Paleogene Floras From the 

Gulf of Alaska Region

By JACK A. WOLFE

GEOLOGICAL SURVEY PROFESSIONAL PAPER 997

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

CECIL D. ANDRUS, Secretary

GEOLOGICAL SURVEY 

V. E. McKelvey, Director

Library of Congress Cataloging in Publication Data

Wolfe,JackA. 1936-

Paleogene floras from the Gulf of Alaska region.

(Geological Survey Professional Paper 997)

Bibliography: p. 94-97

Includes index.

1. Paleobotany-Eocene. 2. Paleobotany-Oligocene. 3. Paleobotany--Alaska--Gulf of Alaska region.

I. Title. II. Series: United States. Geological Survey. Professional Paper 997.

QE926.W64______________560*178_____________________7644847

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

Washington, D.C. 20402 (paper cover)

Stock Number 024-001-03027-6
CONTENTS

Page

Abstract ...................................................................................................... 1

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

Physical stratigraphy ............................................................................ 3

Katalla district .................................................................................. 3

Stillwater and Kushtaka Formations .............................. 3

Katalla Formation .................................................................. 6

Yakataga and Malaspina districts ............................................ 6

Kulthieth Formation .............................................................. 6

Poul Creek Formation ............................................................ 7

Yakataga Formation ........................................................... 7

Unknown unit .......................................................................... 7

Biostratigraphy ........................................................................................ 7

Introduction ...................................................................................... 7

Ravenian Stage ................................................................................ 8

Lower Ravenian ...................................................................... 8

Middle Ravenian ...................................................................... 9

Upper Ravenian .................................................................... 12

Kummerian Stage .......................................................................... 12

Post-Kummerian rocks ................................................................ 13

Angoonian Stage ............................................................................ 13

Definition .................................................................................. 14

Flora .......................................................................................... 14

Age .............................................................................................. 14

Correlation of some other Alaskan Floras ............................ 16

Floristic and vegetational analysis ................................................ 20

"Floristic analysis ............................................................................ 20

Vegetational analysis .................................................................. 21

Introduction ............................................................................ 21

Foliar physiognomy and vegetational analysis ........ 23

Relation of leaf margins to climate ........................ 24

Tropical rain forest ........................................................ 24

Paratropical rain forest .............................................. 25

Subtropical forest .......................................................... 26

Warm temperate forest ................................................ 26

Lower Ravenian ............................................................................ 26

Floristic analysis .................................................................... 26

Vegetational analysis .......................................................... 28

Middle Ravenian ............................................................................ 31

Floristic analysis .................................................................... 31

Floristic and vegetational analysis Continued 

Middle Ravenian Continued

Vegetational analysis .......................................................... 32

Upper Ravenian ............................................................................ 32

Floristic analysis .................................................................... 32

Vegetational analysis .......................................................... 33

Kummerian Stage ......................................................................... 34

Floristic analysis .................................................................... 34

Vegetational analysis .......................................................... 35

Climatic summary ........................................................................ 35

Alaskan Paleogene climates .............................................................. 36

History of the Arcto-Tertiary concept ............................................ 37

A theory of the origins of the Mixed Mesophytic

forest ...................................................................................................... 43

Paleogene tropical migration routes .............................................. 49

Systematics .............................................................................................. 51

Filicales ............................................................................................ 52

Coniferales ........................................................................................ 53

Principes ............................................................................................ 53

Salicales ............................................................................................ 54

Juglandales ...................................................................................... 55

Fagales .............................................................................................. 57

Urticales ............................................................................................ 60

Ranales .............................................................................................. 61

Resales .............................................................................................. 70

Geraniales ........................................................................................ 72

Sapindales ........................................................................................ 74

Rhamnales ........................................................................................ 79

Malvales ............................................................................................ 80

Parietales .......................................................................................... 81

Myrtiflorae ........................................................................................ 83

Umbelliflorae .................................................................................. 85

Ericales .............................................................................................. 85

Tubiflorae .......................................................................................... 85

Incertae sedis .................................................................................. 86

Systematic revisions of the Paleogene species ............................ 87

Locality data ............................................................................................ 93

References ................................................................................................ 94

Index .......................................................................................................... 99

ILLUSTRATIONS

[Plates follow index]

PLATE 1. The Kushtaka Formation on Carbon Mountain.

2-19. Lower Ravenian flora. 

20-25. Middle Ravenian flora. 

26, 27. Upper Ravenian flora. 

28-30. Kummerian flora.

Ill
IV CONTENTS

Page

FIGURE 1. Map of the Gulf of Alaska region ................................................................................................................................................................ 3

2. Generalized geologic map of part of the Katalla district .................................................................................................................... 4

3. Diagram showing stratigraphic position of fossil plant localities in the Katalla district...................................................... 5

4. Generalized geologic map of part of Kootznahoo Inlet........................................................................................................................ 15

5. Generalized stratigraphic section of the type section of the Angoonian Stage .......................................................................... 16

6. Map of Alaska showing location of some fossil plant assemblages.. ........................................................................................... 17

7. Correlation chart of Paleogene formations in the Katalla and Yakataga districts .................................................................. 21

8. Correlation chart of some Alaskan Tertiary formations .................................................................................................................... 22

9-43. Drawings:

9. Marginal venation of Platycarya.................................................................................................................................................. 56

10. Marginal venation of Pterocarya pugetensis............................................................................................................................ 57

11. Marginal venation of Alnus cuprovallis .................................................................................................................................... 58

12. Marginal venation of Alnus martini ........................................................................................................................................... 59

13. Marginal venation of Alnus sp ..................................................................................................................................................... 59

14. Marginal venation of Anamirta..................................................................................................................................................... 62

15. Marginal venation of Diploclisia................................................................................................................................................... 64

16. Hypserpa? sp......................................................................................................................................................................................... 64

17. Marginal venation of Limacia ....................................................................................................................................................... 64

18. Marginal and intercostal venation of Paratinomiscium and Tinomiscium................................................................... 65

19. Venation of Pycnarrhena................................................................................................................................................................. 65

20. Intercostal venation of Knema....................................................................................................................................................... 66

21. Marginal venation of Myristica..................................................................................................................................................... 66

22. Marginal venation of Cananga ..................................................................................................................................................... 66

23. Marginal venation of Prunus ......................................................................................................................................................... 72

24. Marginal venation of Sorbus carbonensis ................................................................................................................................. 72

25. Venation of Caesalpinites sp ......................................................................................................................................................... 72

26. Marginal and intercostal venation of Luvunga....................................................................................................................... 73

27. Marginal and intercostal venation oi Melanorrhoea............................................................................................................. 74

28. Marginal venation of Ilex carbonensis ....................................................................................................................................... 75

29. Marginal and intercostal venation of Goweria alaskana..................................................................................................... 76

30. Intercostal venation of Phytocrene acutissima ....................................................................................................................... 77

31. Marginal venation of Pyrenacantha ........................................................................................................................................... 11

32. Marginal venation of Stemonurus .............................................................................................................................................. 78

33. Marginal venation of Meliosma.................................................................................................................................................... 79

34. Ultimate venation of Plafkeria sp................................................................................................................................................. 81

35. Marginal venation of Saurauia ..................................................................................................................................................... 81

36. Venation of Parashorea ................................................................................................................................................................... 82

37. Marginal and intercostal venation of Barringtonia............................................................................................................... 83

38. Marginal venation of Alangium ................................................................................................................................................... 84

39. Intercostal venation of Eugenia..................................................................................................................................................... 84

40. Venation of Salix and Lithocarpus............................................................................................................................................... 87

41. Venation of Heteromeles and Salix ............................................................................................................................................. 91

42. Venation of Plafkeria, Byttneria, and Alangium ................................................................................................................... 92

43. Venation of Dipterocarpaceae and Rhamnaceae....... ........................................................................................................... 92

TABLES

Page

TABLE 1. Check list of megafossil plants from the Kushtaka, Katalla, Kulthieth, and Poul Creek Formations 10

2. The effect of habitat on leaf margins ...........................................................................-..-..................---.....-.-..-..-.--..... 24

3. Percentages of species that have entire margins in some modern floras ........................................................................................ 25

4. Comparisons between modern and fossil foliar physiognomy ............................................................................................................ 29

5. Distribution of leaf-size classes in the vegetation of eastern Australia............................................................................................ 29

6. Numerical abundance of plant megafossils from the Alaskan lower Ravenian ........................................................................ 30

7. Numerical abundance and diversity of families in the Alaskan lower Ravenian ...................................... ......... ................ 31

8. Specific diversity through time of some families represented in the west American Tertiary ................................................ 47
PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

By JACK A. WOLFE

ABSTRACT

Numerous collections of fossil plants from the Gulf of Alaska 

region were obtained from rocks that are well dated by marine 

mollusks. The mollusks indicate that the oldest possible age for 

the lowest plant assemblage is middle Eocene (Domengine) and 

that the youngest possible age for the highest Paleogene plant 

assemblage is middle Oligocene (Lincoln).

Paleobotanical correlations indicate that the lowest plant 

assemblage is of late middle Eocene (early Ravenian) age, and the 

highest plant assemblage is of early Oligocene (Kummerian) age. 

A new provincial stage the Angoonian is proposed, based on 

assemblages from rocks in southeastern Alaska. The Angoonian 

is thought to be of late Oligocene (early Blakeley) age. Only one 

stratigraphically isolated assemblage from the Gulf of Alaska 

region has been recognized as of Angoonian age. The 

biostratigraphy of the Gulf of Alaska region and of other regions 

in Alaska indicates that the Seldovian, Homerian, and 

Clamgulchian Stages are probably entirely of Neogene age.

The early Ravenian assemblages represent Paratropical Rain 

forest (that is, similar to vegetation of a region bordering the 

Tropical Rain forest); this conclusion is based on the physiognomic analysis of foliage. The most diverse families represented 

are Menispermaceae and Icacinaceae, which are accompanied by 

palms, Annonaceae, Myristicaceae, Dipterocarpaceae, Barringtoniaceae, and Myrtaceae. A minor element is represented by 

broad-leaved deciduous plants. Middle Ravenian assemblage was 

somewhat cooler (subtropical), as indicated by foliar physiognomy, the fewer Tropical Rain forest elements, and the diversity 

of Lauraceae. The late Ravenian assemblage represents a 

temperate forest. The Kummerian assemblages were again 

dominated by Lauraceae, and the physiognomic characters of the 

foliage indicate that a subtropical forest was again represented.

The existence of a broad-leaved evergreen forest in Alaska, such 

as that of the early Ravenian, indicates that extended periods of 

darkness may not have existed at that time. The Alaskan 

Paleogene floras, as well as those from other regions, indicate that 

the earth's axis of rotation may not have had as great an 

inclination in the past.

The history of the development of the Arcto-Tertiary concept 

shows the fundamental lack of evidence for this concept. An 

alternative, more complex, concept is proposed for the development of the Mixed Mesophytic forest. Paleobotanical evidence 

indicates that tolerances of many genera have changed during the 

Tertiary. Many mixed mesophytic genera particularly those 

that are today temperate outliers of basically tropical groups 

may have had a wide distribution in the Paleogene Paratropical 

Rain and subtropical forests and have entered the warm 

temperate forests only during the later Paleogene or Neogene. 

Some mixed mesophytic groups seem to be of later Neogene origin 

and have been significant members of the warm temperate forests 

since that time. Other mixed mesophytic genera apparently were

members of the Paratropical Rain forest only in the earliest 

Paleogene and became adapted to warm temperate climates by the 

late Eocene or perhaps even earlier. The Mixed Mesophytic forest 

may have developed independently in Eurasia and western North 

America.

In the discussion of the systematics of the Gulf of Alaska 

Ravenian and Kummerian assemblages, at least 160 species are 

represented, but the fragmentary nature or poor preservation of 

many does not allow a generic or familial assignment; 126 species 

are discussed and illustrated. They represent at least 45 families 

and 83 genera. Systematic revisions of some species in Paleogene 

floras in conterminous United States are made.

INTRODUCTION

Of all the Tertiary sequences at high northern 

latitudes, the Gulf of Alaska sequence is unique in 

having many plant localities in rock units that also 

contain numerous marine fossils. More significantly, some of the Gulf of Alaska floras of Eocene 

and early Oligocene age are the most northerly floras 

that can be definitely stated to be of that age. The 

significance of these floras, therefore, to floristic and 

vegetational history is apparent: do all these Eocene 

and early Oligocene floras represent a temperate, 

broad-leaved deciduous forest similar to that of the 

supposed Eocene-Oligocene Kenai Formation? If the 

concept of an Arcto-Tertiary Geoflora has validity, 

they must represent this type of vegetation. New 

collections of fossil plants from the Gulf of Alaska 

sequence, coupled with a considerable amount of 

new stratigraphic and molluscan data, allow a far 

better understanding of floristic and vegetational 

history in northern latitudes than has been possible 

before.

The Paleogene floras from the Gulf of Alaska 

section also have an important bearing on the ages 

of other Alaskan floras. Some paleontologists, 

adhering rigorously to the largely unsubstantiated 

concept of homotaxis, have questioned whether 

Paleobotanical correlations can be made between 

floras separated by several degrees of latitude. The 

floras from the Gulf of Alaska section are, however, 

at approximately the same latitude as the floras from 

the Kenai Formation in the Cook Inlet basin, and 

geographically they are only about 240 km (160 mi)
PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

distant. Comparisons between the Gulf of Alaska 

and Kenai assemblages will, therefore, be unaffected 

by latitudinal zonation of vegetation and climate.

Fossil plants from the Gulf of Alaska sequence 

were first collected by U.S. Geological Survey field 

parties in the early 1900's, largely through the efforts 

of G. C. Martin and R. A. Tarr in the Katalla and 

Malaspina districts. Early analysis of these collections by F. H. Knowlton yielded conflicting 

results. Regarding the material from the Malaspina 

district, Knowlton stated (in Tarr, 1906, p. 63): 

"***none of the plants is referable to the Kenai, and, 

if the exigencies of the stratigraphy demand that 

they should be referred to the Pliocene, there is 

nothing to contradict it." The material from Berg 

Lake was considered by Knowlton (in Martin, 1908, 

p. 35) to be somewhat older: "So far as I am able to 

determine, this is not Kenai in age, but just what the 

age is I am uncertain. I should think it ought to be 

Miocene, but without an extensive comparison with 

known Miocene floras its exact position is in doubt." 

Another collection from the same unit as the Berg 

Lake collection was thought by Knowlton (in Martin, 

1908, p. 35) to be considerably older; but we must keep 

in mind that the "Arctic Miocene" was considered by 

Knowlton to be of Eocene age. Knowlton stated: 

"These are well-known forms found in the so-called 

Arctic Miocene and indicate this age for the beds 

whence they came." Martin (1908, p. 40), however, 

who based his conclusions largely on the marine 

mollusks determined by W. H. Dall, stated: "There is 

no doubt that the entire sequence is Tertiary and 

post-Eocene ***."

Shortly after Knowlton's statements were published, Arthur Hollick undertook his studies of 

Alaskan Tertiary plants. The age of the unit 

carrying the Berg Lake flora, the Kushtaka 

Formation, was revised downward by Maddren 

(1914, p. 130): "The Kushtaka carries fossil plants, 

which the recent studies of Arthur Hollick have 

shown to be of Kenai age (Eocene)." In his final 

description of the Alaskan Tertiary plants, Hollick 

(1936) did not specifically discuss the age of the Gulf 

of Alaska plants, but considered that (p. 23) "***the 

general similarity of the Alaska Tertiary flora to that 

of the Eocene in the States proper would appear to be 

demonstrated***." Significantly, however, not 1 of 

the 12 species that Hollick (1936, p. 23) listed as 

occurring in the Alaskan Tertiary as being 

"characteristic of the Fort Union" were listed by 

Hollick as occurring in any of the Gulf of Alaska 

assemblages. P. S. Smith (in Hollick, 1936, p. 26) thus 

remarked concerning the Berg Lake section: "It 

seems reasonably certain, however, that the entire

sequence *** is post-Eocene in age. ***it will be 

observed that the flora of the coal measures 

(Kushtaka formation) does not resemble the type 

Kenai flora, of upper Eocene age, but seems 

younger."

Chaney (1940) considered the Berg Lake flora to be 

of Eocene age, although he cited no new evidence in 

contradicting the age assignment based on the 

stratigraphic and molluscan evidence that was then 

available. Chaney's assignment of the Berg Lake 

flora to his group of "temperate" floras is likewise 

puzzling. Included in Hollick's determinations were 

genera such as Piper, Mohrodendron, Magnolia, 

Cinnamomum, Persea, Malpoenna, Terminalia, and 

Semecarpus none of these genera are indicators of 

a temperate climate.

Mapping of the Gulf of Alaska Tertiary rocks was 

started by D. J. Miller in the 1940's and continued by 

George Plafker in the 1950's. Miller and his 

associates collected few fossil plants, but the 

mapping and the many collections of marine 

mollusks made by Miller and Plafker clarified the 

stratigraphy and revised the ages of most of the older 

plant localities. The stratigraphic interpretations 

adopted in this report are largely those of Miller 

(1957,1961a, 1961b, 1961c), Plafker and Miller (1957), 

and Plafker (1967).

Geologic investigations by the petroleum industry 

in the 1960's has resulted in several new collections 

of fossil plants. In particular British Petroleum 

Company and Shell Oil Company have contributed 

significant collections.

The collections made by the petroleum industry in 

the early 1960's and the older collections partly 

described by Hollick formed the basis for preliminary discussions of the vegetational significance of the Paleogene assemblages from the Gulf of 

Alaska region (Wolfe, 1964, p. N6; 1966, p. B5). These 

assemblages were considered to indicate a subtropical climate. Wolfe and Hopkins (1967, p. 70-73) 

considered the then small mollusk-dated assemblages from the Kushtaka Formation to be of late 

Eocene age and to be subtropical to tropical because 

of the physiognomic characters of the leaves. The 

Tokun and Katalla assemblages were also thought 

to indicate a warm climate, although somewhat 

cooler than the Kushtaka.

The present report is based on the old and a few 

new collections made by U.S. Geological Survey 

geologists, collections submitted by petroleum 

companies, and new collections that I obtained in 

1968. I particularly wish to thank the Humble Oil 

Company for providing logistic support during 1968; 

Drs. Willard Larson and Donald Gunn of Humble
PHYSICAL STRATIGRAPHY

'"62 US'.

11160

60°

143° 142°

FIGURE 1. Map of the Gulf of Alaska region. Numbers represent plant localities.

assisted in making most of the new collections.

The help of George Plafker has been invaluable in 

preparing the stratigraphic discussion. Dr. H. D. 

MacGinitie and H. E. Schorn, University of 

California (Berkeley), have given much time in 

discussing the floristic and vegetational implications of the Gulf of Alaska floras. Dr. Hugh H. Iltis, 

University of Wisconsin, has also made valuable 

suggestions about this report.

PHYSICAL STRATIGRAPHY

KATALLA DISTRICT

Most collections from the Gulf of Alaska Paleogene rocks were made north of latitude 60° N. (see fig. 

1 and also fig. 6). The most complete sequence of 

Paleogene plant assemblages was obtained in the 

western part of the Gulf of Alaska region in the 

Katalla district. Miller (196la) made a detailed 

geologic map of this district, and the map shown here 

(fig. 2) has been adapted from it. The structure in this 

district is complex, but, despite the small-scale 

thrusts and strongly folded beds (see pi. 1), the beds 

can generally be traced for a considerable distance, 

and hence a good section can be pieced together (fig. 

3). The Paleogene formations in the Katalla district 

are, in ascending order, the Stillwater, Kushtaka, 

Tokun, and Katalla Formations.

Most collections from the Yakataga and Malaspina districts, which are east of the Katalla district, 

are small. The oldest named unit, the Kulthieth 

Formation, has yielded most of the Paleogene plants; 

the younger unit, the Poul Creek Formation, has 

yielded almost no plants. The paucity of material 

from the Yakataga and Malaspina districts reflects

the limited time spent searching for localities and 

collecting.

STILLWATER AND KUSHTAKA FORMATIONS

The Stillwater Formation is thought to be the 

oldest of the four units under discussion, although it 

may in part be a lateral equivalent of the Kushtaka 

(MacNeil and others, 1961, p. 1803, 1807). In the 

section exposed northeast of Berg Lakes, in which 

Miller (1961a) did not differentiate between the two 

units, a series of sandstones and siltstones grades 

upward into a sequence of sandstone, siltstone, and 

coal. Although the lower part of the section contains 

a minor amount of very thin coal beds, it is 

lithologically similar to the type section of the 

Stillwater Formation and this lower part is here 

considered to be part of the Stillwater. Mollusks from 

USGS Cenozoic locality M3880 were determined by 

W. O. Addicott as:

Rimella? aff. R. macilenta White

Brachiodontes cowlitzensis (Weaver and Palmer)

Pitar cf. P. californianus (Conrad) 

Addicott (written commun., Oct. 10,1968) assigns a 

middle or late Eocene age to these mollusks. The beds 

from which the mollusks were obtained grade 

westward into the coal-bearing Kushtaka Formation 

and are approximately on strike with plant localities 

3847,11157,11158, and 11159. 1 Plants from localities 

11158 and 11159 are approximately 300 feet 

stratigraphically above locality M3880, and plants

'All fossil plant localities cited by number in this report are in the U.S. Geological 

Survey Paleobotany catalog, Washington, D.C. See also p. 93.
PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

60° 30' ~

01234567 8 KILOMETERS

FIGURE 2. Generalized geologic map of part of the Katalla district. Geology after Miller (1961a). Dashed lines represent ridge crests 

Ts, Stillwater Formation; Tk, Kushtaka Formation; Tsk, Stillwater and Kushtaka Formations, undivided; Tt, Tokun Formation. 

Numbers in italic indicate plant localities; numbers in roman type indicate mollusk localities.

from locality 11166 are approximately 2,000 feet 

higher than locality M3880. The beds at locality 

11166 can be traced in a rough manner westward 

along the flank of Carbon Mountain; these beds 

appear to crop out on the south nose of Carbon 

Mountain about 1,500-2,000 feet stratigraphically 

higher than localities 3847 and 11157, and about 

3,000 feet statigraphically lower than plant locality 

11167 on Carbon Mountain. Localities 9891 and 

11169 are about 50 and 300 feet, respectively, higher 

than locality 11167.

Assuming that no major structure is concealed by 

the alluvium in Canyon Creek, plant localities 11160- 

11162 are estimated to be about 100-1,000 feet higher 

than locality 11159. Locality 11160, which is in the 

coal-bearing Kushtaka Formation, is near the base 

of that unit as mapped by Miller (1961a). Southward 

the Kushtaka grades stratigraphically downward 

into the Stillwater Formation. In the Shockum 

Mountains, mollusks from the Stillwater occur at a 

locality that is probably more than 1,000 feet below 

locality 11160, that is, probably a few hundred feet 

below locality 3847. The mollusks (USGS Cenozoic 

loc. M1207) include, according to F. S. MacNeil 

(written commun., Oct. 24, 1961):

Whitneyella cf. W. sinuata aragoensis Turner 

Tivelina cf. T. vaderensis (Dickerson) 

Eometrix cf. E. martini (Dickerson) 

MacNeil assigns a possible middle Eocene age to 

these mollusks. Collections of mollusks elsewhere in 

the Stillwater Formation are also regarded by 

MacNeil and others (1961, p. 1803) as of middle 

Eocene age. Thus the Stillwater Formation and 

possibly the lowest part of the overlying Kushtaka 

Formation exposed in the vicinity of Berg Lakes as 

well as the Berg Lake fossil plant assemblages (Iocs. 

3847, 11157, 11158, 11159) are, on molluscan and 

stratigraphic evidence, considered to be of late 

middle or early late Eocene age. The Canyon Creek 

assemblages (Iocs. 11160,11161,11162) would also be 

of middle or late Eocene age.

In the upper part of the Kushtaka Formation that 

is exposed along Canyon Creek, plants were 

collected from two localities (Iocs. 3846 and 11164). 

These localities are approximately 700 and 1,000 

feet, respectively, stratigraphically above locality 

11162. A few hundred feet higher than locality 11164 

in the basal part of the Tokun Formation that rests 

conformably on the Kushtaka, marine mollusks 

have been collected. USGS Cenozoic locality 4438
PHYSICAL STRATIGRAPHY

MILLER HILLS, 

NICHIWAK MOUNTAIN

Katalla 

Formation

CHARLOTTE RIDGE, 

LAKE TOKUN

CUNNINGHAM 

RIDGE

CARBON 

MOUNTAIN

Tokun 

Formation

J1168 

M3853

FEET METERS 

4000

2000 

1000

- 500

- ^~^ - -^x

Katalla 

Formation

Tokun 

Formation

Kushtaka 

Formation

^ ~ 

 15791

 D229(T)

 11165 

 9389

 M1215

^ ~~^~~~^~^

Tokun 

Formation

Kushtaka 

Formation

Stillwater 

Formation

 4438

 1 1 164 

 11163

 3846

 11162

 11160

 M1207

Kushtaka 

Formation

Stillwater 

Formation

 11169

 9891 

 11167

 11166

__ 77/53 

 77758

11157 

 3847

 M3880

FIGURE 3. Stratigraphic position of fossil plant localities in the Katalla district. Numbers in italic are 

plant localities; numbers in roman are mollusk localities.

contains Perse washingtonensis (Weaver); this 

species indicates a Tejon (late Eocene) age (W. O. 

Addicott, oral commun., September 1968).

To the west on Charlotte Ridge, the basal Tokun 

has yielded a larger and better preserved fauna. The 

determinations by L. G. Hertlein (written commun., 

July 30, 1954) include:

Acila cf. A. decisa (Conrad)

Gari cf. G. columbiana (Weaver and Palmer)

Pitar cf. P. californianus Conrad

Ficopsis cf. F. cowlitzensis Weaver 

Molopophorus cf. M. tejonensis Dickerson 

Turritella uvasana Conrad 

Whitneyella cf. W. coosensis Turner 

Whitneyella cf. W. sinuata Gabb var. 

This fauna, according to Hertlein, is comparable to 

that of the Tejon "stage" of Weaver and others (1944) 

and is of late Eocene age. Localities 9389 and 11165 

are in the uppermost Kushtaka on Charlotte Ridge; 

these localities are about 300 feet below the
PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Kushtaka-Tokun contact. Farther to the southwest 

on Charlotte Ridge, beds in the Kushtaka that are 

several hundred feet below the Kushtaka-Tokun 

contact have yielded mollusks. The mollusks from 

this locality (USGS Cenozoic loc. M1215, determined 

by F. S. MacNeil, written commun., Oct. 24, 1961) 

include:

Whitneyella cf. W. washingtoniana (Weaver)

Acila cf. A. decisa (Conrad)

MacNeil considers these fossils to be probably late 

Eocene (Tejon) in age. The upper part of the 

Kushtaka Formation on Charlotte Ridge and the 

Charlotte Ridge fossil plant assemblage are thus of 

probable late Eocene age.

The Tokun Formation is not present on Carbon 

Mountain, where the Kushtaka Formation attains 

its greatest thickness. Beds that appear to be 

equivalent on stratigraphic evidence to the uppermost Kushtaka on Charlotte Ridge are overlain by a 

few thousand feet of coal-bearing rocks also 

assignable to the Kushtaka on lithologic evidence. 

Thus at least part of the Kushtaka on Carbon 

Mountain is probably equivalent in age to the Tokun 

Formation. The total age range of the Tokun 

Formation is thought on molluscan evidence to be 

late Eocene through earliest Oligocene, that is, of 

Tejon and Keasey age (MacNeil and others, 1961, p. 

1803). The part of the Tokun that is of Keasey age is 

indicated by the occurrence at USGS Cenozoic 

locality 15791 of a crushed specimen determined as 

Turcicula columbiana Dall? by H. E. Yokes (written 

commun., Jan. 12, 1946). The Kushtaka Formation 

on Carbon Mountain could therefore be at least a 

young as Keasey age.

KATALLA FORMATION

The Katalla Formation seems to be entirely 

marine, and thus only a few localities have yielded 

plant megafossils. Most of these localities are in the 

Split Creek Sandstone Member, the basal member of 

the Katalla. The largest assemblage of plants from 

the Split Creek was obtained at locality 11168 at the 

base of Nichiwak Mountain. A few mollusks were 

also collected at this locality, including a gastropod 

that is closely related to Bruclarkia fulleri Durham, 

an index to the lower part of the "Lincoln Stage" (W. 

O. Addicott, written commun., Oct. 10, 1968). 

Elsewhere in the district, the Split Creek Sandstone 

Member has yielded a large fauna that Yokes 

(written commun., Jan. 12,1946) considered to be the 

same age as the Molopophorus stephensoni zone, 

that is, earliest "Lincoln" (earliest middle Oligocene). Some of the more significant localities are 

listed at the end of this report. A few specimens of

leaves have been obtained from the Split Creek at 

localities 3705, 9552, 9988, 9989, and 9990.

The overlying Basin Creek Member of the Katalla 

has yielded a very few leaves at localities 11181 and 

California Academy of Sciences locality 29181. The 

Basin Creek is obviously at least in part of "Lincoln" 

age, that is, middle Oligocene. This age is confirmed 

by the molluscan fauna, which, according to H. E. 

Yokes (written commun., Jan. 12, 1946) is not 

younger than the Molopophorus gabbi zone; F. S. 

MacNeil (oral commun., October 1968), however, 

considers part of the Basin Creek Member to be as 

young as early Blakeley (late Oligocene).

A summary of the probable age assignments of the 

plant-bearing formations of the Katalla district is 

shown later (p. 21).

YAKATAGA AND MALASPINA DISTRICTS

The geology of these districts has been discussed 

primarily by Miller (1957,1961b, 1961c), Plafker and 

Miller (1957), and Plafker (1967). Three units have 

yielded plant megafossils: the Kulthieth, Poul Creek, 

and Yakataga Formations. An unnamed marine 

siltstone and sandstone sequence in the northwestern part of the Malaspina district is thought to 

intertongue with, and in part underlie, the Kulthieth 

Formation (Plafker and Miller, 1957; MacNeil and 

others, 1961); marine megafossils indicate a probable middle Eocene age for this sequence (MacNeil 

and others, 1961).

KULTHIETH FORMATION

One of the most extensive sections of the Kulthieth 

Formation is exposed on a ridge north of a stream 

that heads near the headwaters of the Kulthieth 

River in T. 18 S., R. 17 E., and flows southeast into the 

Duktoth River; the upper part of this section is 

described in detail by Miller (1957, sec. no. 2). The 

only determinable marine mollusks thus far collected from this section came from the upper part of 

the Kulthieth Formation. USGS Cenozoic locality 

M1368 has yielded a fauna including Acila shumardi 

(Dall) and Boreoscala condoni cf. subsp. oregonense 

Dall. (These were determined by F. S. MacNeil, 

written commun., Dec. 20, 1961.) The Acila clearly 

places this part of the Kulthieth in the Lincoln 

"stage" (middle Oligocene); the species of Boreoscala 

is restricted to the two lower zones of the Lincoln 

"stage" and the subspecies to the lowest zone 

(Molopophorus stephensoni) according to Durham 

(1944). The only determinable plants from this 

section of the Kulthieth are from locality 11170. The 

precise stratigraphic separation of this plant locality 

from the mollusk locality is unknown; Miller and
BIOSTRATIGRAPHY

Plafker measured approximately 9,000 feet of 

Kulthieth in this section, but the plant locality is 

farther downstream than the base of Miller and 

Plafker's measured section. Miller (1957) considered 

that the rocks in the area of locality 11170 "*** may 

be largely or entirely represented in the lower part of 

the section described above." This uncertainty is 

caused by the complex folding and faulting in this 

area. In any case it appears highly probable that 

locality 11170 is at least several thousand feet 

stratigraphically below the lower middle Oligocene 

mollusks and is in the lower part of the Kulthieth 

Formation in this area (George Plafker, oral 

commun., October 1968). Elsewhere in the Yakataga 

District the upper part of the Kulthieth has yielded 

mollusks of late Eocene (Tejon) age (Miller, 1957). 

The available evidence thus indicates that the flora 

from locality 11170 is no younger than late Eocene 

and presumably could be older. The locality, 

however, that has yielded the most diagnostic 

mollusks Turritella uvasana Conrad and Ficopsis 

cowlitzensis Weaver is USGS Cenozoic locality 

16859; it is in a complexly folded and faulted area, 

and the relationship of this locality to the KulthiethPoul Creek boundary has not been determined.

Several small collections of plants have been made 

from the upper part of the Kulthieth Formation in the 

eastern drainage of the Kulthieth River. Localities 

9893 and 9894 are situated at about the KulthiethPoul Creek boundary as mapped by Miller (1961b).

One small collection from locality 9551 in the 

Hanna Lake area is of uncertain stratigraphic 

position, but D. J. Miller (written commun., May 25, 

1961) considered the locality probably to be from the 

upper part of the Kulthieth.

In the Malaspina district, an isolated outcrop of 

coal-bearing rocks on Esker Stream (locality 3879), 

which flows into Yakutat Bay, has provided a small 

collection of plants. Although assigned with 

reasonable certainty to the Kulthieth Formation 

(Plafker and Miller, 1957), the stratigraphic position 

of this locality within the Kulthieth is unknown.

In the Samovar Hills, the Kulthieth Formation has 

yielded marine mollusks of Tejon (late Eocene) age 

(Plafker and Miller, 1957). Stratigraphically well 

below the mollusk-bearing beds, plants were 

obtained at locality 11195; these plants are thus no 

younger than late Eocene and could be of middle 

Eocene age.

POUL CREEK FORMATION

Plant megafossils from the Poul Creek Formation 

have been found in only two localities. One locality

(9895) is apparently in the lower part of Poul Creek, 

but no mollusks have been collected from the area. 

Elsewhere in the region, the Poul Creek contains 

mollusks of late Lincoln (late middle Oligocene) 

through late Blakeley (early Miocene) age. Presumably, therefore, the plants from locality 9895 are of 

late middle or late Oligocene age.

At another locality (11182), a single fruit of 

Juglans was found; the fruit was collected as float, 

but, according to D. J. Miller (written commun., May 

25,1961), the specimen probably came from the Poul 

Creek Formation.

YAKATAGA FORMATION

Although plant fossils are reported to be abundant 

at several localities in the Yakataga Formation, only 

two collections have been made. Locality 11184 from 

the upper part of the Yakataga in the Pinnacle Hills 

is in a sequence that MacNeil (1967) considers to be of 

late Miocene and Pliocene age, based on the 

occurrence of certain pectinids. The second assemblage of plants in the Yakataga (loc. 11183) is 

assigned to the Yakataga Formation because the 

plant-bearing sequence is similar lithologically to 

the marine Yakataga elsewhere (George Plafker, 

oral commun., October 1968). This sequence, which 

occurs as a sliver along a fault, cannot be definitely 

related by field relationships to the Tertiary section 

elsewhere in the Malaspina district.

UNKNOWN UNIT

Plant fossils from locality 11185 and locality 11183 

came from slivers along the same fault. No marine 

fossils occur in this sequence of rocks; although these 

rocks were considered to be part of the Kulthieth 

Formation by Stonely (1967), paleozoological or 

lithologic evidence for the designation is lacking.

BIOSTRATIGRAPHY

INTRODUCTION

The general age assignments of all but one of the 

Paleogene plant localities are known with reasonable certainty because of their stratigraphic relationships to marine mollusk localities (see preceding 

section). Some of the localities, however, are not 

precisely placed stratigraphically relative to mollusk 

localities, and their age assignments must be based 

on the contained plant fossils. The close stratigraphic relationships between some of the mollusk 

and plant localities allows a better understanding of 

the age relationships between the marine megafossil 

chronology and some of the Paleogene stages that 

are based on studies of the plant assemblages of the
8 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Puget Group in Washington (Wolfe, 1968).

Four stages and seven substages were recognized 

in the Puget Group. The Franklinian the lowest 

stage is thought to be equivalent to at least part of 

the Capay "stage" (lower Eocene), and the Fultonian 

is probably equivalent to at least part of the 

Domengine "stage" (middle Eocene). No assemblages yet known from the Gulf of Alaska section 

seem to contain species definitely indicative of these 

two stages. The two upper stages the Ravenian and 

the Kummerian are represented, however.

The time equivalents of the early Ravenian in 

terms of the Pacific Coast marine megafossil 

chronology are not certain. The overlying middle 

Ravenian is almost certainly equivalent in part to 

the upper Eocene (Tejon "stage"), as indicated by the 

occurrence of two assemblages. One assemblage 

discussed later on Charlotte Ridge is bracketed 

stratigraphically by mollusks of Tejon age and is 

also overlain by several hundred feet of strata that 

are also of Tejon age. This evidence indicates that 

the middle Ravenian assemblage falls low in the 

Tejon "stage." A second assemblage that from the 

Mclntosh Formation of Washington is less certainly dated. The foraminifera from the Mclntosh 

indicate that the formation ranges in age from the 

middle Eocene (later part of the Domengine "stage") 

to the early part of the late Eocene (Snavely and 

others, 1958). The assemblage of plants from the 

Mclntosh occurs in strata that cannot be certainly 

related to the foraminiferal localities, but the plantbearing beds are in the upper part of the formation 

and are thus presumably of early late Eocene age. If, 

therefore, the overlying middle Ravenian is of early 

late Eocene age, the early Ravenian is most probably 

of late middle Eocene age, that is, equivalent to part 

of the Domengine "stage."

In the Puget Group of Washington, early Ravenian 

assemblages were obtained from the lower part of the 

Tukwila Formation. Late Fultonian assemblages, 

however, were obtained from the underlying Tiger 

Mountain Formation, which, in turn overlies the 

marine Raging River Formation of middle Eocene 

age (Wolfe, 1968). The lower Ravenian, therefore, is 

most probably the time equivalent of the late 

Domengine and possibly also the "Transition 

stage;" this stage represents a poorly known interval 

between the definite middle and definite late Eocene.

Except for some minor departures of no more than 

zone magnitude, the epochal and subepochal 

boundaries of the marine megafossil geochronology 

on the Pacific Coast will be used in this report. The 

assignments of the various plant stages in terms of

epochs is thus: early Ravenian = late middle Eocene, 

middle Ravenian = early late Eocene, late Ravenian = 

latest Eocene, and Kummerian = early Oligocene. 

The two departures from the marine megafossil 

geochronology are that the lower part of the Keasey 

"stage" is here considered to be of earliest Oligocene 

rather than latest Eocene age, and the lowest zone 

(Molopophorus stephensoni) zone of the Lincoln 

"stage" is here considered to be of latest early rather 

than earliest middle Oligocene age. Some molluscan 

workers, in fact, place the M. stephensoni zone in the 

early rather than the middle Oligocene (H. E. Yokes, 

written commun., Jan. 12, 1946).

The placement of the Eocene-Oligocene boundary 

in this report is purely arbitrary, but it is in 

approximate agreement with both mammalian and 

marine molluscan paleontologists working in western North America (Wolfe, 1968). Foraminiferal 

workers, however, would probably place this 

boundary much higher and include all equivalents of 

the Kummerian Stage and the Keasey and Lincoln 

"stages" in the late Eocene. At least part of the 

problem in these differing correlations stems from 

the lack of agreement on the placement of the 

Eocene-Oligocene boundary in Europe.

The Oligocene-Miocene boundary adopted in this 

report is also purely arbitrary, and it is placed 

between the Angoonian and Seldovian Stages. The 

Angoonian includes assemblages that are known to 

be equivalent in age to the uppermost zone of the 

Lincoln "stage," the lower zone of the Blakeley 

"stage," and probably to the upper zone of the 

Blakeley, that is, latest middle Oligocene through 

earliest Miocene in the marine megafossil geochronology. Foraminiferal workers would place the 

boundary even higher than I have, whereas 

mammalian paleontologists would place the boundary lower. As with other epochal boundaries, the 

varying correlations are in large part a result of lack 

of agreement in varying paleontological disciplines 

on the placement of the epochal boundaries in 

Europe.

RAVENIAN STAGE

LOWER RAVENIAN

In the original definition of the Ravenian Stage 

(Wolfe, 1968, p. 7), the stage was subdivided into two 

informal substages, but here (p. 9, 12) a threefold 

subdivision is proposed. The Kushtaka floras from 

locality 3846 and stratigraphically lower horizons 

are no younger than early Ravenian, as indicated by 

the occurrence at that locality of Cyathea inequaliteralis (Hemitelea pinnata MacG.) and of Goweria
BIOSTRATIGRAPHY 9

dilleri at locality 3847 (see table 1). Locality 3846 has 

also yielded specimens of Litseaphyllum presamarensis, a species that has previously been found in 

the middle Ravenian and younger rocks. Its presence 

thus indicates that this locality is probably near the 

border between the lower and middle Ravenian.

Whether some localities low in the Kushtaka near 

Berg Lake are of late Fultonian or early Ravenian 

age is not clear. Alangium bergensis is apparently 

restricted to the upper Fultonian in the Puget Group, 

but thus far it has been found only at a single locality 

there. Zizyphus alaskanus, on the other hand, is 

known at two lower Ravenian localities, and 

Litseaphyllum similis is known in the Puget Group 

from early Ravenian and younger beds. In addition, 

both Goweria alaskana and Phytocrene acutissima 

are known from the early Ravenian Steel's Crossing 

assemblage of the Puget Group. The Alaskan 

Illicium is not sufficiently well preserved to be 

certain of the identification, but it appears to represent the same species that is known from the Steel's 

Crossing assemblage. The bulk of the evidence indicates that the lower part of the Kushtaka Formation 

exposed at Berg Lakes is probably of early Ravenian 

(late middle Eocene) age.

The lower part of the Kulthieth Formation exposed 

near Duktoth River is also probably early Ravenian. 

Several species (see table 1) are common to the 

assemblage from locality 11170 and the assemblages 

from the lower part of the Kushtaka. In addition 

Anemia gunni is known to range from the 

Franklinian through the early Ravenian in the 

Puget Group (called A. eocenica Berr. by Wolfe, 1968, 

p. 5, 8). Phytocrene sordida has been previously 

recorded from beds of Franklinian age in California 

(MacGinitie, 1941), but the prior fossil record of 

leaves of Icacinaceae is so meager that little age 

significance can be attached to this occurrence in 

Alaska at this time. The age assignment of the 

Duktoth River assemblage to the early Ravenian is 

consistent with the stratigraphic and molluscan 

evidence (see p. 7); this evidence indicates a late 

Domengine or early Tejon age for the lower part of 

the Kulthieth Formation elsewhere in the Yakataga 

district.

The beds on Esker Stream have yielded only a few 

species. The occurrence there of Allantodiopsis 

pugetensis indicates an age no younger than middle 

Ravenian. The occurrences of Celastrus comparabilis and Goweria alaskana also indicate a 

probable correlation with the lower Kushtaka, that 

is, an early Ravenian age.

