the conterminous United 

States no cherry leaves are comparable to this new 

species. Several fragmentary leaves were described 

under the name of P. scotti Heer from Paleocene of 

Greenland (Heer, 1868, 1869b, 1880, 1883) and some 

show close resemblance in secondary venation and 

shape, although the finer venation was not figured. 

Two leaves described as P. scotti from the Mormon 

Creek flora of southwestern Montana (Becker, 1960) 

are different in venation from P. kenaica, although 

somewhat similar in shape.

Among extant species our specimens are similar to 

leaves of P. bicolor Koehne and especially P. vanioti 

Levl., which live in Szechuan.

Holotype.— USNM 208452.

Paratype.—USNM. 208453.

Occurrence. —9856.
38 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Prunus aff. P. pad us Linnaeus

Plate 14, figure 7

Description.—Leaf incomplete, almost half missing, 

broadly ovate in general shape, acute or probably 

acuminate at apex, base unknown; midrib stout, 

slightly arcuate; secondary veins distinct, about 10 

subopposite pairs, irregularly spaced, diverging from 

the midrib at angles of 40 to 50 degrees in the middle 

and upper parts of the lamina, higher angles in the 

lower part, slightly curving upward, then forming distinct, large loops with the next secondary above; some 

slender intersecondary veins leaving the midvein, 

nearly parallel to the secondaries, halfway to the 

margin connecting with the tertiaries; tertiary veins 

branching from the secondaries near the margin, 

forming small marginal loops; fourth order veins 

branching from tertiary vein loops, entering marginal 

teeth; intercostal tertiaries irregularly percurrent or 

branching, enclosing quadrangular or pentagonal 

areoles; areoles intruded by once or twice branching 

veinlets; margin coarsely duplicate serrate with acute 

teeth; texture probably thin; petiole missing.

Discussion.—A single leaf and its counterpart, although incomplete, shows venation typical of Prunus 

and is distinguishable from the P. elliptica in having 

an ovate shape and coarser marginal serrations. 

Among the fossil cherry leaves of North America, our 

specimens are somewhat similar to some leaves of 

Prunus? merriami Knowlton from the Miocene sequence of Oregon (Knowlton, 1902, pi. 11, figs. 6, 7 

(not figs. 2, 3) ) but are different in the marginal looping of the secondaries. Except for this species, no fossil 

cherry leaves are comparable to our specimens; these 

Alaskan specimens are, however, too incomplete to 

form the basis for a new species.

Among extant Prunus, our leaves are most similar 

to P. padus L. of northeastern Asia.

Specimen.—USNM 208454A, B.

Occurrence.—9858.

Genus Sorbaria A. Braun 

Sorbaria hopkinsi (Wolfe) Wolfe et Tanai, comb. nov.

Plate 16, figures 1-2

Spiraea hopkinsi Wolfe, 1966, U.S. Geol. Survey Prof. 

Paper 398-B, p. B24, pi. 8, fig. 2.

Discussion.—The generic determination of this 

species was based on comparisons to the extant 

"Spiraea" lindleyana. This modern species, however, 

is properly placed in Sorbaria, a small genus of northeast Asian shrubs.

Sorbaria has been previously found as fossil in the 

Eocene of Sakhalin (Borsuk, 1956); an undescribed 

species has also been found in the late Oligocene rocks 

of the Alaska Range (loc. 9749 of Wahrhaftig and

others, 1969), and an additional species is known in 

the early Seldovian Capps Glacier flora. The stratigraphic range of S. hopkinsi is now known to be longer 

than indicated by Wolfe, Hopkins, and Leopold (1966). 

This report extends the range downward into the 

upper Seldovian Stage, and collections made in 1967 

from the lower part of the Clamgulchian along 

Kachemak Bay also includes this species. S. hopkinsi 

is, however, much more common and widespread in 

rocks of Homerian age.

Hypotypes.— USNM 208455-208458.

Occurrence.—9858.

Family Leguminosae

Genus Cladrastis Rafinesque

Cladrastis cf. C. aniensis Huzioka

Plate 15, figure 4; plate 16, figures 3, 4

Cladrastis aniensis Huzioka, 1963, Tertiary floras of 

Japan, Miocene floras, p. 205, pi. 35, figs. 5, 6.

Discussion.—Two leaflets, although incomplete, are 

referred to the genus Cladrastis on the basis of venation characters and are closely similar to C. aniensis 

Huzioka from the middle Miocene of Japan. Our leaflets have a lower angle of departure of the secondary veins and have two more secondary veins than 

the type specimens. These differences are not thought 

to be specifically significant, but we hesitate to refer 

them to C. aniensis because our specimens are incomplete. The Alaskan specimens have the fine venation 

preserved and are closely similar to the modern C. 

platycarpa (Maxim.) Mak. living in central and western Honshu and Shikoku, Japan. An incomplete flattened pod including three seeds is linear in shape and 

has irregular networks between the dorsal and ventral ridges. This pod of Cladrastis is additional evidence of the presence of this genus in the Seldovia 

Point flora.

On the basis of well-preserved leaflets, Wolfe (1966) 

reported Cladrastis from the Homerian flora (loc. 9844 

and 9361) and the Capps Glacier flora (loc. 9845). 

These leaves were described as C. japonica by emending Nyssa japonica Tanai and N. Suzuki from the 

Miocene Yoshioka flora of Hokkaido (Tanai and 

Suzuki, 1963) and further by including C. aniensis. 

However, our further investigations reveal that the 

type specimens of "Nyssa japonica" are not Cladrastis 

(although they are also not Nyssa) but that all Alaskan leaflets described as "C. japonica" are rather similar to the modern C. lutea. This second Alaskan 

species of Cladrastis is distinguishable from C. aniensis in shape, number of the secondary veins, and irregular, well-developed, percurrent tertiary veins. Accordingly, these Cladrastis leaflets from the Homerian
SYSTEMATICS 39

and the Capps Glacier locality should be described as 

a new species.

Specimens.— USNM 208459, 208460.

Occurrence. —9858.

Genus Pueraria DeCandolle

Pueraria miothunbergiana Hu et Chancy

Plate 15, figure 3

Pueraria miothunbergiana Hu and Chaney, 1938, 

Palaeont. Sinica, New Ser. A, no. 1, p. 52, pi. 28, 

fig. IDiscussion.—A single leaflet and its counterpart 

compare well with terminal leaflets of the Pueraria in 

shape and venation characters: both the secondary 

and tertiary veins branch outward from the secondaries, gently curve up along the margin, but do not 

form distinct marginal loops; the intercostal tertiaries 

are irregularly percurrent and frequently fork once. 

Our specimens are, although small in size, identical to 

P. miothunbergiana, which has been reported from the 

lower and middle Miocene of east Asia. This Alaskan 

fossil is the first record of the genus Pueraria in North 

America.

Hypotype.— USNM 208461.

Occurrence.—9858.

Family Lythraceae

Genus Decodon J. F. Gmelin

Decodon alaskana Wolfe et Tanai, sp. nov.

Plate 15, figure 6; plate 16, figures 5, 7, 8.

Description.—Leaves simple; shape oval to linear 

oval, base acute, apex acute to acuminate; length 5 to 

more than 10 cm, width 3 to 6 cm; pinnately veined; 

12 to 15 pairs of irregularly spaced secondary veins 

departing from midrib at 80 to 90 degrees, slightly 

curving, sharply brochidodromous; one or two intersecondary veins per intercostal area; tertiary venation 

forming a large, irregular mesh, the veins at a variable angle relative to the secondaries; fourth order 

veins of irregular pattern; a series of marginal loops 

by branches of the secondary veins irregular in size 

and tending to be elongated perpendicular to the midrib; margin entire.

Discussion.—The pronounced series of brochidodromous loops that form an intramarginal vein readily 

differentiate Decodon alaskana from other members of 

the Seldovia Point flora. Such major venation is 

characteristic of members of Myrtales and allied orders (Hickey and Wolfe, 1975). Within most myrtalean families, the intramarginal vein is a continuous 

arc, but in the Seldovia Point leaves—as in leaves of 

the extant D. verticillatus (L.) Ell.—the intramarginal vein is formed by a series of arcs. Further resemblances between the fossils and modern leaves of 

Decodon are in the presence of weak loops abmedial to 

the intramarginal vein, the presence in some intercostal areas of a tertiary vein system that (1) extends directly to the intramarginal vein (intersecondary 

veins) and giving off right-angle branches to the secondary veins, or (2) extends only a short distance from 

the midrib, and the presence of a weak fourth order 

vein system. The fossils differ slightly from modern 

leaves of Decodon in having a higher proportion of intersecondary veins that extend to the intramarginal 

vein and in having the loops at the margin elongated 

perpendicular to the midrib (in D. verticillatus the 

loops are approximately square or are elongated 

parallel to the midrib).

The Seldovia Point specimens of Decodon occur on 

the same slabs as leaves of Salix. Such an association 

is to be expected in view of the marsh to swamp 

habitat of D. verticillatus and many extant species of 

Salix.

Decodon was particularly widespread in the 

Neogene of Eurasia (Dorofeyev, 1963; Eyde, 1972), but 

the Seldovia Point material represents the first occurrence of the genus as a fossil in North America. The 

leaf described as Myrtus oregonensis Lesq., however, 

probably represents Decodon; despite the specific 

epithet proposed by Lesquereux (1883), the leaf actually came from the late Miocene of California (Condit, 

1938). The same or a closely allied species also occurs 

in the Miocene Collawash flora of Oregon (Wolfe, unpub. data).

Holotype.—USNM 208462.

Paratypes.— USNM 208463, 208464.

Occurrence.—9858.

Family Trapellaceae

Genus Hemitrapa Miki

Hemitrapa borealis (Heer) Miki

Hemitrapa borealis (Heer) Miki, 1953, Palaeobotanist,

v. 1, p. 349, text-fig. 2 A. 

Trapa borealis Heer, 1869, Kongl. Svenska Vet.-Akad.

Handl., bd. 8, no. 4, p. 38, pi. 8, figs. 9-14. 

Discussion.—This species was transferred from the 

genus Trapa by Miki (1953) and is characterized by 

having two spindle-shape appendages and a welldeveloped brushy haired apical part. Heer (1869a) 

figured several specimens from the "English Bay" locality, but later collections from this and other Seldovian localities have failed to produce additional 

specimens. Hemitrapa has been described from numerous lower and middle Miocene localities of Japan 

and Sakhalien; the only putative occurrence from the 

conterminous United States (MacGinitie, 1937) is
40 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

based on material properly referable to Trapa. 

Occurrence.—9856 (Heer's material).

Family Aceraceae 

Genus Acer Linnaeus

Five species of Acer were originally thought to be 

represented in the Seldovia Point flora (Wolfe, 1966), 

but our studies indicate the presence of only four. 

Other Seldovian localities, however, have produced 

representatives of two additional species related to the 

extant A. saccharinum and A. pennsylvanicum (or A. 

rufinerve). Presumably a species of Acer related to the 

extant Asian A. mono was also a member of the Seldovian flora, because a closely related species (A. scottiae) is known from the middle and late Miocene of the 

Pacific Northwest. Clearly Acer was an important 

element in the early and middle Miocene flora of 

Alaska.