The lower part of the Kulthieth Formation in the

Samovar Hills has yielded a well-preserved although 

small collection of plants (loc. 11195). The species 

include Platycarya pseudobrauni and Parashorea 

pseudogoldiana, indicating a probable early Ravenian age.

MIDDLE RAVENIAN

The original biostratigraphic definition of the 

upper Ravenian was based on assemblages from 

the middle and upper part of the type section and 

from an assemblage (Cashman) from what was 

thought, on stratigraphic evidence, to be in beds 

equivalent to the lower part of the type section. 

Almost all the species known from the type section of 

the upper Ravenian are also known from the 

Cashman assemblage, but the Cashman is much 

richer than the assemblages from the type section. 

The situation prevails despite the fact that five 

localities in the type section were extensively 

collected. I originally thought that the depauperate 

nature of the assemblages of the type section might 

be the result of a peculiar ecology, but that this 

ecology prevailed during a period in which about 

1,000 feet of coal-bearing rocks were deposited 

appears improbable. Lithologic evidence does not 

indicate that the environment of deposition during 

the deposition of that part of the Puget Group was 

significantly different from the environment during 

the rest of Puget time.

An analysis of other late Eocene to early Oligocene 

assemblages many whose ages are radiometrically 

dated shows that the approximate time of deposition of the middle and upper parts of the upper 

Ravenian represents an interval considerably cooler 

than somewhat older and younger periods (Wolfe 

and Obradovich, unpublished data). In the Green 

River Canyon section of the Puget Group, standard 

biostratigraphic procedures have not indicated a 

distinct separation of the lower from the middle and 

upper parts of the type Ravenian, although many 

species have their highest occurrence in the middle 

Ravenian. I nevertheless propose that, because of the 

strong vegetational difference and because of the 

informal nature of the substages, the lower part of 

the type section of the former upper Ravenian (Wolfe, 

1960, fig. 2) be considered a distinct substage, the 

middle Ravenian. The only known assemblage in the 

Puget Group that represents this interval is the 

Cashman (loc. 9731). The term "upper Ravenian" as 

herein used now refers to beds equivalent in age to 

the beds in the Green River Canyon section from 

about the Gem-Harris horizon to the base of the 

Kummer sandstone bed (see Wolfe, 1968, fig. 2).

Four assemblages in the Kushtaka Formation
10 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

______TABLE 1. Checklist of megafossil plants from the Kushtaka. Katalla, Kulthieth, and Poul Creek Formations

Species

O 0!

Kushtaka Formation Katalla Formation Kulthieth Formation -, g

o o 

CM fc

05

f-OOCTJOC^CD CO -r lO f- CTiOO 00 ^ O

o^o^o^'-^CJi-^cOrHa^aiO^a^'-^O}

Anemia gunni .......................................... -_-_-----_------------X-------_

Anemia1? sp................................................ ____----------X___. ____________

Lygodium larsoni ................................. ----------XX. __________________

Cyathea inequilateralis ........................ _---XX_X_______________________

Allantodiopsis pugetensis .................. -_-X--XX--.X.__________X_______

Dryopteris alaskana .............................. X--__------------------X-------

Woodwardia sp ........................................ --------------X___. __..._______

Glyptostrobus sp .................................... -----------X-X__-----._X_______

Phoenicites sp .......................................... ----------- X___________________

Palmacites sp .......................................... ----X-----X____________________

Palmae, gen. indet ............................... --X--------X__________X_X______

Populus sp ............................................... -------------X_______. _________

Salix carbonensis .................................. -------------X_________________

Safct sp ...................................................... ------- ------X-----------------

Carya cashmanensis ........................... _----------X-__-__.__. _________

Engelhardtia sp-.. ..-.... ...... ~ -----------X---------. -.-.-....

Juglans sp ............................................... ------_------X__-_.____________

Platycarya pseudobrauni .................... X-----X____X__-__-____X_X_----_

Pterocarya pugetensis .......................... ----------XX_X_________________

Alnus cuprovallis .................................. -------------X_________________

Alnus martini .......................................... --X_____X-__--___-__--X________

Alnus scotti .............................................. X-X___________________x_-______

Alnus sp .................................................... ------_-_---_X_________________

Alnus sp .................................................... ------_---__-X__--_____________

Alnus sp .................................................... ------_-------X________________

Betula sp ................................................. ---------------- X ______________

Dryophyllum pugetensis ...................... ----------XXX-__.______________

Fagus sp ................................................... -----------------X---_-_. ______

Girroniera sp ............................................ _-X____________________________

Girroniera sp ................................... ........ ------------_--.-----. X________

Ulmus sp ... .............................................. ----------XX---------______-___

Tetracentron piperoides ............... ..... ----------XX---X______x________

Magnolia reticulata ............................. ------__--_____-X_____ _________

Illicium sp ....................................... ....... _--X___________-.______________

Anamirta milleri ................................... ----------- -__-----___X________

Calkinsia plafkeri .................................. -X-------X_x__--_______________

Cocculus sp .............................................. ------X______________. _________

Cocculus sp .............................................. ------_------X_________________

Diploclisia sp .......................................... --__------------_----. X_. _.--__

Hypserpa? sp .......................................... ---------------- X ______________

Limacia stenophylla ............................ _-XX__X________________________

Paratinomiscium conditionalis ....... -----------------------XX------

Pycnarrhena sp ..................................... ------X________________________

Knema sp .................................................. X---_-_________________________

Myristica sp ........................................... X--------------------. ---------

Cananga sp .............................................. ------_----_--_________x-------

Dasymaschalon? sp ............................. -_X____________________________

Cinnamomophyllum kushtakensis .. ----------XX___________________

Cinnamomophyllum latum ................ XX--X_X________________________

Litseaphyllum carbonensis ................ ---------------X_______________

katallaensis .......................................... -_----_---------X________x_____

praelingue ........................................... -----------X--_---___._________

presamarensis .................................... -X----_X___xx--X------_________

presanguinea ...................................... ----------_---X________________

schorni .................................................. ----------XX---------. _________

Litseaphyllum sp .................................... ---X___________________________

sp .............................................................. ----------__----X-----X________

sp .............................................................. _-___-_____X_________. _________
BIOSTRATIGRAPHY 

TABLE 1. Checklist of megafossil plants from the Kushtaka, Katalla, Kulthieth, and Poul Creek Formations Continued

Kushtaka Formation Katalla Formation Kulthieth Formation

Species __________________ ____________________ ______________ ___________________

 i LO lO OS 35 i Oi

~H 1C LO X X ^H X

-H 05 O; 05 05 05

Litseaphyllum sp ................................ ----------- X . .___._______.___..

sp X

Sp .... ---__---_____- X

Corylopsis sp .................................... ------_--_---X____________.,____

Liquidambar? sp ........ .................... ..-----------X---------__------__

Eucommia sp . ................... ......... _-______-__X______.____________

Platanus comstocki ............ .... ... ..--------------X______________XX

Prunus neuadensis ............. .......... ---------_-_-X-----------------

Sorbus carbonensis ...... ...... . -------------X_________________

Caesalpinites sp ............. . ....... X. -----------------------------

Caesalpinites sp ...................... .. ..... -----------X-------------------

Toona sp . .... ... ....... .................. -----------------------_X______

Euodia alaskana ............................. __X____________________________

Luvunga spatiosa . ... ............ ......... X-----X__. _____________________

Melanorrhoea alaskana ............. x-X----X--------------X____-___

Rhus cf. 7?. mixta ........................... ------------------------ X ______

Anacardiaceae, genus? .......... ..... -----------X-------------------

//fx carbonensis ... ............................... ------------- x -----------------

Celastrus comparabilis .. . .... ........ x-XX--XX______________XX_----__

Goweria alaskana .. ..... .. . .. ......... --X____________________________

Goweria dilleri ............................. ...... X______________________------__

Paleophytocrene elytraeformis ....... ---_---------__-_-_____X_______

Phytocrene acutissima ................... ... X-X___X---__-------------------

Phytocrene sordida .................... .......... --_-_____-------------X-X____._

Pyrenacantha sp .............................. -_--------------------X--------

Stemonurus alaskanus ..................... -- x ----------------------------

Rhamnus sp ............................... ..... -_--___-__-__ X -_-______-----__-

Sageretia sp ......................... _-____X_-_-__---_-_-_-_--------

Zizyphus alaskanus ............. __X--------_-------------------

Allophylus duktothensis ..................... ----------------------X___-____

Allophylus wilsoni .. ........ .. .. ....... ---------------- x X_X_____XX-__-

Allophylus sp .......................................--. ---------X__-_-__-___-____--.

Sapindus? sp .......................................... -----------X___-______---------

Meliosma duktothensis ........ ... ........ ----------------------X-X------

Meliosma kushtakensis ........................ --XX__X ________________________

Vt^'s sp ..................................................... ------ x ---------------X__-_---_

r/7m carbonensis .................................. -------------X------- ----------

Plafkeria sp ...... ..................................... -------------X_________.__.____

Saurauia sp ........... ............... ............ -------X. ______________________

Theaceae, genus? ............ ......... __---_-----X____-______. -------

Parashorea psuedogoldiana ............X_----------------------X_____.

Barringtonia sp ..................................... ----------------------X________

Kandelia wangi ......................... -____________-----------X------

Alangium bergensis ................. ......... x------------------------------

Alangium sp ......................................... __-_--X__________--. ___________

Eugenia sp ............................................. _______--____---------X--------

Eugenia? sp ............................................ ----------- _.X---------. -------

Mastixia irregularis .................... ....... -------X____.___---------------

Clethra sp .............................................. ----------------------X--------

Clerodendrum sp .................................... --X___. ._.___.___---_----------

Helmskioldia speiri? .............................. ----------------*------- -------

Macclintockia pugetensis .................... --------------X-XX-XXX---XXXXXX

Macclintockia sp .................................... ---------X_____-_ ___--_--------

Dicotylophyllum carbonensis . .......... -------------X-----------------

Dicotylophyllum kummerensis .......... ---------X----X----------------
12 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

represent the middle Ravenian. Assemblages from 

localities 3842, 9389, 11164, and 11165 come from 

about the same stratigraphic interval, a short 

distance below the base of the Tokun Formation. 

Critical elements in these assemblages are Allantodiopsis pugetensis, which has its latest occurrence 

in the middle Ravenian, and Dicotylophyllum 

kummerensis, which has its earliest occurrence in 

the middle Ravenian, and Calkinsia ravenensis, 

Carya cashmanensis, Pterocarya pugetensis, and 

Dryophyllum pugetensis, which indicate either a 

middle or late Ravenian age. The vegetational 

analysis (p. 32) indicates a vegetation type more like 

that of the middle than the upper Ravenian. The 

conclusion that this part of the Kushtaka is of middle 

Ravenian age is also supported by the close 

stratigraphic superposition of locality 11164 to a 

lower Ravenian locality (loc. 3846). The fact that 

several thousand feet of Tejon age rocks (lower part 

of the Tokun Formation) overlies the part of the 

Kushtaka containing the middle Ravenian assemblages indicates that the middle Ravenian is 

probably not of latest Eocene age.

UPPER RAVENIAN

The only Alaskan assemblage thought to be of late 

Ravenian age is that from locality 11167 in the 

Kushtaka Formation on Carbon Ridge. The placement of this locality in the Ravenian rather than in 

the overlying Kummerian is not totally satisfactory. 

The depauperate flora from this locality has only one 

species in common with the equally depauperate 

flora of the type section of the upper Ravenian. Two 

species, however, from locality 11167 are also known 

in upland floras in conterminous United States: 

Alnus cuprovallis and Prunus axelrodi. These 

species are known in floras such as the Republic and 

Copper Basin, whose radiometric ages are about 40 

million years (Wolfe and Obradovich, unpublished 

data), that is, probable latest Eocene. Pterocarya 

pugetensis has a known upper limit in the upper 

Ravenian of the Puget Group. The striking 

vegetational and floristic difference between the 

assemblage from locality 11167 and the overlying 

Kummerian assemblages from localities 9891 and 

11168, moreover, indicates that the assemblage from 

locality 11167 probably represents a pre-Kummerian 

stage.

KUMMERIAN STAGE

Macclintockia pugetensis was originally thought 

to be restricted to the upper part of the Kummerian 

Stage and was one of the primary reasons for 

distinguishing an upper from a lower Kummerian

substage. Additional collections, for example from 

the basal part of the Keasey Formation, indicate that 

this species ranges from the base of the Keasey 

through the Molopophorus stephensoni zone of the 

Lincoln "stage." The distinction, therefore, between 

a lower and upper Kummerian substage is of dubious 

value, and I will here not consider allocation of 

individual assemblages to substages within the 

Kummerian.

The rocks in which the Comstock flora of Oregon 

occurs were assigned by me (1968) to the upper 

Ravenian. This assignment probably was an error. 

The occurrence of Calkinsia dilleri and Allantodiopsis pugetensis was the only basis for placing the 

Comstock flora in the Ravenian. As now defined, the 

typical Ravenian species of Calkinsia is C. plafkeri, 

which is morphologically intermediate between the 

Fultonia C. pugetensis and C. dilleri. The specimen I 

determined as Allantodiopsis is sufficiently fragmentary to question the determination. The Comstock assemblage, therefore, could equally well be 

referred to the Kummerian, an age indicated by the 

interpretation of the C. dilleri lineage (see p. 63). The 

base of the Kummerian is thus considered to be 

approximately equivalent to the base of the Keasey 

"stage," and the top of the Kummerian is thought to 

be approximately equivalent to the boundary 

between the Molopophorus stephensoni and M. 

gabbi zones of the Lincoln "stage."

Kummerian assemblages are known from three 

rock units in the Gulf of Alaska region: the 

Kushtaka, Katalla, and Kulthieth Formations. In 

the Kushtaka, Kummerian assemblages are confined to the upper part of the formation exposed on 

Carbon Ridge. Locality 9891, which is 50 to 100 feet 

stratigraphic ally above the late Ravenian locality 

11168, contains Macclintockia pugetensis, Platanus 

comstocki (= Platanus n. sp. of Wolfe, 1968), and 

Dicotylophyllum kummerensis?; this assemblage is 

typical of the Kummerian. Presumably the assemblage from locality 11169, which is 300 feet 

stratigraphically higher than 9891, is also of 

Kummerian age, although, except for Tetracentron 

piperoides, species diagnostic of a Kummerian as 

opposed to an immediately post-Kummerian (Goshen) age are lacking.

The largest assemblage from the Katalla Formation was obtained from the basal member the Split 

Creek Sandstone Member at the base of Nichiwak 

Mountain. Nothing in the assemblage, which 

includes Macclintockia pugetensis, conflicts with 

the molluscan age assignment to the Molopophorus 

stephensoni zone of the Lincoln "stage." The other
BIOSTRATIGRAPHY 13

plant assemblages from the Split Creek Sandstone 

Member have also yielded Macclintockiapugetensis. 

The two small collections from an unknown part of 

the overlying Basin Creek Member also contain 

Macclintockia pugetensis, thus indicating a Kummerian age. As noted previously, MacNeil considers 

the Basin Creek to range from medial Lincoln to 

early Blakeley in age, whereas Yokes considered his 

material from the Basin Creek to be little, if any, 

younger than the Split Creek. Because Macclintockia is elsewhere not known in beds of medial or 

possibly later Lincoln age, I am inclined to agree 

with Yokes that the Basin Creek Member or at least 

that part that has yielded the plants is probably no 

younger than the Molopophorus gabbi zone and 

possibly no younger than the M, stephensoni zone. 

Several localities in the upper part of the Kulthieth 

Formation have yielded small collections of plants. 

Reference to the checklist (table 1) shows that 

Macclintockia pugetensis occurs in most of those 

collections, and hence the upper part of the Kulthieth 

is referred to the Kummerian. This age accords with 

that of the marine megafossils collected from the 

upper Kulthieth.

POST-KUMMERIAN ROCKS

No Paleogene floras of post-Kummerian age from 

the Gulf of Alaska section are yet known in 

association with marine mollusks. Locality 11185 

from rocks near Marvine Glacier in the Malaspina 

district has furnished a small assemblage that is 

clearly distinct from the floras previously discussed. 

The three species present in the Marvine Glacier 

assemblage are: Metasequoiacf. M. glyptostroboides 

Hu and Cheng, Alnus evidens (Holl.) Wolfe, and 

Cercidiphyllum crenatum (Ung,) R. W. Br. Although 

the rocks from which this assemblage has been 

obtained have been assigned to the Kulthieth 

Formation (Stonely, 1967), they could equally well be 

the nonmarine equivalent of the Poul Creek 

Formation (George Plafker, oral commun., October 

1968). None of the species and only one genus (Alnus) 

are known in the Alaskan Ravenian and Kummerian floras. The species of Alnus is particularly 

significant because it is characteristic of the preSeldovian (lowermost) part of the Kenai Group; this 

part of the Kenai is here (p. 13) termed Angoonian. 

The only locality where this species is known in a 

definite relationship to the marine geochronology is 

on Sitkinak Island; there A. evidens is known from a 

locality in nonmarine beds that conformably 

underlie marine beds of late Blakeley (early Miocene) 

age (Wolfe and others, 1966, p. A17; Moore, 1969). A 

reasonable assumption is that the plant-bearing

beds on Sitkinak Island and at Marvine Glacier are 

probably of early Blakeley (late Oligocene) age. This 

age assignment indicates that the beds at Marvine 

Glacier are nonmarine equivalents of some part of 

the Poul Creek Formation.

Three localities from the Gulf of Alaska section 

have yielded leaves of Neogene age. Locality 11183 

occurs in nonmarine beds that, on structural and 

lithologic evidence, may be equivalent to the 

Yakataga Formation (George Plafker, oral commun., October 1968). The Yakataga Formation 

ranges in age from early middle Miocene through 

early Pleistocene (Plafker, 1967). The leaves represent Osmunda sp., Glyptostrobus sp., Metasequoia 

cf. M. glyptostroboides Hu and Cheng, Alnus cappsi 

(Holl.) Wolfe, and Acer sp. The species of Alnus is 

characteristic of the Seldovian Stage of the Kenai 

Group; the species is also known from the Unga 

Conglomerate, which contains mollusks of middle 

Miocene age (Burk, 1965, p. 213). The leaves from 

locality 11183 indicate, therefore, that the enclosing 

beds are no younger than middle Miocene and hence 

are probably from the lower part of the Yakataga 

Formation.

Locality 11184 from the Yakataga Formation in 

the Pinnacle Hills is close stratigraphically to 

marine mollusks of late Miocene age (F. S. McCoy, 

oral commun., April 1968). The one species present is 

Carpinus cobbi Wolfe, a species characteristic of the 

Homerian Stage of the Kenai Group (Wolfe and 

others, 1966).

Locality 11186 occurs in marine rocks in Lituya 

Bay. The age of the mollusks in the same part of the 

section is considered to be late (but not latest) 

Miocene (F. S. McCoy, oral commun., April 1968). 

The single leaf represents a species of Fagus; this 

genus is not known to be present in the Homerian 

and is thought to have become extinct in Alaska 

by the end of the Seldovian Stage. The genus is 

represented, however, in pollen floras from late 

Miocene rocks on the Queen Charlotte Islands 

(Martin and Rouse, 1966). The genus, therefore, may 

have persisted somewhat longer in southeastern 

Alaska than in other areas of the State, or the 

Seldovian Stage may include some rocks that in the 

marine megafossil geochronology are of early late 

Miocene age.

ANGOONIAN STAGE

Fundamental to understanding Alaskan Tertiary 

stratigraphy is a knowledge of the type of 

assemblage represented in the Tsadaka Formation 

and the lowest part of the Kenai Formation in the
14 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Cook Inlet region. This type of assemblage is 

widespread in Alaska; for example, such an 

assemblage is known from St. Lawrence Island, 

various areas of the Alaska Range, the Copper River 

basin, the Matanuska Valley, the Cook Inlet Basin, 

the Alaska Peninsula, the Trinity Islands, the 

Malaspina district, Admiralty Island, and Kuiu 

Island. This type of assemblage is typically 

composed of Alnus evidens and Metasequoia, 

although other species, for example, Cercidiphyllum 

crenatum, are represented at many localities. This 

assemblage was at one time considered to be 

"lower(?) Seldovian" (Wolfe and others, 1966), but it 

has since been excluded from the Seldovian Stage 

(Wahrhaftig and others, 1969).

The only known Alaskan section in which the 

Tsadaka type of assemblage is represented and in 

which recognizably older and younger assemblages 

are also represented is in the Kootznahoo Formation 

on Admiralty Island near the town of Angoon. 

Geologic mapping (Lathram and others, 1965) and 

my own observations made during 10 days of 

field work in 1961 indicate that the section is 

continuous, although some repetition is present 

because of faults of small displacement. This section, 

therefore, is desirable as a type section for a stage 

characterized by the Tsadaka type of assemblage.

DEFINITION

The Angoonian Stage is here designated a 

provincial stage for plant-bearing rocks the same as 

those in the type area in the Kootznahoo Formation. 

The type section is designated as that part of the 

Kootznahoo Formation exposed from the north 

shore of Long Island (see figs. 4 and 5) south (and 

stratigraphically upward) to the rocks exposed on 

the south side of the small island near the west end of 

Kanalku Bay. Thus the rocks at localities 9826 and 

9829 are definitely excluded from the Angoonian.

FLORA

The assemblage in the type section of the 

Angoonian has not been completely studied; some 

items remain indetermined. The species thus far 

determined include: 

Gymnospermae 

Ginkgoales 

Ginkgoaceae 

Ginkgo sp. 

Coniferales 

Taxodiaceae

Metasequoia cf. M. glyptostroboides Hu and 

Cheng

Angiospermae 

Dicotyledonae 

Juglandales 

Juglandaceae 

Juglans magnifica Knowl. 

Pterocarya orientalis (MacG.) Wolfe 

Fagales 

Betulaceae

Alnus evidens (Holl.) Wolfe 

Alnus carpinoides Lesq. 

Fagaceae

Fagus sp. 

Resales

Hamamelidaceae 

Fothergilla sp. 

Sapindales 

Aceraceae

Acer aff. A. glabroides R. W. Br. 

Rhamnales 

Vitaceae

Vitis atwoodi Holl. 

Vitis rotundifolia Newb. 

Malvales 

Tiliaceae

Tilia aspera (Newb.) LaM. 

Plafkeria obliquifolia (Chan.) Wolfe 

Sterculiaceae

Firmiana alaskana (Newb.) Wolfe 

The Angoonian assemblage is floristically and 

physiognomically temperate. None of the species 

and only a few of the genera are known in 

Kummerian assemblages in Alaska or in the Pacific 

Northwest. Most Angoonian species are related to 

extant species that live in temperate climates. The 

percentage of species that have entire-margined 

leaves is 12.

AGE

Only one Alaskan Angoonian assemblage is 

known that is definitely stratigraphically related to 

marine rocks. On Sitkinak Island of the Trinity 

Islands, Alnus evidens, Cercidiphyllum crenatum, 

and Metasequoia occur in nonmarine beds a few 

feet below a conformable contact with overlying 

marine beds; the marine beds are considered to be 

of late Blakeley (earliest Miocene) age. Presumably the plants could be as young as earliest 

Miocene, but I assume that a late Oligocene age is 

more probable. In conterminous United States, 

Angoonian assemblages include the Bridge Creek 

and Willamette floras. Both have radiometric ages of 

about 31 million years (Evernden and James, 1964); 

in the mammalian chronology this is presumably 

very early in the middle Oligocene. The Willamette
BIOSTRATIGRAPHY 15

 57°30'

3 MILES

J

4 KILOMETERS

FIGURE 4. Generalized geologic map of part of Kootznahoo Inlet. Data from Lathram and others (1965). 

Tks (stippled) is the sandstone and shale part of the Kootznahoo Formation; Tkc (lined) is the conglomeratic 

part; Pz, undifferentiated Paleozoic rocks. Heavy lines represent faults. Crosses represent plant localities.

flora, however, comes from beds that overlie with an 

angular unconformity nonmarine beds that have 

yielded the Goshen flora; in turn, the Goshen beds 

conformably overlie marine beds containing mollusks of the Molopophorus gabbi zone (middle 

Lincoln) (Vokes and others, 1951). These data 

indicate that probably the late Oligocene of the 

marine megafossil workers is equivalent, at least in 

part, to the middle Oligocene of the vertebrate 

workers. The Angoonian is considered to be of late 

Oligocene age.

The assemblage from locality 9829, which is 

stratigraphically below the type section of the

Angoonian, includes: Cercidiphyllum crenatum, 

Cinnamornophyllum eocernua, Platanus macginitiei, Fagus oregona, and Plafkeria thornae. The 

Platanus is the same elaborately toothed species 

known in the Goshen, Sweet Home, and Scio floras, 

all of which are of middle Oligocene (middle and 

upper Lincoln) age. Neither the Cinnamomophyllum 

nor the Fagus are known in conterminous United 

States in beds younger than middle Oligocene, 

whereas the Cercidiphyllum and Plafkeria make 

their first appearance in beds of middle Oligocene 

age. The assemblage from locality 9826, which is 

stratigraphically above the type section of the
16 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

FEET METERS 

l 1000

3000

2000

1000 

Plant 

locality

9826

9828

9825

Stage Series

9824

(9822 

 < 9823 

(9827 

9829

Seldovian

Angoonian

Unnamed

(Goshen

equivalent)

stage

03

C

<0

u

o

I

HI

o

0

ra

6

Coal

EXPLANATION

Sandstone and shale Conglomerate

FIGURE 5. Generalized stratigraphic section of the type area of 

the Angoonian Stage.

Angoonian, includes Alnus barnesi, Fagus antipofi, 

Quercus furuhjelmi. This assemblage is thus of 

Seldovian age, which is considered to early and 

middle Miocene.

The Angoonian Stage is separated from the older 

Kummerian by an unnamed stage containing a 

distinctive assemblage. In Washington, for example, 

assemblages similar to the Lower Kootznahoo and 

the Goshen occur in beds demonstrably younger 

than the Kummerian (Wolfe, 1961, 1968). As yet I 

know of no section in which assemblages of the 

Goshen and Lower Kootznahoo type occur above 

Kummerian and below Angoonian assemblages, 

and thus no stage is here proposed for the Goshen 

equivalents.

CORRELATION OF SOME OTHER ALASKAN FLORAS

Chaney (1967, p. 228) has recently expressed the 

opinion that the lower floras the Seldovian and the 

Tsadaka equivalents, here (p. 13) termed Angoonian of the Kenai Group are of early Oligocene age. 

His opinion is based on a proposed correlation of the 

Kenai floras to floras purported to be of early 

Oligocene age from the Port Moller region of the 

Alaska Peninsula. Chaney further considers these 

Port Moller assemblages to be "The nearest

occurrence of well-dated marine sediments with 

associated plant-bearing beds ***" to the Cook Inlet 

basin. The Port Moller assemblages are, of course, 

over 300 miles farther from the Cook Inlet basin than 

are the assemblages from the Katalla district (see fig. 

6). Some of the Port Moller assemblages discussed by 

Chaney, moreover, had been previously shown 

(Wolfe in Burk, 1965, p. 233-236) to be mostly of 

Paleocene age, which conforms with evidence from 

the middle Eocene Foraminifera from the overlying 

beds (Burk, 1965, p. 113). Chaney also included in his 

correlation plant assemblages that, on marine 

molluscan evidence, are of middle Miocene age and 

some that are of late Miocene or early Pliocene age 

(Burk, 1965, p. 116-117). The stratigraphic basis for 

Chaney's proposed correlations is, therefore, invalid. 

The paleobotanical basis for Chaney's (1967) 

correlations, if valid, would be of considerable 

significance, therefore, I will examine the evidence 

he cites in detail. Chaney (1967, p. 228) lists 12 genera 

that he thinks are common to the Seldovian and to 

his purported early Oligocene assemblages from the 

Alaska Peninsula. Chaney also states that many of 

the Seldovian species (although he does not list 

them) have been recorded by Hollick (1936, p. 18-19) 

from Chaney's purported early Oligocene of the 

Alaska Peninsula. The following are the genera 

listed by Chaney:

Acer. Hollick (1936) listed two species of Acer in. 

three assemblages from the Port Moller region. One 

of these two species, Acer arcticum Heer, is based on 

specimens (Hollick, 1936, pi. 78, figs. 7, 8) that have 

the large rounded teeth and venation pattern of 

Dicotylophyllum richardsoni(Heer) Wolfe. The other 

species listed by Hollick is Acer trilobatum 

production (A. Br.) Heer. Hollick, however, was 

following Knowlton (1894) in this listing, and I 

concur with Hollick (1936, p. 135) that "*** the 

identification does not appear to be altogether 

conclusive***." Note that none of the six species of 

Acer listed in the Seldovian (Wolfe and others, 1966, 

p. A16) was listed in the Port Moller assemblages by 

Hollick. Also significant is the fact that the Port 

Moller localities that produced the purported species 

of Acer are now considered to be of Paleocene or early 

Eocene age (Burk, 1965).

Alnus. Hollick (1936) listed a single species of 

Alnus as occurring in the Port Moller assemblages. 

This material of this species, A. kefersteini (Goepp.) 

Heer, has the numerous craspedodrome marginal 

tertiary veins extending from the basal secondary 

veins and the rounded to sharp teeth separated by 

arcuate sinuses of Grewiopsis ariculaecordatus
BIOSTRATIGRAPHY 17

140° 130° 70° 120°

I______i .. I ____ I

T I I I I I

100 200 300 400 500 600 KILOMETERS

FIGURE 6. Map of Alaska showing location of some fossil plant assemblages.

(Holl.) Wolfe. Apparently Chaney (1967, p. 225, 

footnote) thinks that, because I (1966) transferred 

certain of Rollick's material described as Corylus to 

Alnus, all of Hollick's material of Corylus was so 

transferred. However, I specifically stated in the 

synonymies that only part of Hollick's material was 

to be considered as Alnus. Not one of the six 

Seldovian species of Alnus has ever been listed in the 

Port Moller assemblages, except for A. barnesi, 

which is thought to occur in the mollusk-dated 

middle Miocene beds on Unga Island (Burk, 1965, p. 

236).

Betula. Hollick listed two species of Betula from 

the Port Moller assemblages, neither of which was 

listed (Wolfe and others, 1966, p. A16) in the 

Seldovian. I am unsure of the familial relationship of 

the specimens on which the citations are based; at 

least one of the specimens could be betulaceous, but it 

is not the Seldovian species B. sublutea Tan. and 

Suz.

Cercidiphyllum. Hollick did not list any Cercidiphyllum from the Port Moller assemblages.

Presumably Chaney is accepting Brown's transfer of 

certain of Hollick's material listed as Populus arctica 

Heer, P. amblyrnhynca Ward, and P. richardsoni 

Heer to Cercidiphyllum arcticum (Heer) R. W. Br. 

However, these specimens have since been transferred to Cocculus flabella (Newb.) Wolfe and 

Dicotylophyllum richardsoni (Wolfe, 1966, p. B9). 

None of Hollick's material from the Port Moller 

region represents Cercidiphyllum, and none of it has 

ever been referred to the Seldovian species C. 

crenatum (Ung.) R. W. Br.

Fraxinus. A single lamina from the Port Moller 

region was described as F. herendeenensis by 

Knowlton (1894), a species that has never been 

recorded from a Seldovian assemblage. The specimen has tertiary veins that are closely spaced, 

unbranching, oriented at right angles to the 

secondary veins, and of uniform strength throughout their course; in Fraxinus, the tertiary veins are 

typically widely spaced, branching, oriented obliquely to the secondaries, and conspicuously thin. 

The Port Moller specimen, therefore, cannot repre-
18 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

sent Fraxinus. The specimen has the same intercostal tertiary venation pattern and sharp (almost 

spinose) teeth as in Dicotylophyllum flexuosa 

(Newb.) Wolfe, and it probably represents that 

species.

Ginkgo. The same species of Ginkgo apparently 

occurs in both the Port Moller and Seldovian 

assemblages; it also occurs in Paleocene assemblages in the Cook Inlet Region (Wolfe, 1966, p. B2). 

Ginkgo is of almost no stratigraphic value for 

Tertiary correlations, considering that apparently 

the same species ranges throughout the Tertiary 

(Tralau, 1968).

Glyptostrobus. The material of Glyptostrobus 

from the Port Moller region listed by Hollick is that 

from Chignik Bay, which I (Wolfe and others, 1966, 

p. A9) consider to be G. nordenskioldi (Heer) R. W. 

Br.; this species has also been listed (Wolfe and 

others, 1966, p. A9) from the Paleocene of the Cook 

Inlet region. The Seldovian Glyptostrobus is 

considered to represent G. europaeus (Brong.) Heer 

(Wolfe, 1966, p. B3). Again, the Port Moller and 

Seldovian species are distinct and represent a genus 

that ranges throughout the Cenozoic.

Metasequoia. The material from the Port Moller 

assemblages listed by Hollick as Sequoia langsdorfii 

(Brong.) Heer was transferred by Chaney (1951) to 

Metasequoia occidentalis (Newb.) Chaney. Note, 

however, that, although the Port Moller material is 

accepted as that species (Wolfe and others, 1966, p. 

A9), the Seldovian material is thought to represent 

M. glyptostroboides Hu and Cheng (Wolfe, 1966, p. 

B3).

Nyssa. The only Nyssa from the Port Moller 

region is based on the unfigured fructifications 

referred by Lesquereux (1883) to N. arctica Heer from 

middle Miocene marine beds on Unga Island. This 

material was later transferred by Brown (1939) to 

Cercidiphyllum arcticum. Similar, if not conspecific, 

fructifications have also been collected at one of the 

localities near Port Graham; the taxonomic status of 

this type of fructification is, however, in need of 

considerable clarification (Chandler, 1961,1964). In 

any case, there is no basis for the occurrence of 

Nyssa in the Port Moller assemblages.

Populus. All species of Populus listed by Hollick 

from the Port Moller region have been transferred to 

Cercidiphyllum arcticum by Brown (1939) and 

thence to other genera by me (1966). There is no 

taxonomic basis for Chaney's listing of Populus in 

the Port Moller assemblages.

Pterocarya. No species of Pterocarya was listed 

by Hollick from the Port Moller assemblages.

Chaney is apparently assuming that because I 

transferred Heer's Port Graham species Juglans 

nigella to Pterocarya (a species never recorded in the 

Port Moller assemblages), all the material listed as 

Juglans by either Hollick or Heer should also be so 

transferred. This assumption is clearly unfounded. 

The other species of Juglans that Heer described 

from Port Graham was transferred to Salix (Wolfe, 

1966, p. B14). Note also that some material attributed 

to both Juglans and Pterocarya by Hollick (1936) 

was transferred to Dicotylophyllum flexuosa (Newb.) 

Wolfe (Wolfe, 1966, p. Bll). There is no taxonomic 

basis for the inclusion of Pterocarya in the Port 

Moller assemblages of Hollick.

Salix. This genus was recorded from two localities by Hollick (1936) from the Port Moller region. 

Salix minuta Knowlt. from locality 539 is based on a 

single crudely illustrated specimen; the preservation 

of the specimen does not warrant generic assignment. The only valid determination of Salix made by 

Hollick (1936) from the Port Moller assemblages is 

based on the material from locality 5182. This 

material represents two Homerian species, S. 

alaskana Holl. and S. kachemakensis Wolfe, and, as 

has been previously pointed out (Burk, 1965, p. 235; 

Wolfe and others, 1966, p. A20), this material is from 

beds that are of Miocene or Pliocene age based on 

physical stratigraphy and marine mollusks.

Of the 12 genera considered to be common to the 

Seldovian and Port Moller assemblages (Chaney, 

1967), four either have never been recorded from the 

Port Moller region Pterocarya or are based on 

material that has been formally transferred to other 

genera Populus, Cercidiphyllum, and Nyssa. The 

citations of three other genera Acer, Alnus, and 

Fraxinus are either based on fragmentary material 

or can be demonstrated to belong to other genera; 

another genus Betula may be represented in the 

Port Moller assemblages, but clearly not by the 

Seldovian species. At least 3 of the 52 Seldovian 

genera are represented in Port Moller assemblages, 

but in two instances by different and distinct species 

and in the third instance by a stratigraphically 

insignificant Ginkgo. The twelfth genus Salix is 

represented in the Port Moller assemblages but at 

one locality of probable Miocene or Pliocene age. Of 

Chaney's "many" species in common to both the 

Seldovian and the Paleogene Port Moller assemblages, only one the Ginkgo can be demonstrated 

to be valid. The paleobotanical and stratigraphic 

basis for Chaney's correlation of the Kenai floras is, 

therefore, invalid. 

Axelrod (1966a, 1966b, 1968) has discussed his
BIOSTRATIGRAPHY 19

interpretations of the altitudinal factor in interpreting ages of Tertiary floras. His conclusion, which is 

similar to that reached by Chancy (1936, 1940), is 

that in the Eocene, altitude had a pronounced effect 

on floristic and vegetational zonation. Thus Axelrod 

considers the vegetation during the late Eocene of 

coastal southern Alaska to be temperate; this 

temperate forest type ascended altitudinally southward, so that in Nevada it was at an approximate 

altitude of 3,500 feet. Axelrod further considers the 

lowland and more coastal late Eocene vegetation of 

the Pacific Northwest to be warm temperate (1966b, 

p. 42) or subtropical (1966b, p. 25). The Alaskan flora 

that Axelrod considers to be isochronous to the 

temperate late Eocene Copper Basin assemblage 

from Nevada is the Port Graham; the Port Graham 

assemblage, however, is of early Seldovian (early 

Miocene) age (MacGinitie, 1962, p. 88; Wolfe and 

others, 1966; p. 20 of this report). As will be shown 

later, the marginally tropical lowland assemblages of late middle Eocene age from the Pacific 

Northwest represent the same fundamental vegetational type as do isochronous assemblages in 

Alaska. I conclude, therefore, that Axelrod's 

interpretations are fundamentally invalid; as will 

be discussed later, during the Eocene there were 

major climatic fluctuations, and comparisons of two 

assemblages one from a cool interval (Copper 

Basin) and a second from a warm interval (Steel's 

Crossing) can be highly misleading in regard to 

both latitudinal and altitudinal zonation of ancient 

vegetation.

Altitude can, of course, be a factor in the 

interpretation of age of a given assemblage of fossil 

plants, particularly if assemblages are assigned to 

an age largely or totally on their vegetational aspect 

rather than on their specific composition. Vegetationally an upland assemblage of latest Eocene age 

can be similar to a lowland assemblage of Pliocene 

age, that is, both represent conifer or broad-leaved 

deciduous forest. The specific composition of these 

two assemblages if the fossils are examined in 

detail will, however, be totally different (or nearly 

so). Floristic comparisons, on the other hand, 

between two assemblages of the same age in the 

same region typically indicate that some shortranging species are common to the assemblages 

irrespective of altitude. This phenomenon stems in 

part from the fact that some species participate in 

two or more vegetational types; this is particularly 

true of fluviatile and lacustrine species that 

dominate fossil assemblages. During the late Miocene, for example, broad-leaved deciduous forest

occupied the lowlands of the Pacific Northwest 

while conifer forest occupied the uplands (Wolfe, 

1969). Despite the strong physiognomic difference 

between these two vegetational types, species such 

as Salix hesperia (Knowlt.) Cond., Quercus deflexiloba H. V. Sm., and Platanus dissecta Lesq., are 

known in both lowland and upland assemblages. 

The occurrence of these species together in a fossil 

assemblage no matter what the altitude indicates 

a late Miocene age.

The Angoonian and Seldovian assemblages are 

not similar to any assemblages from definite 

Kulthieth, Kushtaka, or Katalla Formations. Families well represented in, and that dominate, the 

Kenai and Tsadaka assemblages are either poorly 

represented in the Gulf of Alaska early and middle 

Ravenian and Kummerian floras for example, 

Betulaceae or are not represented by any taxa for 

example, Salicaceae, Rosaceae, and Aceraceae. 

Conversely, the dominant families of the early and 

middle Ravenian and Kummerian floras are either 

sparsely represented for example, Menispermaceae or are not represented for example, Lauraceae and Icacinaceae in the Kenai and Tsadaka 

floras. The species of the late Ravenian are distinct 

from those of the Angoonian and Seldovian. There 

can be no reasonable doubt that the Kenai and 

Tsadaka floras are younger than the Kummerian 

floras. Both depositional areas were at low altitudes, 

and the fact that the Gulf of Alaska and Cook Inlet 

basins are at the same latitude (in part, the Kenai 

localities are south of the Gulf of Alaska sequence) 

indicates that the differences between the Kenai and 

Ravenian-Kummerian floras are due to a difference 

in age.

An apparent disagreement exists over the age of 

fossil plants collected from the Tolstoi Formation of 

Burk (1965) at Coal Bay on the Alaska Peninsula. I 

considered the plants to be of late Oligocene or early 

to middle Miocene, that is, of Seldovian age as this 

term was used by Wolfe, Hopkins, and Leopold (1966, 

p. 1,17). The marine molluscan fauna is probably of 

early Blakeley (late Oligocene) or possibly of latest 

Lincoln (latest middle Oligocene) age (MacNeil in 

Burk, 1965, p. 222). Roth well, however, considered 

the Foraminifera to be mostly of earliest Oligocene 

age (in Burk, 1965, p. 230). It should be pointed out, 

however, that Roth well's usage of early Oligocene is 

such that he considers part of the Astoria Formation 

to be of late Oligocene age (this is considered by 

molluscan workers to be of middle Miocene age; see 

Moore, 1963) and the lower part of the Blakeley 

"stage" and the Lincoln "stage" to be of early
20 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Oligocene age (W. T. Roth well, oral commun., March 

27, 1964). The discrepancy, if such exists, between 

the Foraminifera, on the one hand, and the plants 

and mollusks, on the other, is not significant. In this 

report I am accepting the geochronology of the 

marine mollusks in regard to the placement of 

epochal boundaries in the western North American 

Tertiary, and thus the Coal Bay plant assemblages 

are considered to be most probably of late Oligocene 

(Angoonian) age. The Coal Bay assemblages include 

Salix, Fagus, Cocculus, and Acer; these assemblages 

probably represent broad-leaved deciduous forest. 

As elsewhere in Alaska, broad-leaved deciduous 

forests of Oligocene age were present only in the later 

part of that epoch.

A flora of the same composition as the Angoonian 

and Seldovian floras does indeed occur in the Gulf of 

Alaska sequence; the Angoonian assemblage occurs 

in rocks of questionable stratigraphic position that 

could, on lithologic and structural grounds, be 

considered the nonmarine equivalent of the Poul 

Creek Formation. This interpretation would indicate 

a latest Lincoln or Blakeley age for the Angoonian 

assemblage. As noted previously (p. 14) a Blakely 

age is indicated for another Angoonian assemblage 

on Sitkinak Island. The available evidence indicates 

that the Angoonian assemblages are of early 

Blakeley (late Oligocene) age.

The Seldovian assemblages are younger than the 

Angoonian assemblages. This relationship can be 

demonstrated both in the Alaska Range section 

(Wahrhafting and others, 1969) and the Kootznahoo 

section. If the Angoonian assemblage is of late 

Oligocene age, as the evidence thus far indicates, 

then the Seldovian floras are probably early Miocene 

at the oldest. Only two small assemblages of 

Seldovian type occur in marine sections in which 

ages independent of the plants can be obtained. One 

is the sequence on Unga Island off the Alaska 

Peninsula. The Unga Conglomerate Member of the 

Bear Lake Formation of Burk (1965) contains marine 

mollusks thought to be of middle Miocene age. The 

plants from this member include Alnus barnesi 

Wolfe, a characteristic Seldovian species. The 

microfossil flora contains Carya, Pterocarya, Fagus, 

Quercus, Ulmus, Liquidambar, and Tilia; this 

assemblage is similar to those reported from the type 

section of the Seldovian (Wolfe and others, 1966). The 

plant megafossil assemblage from possible Yakataga Formation (locality 11183) includes Alnus 

cappsi (Holl.) Wolfe, a species that is known in 

Alaska only from the Seldovian. The increasing 

body of evidence from marine fossils substantiates 

the conclusion of Wolfe, Hopkins, and Leopold (1966,

p. A17) that the Seldovian Stage is of early and 

middle Miocene age and clearly is not consistent 

with Chaney's opinion that the Seldovian Stage is 

of early Oligocene age or Axelrod's opinion that the 

Seldovian assemblage from Port Graham is of 

Eocene age.

The provisional age assignment of the Homerian 

Stage to the upper Miocene was based both on 

specific composition and the small assemblage from 

Herendeen Bay on the Alaska Peninsula. The 

Herendeen Bay assemblage contains characteristic 

Homerian species of Salix and is closely related 

stratigraphically to mollusks of late Miocene or early 

Pliocene age (Burk, 1965). A pollen assemblage 

similar to that of the type section of the Homerian 

has been obtained from rocks in the Bering Sea that 

contain marine diatoms referable to the later half of 

the Miocene (Hopkins and others, 1969). A small 

collection of leaves from the Yakataga Formation 

exposed in the Pinnacle Hills contains the characteristic Homerian Carpinus cobbi Wolfe, and the 

marine megafossils from the same section indicate a 

late Miocene age (F. S. McCoy, oral commun., April 

1968). Again, the increasing body of independent 

evidence indicates a probable late Miocene age for 

the Homerian and contradicts Chaney's (1967) 

opinion that the Homerian is of middle Oligocene 

age. The general epochal assignments of the 

Seldovian, Homerian, and Clamgulchian types of 

assemblages has received considerable confirmation 

from the work of Fotianova (1964, 1967), Sinelnikova, Skiba, and Fotianova (1967), and Chelebaeva (1968), who have noted a similar sequence on 

Sakhalin and Kamchatka.

Although some revisions based on further collecting and study will undoubtedly be made, I do not 

think that the tentative correlations presented in 

figures 7 and 8 will change radically with new 

information. Critics of the age assignments presented here, I think, are obligated to present both 

detailed alternative systematic treatment of the 

plant megafossils and detailed knowledge of the 

physical stratigraphy of the plant-bearing beds.

FLORISTIC AND VEGETATIONAL ANALYSIS

FLORISTIC ANALYSIS

Interpretations of floristic history, as Mason 

(1947) has pointed out, must involve "***piecing 

together genetic lineages." In the Neogene of 

western North America, this piecing together can be 

done with confidence because of our considerable 

knowledge of Neogene assemblages. The lineage, for 

example, that has produced the modern Alnus
FLORISTIC AND VEGETATIONAL ANALYSIS 21

SERIES

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SUBSERIES

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LL

CARBON MOUNTAIN

^_,--^_--^_,^--^ - 

77769 

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Kushtaka

Formation

11166 

11159 

11158 

77757 

3847 

Stillwater

Formation

Not exposed

CUNNINGHAM 

RIDGE

Tok jn

Formation

77764 

77765 

3846 

11162 

11160 

Kushtaka

Formation

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Stillwater

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CHARLOTTE 

RIDGE

Tok uri

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Kushtaka 

77765 

9389 

Formation

Not exposed

NICHIWAK 

MOUNTAIN

Katalla 

Formation

7776S 

Not exposed

MILLER HILLS

Katalla

Formation 

11188 

555? A

3552 

Tokun

Formation

Not exoosed

ROBINSON 

MOUNTAINS

Poul Creek 

Formation

9SS3~}

9893 \^

9894 [^

989SJ

Kulthieth 

Formation

11170 

-^ " "

Unnamed

FIGURE 7. Correlation of Paleogene formations in the Katalla and Yakataga districts. Numbers represent plant localities.

incana can be traced from a rich broad-leaved 

deciduous, forest during the early and middle 

Miocene, into a rich conifer forest during the late 

Miocene, and finally into a depauperate boreal 

conifer forest during the Pliocene (Wolfe, 1966). 

Similarly an extinct lineage of Lyonothamnus was, 

during the early and middle Miocene, a member of a 

warm temperate, summer-wet, broad-leaved deciduous forest in the Pacific Northwest, but by the late 

Miocene the lineage was a member of a conifersclerophyllous broad-leaved forest that lived under a 

summer-dry climate in Nevada (Wolfe, 1964).

Little paleobotanical work has been done on 

Paleogene particularly Eocene assemblages in 

western North America, therefore, the history of 

various lineages is obscure. Although the genera and 

species can be, for the most part, discussed in terms 

of the most closely related living genera and species, 

the detailed history of Paleogene lineages is to a 

great extent speculative. Despite these qualifications, however, the general floristic relationships of 

some Paleogene assemblages to isochronous assemblages elsewhere and to modern floristic regions 

can be discerned.

VEGETATIONAL ANALYSIS

INTRODUCTION

The reliance by many Tertiary paleobotanists on 

the floristic relationships of fossil assemblages in 

arriving at an understanding of past climates and 

vegetational types has led to considerable confusion, 

both in floristic and vegetational history. This 

approach fundamentally relies upon an extremely 

strict application of uniformitarianism to tolerances 

of families, genera, and species. Many workers 

consider that tolerances of members of a particular 

taxonomic category have not changed through time. 

It seems clear, however, that tolerances must have 

changed in many, if not most, groups during time. 

For example, the tolerances of extant members of 

Platycarya and Nipa&re mutually exclusive, and yet 

both genera are abundant in the early Eocene 

London Clay (Reid and Chandler, 1933); Platycarya 

has, on Taiwan and mainland China, the opportunity to extend its range downslope in;o the 

Paratropical Rain forest but has not. Similarly Nipa 

has the opportunity to extend its range northward 

into lowland Taiwan but has not. The conclusion
22 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

SERIES

SUBSERIES

STAGE

MARINE 

MEGAFOSSIL

NENANA 

COAL FIELD PORT MOLLER COOK INLET KATALLA 

DISTRICT

YAKATAGA

AND

MALASPINA 

DISTRICTS

ANGOON

if

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Nenana 

Gravel

HJLJJ-LLU

! 

I

Grubstake 

Formation

Vaqueros

Lignite Creek Fm

Suntrana Fm. x

'/ 'Bear Lake : ;:

 '.- ' Formation-.' '

 of Burk (1965) - Kenai 

Formation

Sanctuary Fm.

Healy Creek 

Formation

£ ?

Yakataga 

Formation

Yakataga 

Formation

':' .' -'S tep ovak .';.': ( 

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 '.' Form

feet .; /.'; 

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Kootznahoo 

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/. Kushtaka 

;/Formation

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VvY/Formation

FIGURE 8. Correlation of some Alaskan Tertiary formations. An (x) denotes approximate stratigraphic position of plant assemblage(s);

stippling denotes rocks that are largely or entirely marine.

appears inescapable that the present tolerances of 

the two genera are mutually exclusive. Some workers 

(Axelrod, 1966b) have suggested that such mixtures 

in fossil assemblages are due mostly, if not entirely, 

to the early Tertiary climates and middle latitudes 

having been "warm temperate," that is, an 

essentially frostless climate that had a high 

equability but a low mean annual temperature. Thus 

Takhtajan (1957) and Axelrod (1966b) have suggested that the distribution of Nipa and certain 

mangroves is controlled mostly by marine, not land, 

temperatures, and their presence in the "warm 

temperate" London Clay flora is a function of marine 

temperatures. This hypothesis seems unreasonable 

in that in equable marine areas today Nipa and 

Platycarya are not associated and no modern 

thanatocoenose is known to contain specimens of 

both genera.

Theoretically it is highly probable that the 

tolerances of lineages have changed through time: 

how else could members of a subclass such as 

Dicotyledones have diversified and occupied widely 

different habitats? If, as is generally thought, the 

more primitive members of various families are 

tropical in distribution, then most, if not all, 

"temperate" genera must be descended from 

lineages that were once tropical. The use of floristic 

relationships to determine vegetational and climatic 

relationships of fossil assemblages has an inherent 

tendency to obscure patterns of changing tolerances 

of given lineages; the fundamental assumption of 

this method is that tolerances have changed little if 

at all. A given association is in a constant state of 

flux when viewed in time; different populations react 

differently to environmental change. If what is 

desired is an analysis of the history of a vegetational
FLORISTIC AND VEGETATIONAL ANALYSIS 23

type such as the Mixed Mesophytic forest or the 

Paratropical Rain forest, then methods largely 

independent of floristic composition must be used to 

identify a fossil assemblage as a particular 

vegetational type (Wolf \ 1971).

FOLIAR PHYSIOGNOMY AND VEGETATIONAL ANALYSIS

One of the most basic methods of analyzing the 

climatic and vegetational indications of a fossil leaf 

flora is by an analysis of the physiognomy of the 

leaves. In his comprehensive analysis of the Tropical 

Rain forest Richards (1952, p. 154) states: 

conclusions as to the climates of Tertiary floras are perhaps more 

firmly based when they are drawn from a statistical study of leaf 

sizes and similar features *** than when *** they *** rest on the 

taxonomic affinities of the fossil flora rather than its 

physiognomy, which appears, at least as far as modern vegetation 

is concerned, to be a very sensitive index of environmental 

conditions.

The strong relationship between foliar physiognomy 

and climate has been previously recognized by Bews 

(1927) and particularly by Bailey and Sinnott (1915, 

1916), who were the first to apply this relationship to 

an analysis of fossil floras. Several paleobotanists 

(for example, Chaney and Sanborn, 1933, MacGinitie 

1941, Wolfe and Hopkins, 1967) have followed the 

lead of Bailey and Sinnott.

Various physiognomic features of foliage are 

probably related to climatic conditions. Features 

such as size can be useful, but the influence of the 

depositional environment on sorting is a difficult 

factor to evaluate; larger leaves will tend to be 

fragmented, and thus the percentages of species in 

the larger size classes will probably be minimal 

figures for fossil assemblages. Tropical rain forests 

have a considerably higher percentage of species 

that have large leaves than do temperate forests. The 

presence or absence of drip-tips is also of significance; this adaptation is particularly common in 

tropical rain forests. The organization (compound 

vs. simple) and major venation pattern (pinnate vs. 

palmate) differs somewhat between tropical and 

temperate vegetation (Bailey and Sinnott, 1916), but 

the lack of an obvious correlation between environment and these features makes application of such 

differences very uncertain. One exception is the 

leaves of lianes; Richards (1952, p. 107) notes: 

The leaves of rain-forest climbers usually fall into the 'mesophylF 

size-class and approximate in texture as well as size to those 

prevailing in the tree stratum which they reach. In shape, 

however, they tend to be different. Simple elliptical or oblonglanceolate leaves are relatively uncommon, and there is a marked 

tendency, as in all climbing plants, towards short leaves broadest 

at the reniform or cordate base.*** The main veins usually diverge 

in a palmate manner *** well-defined drip-tips are not uncommon.***The type of leaf described is so strikingly prevalent 

among climbers, especially twiners, that it is difficult to believe it 

is due to chance.

The correlation between climbers and climate is 

striking. Richards (1952, p. 102-103) again notes: 

Climbing plants *** are far more abundant in the Tropical Rain 

forest than in any other plant formation *** in the West Indies, 

where Rain forest and similar formations are the predominant 

vegetation, woody lianes form about 8% of all the flowering 

plants.*** In Europe climbing plants form less than 2% of the 

flora.*** The great abundance of lianes in the tropics is hardly 

surprising in view of the great advantage of the climbing habit in 

a closed and very tall community such as a rain forest. 

Climbers, moreover, are especially abundant in 

openings in the Tropical Rain forest, particularly 

along rivers (Richards, 1952, p. 103). Richards also 

suggests that most lianes are intolerant of high rates 

of evaporation in the forest undergrowth; the 

abundance a lianes may, therefore, indicate the lack 

of a pronounced hot dry season.

Another physiognomic character of leaves that 

may prove to be useful is the density of venation. 

Tropical leaves have a pronounced tendency for the 

lamina to be supplied with many veins and this 

results in very small areoles; highly branching freely 

ending veinlets are less common than in temperate 

plants. There is clearly genetic control of the 

ultimate venation pattern, because, for example, in 

Tiliaceae, even the temperate members (such as 

Tilid) have small areoles with few freely ending 

veinlets, and in Sapindaceae even the tropical 

members (such as Euphoria) have branching freely 

ending veinlets. In both examples, however, the 

areoles of the tropical members are typically smaller 

than the areoles of temperate members of the same 

family. This relationship of venation to climate is 

probably also connected to the general xeromorphic 

nature of tropical leaves; leaves of plants inhabiting 

xeric regions in the temperate zones also tend to have 

smaller areoles than related species or genera that 

inhabit mesic regions of the temperate zones. As yet 

no quantification of this relationship has been made, 

involving as it does the clearing of leaves of all or 

most woody dicotyledonous species of floras in 

different»climatic regimes. The relationship is, 

however, useful in determining the "tropicality" or 

"temperateness" of a given extinct species or genus 

relative to extant related species and genera.

An additional physiognomic character of leaves 

that is correlated with climate is texture. The texture 

of some fossils is difficult to evaluate in some 

specimens because of the type of preservation. In 

most specimens, however, the texture of a given leaf 

can be approximately determined. A coriaceous 

texture is typical of "mesophyllous" tropical plants,
24 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

as well as the "microphyllous" species of arid and 

frigid climates. In addition, a coriaceous leaf 

indicates that the plant was evergreen. Thin leaves 

are particularly predominant in mesic temperate 

habitats; such leaves indicate a deciduous habit for 

the plants.

Perhaps the most striking correlation between 

climate and foliar physiognomy is the leaf margin. 

As originally pointed out by Bailey and Sinnott 

(1915), leaves that have entire margins are overwhelmingly prevalent in tropical regions, and leaves 

that have nonentire margins are characteristic of 

mesic temperate regions. The exact relationship 

between various environmental factors and the leaf 

margin is unclear; probably physiological aridity is 

in part responsible, as suggested by Bailey and 

Sinnott (1916, p. 36). In any case, the correlation of 

the entire margin on leaves that belong to the 

"mesophyll" size class with the tropical climates is 

striking.

The habitat has a significant effect on leaf 

margins, that is, whether a species is typically a 

slope or a stream- or lake-side species. Table 2 

indicates that vegetation in fluviatile or lacustrine 

habitats, whether trees or shrubs, has consistently 

lower percentages of entire-margined species. This 

fact is particularly significant in the application of 

leaf-margin analyses to fossil assemblages; MacGinitie (1953, p. 46) has noted that "***fossil floras 

represent, almost without exception, the specialized 

streamside or lakeside floras of the time and 

place***" Fossil assemblages may, therefore, yield 

lower precentages of entire-margined species than 

the percentage based on the unknown flora of the 

entire region. In dealing with temperate Neogene 

floras, it might be possible to arrive at a better 

estimate of the regional percentage by, for example, 

weighting the probable woodland and slope species 

more than the probable stream- and lakeside species 

(largely members of Salicaceae and Betulaceae). In 

dealing with Paleogene tropical and subtropical 

assemblages, however, the habitats of many, if not 

most, species are highly conjectural, and habitat 

data from modern tropical and subtropical vegetation are largely lacking. The tendency for lacustrine 

and fluviatile species to be more widespread and 

fewer in number than slope and woodland species is 

a compensating factor if several assemblages of a 

given age and from a given region are considered as 

one. For example, in the late Miocene floras east of 

the Cascade Range, the same species of Salicaceae 

and Betulaceae occur regularly in different floras, 

whereas the greatest floristic difference between

TABLE 2. The effect of habitat on leaf margins

[Data based on Hitchcock and others (1959, 1961, 1964)]

Percentages of species that 

Habitat have entire-margined leaves

Trees Shrubs Ligneous

West of Cascades, below 500 m;

All habitats .........................................

West of Cascades, above 500 m; 

Lacustrine or fluviatile ...................

All habitats .........................................

. 33

. 40

. 37

. 29

. 38

. 33

25

42

36

28

42

36

27

43

36

28

42

35

individual floras occurs in the probable slope and 

woodland species. The larger the flora and the more 

localities represented, therefore, the more valid the 

leaf-margin analysis becomes.

The probable overrepresentation of lacustrine and 

fluviatile plants in megafossil assemblages also 

indicates that consideration of leaf-margin percentages should be based on species rather than on 

specimens. Bailey and Sinnott (1916) note that in 

tropical forests the individual plants that have 

nonentire margins are extremely rare, and that a 

consideration of the percentage of leaves that have 

entire margins would probably produce an even 

stronger percentage gradient between the tropical 

and temperate vegetation than a consideration of the 

percentage of species; in fossil assemblages, the 

percentage should, however, be based solely on 

species.

RELATION OF LEAF MARGINS TO CLIMATE

In order to properly evaluate the climatic 

significance of a leaf margin or other physiognomic 

analyses, terms such as "tropical," "subtropical," 

and "warm temperate," should be related both to 

climatic conditions and leaf-margin percentages. 

These terms have been given rigorous definitions by 

some climatologists and geographers, but there are 

almost as many definitions as climatologists and 

geographers. Similarly, botanists apply terms such 

as "subtropical forest" to widely different vegetational types.

TROPICAL RAIN FOREST

The most commonly accepted definition of tropical 

climates used by climatologists is that the mean 

temperature of the coldest month is not below 18° C. 

If this definition is accepted, the forest of Hong Kong 

and lowland Taiwan that has been termed Tropical 

Rain forest by some botanists (Li, 1963, for example) 

is not truly tropical; these two regions have coldmonth means of less than 16°C. Regions that have
FLORISTIC AND VEGETATIONAL ANALYSIS 25

tropical climates under the meteorological definition 

and for which leaf-margin percentages have been 

compiled are (data after Bailey and Sinnott, 1916; 

Brown, 1919):

Percent

Brazil, Amazon lowland 89 

Malaya ,...,.... 86

Florida Keys 83

Philippine Islands, 200 m - -- 82 

Ceylon, lowland .- ..-.-- 81 

Manilla 81

East Indies 79

Philippine Islands, 450 m 76 

Hawaii, lowland -- 75

Philippine Islands, 700 m 72 

Philippine Islands, 1,100 m 69

The vegetation in most of these regions has 

generally been regarded as Tropical Rain forest. 

Notable exceptions are the oak-laurel forest (termed 

"Submontane Rain forest" by Richards, 1952, p. 87) 

at 700 m in the Philippine Islands, lowland Hawaii 

(presumably Subtropical Rain forest in Richards' 

terminology), and the montane forest (termed 

"Montane Rain forest" by Richards, 1952, p. 87) at 

1,100 m in the Philippine Islands. In terms of the 

climatologist, percentages above 68 could be termed 

"tropical," but to most botanists the Tropical Rain 

forest is indicated by percentages above 75.

PARATROPICAL RAIN FOREST

There is no universally accepted definition of 

subtropical, either in regard to climate or vegetation. 

The use of the term by Richards (1952) includes 

regions that are frost-free, whereas Wang (1961) and 

Li (1963) apply the term exclusively to regions that 

typically receive frost. As used by Richards, the 

terms "Submontane Rain forest," "Montane Rain 

forest," and "Subtropical Rain forest," indicate 

vegetation that is similar to the Tropical Rain 

forest except that the Tropical Rain forest has 

more stories and is thus somewhat more complex. 

Floristically these other vegetational types are 

closely allied to the Tropical Rain forest but are 

typically less rich. Wang (1961) has used the term 

"extratropical rain forest" for the vegetation of 

lowland southern China (the Subtropical Rain forest 

of Richards). Wang's term is preferable to Richards' 

in that, as noted previously, Queensland and 

Hawaii are truly tropical (not subtropical) in the 

commonly accepted meteorological definition of 

tropical. Extratropical, however, has too broad a 

connotation; the temperate conifer rain forest of the 

Olympic Peninsula of Washington is an extratropical rain forest.

The term "Paratropical Rain forest" (para, 

meaning close) is here proposed for the vegetation 

that Richards (1952) termed "Subtropical Rain 

forest" and "Submontane Rain forest" and that 

Wang (1961) termed "extratropical rain forest." This 

type of forest is vegetationally similar to, but simpler 

in structure than, the Tropical Rain forest; floristically, the Paratropical Rain forest is closely allied 

to the Tropical Rain forest. The Paratropical Rain 

forest is not limited poleward by the mere presence of 

frost; for example, the Paratropical Rain forest of 

Taiwan has been subjected to -1° C, and northern 

Hainan has experienced a temperature of -3° C. The 

mean annual temperature ranges from about 20°C 

up to about 25°C. The cold-month temperature has a 

mean of about 13°C-24°C. Precipitation can be 

seasonal, but extended dry periods are lacking. 

These climatic conditions also pertain to the 

Montane and Submontane Rain forests of Richards 

(1952), and, except for the precipitation regime, to 

some monsoonal forests of the tropical region. Leaf 

margin percentages for the Paratropical Rain forest 

vary from 57 up to 75 (table 3); Note that the 

Paratropical Rain forest includes vegetation that is 

predominately broad-leaved evergreen, although 

some deciduous plants may be (and typically are) 

present. Some workers have applied distinct terms 

such as "Semi-deciduous forest" to vegetation of 

which a large partis deciduous; such refinements are 

probably not readily recognizable in relation to fossil 

assemblages.

TABLE 3. Percentages of species that have entire margins in 

some modern floras

Flora Percentage Vegetation

Brazil, lowland .......................

Malaya .....................................

Philippine Islands, 200 m . 

Ceylon, lowland .....................

Manila .....................................

East Indies .............................

Philippine Islands, 450 m . West Indies .............................

Hawaii, lowland ...................

Ceylon, upland .......................

Philippine Islands, 700 m . 

Hong Kong .............................

Hainan, lowland ...................

Philippine Islands, 1,100 m 

Taiwan, 0-500 m ...................

Ceylon, upland .......................

Hawaii, upland .....................

Hainan, upland .....................

Taiwan, 500-2,000 m ...........

Mixed Mesophytic forest, 

China ...................................

88 Tropical Rain forest

86 Do.

82 Do.

81 Do.

81 Do.

79 Do.

76 Do.

76 Do.

75 Paratropical Rain forest

74 Do.

72 Do.

72 Do.

70 Do.

69 Montane Rain forest

61 Paratropical Rain forest

60 Do.

57 Do.

55 Subtropical forest

45 Do.

30 Warm temperate forest
26 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Note that the leaf margin percentages of vegetation of tropical mountains ("Submontane Rain 

forest," "Montane Rain forest") and of the lowlandpart of the Paratropical Rain forest have about the 

same range. This range indicates some similarity of 

environmental conditions (for example, mean annual temperature), at least those conditions that 

affect leaf margins.

Some workers may object to the introduction of the 

term "paratropical" on the grounds that the concept 

to which the term is applied is the same concept as 

subtropical. The term "subtropical," however, covers 

a much broader range of climates and vegetation 

types than does paratropical. As noted previously, 

the term "tropical" is applied to climates in which 

the mean temperature of the cold month is above 

18°C, and subtropical is typically applied to climates 

that have cold-month means between about 6°C 

(Landsberg, 1964, p. 924) and 18°C. Within this 

broad climatic range, the vegetation in mesic regions 

can be readily classified into two major types. The 

first, which I call "paratropical," is physiognomically similar to the Tropical Rain forest in that more 

than one tree story is present, woody lianes are 

diverse and abundant, and buttressing of the tree 

trunks is present. The second, to which I prefer to 

restrict the term "subtropical," is physiognomically 

dissimilar to the paratropical vegetation and 

typically has abundant sclerophylls, lacks a diverse 

and abundant woody liane element, and has no 

buttressing. Despite the fact that both Paratropical 

Rain and subtropical forests are dominantly broad 

leaved evergreen and that both forest types are 

floristically related, the physiognomy of the two 

types differs more than does the physiognomy of the 

Tropical and Paratropical Rain forests.

Sl'B I ROPICAL FOREST

The term "subtropical forest" is reserved here for 

broad-leaved evergreen vegetation that grows under 

a climate in which frosts are present, although never 

severe or of long duration. Some workers might 

prefer the term "warm temperate," but the Mixed 

Mesophytic forest of east central Asia, for example, 

is traditionally considered to be warm temperate, 

and it exists in a region that receives regular hard 

frosts (Wang, 1961). The oak-laurel forest of southern 

China and at moderate altitudes on Taiwan is an 

example of a subtropical forest. Vegetationally, 

therefore, the term "subtropical" indicates a broadleaved evergreen forest (in some instances conifers 

and broad-leaved deciduous plants are important 

constituents of this forest) that exists under

moderate frost. Mean annual temperature ranges 

from about 13° to 18° C and the mean of the coldest 

month ranges from about 0° to 18°C. Leaf-margin 

percentages vary from about 39 up to 55.

WARM TEMPERATE FOREST

As will be shown, only one Gulf of Alaska 

Ravenian or Kummerian assemblage has a leafmargin percentage below 39. The warm temperate 

vegetation thus needs to be mentioned only briefly. 

This modern vegetation is broad-leaved deciduous 

and may have a significant amount of conifers 

intermixed. Broad-leaved evergreens are present but 

not dominant. Freezing temperatures can be 

expected during several months of the year and the 

temperature regime has a pronounced seasonality. 

The mean annual temperature ranges from about 

11° to 13°C, and the mean of the coldest month 

ranges from about -3° to 2°C. Leaf-margin percentages range from about 30 to 38. Under less 

seasonality but the same mean annual temperature, 

the warm temperate broad-leaved deciduous forests 

are typically replaced by conifer forests.

LOWER RAVENIAN

FLORISTIC ANALYSIS

Systematic list of the Alaskan lower Ravenian (late 

middle Eocene) flora

Filicineae 

Filicales 

Schizaeaceae

Anemia gunni Wolfe 

Cyatheaceae

Cyathea inequilateralis (Roll.) Wolfe 

Aspidaceae

Allantodiopsis pugetensis Wolfe 

Dryopteris alaskana (Roll.) Wolfe 

Gymnospermae 

Coniferales 

Taxodiaceae

Glyptostrobus sp. 

Angiospermae 

Monocotyledonae 

Principes 

Palmae

Sabalites sp. 

Dicotyledonae 

Juglandales 

Juglandaceae 

Platycarya pseudobrauni (Roll.) Wolfe
FLORISTIC AND VEGETATIONAL ANALYSIS 27

Angiospermae Continued 

Dicotyledonae Continued 

Fagales 

Betulaceae

Alnus martini Wolfe

Alnus scotti Wolfe 

Urticales 

Ulmaceae

Girroniera sp.

Girroniera sp. 

Ranales

Tetracentraceae

Tetracentron piperoides (Lesq.) Wolfe 

Menispermaceae

Anamirta milleri Wolfe

Calkinsia plafkeri Wolfe

Cocculus sp.

Diploclisia sp.

Limacia stenophylla Wolfe

Paratinomiscium conditionalis (Holl.) Wolfe

Pycnarrhena sp. 

Illiciaceae

Illicium sp. 

Annonaceae

Cananga sp.

Dasymaschalon? sp. 

Myristicaceae

Knema sp.

Myristica sp. 

Lauraceae

Cinnamomophyllum latum (MacG.) Wolfe

Litseaphyllumpresamarensis (Sanb.) Wolfe

Litseaphyllum similis (Knowlt.) Wolfe

Litseaphyllum sp.

Litseaphyllum sp. 

Resales 

Leguminosae

Caesalpinites sp. 

Geraniales 

Meliaceae

Toona sp. 

Rutaceae

Euodia alaskana Wolfe

Luvunga spatiosa (Holl.) Wolfe 

Sapindales 

Anacardiaceae

Melanorrhoea alaskana (Holl.) Wolfe

Rhus cf. R. mixta (Lesq.) MacG. 

Celastraceae

Celastrus comparabilis Holl. 

Icacinaceae

Goweria alaskana Wolfe

Goweria dilleri (Knowlt.) Wolfe

Paleophytocrene elytraeformis (Holl.) Wolfe

Angiospermae Continued 

Dicotyledonae Continued 

Sapindales Continued 

Icacinaceae Continued 

Phytocrene acutissima Wolfe 

Phytocrene sordida (Lesq.) MacG. 

Pyrenacantha sp. 

Stemonurus alaskanus Wolfe 

Sapindaceae

Allophylus duktothensis Wolfe 

Sabiaceae

Meliosma duktothensis Wolfe 

Meliosma kushtakensis Wolfe 

Rhamnales 

Rhamnaceae 

Sageretia sp.

Zizyphus alaskanus Wolfe 

Vitaceae

Vitis sp. 

Parietales 

Actinidiaceae 

Saurauia sp. 

Dipterocarpaceae

Parashorea pseudogoldiana (Holl.) Wolfe 

Myrtiflorae 

Barringtoniaceae

Barringtonia sp. 

Rhizophoraceae

Kandelia wangi Wolfe 

Alangiaceae

Alangium bergensis Wolfe 

Alangium sp. 

Myrtaceae

Eugenia sp. 

Umbelliflorae 

Cornaceae

Mastixia irregularis (Holl.) Wolfe 

Ericales 

Clethraceae

Clethra sp. 

Tubiflorae 

Verbenaceae

Clerodendrum sp.

The known flora of the lower Ravenian in Alaska 

is diverse despite the small size of the collections. 

Altogether at least 71 species are represented, 

although only 60 show sufficient characters to be 

determined to family. At least 47 genera and 31 

families are represented. The most diverse families 

represented are Menispermaceae, Lauraceae, and 

Icacinaceae. The strong floristic relationship of this 

flora to that of the London Clay is apparent in that 23 

families and 10 genera are common to both floras, 

despite the fact that a leaf flora is-being compared to
28 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

a fruit and seed flora and each have their own organ 

genera. The floristic similarities are even more 

pronounced in that the most diverse families of the 

London Clay are also diverse in the Alaskan lower 

Ravenian. The majority (27) of the genera and all the 

families except for Myristicaceae, Barringtoniaceae, 

and Clethraceae, are also known in early or middle 

Eocene floras from western conterminous United 

States. Twenty-one of the lower Ravenian species are 

represented by the same or closely related species in 

the Eocene floras from western conterminous United 

States. Floristically, therefore, the lower Ravenian 

flora of Alaska is closely related to the early middle 

Eocene flora of Eurasia and western conterminous 

United States, as well as to the lower Ravenian flora 

of Washington.

The relationship of the Alaskan lower Ravenian 

flora to extant floras is clear: the Alaskan lower 

Ravenian flora is overwhelmingly paleotropical. Not 

one of the genera or families determined is 

exclusively neotropical, whereas four of the families 

and 23 of the genera are exclusively paleotropical; to 

the generic figure can be added six additional extinct 

genera that have their closest living relatives 

restricted to the paleotropical region. In instances in 

which a genus is found in both old and new worlds, 

the fossil most closely resembles the old-world 

representatives. A few genera are today of warm 

temperate distribution in Asia; the significance of 

these genera is discussed in a later section (p. 43).

Several Alaskan lower Ravenian genera are today 

exclusively members of the Tropical Rain forest, and 

some species of more wide-ranging genera are most 

closely related to members of theTropical Rain forest:

Anamirta Celastrus comparabilis

Limacia Stemonurus

Pycnarrhena Phytocrene

Knema Pyrenacantha

Myristica Parashorea

Euodia Barringtonia

Luvunga

The bulk of the genera and species, however, are 

related to members of the present Paratropical Rain 

forest or the Submontane Rain forest of the 

Paleotropical region.

VEGETATIONAL ANALYSIS

In order to eliminate, at least partially, the effect of 

probable over-representation of stream-side plants, 

the various Lower Ravenian assemblages have been 

treated as one for the leaf-margin analysis. This 

procedure also yields a considerably larger statistical base than if percentages for each assemblage are

calculated. It could be argued that, because the 

assemblages are not from the same precise 

stratigraphic plane, combining the assemblages 

eliminates any possibility of recognizing major but 

brief climatic fluctuations (this indeed happened 

when I computed the percentage for the type 

Ravenian upper Puget that was published by 

Wolfe and Hopkins, 1967). If the individual Alaskan 

assemblages were larger, separate computations 

should indeed be made, but most assemblages 

contain such small numbers of species that 

individual computations are not firmly based 

statistically. If, for example, the percentage for 

locality 3847 (80 percent) is compared with that for 

locality 11166 (57 percent), a considerable deterioration of climate might be construed; such a conclusion 

should, however, be based on a larger body of data, 

that is, at least 20 or more specific entities from a 

given locality.

The percentage of Alaskan lower Ravenian 

dicotyledonous species that have entire margins is 

65; this figure is based on 62 species. In terms of the 

percentages discussed previously, the lower Ravenian assemblage is neither Tropical Rain forest nor 

subtropical forest, but the percentage is intermediate 

between that for the Paratropical Rain forest of 

Hawaii and that for the Paratropical Rain forest of 

Taiwan.

In leaf size and foliar organization (table 4), the 

Alaskan lower Ravenian assemblage compares well 

with the Paratropical (Submontane) Rain forest of 

the Philippine Islands (Brown, 1919), particularly if 

it is kept in mind that leaves of larger sizes are 

underrepresented in fossil assemblages. I have also 

included in table 5 the figures for the early Miocene 

Collawash flora of Oregon, which is a Mixed 

Mesophytic, warm temperate forest. Clearly the 

Alaskan lower Ravenian is unrelated vegetationally 

to the Mixed Mesophytic forest, although there is 

some floristic similarity (p. 44).

Comparisons of the leaf-size classes of the Alaskan 

Early Ravenian assemblage to those of the extant 

Australian rain forests are informative. In a detailed 

study, Webb (1959) has shown significant differences in leaf size between the various forest types, 

ranging from Paratropical Rain forest to temperate 

rain forest types. Based in part on a study of leaf-size 

classes, Webb also formally subdivided the three 

major forest formations of eastern Australia. In 

table 5 some of Webb's data are summarized. The 

notophyll-size class is an addition to the Raunkiaer 

(1934) system of size classes and represents the 

smaller (2,025-4,500 mm2 ) leaves that Raunkiaer 

included in his mesophyll-size class. (As used
FLORISTIC AND VEGETATIONAL ANALYSIS 29

TABLE 4. Comparisons between modern and fossil foliar physiognomy

Flora

Alaskan Kummerian _._._....._.................................................... .

Culled dipterocarp forest, Philippines ........................... .......

Collawash, early Miocene, Oregon ........................................

Number 

of 

species

... ..... . 65

.......................... 37

20

........................... 28

........................... 125

........................... 90

........... ................ 68

. ... ... . 16

........................... 119

Percent 

entire

65

54

20

55

82

76

72

69

25

Percent

less than 

5 cm in width

36

50

86

67

32

32

40

75

66

Percent 

less than 

10 cm in length

42

45

91

53

13

18

18

69

66

elsewhere in this report, mesophyllous is used in 

accordance with Raunkiaer's original definition.) 

Omitted from table 5 are Webb's data for most of the 

subformations representing woodland or thicket 

vegetation; all these have 40 percent or more 

microphyll species and typically a low percentage of 

mesophyll species. The Alaskan early Ravenian 

assemblage in terms of species is: 34 percent 

mesophyll, 50 percent notophyll, and 16 percent 

microphyll. Clearly, the early Ravenian does not 

represent temperate vegetation.

Based on leaf-size classes, the Alaskan early 

Ravenian assemblage is similar to two Australian 

vegetation types: the Complex Notophyll Vine forest 

(Paratropical Rain forest) and the Simple Mesophyll 

Vine forest (Submontane Rain forest). The Simple 

Mesophyll Vine forest is in Australia, as elsewhere, 

tropical in the meteorological definition. The 

Complex Notophyll Vine forest is subtropical in the 

meteorological definition; this forest extends from 

about latitude 21° S. to about 35° S. (Webb, 1959). 

This area of the Complex Notophyll Vine forest lacks 

frost. Webb (1959, p. 552-553) notes that "All the 

Rain forest formations, away from their optimum, 

have smaller leaf sizes, and undergo other physiognomic and structural changes associated with gradients of temperature (altitude or latitude) soil 

properties (moisture, drainage, fertility), and exposure." Because the Alaskan early Ravenian assemblage has a somewhat larger percentage (34 

compared to 30) of mesophylls than Webb (1959) 

recorded for any plots in the Complex Notophyll 

Vine forest, the early Ravenian may represent the

optimal development of that forest or may represent 

simple Mesophyll Vine forest. This suggestion has 

even more validity if, as suggested previously, the 

larger size classes are underrepresented in fossil 

assemblages.

The fragmentary nature of many of the fossils does 

not allow a complete analysis of the percentage of 

dicotyledonous species that have drip-tips. Obvious 

drip-tips are, however, present in many species, for 

example, Girroniera spp., Limacia stenophylla, 

Luvunga spatiosa, Phytocrene spp., Parashorea 

pseudogoldiana, and Alangium bergensis. The 

climatic conditions that are responsible for the 

natural selection of drip-tips in tropical lineages 

were probably present in Alaska during the early 

Ravenian.

The length of the drip-tip and its absence or 

presence provide clues as to both the vegetational 

type, as noted earlier, and the position of the 

particular species in the vegetation. Richards (1952, 

p. 84-87) notes that drip-tips are uncommon on 

leaves from the upper stories in comparison to leaves 

from the lower stories, including undergrowth. 

Elongated drip-tips on leaves of Girroniera spp., 

Alangium, bergensis, and Phytocrene acutissima, 

indicate that these plants were most probably 

members of the lower stories of the forest. Not so 

elongated are the drip-tips of Allophylus duktothensis, Luvunga spatiosa, Stemonurus alaskanus, 

Phytocrene sordida, and "Rhododendron crassum"; 

these species could presumably have been members 

of upper stories. Species such as Euodia alaskana 

would more likely have been members of the upper

TABLE 5. Distribution of leaf-size classes in the vegetation of eastern Australia

__________________________[Data from Wefab (1959)]____________ _________

Percent of species that are:

Paratropical Rain forest ..........

Monsoon forest ....... .......... ........

Submontane Rain forest ..........

Submontane Rain forest ..........

Montane Rain forest ................

Subtropical forest ......................

Sublormation tarter weoD)

... .........Simple Mesophyll Vine forest ....... .. . ..

............Simple Notophyll Vine forest . ................

............Microphyll Mossy thicket ....... . . ....... ........

............Complex Notophyll Vine forest ... ... ...... ...........

Mesophyll

50-70

30

30-50

0-30

0

15-30

Notophyll

30-40

40

40-50

55-70

0

50-70

Microphyll

A K.

30

0-15

0-40

100

10-20
30 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

story, and the emarginate apex of Caesalpinites sp. 

clearly indicates exposure to the hotter conditions of 

the upper story.

The percentage of probable lianes in the Alaskan 

lower Ravenian is very high (about 25 percent of the 

species). Species that have leaves that clearly 

conform to the morphology of liane leaves mentioned 

by Richards (1952, p. 107; p. 23 of this report) are 

Anamirta milleri, Cocculus sp., Diploclisia sp., 

Paratinomiscium conditionalis, Phytocrene acutissima, and P. sordida. Other species of genera whose 

extant members are exclusively lianes are Limacia 

stenophylla, Pycnarrhena sp., Luvunga spatiosa, 

Celastrus conditionalis, Pyrenacantha sp., and 

Vitis sp. The tribe Phytocreneae is also today 

composed exclusively of climbers, and hence 

Goweria alaskana, G. dilleri, and Paleophytocrene 

elytraeformis may also be members of the liane 

element. The diversity of probable lianes is another 

indication that the Alaskan lower Ravenian 

vegetation was, in the broad sense, tropical. The 

coriaceous texture of most of the leaves (see, for 

example, pi. 5, fig. 7; pi. 6, fig. 8, pi. 8, fig. 4) is a strong 

indication that the plants that bore them were 

evergreen.

The Alaskan early Ravenian assemblages contain

a probable strand element. Barringtonia today 

grows behind beaches in the South Pacific; the beds 

from which the Alaskan specimens were obtained 

are both under- and overlain by beds containing 

brackish-water mollusks. The mangrove Kandelia 

today extends along shores from southeast Asia 

north to southern China; I suggest that such a highly 

specialized habitat as the mangroves of Rhizophoraceae have today is an ancient adaptation and 

that the Alaskan Kandelia was also a mangrove.

Some workers consider that the numerical 

abundance of plant megafossils closely reflects the 

relative abundance of organisms of various species 

in the forest of the area of the site of deposition. For 

those workers, tables 6 and 7 are presented. The 

correlation, however, between representation of 

large organs and representation of species in the 

forest has yet to be demonstrated; the supposed 

correlation is an unfounded assumption. Quaternary 

pollen workers, on the other hand, are of almost 

unanimous opinion that megafossil assemblages 

generally represent the vegetation near the immediate site of deposition and thus do not adequately 

reflect the vegetation of the region (see Faegri and 

Iversen, 1964, p. 39). This opinion is based on 

numerous studies of modern pollen rain in a region

TABLE 6. Numerical abundance of plant megafossils from the Alaskan lower Ravenian

Species

Limacia stenophylla ...........................

Celastrus comparabilis .......................

Allophylus duktothensis .....................

Alnus martini ........................................

Platycarya pseudobrauni ...................

Meliosma duktothensis .......................

Sabalites sp .............................................

Vitis sp .....................................................

Phytocrene acutissima .......................

Eugenia sp .................... .........................

Dryopteris alaskana .. ..... .

Clethra sp ...............................................

Litseaphyllum similis

Parashorea pseudogoldiana ...............

Alnus scotti ................. ........................

Clerodendrum sp ...................................

Meliosma kushtakensis .......................

Allantodiopsis pugetensis .................

Melanorrhoea alaskana .......................

Litseaphyllum presamarensis .........

Glyptostrobus sp ...................................