Acer ezoanum Oishi et Huzioka

Plate 17, figures 1-3, 5; plate 19, figures 2, 5

Acer ezoanum Oishi and Huzioka, 1943, Hokkaido 

Univ. Fac. Sci. Jour., ser. 4, v. 7, p. 89, pi. 10, 

figs. 1-4; pi. 11, figs. 1-4; pi. 12, fig. 2. 

Tanai and N. Suzuki, 1969, Hokkaido Univ. Fac. 

Sci. Jour., ser. 4, v. 10, p. 556, pi. 1, figs. 1, 2; pi. 

2, figs. 1, 2; pi. 3, figs. 1-4; pi. 9, figs. 20-25 [see 

synonymy for references through I960.]. 

Tanai, 1961, Hokkaido Univ. Fac. Sci. Jour., ser.

4, v. 11, p. 356, pi. 26, figs. 1, 2, 5. 

Tanai and N. Suzuki, 1963, Tertiary Floras Japan, v. 1, p. 137. 

Huzioka, 1964, Akita Univ. Mining Coll. Jour.,

ser. A, v. 3, no. 4, p. 89, pi. 14, figs. 6-9. 

Wolfe, 1966, U.S. Geol. Survey Prof. Paper

398-B, p. B25, pi. 8, fig. 6. 

Ishida, 1970, Kyoto Univ. Fac. Sci. Mem., Geology

and Mineralogy Ser., v. 37, p. 95. 

Huzioka, 1972, Akita Univ. Mining Coll. Jour.,

ser. A, v. 5, no. 1, p. 63, pi. 8, fig. 2.

Tanai and N. Suzuki, 1972, Hokkaido Univ. Fac.

Sci. Jour., ser. 4, v. 15, p. 335, pi. 7, figs. 1-5.

Discussion.—A number of leaves are referred to A.

ezoanum, which was originally described from the

lower Miocene of Hokkaido and Sakhalin. Several

winged fruits with expanded seeds are also found from

the same locality and are also referred to A. ezoanum.

Our Alaskan leaves are more closely related to those

of the modern Japanese maple, A. miyabei Maxim.,

than to any of the North American maples. Leaves of

A. ezoanum closely resemble in general appearance

those of the modern A. macrophyllum Pursh. but distinctly differ in fine venation. The quadrangular or 

pentagonal areoles are intruded by simple or once 

branched veinlets in A. ezoanum as well as in A. 

miyabei, but the irregularly four-sided areoles are intruded by twice or more branched veinlets in A. macrophyllum.

The only other known Alaskan occurrence of Acer 

ezoanum is a leaf from locality 9848 along Beluga 

River (Wolfe, 1966). This locality is apparently also of 

middle Miocene age (see p. 8).

Hypotypes.— USNM 208465-208470.

Occurrence.—9856 (fruits), 9858 (fruits and leaves).

Acer glabroides R. W. Brown

Plate 17, figures 7, 8

Acer glabroides Brown, 1937, U.S. Geol. Survey Prof. 

Paper 186-J, p. 180, pi. 58, figs. 13-15.

Discussion.—Two samaras are similar to both 

Brown's type material and to the extant Acer rubrum. 

No leaves of the rubrum type have yet been collected 

from Seldovia Point, but such leaves occur at other 

Seldovian localities (for example, loc. 9867 on Cache 

Creek).

Hypotypes.— USNM 208471, 208472.

Occurrence.—9858.

Acer grahamensis Knowlton et Cockerell

Plate 17, figure 6; plate 18, figure 5; plate 19, figures 1, 4

Acer grahamensis Knowlton and Cockerell, 1919, U.S.

Geol. Survey Bull. 696, p. 50. 

Acer macropterum Heer, 1869, Flora Fossilis Arctica,

v. 2, pt. 2, p. 37, pi. 9, figs. 8, 9 (excluding fig. 7). 

Discussion.—Two samaras are identical to Acer 

macropterum in their large size and venation of the 

wing; this species was originally described from the 

"English Bay" locality by Heer (1869a). The epithet 

instituted by Heer is, however, a junior homonym, 

and Knowlton and Cockerell (in Knowlton, 1919) proposed the name A. grahamensis for this material. Although our samaras are also incomplete as in the case 

of Heer's specimens, one of ours has an elliptical seed 

and suggests that this species may be related to the 

extant A. macrophyllum Pursh. living in the western 

United States. A fragmentary leaf was included in A. 

macropterum by Heer, but that leaf does not appear to 

be similar to A. macrophyllum. Two deeply lobed 

leaves of Acer from Seldovia Point are, although 

fragmentary, distinctly different in the secondary venation looping and the pattern of fine veinlets from 

foliage of A. ezoanum; especially the veinlets usually 

branch more than twice and are closely similar to 

those of A. macrophyllum. Accordingly, these two incomplete specimens may represent leaves of A.
SYSTEMATICS 41

grahamensis. This species is closely similar to A. 

oregonianum Knowlton from the Miocene of the conterminous western United States. Chelebaeva (1968) 

assigned samaras from the middle Miocene of Kamchatka to Acer cf. grahamensis. Although in features 

pereserved her specimens are similar to A. grahamensis, we concur with Chelebaeva that the lack of seeds 

attached to the samaras makes the specific determination uncertain.

Hypotypes.—USNM 208473-208475.

Occurrence.— 9856, 9858.

Acer heterodentatum (Chancy) MacGinitie

Plate 16, figure 6; plate 17, figure 4; 

plate 18, figures 2-4; plate 19, figure 3

Acer heterodentatum (Chaney) MacGinitie, 1953, Carnegie Inst. Washington Pub. 599, p. 140, pi. 57, 

fig. 3 [see synonymy and discussion].

Discussion.—This species is represented by two 

terminal and several lateral leaflets that show a close 

resemblance to the modern Acer negundo L. and A. 

henryi Pax. Leaflets of this type are uncommon in the 

Tertiary of the conterminous western United States, 

but fruits of the negundo type have been commonly 

reported. Our Alaskan leaflets are generally larger 

than the type specimens: the terminals are 9 to 12 cm 

(estimated) long and 8 to 9.5 cm wide, and the laterals 

are 9 to 18 cm long and 3 to 8 cm wide. These leaflets 

are more closely related in venation to A. negundo 

than to the East Asiatic A. henryi or A. cissifolium 

(Sieb. and Zucc.) K. Koch.: the fourth order veins form 

irregularly quadrangular or pentagonal meshes, 

which are intruded by unbranched or once branched 

veinlets. However, two samaras from the same locality closely resemble those of A. cissifolium in their 

shape and size of seeds.

There have been several different opinions regarding the specific name for this Tertiary boxelder. As 

already noted by MacGinitie (1953, p. 141), two types 

of Rulac crataegifolium and Phyllites bifurcies are too 

poorly preserved to serve as type specimens, though 

they resemble the modern boxelder leaflets. Although 

Rulac crataegifolium was transferred to Acer by 

LaMotte (1952, p. 53), the new combination in any 

case is homonymous for the extant A. crataegifolium 

Sieb. et Zucc. Chaney and Axelrod (1959) nominated 

"Acer minor" but the type specimen figured by 

Knowlton (1902) lacks the seed and is not an adequate 

name-bearing specimen. Thus we concur with MacGinitie (1953) and accept A. heterodentatum as the 

oldest valid combination for this species.

Hypotypes.— USNM 208476-208481.

Occurrence.—9858.

Family Nyssaceae

Genus Nyssa Linnaeus

Nyssa cf. N. knowltoni Berry

Plate 18, figure, 1

Nyssa knowltoni Berry, 1929, U.S. Geol. Survey Prof. 

Paper 154-H, p. 261, pi. 59, fig. 7.

Discussion.—One leaf, although fragmentary, is assigned firmly to Nyssa on the basis of its venation 

characters and irregularly large teeth. It closely resembles the leaves described as N. knowltoni from the 

Miocene Latah flora of Oregon (Berry, 1929; Brown, 

1937) but is too fragmentary for specific determination. Brown (1937) and Chaney and Axelrod (1959) 

transferred many Tertiary fossil leaves from the western United States resembling the modern N. aquatica 

to N. hesperia, which was originally defined on the 

basis of seeds (Berry, 1931). However, as already 

pointed out by Eyde (1963), it can be difficult to distinquish the modern N. aquatica and N. sylvatica by 

only external features of the seeds. Furthermore, 

some leaves of N. sylvatica have large irregularly 

spaced teeth, similar to those of N. aquatica. Until, 

therefore, venation characters are thoroughly investigated in west American leaves of Nyssa, the 

synonymy of N. hesperia and N. knowltoni cannot be 

accepted without scepticism.

Specimen.—USNM 208482.

Occurrence.—9858.

Family Alangiaceae

Genus Alangium Lamarck

Alangium mikii Wolfe et Tanai, sp. nov.

Plate 20, figures 1, 2; plate 21

Marlea aequalifolia (Goeppert) Oishi, 1950, Illust.

Catalogue of East Asiatic Fossil Plants, p. 171,

pi. 50, fig. 1.

Tanai, 1952, Japanese Jour. Geology and Geography, v. 22, p. 132, pi. 5, fig. 3. 

Huzioka, 1955, Illust. Fossil Catalogue in Fukui

Pref., no. 6, p. 8, pi. 3, figs. 6-8. 

Alangium aequalifolium (Goeppert) Kryshtfovich and

Borsuk, 1939, Problems Palaeont., no. 5, p. 390,

pi. 5, figs. 1-8; pi. 6, fig. 12. 

Tanai, 1961, Hokkaido Univ. Fac. Sci. Jour., ser.

4, v. 11, no. 2, p. 371, pi. 30, fig. 1; pi. 31, fig. 9. 

Huzioka, 1963, Tertiary Floras of Japan, Miocene

Floras, p. 212, pi. 38, fig. 9. 

Matsuo, 1963, Tertiary Floras of Japan, Miocene

Floras, p. 242, pi. 52, fig. 6; pi. 53, figs. 3-5

(excluding pi. 53, fig. 1). 

Huzioka, 1964, Akita Univ. Mining Coll. Jour.,

ser. A, v. 3, no. 4, p. 96, pi. 17, fig. 1. 

Huzioka, 1972, Akita Univ. Mining Coll. Jour.,

ser. A, v. 5, no. 1, p. 70, pi. 14, fig. 4. 

?Marlea iragawaense Tanai, 1952, Japanese Jour.
42 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

Geology and Geography, v. 22, p. 133, pi. 5, fig. 

5.

Description.—Leaves simple, actinodromous; shape 

symmetrical to asymmetrical and broadly oval, apex 

acute, base cordate and typically asymmetrical; 

length 10 to more than 22 cm, width 10 to more than 

22 cm; four or five pairs of lateral primaries, the first 

pair (from the midrib) paralleling the medial secondary veins and extending into the apical half of the 

lamina, five to seven lateral secondaries on basal side; 

second lateral pair of primaries paralleling secondaries of first lateral primary, also with five to seven 

pairs of lateral secondaries on basal side (these are 

absent or reduced on smaller side of highly asymmetrical laminae); third lateral primary veins conspicuous only on enlarged side of highly asymmetrical 

laminae, with three or four lateral secondary veins; 

medial secondary veins five to seven pairs, departing 

at an angle of 60 to 70 degrees, straight to broadly 

curving, brochidodromous, looping close to margin; intersecondary veins typically absent; tertiary veins 

regularly and closely spaced, perpendicular to secondaries, branched or percurrent, slightly arched abmedially; fourth order veins numerous, closely spaced, 

perpendicular to tertiaries, branching acutely; fifth 

order veins forming a polygonal areolation that is intruded by profusely branching freely ending veinlets; 

margin entire; petiole thick, incomplete but more 

than 3 cm long.

Discussion.—Considerable uncertainty and confusion exist concerning the nomenclatural and systematic position of Alangium-like foliage from the Tertiary 

of Eurasia. Many Alangium-like leaves have previously been referred to either A. aequifolium (Goepp.) 

Krysht. et Bors. or A. tiliaefolium (Ung.) Krysht.; both 

species are based on material from the European Tertiary record. Knoblock and Kvacek (1964), however, 

synonymized these two species and further considered 

them to represent a sterculiaceous plant, 

Byttneriophyllum tiliaefolium (A. Braun) Knobl. et 

Kvac. This reassignment is based on examination of 

nontype material from Tertiary rocks of Czechoslovakia, which showed that this material possessed 

compound trichomes in contrast to the uniformly simple trichomes in extant Alangium.

We emphasize that the megascopic characters of the 

leaves from which Knobloch and Kvacek obtained 

cuticular material have not been illustrated (Knobloch 

and Kvacek, 1964, p. 162). In Byttneria, as in most 

woody Malvales, the fourth and higher order venation 

is organized into pronouncedly orthogonal patterns 

(that is, the branching of any particular order is at 

approximately right angles), and the various orders of 

venation are well differentiated in size from one 

another (see Knobloch and Kvacek, 1964, pi. IV). In

Alangium, the fourth and higher order venation 

branches at definite acute angles (a Y-shaped pattern 

predominates in the fourth order venation), and the 

veins thin so gradually that the fourth and higher 

order veins are not clearly differentiated in size.

Examination of topotypic material of Byttneriophyllum tiliaefolium (=Cordia tiliaefolia A. Braun) from 

Oeningen (material in the U.S. Geol. Survey collections) substantiates Knoblock and Kvacek's (1964) 

opinion. This material has an orthogonal pattern in 

the fourth and higher order venation. Material from 

Sotzka, on which Goeppert based Dombeyopsis 

aequifolia, has not been examined; Goeppert's (1852) 

illustrations of this species are inadequate for determination, but, if accurately drawn, the specimen on 

the same slab assigned to Dombeyopsis grandifolia 

would also be malvalean because of the predominately 

orthogonal fourth order venation.

The east Asian Neogene leaves reassigned by Knobloch and Kvacek (1964) to Byttneriophyllum 

tiliaefolium, however, have intercostal venation of 

Alangium. These leaves represent the same species as 

that found at Seldovia Point and the Capps Glacier 

Seldovian. The Capps Glacier material, moreover, 

shows that the trichomes are all simple (pi. 20, fig. 2), 

leaving no reasonable doubt that Alangium is represented. The epithet iragawaense is rejected for these 

leaves because it is based on poorly preserved material. We have, therefore, established a new species, A. 

mikii, to include the Alaskan and east Asian Neogene 

leaves previously assigned to A. aequifolium.

Although Alangium was a conspicuous element in 

some Paleogene floras in North America (MacGinitie, 

1969; Eyde and others, 1969), this is the first validated occurrence for the genus in the Neogene of 

North America. Pollen of the A. barghoornianum type 

(Traverse, 1955) has, however, been found in the middle Miocene part of the Kirkwood Formation of New 

Jersey (Wolfe, unpub. data). A. barghoornianum, A, 

riparius (MacG.) MacG., and A. mikii all represent the 

section Marlea. The species of Alangium from the 

Eocene of Alaska (Wolfe, 1977), however, is a member 

of the section Alangium.

We take pleasure in naming this species in recognition of the contributions of the late Professor Shiguru 

Miki to the fossil history of the Cornales.

Hototype.—USNM 208483.

Paratypes.—USNM 208485 (loc. 11091), 208484 

(loc. 9858).

Occurrence.—9858, 11091 (Capps Glacier).

Family Araliaceae

Genus Kalopanax Miguel

Kalopanax n-suzukii Wolfe et Tanai, sp. nov.

Plate 22, figure 4

Kalopanax acerifolium auct. non (Nathorst) Hu and
SYSTEMATICS 43

Chaney. Hu and Chaney (nontypic), 1940, Carnegie Inst. Washington Pub. 507, p. 70, pi. 47, 

figs. 3, 5. 

Okutsu, 1955, Tohoku Univ. Sci. Repts., ser. 2, v.

26, p. 110, pi. 5, fig. 1. 

Tanai and Onoe, 1960, Japan Geol. Survey Kept.

187, p. 285, pi. 7, fig. 4. 

Tanai, 1961, Hokkaido Univ. Fac. Sci. Jour., ser.

4, v. 11, p. 377. 

Tanai and Suzuki, 1965, Palaeont. Soc. Japan

Spec. Paper 10, p. 43, pi. 20, fig. 2. 

Huzioka and Uemura, 1973, [Tokyo] Natl. Sci.

Mus. Bull., v. 16, p. 723, pi. 18, fig. 1. 

Description.—See Hu and Chaney (1940, p. 70- 

71).

Discussion,—Examination of Nathorst's type of 

Acanthopanax acerifolium indicates that these represent Acer and not any genus of Araliaceae. Hu and 

Chaney's (1940) Shanwang material, however, does 

represent Kalopanax and the same species that occurs 

in the Seldovia Point flora. We accept Hu and 

Chaney's (1940) description, but the unavailability of 

the specimens on which that description was based 

make it desirable to designate as types specimens that 

are in existence. We thus designate as holotype University of Hokkaido Museum of Paleontology 25727 

(figured by Tanai and Suzuki, 1965, pi. 20, fig. 2) and 

as paratypes 25726 and 25729; all these specimens are 

from the Shanabuchi locality.

Kalopanax is known from several localities in eastern Asia and is known to range from the early 

Miocene through the Holocene (Tanai, 1972). The Seldovia Point occurrence is the first known in North 

America.

We take pleasure in naming this species for Mr. 

Nobuo Suzuki, in recognition of his contributions to 

the Tertiary floras of Japan. 

Hypotype.— USNM 208486. 

Occurrence.—9856, 9858.

Family Vitaceae

Genus Vitus Linnaeus

Vitis seldoviana Wolfe et Tanai, sp. nov.

Plate 22, figure 1; plate 23, figures, 1, 3

Description.—Leaves incomplete, pentagonal in 

general outline, palmately five lobed, broadly cordate 

at base; midvein thick, nearly straight, giving off 

more than four pairs of secondaries, which are nearly 

parallel to the inner primaries; the inner pair of the 

primaries well defined, making angles of 40 to 60 degrees with the midvein, giving off about six pairs of 

secondaries which end in large teeth, the intersecondary veins leaving midrib, nearly parallel to the secondaries, camptodromous; the lowest pair of the

primaries somewhat slender, nearly at right angles to 

the midvein, with five or more secondaries; the intercostal tertiary veins thin but distinct, irregularly percurrent, nearly perpendicular to the secondaries; the 

tertiaries near the margin branching from the 

secondaries, forming loops with the intersecondaries; 

fourth and fifth order veins forming quadrangular or 

pentagonal areoles which are penetrated by once or 

twice branching veinlets; margin coarsely dentate, 

with large deltoid teeth; petiole thick, more than 2.2 

cm long.

Discussion.—Several incomplete specimens represent leaves of Vitis on the basis of their marginal dentation and vein characters. No fossil leaves of Vitis 

from North America are comparable to our Alaskan 

materials. Although all the specimens are fragmentary the general character of the foliage can be reconstructed. These leaves superficially resemble V. 

naumanni (Nathorst) Tanai, which was reported from 

the lower to upper Miocene of Japan, but differ distinctly in having larger teeth and well-defined intercostal tertiary veins.

Vitis seldoviana also superficially resembles V. alia 

Holl. from the Miocene of British Columbia (Hollick, 

1936). The latter species, however, has narrower and 

more pronounced teeth, and the tertiary veins extend 

perpendicularly from the midrib; in V. seldoviana, 

homologous tertiary veins are oriented at an acute 

angle to the midrib.

Holotype.— USNM 208487.

Paratypes.—USNM 208488, 208489.

Occurrence. —9856.

Family Oleaceae

Genus Fraxinus Linnaeus

Fraxinus kenaica Wolfe et Tanai, sp. nov.

Plate 24, figures 1-3

Description.—Presumably a leaflet; venation pinnate; shape oval; apex not preserved, base acute; 

length at least 17 cm, width 9 cm; about 13 pairs of 

secondary veins, irregularly spaced, departing at angles of 70 to 80 degrees (higher on one side of lamina), 

slightly curving, irregularly brochidodromous; intersecondaries numerous, strong; tertiary veins thin, irregularly spaced, oblique to more apical secondary 

veins and orthogonal to more basal secondary veins, 

branching irregularly; fourth order veins forming a 

thin reticulum of irregular shape and size; loops 

abmedial to secondary loops of irregular shape and 

size, one side formed by veins that extend toward 

sinuses, these veins either ending in sinuses or extending along apical side of teeth; margin coarsely 

and irregularly serrate; teeth tending to have convex 

basal and apical sides; petiolule(?) about 2 cm long.

Discussion.—The specimen gives no certain indica-
44 MIOCENE SELDOVIA POINT FLORA FROM THE KENAI GROUP, ALASKA

tion whether it is a leaf or leaflet, although— 

assuming the validity of the generic assignment—the 

lack of an inflated area at the base of the petiolule is 

consistent with the morphology of terminal leaflets of 

Fraxinus, which typically are conspicuously stalked 

and lack an inflated area. Lateral leaflets may also be 

stalked in Fraxinus but not as elongated as in the fossil.

The assignment to Fraxinus is based on characters 

of intercostal and marginal venation. In extant 

Fraxinus the intercostal areas are typically highly irregular in size (that is, the secondaries are very irregularly spaced), the tertiary and higher order veins 

are also inconsistent in spacing, and the areoles, 

which are formed by sixth order venation, are intruded by profusely branching veinlets. The secondary 

veins are consistently camptodromous but give off external branches that either bifurcate, sending one 

branch into the sinus or apically along the margin, or 

rarely enter the teeth centrally.