Alangium bergensis .............................

Cyathea inequilateralis .......................

Myristica sp ...........................................

Anamirta milleri ...................................

Cinnamomophyllum latum ...............

Luvunga spatiosa .................................

Euodia alaskana ...................................

Barringtonia sp .....................................

Others .......................................................

Totals .............................................

11157

3847

....... 4

6

1

10

....... 4

9

8

1

1

3

3

2

9

....... 61

11159

11158

46

20

22

4

13

1

1

8

5

4

3

3

21

151

11160

11161 11166 3846

.............. 2 ..............

.............. 5 1

.............. 1 ..............

14 .............. ..............

.............. 19 ..............

.............. 5 ..............

.............. .............. 1

.............. 2 1

9 .............. ..............

............. 3 ..............

.............. 1 1

.............. .............. 2

.............. .............. 5

.............. 2 ..............

2 .............. 1

.................. 1 ..............

1 10 2

26 51 14

11170

17

39

17

31

28

1

1

19

10

1

3

3

16

181

3879 11195 Total

.............. .............. 48

1 .............. 48

.............. .............. 40

.............. .............. 39

.............. 1 38

.............. 1 29

.............. 5 24

.............. .............. 20

.............. .............. 19

.............. .............. 19

6 .............. 17

.............. .............. 14

.............. .............. 13

.............. 1 10

.............. .............. 9

.............. .............. 8

.............. .............. 8

1 .............. 7

.............. .............. 7

.............. .............. 5

4 .............. 4

1 .............. 4

.............. .............. 3

.............. .............. 3

.............. .............. 3

.............. .............. 3

.............. .............. 3

.............. .............. 3

.............. .............. 3

3 7 70

16 15 515

Percent

9.3

9.3

7.5

7.3

7.1

5.6

4.9

3.8

3.7

3.7

3.3

2.7

2.5

1.9

1.7

1.5

1.5

1.3

1.3

1.0

0.8

.8

.6

.6

.6

.6

.6

.6

.6

12.6

99.3
FLORISTIC AND VEGETATIONAL ANALYSIS 31

TABLE 7. Numerical abundance and diversity of families in the 

Alaskan lower Ravenian

Family Number

Number Number 

Percent of genera of species

Menispermaceae ............

Betulaceae ........................

Celastraceae ...................

Sapindaceae ...................

Juglandaceae ..................

Sabiaceae ..........................

Icacinaceae .....................

Palmae ..............................

Lauraceae ..........................

Vitaceae ...........................

Myrtaceae .......................

Clethraceae ......................

Aspidaceae ........................

Dipterocarpaceae ...........

Verbenaceae ....................

Anacardiaceae ................

Rutaceae ............................

Myristicaceae .................

...... 59

...... 48

...... 48

40

...... 37

...... 37

...... 26

...... 25

....... 23

...... 20

...... 19

...... 14

24

...... 10

8

7

6

5

11.6

9.3

9.3

7.8

7.2

7.2

5.1

4.9

4.5

3.9

3.7

2.7

4.7

1.9

1.6

1.4

1.2

1.0

7

1

1

1

1

1

5

2?

2

1

1

1

2

1

1

1

2

2

7

2

1

1

1

2

7

2?

5

1

1

1

2

1

1

1

2

2

as compared to the vegetation, and the application of 

these data to fossil Quaternary pollen and megafossil assemblages. I agree that the representation of 

leaves is largely a reflection of the existence of 

certain species near the site of deposition and that 

leaf counts are largely meaningless in determining 

the vegetation of the region of the depositional basin. 

The general floristic relationships, the presence of 

large-leafed ferns (including a genus Hemitelia 

whose extant members are tree ferns), palms, diverse 

probable lianes, a high percentage of species that 

have entire-margined leaves, many large-leafed 

species, many species that have drip-tips, and many 

species that had a coriaceous texture all these are a 

strong indication that the Alaskan lower Ravenian 

vegetation should be considered as Paratropical 

Rain forest.

MIDDLE RAVENIAN

FLORISTIC ANALYSIS

Systematic list of the Alaskan middle Ravenian 

(lower upper Eocene) flora

Filicineae 

Filicales 

Schizaeaceae

Lygodium larsoni Wolfe 

Aspidaceae

Allantodiopsis pugetensis Wolfe 

Gymnospermae 

Coniferales 

Taxodiaceae

Glyptostrobus sp. 

Angiospermae 

Monocotyledones

Angiospermae Continued 

Monocotyledones Continued 

Principes 

Palmae

Phoenicites sp. 

Sabalites sp. 

Dicotyledones 

Juglandales 

Juglandaceae

Carya cashmanensis Wolfe 

Engelhardtia sp.

Platycarya pseudobrauni (Holl.) Wolfe 

Pterocarya pugetensis Wolfe 

Fagales 

Fagaceae

Dryophyllum pugetensis Wolfe 

Urticales 

Ulmaceae

Ulmus kushtakensis Wolfe 

Ranales 

Tetracentraceae

Tetracentron piperoides (Lesq.) Wolfe 

Menispermaceae

Calkinsia plafkeri Wolfe 

Lauraceae

Cinnamomophyllum kushtakensis Wolfe 

Litseaphyllum kushtakensis Wolfe 

Litseaphyllum presamarensis (Sanb.) Wolfe 

Litseaphyllum schorni Wolfe 

Litseaphyllum sp. 

Litseaphyllum sp. 

Resales 

Hamamelidaceae

Liquidambar? sp. 

Eucommiaceae

Eucommia sp. 

Leguminosae

Leguminosites sp. 

Sapindales 

Anacardiaceae 

Genus indet. 

Sapindaceae 

Allophylus sp. 

Sapindus? sp. 

Parietales 

Theaceae

Ternstroemites sp. 

Incertae sedis

Artocarpoides kummerensis Wolfe 

Macclintockia sp.

The total number of species known from the 

Alaskan middle Ravenian is smaller than for the 

early Ravenian. This apparent lesser diversity may
32 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

be a function of the fewer localities of middle 

Ravenian age from which fossils were collected. 

Locality 11165 has produced representatives of 37 

species, although not all could be determined to 

family because of poor preservation. This assemblage is the richest known from a single Alaskan 

Eocene locality.

In contrast to the lower Ravenian assemblages, 

Lauraceae are proportionally better represented; 

Menispermaceae are poorly represented and Icacinaceae are apparently absent. Juglandaceae are also 

better represented in the middle than in the lower 

Ravenian. Most middle Ravenian families and 

genera are known in the lower Ravenian, either in 

Alaska or conterminous United States. Many species 

are also common to the two substages, and it is clear 

that the bulk of the middle Ravenian flora is closely 

allied floristically to the lower Ravenian.

The relationships of the middle Ravenian flora to 

extant floristic regions is Asian, as in the lower 

Ravenian. The floristic similarities of the middle 

Ravenian to the Indomalaysian region is, however, 

much less pronounced. The closest relationship of 

the middle Ravenian appears to be with the 

Notophyllous Broad-leaved Evergreen forest. The 

representation of Juglandaceae (exclusive of Pterocarya), Fagaceae, Lauraceae, Liquidambar?, and 

Theaceae, gives the Alaskan middle Ravenian flora 

a taxonomic aspect that is most similar to that of the 

notophyllous, subtropical, oak-laurel forest of eastern Asia.

The mixed mesophytic element, although not 

dominant, is more pronounced in the middle than in 

the lower Ravenian. The Juglandaceae (exclusive of 

Engelhardtia), Ulmus, Tetracentron, Eucommia, 

and Liquidambar?, are members of this element. 

Note, however, that five of the seven mixed mesophytic genera in the Alaskan middle Ravenian are 

known in correlative assemblages in the Puget 

Group, and a sixth Platycarya is a common 

member of early and middle Eocene assemblages 

in conterminous United States.

VEGETATIONAL ANALYSIS

The percentage of middle Ravenian species that 

have entire margins is 54, compared to 65 for the 

lower Ravenian. The middle Ravenian figure is 

based on 36 dicotyledonous species. Comparisons 

with modern vegetation (table 4) indicate that the 

middle Ravenian is only marginally Paratropical 

Rain forest or more probably in the warmer part of 

the notophyllous forest. The leaf-margin analysis is 

corroborated by the proportionately fewer coriaceous 

and hence probably broad-leaved evergreen species,

the sparse representation of probable lianes (the only 

probable liane is Calkinsia), and the almost total 

lack of leaves that have drip-tips.

The contrast between the lower and middle 

Ravenian in both vegetation and flora is significant 

in that both kinds of evidence indicate a cooler 

climate for the middle Ravenian. In Alaska, light 

frosts probably characterized the middle Ravenian 

climate, whereas they were probably lacking in the 

early Ravenian. In the middle Ravenian of the Puget 

Group, the physiognomic characters of the foliage 

indicate the persistence in the conterminous United 

States of Paratropical Rain forest and hence 

typically frost-free climate. The middle Ravenian 

floras of the Puget Group, however, also indicate a 

climate cooler than that of the lower Ravenian, as 

shown by a decline in the leaf-margin percentages 

from 75 to 62 (Wolfe, 1971).

UPPER RAVENIAN

FLORISTIC ANALYSIS

Systematic list of the Alaskan upper Ravenian 

uppermost Eocene) flora

Gymnospermae 

Coniferales 

Taxodiaceae

Glyptostrobus sp. 

Angiospermae 

Dicotyledones 

Salicales 

Salicaceae 

Populus? sp. 

Salix carbonensis Wolfe 

Salix sp. 

Juglandales 

Juglandaceae 

Juglans (Cardiocaryori) sp. 

Pterocarya pugetensis Wolfe 

Fagales 

Betulaceae

Alnus cuprovallis Axelr. 

Alnus sp. 

Alnus sp. 

Ranales

Menispermaceae

Cocculus sp. 

Resales 

Hamamelidaceae

Corylopsis sp. 

Rosaceae

Prunus axelrodi Wolfe 

Sorbus carbonensis Wolfe
FLORISTIC AND VEGETATIONAL ANALYSIS 33

Angiospermae Continued 

Dicotyledones Continued 

Sapindales 

Aquifoliaceae

Ilex carbonensis Wolfe 

Rhamnales 

Rhamnaceae

Rhamnus? sp. 

Malvales 

Tiliaceae

Tilia carbonensis Wolfe 

Plafkeria sp. 

Myrtiflorae 

Myrtaceae

Eugenia? sp. 

Incertae sedis

Dicotylophyllum carbonensis Wolfe

The Alaskan upper Ravenian is, unfortunately, 

known from only one locality; the preservation of the 

leaves, however, is the best for any of the Alaskan 

Eocene localities. At least 21 specific entities are 

represented, but only 19 were sufficiently complete to 

determine to family, and one of these (Dicotylophyllum carbonensis), although the most abundantly represented species, is thought to represent 

an extinct genus.

The upper Ravenian assemblage is not closely 

allied floristically to the lower or middle Ravenian. 

Only 5 of the 15 genera are known from the earlier 

Ravenian assemblages. It is also significant that 

Lauraceae, Rutaceae, and Dipterocarpaceae are 

unrepresented in the upper Ravenian. Palynologic 

evidence, however, indicates that some upper 

Ravenian genera, for example Salix, Juglans, and 

Tilia, were represented in subtropical or Paratropical 

Rain forests in conterminous United States during 

the earlier Eocene and leaves of Populus are known 

in subtropical forests in the middle Eocene (Leopold 

and MacGinitie, 1972). One element, represented by 

Corylopsis and Dicotylophyllum carbonensis, seems 

to be derived from high-latitude warm temperate and 

subtropical Paleocene assemblages, although Corylopsis was also a member of Paratropical Rainforest 

during at least the early Eocene (Chandler, 1964). 

Part of the Alaskan upper Ravenian flora may, 

therefore, have been derived from elements that were 

present but rare in the Paratropical Rain and 

subtropical forests of the early and middle Eocene.

The Alaskan upper Ravenian also has two species 

that occur in upper Ravenian upland assemblages in 

conterminous United States. Alnus cuprovallis and 

Prunus axelrodi are also known in assemblages such 

as that from Copper Basin (Axelrod, 1966b). The

lowland upper Ravenian assemblages from conterminous United States (Puget Group, John Day Gulch 

assemblage of the Clarno Formation) has one 

species, Pterocarya pugetensis, that also occurs in 

the Alaskan upper Ravenian. The Alaskan upper 

Ravenian rosaceous genera have not been validly 

recorded from earlier Paleogene assemblages; I 

suggest that the strong representation of the family 

in upper Ravenian floras throughout western North 

America is a reflection of the time of diversification 

of Rosaceae. Note that Chandler (1964) has only 

recorded one rosaceous genus, the partially tropical 

Rubus, in the British Eocene.

The relationships of the Alaskan upper Ravenian 

to modern floristic regions are not clear. The taxa 

probably derived from the main elements of the 

Alaskan early and middle Ravenian Cinnamomophyllum, Plafkeria, Eugenia? sp., and Glyptostrobus are, of course, in a broad sense paleotropical. Corylopsis and Pterocarya are found today only 

or primarily in the Mixed Mesophytic forest, and all 

but four upper Ravenian genera also range into that 

forest type. In general the Alaskan upper Ravenian 

flora has its closest relationship to the broad-leaved 

deciduous forest of eastern Asia.

VEGETATIONAL ANALYSIS

The leaves of the dicotyledonous species represented in the Alaskan upper Ravenian have margins 

that are overwhelmingly nonentire. Of the 20 

dicotyledons known, only 20 percent have entire 

margins. The leaf margin percentage is about 

midway between that for mesic temperate forests, 

such as New England, and that for mesic warm 

temperate forests, such as the Mixed Mesophytic 

forest. If, as discussed previously, stream-side types 

are overrepresented (this is a particularly reasonable 

assumption for a small assemblage), the upper 

Ravenian vegetation is more comparable to the 

Mixed Mesophytic forest. The dicotyledons represented are predominantely deciduous; five species 

the Cinnamomophyllum, Plafkeria, Eugenia?, Dicotylophyllum, and a fragmentary, generically 

indetermined species have a coriaceous texture and 

thus were probably evergreen.

The small size of the Alaskan upper Ravenian 

assemblage, however, does not yield a firm 

statistical base for a precise leaf margin analysis. 

The fact that 25 percent of the dicotyledons known 

were broad-leaved evergreen indicates that, if the 

forest was broad-leaved deciduous, it would have 

fallen in the highly warm-temperate part, that is, in 

the warmer part of the Mixed Mesophytic forest 

where broad-leaved evergreens are significantly
34 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

represented. Although the assemblage clearly 

represents vegetation cooler than that of the middle 

Ravenian, just as clearly more collections are needed 

to determine beyond reasonable doubt the exact 

vegetational type represented.

Vegetationally and floristically the Alaskan upper 

Ravenian assemblage is most similar to the extant 

Mixed Mesophytic forest of Asia. To consider, 

however, the forest represented by the Alaskan 

upper Ravenian as a lineal predecessor floristically 

or vegetationally of the Mixed Mesophytic forest 

may not be valid. At least four of the upper Ravenian 

species Pterocarya pugetensis, Alnus cuprovallis, 

Alnus sp., and Sorbus carbonensis appear to 

represent lineages that survived in Alaska after the 

late Ravenian; during the early and middle Miocene 

(Seldovian) these lineages participated in the 

Alaskan Mixed Northern Hardwood forest, but in 

the late Miocene (Homerian) the lineages had 

adapted to a predominately coniferous forest. One 

lineage has since become extinct, but the lineages of 

Alnus are still represented in the extant Boreal 

forest. Other genera, as noted previously, are 

unrepresented in either the Neogene or extant Mixed 

Mesophytic forest, for example, Plafkeria, Eugenia?, 

and the genus represented by Dicotylophyllum 

carbonensis; Plafkeria, however, was a member of 

broad-leaved deciduous forest in the Pacific Northwest during the late Oligocene, but after that the 

lineage became extinct. In lineages in which the 

fossil record is even moderately known, it is apparent 

that vegetational types have undergone much 

floristic change during the Tertiary.

The particular forest represented by the Alaskan 

upper Ravenian probably is not the lineal predecessor of the extant Mixed Mesophytic forest. As 

shown later, the younger (Kummerian) assemblages 

indicate a return to at least subtropical conditions; 

what became of the forest zone represented by the 

upper Ravenian is not clear. This zone was probably 

restricted to cool upland areas; in conterminous 

United States, early Oligocene assemblages from the 

northern Rocky Mountains such as the Ruby 

(Becker, 1961) are vegetationally similar to upper 

Ravenian assemblages that apparently grew at 

moderate elevations in the Pacific Northwest. How 

extensive these cool upland areas were during the 

early Oligocene is unknown.

KUMMERIAN STAGE

FLORISTIC ANALYSIS

Systematic list of the Alaskan Kummerian (lower 

Oligocene) flora

Filicineae 

Filicales 

Schizaeaceae

Anemia? sp. 

Blechnaceae

Woodwardiq sp. 

Angiospermae 

Dicotyledones 

Fagales 

Betulaceae

Alnus aff. A. martini Wolfe 

Betula sp. 

Fagaceae

Fagus sp. 

Ranales 

Tetracentraceae

Tetracentron cf. T. piperoides (Lesq.) Wolfe 

Magnoliaceae

Magnolia reticulata Chan. and Sanb. 

Menispermaceae 

Hypserpa? sp. 

Lauraceae

Litseaphyllum katallensis Wolfe 

Litseaphyllum presanguinea (Chan. and

Sanb.) Wolfe

Litseaphyllum presamarensis (Sanb.) Wolfe 

Litseaphyllum raminervum Potb. 

Litseaphyllum carbonensis Wolfe 

Litseaphyllum sp. 

Litseaphyllum sp. 

Resales

Platanaceae

Platanus comstocki (Sanb.) Wolfe 

Sapindales 

Sapindaceae

Allophylus wilsoni Chan. and Sanb. 

Tubiflorae 

Verbenaceae

Holmskioldia speiri (Lesq.) MacG.? 

Incertae sedis

Macclintockia pugetensis Wolfe 

Artocarpoides kummerensis Wolfe

The Alaskan Kummerian assemblages, although 

not diverse, give clear evidence of old-world 

affinities. All the laurels represent foliar types 

exemplified by the largely paleotropical Lindera, 

Litsea, and Cryptocarya. Other paleotropical elements include Hypserpa? and Holmskioldia? The 

Alaskan Kummerian has stronger relationships to 

the subtropical forest than to the Paratropical Rain 

forest; as noted in the discussion of the middle 

Ravenian assemblages, Lauraceae have their great-
FLORISTIC AND VEGETATIONAL ANALYSIS 35

est abundance and diversity in the subtropical 

forests.

The historical floristic relationships of the 

Alaskan Kummerian assemblages are somewhat 

obscure; this is largely the result of the high 

representation of Lauraceae (90 percent of the 

specimens at loc. 11169 represent Lauraceae) a 

family in which foliar taxonomy is difficult to 

determine and of the small size of the assemblages. 

Some Kummerian laurels, for example Litseaphyllum presamarensis, and Tetracentron appear 

to represent species known in the lower and middle 

Ravenian; whether such lineages adapted to the cool 

late Ravenian climate and were minor elements of 

the Alaskan vegetation at that time is unknown. It is 

equally possible that such widespread lineages were 

totally eliminated from Alaska during the late 

Ravenian and spread northward again during the 

Kummerian. Lacking more information, I assume 

that many Alaskan Kummerian lineages followed 

this pattern.

Some lineages, for example those represented by 

the Betula and the Fagus, are not known in the 

subtropical and Paratropical Rain forests during the 

middle Ravenian and earlier times, either in Alaska 

or the Pacific Northwest. Both genera are represented in the Clarno nut bed assemblage (Scott in 

Chandler, 1964, p. 58), which represents Paratropical 

Rain forest. Both genera, however, are also 

represented in the temperate forest of the upper 

Ravenian; Betula is known in both the John Day 

Gulch and Republic assemblages, and Fagus is 

known in the presumably upper Ravenian assemblage from Joseph Creek, British Columbia. Both 

genera could presumably have been represented in 

the Alaskan upper Ravenian, and the Kummerian 

representatives could belong to the same lineages.

The floristic relationships of the Alaskan Kummerian to the Kummerian assemblages from the 

Pacific Northwest are strong; of the 12 specifically 

determined Alaskan species, 10 are known in the 

Kummerian of the Pacific Northwest. Alaska again 

was part of the same floristic province as the lowland 

Pacific Northwest, as it was in the early and middle 

Ravenian.

VEGETATIONAL ANALYSIS

The leaf-margin percentage for the Alaskan 

Kummerian assemblage is 55; this figure is based on 

the items in the systematic list plus nine fragmentary or poorly preserved, indetermined entities. 

The Kummerian percentage is the same as that for 

the middle Ravenian and is indicative of subtropical 

forest. Most species have coriaceous leaves, thus

indicating an evergreen habit. The texture of the 

leaves in continental deposits (loc. 11169, for 

example) is predominantly coriaceous as it is at 

localities in marine rocks (loc. 11168, for exam pie), in 

which coriaceous leaves would presumably have a 

higher probability of being preserved. The size of the 

leaves is small in the collections from marine rocks; 

the small size is probably a result of destruction of 

larger leaves in marine currents. Most of the leaves 

do not have pronounced drip-tips, although such tips 

are present on at least two species of Lauraceae. The 

liane element is poorly represented, the Hypserpa? 

sp. being the only probable representative. The 

vegetational analysis thus corroborates the floristic 

analysis in indicating that the Kummerian assemblage represents subtropical forest.

The Alaskan Kummerian assemblages, although 

indicating a strong floristic relationship to the 

correlative assemblages in the Pacific Northwest, 

apparently indicate a cooler climate. The Clarno nut 

bed and the Comstock assemblages of Oregon, and 

the LaPorte assemblage of northern California 

contain numerous representatives of Menispermaceae, Annonaceae, Icacinaceae, and Dipterocarpaceae. Leaf-margin percentages range from 64 to 67, 

indicating definite Paratropical Rain forest.

An interesting aspect of the Alaskan Kummerian 

assemblages is the presence of a probable strand 

element. The extinct genus Macclintockia seems to 

have been a strand plant. Although the genus is 

represented in nonmarine rocks of the Puget Group, 

the sea was presumably not far distant from the 

Puget basin of deposition. The common occurrence of 

Macclintockia is, however, in marine rocks or 

nonmarine rocks that interfinger a short distance 

laterally with marine rocks. The genus is known in 

such situations at three localities in the Pacific 

Northwest and at 13 localities in the Alaskan 

Kummerian. Koch (1963) has noted a similar 

situation for the Paleocene species of Macclintockia. 

Whatever family Macclintockia represents, it seems 

to have favored a lowland habitat near the sea. 

Another possible strand plant is Allophylus wilsoni. 

The occurrence of this species in the postKummerian of Oregon (Chaney and Sanborn, 1933) 

is in rocks that were probably laid down near the 

margin of a withdrawing embayment (Yokes and 

Snavely, 1948), and the five Alaskan occurrences 

all with Macclintockia are in marine rocks.

CLIMATIC SUMMARY

The early Ravenian (late, middle Eocene) vegetation of Alaska, as interpreted here, indicates that 

this was the warmest known period in the Alaskan
36 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Tertiary; comparisons cannot be made, however, to 

the earlier Eocene, because assemblages of that age 

are not yet known in Alaska. The climate during the 

early Ravenian is indicated to have been paratropical, that is, it was characterized by a probable 

lack of frost, a mean annual temperature of 20°- 

25°C, abundant precipitation throughout the year, 

and little seasonality of temperature or precipitation. 

The mean temperature of the coldest month could 

have been as low as about 13°C, and thus the climate 

would, in that case, not be considered to be strictly 

tropical in the sense this term is used by most 

climatologists (see p. 24).

The middle Ravenian (early late Eocene) assemblages indicate a somewhat cooler climate, probably 

in the warmer range of subtropical climates. Frosts 

occurred, although they may have been mild and of 

short duration. Extant vegetation similar to that of 

the middle Ravenian lives under climates in which 

the temperature drops as low as -14°C, the mean 

annual temperature ranges from 13° to 20°C, and the 

mean temperature of the coldest month varies from 

1° to 18° C. Precipitation would again have been 

abundant and distributed throughout the year.

The late Ravenian (latest Eocene) assemblage 

indicates considerably cooler temperatures than 

either the early or middle Ravenian. The climate was 

temperate, that is, temperatures probably at least as 

low as -14°C, a mean annual temperature in the 

range of 11° to 13°C, and the mean temperature of 

the coldest month in the range of -3° to 2°C.

By the Kummerian (early Oligocene), climatic 

conditions were again subtropical. The climate was 

probably about the same as that of the middle 

Ravenian. The probable post-Kummerian assemblages from Admiralty Island in southeastern 

Alaska indicate a subtropical climate which persisted into the middle Oligocene, but at the end of 

that time another major deterioration occurred that 

resulted in the cool temperate climate of the 

Angoonian cooler than even the late Ravenian. 

During the warm periods of the early and middle 

Ravenian and Kummerian, the vegetation and 

hence the climate showed little latitudinal zonation 

in comparison to the cool periods of the late 

Ravenian and the Angoonian, when considerable 

zonation of vegetation is apparent.

ALASKAN PALEOGENE CLIMATES

The salient feature of the Alaskan lower and 

middle Ravenian and Kummerian assemblages is 

that they are broad-leaved evergreen forest; from the 

standpoint of foliar physiognomy and floristic

analysis the lower Ravenian vegetation is similar to 

the extant Paratropical Rain and Submontane Rain 

forests. The assumption is here made, that the 

Alaskan Paratropical Rain forest (the geographic 

and topographic position demand this appellation) 

grew under climatic conditions common to these two 

extant forest types. Such conditions are:

(1) a lack of frost,

(2) a mean annual temperature of 20°-25°C,

(3) abundant precipitation, and

(4) no pronounced dry season.

Compare these conditions with the climatic conditions at Cordova in the Alaska region today:

(1) minimal temperature -28°C,

(2) a mean annual temperature of 5°C,

(3) annual precipitation of 3,700 mm, and

(4) precipitation of driest month 154 mm. 

In regard to precipitation, no change is needed; the 

Paratropical Rain forest exists under less precipitation, both yearly and dry month, than Cordova 

receives today. Clearly, however, the temperature 

regime during the early Ravenian in Alaska must 

have been very different than it is today.

The possibility that the Alaskan Paleogene 

assemblages described in this report may not be at 

the latitude at which the enclosing beds were laid 

down is an important consideration. Some proponents of large-scale continental drift during the 

Tertiary have in fact suggested that the warm 

character of the Gulf of Alaska Eocene floras 

indicates that they lived at more southerly latitudes 

and the containing rocks have since drifted 

northward (Hamilton, 1968). The movement to the 

northwest of the northeastern Pacific plate has been 

apparently about 2-3 cm per year, but this movement 

is thought to have started only about 8-10 million 

years ago (Vine, 1966; Tobin and Sykes, 1968). More 

significant is that, insofar as present geologic 

evidence indicates, the Tertiary rocks of the Katalla 

and Yakataga districts are thought to be part of the 

upper thrust plate, that is, the rocks were about at 

their present geographic position 10 million years 

ago. Whether 35 million years previous to that the 

rocks were at about their present position is 

unknown. Grantz (1965), however, suggests that, 

during the earlier Tertiary, rotation of the active 

west limb of Carey's (1958) postulated orocline (the 

Katalla and Yakataga districts are on the east limb) 

brought western Alaska into a more southerly 

position relative to the east limb; the east limb of the 

postulated orocline appears to have been relatively 

stable during the rotation. What is significant here is 

that the presently available evidence of large-scale 

earth movements during the Tertiary indicates that
HISTORY OF THE ARCTO-TERTIARY CONCEPT 37

the Eocene rocks of the Gulf of Alaska district are at 

approximately the same latitude at which they were 

deposited.

The assumption has generally been made that 

paleomagnetic data that approximately locate past 

magnetic poles also indicate the position of the 

rotational pole. Six Late Cretaceous pole positions 

have been obtained from North America; these data 

indicate that the pole was in the Chukchi Sea north 

of Bering Strait. With one exception, the Paleogene 

pole positions are located intermediate between the 

Late Cretaceous pole and the present pole. The 

pertinent paleomagnetic evidence thus indicates 

that during the Ravenian, North America was perhaps closer to the magnetic pole than at present. 

Given the basic assumption previously mentioned, 

the Alaskan Ravenian vegetation grew at latitude 

60° or somewhat closer poleward.

A problem that must be discussed with that of 

temperature is light. Mason (1947, p. 206) has pointed 

out that under the present distribution of land 

masses relative to the poles and the tilted relation of 

the axis of rotation to the earth's orbit, no broadleaved evergreen forest could possibly exist at high 

latitudes:

To raise the temperature in such an area and not change the long 

periods of darkness would only aggravate the situation because 

the increased temperatures would increase the respiration rate to 

the point that in evergreen species the reserve food supply would 

soon be depleted***

Dicotyledons have adapted to polar nights in three 

ways: as low evergreen shrubs that have small 

leaves, as annual herbaceous forms, and as 

deciduous trees or shrubs. Evergreen plants have, of 

course, adapted to long polar nights. The leaves of 

such plants are, however, of extremely small size in 

comparison to the probable evergreen species of the 

early Ravenian. The early Ravenian species almost 

certainly belong to neither of the first two categories. 

As mentioned previously, the evergreen nature of 

most of the species is attested to by the coriaceous 

texture of their leaves; a deciduous palm, moreover, 

would be a most curious sight. The idea that broadleaved evergreen plants in the mesophyll and 

megaphyll size classes could become dormant during 

winters in the Arctic is almost certainly invalid; the 

broad-leaved evergreen species such as palms are 

limited by the lowest temperature at which 

photosynthesis can be carried on, that is, the plants 

must carry on life functions at all times. Also 

arguing against dormancy is the fact that lineages 

representing many different orders, families, and 

genera would have had to adapt to such conditions. 

Natural selection should yield a forest of deciduous

plants at high latitudes if the climate were warmer 

but the inclination of the earth's axis were the same 

as at present because these are the plants best able to 

become dormant for extended periods.

Some tropical plants are capable of attaining 

dormancy under short days, as shown by the 

experiments of Peacock and McMillan (1968) on 

Prosopis and Acacia. The experimental conditions 

these investigators used, however, were not comparable to the present light conditions at latitude 60°, 

because the shortest day period in the experiments 

was 8 hours (compared to 5 hours at latitude 60°), and 

the light intensity was considerably higher than the 

winter light intensity at 60°; note also that the 

species investigated are deciduous, and dormancy 

mechanisms are well developed in deciduous plants. 

Although experimental work on tropical, broadleaved evergreen plants needs to be done, I think that 

Mason's (1947) point is well taken: the high 

temperatures at high latitudes during the Paleogene 

would, if the light regime was that of today, not have 

allowed the existence there of a mesophyllous, broadleaved evergreen forest.

If, as seems probable, the Alaskan early Ravenian 

assemblages lived approximately at latitude 60° N., 

if, as has been shown previously, they represent a 

mesophyllous broad-leved evergreen forest, and if, as 

what is known of plant physiology indicates, such a 

forest could not live under the present light 

conditions at latitude 60° N., then the light conditions during the early Ravenian must have been 

significantly different from today. This line of 

reasoning leads me to the conclusion that the axis of 

rotation of the earth may have had much less 

possibly no inclination.

HISTORY OF THE ARCTO-TERTIARY CONCEPT

The Arcto-Tertiary concept has been the primary 

contribution of paleobotany to the field of plant 

geography. The concept fundamentally envisions 

that a broad-leaved deciduous forest of a particular 

floristic composition evolved in the Arctic during the 

Cretaceous, persisted there during the early Tertiary, 

and, in response to gradually cooling climate, 

migrated southward during the middle Tertiary into 

middle latitudes where the forest persists today in 

east-central Asia and southeastern North America. 

During its migration, the Arcto-Tertiary flora (or 

Geoflora) supposedly underwent little change in 

floristic composition. The fundamental significance 

of this concept to plant geography is clear; the 

concept is accepted as fact in standard textbooks on 

plant geography such as Cain (1944) and Good 

(1953). Good, in fact, bases one of his six principles of
38 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

his theory of tolerance on the geofloral concept (1953, 

p. 361): "Great movements of species and of floras 

have taken place in the past and are apparently still 

continuing." As outlined in the present report and 

elsewhere (Wolfe, 1972), the concept of an ArctoTertiary Geoflora is not supported by our current 

knowledge of the Alaskan Tertiary vegetational and 

floristic history. I will, therefore, examine in 

considerable detail the history of the Arcto-Tertiary 

concept; it is, I think, interesting to consider how 

such a concept approximately 100 years old could 

gain such wide acceptance as fact.

The concept of homotaxis is the fundamental 

concept of which the Arcto-Tertiary and other 

geofloral theories are simply applications. In 

enunciating his theory of homotaxis, Thomas 

Huxley stated (1870, p. xlii): "It is possible that 

similar, or even identical, faunae and florae in two 

different localities may be of extremely different 

ages***" This fashionable theory was then adopted 

by two men the paleobotanist Gardner and the 

neobotanist Gray and the adoption had a profound 

effect on botanical thinking for the next 90 years.

Gray's adoption of homotaxis was an effort to 

explain the close floristic relationship between 

temperate eastern Asia especially Japan and 

temperate eastern North America. Gray (1878, p. 192- 

-193) remarked:

The [polar] lands***were once the home of those trees, where they 

flourished in a temperate climate. The cold period which followed, 

and which doubtless came on by very slow degrees during ages of 

time, must [have] long before its culmination have brought down 

to our latitudes, with the similar climate, the forest they possess 

now, or rather the ancestors of it***Geologists give the same name 

[Miocene Tertiary] to these beds, in Greenland and Southern 

Europe, because they contain the remains of identical and very 

similar species of plants; and they used to regard them as of the 

same age on account of this identity. But in fact this identity is 

good evidence that they cannot be synchronous. The beds in the 

lower latitudes must be later, and were forming when Greenland 

probably had very nearly the climate which it has now.

Huxley's "possible" was transferred to "good 

evidence" without any additional data. Note also 

that Gray's attitude involves little or no evolution of 

the plant lineages during their supposed migrations. 

Gardner's writings have similarly affected paleobotanical thought. Discussing the Tertiary, and 

especially the Eocene, floras of Europe, Gardner (in 

Gardner and Ettingshausen, 1879, p. 8) stated:

There is no great break in passing from one to the other [Eocene to 

Miocene floras] when we compare them over many latitudes, and 

but little change beyond that brought about by altered 

temperature or migration. But if Tertiary floras of different ages 

are met with in one area, great changes on the contrary are seen, 

and these are mainly due to progressive modifications in climate, 

and to altered distribution of land. Imperceptibly, too, the tropical 

members of the flora disappeared; that is to say, they migrated, for

most of their types, I think, actually survive at the present day, 

many but slightly altered. Then the subtropical members 

decreased, and the temperate forms, never quite absent even in the 

Middle Eocenes, preponderated. As decreasing temperature drove 

the tropical forms south, the more northern must have pushed, 

from their home in the far north, more and more south as climates 

chilled, and at last, in the Miocene time, occupied our latitudes. 

The relative preponderance of these elements, I believe, will assist 

in determining the age of Tertiary deposits in Europe more than 

any minute comparison of species. Thus it is useless to seek in the 

Arctic regions for Eocene floras, as we know them in our latitudes, 

for during the Tertiary period the climate conditions of the earth 

did not permit their growth there. Arctic floras of temperate, and 

therefore Miocene aspect, are in all probability of Eocene age, and 

what has been recognized in them as newer or Miocene facies is 

due to their having been first studied in Europe in latitudes which 

only became fitted for them in Miocene times.

Gardner, as did Gray, thus had a concept of static 

composition of plant communities and of little 

morphologic evolution. Gardner's use, however, of 

"in all probability" indicates that he did not at that 

time consider the theory proved. Four years later, 

Gardner (1883, p. 3) still adhered to homotaxis:

It is obvious***that floras from the south of England, supposed to 

be of the same age, are too widely separated in latitude from those 

of Greenland to permit any useful comparison being made directly 

between their species.

Gardner was also the first paleobotanist to direct 

himself to the significant problem of what flora or 

vegetation existed in the north when temperate 

floras were present at middle latitudes:

***the plant-beds below the London Clay [Paleocene of today] are 

filled with Plane and forms resembling Hazel and Lime, which 

represent a climate so nearly approximating to that of these 

latitudes at the present day that no forest of leafy trees could 

possibly have co-existed with them in the Arctic Circle (Gardner, 

1883, p. 7).

Four years later, however, Gardner (1887a, p. 299) 

made a statement that the proponents of the ArctoTertiary Geoflora have curiously neglected:

These determinations [of Miocene species in the Upper Atanekerdluk flora of Greenland] are for the most part based upon specimens 

which I regard, under any circumstances, as too fragmentary to be 

of any value***it would***have been just as easy to have 

identified them with either Cretaceous, Eocene, or living species 

as with Miocene. It is not a little significant that none of the finer 

and most distinctive plants of the Mull, the Antrim, or the 

Atanekerdluk floras in question were, or could be, identified with 

Miocene species, but were all admitted to be peculiar. Among 

these, we have the Pterospermites***from Atanekerdluk*** 

recalling the larger leaves of Sezanne [a flora of the type section of 

the Paleocene]***But the best marked and most thoroughly 

characteristic leaves of the whole are the Daphnogene Kanii, Hr., 

and the MacClintockia trinervis, Hr., forms which are equally 

represented in the Thanet flora of Gelinden [a Belgian Paleocene 

flora] and in the Antrim floras***they evidently existed in one 

definite stage and no other, and afford paleontological evidence 

that should be conclusive.

This statement clearly contradicts Gardner's previous ideas and by inference brings into question the 

concept of homotaxis as applied to floras. Gardner
HISTORY OF THE ARCTO-TERTIARY CONCEPT 39

was correlating floras because of similar or identical 

species, that is, he was using the same fundamental 

basis for correlation as did Oswald Heer, but he 

disagreed only with Heer's taxonomy. Note that 

Gardner did not consider any species from the 

Atanekerdluk to be Miocene species. Gardner had 

apparently arrived at the conclusion that "minute 

comparison of species" was, after all, the basis for 

paleobotany.

Not only had Gardner decided that European 

Paleogene plant assemblages could be correlated 

with high-latitude assemblages by means of standard biostratigraphic methods, he also thought that 

some assemblages described by Heer from the Arctic 

were as Heer thought of Neogene age. "The first 

acid eruptions were Miocene, as shown by the floras 

preserved in Iceland." (Gardner, 1887b, p. 91). Not 

only has Gardner's reversal in his theories been 

overlooked (or ignored) by the proponents of the 

Arcto-Tertiary concept, even Friedrich (1966), in his 

revision of the flora from one of the classic Icelandic 

Tertiary localities, has apparently been unaware of 

Gardner's opinion on the Icelandic assemblages.

The term "arcto-tertiary" was first coined by 

Engler (1882) as the "arcto-tertiare Element." Engler 

(1882, p. 327) stated (freely translated) that this 

element was "***distinguished by numerous conifers and the numerous genera of trees and shrubs 

that now dominate in North America or in 

extratropical east Asia and in Europe." Engler (1879, 

p. 2-3) was aware that Gardner considered at least' 

some of the arctic Tertiary assemblages to be of 

Eocene age; Engler, however, accepted Heer's age 

assignment of the "Arctic Miocene" as valid, and 

noted that the "Arctic Miocene" floras differed 

sufficiently from the subtropical floras of the 

southern European Miocene for the "Arctic Miocene" 

of Heer and the southern European Miocene to be 

isochronous. Engler's hypothesis has, in large part, 

been substantiated by the paleobotanical research of 

this decade.

The American paleobotanist J. S. Newberry, 

although accepting Gardner's 1879 determination of 

the Atanekerdluk flora as Eocene, did not accept the 

original homotaxial basis for that determination. 

Newberry (1890, p. 8) stated:

"I called the Fort Union Group Miocene because I identified it with 

the plant-bearing beds of MacKenzie River, Disco Island, 

Greenland, etc.***This flora***has since been shown by Mr. J. 

Starkie Gardner to be Eocene. The Fort Union flora has many 

species in common with the Eocene beds of the Island of Mull, 

Bournemouth, etc., and holds undoubtedly the same position."

Newberry's basis for correlation was clearly not any 

concept of homotaxis but rather was on the 

community of species between any two floras. The

Canadian Sir William Dawson (1888), who had read 

Gardner's (1887a) paper cited above, clearly concurred with Gardner. Dawson (p. 36) further 

discussed the several species that were common to 

various floras of middle and north latitudes that we 

now, for the most part, call Paleocene. Dawson, 

however, considered that some of the typical 

Paleocene species occurred in the Alaskan Tertiary 

flora described by Heer (1869) and thus he also 

thought the Alaskan Tertiary flora to be "Lower 

Eocene" or Paleocene.

Dawson's treatment was followed by that of F. H. 

Knowlton, who was the next paleobotanist to work 

on Alaskan Tertiary plants. Knowlton (1894, p. 238) 

remarked that "***the fossil flora of Alaska is 

inseparably connected with that of the Disco Island 

and Atanekerdluk beds of Greenland and the socalled Arctic Miocene of Spitzbergen and Sachalin. 

Whatever is decided concerning them must apply 

with equal force to Alaska." Knowlton's basis for 

considering the Alaskan Tertiary plants to be of 

Eocene age was thus more the fact that he thought 

they represented the same species that were also 

found in other "Eocene" (now Paleocene) floras both 

at high and middle latitudes. Knowlton (1894, p. 238), 

however, apparently still considered homotaxis to be 

probable, because he quoted Gardner (1879, p. 8) and 

remarked "This change of view as to the age of the socalled Arctic Miocene, as proposed by Gardner, has 

already received considerable confirmation from 

American paleobotanists, and while it can hardly be 

regarded as settled, it may be accepted as extremely 

probable." Thus, while stating a belief in the 

significance of homotaxis, Knowlton in practice 

rejected such a concept for purposes of correlation.

Some substantiating data for the "Eocene" age of 

the Alaskan Tertiary plants was apparently given 

by supposed stratigraphic relationships of some of 

the plant beds to beds containing marine mollusks. 

On the Alaska Peninsula, plants were collected from 

beds that were thought by Dall and Harris (1892, p. 

251) to underlie conformably Miocene marine beds 

(the term Oligocene was not then used by Dall). 

Dall and Knowlton, moreover, based the assignment 

of the Alaskan Tertiary plants to the Eocene on two 

assumptions: (1) that all plant-bearing beds in 

Alaska were essentially synchronous and were part 

of their Kenai Group, and (2) that the marine 

lithologic unit on the Alaska Peninsula was the same 

unit as that found at Astoria, Oregon, some 

thousands of miles distant. Later Knowlton (in Dall, 

1896, p. 842) concluded that these stratigraphic 

relationships were valid and that "***it would seem 

that we are justified in referring the Kenai group to
40 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

the horizon of the Oligocene of European geologists." 