Fraxinus kenaica appears to be most closely related to the extant F. hopeinsis Tang from northern 

China. Particularly notable similarities are in the 

sharpness and large size (for Fraxinus) of the teeth. In 

this extant species, however, the secondary veins are 

considerably more apically directed, and there are 

fewer centrally entered teeth.

Holotype.—USNM 208490A, B.

Occurrence—9858.

Family Caprifoliaceae 

Genus Lonicera Linnaeus

Lonicera sp. 

Plate 25, figure 2

Discussion.—This single, specimen has an entire 

margin except for a single small tooth along the lower 

left-hand side. The secondary venation forms irregularly shaped intercostal areas that contain irregularly 

spaced branching tertiary veins. Abmedial to the secondary loops is a single series of loops that are also 

irregular in size and shape. Such features indicate a 

relationship to certain genera of Caprifoliaceae, but 

the Seldovia Point specimen is too incomplete and 

poorly preserved to make generic determination. In 

characters preserved, this specimen appears conspecific with undescribed leaves from the overlying 

Homerian assemblages; these younger leaves have the 

morphology of Lonicera.

Specimen.— USNM 208491.

Occurrence.—9858.

Family Alismataceae

Genus Alisma Linnaeus

Alisma seldoviana Wolfe et Tanai, sp. nov.

Plate 25, figures 3, 6

Description.—Leaves elliptic-lanceolate in shape,

7.2 to 15 cm (estimated) long and 2.2 to 5.5 cm wide; 

apex gradually narrowed, somewhat acuminate; base 

rounded to slightly cordate, sometimes slightly 

twisted; nine principal veins leaving the very base, 

acrodromous; mid vein stout and thick, nearly 

straight; two pairs of lateral primaries rather thick, 

but more slender than the midvein, making initial 

angles of about 20 to 50 degrees with the midvein, 

then gently arched, nearly parallel to the margin; 

outer two pairs of the lateral primaries thinner and 

more slender than the inner primaries, running along 

the margin; the tertiaries thin, obliquely transverse to 

the primaries with the angles of about 40 to 70 degrees, irregularly spaced; fourth order veins very fine, 

crossing with the tertiary veins, nearly parallel to 

the primaries, once or twice bifurcating to form elongate reticulation; margin entire; texture thin; petiole 

missing.

Discussion.—These fossils superficially resemble 

leaves of Smilax, but in that genus the fourth order 

venation is randomly reticulate. In Alisma, however, 

the fourth order veins are, as in the fossils, consistently perpendicular to the tertiary veins. Alisma seldoviana closely resembles the extant A. plantagoaquatica L., a species (or species complex) of wide 

distribution throughout the Northern Hemisphere.

Holotype.—USNM 208492.

Paratype.— USNM 208493.

Occurrence.—9858.

Family Najadaceae

Genus Potamogeton Linnaeus

Potamogeton alaskanus Wolfe et Tanai, sp. nov.

Plate 22, figures 2, 3; plate 25, figures 1, 7

Description.—Floating leaves variable in size and 

shape, 0.5 to 3.5 cm wide and 1.4 to 6.5 cm (estimated) 

long, elliptic to broadly elliptic in shape; base cuneate 

or rounded; apex incomplete but probably obtuse; 

petiole rather thick; midvein thick, nearly straight; 

lateral primary veins, three or four on both sides of 

midvein, acrodromous; thin and slender lateral subprimaries running nearly parallel to lateral primaries, one or two among each interlateral space; 

numerous secondary veins very thin, crossing somewhat irregularly to the primaries; margin entire. 

Submerged leaves narrowly linear, 2 or 3 mm wide, 

more than 5 cm long; base sessile; apex unknown; 

veins three, lateral veins inconspicuous.

Fruits obliquely suborbicular or obovoid, 1.5 to 2 cm 

long and 1.2 to 2 cm wide; the side flat; dorsal keel 

prominent, alate, undulate and with sutures; lateral 

keel low but evident, slightly detate; beak prominent, 

straight or slightly incurved, 0.8 mm long; apex of 

seeds pointing slightly above basal end; embryo 

coiled.
REFERENCES CITED 45

Discussion.—A number of leaves and fruits referred 

to Potamogeton occur at the Seldovia Point locality, 

and it is reasonably supposed that the plant-bearing 

rocks were mainly deposited in fresh, slowly moving 

water. Many leaves of Potamogeton have been reported from the Tertiary of the conterminous United 

States, but our Alaskan specimens are not identical 

with any species described before. P. alaskanus somewhat resembles P. hetrophylloides Berry from the 

Latah flora of eastern Washington (Brown, 1940) but 

differs in having the lateral subprimary veins. 

Dorofeyev (1963) described many species of Potamogeton on the basis of fruits from the Miocene of western 

Siberia, of which species P. alaskanus is most similar 

to P. tertiarius Dorof. in characters of the dorsal and 

lateral keels.

Holotype.—USNM. 208494.

Paratypes.— USNM 208495-208497.

Occurrence.—9856, 9858.

Mocotyledonae Incertaesedis

Genus Monocotylophyllum Chandler

Monocotylophyllum alaskanum (Heer) Wolfe et Tanai, comb. nov.

Plate 23, figure 2; plate 25, figure 8

Phragmites alaskana Heer, 1869, Kongl. Svenska 

Vet.-Akad. Handl., v. 8, no. 4, p. 24, pi. 1, fig. 12.

Discussion.—A number of fragmentary linear 

leaves with many parallel veins are similar to 

Phragmites alaskana, which are originally described 

from the "English Bay" locality. This generic reference may, however, not be valid, because the monocot 

leaves of Gramineae and other monocotyledonous 

families are too simply nerved to determine their 

generic status. Our specimens are from 1.2 to 4 cm in 

width, but their length is unknown.

Hypotypes.—USN'M. 208498, 208499.

Occurrence.— 9856, 9858.

Monocotylophyllum spp. 

Plate 25, figures 4, 5

Poacites tenuestriatus Heer, 1869, Kongl. Svenska 

Vet.-Akad. Handl., v. 8, no. 4, p. 24, 11. 1, fig. 12.

Discussion.—These species are represented by several fragmentary leaves which are linear with about 

17 to 30 mm width and contain numerous indistinct, 

parallel veins. The genus Poacites has been used for 

the grass leaves, and it is uncertain whether or not 

our specimens belong to the Gramineae. Poacites was 

originally proposed by Schlotheim for the Carboniferous grasslike specimens, but, as pointed out by Andrews (1970), the original specimens are not grass but 

are lycopod.

Specimens.— USNM 208500-208503.

Occurrence.— 9856, 9858.

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Tralau, Hans, 1968, Evolutionary trends in the genus Ginkgo: 

Lethaia, v. 1, p. 63-101.

Traverse, Alfred, 1955, Pollen analysis of the Brandon lignite of 

Vermont: U.S. Bur. Mines Rept. Inv. 5151, 107 p.

Trelease, William, 1924, The American oaks: Natl. Acad. Sci. Mem., 

v. 20, 255 p.

Triplehorn, D. M., Turner, D. L., and Naeser, C. W., 1977, K-Ar and 

fission-track dating of ash partings in coal beds from the Kenai 

Peninsula, Alaska: A revised age for the Homerian StageClamgulchian Stage boundary: Geol. Soc. America Bull., v. 88, 

p. 1156-1160.

Wahrhaftig, Clyde, Wolfe, J. A., Leopold, E. B., and Lanphere, 

M. A. 1969, The coal-bearing group in the Nenana coal field, 

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U.S. Geol. Survey Prof. Paper 398-B, 32 p.

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Geol. Survey open-file rept., 114 p.

———1969b, Neogene floristic and vegetational history of the 

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INDEX

[Italic page numbers indicate both major references and descriptions]

Page 

Abies ...... .....................................15

Abura flora ......................................20

Acer ........... .................... ..15, 16, 20,40

barghoonianum ............................ .42

bendirei .............................. .... .29

chaneyi ......................................8

circinnatum ................................. .18

cissifolium .................................. .41

crataegifolium ........................... .25, 41

ezoanum ... .8, 9, 10, 14, 16, 18, 25, 40; pis. 17, 19

fatisiaefolia ................................. .25

glabroides ........................ .10, 40; pi. 17

grahamensis .... .10, 16, 18, 25, 40; pis. 17, 18, 19

henryi ...................................18, 41

heterodentatum . .10, 18, 25, 41; pis. 16, 17, 18, 19 

japomcum .................................. .18

kushiroanum ............................. .9, 14

macrophyllum .................... .12, 15, 18, 40

macropterum ............................ .25, 40

merriami . ................................. .29

minor ......................... .............41

miyabei ............................. .14, 18, 40

mono .................................13, 18, 40

negundo .................................18, 41

oregonianum ................................ .41

pennsylvanicum ......................... .24, 40

riparius ......................... ...........42

rubrum .............................. .13, 15, 40

rufinerve .....................................40

saccharinum ...................... .13, 15, 24, 40

scottiae ............ ................. ... .18, 40

subpictum .................................. .25

sp ..... .....................................25

Aceraceae ......... ................10, 11, 20, 24, 40

acerifolius, Kalopanax .......... . . .11, 25, 42; pi. 22

aceroides, Platanus .............................. .29

acutilobum, Liqmdambar .......... ............. .28

aequalifolia, Marlea . ........................... .41

aequalifolium, Alangium ..................... .41, 42

aequifolia, Dombeyopsis .... ................. ... .42

Aesculus majus ................................. .24

Age .... .. .. ...................................7

Alangiaceae .... .............. .. .......10, 11, 41

Alangium .................... .. .....12,13,19,41

aequalifohum ............................ .41, 42

mikii ..... ... .8, 10, 15, 22, 24, 41; pis. 20, 21

riparius . . . ........ .............. ....... .42

tiliaefolium ............... ................. .42

Alaska Range ......... ................. .... .8, 28

alaskana, Decodon ...... .10, 16, 18, 25, 39; pis. 15, 16

Phragmites ...... .. . . .................... .25

Potamogeton ............. .11, 25, 44; pis. 22, 25

Tilia ...... ................. ... .......... .33

alaskanum, Artocarpidium ..... ................ .32

Ceridiphyllum . . .10, 15, 18, 24, 25, 27; pis. 2, 11 

Monocotylophyllum . ..... . ... .11,25; pis. 23, 25

Alchornea sp .............. ........... ......... .25

aleutica, Phyllodoce .. ............................9

aha, Vitas .. ........ ...... . ..... . . .... ....43

Ahsma ...... .. ...................... . . ..... .44

plantagoaquatica . ..... .................... .44

seldoviana ..................... .11, 25, 44; pi. 25

Alismataceae ........... .... .......... . 11,44

Alnus ............ .... .. ....... .14, 15, 18, 20,32

barnesi ............ .... ................... .24

cappsi ....... ..... .8, 10, 15, 16, 24, 25, 32; pi. 8

Alnus (continued) Page 

evidens ..................................... .32

fairi ...................... .10, 24, 25, 32; pi. 8, 9

healyensis .............. .8, 10, 24, 25, 32; pi. 8, 9

kluckingi ................................... .32

relatus ..................................... .32

sp .................................15, 16,; pi. 9

andersoni, Spiraea .............................. .25

Aniai flora ........................................8

aniensis, Cladrastis .................... .10, 38; pi. 15

antipofi, Fagus ... .8, 10, 15, 16, 20, 25, 31; pis. 5, 6, 7

Appcarya ....................................... .34

aquatica, Nyssa ................................. .41

Araliaceae ...................................11, 42

Arbutus .........................................22

matthesii ................................... .34

Arisaema hesperia ............................... .28

Artocarpidium alaskanum ....................... .32

aspera, Platanus .................................29

Aspidiaceae ...................................9, 25

asymmetrosa, Pterocarya ......................... .35

auriculata, Cocculus .... .8, 10, 15, 18, 25, 26, 27; pi. 2

Hedera ......................................26

B

Barabara Point ................................ .5, 6

barghoornianum, Acer ........................... .42

barnesi, Alnus .................................. .24

Beluga Formation .................................5

Beluga River ....................................40

bendirei, Acer ................................... .29

Carya .............. .10, 15, 17, 25, 33; pis. 10, 11

Platanus ........... .9, 10, 16, 18, 25, 28; pis. 3, 4

Rhus ....................................... .33

Berchemia ...................................... .19

Betula ...................................... .15, 32

fami ....................................... .32

sublutea ............................10,32; pi. 9

^or ......................................... .8

sp .........................................pi. 9

Betulaceae ..................10, 11, 14, 15, 20, 24, 32

bicolor, Prunus ...................................37

bifurcies, Phylhtes ............................... .41

biloba, Ginkgo ....................... .9, 12, 26; pi. 1

boreahs, Hemitrapa ......... ............. .10, 25, 39

TVapa ...................................... .39

bretzi, Quercus ......... ........................ .25

Bridge Creek flora ................................32

browni, Zelkoua . ................ .10, 30, 31; pis. 5, 6

Byttneriophyllum tiliaefolium .................... .42

C

Cache Creek .....................................15

californicum, Liquidambar .. ................ ... .28

Camellia protojaponica .......................... .20

camtschaticum. Rhododendron .....................9

Cape Blanco, Oreg ....................... .. ... .22

Capps Glacier ..... .3, 4, 7, 8, 24, 28, 32, 36, 38, 39, 42

cappsensis, Carpinus . . . ..... ................. .24

Sahx ........ .10, 16, 24, 25, 35, 36; pis. 12, 13, 14

cappsi, Alnus ........ .8, 10, 15, 16, 24, 25, 32; pi. 8

Crataegus ....... .............. ...........32

Caprifoliaceae ............. .............. .11, 15, 44

carpmifolia, Zelkova ... .............. .......... .30

Carpinus ................ .. ..... ..... .15,16,33

Carpinus ( continued) Page 

cappsensis .................................. .24

cordata ..................................... .33

seldoviana .............. .10, 16, 25, 33; pis. 9, 10

subcordata .................................. .33

Carya ................................ 12 , 16, 17, 33

bendirei ............ .10, 15, 17, 25, 33; pis. 10, 11

egregia ..................................... .34

miocathayensis .............................. .34

sessilis ..................................... .25

simulata .................................... .34

sp. .....................................8, 12, 25

Castanea .................................... .20, 31

ungeri .......................................31

Cebatha hetermorpha ............................ .26

multiformis ................................. .26

Cedrela ......................................... .20

Cedrus ..........................................20

Celtis ....................................... .10,29

kansana .....................................29

sp .................................... 29; pi. 4

Cercidiphyllaceae ............................ .10, 27

Cercidiphyllum ........................... .11, 19, 27

alaskanum .. . .10, 15, 18, 24, 25, 27, 28; pis. 2, 11

crenta ...................................... .27

crenatum ............................ .25, 27, 28

eojaponicum ............................. .27, 28

japonicum .................................. .27

Chamaecyparis sp ............................... .24

chamisonii, Crataegus .............. .10, 25, 37; pi. 14

Quercus .................................... .37

chaneyi, Acer .....................................8

chinensis, Corylus ................................33

chuitensis, Salix .............................. . . 36

Chuitna River ............................... 5, 7

ciliata, Populus ................................. 25

circinnatum, Acer ............................... .18

Cissampelos dubiosa ............................. .26

cissifolium, Acer ................................ .41

Cladrastis ................................... .12, 38

aniensis .......................... .10, 38; pi. 15

japonica .................................... .38

/utea ................................. .8, 24, 38

platycarpa .................................. .38

rafinesque .................................. 38

sp .................................. .pis. 15, 16

Clam Gulch .......................................5

Coal Cove ..................................... .5, 6

Cocculus ...................... ...... .12, 19, 24,26

auriculata ......... .8, 10, 15, 18, 25, 26, 27; pi. 2

ezoensis ................................. . . .27

/Za6e//a .................................... .27

heteromorpha .............................. .27

Collawash flora .......................... .30, 32, 34

Communities ............... ................... .16

Comptonia naumannii ........................... .24

Cook Inlet ........................ .......1, 4, 8, 24

Copper Basin Nevada ............................11

cordata, Carpinus ............................... .33

Cordia tiliaefolia ................................ .42

Corylus .................................. .12, 15, 33

chinensis ................................ .. .33

rostrata ................ .. . ............... .33

sp ......... ........ ........ . . .10, 12,33; pi. 9

crataegifolium, Acer ................ ......... .25, 41

Rulac .......................................41

49
50 INDEX

Crataegus ................................... .12, 37

cappsi ...................................... .32

chamisonii .................... .10, 25, 37; pi. 14

douglasii ....................................37

sp .......................................24, 25

crenata, Cercidiphyllum ......................... .27

Fa#us ............................ .10, 25; pi. 6

Vitis .......................................35

crenatum, Cercidphyllum ................. .25, 27, 28

crispula, Quercus .................................32

Cunninghamia .................................. .13

Cyclocarya ........................ .12, 13, 16, 24, 34

ezoarca ..........................8, 10, 15, 25, 34

paliurus ..................................16, 34

(Cydoptera), Pterocarya .......................... .25

Cyperacites sp ....................................25

Cyttneria ........................................42

D

decipiens, Dryopteris ............................. .25

Decodon ........................................ 39

alaskana ........... .10, 16, 18, 25, 39; pis. 15, 16

verticillatus ..................................39

dentata, Hicoria ..................................33

dissecta, Platanus ............................... .29

distichum, Taxodium ............................ .25

Distribution .....................................75

Dombeyopsis aequifolia .......................... .42

grandifolia ................................. .42

douglasii, Crastaegus .............................37

Dryopteris ...................................... 25

decipiens ................................... .25

guyotti ......................................25

idahoensis .................................. .25

tokyoensis .................................. .25

sp ...................................9, 25; pi. 1

dubiosa, Cissampelos ............................ .26

Dui Flora ........................................20

E

e&oe, Nuphar ................................. .pi. 1

Nymphar .................... .8, 10, 25, 26; pi. 1

egregia, Carya .................................. .34

Juglans .....................................34

Eleagnus ....................................... .15

elliptica, Prunus ............................. .37, 38

engelhardti, Salix ................................33

English Bay ............................ ... .1, 6, 30

eojaponicum, Cercidiphyllum ................. .27, 28

eotremuloides, Populus ............................ .8

Equisetum sp ....................................25

Ericales ........................................ 15

Esutoru flora ...................................20

etymodrys, Quercus .............................. .31

Eucommia ...................................15, 29

montana ........................... .10, 29; pi. 4

ulmoides ................................... .29

Eucommiaceae ...............................10,29

europaeum, Liquidambar ........................ .28

patulum, Liquidambar ....................... .28

europaeus, Glyptostrobus ...........9, 24, 25, 26; pi. 1

evidens, Alnus ...................................32

Exbucklandia ....................................22

ezoana, Cyclocarya ........... .8, 10, 15, 25, 34; pi. 10

Pterocarya .................................. .34

ezoanum, Acer .. .8, 9, 10, 14, 16, 18, 25, 40; pis. 17, 19 

ezoensis, Cocculus ................ ............. .27

Fagaceae ......................... 10, 14, 22, 24, 37

Fagus ........................ .7, 8, 11, 15, 16, 20, 37

antipofi . ... ..8, 10, 15, 16, 20, 25, 37; pis. 5, 6, 7

crenata ......................... .10, 25, 37; pi. 6

macrophylla ................................ .31

palecrenata ................................. .25

sp ............ ....................... .25; pi. 6

Page 

/aim, Atous ...................10, 24, 25, 32; pi. 8, 9

Betula ...................................... .32

Populus .................................... .26

fatisiaefolia, Acer ................................ .25