Still later, however, the Kenai Group (i.e., all plantbearing beds in Alaska) was assigned to the upper 

Eocene. This assignment was based on further 

collections of marine mollusks from the Alaska 

Peninsula and their supposed stratigraphic relationships to fossil plants. Beds containing the 

mollusks were assigned by the geologist Atwood 

(1911) to the "Kenai group," and the mollusks were 

determined by Ball (1904 and in Atwood, 1911, p. 55) 

to be of Eocene age. Atwood (1911, p. 58) further 

stated that the invertebrates occurred "***with 

typical Kenai plants, in some places on the same slab 

of rock***" although none of the fossil plant or 

mollusk localities as given by Atwood were identical. 

The next paleobotanist to work on the Alaskan 

tertiary plants was Arthur Rollick. He (in Martin 

and others, 1915, p. 88), in discussing the age of the 

Kenai Formation proper of the Cook Inlet region, 

considered the Kenai to be of Eocene age because 

"***the so-called 'Arctic Miocene' of the north 

circumpolar region*** [is] now generally recognized 

as Eocene in age." The Kenai in particular was of 

late Eocene age because some of the species were 

found in the Fort Union flora and some in Oligocene 

and Miocene floras, and thus a late Eocene age was a 

compromise. Later, however, Rollick decided that 

the closest relationship of the Alaskan Tertiary 

plants was with the Fort Union flora; he listed 12 

Alaskan species that were "***characteristic or 

index fossils of the Fort Union formation." (Rollick, 

1936, p. 22). Moreover, "In view of the facts above set 

forth the general similarity of the Alaska Tertiary 

flora to that of the Eocene in the States proper would 

appear to be demonstrated***" (Rollick, 1936, p. 23). 

Curiously, Rollick's thinking had the same internal 

logical schism as did Knowlton, because Rollick 

went on to quote Gardner's (in Gardner and 

Ettinghausen, 1879) argument for floral homotaxis. 

Also curious is the fact that Rollick listed about 30 

species from rocks that the geologist P. S. Smith (in 

Rollick, 1936, p. 26) was "reasonably certain" on 

geologic and molluscan evidence were of "postEocene" age. Rollick (1936, p. 12) did not try to 

determine whether several different ages were 

represented in his collections; he considered that the 

province of the geologists. The failure of Rollick and 

most other paleobotanists to attempt to differentiate 

the various fossil plant assemblages from Alaska or 

elsewhere in the Arctic is puzzling. Perhaps the most 

astute observation was that of Chaney (1927, p. 98): 

"The Arctic floras of Greenland and Spitzbergen 

have been called Miocene in age, but it seems 

probable that***several horizons are represented, of

which the oldest may be Eocene or Oligocene."

After publication of Rollick's work, there was no 

futher research on Alaskan Tertiary floras until the 

present decade. Most Arctic Tertiary floras, indeed, 

were neglected until about 1955. In the meantime, the 

Arcto-Tertiary concept of homotaxis was almost 

unchallenged. Published originally as a theory, it 

became increasingly accepted as a fact by almost all 

paleobotanists (Krystofovich, 1929; Berry, 1930b; 

Chaney, 1936). The most thorough discussion of the 

Arcto-Tertiary theory was by Chaney (1936; 1938; 

1940), who remarked (1940, p. 474): "Only the 

evidence of their stratigraphic occurrence has 

subsequently demonstrated that the 'Arctic Miocene' 

flora of Heer comes from lower levels than the Bridge 

Creek and other Middle Tertiary floras from 

temperate latitudes." What, in fact, was the 

"evidence"? From the preceding discussion, in 1940 

only two floras in Alaska were known to have some 

independent evidence as to age. One of these floras, 

from the Alaska Peninsula, was considered to be 

Eocene, but the other, from the Bering River coal 

field (Berg Lake flora of Chaney), was considered to 

be Oligocene or Miocene. Each of these two floras 

contained about 30 species. These data indicate that 

the "evidence of their stratigraphic occurrence" was, 

in fact, contradictory.

Chaney's concept of the Arcto-Tertiary Geoflora 

clearly has its roots in Clementsian ecology: "The 

differentiation of Tertiary vegetation into a series of 

climaxes, or a clisere, in response to differences in 

latitude and altitude was suggested by Clements 

almost twenty years ago" (Chaney, 1936, p. 62). The 

Clementsian ecology was coupled with Huxley's 

homotaxis: "It might even be said that if a flora from 

Oregon was closely similar in composition to one 

from Alaska, the age of the two must be different" 

(Chaney, 1936, p. 319). The confusion of vegetation 

and flora is clearly apparent in the more sophisticated Arcto-Tertiary concept of Chaney: "A separation in latitude of 20 degrees may be expected to have 

involved, in the past as at the present time, the 

development of plant assemblages of wholly 

distinctive composition" (Chaney, 1936, p. 319). It is, 

of course, true that there is some vegetational 

difference between the coastal conifer forest of, for 

example, Cordova, and the lowland forest of the 

Pacific Northwest today; floristically, however, the 

difference is much less. Approximately 70 percent of 

the species of woody plants near Cordova extend 

southward into the Pacific Northwest and about 

40 percent are found at low altitudes. The present 

strong latitudinal zonation of vegetation has not 

only been extended back in time by the proponents of
HISTORY OF THE ARCTO-TERTIARY CONCEPT 41

the Arcto-Tertiary concept, but they have also 

equated erroneously I think vegetational zonation with floristic zonation.

One of the main points of Chaney's (1940) paper 

was that the Arctic Tertiary floras showed clear 

evidence of climatic zonation and that no subtropical 

floras had been discovered there. A curious aspect of 

the development of the theory of an Arcto-Tertiary 

Geoflora is the vegetational treatment by the 

proponents of this theory of the so-called Berg Lake 

flora. This assemblage (loc. 3847) was originally 

published by Rollick, who listed 18 species (Rollick, 

1936, p. 15). Although temperate genera were listed 

(Populus, Juglans, Planera, Ulmus, Rhamnus, 

Cornus, Rhododendron, and Fraxinus), an equal 

number of tropical elements were listed (Artocarpidium, Mohrodendron, Magnolia, Cinnamomum, Persea, Malpoenna, Terminalia, and Semecarpus). Without any revision of Rollick's taxonomy, 

Chaney (1940) considered the Berg Lake flora to be 

temperate, and he reiterated the same opinion later 

(Chaney, 1964). The simple utilization of leaf margin 

percentages (71 percent of Rollick's "species" have 

entire margins) would have indicated that this 

assemblage could not be temperate. Note that the 

systematic revision of Rollick's illustrated Berg 

Lake material indicates the presence of: Dryopteris, 

Platycarya, Alnus, Knema, Myristica, Cinnamomophyllum, Luuunga, Melanorrhoea, Celastrus, 

Parashorea, and Alangium. Even without additional collections, it should have been apparent that 

the Berg Lake assemblage floristically was closely 

allied to the modern tropical flora. Further, the 

"climatic zonation" was based on "Eocene" floras 

that were either small or of other ages. Three 

examples will illustrate this point: (1) The "temperate" Kobuk River flora was known to contain 

only one species, a Gm&go(Knowltonin Mendenhall, 

1902) one species is hardly sufficient to determine 

climate or age; (2) the "temperate Eocene" Norton 

Sound assemblage had been shown to be of medial 

Cretaceous age (Martin in Rollick, 1930, p. 21); (3) the 

"Eocene" Simonova flora was thought to be of Late 

Cretaceous age (Krystofovich, 1929, p. 304). The 

supposed climatic zonation thus loses its validity, at 

least in regard to high latitude floras as they were 

then known.

The supposed strong resemblance mentioned by 

Chaney (1936) between middle Tertiary floras such 

as the Bridge Creek and the "Kenai flora" has never 

been documented. Only five species have ever been 

listed (Chaney, 1952, p. 107) as conspecific between 

Alaskan "Eocene" and the Bridge Creek; this 

number of species is not impressive in view of the fact

that Rollick listed over 300 species from the Alaskan 

Tertiary.

Opposition to the Arcto-Tertiary concept began on 

the theoretical level in 1947. Mason (1947, p. 205) 

argued:

It is difficult***to envisage such floristics as an Arcto-Tertiary 

flora (Chaney, 1936) in contrast to a Madro-Tertiary flora 

(Axelrod, Mss.) as accounting for floristic sources and centers of 

origin during Tertiary time. Such concepts of floristic organization and development demand unity and stability of 

communities in time and space beyond what is possible in the 

light of the nature of floristic dynamics such as are bound up with 

the genetics of the population, the physiology of the individual, 

and the diversity and fluctuation of the environment. 

Mason's reasoning was apparently ignored by 

proponents of the Arcto-Tertiary concept. Preliminary studies of the Alaskan Tertiary floras 

indicated that the Arcto-Tertiary concept was 

invalid. We (MacNeil and others, 1961, p. 1801) 

stated that:

"***evaluation by Wolfe of new and old plant collections from 

Alaskan rocks whose age is determined by marine invertebrates 

clearly shows that floras of the same age in Alaska and in Oregon 

or Washington are similar on the specific level."

MacGinitie (1962, p. 87) "***questioned whether 

any flora, as a unit, migrated during the Tertiary*** 

The terms 'Arcto-tertiary,' 'Madro-Tertiary,' and the 

like imply extremely useful concepts if we do not 

think of these terms as representing areas or centers 

from which mass migrations occurred. They picture 

to us in a general way the vegetation occupying an 

area***." MacGinitie (1962, p. 88), moreover, 

concurred that the Port Graham assemblage (early 

Seldovian of present usage) was of earlier Miocene 

age.

Preliminary evaluation of the earlier Paleogene 

floras of Alaska (Wolfe, 1964, p. N6) indicated that 

they contained "***abundant cycads, palms, Lauraceae, Menispermaceae, Alangium, and other taxa 

indicative of subtropical if not tropical climates." 

In connection with a discussion of the Kenai 

assemblages, it was noted (Wolfe, 1966, p. B4):

The general aspect of the Chickaloon flora is so greatly dissimilar 

to the overlying Seldovian flora that it is evident that, as a floristic 

type, the Seldovian flora did not exist in the Kenai region during 

the Paleocene. The nearest well-dated Eocene floras are those of 

middle and late Eocene age at the head of the Gulf of 

Alaska***these Eocene floras look very much like those in the 

Eocene at middle latitudes.

At about the same time that revision of Alaskan 

floras was undertaken, advances were also being 

made in Siberia. Most of the Siberian work has been 

on Neogene assemblages. The classic Upper Dui 

flora of Sakhalin, for example, had been described by 

Heer (1878) as part of his "Arctic Miocene." Chaney 

(1940, 1964) had considered the Upper Dui as an 

Eocene flora, but marine mollusks indicate a middle
42 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Miocene age for this assemblage (Krystofovich, 

1960, 1964). Fotianova (1964) described a late 

Miocene assemblage from mollusk-dated rocks on 

Sakhalin, and she (1967) has also discussed the 

significant aspects of the Upper Dui assemblage. 

The Pliocene plants from the predominantly marine 

Enemtensky Formation on Kamchatka were described by Fotianova (in Sinelnikova and others, 

1967); she remarked on the vegetational similarity of 

the Enemtensky and Clamgulchian assemblages 

and the general similarity of Neogene vegetational 

history at northern latitudes.

In 1967 Chaney attempted to defend the ArctoTertiary concept; he claimed that Neogene assemblages were unknown north of Hokkaido and 

Washington. Chaney was apparently unaware of the 

Siberian paleobotanical work (his most recent 

citation concerning Siberian paleobotany is a 1934 

paper) and the stratigraphic work in Alaska 

published by Burk (1965), which included lists of 

plants from Neogene, mollusk-dated rocks.

Some workers might argue that the probable lack 

of Arctic waters pouring into the North Pacific in 

combination with a warmer Japan Current would 

lead to warmer conditions along the Alaskan coast. 

Paratropical Rain or subtropical forest could then 

extend along a narrow coastal strip, but in areas of 

interior Alaska, interior northern North America, 

and interior northern Siberia, an Arcto-Tertiary 

Geoflora could have persisted during the Paleogene. 

Such an hypothesis has no more factual foundation 

than the previously discussed "evidence" for the 

Arcto-Tertiary concept. As noted elsewhere (p. 54), a 

fan palm is known from interior Alaska (lat 65V2 0 N.); 

the age of this palm is unknown, but its occurrence 

indicates that warm conditions extended at times 

into central Alaska. Insofar as I am aware, no 

definite Eocene or Oligocene assemblages are known 

from the Canadian shield area; analysis of the little 

known Tertiary plant assemblages from the Arctic 

Islands north of Canada may shed considerable 

light on the floristic and vegetational history of that 

region, but neither stratigraphic nor paleobotanical 

evidence is sufficient at present.

The existence of Paratropical Rain forest at 

latitude 60° N. in fact probably indicates that 

temperate broad-leaved deciduous forest was not 

present in Alaska during the early Ravenian. 

Because the known early Ravenian assemblages are 

not marginally Paratropical Rain forest, they 

indicate that more of this vegetation extended north 

from the Gulf of Alaska region. North from the 

northern limit of the Paratropical Rain forest, it is 

reasonable to expect the development of subtropical

vegetation. Considering the generally slight zonation of climate during warm periods of the 

Paleogene, it is possible that all Alaska was occupied 

by Paratropical Rain and subtropical forests during 

the early Ravenian. Conceivably montane areas 

could have supported temperate vegetation, but on 

Taiwan today the Paratropical Rain forest is 

separated altitudinally from the temperate forest by 

1,500 m. Mountains of that height are not known to 

have existed in Alaska during the Paleogene. Large 

mountain masses, such as the central part of the 

Alaska Range and the Wrangell Mountains, are 

largely or entirely of later Neogene origin. The 

deposition of the Nenana Gravel marks the 

beginning of the uplift of the central part of the 

Alaska Range, and this gravel is probably of 

Pliocene age (Wahrhaftig and others, 1969). The 

Wrangell Lava, which forms the bulk of the high 

country of the Wrangell Mountains, overlies the 

Frederika Formation of Miocene age (MacKevett, 

1970). If, therefore, temperate broad-leaved deciduous forest existed during warm periods such as the 

early Ravenian, the forest would probably have 

grown as isolated pockets in montane areas of the 

northern and interior parts of the northern 

continents.

In Siberia, much the same overall vegetational 

development occurred as in Alaska. In the Paleogene: "The Eocene was the warmest epoch to occur 

during this time span. In places, the temperate forest 

zone moved northward, beyond latitude 50°-60° N. 

Proof of this shift is the fact that in the south Urals 

and in West Kazakhstan Late Cretaceous Platanaceae forests were succeeded by narrow-leaf, subtropical vegetation, including palms." (Vakhrameyev, 

1966, p. 16). The important point is that Vakhrameyev infers that temperate vegetation must have been 

present north of latitude 50°-60° N., although, 

insofar as I am aware, no temperate Eocene 

assemblages are known from Siberia. The presence 

in western Siberia of the extensive embayment that 

lasted until the early Oligocene would also have 

favored the development in Siberia of tropical or 

subtropical vegetation. The forests of Siberia during 

the Eocene and Oligocene represent, insofar as 

known, dominately broad-leaved evergreen vegetation (Iljinskaja, 1963).

Also significant is the work of Dorofeyev (1963), 

who has described large fruit and seed assemblages 

from localities that are at latitudes 55°-60° N. in 

western Siberia. The localities are in rocks that rest 

on earlier Oligocene marine rocks, and Dorofeyev 

assigns some assemblages to the Oligocene and 

some to the Neogene. The assemblages are floris-
A THEORY OF THE ORIGINS OF THE MIXED MESOPHYTIC FOREST 43

tically allied to the broad-leaved deciduous forests. 

The lack of leaf assemblages makes it necessary to 

rely on the floristic relationships to determine the 

vegetation; the occurrence of genera such as 

Magnolia, Sinomenium, and Leitneria indicates a 

warm temperate forest. The floristic relationships of 

the various assemblages indicate that Mixed 

Mesophytic forest persisted in central Eurasia at 

northern latitudes well into the Miocene, just as this 

vegetational type did in Alaska. Some genera, for 

example Magnolia, that persisted into the Miocene 

in central Eurasia are not known in the Alaskan 

Miocene; this may indicate that the warmer 

summers of an interior location allowed some genera 

that are sensitive to this parameter of tolerance to 

persist longer in the interior of continental regions. 

Dorofeyev (1963) notes that continentality increased 

from the Oligocene into the Miocene.

Also significant is the recognition by Russian 

workers of what is probably the same Oligocene 

climatic deterioration as detected in western North 

America. Zhilin (1966, p. 2 of English translation) 

remarked that the floristic data from Kazakhstan 

"***indicate a considerable difference of climate 

between the early Oligocene and the second half of 

the Oligocene in West Kazakhstan. The Early 

Oligocene climate was transitional from subtropical 

to warm temperate***The climate at the end of the 

Late Oligocene can be described as warm temperate." Zhilin, of course, did not have the radiometric 

data available that indicate that this climatic shift 

was rapid (Wolfe and Hopkins, 1967).

The large late Oligocene leaf assemblage described 

by Krystofovich (1956) and his colleagues from 

latitude 48° N. in Kazakhstan also indicates that the 

interior vegetation was not significantly different 

from the Alaskan vegetation at that time. The leafmargin percentage for this Eurasian assemblage is 

18 as compared to 6-22 for the Alaskan late 

Oligocene assemblages. If, during the late Oligocene 

and most of the Miocene, continentality did not play 

a significant role in the differentiation of vegetation, 

I do not think continentality was a significant factor 

during the early Paleogene. During the Paleogene 

neither latitude nor continentality seem to have been 

as significant in the differentiation of vegetation as 

they are today.

The conclusion is, I think, inescapable that the 

Arcto-Tertiary concept has never had a satisfactory 

stratigraphic foundation. The older theories of the 

relationships between plants and their environments allowed, if not demanded, the acceptance of a 

Arcto-Tertiary Geoflora. As knowledge of genetics 

and physiology increased, it should have been

apparent, as Mason (1947) pointed out, that the 

Arcto-Tertiary concept was invalid. The discarding 

of this concept, which is indicated by the Alaskan 

and Siberian assemblages of fossil plants, is 

fundamental to an understanding of floristic and 

vegetational history.

A THEORY OF THE ORIGINS OF THE MIXED 

MESOPHYTIC FOREST

The Mixed Mesophytic forest is a complex floristic 

and vegetational type (Wang, 1961). The vegetation, 

which is characteristic of a warm temperate climate, 

is floristically diverse, notably in families such as 

Salicaceae, Juglandaceae, Betulaceae, Fagaceae 

(including the "lobed" oaks), Ulmaceae, Hamamelidaceae, Rosaceae, Aceraceae, Rhamnaceae, and 

Ericaceae. Notable also is the presence of numerous 

monotypic and oligotypic genera, for example, the 

woody ranalean Cercidiphyllum, Euptelea, 

Trochodendron, and Tetracentron. No single genus 

or family dominates the canopy of the forest; next to 

the tropical forest types, the Mixed Mesophytic forest 

is the most complex and most diverse vegetational 

type extant.

The origins of this forest have attracted the 

attentions of both botanists and paleobotanists for 

over a century. As noted in the preceding section of 

this report, the widely accepted explanation of the 

origin of this forest the concept of an ArctoTertiary Geoflora is contradicted by what is known 

of the Tertiary stratigraphy at high northern 

latitudes (see previous sections). The vegetational 

and floristic history of the Northern Hemisphere has 

been confused in large part because of the apparent 

confusion by many paleobotanists of vegetation and 

flora (see also p. 40). This confusion is inherent in 

such a concept as a "Geoflora," which includes both 

taxonomic composition and vegetational type. 

Similarly it must be emphasized that the Mixed 

Mesophytic forest is recognized by both taxonomic 

composition and vegetational type. Only by keeping 

distinct the history of warm temperate vegetation 

from the history of lineages can the origins of the 

Mixed Mesophytic forest be understood. Warm 

temperate vegetation has probably always been 

present on the earth since the time of origin of land 

plants. Clearly, however, the composition of warm 

temperate vegetation prior to the origin of the 

angiosperms and their assumption of dominance in 

the Late Cretaceous is of little value to this 

discussion. Although some workers have stated that 

the "Arcto-Tertiary Geoflora" should be called the 

"Arcto-Cretaceo-Tertiary Geoflora," the evidence for 

the presence of most mixed mesophytic genera in the
44 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Cretaceous has never been documented. Considering 

the nature of Late Cretaceous pollen floras (see, for 

example, Gdczan and others, 1967), in which almost 

no extant dicotyledonous genera have been detected, 

it is clear that Late Cretaceous vegetation could not 

have represented a Mixed Mesophytic forest. 

Similarly, Paleocene floras, although somewhat 

more modern in taxonomic composition, contain 

numerous extinct genera. Some mixed mesophytic 

genera can be recognized, for example, Alnus, 

Corylopsis, and Sinowilsonia, but the inclusion in 

the same assemblages of genera such as Fagopsis, 

Pterospermites, Grewiopsis, Macclintockia, and 

numerous species of Dicotylophyllum that cannot be 

relegated to any extant family indicates that none of 

the Paleocene forests can properly be classed as 

Mixed Mesophytic. Indeed the fact that a few species, 

notably Cocculus flabella, Fagopsis groenlandica, 

Dicotylophyllum flexuosa, and D. richardsoni, 

appear to have dominated the warm temperate 

vegetation over very broad areas and that the 

Paleocene assemblages are not particularly diverse 

indicates also that vegetation resembling the Mixed 

Mesophytic forest was not in existence during the 

Paleocene. Not until the Eocene did significant 

numbers of mixed mesophytic genera become 

recognizable.

In the following discussion I have used in many 

instances only generic records. My reason for this in 

part stems from the fact that the occurrences of 

many mixed mesophytic genera are documented by 

diverse organs fructifications, leaves, and pollen 

and it cannot, of course, be determined in most 

instances whether a particular seed belongs to the 

same lineage as a particular leaf species. The 

assumption is made, for example, that, if a particular 

mixed mesophytic genus is known from non-foliar 

organs only from tropical types of vegetation in the 

Eocene, an Oligocene leaf species from temperate 

vegetation was probably derived from a tropical 

lineage. The record of lineages known from leaf 

species is more complete from the late Eocene and 

younger horizons that in pre-late Eocene time. Note 

that in the following discussion the unqualified use 

of the word tropical in reference to vegetation 

denotes both Tropical and Paratropical Rain forest.

The Alaskan early and middle Ravenian (late 

middle and early late Eocene) and Kummerian (early 

Oligocene) assemblages represent, respectively, 

Paratropical Rain forest and the last two subtropical 

forest (p. 35). These assemblages contain a significant number of mixed mesophytic genera and lineages, including:

Carya (bendirei type)

Platycarya

Pterocarya

Alnus

Betula sp.

Fagus sp.

Ulmus (newberryi type)

Tetracentron piperoides

Cocculus

Eucommia

Liquidambar?

Platanus (bendirei type)

Toona

Euodia

Rhus

Celastrus

Sageretia

Zizyphus

Vitis

Alangium

Clethra

Most of these genera and many others have been 

recorded from tropical Eocene and early Oligocene 

floras in the Pacific Northwest (Wolfe, 1968, and 

unpublished data; Scott in Chandler, 1964):

Carya

Juglans

Alnus

Betula

Fagus

Quercus

Calycocarpum

Euptelea

Cercidiphyllum

Tetracentron

Liriodendron

Magnolia

Hydrangea

Fothergilla

Hamamelis

Liquidambar (pachyphylla type)

Platanus

Cladrastis (japonica type)

Sageretia

Vitis

Alangium

Nyssa

Additional mixed mesophytic genera have been 

recorded from various subtropical or tropical 

Paleogene assemblages in Eurasia and North 

America:
A THEORY OF THE ORIGINS OF THE MIXED MESOPHYTIC FOREST 45

Populus

Salix

Comptonia

Myrica

Carpinus

Celtis

Zelkova

Cinnamornum

Exbucklandia

Fortunearia

Sinowilsonia

Rubus

Pachysandra

Ailanthus

Cedrela

Ilex

Rhus

Toxicodendron

Acer

Ampelopsis

Paliurus

Tilia

Actinidia

Eurya

Gordonia

Cornus

Diospyros

Symplocos

Halesia

Styrax

Ehretia

Catalpa

Sambucus

The significant fact is that these genera occur with 

regularity in fossil assemblages that on 

physiognomic criteria and overall floristic 

composition represent tropical or subtropical 

vegetation. Significant also is the fact that some 

of these genera (Platycarya, Tetracentron, Trochodendron, Euptelea, Calycocarpum, Fortunearia), 

although apparently widespread in the Paleogene 

tropical forests, have not been recorded from the 

Mixed Mesophytic forest as represented in the 

Neogene of western North America. The evidence 

thus indicates that some of the most characteristic 

genera now endemic to the Mixed Mesophytic 

forest attained their widest distribution in the 

Paleogene tropical forests and not in the Neogene 

Mixed Mesophytic forest. The modern distribution 

of many mixed mesophytic genera and alliances, 

in fact, indicates that many are fundamentally 

tropical or subtropical today, the mixed mesophytic 

representatives being only warm temperate outliers; 

the following genera exemplify this distribution:

Quercus

Akebia

Cocculus

Calycocarpum

Euodia

Vitis

Liriodendron

Magnolia

Liquidambar

Celastrus

Sageretia

Zizyphus

Alangium

Clethra

It is, therefore, not surprising that paleobotanical 

evidence also indicates that many mixed mesophytic genera adapted to warm temperate conditions 

after attaining a wide distribution in tropical 

forests during the Paleogene.

The late Ravenian (latest Eocene) assemblages 

from Alaska, British Columbia, and conterminous 

United States record a pronounced interval during

which most of the region was occupied by broadleaved deciduous or at least subtropical forest that 

had a diverse deciduous element (p. 33). By the latest 

Eocene, some taxa that, earlier in the Tertiary, were 

distributed in the Paratropical Rain and subtropical 

forests had adapted to more temperate climates. The 

Alaskan late Ravenian assemblage includes:

Populus?

Salix

Juglans (Cardiocaryon)

Pterocarya pugetensis

Alnus (cappsi type)

Corylopsis

Rhamnus

Tilia

To this list can be added genera known from late 

Ravenian floras farther south that display a similar 

distribution pattern:

Fagus

Betula (papyrifera type)

Comptonia

Carya (bendirei type)

Ulmus

Zelkova

Tetracentron (piperoides type)

Acer (rubrum type)

Vitis

These warm temperate forests were apparently 

taxonomically depauperate in comparison with the 

Mixed Mesophytic forest of the Neogene. Note that 

Hamamelidaceae are almost absent, in contrast to 

the widespread occurrences of Liquidambar and 

Exbucklandia in the middle Miocene of the Pacific 

Northwest. Cercidiphyllum, Euptelea, Liriodendron, 

Magnolia, and Platycarya are also absent from these 

late Ravenian temperate assemblages, although 

these genera are present in the early Oligocene 

tropical assemblages in the Pacific Northwest. Some 

genera had apparently not achieved a distribution 

outside the tropical vegetation during the latest 

Eocene and probably extended into the temperate 

regions only at a later time. Also of significance is 

that genera such as Acer are represented by only a 

few species in the late Ravenian but are well 

represented in later temperate vegetation, and, of 

course, "lobed" oaks are totally lacking in the later 

Ravenian assemblages. Although vegetationally the 

late Ravenian temperate forest is probably 

somewhat similar to the extant Mixed Mesophytic 

forest and although some floristic similarities exist, 

in general the late Ravenian temperate forest is not a 

Mixed Mesophytic forest. 

During the early to middle Oligocene, temperate
46 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

vegetation was apparently restricted to upland 

regions in the interior of the continent. Such 

assemblages as the Ruby of Montana (Becker, 1961) 

and, in part, the Florissant of Colorado (MacGinitie, 

1953) represent this type of vegetation. Various 

lineages and genera are represented in these two 

assemblages that indicate a floristic relationship to 

late Ravenian vegetation. Species of Acer, 

Dipteronia, Fagopsis, Alnus, Rosaceae, and 

Ulmaceae were represented in the early and middle 

Oligocene upland and late Ravenian assemblages by 

identical or closely related species. Note, however, 

that most characteristic mixed mesophytic genera 

lacking in the late Ravenian are also lacking in the 

early and middle Oligocene temperate vegetation but 

are present at that time in tropical vegetation. Some 

groups, for example, the "lobed" oaks and most 

Ericaceae, are not known in any early and middle 

Oligocene assemblages. As with the late Ravenian 

assemblages, the upland early and middle Oligocene 

assemblages represent warm temperate vegetation 

but not the mixed mesophytic forest.

The rapid climatic deterioration of the late 

Oligocene (the deterioration is probably of latest 

middle Oligocene age in the marine megafossil 

chronology but is apparently of early middle 

Oligocene age in the vertebrate chronology, that is, 

about 32 million years ago) led to wide lowland areas 

of warm to cool temperate climate. Assemblages 

such as those of Angoonian age in Alaska (Tsadaka) 

and Oregon (Bridge Creek, Willamette) represent 

broad-leaved deciduous forest; in Oregon broadleaved evergreens such as Cinnamomum, Paleophytocrene, and Willisia were a minor element in the 

vegetation (Wolfe, in Peck and others, 1964), but this 

element is represented only by Willisia in southeastern Alaska and is lacking in the Cook Inlet and 

other regions of Alaska. Floristically this vegetation 

is closely related to that of the uplands of the early 

and middle Oligocene, as shown by numerous closely 

related species of Alnus, Betula, Rosaceae, Acer, and 

Dipteronia. Added to the Angoonian assemblages in 

Oregon, however, are the following genera or 

lineages that have not been recorded from older 

upland assemblages but were found in the lowland 

early and middle Oligocene:

Engelhardtia Liquidambar

Cercidiphyllum Platanus (dissecta type)

Cinnamomum Toxicodendron

Exbucklandia

This element appears to have been added to the 

temperate vegetation during, or shortly after, the 

major climatic deterioration. In Alaska, but not in 

Oregon, Salicaceae played a major role in the

vegetation. Some mixed mesophytic genera that had 

been members of the temperate or the tropical forest 

of the early and middle Oligocene apparently 

became extinct in western North America at about 

the time of the deterioration; these include Platycarya, Tetracentron, Trochodendron, Euptelea, and 

Calycocarpum. Renewed warming in the latest 

Oligocene resulted in marginally subtropical (leaf 

margin percentages are about 40) assemblages in at 

least coastal Oregon, and marginally warm temperate assemblages in Alaska. In Oregon, Lauraceae 

were represented by several species, but broadleaved deciduous species were still apparently 

dominant members of the vegetation; most of these 

species are either conspecific with, or closely related 

to, species of the older late Oligocene temperate 

assemblages. In Alaska, the flora was more diverse 

during late Oligocene than during the earlier and 

cooler part of the Angoonian; broad-leaved evergreens are represented by Engelhardtia (characteristic involucres have been found in late Angoonian 

rocks in the central Alaska Range), Ilex, and 

Alangium. Pollen records include Cedrela/Melia. 

Rosaceae also were apparently more diverse in 

Alaska than previously and include Duchesnea or 

Fragraria, Rubus, Sorbus, and Spiraea. The temperate vegetation seems to have been enriched, partly 

through the adaptation of tropical lineages to 

temperate climates and partly through the diversification of lineages previously present in temperate 

vegetation.

The early Miocene (early Seldovian) seems to have 

been cooler than the latest Oligocene. Leaf-margin 

percentages in Oregon for early Miocene assemblages are about 25 and in Alaska are about 12. 

Despite the cooling, the broad-leaved deciduous 

forest seems to have been taxonomically more 

diverse than anytime in the Angoonian. In Oregon, 

the Collawash assemblage contains representatives 

of 140 megafossil and about an additional 20 

microfossil entities; this assemblage is the richest 

Neogene assemblage known in North America, 

although the middle Miocene Latah flora of 

Washington approaches the Collawash in diversity. 

The inclusion in the Collawash of diverse Salicaceae, 

Juglandaceae, Betulaceae, Fagaceae (including 

"lobed" oaks), Ulmaceae, Hamamelidaceae, Rosaceae, Aceraceae, and Ericaceae, along with some 

broad-leaved evergreens such as Lauraceae, indicates that floristically the warm temperate vegetation was a Mixed Mesophytic forest for the first time. 

This vegetation in Oregon, however, lacked some 

genera that were present earlier in the Tertiary in 

North America and today are members of the Asian
A THEORY OF THE ORIGINS OF THE MIXED MESOPHYTIC FOREST 47

or east American Mixed Mesophytic forest, such as, 

Platycarya, Euptelea, Trochodendron, Tetracentron, Calycocarpum, Corylopsis, Sinowilsonia, Fortunearia, Eucommia, and Dipteronia. Some of these 

genera apparently did not adapt to temperate 

conditions in western North America, although 

some, for example, Dipteronia, have a record in the 

temperate vegetation of this region from at least the 

late Eocene through the late Oligocene. These genera 

were following a different pattern in western North 

America than in Eurasia.

Families such as Betulaceae, Salicaceae, Rosaceae, and Aceraceae, and genera such as Fagus and 

Castanea, that were taxonomically depauperate in 

the Paleocene and Eocene floras, although present in 

the Paleogene subtropical and tropical forests, were 

able to adapt readily to the new temperate climates 

and diversified. From both described and undescribed material from the Tertiary of western North 

America, table 8 has been compiled. Although the 

table may reflect in part the lesser knowledge of 

Eocene and Pliocene floras as compared to other 

epochs, the table probably reflects primarily the 

relative diversity of the families. Ericalean pollen is 

known from earlier Paleogene rocks but is rare. 

Lineages of some Rosaceae and Aceraceae can be 

traced to the upland floras of the early middle 

Oligocene, but in the instances of Salicaceae and 

Betulaceae, new morphologic types appeared, apparently suddenly, and diversified rapidly. The 

diversification of Populus, Betulaceae, Rosaceae (at 

least in part), and Aceraceae, seems to have reached 

its maximum in the early and middle Miocene in the 

Mixed Mesophytic forest as far as the west American 

record is concerned. Note that the diversity of 

Betulaceae and Aceraceae was greater in the Mixed 

Mesophytic forest of the Miocene of Asia (Tanai, 

1961), as it is today in Asia, than in North America.

Both Salix and Ericaceae, although present in 

considerable diversify in the Miocene Mixed Mesophytic forest, continued to diversity in western North 

America after the decline of the Mixed Mesophytic 

forest here; Ericaceae continued to diversify also in 

the Mixed Mesophytic forest of Asia, as evidenced by 

the richness of genera such as Rhododendron. Such 

diversification is unparalleled in North America. 

Note that warm temperate rosaceous and ericaceous 

genera display the greatest degree of endemism in 

the extant Mixed Mesophytic forest. This endemism 

probably indicates that many of these genera are of 

more recent origin; by the time they had evolved, the 

Alaskan forest definitely indicates a cooler climate 

than was present in conterminous United States 

(Wolfe and Leopold, 1967) and would thus inhibit the 

interchange between Asia and North America of 

more newly evolved warm temperate genera. An 

additional barrier to the dispersal of some taxa 

during the Miocene may have been precipitation: the 

Mixed Mesophytic forest of the Pacific Northwest 

includes numerous genera that have, at least during 

the later Tertiary, displaced a proclivity for summer 

dry climates. Included are:

Umbellularia Ceanothus 

Cercocarpus Colubrina 

Holodiscus Karwinskia 

Heteromeles Xylonagra 

Lyonothamnus Garrya 

Peraphyllum Arbutus

Arctostaphylos

The mechanics of the atmospheric circulation of the 

earth demand that the west coasts of major land 

masses be winter wet; although the Miocene Mixed 

Mesophytic forest in the Pacific Northwest probably 

had much more summer precipitation than at 

present, the climate probably could not have been as 

summer-wet as in the regions where the Mixed

TABLE 8. Specific diversity through time of some families represented in the west American Tertiary

[The term "many" indicates numerous species, but the taxonomic status is not satisfactory]

Eocene

Family or genus

Pinaceae .........................................

Salicaceae ......................................

Juglandaceae (excluding 

Englehardtia) ............................

Betulaceae .....................................

Ulmaceae .......................................

Trochodendraceae (s.l.) .............

Magnoliaceae ...............................

Aceraceae .......................................

Ericaceae .......................................

Paleocene

4

?

1

3

2

4?

5

?

6

2?

2

0

tropical

3 

2 

3

13 

4 

11 

5 

4 

many 

many 

6 

2 

2? 

0

temperate

3

7 

3

1 

4 

2 

1 

0 

3 

0 

1 

8 

3 

1

early

tropical

3 

0 

0

5 

3 

3 

4 

4 

many 

many 

3 

1 

0 

0

Oligocene

early

upland

3 

9

8

3 

2 

3 

1 

0 

2 

0 

0 

17 

5 

0

Miocene

late

5

13 

10

8 

15 

8 

1 

2 

7 

2 

4 

12 

6 

2

early and 

middle

5 

13

25

10 

12 

6 

1

2 

7 

1 

4 

20 

11 

4

late

2 

21

35

6 

12 

3 

1 

1 

1 

0 

1 

28 

6 

12

Pliocene

2 

23 

many

3 

4 

1 

0 

0 

1 

0 

0 

28 

4 

14
48 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Mesophytic forest still survives. Alaska receives 

more summer precipitation today than the Pacific 

Northwest, and the same relationship probably 

prevailed in the Miocene. Thus, the cooler climate in 

combination with the wetter summers may have 

acted as an effective barrier to the exchange of many 

genera between Eurasia and western North America. 

The Mixed mesophytic forest of the Miocene of 

western North America thus seems to have had three 

major sources:

1. Genera derived directly from the Paleogene 

tropical and subtropical forests,

2. genera that had adapted by the late Eocene or 

early to middle Oligocene to interior warm 

temperate uplands, and

3. genera that evolved during the Oligocene and 

Miocene from lineages that were associated 

with either group 1 or group 2.

The history of the Mixed Mesophytic forest has 

involved little if any significant amount of migration 

of the association on a maj or scale. As a recognizable 

floristic and vegetational unit, the Mixed Mesophytic forest developed in the early and middle Miocene 

from the depauperate temperate vegetation that 

resulted from the middle late Oligocene climatic 

deterioration. The progressive enrichment of the 

Mixed Mesophytic forest in Asia continued during 

the Neogene, probably both by diversification of 

lineages already part of the forest and by extensions 

into the forest of subtropical and tropical lineages. In 

western North America, the Mixed Mesophytic 

forest was eliminated by increasingly pronounced 

summer-dry conditions by the end of the middle 

Miocene, although individual lineages of the Mixed 

Mesophytic forest persist to this day in western 

North America.

In eastern North America, the Mixed Mesophytic 

forest does not seem to have been as rich at any point 

in time as the Mixed Mesophytic forests of either 

eastern Asia or western North America. Almost all 

woody genera now present in the eastern Mixed 

Mesophytic forest were also present in the west 

American forest. Although the Neogene fossil record 

in eastern North America is scanty, megafossil and 

microfossil assemblages from the middle and late 

Miocene of the central Atlantic coast states indicate 

that numerous Mixed Mesophytic genera were 

apparently not present there that were in western 

North America:

Gingko Schoepfia 

Keteleeria Mahonia 

Pseudolarix Litsea 

Metasequoia Lindera 

Zelkoua Umbellularia

Cinnamomum

Cercidiphyllum

Exbucklandia

Cercocarpus

Holodiscus

Lyonothamnus

Peraphyllum

Cedrela

Euodia

Pistacia 

Colubrina 

Ceanothus 

Karwinskia 

Zizyphus 

Xylonagra 

Garrya 

Arbutus 

Clerodendrum 

Idesia

Although many of these genera are apparently less 

tolerant of summer moisture, others are typical of 

regions of summer-wet climate today. Certain of the 

genera would be expected in eastern North America 

if there had been an Arcto-Tertiary Geoflora that 

gradually migrated southward. The apparent depauperateness of the eastern Mixed Mesophytic 

forest may be the result of four major factors:

1. The middle to late Oligocene climatic deterioration that probably would have left as it 

did in northwestern North America a 

depauperate flora,

2. the elimination at the same time of the tropical forest entirely or in large part from 

southeastern United States, thus making it 

more difficult for a progressive enrichment 

of the temperate forest from the tropical 

forest,

3. the essential isolation of eastern North 

America both from Europe and western 

North America, and

4. the little topographic diversity of eastern

North America.

The first two factors may account for the poor 

development of fundamentally tropical families in 

the eastern Mixed Mesophytic forest. At present, 

only two genera of Lauraceae Persea and Sassafras are represented in the eastern forest, as 

compared to the Miocene of western North America 

where six genera (Cinnamomum, Lindera, Litsea, 

Persea, Sassafras, and Umbellularia) were represented, and to the forest of eastern Asia where six 

genera have been, and are now, represented (Cinnamomum, Lindera, Litsea, Neolitsea, Persea, and 

Sassafras). In western north America, the Miocene 

forest, in addition to Lauraceae, contained many 

genera of fundamentally subtropical or tropical 

relationships: Cedrela, Clerodendrum, Cocculus, 

Colubrina, Diospyros, Exbucklandia, Gordonia, 

Idesia, Karwinskia, Magnolia, Nyssa, Oreamunoa, Oreopanax, Pistacia, Schoepfia, Vitis, Zizyphus; many of these same genera plus many other 

genera were and are well represented in the Asian 

Mixed Mesophytic forest. In eastern North America,
PALEOGENE TROPICAL MIGRATION ROUTES 49

the comparatively few genera of this type are: 

Berchemia, Calycocarpum, Cocculus, Gordonia, 

Magnolia, Nyssa, Symplocos, and Vitis. Genera of 

Lauraceae, Leguminosae, and Sapotaceae that were 

abundant in the tropical Paleogene forest of eastern 

North America are, for the most part, unrepresented 

by species in the Mixed Mesophytic forest Tertiary 

or extant of the same region.

The fourth factor may account for the little generic 

diversification that took place in families such as 

Rosaceae and Ericaceae, and the third factor 

isolation prevented any great amount of floristic 

interchange with regions that had topographic 

diversity. It should be noted that the Paleogene 

tropical forest of eastern North America apparently 

had few genera in common with the Paleogene forest 

of western North America and Eurasia; the floristic 

isolation of eastern North America seems to have 

been ancient (MacGinitie, 1969). Most of the 

vicarious species between eastern North America 

and eastern Asia today belong to genera that are 

found in Paleogene tropical forests and that have 

in western North America and Eurasia shown a 

definite tendency to become readily adapted to 

temperate conditions; these same genera have in 

some instances been recorded from the Paleogene 

tropical forest of eastern North America (Gray, 

1960). Historical factors have, therefore, probably 

been responsible for the independent development of 

the Mixed Mesophytic forest in Eurasia, western 

North America, and eastern North America, as well 

as for the greater similarity between the forest in the 

Miocene of western North America and Eurasia.

As has been emphasized previously (Wolfe, 1972), 

the lineages that are represented in the Mixed 

Mesophytic forest whether that extant in eastern 

Asia and eastern North America or that of the 

Neogene of western North America are in this 

forest coincidentally. The lineages have, in some 

instances, followed similar patterns of distribution, 

but as the history of the lineages has become clearer 

through paleobotanical work, several different 

patterns have emerged. I think that as more work is 

done, many more patterns will become apparent. 

Some lineages have followed a general pattern for a 

longer period than have other lineages, for example, 

some lineages have been adapted to temperate 

climates for a longer period. Some lineages have 

participated in the temperate vegetation of North 

America but became extinct before the optimal 

development of the Mixed Mesophytic forest. The 

warm temperate forest of the Northern Hemisphere 

has been and will continue to be in a continual state 

of flux.

Lineages have entered and departed from this vegetation at 

different times in different places in response to different and 

changing parameters of the tolerances of the individual members 

of the lineages. Various lineages have diversified in the warm 

temperate vegetation at different times and in different regions. 

The history of the Mixed Mesophytic forest is complex and can be 

understood best by reconstructing the patterns of distribution in 

time and space of the component lineages (Wolfe, 1972, p. 231).

PALEOGENE TROPICAL MIGRATION ROUTES

The existence in Alaska during parts of the 

Paleogene of Paratropical Rain forest has an 

important bearing on the times and routes of 

migration of tropical lineages (Wolfe, 1975). In a 

significant paper, van Steenis (1962) has discussed 

in detail the significance of trans-Pacific angiospermous genera and families, and he concludes that 

the only means of attaining such a bicontinental 

distribution for a tropical group is by means of 

former trans-Pacific land bridges that were within 

present tropical latitudes.

The disjunctions of northern warm temperate or 

subtropical genera between North America and 

Eurasia are, as van Steenis (1962, p. 243) points out, 

in many instances involved with the disjunctions of 

tropical groups. The probable phytogeographic 

history of many warm temperate or subtropical 

disjunct groups has been discussed in the preceding 

section of this report; such disjunctions have, in 

many instances, the same basic historical pattern as 

the disjunct tropical groups.

Van Steenis (1962) is primarily concerned with the 

"Amphi-transpacific" genera and families that are 

megatherms (that is, "adapted to tropical climate"; 

van Steenis, 1962, p. 239). He logically concludes (p. 

290) "***that there is no reason to assume significant 

changes during the geological past in the thermoecology of the majority of the tropical genera 

concerned." Tropical lineages could not, therefore, 

have adapted to cool conditions to cross, for example 

Beringia, and then become readapted to tropical 

conditions. Van Steenis (p. 271-322) also discusses 

seven other explanations for the present distribution of tropical groups, some of which, for example, 

polyphylfetic origins, are almost certainly improbable. The only explanation that van Steenis 

considers credible is that of transoceanic land 

bridges at tropical latitudes.

The data presented here indicate that van Steenis' 

conclusion is, at least in regard to some tropical 

disjunctions, not necessary. He was led to disregard 

the significance of the Beringian route for tropical 

genera for two major reasons. First, he assumed 

(1962, p. 271) a "steady state" framework for the 

earth, that is, there has been little or no continental
50 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

drift, the inclination of the axis of rotation has been 

unchanged, and latitudinal climatic zonation has 

changed little. From these assumptions he concludes 

(p. 271) that the areas between latitudes 50°-70° N. 

"***might have been warm-temperate to subtropical 

and the majority of the trees must have been 

deciduous." Second, van Steenis (p. 271-281) accepts 

without question the concept of the Arcto-Tertiary 

Geoflora and as well considers the London Clay flora 

to be atypical of Eocene vegetation at latitude 50° N. 

Chandler (1964, p. 86-89) has adequately invalidated 

van Steenis' conclusions regarding the London Clay; 

that assemblage almost certainly represents warm 

Paratropical to marginally Tropical Rain forest 

(Richards, 1952, p. 154).

Van Steenis (1962, p. 273) recognizes the physiological problems encountered if tropical plants had 

grown at high northern latitudes: "Though at 

present low temperature is more important than 

light, increase of the world's temperature will, at 

arctic latitudes, very soon find its bottleneck in the 

form of this same light." He thus considers 

reasonable the existence during the Paleogene of 

broad-leaved deciduous and conifer forests at high 

northern latitudes. Discounting the occurrence of 

palms in the Greenland Tertiary, van Steenis (p. 272) 

states:

***palms have never been found beyond about 50° latitude. Their 

distribution forms a most important argument for the conservative view of only slight climatic changes in zonation at medium 

latitudes and a full confirmation of the steady state principle. 

No mention is made of the well-documented 

occurrences of fan palms at latitudes 57° and 61.5° N. 

in Alaska (Rollick, 1936).

The data given in the present report clearly 

indicate that tropical forests (in van Steenis' concept 

of tropical vegetation, the Paratropical Rain forest is 

included) indeed existed in Alaska. The Alaskan 

early Ravenian (late middle Eocene) assemblages 

include some of van Steenis' "Amphi-transpacific" 

genera: Sageretia, Meliosma, and Saurauia. Van 

Steenis (1962, p. 268), moreover, considers that more 

significant than genera are the various exclusively 

"pantropical larger megatherm families" and the 

"tropical pro majore parte" pantropical families. In 

the first category are the early Ravenian Annonaceae, Myristicaceae, and Lecythidaceae (including 

Barringtoniaceae), and in the second category are 

the early Ravenian Palmae, Menispermaceae, 

Lauraceae, Rutaceae, Anarcadiaceae, Icacinaceae, 

Sapindaceae, Vitaceae, Theaceae, Myrtaceae, and 

Verbenaceae. Added to these are the early Ravenian 

groups that are exclusively paleotropical (p. 28). 

Considering the small size of the known Alaskan

early Ravenian flora, the families listed above are 

impressive evidence that many pantropical or 

"Amphi-transpacific" genera and families could 

have attained such a distribution by way of 

Beringia. By analogy, some tropical groups may 

have been distributed by way of a high latitude 

Southern Hemisphere migration route, although 

paleobotanical documentation for such a route is 

lacking.

Considering that many fundamentally tropical 

groups were distributed through Beringia and into 

middle latitudes of western North America during 

the Tertiary, it is significant that the present tropical 

floras of the Americas and southeastern Asia do not 

have more genera in common today. The known 

Paratropical Rain forest of the Paleogene of western 

United States contains the following now exclusively paleotropical groups:

Menispermaceae: Anamirta, Diploclisia, Hypserpa, Limacia, Pycnarrhena, Tinomiscium, 

Tinospora

Myristicaceae: Knema, Myristica

Annonaceae: Cananga

Olacaceae: Erythropalum, Olax

Hamamelidaceae: Altingia

Rutaceae: Euodia, Luvunga

Anacardiaceae: Dracontomelon, Poupartia

Icacinaceae: lodes, Miquelia, Phytocrene, Pyrenacantha, Stemonurus

Vitaceae: Tetrastigma

Barringtoniaceae: Barringtonia

Dipterocarpaceae: Parashorea

Alangiaceae: Alangium

Cornaceae: Mastixia

The fact that these genera have not survived in the 

American tropics may be the result of the major 

climatic change during the Oligocene coupled with 

the small land area of humid tropical and 

paratropical climates in North America (Wolfe, 

1975).

Disjunction of tropical genera and families, 

therefore, may not be the result of migrations of 

lineages across tropical latitude land bridges that 

have subsequently disappeared. Some of the 

migrations were probably by way of Beringia. The 

times of such migrations could have been during the 

Paleocene because Eohypserpa, Macaranga, Melanolepis, and Sabalites are known from the Alaskan 

Paleocene (Wolfe, 1966; Burk, 1965), during early 

Eocene when warming in Alaska is assumed to have 

occurred simultaneous with that known in middle 

latitudes, or during early late Eocene or the early 

Oligocene when the very warm subtropical forest 

indicates that locally warmer areas might have
SYSTEMATICS 51

allowed some tropical lineages to have migrated 

through Alaska. Chance dispersal through tropical 

or middle latitudes across various islands should 

also not be overlooked as a means of attaining 

pantropical or "Amphi-transpacific" distribution; 

such migrations are "land-bridge dispersals" of van 

Steenis (1962). It is doubtful if each tropical 

disjunction can be explained in terms of precisely the 

same pattern in regard to both time and space. I 

think that various patterns, as discussed in reference 

to the Mixed Mesophytic forest, will be found to be 

valid in explaining tropical disjunctions.

SYSTEMATICS

The preservation of many specimens is not so good 

as could be desired. In most, venation below the 

fourth order is not preserved. Sufficient characters 

are preserved, however, to allocate most of the 

material to genera. If the specimen is fragmentary 

but well preserved, no specific epithet is assigned in 

most instances. I emphasize that the apparently 

modern aspects of the floras at the generic level may 

be a reflection of the lack of preservation of fifth and 

sixth order venation.

Uncleared herbarium specimens of foliage of 

tropical plants are typically difficult to compare to 

fossil leaves. The thick texture of most tropical 

leaves obscures the venation pattern so that in most 

instances venation below the level of secondaries or 

tertiaries cannot be clearly seen. The use of cleared 

material is almost mandatory for the study of 

assemblages such as that of the early Ravenian in 

Alaska. To a certain extent, therefore, the determinations made here are a reflection of the extent of the 

U.S. Geological Survey's reference collection of 

cleared leaves. At this time (May 1971) this collection 

comprises about 2,000 species representing about 

1,200 genera; over 1,000 of these species are of 

tropical distribution.

The systematic lists of Eocene assemblages such 

as the early Ravenian differ markedly in one 

important aspect from modern floras. Members of 

Sympetalae are notably scarce or lacking in Eocene 

and older assemblages. Families of Sympetalae that 

are diverse in the tropical regions today Rubiaceae, 

Boraginaceae, and Bignoniaceae, for example are 

proportionately underrepresented in most Eocene 

assemblages in comparison with their present 

diversity; this observation applies to both leaf and 

seed floras. I suggest that the diversification of many 

families of Sympetalae was largely post-Eocene.

With the exception of a few specimens in the 

collections of the California Academy of Sciences 

(here abbreviated CAS) and some comparative

material from the collections of the University of 

California Museum of Paleontology (UCMP), all 

figured or cited specimens are deposited in the 

United States National Museum (USNM). Rather 

than retouch photographs, which can be highly 

misleading, I have included camera lucida drawings 

of most critical features.

The material from the Gulf of Alaska Coastal 

Province illustrated by Hollick (1936) includes some 

specimens that, although determined to species by 

Hollick, are inadequate for either generic or specific 

assignment. Species that are here relegated to 

nomina dubia are:

Piper disputabilis Hollick. Although the specimen matches well the specimen of Cornus 

irregularis Holl. from the same locality, the 

type of P. disputabilis is extremely fragmentary and is not a satisfactory name-bearing 

type.

Mohrodendron inopinum Hollick. Hollick's specimen lacks both base and apex, and is, therefore, not a satisfactory name-bearing specimen. 

Rhododendron crassum Hollick. the venation

below the level of secondaries is lacking. 

Hollick (1936) also based citations of previously 

described species on poor material. The following 

citations should not be considered valid and cannot 

be satisfactorily synonymized to other species:

Canavalia eocenica Berry 

Rhamnus decheni Weber

Hollick (1936, p. 130, pi. 93, fig. 3) cited Tetrapteris 

harpyiarum Ung. as occurring at locality 3879; in 

fact, the specimen came from locality 5892 in the 

Chickaloon Formation of the Matanuska Valley. 

This specimen belongs to Dicotylophyllum flexuosa 

(Newb.) Wolfe (1966, p. Bll). Name-bearing specimens of Hollick (1936) from the Gulf of Alaska region 

not discussed above are:

Asplenium alaskanum Holl. = Dryopteris alaskanum (Holl.) Wolfe 

Pteris inaequilateralis Holl. = Cyathea inaequilateralis (Holl.) Wolfe 

Celastrus comparabilis Holl., determination

valid 

Piper concavum Holl. = Cryptocarya presamarensis Sanb. 

Ulmus pseudobraunii Holl. = Platycarya pseudobraunii (Holl.) Wolfe

Planera aquaticiformis Holl. = Platycarya pseudobraunii (Holl.) Wolfe 

Hampea conditionalis Holl. = Paratinomiscium

conditionalis (Holl.) Wolfe
52 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Persea spatiosa Holl. = Luvunga spatiosa (Roll.) 

Wolfe

Semecarpus alaskana Holl. = Melanorrhoea 

alaskana (Holl.) Wolfe

Rhamnus pseudogoldianus Holl. = Parashorea 

pseudogoldiana (Holl.) Wolfe

Carpolithes elytraeformis Holl. = Paleophytocrene elytraeformis (Holl.) Wolfe

Cornus irregularis Holl. = Mastixia irregularis

(Holl.) Wolfe

Transfers of citations by Hollick of previously 

described species are:

Cinnamomum cinnamomeum of Hollick (non 

Phyllites cinnamomeus Rossm., 1840) = Cinnamomophyllum latum (MacG.) Wolfe

Malpoenna magnified of Hollick (pi. 66, fig. 5 

only) = Alangium bergensis Wolfe

Glyptostrobus europaeus (Brongn.) Heer = 

Glyptostrobus sp.

Magnolia wormskioldi Heer = Cananga sp.

Daphnogene kanii Heer = Macclintockia pugetensis Wolfe

Dryophyllum stanleyanum Daws. = Melanorrhoea alaskana (Holl.) Wolfe

Cornus buchii Heer = Laurophyllum presamarensis (Sanb.) Wolfe

Cornus hyperborea Heer? = Parashorea pseudogoldiana (Holl.) Wolfe

Populus balsamoides Goepp. = Celastrus comparabilis Holl.

Juglans thermalis Lesq. = Goweria dilleri 

(Knowl.) Wolfe

Artocarpidium alaskanum Holl. (nontypic) = 

Alnus scotti Wolfe

Magnolia ovalis Lesq. = indet. leaf

Rhamnus marginatus Lesq. = Parashorea pseudogoldiana (Holl.) Wolfe

Terminalia? sp. of Holl. = Myristica sp.

Fraxinus lateralis Holl. (pi. 105, fig. 6) = Platycarya pseudobraunii (Holl.) Wolfe

Order FILICALES

Family SCHIZAEACEAE

Genus Anemia Swartz 

Anemia gunni Wolfe, new species

Plate 2, figure 3

Diagnosis. Pinna pinnatifid, apparently ovate, 

at least 25 cm long and 26 cm wide; at least 8 

pinnules, decurrent along midrib, reflexed basally, 

decidedly slender and acuminate; secondary veins 

thinning towards apices of pinnules; tertiary veins 

simple to twice-branching, typically basal dichotomy dichotomizing again and apical dichotomy

unbranching, craspedodrome; margin finely serrate; 

texture coriaceous.

Discussion. The single specimen is an incomplete pinna, because the fertile pinnules are lacking; 

in Anemia, the basal pinnules are typically fertile. 

Among described species, the Kulthieth specimen 

most closely resembles A. occidentalis Knowlt. in the 

shape of the pinna and pinnules, but Knowlton's 

species is described as having once-dichotomizing 

tertiary veins (Knowlton, 1917, p. 285). Both A. 

eocenica Berr. and A, elongata (Newb.) Knowlt. have 

pinnules that are deeply incised or "lobed." The 

tertiary venation pattern, as well as texture, however, of A. gunni are similar to A. eocenica. A. 

gunni also occurs in the Puget Group; I (1968) had 

included the fragmentary Puget material in A. 

eocenica.

This species is named for Dr. Donald Gunn, 

Humble Oil and Refining Corporation, who assisted 

in collecting the material.

Holotype. USNM 43246.

Occurrence. Kulthieth Formation, loc. 11170.

Anemia? sp. 

Plate 28, figure 5

Discussion. A fragment, apparently representing the apex of a pinnule of Anemia, has a serrate 

margin and once- or twice-dichotomizing veins.

Specimen. USNM 43247.

Occurrence. Kushtaka Formation, loc. 9891.

Genus Lygodium Swartz 

Lygodium larsoni Wolfe, new species

Plate 20, figures 1, 2

Diagnosis. Sterile pinnules palmately two-lobed; 

one lobe larger and more elongated than the other; 

short lobe up to 5 cm in length, long lobe up to 7 cm in 

length; lobes about 1.2-1.5 cm in width; base broadly 

rounded to shallowly cordate; apex narrowly 

rounded; primary veins strong basally but thinning 

apically and about two-thirds the distance to the 

apex becoming the same strength as the secondary 

veins; secondary veins departing at low angles, 

dichotomizing two to three times; margin entire.

Discussion. All 16 nearly complete specimens are 

bilobed, although one fragment apparently has a 

weakly developed third lobe. The widespread Eocene 

species Lygodium kaulfussi includes material that is 

bilobed, but the leaves of that species typically have 

three lobes. L. bineruatum (Lesq.) Berr. has bilobed 

leaves, but the lobes are very blunt and short.

This species is named for Dr. Willard Larson, 

Humble Oil and Refining Corporation, who assisted 

in collecting the material.

Holotype. USNM 43248.
SYSTEMATICS 53

Paratype. USNM 43249.

Occurrence. Kushtaka Formation, Iocs. 9389, 

11165.

Family Cyatheaceae

Genus Cyathea Smith 

Cyathea inequilateralis (Hollick) Wolfe, new combination

Plate 3, figure 3

Pteris inequilateralis Hollic, 1936, U.S. Geol. Survey Prof. Paper

182, p. 38, pi. 5, fig. 1. 

Hemitelea pinnata MacGinitie, 1941, Carnegie Inst. Washington

Pub. 534, p. 97, pi. 10, fig. 1. 

Cyathea pinnata (MacGinitie) LaMotte, 1951, Geol. Soc. America

Mem. 51, p. 140. 

Pabst, 1968, California Univ. Pubs. Geol. Sci., v. 76, p. 36, pi. 2,

fig. Ic; pi. 3, fig. 2; pi. 4, fig. 2; pi. 5.

Discussion. The asymmetry displayed by Hoilick's specimen can also be seen in the material 

figured by MacGinitie (1941) and Pabst (1968). The 

epithet proposed by Hollick is clearly the senior 

synonym. The material of Asplenium magnum 

Knowl. from Eocene beds in Yellowstone Park is, as 

MacGinitie (1941, p. 98, stated, not so deeply dissected as in Cyathea inequilateralis.

Holotype. USNM 38665.

Hypotype. USNM 43250.

Occurrence. Kushtaka Formation, Iocs. 3846, 

11160, 11162.

Family ASPIDACEAE

Genus Allantodiopsis Knowlton and Maxon 

Allantodiopsis pugetensis Wolfe

Plate 2, figures 1, 2; plate 20, figure 4

Allantodiopsis pugetensis Wolfe, 1968, U.S. Geol. Survey Prof. 

Paper 571, p. 14, pi. 1, fig. 6.

Discussion. The Alaskan specimens have the 

same linear shape, serrate margin, and simple or 

once-branching veins as in the types of Allantodiopsis pugetensis.

Hypotypes. USNM 38957, 43251, 43252.

Occurrence. Kushtaka Formation, Iocs. 3846, 

11159, 11165, 11166; Kulthieth Formation, loc. 3879. 

Genus Dryopteris Adanson

Dryopteris alaskana (Hollick) Wolfe, new combination 

Plate 3, figures 1, 4

Asplenium alaskanum Hollick, 1936 (part), U.S. Geol. Survey 

Prof. Paper 182, p. 38, pi. 3, figs. 1, la, lax; pi. 4, figs, la, lax, 

Ib, Ibx, 2-6, 6a; pi. 5, fig. 5 only.

Discussion. The diagnostic characters of sterile 

specimens of Dryopteris gibbsi listed by Pabst (1968, 

p. 39-40), namely the arched, falcate pinnules and 

alternate venation, are also present in Hollick's 

material of Asplenium alaskanum. The sori of D. 

gibbsi are, however, round, whereas the sori of D. 

alaskana are decidedly oval.

Lectotype. USNM 38664.

Syntypes. USNM 38662, 245724. 

Occurrence. Kushtaka Formation, loc. 3847; 

Kulthieth Formation, loc. 3879.

Family BLECHNACEAE

Genus Woodwardia J. E. Smith

Woodwardia sp.

Plate 28, figure 1

Discussion. The specimen is a fertile frond and 

has kidney-shaped sori next to the rachillae; the 

pinnae are confluent along the rachis. These 

characters indicate that the specimen represents 

Woodwardia, although the assignment should be 

considered tentative because of the poor preservation of the venation.

Specimen. USNM 43253.

Occurrence. Kushtaka Formation, loc. 9891.

Order CONIFEBALES 

Family TAXODIACEAE

Genus Glyptostrobus Endlicher

Glyptostrobus sp. 

Plate 3, figure 2; plate 20, figures 3, 6; plate 21, figure 2a

Discussion. Lacking fertile material I cannot 

determine which species is represented or indeed if 

more than one species is represented. The strongly 

dimorphic character of the foliage at the Charlotte 

Ridge locality (pi. 21, fig. 2a) indicates that this 

species probably does not represent Glyptostrobus 

nordenskioldi (Heer) R. W. Br.

Glyptostrobus is today the member of Taxodiaceae 

that consistently inhabits a paratropical climate. 

Despite this modern tolerance, it is certain that the 

genus contained members of widely differing tolerances. The genus was still represented in the Clamgulchian, for example, when Alaska had a paratemperate climate (Wolfe, 1966). Glyptostrobus is 

the only genus of its family known to be represented 

in every epoch of the Tertiary in Alaska. Although 

much has been made of the former wide distribution 

of Metasequoia particularly at high latitudes it 

seems that Glyptostrobus is even more characteristic 

of high latitude Tertiary assemblages.

Specimens. USNM 38724, 43254-43256.

Occurrenee. Kushtaka Formation. Iocs. 11165, 

11167; Kulthieth Formation, loc. 3879. 

Order PRINCIPES 

Family PALMAE

Genus Phoenicites Brongnairt

Phoenicites sp.

Plate 20, figure 5

Discussion The specimens represent a species of 

feather palm. The rachis does not appear to have 

been spinose, and no spines can be detected on the 

margins of the pinnules. Both specimens have the
54 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

pinnules arranged alternately. The midrib of the 

pinnules is very thick; several lateral veins are 

present, and some are thicker than others. In method 

of attachment of the pinnules, which narrow and are 

not folded at the point of attachment, and in 

venation, the fossils resemble the sago palm, 

Metroxylon. This genus, however, typically has 

widely spaced spines on the margins of the pinnules, 

although spines are lacking on some pinnules. 

Ptycoraphis is also similar to the fossils. Considering the fragmentary nature of the specimens, 

however, no generic assignment other than to the 

organ genus Phoenicites is warranted.

Specimen. USNM 43257.

Occurrence. Kushtaka Formation, loc. 11165.

Genus Palmacites Brongniart 

Palmacites sp.

Plate 4, figure 5

Discussion. The generic determination of leaves 

of fan palms is highly problematic; some probably 

can be determined, but the type known in the 

Kushtaka Formation is too generalized to determine 

to genus. Palms are especially abundant at locality 

11160 and can be seen on numerous blocks in this 

slumped area. A large palm leaf that has an 

elongated hastula (and would thus represent 

Sabalites) was seen on a large block near locality 

11158 but was not collected.

The Ravenian occurrences of palms are not the 

most northerly known. Rollick (1936, pi. 21) 

illustrated an obvious fan palm from Paleocene rocks 

in the Matanuska Valley at about lat 61.5° N., and 

new collections from Paleocene rocks at about lat 62° 

N. contain a species different from Rollick's. A 

collection of specimens of apparent Eocene age from 

lat 65.5° N. along the Yukon River also contains a 

fan palm. Koch (1963) considers some seeds from the 

Paleocene of Greenland at about lat 70°N. to represent palm; Chancy (1967, p. 221) considers these 

seeds to be "***Of rather doubtful status," but 

inasmuch as Chancy has not stated the morphologic 

basis for his conclusion, I think Koch's determination should be accepted.

Specimen. USNM 43258.

Occurrence. Kushtaka Formation, Iocs. 9389, 

11160.

Genus and species indetermined

Discussion. Numerous fragments of palm leaves 

occur at several localities. None of these fragments 

are attached to a leaf-base, and thus are generically 

indeterminable.

Specimen. USNM 43279.

Occurrence. Kushtaka Formation, Iocs. 11158, 

11165; Kulthieth Formation, Iocs. 11170, 11195.

Order SALICALES 

Family SALICACEAE

Members of this family are lacking in the 

Ravenian and Kummerian except for the occurrence 

of Populus and Salix in the one late Ravenian 

assemblage. Whether the Paleocene occurrences of 

Salix are valid is highly questionable. In describing 

the Paleocene S. aquilina, Brown (1962, p. 55) stated: 

"These fragmentary leaves are referred to Salix with 

some hesitation." The fragmentary leaves are 

neither sufficiently well described nor illustrated to 

substantiate the generic assignment. Insofar as I am 

aware, the oldest valid species of either Populus or 

Salix is in the subtropical assemblages from the 

Wind River Formation (latest early Eocene) of 

Wyoming (MacGinitie, in press); MacGinitie (oral 

commun., February 1968) also reports that the 

extant P. dimorpha Brandg. is common along 

streams in the lowland tropical area near Mazatlan, 

Mexico. In western North America, the members of 

the family appear to have diversified primarily in the 

upland areas of the Eocene and Oligocene ancestral 

Rocky mountains and later in the late Oligocene and 

Neogene of Alaska. Few members of the family 

apparently ever adapted to the Paratropical or 

Tropical Rain forests; I suspect that the group that 

gave rise to Salicaceae presumably in the Paleocene or early Eocene had an extended history in 

temperate vegetation. The entry of members of the 

family into temperate vegetation in any diversity 

appears to have occurred in the early Miocene; 

Salicaceae are notably rare or lacking in the various 

assemblages of the late Oligocene Bridge Creek flora 

and coeval assemblages in the Pacific Northwest. In 

Alaska, however, several species are known in the 

type Angoonian, Tsadaka, and Rex Creek assemblages. Similarly the family is more diverse in 

Seldovian assemblages from Alaska than in the 

assemblages from the Pacific Northwest. Salicaceae 

seem to be one of the few families of woody plants 

that can be considered truly boreal.

Genus Populus Linnaeus

Populus sp. 

Plate 26, figure 1

Discussion. The single fragmentary specimen 

has palmate venation, medial secondary veins 

parallel to the lateral primaries, camptodrome 

lateral secondary veins, closely spaced and sinuous 

intercostal tertiary veins, irregularly shaped areoles 

intruded by branching freely ending veinlets, and a
SYSTEMATICS 55

nonentire margin. The combination of these features 

indicates that the specimen represents Populus, 

although the determination should be considered 

tentative until more complete material is found.

Specimen. USNM 43259.

Occurrence. Kushtaka Formation, loc. 11167.

Genus Salix Linnaeus

Salix carbonensis Wolfe, new species

Plate 26, figures 11, 13

Diagnosis. Leaves simple; venation pinnate; 

shape broadly oval; base rounded, apex acute; six or 

seven pairs of secondary veins, broadly curving, 

camptodrome; marginal tertiary veins craspedodrome; intercostal tertiary veins numerous, closely 

spaced, slightly oblique to secondaries, typically 

branching; many closely spaced, small, narrowly 

rounded teeth; sinuses angular.

Discussion. The fossils resemble the leaves of the 

extant Salix wilsoni Seem, in having a broad shape 

and numerous small teeth. S. wilsoni, however, has 

more widely spaced secondary vedns and numerous 

intersecondary veins. The fossils also resemble in 

venation leaves of S. richardsoni Hook., but the 

extant species has sharp sickle-shaped teeth.

Holotype. USNM 43260.

Paratype.-USNM 43261.

Occurrence. Kushtaka Formation, loc. 11167.

Salix sp. 

Plate 26, figure 2

Discussion. The single fragmentary specimen 

has numerous, sharp, glandular teeth; secondary 

veins that depart at a low angle and are 

camptodrome; marginal tertiary veins that are 

craspedodrome and enter the teeth centrally; 

intercostal tertiary veins that are numerous and 

oriented almost perpendicular to the midrib; and 

fourth-order venation that is highly irregular. These 

features particularly those of the margin and the 

intercostal venation are characteristic of Salix. 

Considering, however, the fragmentary nature of the 

specimen, the generic determination should be 

considered tentative.

Specimen. USNM 43262.

Occurrence. Kushtaka Formation, loc. 11167.

Order JUGLANDALES 

Family JUGLANDACEAE

Genus Carya Nuttall 

Carya cashmanensis Wolfe

Plate 20, figure 8

Carya cashmanensis Wolfe, 1968, U.S. Geol. Survey Prof. Paper 

571, p. 15, pi. 1, figure 2; text figure 5.

Discussion. The Kushtaka leaflet compares well

with the type material from the Puget Group. 

Particularly significant are the conspicuously 

forking secondary veins and the numerous, sharp 

teeth.

Hypotype. USNM 43263.

Occurrence. -Kushtaka Formation, loc. 11165.

Genus Engelhardtia Leschenault 

Engelhardtia sp.

Plate 20, figure 7

Discussion. Although the base is missing, the 

specimen is thought to represent a leaflet; the angle 

of divergence of the secondary veins is considerably 

different on the two lateral halves of the lamina, 

particularly in the more basal part of the specimen. 

The counterpart of the figured specimen, moreover, 

appears to have an asymmetric shape in the more 

basal part. Leaflets that have similar numerous, 

closely spaced, and rounded teeth occur in both 

Juglandaceae (Engelhardtia) and Burseraceae (Canarium). In the latter genus, however, the leaflets 

have smoothly curving secondary veins that do not 

fork and that have small loops apical to the juncture 

of any two secondary veins; the secondary veins 

forming the basal part of such a loop would, if 

extended to the margin, form about a 45° angle with 

the margin, in both Engelhardtia and the fossil, the 

secondary veins fork conspicuously and form 

comparatively large loops; the vein forming the 

basal side would intersect the margin at about an 80° 

angle. In addition, the toothed leaves of Canarium 

have apical secondary veins that do not extend far 

along the margin, in contrast to the apical secondary 

veins of Engelhardtia and the fossil, which extend 

for a considerable distance. Among extant species of 

Engelhardtia, the fossil most closely resembles the 

leaflets of E. apoensis Elm., particularly in having 

numerous teeth. Most species of Engelhardtia 

(Engelhardtia) and all species of E. (Oreamunoa) 

have entire-margined leaflets.

The fossil is the only foilage of Engelhardtia from 

North America to be validly referred to the genus. 

The fossil record of the genus in North America is 

comprised, except for the Kushtaka specimen, of the 

characteristic involucres. Most of the involucres 

have venation that is palmate and characteristic of 

Oreamunoa, that is, in the lobes of the involucre are 

typically two veins that flank the midrib and extend 

at least half way to the apex without looping. In the 

subgenus Engelhardtia, the lobes lack the flanking 

veins. Despite having the venation of Oreamunoa, 

most west American Tertiary involucres lack the 

large posterior lobe characteristic of Oreamunoa and 

in this respect are similar to Engelhardtia. These
56 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

1 cm

FIGURE 9. Marginal venation of Platycarya. A, P.pseudobrauni, 

hypotype, USNM 43267, loc. 11170. B, P. strobilacea, USGS 

Ref. Colln. 253.

involucres may represent an extinct subgenus (or 

genus if Oreamunoa is considered generically 

distinct from Engelhardtia). That Engelhardtia 

(Engelhardtid) was present in the American Tertiary 

is indicated, however, both by the Kushtaka leaflet 

and some involucres from the West Branch Creek 

locality of the Clarno Formation. During the 

Angoonian, the genus Engelhardtia is represented 

in western North America by the probably extinct 

subgenus mentioned previously; the genus had 

almost certainly adapted to temperate climates, as 

indicated by the presence of characteristic involucres in the Angoonian rocks of the central Alaska 

Range.

Specimen. USNM 43264.

Occurrence. Kushtaka Formation, loc. 11165.

Genus Juglans Linnaeus

Juglans sp. 

Plate 26, figure 3

Discussion, Although fragmentary, the specimen is well preserved. The reference to Juglandaceae 

is certain because of the intercostal tertiary veins, 

which are uniformly spaced perpendicular to the 

secondary veins, and the forking craspedodrome 

secondary veins. The broad mucronate teeth that are 

elongated basally and the craspedodrome secondary 

veins that enter the teeth centrally indicate that the 

specimen represents a species of the section 

Cardiocaryon of Juglans.

Specimen. USNM 43265.

Occurrence. Kushtaka Formation, loc. 11167.

Genus Platycarya Siebold and Zuccarrini 

Platycarya pseudobrauni (Hollick) Wolfe, new combination

Plate 4, figure 4; plate 5, figures 1,2; plate 21, figure 2b; text figure 9 

Ulmus pseudobraunii Hollick, 1936, U.S. Geol. Survey Prof. Paper

182, p. 105, pi. 58, figs. 1-3. 

Planera aquaticiformis Hollick, 1936, U.S. Geol. Survey Prof.

Paper 182, p. 107, pi. 57, fig. 7. 

Fraxinus lateralis Hollick, 1936 (part), U.S. Geol. Survey Prof.

Paper 182, p. 164, pi. 105, fig. 6 only.

Discussion. Pollen of Platycarya is of wide 

occurrence in the Eocene of North America (Leopold 

and MacGinitie, 1972), although no North American 

fossil leaflets have been previously referred to the 

genus. Hollick's material listed in the synonymy has 

considerably broader teeth than is typical of the one 

extant species of the genus, but specimens from one 

locality (11170) show an intergradation from broad 

teeth to the sharp teeth as in P. strobilacea Sieb. and 

Zucc. Considering the variation displayed by the 

Alaskan material, it is possible that the leaflets 

referred to a new genus, Vinea (Wolfe, 1968), may 

eventually prove to belong to Platycarya. Platycarya 

was almost certainly adapted to much warmer 

conditions during the Tertiary; the fruits of the genus 

are abundant in the London Clay assemblage (Reid 

and Chandler, 1933) in association with abundant 

Nipa, Menispermaceae, and Icacinaceae. Although 

predominately a member of the Mixed Mesophytic 

forest today, Platycarya also forms a minor element 

in the subtropical broad-leaved evergreen forest of 

China (Wang, 1961, p. 145). I do not think that the 

present tolerance, however, of the species of a 

monotypic genus such as Platycarya can be 

extended to include extinct species; it would in fact be 

remarkable if an extinct species had the same 

tolerances as an extant species of the same genus. 

The numerous occurrences of Platycarya pollen in 

Eocene assemblages that represent subtropical to 

Paratropical Rain forest is a clear indication that 

some extinct species of the genus tolerated considerably warmer climates than does the extant species.

Lectotype. USNM 38895 (Hollick's pi. 58, figs. 1, 

2).

Hypotypes. USNM 38893, 39063, 43266, 43267, 

245726.

Occurrence. Kushtaka Formation, Iocs. 3847, 

11165, 11166; Kulthieth Formation, Iocs. 11170, 

11195.

Genus Pterocarya Kunth 

Pterocarya pugetensis Wolfe 

Plate 21, figure 4; text figure 10

Pterocarya pugetensis Wolfe, 1968, U.S. Geol. Survey Prof. Paper 

571, p. 16, pi. 1, figs. 3-5; text fig. 8.

Discussion. The specimen figured in this report 

has the smoothly looping secondary veins and the 

numerous, closely spaced, and sharp teeth of 

Pterocarya pugetensis. The lineage to which this 

species is thought to belong is well represented in the 

Mixed Mesophytic forest of the Neogene of both the 

Pacific Northwest and Alaska (Wolfe, 1966, p. B15; 

1968, p. 17).

A single incomplete specimen from locality 11167 

has many small, rounded, broad teeth, tertiary veins
SYSTEMATICS 57

1 cm

FIGURE 10. Marginal venation of Pterocarya pugetensis. Hypotype, USNM 43269, loc. 11167.

that are craspedodrome, secondary veins that fork 

and are camptodrome, and uniformly intercostal 

tertiary veins that are oriented perpendicular to the 

secondary veins. These features indicate that the 

specimen represents Pterocarya and the spacing and 

shape of the teeth is the same as in P. pugetensis.

Hypotype. USNM 43268, 43269.

Occurrence. Kushtaka Formation, Iocs. 9389, 

11165, 11167.

Order FAG ALES 

Family BETULACEAE

This family is typically thought of as a boreal 

temperate family. The fossil occurrences of the 

family, however, indicate that the family has a long 

history in the Paratropical and subtropical rain 

forests. From palynologic data, the earliest definite 

record of the family is the occurrence of Alnus in 

Maestrichtian rocks of the Rocky Mountain region 

(R. Tschudy, written commun., January l969).Alnus 

is also known in the Paleocene, but the pollen is 

scarce in the material described from basal 

Paleocene rocks on Greenland; Manum (1962, p. 119) 

suggests that Alnus was not a significant part of the 

vegetation in that area at that time. Leaves of certain 

betulaceous character that have been referred to 

Corylus or Corylites and Betula have been recorded 

from a number of localities in Paleocene rocks in 

Alaska (Wolfe, 1966) and the high plains region of 

Canada and conterminous United States (Brown, 

1962), as well as from Paleocene rocks of Greenland 

(Koch, 1963) and Great Britain. The diversity of the 

family during the Paleocene does not appear to have 

been great. The Paleocene occurrences are in

assemblages that, on physiognomic characters of 

the foliage, represent warm temperate to subtropical 

forests. The Eocene occurrences of the family are 

primarily those of Alnus; leaves of this genus have 

been recorded from a number of assemblages in 

western North America that represent subtropical or 

Paratropical Rain forests (MacGinitie, 1941; Wolfe, 

1968). Chandler (1964) similarly records Alnus from 

British Eocene assemblages that represent Paratropical or Tropical Rain forests. The leaves are the 

types that have simple teeth, that is, the teeth are not 

grouped into conspicuous "lobes." The secondary 

veins of many of the leaves tend to be camptodrome. 

These types of alder leaves are today more southern 

in distribution than the "lobed" types. Note, 

however, that Alnus does not appear to have been 

diverse in the Paratropical Rain forest; a few 

lineages apparently were represented. Distinctive 

types, such as the latest Eocene A. cuprovallis Axelr., 

have a pronounced tendency towards the grouping of 

the teeth into "lobes." The dominant types of alders, 

however, in the late Eocene and early to middle 

Oligocene cool assemblages are related to the alders 

of the Paratropical Rain forest. The genus persisted 

in the Paratropical Rain forest in western North 

America through at least the Kummerian. The 

Angoonian assemblages both in the Pacific Northwest and Alaska display a diversity of Betulaceae, 

particularly Alnus, that is unmatched in the earlier 

Paleogene. The groups that have "lobed" types of 

leaves became increasingly diverse. The genus 

reached its acme in western North America during 

the early and middle Miocene.

The other genera of Betulaceae have a poorly 

documented record. Betula steuensoni Lesq. represents, at least in part, a species of Ostrya or 

Carpinus, as indicated both by the lack of branching 

freely ending veinlets and the sharp apical bend of 

the secondary veins within the teeth. Carpinus has 

been recorded from the British Eocene tropical 

assemblages. Betula has been described from the 

Claiborne flora, but, without knowing the ultimate 

venatiori pattern, it is not possible to determine 

whether Betula or Ostrya-Carpinus is represented. 

In the later Eocene, Betula is known both from the 

upland Republic assemblage of Washington and the 

lowland John Day Gulch assemblage of Oregon. In 

the early Oligocene, Betula wood is known from the 

Clarno nut bed assemblage (Chandler, 1964); 

although it is possible that the wood was derived 

from more upland vegetation, I think that it is 

improbable that the upland vegetation bordering the 

Clarno basin was any cooler than subtropical. By the 

late Oligocene, Betula was apparently more diverse
58 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

than previously, but the diversity attained was not of 

as great a magnitude as that of Alnus. Ostrya and 

Carpinus were apparently of little diversity in the 

Mixed Mesophytic forest of western North America, 

in contrast to their considerable diversity in the 

Mixed Mesophytic forest of the Japanese Cenozoic.

Corylus has an extremely poor record. Some 

Paleocene leaves of Corylites may represent Corylus 

or a closely allied genus, but Eocene and younger 

records of the genus are generally poor. Material 

from the late Oligocene and late Miocene of Alaska 

referred to Corylus harrimani Knowlt. and C. 

chuitensis Wolfe, respectively, have small areoles 

intruded by typically unbranching veinlets; the 

lineage represented probably represents Ostryopsis, 

which today is a monotypic genus inhabiting the 

northern, more temperate regions of China.

The Betulaceae, therefore, display several distribution patterns. The best documented pattern is that 

of Alnus, which during the Paleogene particularly 

the Eocene was a little diversified genus of the 

Paratropical and subtropical Rain forests. The 

cooling of the middle late Oligocene opened wide 

areas of temperate climate into which Alnus 

apparently extended and diversified rapidly. Some 

of the other genera were also represented in the 

Paratropical Rain forest of the Paleogene but poorly 

in comparison to alders. These genera also apparently diversified from the late Oligocene through the 

later Cenozoic, but a genus such as Carpinus never 

attained the diversity in western North America that 

it did in Asia. Ostryopsis, however, from its first 

appearance in the late Oligocene of Alaska, did not 

have any tendency to diversify and, at least from the 

late Oligocene, has been a member of temperate 

vegetation.

Genus Alnus Linnaeus

Alnus cuprovallis Axelrod

Plate 26, figure 7; text figure 11

Alnus cuprovallis Axelrod, 1966, California Univ. Pubs. Geol. Sci., 

v. 59, p. 66, pi. 11, figs. 1-13.

Discussion. Axelrod (1966b, p. 66) states that the 

leaves of Alnus cuprovallis and A. crispa differ 

"***only slightly in that the secondaries in the 

fossils tend to be more curved distally." Note, 

however, that in A. cuprovallis the teeth are broad, 

the maximum number of teeth per secondary vein is 

seven, the intercostal tertiary veins are spaced about 

1.5 mm apart, and the areoles are formed by fourthorder venation and are about 1 mm across. In A. 

crispa the teeth are elongated and sharp, any given 

leaf typically has a secondary vein that feeds at least 

9-10 teeth, the intercostal tertiary veins are spaced 

about 0.5 mm apart, and the areoles are formed by

1 cm

FIGURE 11. Marginal venation of Alnus cuprovallis. A, hypotype, USNM 43270, loc. 11167. B, paratype, UCMP 20003, 

Copper Basin flora.

fifth-order venation and are about 0.2-0.3 mm 

across. Any relationship between A. crispa and A. 

cuprovallis is distant. The leaves of A. incana, 

however, are more closely related to A. cuprovallis. 