Fish Creek flora ..................................22

flabella, Cocculus ................................ .27

Floristic composition ..............................9

Floristics ........................................76

formosana, Liquidambar ......................... .12

fraxinifolia, Pterocarya .......................... .35

Fraxinus ....................................... .43

hopeinsis ................................... .44

kenaica ...................... .11, 25, 43; pi. 24

seldoviana .................................. .44

sp .......................................... .25

furuhjelmi, Quercus ..... .8, 10, 15, 16, 24, 25; pi. 7, 8

fusca, Malus ......................................9

G

gale, Myrica ..................................... .9

garryana, Quercus ............................... .13

Geologic occurrence .............................. 3

Ginkgo ...................................... .12,26

biloba .........................9, 12, 25, 26; pi. 1

Ginkgoaceae .................................. .9,26

glabroides, Acer ....................... .10, 40; pi. 17

Gleditschia ......................................20

glyptostroboides, Metasequoia . . .9, 12, 15, 24, 25; pi. 1 

Glyptostrobus ............................ .12, 25,26

eurpaeus ......................9, 24, 25,26; pi. 1

oregonensis ................................. .26

pensilis ..................................... .12

grahamense, Acer .................... .pis. 17, 18, 19

grahamensis, Acer ................. .10, 16, 18, 25, 40

Grand Coulee flora ...............................22

grandifolia, Dombeyopsis ........................ .42

grosseserrata, Quercus mongolica ................. .32

guyotti, Dryopteris ............................... .25

H

Hamamelidaceae ............................ .10, 28

Harriet Creek .................................... .5

Healy Creek Formation ........................ .7, 8

healyensis, Alnus .......... .8, 10, 24, 25, 32; pis. 8, 9

Hedera aunculata ............................... .26

heeriana, Vitis .................................. .35

Hemitrapa ...................................... 39

borealis ...... ....................... .10, 25, 39

Hemlock Conglomerate ........................... .4

henryi, Acer ................................. .18, 41

hesperia, Arisaema .............................. .28

Nyssa .......................................41

Onoclea .....................................25

Heterobalanus .................................. .32

heterodentatum, Acer .10, 18, 25, 47; pis. 16, 17, 18, 19 

heteromorpha. Cebatha ................... ...... .26

Cocculus ..... ...............................27

Populus .................................... 26

hetrophylloides, Potamogeton ..................... .45

Hicoria dentata .................... ............ .33

orientahs ....................................34

pecanoides .................................. .34

Honshu flora .................................26, 27

hopeinsis, Fraxinus .............................. .44

hopkinsi, Salix ............. .10, 16, 35, 36; pis. 12, 13

Sorbaria .................... ..... .10, 38; pi. 16

Spiraea ............... ..................... .38

horndum, Oplopanax ..............................9

Hydrangea ....... .... ......................... 37

sp .......... ............ ..10, 24, 25, 37: pi. 14

Hydrangeaceae .................. ..... ..... .10, 37

inquirenda, Salix .................................25

insignis, Ilex .................................... .37

Introduction ......................................7

iragawaense .................................... .42

Marlea ......................................41

Ishikari flora ....................................27

japomca, Cladrastis ..............................38

Nyssa .......................................38

japonicum, Acer ..................................18

Cerecidiphyllum ..............................27

Juglandaceae ..........................10, 14, 24,33

Juglans ........................................ .15

egregia ......................................34

nigella ......................................35

oregoniana .................................. .35

picroides .....................................36

K

Kachemak Bay ..............................2, 5, 38

Kalopanax ...................................... -42

acerifolium ..................................42

acerifohus ........................ .11, 25; pi. 22

n-suzukii ................................... .42

sp .......................................... .25

Kamchatka ..................................... -16

kansana, Celtis ...................................29

Kashihi flora ....................................20

Kenai Peninsula ................................. .1

kenaiana. Populus ..... .8, 10, 15, 24, 25, 35; pi. 11, 12

kenaica, Prunus ................... 10, 25, 37 pi. 15

kenaica, Fraxinus ................. .11, 25, 43; pi. 24

Keteleerm sp ................................... .24

Kirkwood Formation ............................42

kluckingi, Alnus .................................32

knowltoni, Nyssa ................... .10, 25, 47; pi. 18

Ulmus ......................... .10, 25, 30; pi. 5

Kodiak Island .....................................6

Kuiu Island .......................................1

Kukak Bay ......................................36

kushiroanum, Acer .............................9, 14

Larix ............................................15

Latah flora ...............................22, 41, 45

Lauraceae .......................................22

Leguminosae ................................... 38

Leucobalanus .... .............................. .32

leucophylla, Populus ............................. .31

Lindera ........................................ .19

hndgreni, Populus ............................... .35

hndleyana, Spiraea .............................. .37

Liquidambar ....... .7, 8, 12, 13, 14, 15, 17, 19, 24, 28

acutilobum ..................................28

californicum ................................ .28

europaeum .................................. .28

patulum ................................ .28

formosana .................................. .12

mwformosana ........................... .25, 28

pachyphylla .......... .10, 16, 17, 25, 28; pis. 2, 3

protensum .................................. .28

straciflua ................................ .12, 16

vachyphylla ................................. 28

Lonicera ................................. .12, 15, 44

sp ..... ........................ ...11, 44; pi. 25

longifilia, Ulmus .................................25

lutea, Cladrastis ........................... .8, 24, 38

lyrata, Quercus ................................. .13

Lythraceae ...................................10, 39

M
INDEX 51

Page 

macropterum, Acer ........................... .25, 40

Magnolia ........................................22

ingelfieldi ....................................29

magus, Aesculus .................................24

Mallotus sp ......................................25

Malus fusca .......................................9

Malvaceae .......................................15

Marlea aequalifolia ...............................41

iragawaense .................................41

matthesii, Arbutus ...............................34

Megafossils ......................................11

Menispermaceae ..............................10, 26

merriami, Acer ...................................29

Prunus .................................. .34, 38

Metasequoia .......................... .12, 18, 25, 26

glytostroboides ..........9, 12, 15, 24, 25, 26; pi. 1

occidentalis ..................................26

sp ...................................8, 15; pi. 1

mizrocrispula, Quercus .......................... .32

mikii, Alangium ......8, 10, 15, 22, 24, 41; pis. 20, 21

minor, Acer ......................................41

mwcathayensis, Carya ............................34

miocenica, Ptelea .................................34

mioformosana, Liquidambar .................. .25, 28

miothunbergiana, Pueraria ............. .10, 25; pi. 15

mixta, Pterocarya ............................ .25, 35

miyabei, Acer .............................14, 18, 40

mongohca, Quercus ............................... 13

grosseserrata, Quercus ....................... .32

mono, Acer ...............................13, 18, 40

Monocotylophyllum ............................. 25

alaskanum ................... .11, 25; pis. 23, 25

sp ........................... ......11, 25; pi. 25

montana, Eucommia ..................... 10, 29; pi. 4

Mormon Creek flora ..............................37

multiformis, Cebatha ............................ .26

Myrica gale .......................................9

sp ...........................................24

Myrtus oregonensis .............................. .39

N

Naihoro flora . ..................................20

Najadaceae ..................................... .44

naumanni, Comptonia ............................24

Vitas ............... ........................43

negundo, Acer ............................... .18, 41

Nenana coalfield ...................... .3, 7, 8, 27, 36

newberryi, Ulmus ............................... .25

nigella, Juglans ................................. .34

Pterocarya ......8, 10, 15, 18, 24, 25, 34, 35; pi. 11

nobihs, Platanus ................................ .29

North Pacific Basin ............................ .24

Noxapaga Formation .............................15

n-suzukii, Kalopanax .............................42

Nuphar ..................................... .16, 26

ebae ................................... . .pi. 1

sp ...........................................25

Nymphaea ...................................... .26

Nymphaeaceae .............................8, 10, 26

Nymphar .................. ............6, 11, 16, 26

ebae ............. ..................8, 10, 25, 26

Nyssa ............ ................... .12, 15, 38, 41

aquatica ...................... ............. .41

hesperia .................................... .41

japonica .....................................39

knowltoni ...... .............. .10, 25, 41; pi. 18

sylvatica ............................ ....12, 41

sp . ..... ....................... ... .25; pi. 18

Nyssaceae ................................. .10, 41

O

occidentalis, Metasequoia ....... ................ .26

Platanus ................... .................12

Odasu flora ......................................20

Oleaceae ...... ..............................11, 43

Onoclea ........................................ 25

hesperia .....................................25

senibilis ............................ .9, 25; pi 1

Page 

Oplopanax horridum ............................. .9

oregona, Tilia ................................... .37

oregonensis, Glyptostrobus ....................... .26

Myrtus ......................................39

oregoniana, Juglans ..........................35

Osirya ............................. .10, 33; pi. 9

Quercus .................................... .32

Zelkova ................................. .25, 30

oregonianum, Acer ...............................41

orientalis, Hicoria ............................... .34

Ostrya .......................................15,33

oregoniana ..........................10, 33; pi. 9

sp .........................................pi. 9

owyheensis, Ulmus ................ .10, 25, 30; pi. 4, 6

pachyphylla, Liquidambar .. .10, 16, 17, 25, 28; pi. 2, 3 

padus, Prunus ............................... .10, 3$

Paleoclimatology ................................ J9

paleocrenata, Fagus ..............................25

paliurus, Cyclocarya ......................... .16, 34

Pterocarya ...................................34

pandurata, Quercus ..............................31

patulum, Liquidambar europaeum ................ .28

paucidentata, Platanus .......................... .29

Wmus .......................................30

pecanoides, Hicoria .............................. .34

pennsylvanicum, Acer ........................ .24, 40

pensilis, Glyptostrobus ............................12

Phragmites alaskana .............................25

Phyllodoce aleutica ................................9

Picea .....................................14, 15, 20

picroides, Juglans ............................... .36

Safe ............10, 16, 24, 25, 35, 36; pis. 12, 13

Pinaceae ..............................14, 15, 16, 20

Pinus ....................................... .15, 20

Pistacia .........................................20

Planera ungeri ...................................30

plantagoaquatica, Alisma ........................ .44

Platanaceae ..................................10, 28

Platanus ...................... .12, 15, 18, 19, 24, 28

aceroides ................................... .29

aspera ...................................... .29

bendirei .............9, 10, 16, 18, 25, 28; pis. 3, 4

dissecta ..................................... .29

nobilis ...................................... .29

occidentalis ..................................12

paucidentata .................................29

racemose .............................17, 18, 29

raynoldsu .................................. .29

youngii ..................................... .29

platycarpa, Cladrastis ........................... .38

Platyptera .......................................35

plurinervis, Ulmus ...............................30

Poacites .........................................25

tenuistriatus ................................ .25

Point Pogibshi ....................................6

Pollen .......................................... 74

Populus ............................15, 16, 20, 27,35

ciliata .......................................25

eotremuloides .................................8

/aim ........................................26

heteromorpha ................................26

kenaiana ......... .8, 10, 15, 24, 25, 35; pi. 11, 12

leucophylla ................................. .31

lindgrem ................................... .35

reniformis ............... ................. .25

trichocarpa ................................. .9

sp .......................10, 15, 24, 25, 35; pi. 11

Port Graham .........................5, 7, 14, 16, 33

Potamogeton ...............................6, 16,44

alaskana ...................11, 25,44; pis. 22, 25

hetrophylloides .............................. .45

tertiarius ........................... ........45

sp ........... ...............................25

Potamogetonaceae ................................11

prinoides, Quercus ...................... ........32

protensum, Liquidambar ..... ....................28

Page 

Prunus ......................................... 37

bicolor .......................................37

elliptica ................................. .37, 38

kenaica ........................10, 25,37; pi. 15

merriami .................................34, 38

padus ....................................10,35

scotti ........................................37

vanioti ......................................37

sp ....................................25; pi. 14

pseudomonticola, Salix .......................... .36

Pseudotsuga .................................... .15

Pseudocastanea, Quercus ......................... .32

Ptelea miocenica ................................ .34

Pterocarya ................. .11, 15, 16, 19, 24, 34, 35

asymmetrosa ................................ .35

ezoana ...................................... .34

fraxinifolia ................................. .35

miifa ....................................25, 35

nigetta .........8, 10, 15, 18, 24, 25, 34,35; pi. 11

paliurus .................................... .34

pugetensis .................................. .18

rhoifolia ............................. .12, 18, 35

(Cycloptera) sp .............................. .25

sp .......................................... 35

Pueraria .....................................19, 39

miothunbergiana ............... .10, 25, 39; pi. 15

Puget Group .................. ..............18, 27

pugetensis, Pterocarya ........................... .18

Phyllites bifurcies ............................... .41

Q

Quercus ................................. .16, 3i, 37

6retei ....................................... .25

chamissoni ..................................37

crispula .....................................32

elliptica .....................................20

etymodrys ...................................31

furuhjelmi .......8, 10, 15, 16, 24, 25, 3i; pis. 7, 8

garryana ................................... .13

lyrata .......................................13

macrocarpa ................................. .13

microcrispula ............................... .32

mongolica .................................. .13

grosseserrata ............................ .32

oregoniana ...................................32

pandurata .................................. .31

prinoides ....................................32

pseudocastanea ...............................32

sadleriana ...................................32

R

racemosa, Platanus ...........................17, 18

raynoldsu, Platanus ..............................29

References cited ..................................45

relatus, Alnus ................................... .32

reniformis, Populus .............................. .25

Rhododendron camtschaticum ......................9

rhoifolia, Pterocarya ...................... .12, 18, 35

Rftus bendirei ....................................33

richardsonii, Salix ...............................36

riparius, Alangium .............................. .42

Rosaceae .................................10, 15, 37

rostrate, Corylus ................................ .33

rubrum, Acer ............................ .13, 15, 40

rufunerve, Acer ................................. .40

Rulac crataegifolium .............................41

S

saccharinum, Acer .................... .13, 15, 24, 40

sadleriana, Quercus ............................. .32

St. Eugene flora .... .............................22

Salicaceae .............................10, 20, 24,35

Safe ..............................15, 16, 18, 35, 39

cappsensis .. 10, 16, 24, 25, 35, 36; pi. 12, 13, 14 

chuitensis ................................... .36

engelhardti ................................. .33
52 INDEX

Salix {continued} Page 

hopkinsi ............... .10, 16, 35, 36; pis. 12, 13

inqmrenda .............................. .25, 35

picroides ........ .10, 16, 24, 25, 35,36; pis. 12, 13

pseudomonticola .............................36

richardsonii ................................ .36

seldoviana ............. .10, 16, 35, 36; pis. 12, 14

various .................................. .35, 36

sp ..............................8, 15, 16, 24, 25

Sanctuary Formation ........................7, 8, 15

Schisandra ..................................... .19

scotti, Prunus ................................... .37

scottiae, Acer ................................ .18, 40

seldoviana, Ahsma ................. .11, 25, 44; pi. 25

Carpinus ............... .10, 16, 25, 32; pis. 9, 10

Fraxinus ................................... .44

Salix .................. .10, 16, 35, 36; pis. 12, 14

Vitis ......................... .11, 25; pis. 22, 23

sensibilis, Onoclea ....................... .9, 26; pi. 1

serrata, Zelkova ................................. .31

sessilis, Carya ................................... .25

Seward Peninsula ................................ 15

simulata. Carya ................................. .34

sinica, Zelkova ...................................31

Skull Springs, Oreg ..............................18

Smilax ..........................................44

sp ...........................................25

Sorbaria .....................................12,38

hopkinsi .......................... .10, 38; pi. 16

sp .......................................... .24

speciosa, Ulmus ........................ .10, 30; pi. 4

Spiraea andersom ................................25

hopkinsi .................................... .38

lindleyana .................................. .38

weaveri ..................................... .24

Sterling Formation ................................5

styraciflua, Liquidambar ..................... .12, 16

subcordata, Carpinus .............................33

sublutea, Betula ........................ .10, 32; pi. 9

subnobilis, Tilia ....................... .10, 25; pi. 15

subpictum, Acer ................................. .25

Sucker Creek flora .............................. .30

Suntrana Formation .....................7, 8, 15, 28

sylvatica, Nyssa .............................. .12, 41

Page 

Symphoricarpos ................................. .16

sp ...........................................25

Systematics ..................................... 24

Taxodiaceae ...................................9,26

Taxodium distichum ............................ .25

tenuistriatus, Poacites ........................... .25

tertiarius, Potamogeton .......................... .45

thor, Betula .......................................8

Ti/ia ....................................7, 8, 15,36

alaskana ....................................33

oregona ..................................... .37

subnobilis ..................... .10, 25, 36; pi. 15

sp ...........................................25

Tiliaceae .................................10, 11, 36

tiliaefolia, Cordia ............................... .42

tiliaefolium, Alangium ............................42

Byttneriophyllum ............................ .42

tokyoensis, Dryopteris ............................ .25

Trapa .......................................... .39

borealis ......................................39

Trapaceae ....................................10, 11

Trapellaceae .................................... 39

trichocarpa, Populus .............................. .9

Tsuga ....................................14, 15, 20

Tyonek Formation ..............................4, 5

U

Ulmaceae ..........................10, 11, 20, 24, 29

ulmoides, Eucommia ............................ .29

Ulmus .....................7, 8, 10, 15, 16, 20, 24, 30

knowltoni ...................... .10, 25, 30; pi. 5

longifolia ................................... .25

newberryi ................................... .25

owyheensis ................... .10, 25, 30; pi. 4, 6

paucidentata ................................ .30

plurinervis .................................. .30

speciosa ............................. 10, 30; pi. 4

sp ........................ .8, 10, 15, 20, 24; pi. 5

ungeri, Castanea ................................ .31

ungeri, Castanea (continued> Page 

Planera ......................................30

Ze/fewa ...................10, 25, 30, 3^; pis. 5, 6

Upper Healy Creek ...............................32

Utto flora ........................................22

V

vanioti, Prunus ................................. .37

varians, Salix ............................... .35, 36

Vegetation ...................................... .11

verticillatus, Decodon ............................ .39

Vitidaceae ...................................11, 43

Vitis .........................................12,43

alia .........................................43

crenata ..................................... .35

heeriana .....................................35

naumanni .................................. .43

seldoviana ................. .11, 25, 43; pi. 22, 23

sp ...........................................25

W

weaveri, Spiraea ................................ .24

West Foreland Formation .......................3, 4

Wishkaw River, Wash ........................... .22

Wrangell Mountains .............................15

Yakataga District .................................6

Yoshioka flora ...................................20

youngii, Platanus ............................... .29

Yukon River ......................................1

Zelkova .................................. .16, 24, 30

browni ....................10, 25,30, 31; pi. 5, 6

carpinifolia ................................. .30

oregomana .................................. .25, 30

serrata ..................................... .31

sinica ...................................... .31

ungeri ................... .10, 25, 30, 31; pis. 5, 6

sp .......................................... .16
PLATES 1-25

[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

[All figures natural size unless otherwise stated]

FIGURE 1. Ginkgo biloba Linnaeus, (p. 26 ).

Hypotype, USNM 208356; loc. 9857.

2, 6, 7, 10. Metasequoia sp. cf. M. glyptostroboides Hu et Cheng. (p. 26 ). 

2, 10. USNM 208352, 208355; loc. 9858. 

6, 7. USNM 208353, 208354; loc. 9857. 

3, 4. Onoclea sensibilis Linnaeus, (p. 25 ).

3, 4. Hypotypes USNM 208349, 208350; loc. 9858. 

5, 9. Dryopteris sp. (p. 25 ).

USNM 208348A, B; loc. 9858. 

8. Glyptostrobus europaeus (Brongniart) Heer. (p. 26).

x2. Hypotype, USNM 208351; loc. 9858. 

11-14. Nymphar ebae (Huzioka) Ozaki (p. 26).

Hypotypes, USNM 208357-208360; loc. 9858. Fig. 13, x2.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 1

13

14

FILICALES, CONIFERALES, MAGNOLHDAE
PLATE 2

[All figures natural size unless otherwise stated]

FIGURES 1-4. Cercidiphyllum alaskanum Wolfe et Tanai. (p. 27 ).

1. 4. Holotype, USNM 208361A, B; loc. 9856.

2. Paratype, USNM 208362; loc. 9858.

3. Showing fine veinlets (enlargement of holotype specimen). 

5, 6. Liquidambar pachyphylla Knowlton. (p. 28 ).

5, 6. Hypotypes USNM 208364, 208365; loc. 9858. 

7. Cocculus auriculata (Heer) Wolfe. (p. 26). 

Hypotype USNM 208369; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 2

HAMAMELIDIDAE, RANUNCULIDAE
PLATE 3

[All natural size|

FIGURE 1-3. Liquidambar pachyphylla Knowlton. (p. 28 ).

Hypotypes USNM 208366-208368; loc. 9858. 

4. Platanus bendirei (Lesquereux) Wolfe. (p. 28 ). 

Hypotype USNM 208370; loc. 9858.
GEOLOGICAE SURVEY PROFESSIONAL PAPER 1105 PLATE 3

HAMAMELIDIDAE
PLATE 4

[All natural size]

FIGURES 1, 2, 4. Platanus bendirei (Lesquereux) Wolfe. (p. 28 ). 

Hypotypes USNM 208371-208373; loc. 9858. 

3, 5. Ulmus speciosa Newberry. (p. 30 ).

Hypotypes USNM 208381, 208382; loc. 9858.

6. Eucommia cf. E. montana R. W. Brown, (p. 29 ). 

USNM 208374; loc. 9858.

7. Celtis sp. (p. 29 ).

USNM 208375; loc. 9858.

8. Ulmus owyheensis H. V. Smith, (p. 30 ). 

Hypotype USNM 208376; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 4

HAMAMELIDIDAE
PLATE 5

[All figures natural size unless otherwise stated]

FIGURES 1, 2. Ulmus knowltoni Tanai et Wolfe. (p. 30 ).

Hypotypes USNM 208376, 208377; loc. 9856. 

3, 6, 8a, 8c. Zelkova browni Tanai et Wolfe. (p. 30 ).

3, 6. Hypotypes USNM 208384-208385; loc. 9858. 

8a, 8c. Hypotypes USNM 208386-208388; loc. 9856. 

4, 7. Zelkova ungeri Kovats. (p. 31 ).

Hypotypes USNM 208389, 208390; loc. 9858. 

5. Ulmus sp.

USNM 208383; loc. 9856. 

8b. Fagus antipofi Heer. (p. 31 ).

Hypotype, USNM 208394; loc. 9856.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATES

HAMAMELIDIDAE
PLATE 6

[All figures natural sizel

FIGURES 1, 2, 6. Zelkova ungeri Kovats. (p. 31 ).

Hypotypes, USNM 208391, 208392, 208393; loc. 9858. 

3, 4. Fagus antipofi Heer. (p. 31 ).

Hypotypes USNM 208395, 208396; loc. 9858. 

5. Fagus sp. aff. F. crenata Blume. (p. 31 ).

Hypotype USNM 208400; loc. 9856. 

7a, 7b. Ulmus owyheensis H. V. Smith, (p. 30 I.

Hypotypes USNM 208377A, 208377B; loc. 9856. 

8. Zelkova browni Tanai et Wolfe. (p. 30 ). 

Hypotype USNM 208388; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 6

HAMAMELIDIDAE
PLATE 7

(All figures natural size)

FIGURES 1, 5, 6. Fagus antipofi Heer. (p. 31 ).

Hypotypes USNM 208397, 208398, 208399; loc. 9858. 

2-4. Quercus furuhjelmi Heer. (p. 31 ).

Hypotypes USNM 208401, 208402, 208403 loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 7

\ ># 19

HAMAMELIDIDAE
PLATE 8

[All figures natural size]

Figures 1-3. Quercus furuhjelmi Heer. (p. 31 ).

Hypotypes USNM 208404, 208405, 208406; loc. 9858.

4. Alnus fairi (Knowlton) Wolfe. (p. 32 ). 

Hypotype USNM 208408; loc. 9858.

5. Alnus cappsi (Hollick) Wolfe. (p. 32 ). 

Hypotype USNM 208407; loc. 9858.