This is indicated by the following characters of the 

extant species: broad teeth that can be mucronate; 

typically 3-6 teeth per secondary vein; the intercostal tertiary veins that are spaced about 1 mm 

apart; and the areoles that are formed by fourthorder venation and are about 0.5 mm across. A. 

cuprovallis appears to be the oldest known member 

of the A. incana lineage, which is well represented in 

Alaska in the late Oligocene and Neogene.

Hypotype. USNM 43270.

Occurrence. Kushtaka Formation, loc. 11167.

Alnus martini Wolfe, new species 

Plate 5, figures 3, 5: plate 21, figure 3; text figure 12

Diagnosis. Leaves simple; venation pinnate; 

base acute to typically broadly rounded; apex acute; 

shape oval; 7-11 pairs of broadly curving secondary 

veins; camptodrome or craspedodrome and entering 

the teeth centrally; intercostal tertiary veins 

uniformly spaced, approximately perpendicular to 

secondary veins; marginal tertiary veins craspedodrome; typically entering teeth along apical sides; 

typically 3 to 4 sharp to narrowly rounded and small 

teeth per secondary vein.

Discussion. These leaves are related to the Asian 

alders, such as Alnus japonica Sieb. and Zucc. and 

its allies, in having some camptodrome secondary
SYSTEMATICS 59

FIGURE 12. Marginal venation of Alnus martini. Holotype, 

USNM 43271, loc. 11158.

veins and some marginal tertiary veins that enter 

the teeth along the apical side. In A. japonica and its 

allies, however, the craspedodrome secondary veins 

typically enter the teeth along the apical sides; an 

exception to this is A. fauriei Lev., but the cuneate 

base, obovate shape, and irregular spacing of the 

intercostal tertiary veins readily distinguish A. 

fauriei from A. martini. Neither A. martini nor A. 

scotti appear to be closely related to any extant 

species.

Alnus martini is named in recognition of G. C. 

Martin's work in the Katalla district.

Holotype. USNM 43271.

Paratypes. 43272, 43279.

Occurrence. Kushtaka Formation, Iocs. 11158, 

11163; Kulthieth Formation, loc. 11170.

Alnus scotti Wolfe, new species 

Plate 4, figures 1-3

Artocarpidium alaskanum Hollick, 1936 (part, nontypic), U.S. 

Geol. Survey Prof. Paper 182, p. 108, pi. 59, figs. 1-4.

Diagnosis. Leaves simple; pinnately veined; 

base rounded, typically slightly asymmetric; apex 

acuminate; 6-9 pairs of craspedodrome secondary 

veins; intercostal tertiary veins oriented obliquely to 

secondary veins; marginal tertiary veins craspedodrome; teeth large, 2-3 per secondary vein, typically 

separated by arcuate sinuses.

Discussion. Most alders, unlike Alnus scotti, 

have leaves that have intercostal tertiary veins 

oriented perpendicular to the secondary veins, 

although the tertiaries of A. sinuata Rydb. are also 

oblique to the secondary veins. Also unusual are the 

arcuate sinuses, but a few species, for example A. 

nitida Endl., have arcuate sinuses. The combination, 

however, of large teeth, oblique tertiary veins, and 

arcuate sinuses is unique.

This species is named in recognition of the work of 

R. A. Scott on the Paleogene floras of western North 

America.

Holotype. USNM 38899.

1 cm

FIGURE 13. Marginal venation of Alnus sp. USNM 43277, loc.

11167.

Paratypes.-USNM 43273, 43274. 

Occurrence. Kushtaka Formation, Iocs. 

11158; Kulthieth Formation, loc. 11170.

3847,

Alnus sp. 

Plate 26, figures 4, 10

Discussion. Fragmentary specimens from the 

late Ravenian locality represent a species of Alnus 

that has a cordate base and slight indications of 

"lobing." The rounded teeth, as well as these other 

features, indicate a relationship to the strongly 

"lobed" A. barnesi Wolfe.

Specimens. USNM 43275, 43276a.

Occurrence. Kushtaka Formation, loc. 11167.

Alnus sp. 

Text figure 13

Discussion. One specimen of an alder leaf has 

very sharp teeth reflexed basally; although insufficient for specific determination, it does not belong to 

either of the other two species of Alnus represented in 

the same collection. The cuneate base and tertiary 

veins oriented obliquely to the secondary veins 

indicate a possible relationship to A. healyensis 

Wolfe; the Seldovian species, however, has considerably more teeth per secondary vein.

Specimen.-USNM 43277.

Occurrence. Kushtaka Formation, loc. 11167.

Alnus sp. 

Plate 28, figure 3

Discussion. Two specimens from the Kummerian 

have small, rounded to sharp teeth and marginal, 

tertiary veins that enter the teeth along the apical 

sides. These leaves thus resemble A. martini, but the
60 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Kummerian species has a base apparently decurrent 

along the petiole and fewer (5-6) secondary veins.

Specimen. USNM 43278.

Occurrence. Kushtaka Formation, loc. 9891.

Genus Betula Linnaeus 

Betula sp.

Plate 28, figure 4

Discussion. A single battered specimen has a 

sharply serrate margin and craspedodrome secondary veins that bend sharply apically within the 

teeth. The ultimate venation is poorly preserved, but 

the areoles appear to be intruded by branching 

veinlets, and thus the specimen probably represents 

Betula.

Specimen. USNM 43280.

Occurrence. Katalla Formation, Split Creek 

Sandstone Member, loc. 11168.

Family FAGACEAE 

Genus Dryophyllum Debey 

Dryophyllum pugetensis Wolfe 

Plate 21, figures 1, 5, 6

Dryophyllum pugetensis Wolfe, 1968, U.S. Geol. Survey Prof. 

Paper 571, p. 18, pi. 3, figs. 4, 8.

Discussion. Two fundamental types of leaves 

have been assigned to Dryophyllum. One type, as 

represented by D. pugetensis, has marginal venation 

that is typical of genera such as Castanopsis, 

Lithocarpus, and Quercus; in this type, the apical 

tertiary branch that is nearest the margin is 

aparallel to the intercostal tertiary veins and 

approximately parallels the margin. In the second 

type, as represented by D. tennesseensis Berr., the 

marginal venation is similar to Fagus, that is, the 

apical tertiary branch nearest the margin typically 

is parallel to the intercostal tertiary veins. This is a 

fundamental difference in Fagaceae and indicates to 

me that two different genera are represented. I have, 

however, not seen material of the type species of 

Dryophyllum and thus cannot determine whether D. 

pugetensis is validly assigned to that genus. 

Whether Dryophyllum contains ancestors of other 

fagaceous genera is unknown. The extremely thick 

texture of the leaves of D. pugetensis indicates a 

probable evergreen habit.

Hypotypes.-USNM 43281-43283.

Occurrence. Kushtaka Formation, Iocs. 3842, 

9389, 11165.

Genus Fagus Linnaeus

Fagus sp. 

Plate 28, figure 2

Discussion. A fragmentary leaf has numerous 

straight secondary veins that end in small teeth.

What may represent the same species also occurs in 

rocks assigned to the basal part of the Poul Creek 

Formation.

Specimen. USNM 43284.

Occurrence. Katalla Formation, Split Creek 

Sandstone Member, loc. 9552.

Order URTICALES

Family ULMACEAE

Genus Girroniera Gaudich

Girroniera sp.

Plate 5, figure 7

Discussion. The specimen has an entire margin 

except for a few rounded small teeth a short distance 

from the apex. The secondary veins are camptodrome and the teeth are entered by tertiary veins. 

The leaf also has a pronounced drip-tip. Similar 

leaves are present in extant species of Girroniera; 

these leaves can have an entire margin or a few 

rounded teeth near the apex.

Specimen. USNM 43285.

Occurrence. Kushtaka Formation, loc. 11158.

Girroniera sp. 

Plate 5, figure 8

Discussion. The leaf is similar to the specimen of 

Girroniera from the Kushtaka Formation, but differs 

in having sharper teeth and almost twice as many 

secondary veins. The secondaries also loop closer to 

the margin in the Kulthieth specimen.

Specimen. USNM 43286.

Occurrence. Kulthieth Formation, loc. 11170.

Genus Ulmus Linnaeus 

Ulmus sp.

Plate 23, figure 1

Discussion. Four specimens have an asymmetric 

base, numerous and almost straight secondary veins 

that are craspedodrome, craspedodrome marginal 

tertiary veins that depart from the basal sides of the 

secondaries, and a finely compound serrate margin. 

These features indicate that the specimens represent 

Ulmus, although the lack of preservation of the 

intercostal venation does not allow application of a 

specific epithet. In characters that are preserved, the 

specimens are similar to leaves of U. newberryi 

Knowlt., except that the latter has considerably 

larger teeth.

Ulmus has a long history in the subtropical forest; 

ulmaceous pollen grains (these could represent 

Ulmus, Zelkova, or possibly an ancestral genus) are 

first known in Paleocene rocks in warm temperate 

and subtropical forests. In the Eocene, representatives of the genus are found in both Paratropical
SYSTEMATICS 61

Rain and subtropical forests; in the Eocene the genus 

is also known in temperate forests such as 

represented by the Republic assemblage. In the early 

to middle Oligocene, the genus is represented 

primarily in the upland forests, and from the late 

Oligocene on it has been a significant element of the 

warm temperate forests.

Specimen. USNM 43287.

Occurrence. Kushtaka Formation, Iocs. 9389, 

11165.

Order RANALES 

Family TETRACENTRACEAE

Genus Tetracentron Oliver

Tetracentron piperoides (Lesquereux) Wolfe,

new combination

Plate 5, figure 4; plate 22, figure 3; plate 28, figure 7

Zizyphus piperoides Lesquereux, 1878, Harvard Coll. Mus. Comp.

Zoology Mem., v. 6, no. 2, p. 28, pi. 8, figs. 10, 11. 

Zizyphus microphyllus Lesquereux, 1878, Harvand Coll. Mus.

Comp. Zoology Mem., v. 6, no. 2, p. 28, pi. 8, fig. 9. 

Zizyphus californica Knowlton and Cockerell, 1919, U.S. Geol.

Survey Bull. 696, p. 659. 

Ceanothus idahoensis Brown, 1935, Journ. Paleontology, v. 9,

p. 581, pi. 69, fig. 8.

Cercidiphyllum elongatum Brown, 1939 (part), Jour. Paleontology, v. 13, p. 494, pi. 55, figs. 1, 2, 7 only; 56, fig. 9 only. 

MacGinitie, 1941 (part), Carnegie Inst. Washington Pub. 534,

p. 112, pi. 12, fig. 4; pi. 13, fig. 4 only; pi. 14, figs. 1, 2 only. 

Becker, 1961 (part), Geol. Soc. America Mem. 82, p. 66, pi. 10,

fig. 8; pi. 19, figs. 1-5 only. 

Cercidiphyllum piperoides (Lesquereux) LaMotte, 1952, Geol. Soc.

America Mem. 51, p. 120.

Wolfe, 1968, U.S. Geol. Survey Prof. Paper 571, p. 19. 

Trochodendroides zaddachi Sanborn non Populus zaddachi Heer.

Sanborn, 1935, Carnegie Inst. Washington Pub. 465, p. 16,

pi. 3, fig. 2. 

Populus smilacifolia auct. non Newberry. Jennings, 1920,

Carnegie Mus. Mem., v. 8, p. 409, pi. 26, fig. 4. 

Populus glandulifera auct. non Heer. Knowlton, 1899, U.S. Geol.

Survey Mon. 32, p. 694, pi. 84, fig. 1. 

Populus daphnogenoides auct. non Ward. Knowlton, 1899, U.S.

Geol. Survey Mon. 32, p. 695, pi. 84, fig. 2.

Populus speciosa auct. non Ward. Knowlton, 1899, U.S. Geol. Survey Mon. 32, p. 694, pi. 84, fig. 3. 

Populus xantholithensis Knowlton, 1899, U.S. Geol. Survey Mon.

32, p. 695, pi. 85, figs. 1, 2. 

Zizyphus serrulata auct. non Ward. Knowlton, 1899, U.S. Geol.

Survey Mon. 32, p. 740, pi. 101, figs. 4, 5. 

Cercidiphyllum crenatum Brown non Grewia crenata Unger.

Brown, 1935 (part), Journ. Paleontology, v. 9, p. 575, pi. 68,

figs. 1, 6, only. 

Brown, 1939 (part), Jour. Paleontology, v. 13, p. 494, pi. 56,

fig. 7. 

Trochodendroides arctica Berry non Populus arctica Heer. Berry,

1926, Canada Geol. Survey Bull. 42, p. 109, pi. 13, figs. 1-4. 

Populus richardsoni auct. non Heer. Lesquereux, 1878, U.S. Geol.

Survey Terr. Kept., v. 7, p. 177, pi. 22, figs. 10-12. 

Populus rotundifolia auct. non Newberry. Dawson, 1895, Royal

Soc. Canada Trans., 2nd ser., v. 1, sec. 4, p. 147, text fig. 10.

Discussion. Few previous citations of the occurrence of Cercidiphyllum in the Tertiary of North 

America have proved to be valid. The Paleocene 

specimens assigned to this genus represent several 

different genera and species, primarily Cocculus 

flabella (Newb.) Wolfe, Trochodendroides serrulata 

(Ward) Wolfe, and Dicotylophyllum richardsoni 

(Heer) Wolfe (Wolfe, 1966, p. B9-B10). A few 

unfigured specimens from the Fort Union Formation 

do, however, represent Cercidiphyllum. Similarly 

the citations of Cercidiphyllum in the Eocene have 

proved to be invalid, although unfigured material 

from probable early Eocene rocks in north-central 

Washington does represent Cercidiphyllum, and 

Scott (in Chandler, 1964, p. 58) reported wood of the 

genus from the Clarno nut bed (latest Eocene). The 

late Oligocene and Miocene occurrences of Cercidiphyllum crenatum (Ung.) R. W. Br. also are valid 

citations of the genus.

The leaves of extant Tetracentron and Cercidiphyllum are indeed difficult to distinguish. Leaves of 

Tetracentron are, of course, typically more elongated 

than those of Cercidiphyllum; on this basis alone, 

the Eocene and early Oligocene specimens synonymized here are assignable to Tetracentron. In addition, 

the teeth of Tetracentron are typically larger and 

more equal sided than those of Cercidiphyllum; the 

items in the synonymy are again closer to 

Tetracentron. Perhaps most significant, however, is 

that in Tetracentron veins connect the glandular 

region of the tooth to the adjacent sinuses, but in 

Cercidiphyllum such veins are absent; wellpreserved fossil material, including topotypic material from Chalk bluffs, indicates that "Zizyphus" 

piperoides is Tetracentron. Sanborn (1935) also 

compared her specimens of "Trochodendroides 

zaddachi" to Tetracentron.

The various seeds and fruits associated at some 

localities with these fossil leaves were thought to 

represent Cercidiphyllum. Chandler (1961, p. 84), 

however, has demonstrated that the English 

material thought by Brown (1962) to be Cercidiphyllum is of unknown affinities.

MacGinitie (1941, p. 112) noted that the early 

Eocene specimens of this taxon are more elongated 

than the late Eocene specimens, and that the early 

Eocene material has a regularly and more finely 

crenate margin whereas the late Eocene material 

has an irregularly and more coarsely crenate 

margin. Two species may indeed be represented, but 

the almost complete transition of suites of these 

leaves in the Puget Group from the early Eocene 

through the early Oligocene makes it difficult to delineate more than one taxon. The Alaskan material 

collected thus far is closer to the Puget material from
62 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

the Fultonian and Ravenian than to Franklinian 

leaves of the species.

Hypotypes. USNM 43288-43290.

Occurrence. Kushtaka Formation, Iocs. 9389, 

11165, 11169; Kulthieth Formation, loc. 11170.

Family MAGNOLIACEAE

Genus Magnolia Linnaeus

Magnolia reticulata Chaney and Sanborn

Plate 30, figure 7

Magnolia reticulata Chaney and Sanborn, 1933, Carnegie Inst. 

Washington Pub. 439, p. 70, pi. 13, figs. 1, 2, 4.

Discussion. The fossil has numerous secondary 

and intersecondary veins that depart from the 

midrib at a high angle; the secondaries extend with 

little or no curvature towards the margin, and loop 

sharply. The intercostal venation, which is considerably weaker than the secondaries, forms a uniform 

quadrangular mesh. In shape and venation, therefore, the fossil falls within the range of variation of 

the Goshen species.

An undescribed species of Magnolia from the early 

Miocene Collawash assemblage of Oregon has 

venation that is similar to the Goshen and Katalla 

species. The Collawash species has fewer intersecondary veins and tends toward an obovate shape, 

but otherwise it is similar to M. reticulata.

Hypotype. USNM 43291.

Occurrence. Katalla Formation, Split Creek 

Sandstone Member, loc. 11168.

Family ILLICIACEAE

Genus Illicium Linnaeus

Illicium sp.

Plate 15, figure 3

Discussion. The single specimen, which lacks the 

basal part of the lamina, has several features that 

indicate an assignment to Illicium: (1) widely spaced 

secondary veins that loop sharply at the margin to 

form a trapezoidal intercostal area, (2) an extensive 

series of loops of marginal tertiary veins that are 

elongated parallel to the midrib, (3) widely and 

irregularly spaced, rounded, small teeth, and (4) an 

attenuated apex. The fossil is particularly similar to 

the extant /. lanceolatum A. C. Sm. from southern 

China.

Specimen. USNM 43292.

Occurrence. Kushtaka Formation, loc. 11159.

Family MENISPERMACEAE

Genus Anamirta Colebrooke

Anamirta milleri Wolfe, new species

Plate 6, figure 6; text figure 14

Diagnosis. Leaves simple; venation palmate; 

base cordate, apex acuminate; seven primary veins, 

first lateral pair extending about half the distance to

B

1 cm

FIGURE 14. Marginal venation of Anamirta. A, A. 

milleri, holotype, USNM 43293, loc. 11170. B, A. cocculus, USGS Ref. Colln. 491.

apex; second pair parallel to lateral secondaries of 

first lateral primaries, forming a series of 5-7 loops; 

lateral primaries and all secondary and marginal 

tertiary veins camptodrome; intercostal tertiary 

veins spaced about 0.4 cm apart, irregularly 

branching, oriented approximately perpendicular to 

midrib; marginal vein present.

Discussion. The prominent marginal vein, irregularly branching intercostal tertiaries, palmate 

venation, and cordate base, ally these fossils to 

Menispermaceae. The lack of craspedodrome secondary or tertiary veins, as well as the cordate base, 

indicates that the most closely allied genera are 

Anamirta, Tinomiscium, and Tinospora. In both 

Tinomiscium and Tinospora the intercostal tertiary 

veins are widely spaced; the spacing of these veins is, 

however, similar in the fossils and in Anamirta. A. 

cocculus Wight and Arn. has leaves very similar to 

the fossils, but the fossils are widest at the base 

whereas the leaves of A. cocculus are widest above 

the base.

Holotype. USNM 43293.

Occurrence. Kulthieth Formation, loc. 11170.

Genus Calkinsia Wolfe 

Calkinsia plafkeri Wolfe, new species 

Plate 6, figure 8; plate 23, figures 3, 4, 6, 7b 

Hyperbaena dilleri (Knowlton) Wolfe, 1968 (part, nontypic), U.S. 

Geol. Survey Prof. Paper 571, p. 21, specimens listed from 

Iocs. 8640, 9680, 9731, 9738.

Diagnosis. Leaves simple; venation palmate; 

shape oval; base acute, apex acuminate; length 2-12 

cm, width 1.5-7 cm; three or four pairs of broadly 

convex medial secondary veins, departing at an 

angle of 20°-30°; lateral secondaries departing at an 

angle of 40°-80°; tertiary veins departing from 

midrib at an angle of 70°-80°.
SYSTEMATICS 63

Discussion. As noted previously (Wolfe, 1968, p. 

21), the fossils have conspicuously branched 

intercostal tertiary veins; the spacing of these veins, 

however, is much wider in extant species of 

Hyperbaena than in the fossils. Leaves of Hyperbaena, moreover, typically do not have such a broad 

oval shape. In the spacing of the nervilles and broad 

shape, the fossils can be matched most closely by 

Calkinsia franklinensis Wolfe, the type species of the 

genus. C. franklinensis, however, typically has a 

broader and obovate shape and the apex is narrowly 

rounded. The tertiaries, moreover, typically depart 

from the midrib at an angle of 90° in C. 

franklinensis.

Morphologically, therefore, Calkinsia franklinensis and C. plafkeri are closely related and probably 

represent the same phylad. C. franklinensis is 

known in the Franklinian and lower Fultonian and 

C. plafkeri is known in the lower through upper 

Ravenian. The tendency towards a narrower shape 

is continued in this lineage by C. dilleri, which is 

known in the Comstock and Upper Clarno assemblages of Oregon.

Calkinsia has been likened to the extant genus 

Hypserpa; this resemblance is enhanced by the 

configuration of the tertiary veins between the first 

lateral primaries and the most basal medial 

secondaries. Material of Hypserpa studied at the 

time the original diagnosis of Calkinsia was written 

did not have lateral primaries extending into the 

apical half of the lamina. A specimen of Hypserpa 

nitida Miers, since added to the University of 

California Herbarium (sheet no. 1011338), bears 

leaves that have the first lateral primaries extending 

slightly more than half way to the apex. Calkinsia, 

therefore, is probably most closely related to 

Hypserpa. Seeds of Eohypserpa have been found in 

the late Eocene Clarno nut bed (Scott in Chandler, 

1964, p. 58), where I have collected leaves of C. dilleri. 

Calkinsia and Eohypserpa may possibly represent 

organs of the same genus.

This species is named in recognition of George 

Plafker's invaluable help in the preparation of this 

report.

Holotype. USNM 43296.

Paratypes. USNM 43294-43295, 43297, 43298, 

43312.

Occurrence. Kushtaka Formation, Iocs. 11157, 

11164, 11165.

Genus Cocculus Miers 

Cocculus sp.

Plate 6, figure 5

Discussion. The seven primary veins, camptodrome lateral secondary veins that form angular

loops, the craspedodrome lateral tertiary veins that 

fuse with the marginal vein, the widely spaced and 

branching intercostal tertiaries, the cordate base, 

and the rotund shape of the fossil can be matched by 

an undescribed species from the early Ravenian of 

the Puget Group. The upper margin of the Alaskan 

specimen appears to have been untoothed, but the 

Puget Group specimens vary from having an entire 

to a toothed margin. In shape and overall venation 

pattern, the early Ravenian species is similar to the 

Paleocene Cocculus flabella (Newb.) Wolfe and to the 

Neogene or extant species such as C. auriculata 

(Heer) Wolfe and C. trilobus DC. The early Ravenian 

species, however, has areoles that are about 1 mm or 

slightly more in width, but the areoles of C. 

auriculata average about 2 mm or slightly more in 

width. The early Ravenian species, which inhabited 

Paratropical Rain forest, is probably in the ancestry 

of the Neogene and Quaternary species of the Mixed 

Mesophytic forest.

Specimen. USNM 43299.

Occurrence. Kushtaka Formation, loc. 11166.

Cocculus sp. 

Plate 26, figure 5

Discussion. The fragmentary specimen has palmate venation, lateral secondary veins that are 

camptodrome, lateral tertiary veins that are craspedodrome into the marginal vein, and areoles 

intruded by veinlets that branch two or three times. 

The species represented probably does not belong to 

the group that produced the toothed, deeply cordate 

species of Cocculus that were or are members of the 

Mixed Mesophytic forest. The mixed mesophytic 

group is represented in the early Ravenian Steel's 

Crossing assemblage of Washington and in the 

Alaskan early Ravenian by a species that has 

areoles about twice as large as the species under 

discussion. It is more probable that the late 

Ravenian species was separately derived from 

another tropical group of the genus.

Specimen. USNM 43300.

Occurrence. Kushtaka Formation, loc. 11167.

Genus Diploclisia Miers 

Diploclisia sp.

Plate 6, figure 7; text figure 15

Discussion. The conspicuously forking lateral 

primary veins and forking, craspedodrome lateral 

secondary veins indicate that this fossil is a member 

of Diploclisia. The incompleteness of the specimen, 

however, does not allow a satisfactory specific 

diagnosis or description.

Specimen. USNM 43301.
64 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

FIGURE 15. Marginal venation of Diploclisia. A, D. sp., 

USNM 43301, loc. 11170. B, D. chinensis, USGS Ref. Colln. 

242.

Occurrence. Kulthieth Formation, loc. 11170. 

Genus Hypserpa Miers 

Hypserpa? sp.

Text figure 16

Discussion. A fragmentary leaf has a marginal 

vein, apparently palmate venation (the base is 

missing), only one pair of conspicuous presumed

1 cm

FIGURE 16. Hypserpa? sp. USNM 43302, loc. 11168.

1 cm

FIGURE 17. Marginal venation of Limacia. A, L. 

stenophylla, paratype, USNM 43305, loc. 11159. B, L. 

oblonga, USGS Ref. Colln. 1803.

lateral primaries, craspedodrome marginal tertiary 

veins, and a linear shape. Although clearly a 

member of Menispermaceae, the generic reference is 

questionable because of the incompleteness of the 

specimen.

Specimen. USNM 43302.

Occurrence. Katalla Formation, Split Creek 

Sandstone Member, loc. 11168.

Genus Limacia 

Limacia stenophylla Wolfe, new species

Plate 6, figures 1-3; plate 7, figures 4, 7; text figure 17

Diagnosis. Leaves simple; venation palmate; 

shape ovate, linear; base narrowly rounded, apex 

acute; length 6-12 cm, width 2-4 cm; a single pair of 

conspicuous lateral primaries extending less than a 

third the distance to the apex; secondaries and 

lateral primaries camptodrome; inconspicuous lateral secondaries; 7-10 pairs of medial secondaries, 

departing at an angle of 40°-70°, straight to broadly 

curving, looping sharply near margin; numerous 

intersecondaries; intercostal tertiary veins branching, approximately perpendicular to secondary 

veins, widely spaced.

Discussion. The fossils show the folio wing points 

of resemblance to leaves of Menispermaceae: 

palmate venation, a strong marginal vein, and 

craspedodrome lateral secondary veins near the 

apex that join the marginal vein. Among extant 

Menispermaceae, the fossils most closely resemble 

Limacia. This resemblance is particularly strong in 

(1) several widely spaced medial secondary veins, as 

in L. velutina Miers and L. oblonga Miers; (2) 

craspedodrome lateral secondary veins near the 

apex, as in L. velutina and L, scandens Lour.; (3) 

camptodrome medial secondary veins, as in L. 

velutina and L. oblonga; (4) widely spaced and
SYSTEMATIC S 65

FIGURE 18. Marginal and intercostal venation of Paratinomiscium and Tinomiscium. A, P. conditionalis, 

holotype, USNM 39117, loc. 3879 (A, marginal; B, intercostal venation). C, T. petiolare, USGS Ref. Colln. 1808.

branching tertiary veins connecting the lateral 

primaries to the midrib, as in all extant species 

examined; (5) only one pair of lateral primaries, as in 

all extant species examined; and (6) a narrowly 

rounded base, as in L. oblonga. The major features of 

the fossil can, therefore, be found in Limacia, 

although no single species of Limacia combines all 

these features. L. stenophylla, in addition, has a 

more elongated shape than in the extant species 

examined.

Holotype. USNM 43303.

Paratypes. USNM 43304-43307.

Occurrence. Kushtaka Formation, Iocs. 11158, 

11159, 11166.

Genus Paratinomiscium Wolfe, new organ genus

Diagnosis. Leaves simple; venation palmate; 

base broadly rounded; five primary veins, first pair 

extending into apical half of lamina, camptodrome; 

lateral secondary veins camptodrome, forming 

loops; a few pairs of medial secondary veins, 

camptodrome, forming loops; marginal tertiary 

veins craspedodrome, joining conspicuous marginal 

vein; intercostal tertiary veins approximately perpendicular to secondary veins, branching or unbranching, closely spaced; quartary mesh irregularly polygonal; petiole long.

Type species. P. conditionalis (Hollick) Wolfe 

[Hampea].

Discussion. The fossils apparently represent a 

new genus of Menispermaceae. Among extant 

genera, the fossils are most similar to Tinomiscium 

in the broadly rounded base, lateral primary veins 

extending about two-thirds the distance to the apex, 

conspicuous marginal vein, and long petiole. In

1 cm B

FIGURE 19. Venation of Pycnarrhena. A, P. sp., USNM 

43308, loc. 11166. B, P. celebica, USGS Ref. Colln. 1806.

Tinomiscium, however, the intercostal tertiary veins 

are spaced about twice the distance apart as in 

Paratinomiscium and the marginal tertiary veins 

form a series of loops.

Paratinomiscium conditionalis (Hollick) Wolfe 

Plate 7, figure 5; text figure 18

Hampea conditionalis Hollick, 1936, U.S. Geol. Survey Prof. Paper 

182, p. 150, pi. 118, fig. 1.

Holotype. USNM 39117.

Occurrence. Kulthieth Formation, Iocs. 3879, 

11195.

Genus Pycnarrhena Miers 

Pycnarrhena sp.

Plate 6, figure 4; text figure 19

Discussion. The fossil leaf is palmately veined, 

although the lateral primaries extend a short 

distance above the top of the petiole parallel to the 

midrib. The first pair of lateral primaries is 

prominent, and extend about half the distance to the 

apex. The area formed by the midrib and the looping 

secondary veins are almost square. The tertiary 

venation has a strong tendency to extend in an 

abmedial direction and disappear, presumably 

thinning into fourth order venation, which is not 

preserved. Although the leaves of the two extant 

species of Pycnarrhena examined both have more 

trapezoidal areas formed by the secondary veins, the 

thinning tertiary venation, closely spaced tertiary 

veins within the marginal loops, and the presence of 

a series of tertiary loops; these characteristics 

indicate a close relationship to the fossil.

Specimen. USNM 43308.

Occurrence. Kushtaka Formation, loc. 11166.

Family MYRISTICACEAE 

Genus Knema Loure
66 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

B

1 cm

FIGURE 20. Intercostal venation of Knema. A, K. sp., USNM 

43309, loc. 3847. B, K. glauca, USGS Ref. Colln. 1827.

Knema sp. 

Plate 8, figures 3, 4; text figure 20.

Rhamnus marginatus auct. non Lesquereux. Hollick, 1936, U.S. 

Geol. Survey Prof, paper 182, p. 140, pi. 78, figs. 1, 2.

Discussion. The numerous secondary veins that 

loop close to the margin, unequally spaced intercostal tertiary veins, and fourth-order venation that 

is irregularly branching and typically set at oblique 

angles to the intercostal tertiary veins are features 

that ally these fossils to Knema. Although some 

extant species of the genus have thick veins, I have 

not seen any quite so thick as in the fossil leaves. 

Among extant species, the fossil most closely 

resembles K. glauca (Blanco) Merr. in shape and 

extensive secondary looping and K. korthalsii Warb. 

in the spacing of the intercostal tertiary veins.

Specimens. USNM 38915, 43309.

Occurrence. Kushtaka Formation, loc. 3847.

Genus Myristica Gronov

Myristica sp.

Plate 5, figure 6; plate 8, figure 1; text figure 21 

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

p. 155, pi. 93 fig. 6.

Magnolia oualis auct. non Lesquereux. Hollick, 1936 (part), U.S. 

Geol. Survey Prof. Paper 182, p. 114, pi. 63, fig. 1 only.

Discussion. The extensive and irregular secFIGURE 21. Marginal venation of Myristica. A, M. sp., USNM 

38667, loc. 3847. B, M. guaterrifolia, USGS Ref. Colln. 1833.

1 cm

FIGURE 22. Marginal venation of Cananga. A, C. sp., USNM 

38911, loc. 3879. B, C. odorata, USGS Ref. Colln. 1632a.

ondary and tertiary loops, closely spaced and 

undulatory intercostal tertiary veins, and branching 

fourth-order venation indicate that the fossil 

represents a species of Myristica. The secondary 

loops are more elongated perpendicular to the midrib 

than in any extant species examined.

Specimens. USNM 38667, 39019.

Occurrence. Kushtaka Formation, loc. 3847.

Family ANNONACEAE 

Genus Cananga (DeCandolle) Hooker f. and Thomas

Cananga sp. 

Plate 8, figure 6; text figure 22

Magnolia wormskioldi auct. non Heer. Hollick, 1936, U.S. Geol. 

Survey Prof. Paper 182, p. 115, pi. 63, fig. 3.

Discussion. The fossil appears to represent 

Annonaceae and in particular resembles Cananga 

odorata Hook. f. This resemblance is in the very 

broadly rounded base, curvature of the secondary 

veins, thick midrib but thin secondary veins, 

angular looping of the secondary veins and 

orientation of the apical side of the loop, slightly 

irregular looping of the marginal tertiary veins, 

orientation and spacing of the intercostal tertiary 

veins, irregularly branching quartary veins, and 

entire margin lacking a marginal vein.

Specimen. USNM 38911.

Occurrence. Kulthieth Formation, loc. 3879.

Genus Dasymaschalon (Hooker f. and Thomas) Dalla Torre

and Harms

Dasymaschalon? sp.

Plate 8, figure 5

Discussion. The combination of a broadly rounded cordate base, pinnate venation, and entire 

margin, is unusual. Some Annonaceae, however,
SYSTEMATICS 67

display such a combination of features. Broadly 

arching and thin secondary veins that extend a 

considerable distance towards the apex are found in 

Dasymaschalon trichophorum Merr., as is a broadly 

cordate base. Because the tertiary venation of the 

fossil is obscured by the coriaceous texture, the 

generic reference should not be considered certain.

Specimen. USNM 43310.

Occurrence. Kushtaka Formation, loc. 11158.

Family LAURACEAE

Foliage of Lauraceae is difficult to determine as to 

genus under the best of circumstances. Many species 

in the western Tertiary particularly the Paleogene have been assigned to extant genera simply 

on the basis that a particular fossil species most 

closely resembles a particular species; that is, the 

generic assignment has no morphologic basis in that 

the fossil species is not stated to display the 

characters diagnostic of the particular genus. In fact 

few extant genera of Lauraceae can be diagnosed on 

the features of the leaves particularly noncuticular 

features. This circumstance should not be surprising 

in view of the fact that Reid and Chandler (1933) 

were unable to clearly delimit the genera of 

Lauraceae despite having well-preserved fruits. An 

approach to lauraceous leaves that has morphologic 

validity is to use organ genera that represent 

morphologic types except in the few instances (for 

example, Sassafras) that assignments to extant 

genera can be confidently made.

One grouping that can be made of lauraceous 

leaves, as pointed out by Krausel and Weyland 

(1951), is of those leaves that have secondary veins 

that are approximately parallel and of approximately the same strength; the name Litseaphyllum 

is applicable to such leaves. This organ genus, in 

addition, should be emended so as to include leaves 

that have few or no freely ending veinlets; if freely 

ending veinlets are present, they are unbranching. 

An additional character of the genus is the presence 

of a continuous marginal vein. Litseaphyllum thus 

includes leaves ofLaurus, Machilus, Persea, Lindera 

(part), and Litsea; definitely excluded because of the 

presence of branching veinlets are Endiandra, 

Ocotea, Nectandra, Licaria, Phoebe, and Umbellularia. The various species from the west American 

Tertiary that have been attributed to Nectandra are 

all Litseaphyllum on the basis of ultimate venation.

A second grouping of lauraceous leaves are those 

that have a pseudo-palmate venation, that is, one 

pair of secondary veins typically departing a short 

distance above the top of the petiole are aparallel 

to, and typically thicker than, the other secondary

veins. Additionally, a continuous marginal vein is 

present and the areoles have few or no freely ending 

veinlets; if present, the veinlets are unbranching. 

Leaves of this type are referred to Cinnamomophyllum (Krausel and Weyland, 1951). Included in 

Cinnamomophyllum are leaves of Cinnamomum, 

Cryptocarya, Lindera (part), and Neolitsea; excluded 

because of the presence of branching veinlets are 

Ocotea and Sassafras. Note that Cinnamomophyllum only includes leaves of genera that are 

today old world in distribution. Significantly, none 

of the species from the west American Tertiary that 

have been attributed to Ocotea represent that genus; 

without exception they have the ultimate venation of 

Cinnamomophyllum.

Genus Cinnamomophyllum Krausel and Weyland

Cinnamomophyllum kushtakensis Wolfe, new species

Plate 23, figures 5, 7a

Diagnosis. Leaves simple; venation pinnate; 

shape ovate, linear; base acute; five or six pairs of 

conspicuous secondary veins, the lowest pair more 

pronounced, extending about half the distance to 

apex, aparallel to other secondary veins; other 

secondary veins departing at an angle of about 40°- 

60°, curving apically, looping; intercostal tertiary 

veins widely spaced, branching; areoles intruded by 

unbranching or typically no veinlets; margin entire, 

with marginal vein.

Discussion. The description of the ultimate 

venation is based on a specimen from locality 9731 in 

the Puget Group; this specimen in other respects 

matches well the Katalla specimens. Cinnamomophyllum kushtakensis can be separated from other 

described species of the genus primarily by the 

linear shape. C. kushtakensis may be closely related 

to C. latum.

Holotype. USNM 43311.

Paratypes. USNM 43312-43313 (loc. 9732, Puget 

Group).

Occurrence. Kushtaka Formation, Iocs. 9389, 

11165.

Cinnamomophyllum latum (MacGinitie) Wolfe,

new combination 

Plate 8, figure 2; plate 9, figures 3, 10

Neolitsea lata MacGinitie, 1941, Carnegie Inst. Washington

Pub. 534, p. 118, pi. 20, figs. 2, 3. 

Cinnamomum cinnamomeum auct. non Phyllites cinnamomeus

Rossmaessler. Rollick, 1936, U.S. Geol. Survey Prof. Paper

182, p. 115, pi. 64, fig. 2.

Discussion. MacGinitie's holotype has a distinct 

marginal vein and almost no freely ending veinlets 

and is thus transferred to Cinnamomophyllum.

Hypotypes. USNM 38914, 43314-43315.
68 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Occurrence. Kushtaka Formation, Iocs. 3847, 

11157, 11160, 11166.

Genus Litseaphyllum Wolfe, new name 

Laurophyllum Goeppert, 1853, Neues Jahrb., p. 434.

Discussion. The name "Laurophyllum" is an 

orthographic variant of the anacard Laurophyllus 

Thunb., and this extant genus has clear priority. A 

new name for the organ genus is thus needed.

Litseaphyllum carbonensis Wolfe, new species 

Plate 28, figures 6, 9

Diagnosis. Leaves simple; venation pinnate; 

shape oval; base acute to narrowly rounded, apex 

acute; three or four pairs of conspicuous secondary 

veins, departing at low angles, curving sharply 

apically, looping; second pair of secondary veins 

extending about three-fourths distance to apex, third 

pair extending to apex; intercostal tertiaries oriented 

approximately perpendicular to midrib, closely and 

uniformly spaced; areoles intruded by unbranching 

or typically no veinlets; margin entire.

Discussion. This species closely resembles 

Laurophyllum presamarensis in general aspect, but 

in L. carbonensis the secondary veins curve more 

sharply and extend farther towards the apex.

Holotype. USNM 43317.

Paratype. USNM 43316.

Occurrence. Kushtaka Formation, loc. 11169.

Litseaphyllum katallaensis Wolfe, new species 

Plate 29, figures 1,3

Diagnosis. Leaves simple, venation pinnate; 

shape oval to ovate; base acute, apex acute; seven to 

nine pairs of secondary veins, curving apically, 

extending almost to margin and looping weakly; 

weak basal secondary veins, intercostal tertiary 

veins spaced uniformly about 0.5 cm apart, 

branching, approximately at right angles to secondary veins; areoles with unbranching veinlets; 

margin entire.

Discussion. This species resembles in shape and 

secondary venation "Laurophyllum" ramineruum 

Potb., but that species has areoles intruded by 

branching veinlets (Potbury, 1935, p. 67) and cannot 

represent Litseaphyllum as this genus is conceptualized in this report.

Holotype. USNM 43319.

Paratype. USNM 43318.

Occurrence. Katalla Formation, Split Creek 

beds, loc. 11168; Kulthieth Formation, loc. 9551.

Litseaphyllum praelingue (Sanborn) Wolfe, new combination 

Plate 22, figure 6

Persea praelingue Sanborn, 1935, Carnegie Inst. Washington

Pub. 465, p. 20, pi. 6, fig. 1. 

Magnolia californica auct. non Lesquereux. Sanborn, 1935 (in

part), Carnegie Inst. Washington Pub. 465, p. 17, pi. 2,

figs. 3-5.

Discussion. Sanborn's holotype of Persea praelingue appears to be an aberrant leaf of the species 

she also called Magnolia californica. In Litseaphyllum praelingue, the intercostal tertiary veins 

are oblique to the secondary veins, the intramarginal 

loops have a pronounced trapezoidal shape, the 

midrib is thick, and the petiole is thick.

Hypotype.-USNM 43324.

Occurrence. Kushtaka Formation, loc. 11165.

Litseaphyllum presamarensis (Sanborn) Wolfe, 

new combination

Plate 9, figure 2; plate 22, figures 1, 2; plate 28, figures 8b, lOa, b 

Cryptocarya presamarensis Sanborn, 1935, Carnegie Inst. 

Washington Pub. 465, p. 19, pi. 5, figs. 4, 6.

Discussion. Although these leaves could presumably represent Cryptocarya, they cannot be 

satisfactorily distinguished from leaves of several 

other genera of Lauraceae. Ultimate venation of 

leaves from locality 9731 in the Puget Group provide 

the basis for referring this species to Litseaphyllum.

Hypotypes.-USNM 43325-43330.

Occurrence. Kushtaka Formation, Iocs. 3842, 

3846, 11157, 11165, 11169.

Litseaphyllum presanguinea (Chaney and Sanborn) Wolfe, 

new combination

Plate 29, figure 9

Nectandra presanguinea Chaney and Sanborn, 1933, Carnegie 

Inst. Washington Pub. 439, p. 73, pi. 18, figs. 1, 2, 5, 7.

Discussion. The ultimate venation of the type of 

this species indicates that the specimen cannot 

represent Nectandra but is related to various Asian 

laurels (see p. 67). The undulatory secondary veins 

that loop well within the margin, numerous 

intersecondary veins, and linear shape characterize 

this species.

Hypotype. USNM 43331.

Occurrence. Kushtaka Formation, loc. 9891.