6. Alnus healyensis Wolfe. (p. 32 ).

Hypotype USNM 208410; loc. 9858.
GEOLOGICAL SURVEY

*

PROFESSIONAL PAPER 1105 PLATE 8

•»

HAMAMELIDIDAE
PLATE 9

[All figures natural size unless otherwise stated]

FIGURE I. Alnus healyensis Wolfe. (p. 32 ).

Hypotype USNM 208411; loc. 9858.

2. Alnus fairi (Knowlton) Wolfe. (p. 32 ;. 

Hypotype USNM 208409; loc. 9858.

3. Corylus sp. (p. 33 ).

USNM 208419; loc. 9858.

4-6. Ostrya sp. cf. O. oregoniana Chaney. (p. 33 )• 

USNM 208420-208422; loc. 9858. x2.

7. Carpinus seldoviana Wolfe. (p. 33 ). 

Hypotype USNM 208414; loc. 9858.

8. Betula sp. cf. B. sublutea Tanai et N. Suzuki, (p. 32). USNM 208413; loc. 9858.

9. Alnus sp.

USNM 208412; loc. 9858.
PROFESSIONAL PAPER 1105 PL ATE 9

•A.

HAMAMELIDIDAE
PLATE 10

[All figures natural size unless otherwise stated]

FIGURE 1-3. Carpinus seldoviana Wolfe. (p. 33).

Hypotypes USNM 208415, 208416, 208417; loc. 9858.

(fig. 3, x2). 

4-6, 8, 9a, 9b. Cyclocarya ezoana (Tanai et N. Suzuki) Wolfe et Tanai. (p. 34 ).

4, 5, 8, 9a, 9b. Hypotypes USNM 208425, 208426, 208427, 208428, 208429; 

loc. 9858.

(fig. 5, x2; fig . 9, x3). 

6. Hypotype USNM 208426; loc. 9856.

7. Carya bendirei (Lesquereux) Chaney and Axelrod. (p. 33 ). 

Hypotype USNM 208424; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 10

HAMAMELIDIDAE
PLATE 11

[All figures natural size unless otherwise stated]

FIGURES 1, 2. Carya bendirei (Lesquereux) Chaney et Axelrod. (p. 33 ).

1, 2. Hypotype USNM 208423; loc. 9856 (fig. 2, x3). 

3, 5. Cercidiphyllum alaskanum Wolfe et Tanai. (p. 27 ).

Paratypes USNM 208363A, B; loc. 9858. 

4, 6, 7. Pterocarya nigella (Heer) Wolfe. (p. 35 ).

4, 6. Hypotypes USNM 208430, 208431; loc. 9858. 

7. Hypotype USNM 208432; loc. 9856. x2.

8. Populus sp. (p. 35 )-

USNM 208434; loc. 9858.

9. Populus kenaiana Wolfe. (p. 35 ).

Hypotype USNM 208433; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 11

HAMAMELIDIDAE, DILLENHDAE
PLATE 12

[All figures natural size unless otherwise stated]

FIGURES 1. Populus kenaiana Wolfe. (p. 35 ).

Hypotype USNM 42264B; loc. 9858. 

2, 3. Salix seldoviana Wolfe et Tanai. (p. 36 ).

2, 3. Holotype USNM 208446; loc. 9858. Fig. 2, x3. 

4. Salix hopkinsi Wolfe et Tanai. (p. 36 ).

Holotype USNM 208440; loc. 9858. 

5-7. Salix picroides (Heer) Wolfe. (p. 36 ).

5. 7. Hypotypes USNM 208442, 208443; loc. 9858.

6. Showing fine venation and margin of USNM 208442 (fig. 5). Ca. x3. 

8. Salix cappsensis Wolfe. (p. 36 ).

Hypotype USNM 208435; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 12

6 7 

DILLENIIDAE
PLATE 13

[All figures natural size unless otherwise stated]

FIGURES 1, 2, 4. Salix cappsensis Wolfe. (p. 36 ).

1, 2, 4. Hypotypes USNM 208436, 208437, 208438; 

loc. 9858. Fig. 2, x3. 

3, 7. Salix picroides (Heer) Wolfe. (p. 36 ).

Hypotypes USNM 208444, 208445; loc. 9858. 

5, 6. Salix hopkinsi Wolfe et Tanai. (p. 36 ).

5, 6. Paratypes USNM 208441A, 208441B; loc. 9858. 

Fig. 6, x3.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 13

DILLENIIDAE
PLATE 14

[All figures natural size]

FIGURES 1. Hydrangea sp. (p. 37 ).

USNM 208449A; loc. 9858. 

2. Salix seldoviana Wolfe et Tanai. (p. 36 ).

Paratype USNM 208447; loc. 9858. 

3, 4. Salix cappsensis Wolfe. (p. 36 ).

Hypotypes USNM 208439A, 208439B; loc. 9858. 

5, 6. Crataegus chamissoni (Heer) Wolfe et Tanai. (p. 37 ).

Hypotypes USNM 208450, 208451; loc. 9858. 

7. Primus sp. (p. 38 ).

USNM 208454A, B; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 14

DILLENHDAE, ROSIDAE
PLATE 15

[All figures natural size unless otherwise stated]

FIGURES 1. Tilia subnobilis Huzioka. (p. 36 ).

Hypotype USNM 208448; loc. 9856. 

2, 5. Prunus kenaica Wolfe et Tanai. (p. 37 ). 

2. Holotype USNM 208452; loc. 9856. 

5. Paratype USNM 208453; loc. 9856.

3. Pueraria miothunbergiana Hu et Chaney. (p. 39 ). 

Hypotype USNM 208461; loc. 9858.

4. Cladrastis sp. cf. C. aniensis Huzioka. (p. 38 ).

Hypotype USNM 208459; loc. 9858. x2. 

6. Decodon alaskana Wolfe et Tanai. (p. 39 ).

Paratype USNM 208464; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 15

DILLENIIDAE, ROSIDAE
PLATE 16

[All figures natural size unless otherwise stated]

FIGURES 1, 2a, b, c. Sorbaria hopkinsi (Wolfe) Wolfe et Tanai. (p. 38 ).

Hypotypes USNM 208455-208458; loc. 9858. 

3, 4. Cladrastis sp. cf. C. aniensis Huzioka. (p. 38 ).

3, 4. USNM 208460; loc. 9858. Fig. 4, x3. 

Decodon alaskana Wolfe et Tanai. (p. 39 ). 

5. Paratype USNM 208463; loc. 9858. 

7, 8. Holotype USNM 208462; loc. 9858. Fig. 8, x3. 

6. Acer heterodentatum (Chaney) MacGinitie. (p. 41 ). 

Hypotype USNM 208476; loc. 9858.
PROFESSIONAL PAPER 1105 PLATE 16

, .

ROSIDAE
PLATE 17

[All figures natural size]

FIGURES 1-3, 5. Acer ezoanum Oishi et Huzioka. (p. 40 ).

Hypotypes USNM 208466, 208467, 208468; loc. 9858. 

4. Acer heterodentatum (Chaney) MacGinitie. (.p. 41 ).

Hypotype USNM 208477; loc. 9858. 

6. Acer grahamensis Knowlton et Cockerell. (.p. 40).

Hypotype USNM 208473; loc. 9858. 

7, 8. Acer glabroides Brown, (p. 40 ).

Hypotypes USNM 208471, 208472; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 17

ROSIDAE
PLATE 18

[All figures natural size]

FIGURE 1. Nyssa sp. cf. N. knowltoni Berry, (p. 41).

USNM 208482; loc. 9858. 

2-4. Acer heterodentatum (Chaney) MacGinitie. (p. 41 ).

Hypotypes USNM 208478, 208479, 208480; loc. 9858. 

5. Acer grahamensis Knowlton et Cockerell. (p. 40). 

Hypotype USNM 208474; loc. 9856.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 18

ROSIDAE
PLATE 19

[All figures natural size]

FIGURES 1, 4. Acer grahamensis Knowlton et Cockerell. (p. 40).

Hypotype USNM 208475; loc. 9856. 

2, 5. Acer ezoanum Oishi et Huzioka. (p. 40 ).

Hypotypes USNM 208469, 208470; loc. 9858. 

3. Acer heterodentatum (Chaney) McGinitie. (p. 41 ). 

Hypotype USNM 208481; loc. 9858.
GEOLOGICAL SURVEY 

T,

PROFESSIONAL PAPER 1105 PLATE 19

ROSIDAE
PLATE 20

FIGURE 1, 2. Alangium mikii Wolfe et Tanai. (p. 41).

1. Holotype, USNM 208483; loc. 9858. xl.

2. Paratype USNM 208485; loc. 11091 (Capps Glacier), ca. x.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 20

ROSIDAE
PLATE 21

[Natural size]

Alangium mikii Wolfe et Tanai. (p. 41 ). 

Paratype USNM 208484; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 21

ROSIDAE
PLATE 22

[All figures natural size]

FIGURES 1. Vitis seldoviana Wolfe et Tanai. (p. 43 ).

Paratype USNM 208488; loc. 9856. 

2, 3. Potamogeton alaskanus Wolfe et Tanai. (p. 44).

2. Paratype USNM 208495; loc. 9858

3. Holotype USNM 208494; loc. 9858. 

4. Kalopanax n-suzikii Wolfe et Tanai (p. 42). 

Hypotype USNM 208486; loc. 9856.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 22

ROSIDAE, MONOCOTYLEDONES
PLATE 23

[All figures natural size]

FIGURES 1, 3. Vitis seldoviana Wolfe et Tanai. (p. 43 ). 

1. Paratype USNM 208489; loc. 9856. 

3. Holotype USNM 208487; loc. 9856.

2. Monocotylophyllum alaskanum (Heer) Wolfe et Tanai. (p. 45 ). 

Hypotype USNM 208498; loc. 9858.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 23

ROSLDAE, MONOCOTYLEDONES
PLATE 24

[All figures natural size unless otherwise stated]

FIGURES 1-3. Fraxinus kenaica Wolfe et Tanai. (p. 43 ).

Holotype USNM 208490 A, B; loc. 9858. Fig. 3, x3.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 1105 PLATE 24

ROSIDAE
Plate 25

[All figures natural size unless otherwise stated]

FIGURES 1, 7. Potamogeton alaskanus Wolfe et Tanai. (p. 44). 

1. Paratype USNM 208496; loc. 9858. 

7. Paratype USNM 208497; loc. 9858. x5. 

2. Lonicera sp. (p. 44).

USNM 208491; loc. 9858.

3, 6. Alisma seldoviana Wolfe et Tanai. (.p. 44 ). 

3. Paratype USNM 208493; loc. 9858. 

6. Holotype USNM 208492; loc. 9858.

4. Monocotylophyllum sp. a. (p. 45 ). 

USNM 208500, loc. 9858.

5. Monocotylophyllum sp. a. (p. 45 ).

USNM 208503, loc. 9858. 

8. Monocotylophyllum alaskanum (Heer) Wolfe et Tanai. (p. 45 ).

Hypotype USNM 208499; loc. 9858.
PROFESSIONAL PAPER 1105 PLATE 25

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ASTERIDAE, MONOCOTYLEDONES