Litseaphyllum schorni Wolfe, new species 

Plate 25, figures 1, 2, 6

Diagnosis. Leaves simple; venation pinnate; 

shape oval to obovate; base cuneate, apex acuminate; weak pair of basal secondary veins that extend 

a short distance along margin; 11-13 pairs of 

secondary veins, uniformly spaced, broadly curving 

towards margin, curving sharply and looping near 

margin; short series of small, angular loops 

extending apically to about where next most apical 

secondary loops; intercostal tertiary veins numer-
SYSTEMATICS 69

ous, straight or branching, perpendicular to secondaries; margin entire; petiole thick and long.

Discussion. Well-preserved leaves from the middle Ravenian of the Puget Group appear to have the 

diagnostic features of Litseaphyllum schorni; the 

Puget leaves have small areoles, most of which lack 

freely ending veinlets. The fossils closely resemble 

Cryptocarya infectoria (Bl.) Miq. of the Philippine 

Islands. The resemblance is particularly strong in 

the secondary veins, which are strongly, smoothly, 

and gradually curving and sharply looping near the 

margin, and in the spacing and strength of the 

intercostal tertiary veins. The fossils, however, have 

more numerous and closely spaced secondary veins 

and a longer petiole than does C. infectoria.

This species is named for H. E. Schorn in 

recognition of his assistance in discussing the 

vegetational and floristic aspects of this report.

Holotype. USNM 43333.

Paratypes. USNM 43332, 43334.

Occurrence. Kushtaka Formation, Iocs. 9389, 

11165.

Litseaphyllum similis (Knowlton) Wolfe, new combination

Plate 9, figures 1, 4; plate 23, figure 2; plate 29, figure 2 

Laurus similis Knowlton, 1900, U.S. Geol. Survey 20th Ann. Kept.,

pt. 3, p. 48, pi. 5, figs. 1-4. 

Persea pseudocarolinensis auct. non Lesquereux. Sanborn, 1935,

Carnegie Inst. Washington Pub. 465, p. 20, pi. 5, fig. 3.

Discussion. Laurus similis was synonymized to 

Persea pseudocarolinensis by Sanborn (1935). In 

Persea, however, the veins that depart from the 

midrib basal to the first pair of conspicuous 

secondaries are stronger than the tertiary veins and 

typically are perpendicular to the midrib. Laurus 

similis does not represent a Persea-Machilus type of 

leaf, but appears to be related to laurels such as 

Laurus, Lindera, and Litsea.

Hypotypes. USNM 43320-43323.

Occurrence. Kushtaka Formation, Iocs. 11157, 

11165, 11169.

Litseaphyllum spp.

The species of Litseaphyllum discussed on the 

following pages do not, in most instances, have the 

ultimate venation preserved. Most of the species, 

although having the weak basal secondary veins 

characteristic of Lauraceae, are therefore only 

tentatively referred to Litseaphyllum; I am thus 

using Litseaphyllum in a broader sense than for the 

named species previously discussed.

Litseaphyllum sp.

Plate 9, figure 7

Discussion. The specimen has an entire margin 

and a highly coriaceous texture. Weak, basal

secondary veins are present. The few secondary 

veins give off strong branches near the margin, 

unlike the secondaries of Litseaphyllum presamarensis and L. similis. The quartary mesh is 

formed of almost quadrangular areoles, and thus the 

fossil resembles leaves of some species of Lindera 

and Litsea.

Specimen.-USNM 43335.

Occurrence. Kushtaka Formation, loc. 11159.

Litseaphyllum sp. 

Plate 29, figure 5

Discussion. These small leaves have a coriaceous 

texture. The secondary veins in the basal part of the 

lamina depart from the midrib almost at a right 

angle; this feature is also found in leaves ofMachilus 

and Persea.

Specimen.USNM 43336.

Occurrence. Kulthieth Formation, loc. 11170; 

Katalla Formation, Split Creek beds, loc. 11168.

Litseaphyllum sp. 

Plate 22, figure 5

Discussion. The specimen has weak secondary 

veins departing from the midrib at an angle of about 

90° in the basal part of the lamina; as the preceding 

species, this species is apparently allied to Machilus 

or Persea (considered by some neobotanists to be the 

same genus). The more numerous secondary veins 

that do not curve as sharply indicate that the 

specimen from the Kushtaka does not represent the 

same species as the preceding form.

Specimen. USNM 43337.

Occurrence. Kushtaka Formation, loc. 11165.

Litseaphyllum sp.

Plate 22, figure 4

Discussion. This leaf has a pronounced asymmetric, ovate shape and broadly arching secondary 

veins; these features distinguish the species from 

Litseaphyllum praelingue. The secondary veins give 

off strong branches that loop with the next most 

basal secondary vein; the loops thus formed are 

angular. The intercostal tertiary venation is obscure, 

but the tertiaries appear to be evenly and widely 

spaced and approximately perpendicular to the 

secondary veins.

Specimen. USNM 43338.

Occurrence. Kushtaka Formation, loc. 11165.

Litseaphyllum sp. 

Plate 28, figure 8a

Discussion. This leaf is distinguished from the 

other laurels discussed in this report by the linear,
70 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

ovate shape and the numerous secondary veins. 

Specimen. USNM 43339. 

Occurrence. Kushtaka Formation, loc. 11169.

Litseaphyllum sp. 

Plate 29, figure 8

Discussion. This linear leaf has fewer and more 

widely spaced secondary veins that the preceding 

form. The shape, in addition, is oval rather than 

ovate.

Specimen. USNM 43340.

Occurrence. Kushtaka Formation, loc. 9891.

Order ROSALES

Family HAMAMELIDACEAE

Genus Corylopsis Siebold and Zuccarrini

Corylopsis sp.

Plate 26, figure 6

Discussion. The asymmetric shape, palmateappearing venation, and nonentire margin are 

characters indicative of the subfamily Hamamelioideae. The specimen has numerous, sharp almost 

spinose teeth, smoothly curving secondary venation, and a cordate base; a species of Corylopsis is, 

therefore, represented. The only other almost spinose 

leaves in the subfamily belong to Fc^tunearia and 

Sinowilsonia; in both these genera, however, the 

secondary venation is undulatory and conspicuously 

forking.

Koch (1963) has described leaves from Greenland 

as Corylopsiphyllum; I can see no morphologic basis 

for referring these leaves to any genus other than 

Corylopsis. Various species of Hamamelis and 

Hamamelites have been described from Paleocene 

and Eocene assemblages throughout the northern 

hemisphere. Most of these leaves have the diagnostic 

features of the subfamily, although some of the 

species do not appear to be clearly referrable to any 

extant genus. Together with the occurrence of 

Corylopsis in the tropical London Clay assemblage 

(Chandler, 1964), Fothergilla in the tropical middle 

Eocene assemblages of the Puget Group (Wolfe, 

1968), and of Fortunearia in the subtropical early 

basic breccia assemblage (see p. 89), the Paleogene 

occurrences of the subfamily indicate an adaptation 

in the early Tertiary to the tropical vegetation types. 

The fact that some extant members of the subfamily 

are winter-flowering a peculiar adaptation for a 

temperate, deciduous plant could be explained in 

terms of a former wide distribution in tropical 

vegetation types, in which the time of flowering 

would not have been significant, the winterflowering may thus be an anachronism.

Specimen. USNM 43341A, B.

Occurrence. Kushtaka Formation, loc. 11167.

Genus Liquidambar Linnaeus 

Liquidambar? sp.

Plate 24, figure 1

Discussion. The margin and venation of this 

specimen is not well preserved, but the deeply cut 

lobes and apparently finely serrate margin combined with the camptodrome secondary veins, 

indicate that the specimen is Liquidambar. In 

species of Acer that have nonentire margins, the 

secondary veins are typically craspedodrome.

Specimen. USNM 43342.

Occurrence. Kushtaka Formation, loc. 11165.

Family EUCOMMIACEAE 

Genus Eucommia Oliver

Eucommia sp. 

Plate 24, figure 7

Discussion. The specimen has widely and irregularly spaced secondary veins, the apical ones 

curving more sharply than the more basal ones. The 

secondaries are camptodrome; two sets of inframarginal loops are present, and thus the quartary 

veins are craspedodrome. The teeth are numerous 

and are separated by either arcuate or angular 

sinuses. The intracostal tertiary veins are obscure, 

but they appear to be widely spaced and slightly 

oblique to the secondaries. In the features preserved, 

the specimen is assignable to Eucommia.

Brown (1962) has assigned some Paleocene leaves 

to Eucommia. These leaves, however, have both 

craspedodrome and camptodrome secondary veins, 

craspedodrome tertiary veins, and apical secondary 

veins that are typically approximately parallel to the 

more basal secondaries; these leaves do not represent 

Eucommia. The seeds from the Claiborne assemblage that Brown (1940) referred to Eucommia also 

do not appear to represent that genus. In Eucommia, 

a conspicuous vascular strand extends from the base 

of the samara alongside the seed, splits at the apical 

cleft and extends to the apex of the samara along 

either side of the cleft. It is clear from Berry's 

illustration (1930a, pi. 44, figs. 15,16) that such a vein 

and an apical cleft are not present. The other 

Claiborne specimen assigned to Eucommia by 

Brown (1940) similarly is not Eucommia', Berry's 

illustrations (1930a, pi. 33, figs. 5,6) indicate that the 

seed was basal and an expanded samara extended 

apically.

Eucommia montana R. W. Br. and E. browni Beck, 

both appear to be valid representatives of the genus. 

The three known North American representatives of 

Eucommia indicate that the genus was present in
SYSTEMATICS 71

both lowland subtropical forest in the late Eocene 

and upland temperate forest in the early to middle 

Oligocene. The genus, however, apparently did not 

survive in North America to become a member of the 

Neogene Mixed Mesophytic forest.

Specimen. USNM 43343.

Occurrence. Kushtaka Formation, loc. 11165.

Family PLATANACEAE

Genus Platanus Linnaeus

Platanus comstocki (Sanborn) Wolfe, new combination

Plate 29, figure 7 

Mallotus comstocki Sanborn, 1935, Carnegie Inst. Washington

Pub. 465, p. 23, pi. 6, fig. 9. 

Platanus aceroides auct. non Goeppert. Sanborn, 1935, Carnegie

Inst. Washington Pub. 465, p. 21, pi. 6, fig. 10.

Discussion. Mallotus, as do many Euphorbiaceae, has leaves in which the quartary veins are 

perpendicular to the tertiary veins; this situation is 

clearly not present in Sanborn's Mallotus comstocki. 

Her interpretation of the leaf as being subpeltate is 

due to the fusion of the acropetiolar stipule to the 

lamina, which is typical of Platanus. P. aceroides is a 

strongly toothed and "lobed" species, unlike P. 

comstocki.

Hypotype. USNM 43344.

Occurrence. Kushtaka Formation, loc. 9891; 

Kulthieth Formation, loc. 11189.

Family ROSACEAE

The geologic history of Rosaceae is largely 

confined to the latest Eocene, Oligocene, and 

Neogene. In contrast to the diversity of the family in 

the warm temperate assemblages of the Neogene, the 

family is apparently lacking in warm temperate 

Paleocene assemblages. Brown (1962) has attributed 

some leaves from the Rocky Mountain Paleocene to 

Prunus, but none of these leaves has the acropetiolar 

glands or glandular teeth characteristic of Prunus. 

The fructifications assigned by Brown (1962) to 

Prunus should be reinvestigated; they have obvious 

similarities to the fruits of Icacinaceae. Chandler 

(1964) recorded seeds of Rubus from the early Eocene, 

and MacGinitie (1941) has recorded Vauquelinia in 

the early Eocene. The family does not appear to have 

been well represented in subtropical or Paratropical 

Rain forest during the Paleogene a situation that 

still obtains today in these vegetational types. In the 

cool period of the latest Eocene, Rosaceae appear for 

the first time in diversity. Genera that appear to be 

validly recorded from beds of this age are: 

Amelanchier, Crataegus, Chamaebatiaria, Potentilla, Prunus, Pyrus, Rubus, and Sorbus. Note that all 

four subfamilies were represented. Most of these 

genera occur in the more temperate vegetation in

lowland Alaska and upland Pacific Northwest, but 

one genus Rubus is known in the John Day Gulch 

assemblage, which is warm temperate to subtropical. 

The upland floras of the early to middle Oligocene 

(Ruby, Florissant) display a considerable diversity 

of Rosaceae. In addition to the genera that first 

appeared in the latest Eocene, several other genera, 

such as Cercocarpus, are represented; more significantly, the specific diversity within each genus 

appears to have been greater during the early to 

middle Oligocene than during the latest Eocene. 

From the later Oligocene on, the family appears to 

have increased in both generic and specific diversity. 

The early Miocene Collawash assemblage from 

Oregon representing the Mixed Mesophytic forest contains numerous Rosaceae: Amelanchier, 

Cercocarpus, Crataegus (three species), Holodiscus, 

Lyonothamnus, Peraphyllum (or a closely related 

genus), Prunus (three species), Pyrus, Rosa, and 

Rubus. By the Miocene, almost all woody genera of 

Rosaceae now native to western North America had 

appeared, and several of the lineages represented 

appear to be the same as those that have survived to 

the present time.

Genus Prunus Linnaeus 

Prunus axelrodi Wolfe, new name

Plate 26, figure 9; text figure 23 

Euonymus nevadensis Axelrod, 1966b, California Univ. Pubs.

Geol. Sci., v. 59, p. 75, pi. 18, figs. 7, 8.

Prunus chaneyi auct. non Condit. Axelrod, 1966b (part), California Univ. Pubs. Geol. Sci., v. 59, p. 72, pi. 17, fig. 1 only. 

Prunus harneyensis auct. non Axelrod 1944. Axelrod, 1966b, California Univ. Pubs. Geol. Sci., v. 59, p. 72, pi. 17, figs. 4-7.

Discussion. The holotype of Euonymus nevadensis has large glandular teeth just apical from the 

top of the petiole and is thus clearly Prunus. The 

flowing intercostal venation indicates a relationship 

to P. serotina Ehren. The Alaskan fossil tentatively 

referred to P. axelrodi is fragmentary, but it has 

about the same spacing and type of teeth and the 

same tertiary venation as does P. axelrodi; in 

addition, the spacing of the secondary veins, their 

angle of departure, and their curvature in the 

Alaskan fossil is the same as in P. axelrodi.

The epithet "nevadensis" has been previously 

applied to a species of Prunus, and thus a new name 

is required. This species is named for the original 

describer, D. I. Axelrod.

Hypotype. USNM 43345A, B.

Occurrence. Kushtaka Formation, loc. 11167.

Genus Sorbus Linnaeus

Sorbus carbonensis Wolfe, new species

Plate 27, figures 3, 4; text figure 24

Diagnosis. Leaf compound; venation of leaflet
72 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

A B C

1 cm

FIGURE 23. Marginal venation of Prunus. A, P. axelrodi, hypotype, USNM 43345B, loc. 11167. B, P. 

axelrodi, holotype, UCMP 20111A, Copper Basin flora. C, P. serotina, USGS Ref. Colln. 1202.

pinnate; shape linear, oval; base asymmetric; at 

least 13 pairs of secondary veins, straight to slightly 

curving, craspedodrome; marginal tertiary veins 

craspedodrome; intercostal tertiary veins oriented 

perpendicular to secondaries, branching; quartary 

venation forming irregularly polygonal areoles 

intruded by profusely branching veinlets; teeth very 

sharp, about two per secondary vein.

Discussion. The single leaflet has the finely and 

compoundly serrate margin of leaflets ofSorbus. The 

most closely related species to S. carbonensis is an 

undescribed species from the type section of the 

Seldovian Stage; this undescribed species typically 

has three or four teeth per secondary vein, but it has 

the same intercostal venation as does S. carbonensis.

Holotype. USNM 43346A, B.

Occurrence. Kushtaka Formation, loc. 11167.

Family LEGUMINOSAE 

Genus Caesalpinites Saporta

Caesalpinites sp. 

Plate 9, figure 6; text figure 25

Discussion. The single leaflet has features of a 

number of genera of legumes. In my opinion, 

however, most isolated leaflets of legumes cannot be 

satisfactorily assigned to a genus because of the 

overlap of characters between genera.

1 cm

FIGURE 24. Marginal venation ofSorbus carbonensis. Holotype, 

USNM 43346B, loc. 11167.

Specimen. USNM 43347.

Occurrence. Kushtaka Formation, loc. 3847.

Caesalpinites sp. 

Plate 25,. figure 5

Discussion. The leaflet has secondary veins that 

extend directly to the margin; as with the preceding 

species, assignment to other than an organ genus is 

not warranted.

Specimen. USNM 43348.

Occurrence. Kushtaka Formation, loc. 11165.

Order OERANIALES 

Family MELIACEAE 

Genus Toona Roemer

Toona sp. 

Plate 19, figure 4

Discussion. A single leaflet resembles the leaflets 

of some species of Toona. The fossil is petiolulate, 

linear. Camptodrome secondary veins curve close to 

the margin. Widely spaced, branching tertiary veins 

are perpendicular to the secondary veins. A revolute 

margin has two irregularly spaced notches. These 

features are also found in some leaflets of Toona, 

particularly T. sureni (Blume) Merr.; this species

FIGURE 25. Venation of Caesalpinites sp. USNM 43347, 

loc. 3847.
SYSTEMATICS 73

B

FIGURE 96. Marginal and intercostal venation of Luvunga. A, L. spatiosa, hypotype, USNM 43352, loc. 3847. B, D, L. latifolia,

USGS Bef. Colln. 1696. C, L. spatiosa, holotype, USNM 38920, loc. 3847.

inhabits the Paratropical Rain forest of southern 

China.

Specimen. USNM 222842.

Occurrence, Kulthieth Formation, loc. 11195.

Family RUTACEAE

Genus Euodia J. R. and G. Forster

Euodia alaskana Wolfe, new species

Plate 9, figures 5, 8, 9

Diagnosis. Leaf compound; venation of leaflets 

pinnate; shape oval to ovate; base cuneate, slightly 

asymmetric; apex narrowly rounded; 12-15 pairs of 

secondary veins, extending at high angles straight 

or undulatory, thick, camptodrome, forking conspicuously, some loops extending to margin; intercostal 

tertiary veins uniformly spaced, forking; intersecondary veins absent or no more than one per secondary 

vein; margin entire, undulatory; texture coriaceous.

Discussion. The fossil leaflets have conspicuously forking secondary veins, highly irregular 

"flowing" intercostal venation (which tends to 

extend abmedially, recurve, and extend admedially), 

and a highly undulate, almost crenulate margin; 

these characters are those of leaflets of Euodia. 

Among extant species, E. crassifolia Merr. from the 

Philippine Islands has leaflets most similar to the 

fossil. This similarity is particularly displayed in the 

marginal venation; some of the secondary veins fork

and extend straight to the margin. E. alaskana has 

but one or no intersecondary vein per intercostal 

area, whereas E. crassifolia has two or more, one of 

which typically extends abmedially almost to the 

margin.

Holotype. USNM 43349.

Paratypes. USNM 43350, 43351.

Occurrence. Kushtaka Formation, loc. 11158.

Genus Luvunga 

Luvunga spatiosa (Hollick) Wolfe, new combination

Plate 10, figures 1, 4; text figure 26

Persea spatiosa Holick, 1936, U.S. Geol. Survey Prof. Paper 182, 

p. 116, pi. 65, fig. 1.

Discussion. The pronounced asymmetry of Hoilick's specimen and of the additional specimen 

referred here to this species indicate that they were 

leaflets, not leaves as in Lauraceae. The widely 

spaced and irregularly branching intercostal tertiary veins are also not lauraceous. The leaflets of 

Rutaceae, particularly those of Luvunga, are very 

similar to the fossils. These similarities are: (1) an 

entire margin, (2) broadly curving secondary veins, 

(3) extensive and large loops, some of which are 

elongated parallel to the midrib, (4) numerous 

intersecondary veins that parallel the secondary 

veins and that extend at least half way to the 

margin, (5) widely spaced and irregularly branching 

intercostal tertiary veins some of which join to form
74 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

FIGURE 27. Marginal and intercostal venation of Melanorrhoea. A, M. alaskana, holotype, USNM 39020, loc. 3847. B, M. macrocarpa, USGS Ref. Colln. 1769. C, M. alaskana, 

hypotype, USNM 43353, loc. 11158.

a chevron pointing abmedially, and (6) an abruptly 

acute apex forming a drip-tip. L. spatiosa particularly resembles L. latifolia Tan. and L. crassifolia 

Tan., both of which are scandent vines of Melanesia.

Holotype.-USNM 38920.

Hypotype.-USNM 43352.

Occurrence. Kushtaka Formation, Iocs. 3847, 

11166.

Order SAPINDALES 

Family ANACARDIACEAE 

Genus Melanorrhoea Wallich

Melanorrhoea alaskana (Hollick) Wolfe, new combination 

Plate 10, figures 2, 3, 5; text figure 27

Semecarpus alaskana Hollick, 1936, U.S. Geol. Survey Prof. Paper

182, p. 130, pi. 93, fig. 7. 

Dryophyllum stanleyanum auct. non Dawson. Hollick, 1936

(part), U.S. Geol. Survey Prof. Paper 182, p. 103, pi. 56, fig. 4

only.

Discussion. Hollick's assignment of this species 

to Anacardiaceae is considered valid. The significant characteristics of this species are the forking 

and change in orientation of the intercostal tertiary 

veins midway between the secondaries and the 

conspicuous thinning of the secondary veins near 

the margin. The fossils have about the same shape as 

the leaflets of some species of Toxicodendron, but in 

that genus the intercostal tertiaries thin to become 

fourth order veins. In Semecarpus the leaflets have a 

pronounced obovate shape, the tertiary veins do not 

change direction, and the intramarginal loops are 

strong. The fossils most closely resemble the leaflets 

of Melanorrhoea, particularly in regard to shape of 

the intercostal area, pattern and size of the third 

order mesh, and the presence of a series of loops 

formed by the marginal tertiary veins.

Holotype.-USNM 39020.

Hypotypes. USNM 38885, 43353.

Occurrence. Kushtaka Formation, Iocs. 3846, 

3847, 11158; Kulthieth Formation, loc. 11170.

Genus Rhus Linnaeus

Rhus cf. R. mixta (Lesq.) MacGinitie

Plate 19, figure 3

Discussion. The single specimen is fragmentary, 

but it displays the diagnostic features of leaflets of 

Rhus, that is, it has broad teeth that have elongated 

basal flanks, craspedodrome secondary veins that 

may or may not fork, numerous intersecondary veins 

that may or may not extend to the sinuses, and 

intercostal tertiary veins that are oblique to the 

secondary veins and that thin conspicuously 

midway between secondary veins. MacGinitie (1941) 

has illustrated many leaflets from the early Eocene 

of California that are similar to the Alaskan fossil, 

but its fragmentary nature does not allow specific 

assignment.

Specimen. USNM 222840.

Occurrence. Kulthieth Formation, loc. 11195.

Genus and species indetermined 

Plate 25, figure 4

Discussion. The specimen has secondary veins 

that thin conspicuously as they approach the 

margin. Although this feature indicates an assignment to Anacardiaceae, the preservation of the 

venation below the level of secondaries is too poor for 

generic assignment.

Specimen. USNM 43354.

Occurrence. Kushtaka Formation, loc. 11165.

Family AQUIFOLIACEAE

Genus Ilex Linnaeus 

Ilex carbonensis Wolfe, new species

Plate 26, figure 12; text figure 28
SYSTEMATICS 75

1 cm

FIGURE 28. Marginal venation of Ilex carbonensis. Holotype, 

USNM 43355M, loc. 11167.

Diagnosis. Leaf apparently pinnate (base missing); venation pinnate; shape oval; at least six pairs 

of secondary veins, widely spaced, curving broadly, 

forking to form angular loops; marginal tertiary 

veins camptodrome; marginal quartary veins craspedodrome, entering along apical side of teeth; 

intercostal tertiary veins few, widely and irregularly 

spaced, irregularly branching; fourth order mesh 

irregularly polygonal, forming areoles intruded by 

profusely branching veinlets; margin with three to 

four teeth per secondary vein; teeth small, separated 

by arcuate sinuses; margin thickened.

Discussion. The arcuate sinuses, the irregular 

intercostal tertiary pattern, and the profusely 

branching veinlets indicate an assignment to Ilex. 

In size of areoles, I. carbonensis is most similar to I. 

geniculata Maxim, of Japan; in marginal venation 

and size of the teeth, the fossil is more similar to I. 

micrococca Maxim, from the temperate upland 

region of southern China.

Holotype. USNM 43355A, B.

Occurrence. Kushtaka Formation, loc. 11167.

Family CELASTRACEAE

Genus Celastrus Linnaeus

Celastrus comparabilis Hollick

Plate 11, figures 1, 4-6; plate 18, figure 4

Celastrus comparabilis Hollick, 1936 (part), U.S. Geol. Survey

Prof. Paper 182, p. 132, pi. 75, fig. la only. 

Populus balsamoides auct. non Goeppert. Hollick, 1936 (part), U.S.

Geol. Survey Prof. Paper 182, p. 65, pi. 23, fig. 4 only. 

Discussion. Rollick's (1936) determination of 

this species as a member of Celastrus is valid. The

characters of the fossils that indicate such a 

relationship are: a uniformly, closely serrate margin; 

several widely spaced, camptodrome secondary 

veins; two series of loops near the margin; intercostal 

tertiary veins that bifurcate and that are typically 

almost perpendicular in orientation to the midrib; 

branches from the tertiary veins that tend to thin 

and form a highly irregular pattern of fourth order 

venation; some tertiary veins that depart from the 

midrib and fork more than once, each fork joining 

the next most basal secondary vein. These characteristics also indicate a close relationship to extant 

members of the subgenus Celastrus, which is 

distributed in Madagascar, southeast Asia, Melanesia, and eastern United States; the subgenus, 

however, has its current center in the tropical and 

subtropical region of southeastern Asia (Hou, 1955, 

p. 223). The fossils appear to be most closely related 

to C. monospermoides Loes., which is widely 

distributed from New Guinea to the Philippine 

Islands and Malaya.

Lectotype. USNM 38956.

Hypotypes. USNM 38744, 43356, 43357, 245728.

Occurrence. Kushtaka Formation, Iocs. 3846, 

3847, 11158, 11159, 11166; Kulthieth Formations 

Iocs. 3879, 11170.

Family ICACINACEAE

Genus Goweria Wolfe, emended

Goweria Wolfe, 1968, U.S. Geol. Survey Prof. Paper 571, p. 20.

Discussion. Leaves of this organ genus were 

stated to have a marginal vein and were placed in 

Menispermaceae. Reexamination of the type species, 

Goweria dilleri, from the Puget Group, indicates that 

I had misinterpreted a thickened margin as a vein. 

One specimen from the early Ravenian of the Puget 

clearly displays the small marginal protuberances 

typical of Phytocrene. Goweria, however, differs 

from Phytocrene in a few important characteristics: 

Goweria has only five primary veins, and the fourth 

order mesh is composed of polygons elongated 

parallel to the tertiary veins. In Phytocrene, there are 

seven primary veins and the fourth order mesh is 

approximately quadrangular. Four species of Goweria are now known: Goweria linearis, an undescribed 

species related to G. alaskana from the early 

Fultonian of the Puget Group, G. alaskana, and G. 

dilleri. Goweria may represent the same genus as 

Paleophytocrene; both organ genera are represented 

in a very small assemblage from the Kulthieth 

formation (loc. 3879).

Goweria alaskana Wolfe, new species

Plate 13, figure 3; text figure 29 

Diagnosis. Leaves simple, pinnately veined; five
76 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

1 cm

FIGURE 29. Marginal and intercostal venation of Goweria alaskana. A, B, Holotype,

USNM 43358A, loc. 11158.

primary veins; first lateral pair of primaries 

extending apparently about half the distance to the 

apex, subparallel to medial secondary veins, 

camptodrome, apical sides of loops approximately 

perpendicular to midrib; second pair of lateral 

primaries weak; at least two pairs of medial 

secondary veins; intercostal tertiary veins closely 

spaced, extending perpendicular to midrib or 

slightly apically, forking or unforking, about half 

the thickness of the secondary veins; marginal 

tertiary veins camptodrome or craspedodrome and 

ending in irregularly spaced marginal bumps.

Discussion. The apparently few pairs of medial 

secondary veins and the extension of the first lateral 

primaries into the apical half of the lamina 

distinguish this species from other described species 

of Goweria. An undescribed species from the early 

Fultonian of the Puget Group closely resembles G. 

alaskana, but the Fultonian species is considerably 

more rotund and the medial secondaries are more 

closely spaced than in G. alaskana.

Holotype. USNM 43358A.

Occurrence. Kushtaka Formation, loc. 11158.

Goweria dilleri (Knowlton) Wolfe 

Plate 11, figure 3

Goweria dilleri (Knowlton) Wolfe, 1968, U.S. Geol. Survey Prof.

Paper 571, p. 20, pi. 7, figs. 1, 2; text fig. 13. 

Benzoin dilleri Knowlton, 1900, U.S. Geol Survey 20th Ann. Kept.,

pt. 3, p, 46, pi. 4, fig. 3. 

Juglans thermalis auct. non Lesquereux. Hollick, 1936, U.S. Geol.

Survey Prof. Paper 182, p. 79, pi. 60, fig. la.

Discussion. The single fragmentary specimen 

was probably palmately veined; this is inferred from 

the strong difference in orientation between the two 

lowest major veins on the left side of the specimen. 

The specimen has the marginal protuberances,

broad shape, and slightly sinuous intercostal 

venation of G. dilleri.

Hypotype. USNM 38901.

Occurrence. Kushtaka formation, loc. 3847.

Genus Paleophytocrene Reid and Chandler

Paleophytocrene elytraeformis (Hollick) Wolfe, 

new combination

Plate 13, figure 2

Carpolithes elytraeformis Hollick, 1936, U.S. Geol. Survey Prof. 

Paper 182, p. 171, pi. 120, figs. 3, 4.

Discussion. The specimen illustrated by Hollick 

does not have the pits arranged as precisely in rows 

as Hollick or his illustrator show. The pits actually 

are in irregular rows. The specimen was broken 

before fossilization, but it is an almost complete 

impression. The pits are circular or slightly oval in 

cross section; together with their somewhat irregular 

arrangement, these kinds of pits indicate that the 

specimen represents Paleophytocrene. P. elytraeformis resembles some specimens of P. foveolata 

Reid and Chandl., but the Alaskan species has about 

sixteen pits lengthwise and the British species 

typically has only 12. R. A. Scott of the U.S. 

Geological Survey first suggested to me that 

Hollick's species represented Paleophytocrene.

Holotype. USNM 39127A, B.

Occurrence. Kulthieth Formation, loc. 3879.

Genus Phytocrene Wallich

Phytocrene acutissima Wolfe, new species

Plate 12, figures 1-6; plate 13, figure 1; text figure 30

Diagnosis. Leaves simple; palmately veined; 

shape ovate; base broadly rounded to cordate; apex 

pronouncedly acuminate, forming a drip-tip; seven 

primary veins, the most basal pair weak; lateral and 

medial secondary veins camptodrome, widely spaced;
SYSTEMATICS 77

1 mm.

FIGURE 30. Intercostal venation of Phytocrene acutissima, Holotype, USNM 43362B, loc. 11158.

intercostal tertiary veins sinuous; fourth order mesh 

quadrangular; marginal tertiary veins camptodrome or craspedodrome and ending in small, 

irregularly spaced marginal protuberances.

Discussion. The extremely elongated apex distinguishes this species from other species of 

Phytocrene that have been examined. Other species 

have, in addition, a consistently cordate base. The 

characters of the fossils that indicate that they are 

Phytocrene are, except for the base, the same as 

listed under P. sordida.

Holotype. USNM 43362A, B (figs. 4 and 6 on pi. 

13 are counterparts),

Paratypes. USNM 43359-43361, 43363, 43364.

Occurrence. Kushtaka Formation, Iocs. 3847, 

11158, 11166.

Phytocrene sordida (Lesquereux) MacGinitie 

Plate 13, figure 4

Phytocrene sordida (Lesquereux) MacGinitie, 1941, Carnegie Inst.

Washington Pub, 534, p. 140, pi. 35, fig. 2; pi. 36, fig. 3. 

Ficus sordida Lesquereux, 1878, Harvard Mus. Corap. Zoology 

Mem., v. 6, no. 2, p. 17, pi, 4, figs. 6, 7.

Discussion. The seven primary veins, cordate 

base, and few craspedodrome marginal tertiary 

veins that end as small protuberances at the margin, 

place these leaves in Phytocrene. The spacing of the 

intercostal tertiary veins and the angle of departure 

of the medial secondary veins in the fossils are 

similar to that of P. blancoi (Aza.) Men*., but this 

species does not have craspedodrome tertiary veins. 

The marginal characteristics of the fossils are 

similar to those of P. bracteata Wall., but the 

intercostal tertiary veins are more widely spaced and 

the medial secondaries in the apical region depart 

almost perpendicularly from the midrib.

Hypotype.-USNM 43365.

Occurrence. Kulthieth Formation, Iocs. 11170, 

11195.

Genus Pyrenacantha Wight 

Pyrenacantha sp.

Plate 11, figure 2; text figure 31A

Discussion. The fossil leaf has several characteristics that indicate a close relationship of Pyrenacantha: pinnate venation, a weak pair of basal 

secondaries that form marginal loops whose apical 

sides are oriented basally, the next more apical 

secondaries are stronger than the basal pair, some 

tertiary veins that depart and extend basally from 

the midrib, widely spaced and irregularly branching 

intercostal tertiary veins, and marginal tertiary 

veins that extend from loops of the secondary veins

1 cm

FIGURE 31. Marginal venation of Pyrenacantha. A, Pyrenacantha sp., USNM 43366, loc. 11170. 

B, P. repanda, USGS Ref. Colln. 1873. C, P. volubilis, USGS Eef. Colln. 1052a.
78 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

B

1 cm

FIGURE 32. Marginal venation of Stemonurus. A, S. 

alaskanus, holotype, USNM 43368, loc. 11158. B, S. 

scorpiodes, USGS Ref. Colln. 1874.

to a few widely spaced glands on the margin. The 

Kulthieth species is particularly close to P. volubilis 

Wight from Hainan; the fossil differs from leaves of 

this extant species in having the loops of the 

secondary veins placed much closer to the margin.

Specimen. USNM 43366.

Occurrence. Kulthieth Formation, loc. 11170.

Genus Stemonurus Blume

Stemonurus alaskanus Wolfe, new species

Plate 13, figures 5, 6; text figure 32A

Diagnosis. Leaves simple; venation pinnate; 

shape oval; base acute, apex abruptly acute with 

drip-tip; 9-12 pairs of secondaries broadly curving or 

straight, the most basal pair aparallel to the others, 

looping sharply near the margin; marginal tertiary 

veins forming a series of small, angular loops; 

intersecondaries numerous, extending almost to 

major loops, forking; intercostal tertiary veins

irregularly spaced, few, irregularly forking; margin 

entire; texture coriaceous; petiole thick.

Discussion. The difference in orientation between the most basal pair of secondaries and the 

remaining secondaries, the numerous intersecondaries, the irregular tertiary venation, and the 

angular looping of the fossils indicates a close 

relationship to Stemonurus, and particularly to S. 

scorpiodes Becc. The fossils differ from leaves of this 

extant Indonesian species in the broader shape and 

slight curvature of the secondary veins of the fossils.

Holotype.-USNM 43368.

Pamtype.-USNM 43367.

Occurrence. Kushtaka Formation, loc. 11158.

Family SAPINDACEAE

Genus Allophylus Linnaeus 

Allophylus duktothensis Wolfe, new species

Plate 14, figures 3, 4

Diagnosis. Leaf compound; venation of leaflets 

pinnate; shape oval to ovate, broad; base acute, apex 

acuminate; six to eight pairs of secondary veins, 

straight to slightly curving, craspedodrome, entering teeth along apical sides or ending in sinuses or 

forking, with one fork extending apically and the 

other entering the teeth or sinus; some marginal 

tertiary veins craspedodrome; intercostal tertiary 

veins forming a thin, irregularly forking, large mesh; 

teeth typically one or two per secondary vein; sinuses 

arcuate.

Discussion. Toothed leaves of most Sapindaceae 

have secondary veins that tend to enter teeth along 

apical sides or to end in the sinuses. This feature, in 

combination with the irregular intercostal tertiary 

venation pattern, indicates that the fossils represent 

Allophylus. Among extant species, A. duktothensis 

most closely resembles A. peduncularis Radlk. in 

shape and intercostal venation; this extant Philippine species, however, has but one tooth per 

secondary vein and the sinusus are arcuate or 

angular.

Holotype.-USNM 43372.

Paratype. USNM 43367.

Occurrence. Kulthieth Formation, loc. 11170.

Allophylus wilsoni Chaney and Sanborn 

Plate 29, figures 4, 6

Allophylus wilsoni Chaney and Sanborn, 1933, Carnegie Inst. 

Washington Pub. 439, p. 81, pi. 24, figs. 1-4, 6.

Discussion. Some Alaskan specimens are narrower than the Goshen specimens of this species, but 

other Alaskan specimens are just as broad.

Hypotypes. USNM 43374A, B, 43375.

Occurrence. Katalla Formation, Split Creek beds,
SYSTEMATICS 79

B

1 cm

FIGURE 33. Marginal venation of Meliosma. A, M. 

duktothensis, holotype, USNM 43378, loc. 11170. 

B, M. pungens, USGS Ref. Colln. 100.

Iocs. 9552, 9889, 11168; Kulthieth Formation, Iocs. 

9551, 9553.

Allophylus sp. 

Plate 24, figure 3

Discussion. The asymmetry of the basal part of 

the lamina indicates that this specimen represents a 

leaflet. The marginal venation is typically sapindaceous, and the teeth about one per secondary vein 

also resemble the teeth of Allophylus. The poor 

preservation of the tertiary venation, however, 

makes the generic reference tentative.

Specimen. USNM 43376.

Occurrence. Kushtaka Formation, loc. 11165.

Genus Sapindus Linnaeus 

Sapindus? sp.

Plate 24, figure 9

Discussion. The strongly falcate shape, thick 

primary vein, entire margin, and camptodrome 

secondary venation indicate a relationship to 

Sapindus. The thick texture of the specimen has 

obscured the tertiary venation pattern, and thus the 

generic reference is not secure.

Specimen. USNM 43377.

Occurrence. Kushtaka Formation, loc. 11165.

Family SABIACEAE

Genus Meliosma Blume

Meliosma duktothensis Wolfe, new species

Plate 14, figures 6-8; text figure 33

Diagnosis. Leaves simple; venation pinnate; 

shape narrowly oval to obovate; base cuneate; 8-12 

pairs of broadly curving secondary veins, departing

at an angle of about 45°, forking near margin, apical 

fork extending apically and the basal entering tooth; 

intercostal tertiary veins widely spaced, branching, 

approximately perpendicular to secondaries; margin 

typically with one or two teeth per secondary vein; 

teeth small, almost spinose, separated by arcuate 

sinuses; petiole long.

Discussion. The fossil leaves have the arcuate 

sinuses and sharp teeth characteristic of Meliosma', 

the forking of the secondary veins near the margin is 

characteristic of some species of Meliosma, for 

example M. simplicifolia (Roxb.) Walp. (van Beusekom, 1971, p. 407). This extant subtropical Asian 

species also resembles M. duktothensis in having 

one or two teeth per secondary vein. M. simplicifolia, 

however, has a short petiole and the secondary veins 

fork farther from the margin than in M. duktothensis.

Holotype.-USNM 43378.

Paratypes. USNM 43379, 43380.

Occurrence. Kulthieth Formation, Iocs 11170, 

11195.

Meliosma kushtakensis Wolfe, new species 

Plate 14, figures 1, 2

Diagnosis. Leaves simple; venation pinnate; 

shape oval; base cuneate; 13 or 14 pairs of secondary 

veins, broadly curving, departing at an angle of 

approximately 60°, typically craspedodrome and 

branching; intercostal tertiary and quartary veins 

approximately perpendicular to secondaries and 

alternating, closely spaced; typically two teeth per 

secondary vein; teeth small, almost spinose, separated by arcuate sinuses; petiole long.

Discussion. The character of the intercostal 

venation indicates a relationship to the extant 

Meliosma simplicifolia. The intercostal venation, 

tendency towards craspedodromy, and higher angle 

of departure of the secondary veins distinguishes M. 

kushtakensis from M. duktothensis. Although both 

fossil species are related to the same extant species, 

van Beusekom (1971) considered them to resemble 

different subspecies, which some systematists 

considered to be of specific rank.

Holotype.-USNM 43382.

Paratype. USNM 43381.

Occurrence. Kushtaka Formation, Iocs. 11158, 

11159, 11166.

Order RHAMNALES

Family RHAMNACEAE

Genus Rhamnus Linnaeus

Rhamnus sp. 

Plate 27, figure 2

Discussion. The fossil has a finely serrate
80 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

margin, smoothly curving and parallel secondary 

veins that extend along the margin, and branching 

tertiary veins that are oriented obliquely to the 

secondaries. The specimen appears, therefore, to 

represent a species of Rhamnus.

Specimen. USNM 43369.

Occurrence. Kushtaka Formation, loc. 11167.

Genus Sageretia Brongnairt

Sageretia sp. 

Plate 15, figure 1

Discussion. Leaves of Sageretia typically have: 

many small and closely spaced, rounded teeth; 

smoothly curving, camptodrome secondary veins; a 

pair of secondary veins that depart at or near the 

base, giving the leaf an almost palmate venation; a 

broadly rounded base; and numerous, closely spaced, 

and anastomosing intercostal tertiary veins that are 

oriented perpendicular (or nearly so) to the midrib. In 

all these features, a few fragmentary specimens from 

the Kushtaka Formation qualify as Sageretia.

Sageretia is also known in the early Eocene Chalk 

Bluffs assemblage from California (see p. 91) and the 

middle Miocene Latah assemblage from Washington.

Specimen. USNM 43370.

Occurrence. Kushtaka Formation, loc. 11166.

Genus Zizyphus Linnaeus 

Zizyphus alaskanus Wolfe, new species

Plate 14, figures 5, 9

Diagnosis. Leaf simple; venation palmate, with 

five primary veins; shape pronouncedly ovate; base 

broadly rounded, apex acuminate; first pair of lateral 

primaries extending about three-quarters distance to 

apex and looping; second pair confined to basal 

quarter of lamina; lateral secondary veins numerous, camptodrome; lateral tertiary veins craspedodrome; two pairs of medial secondary veins, 

camptodrome; numerous, closely spaced intercostal 

tertiary veins, oriented perpendicular to midrib; 

numerous small teeth, rounded.

Discussion. The fossil leaf superficially resembles the leaves of Tetracentron piperoides (Lesq.) 

Wolfe, but the closely spaced tertiary veins that 

connect the lateral primaries to the midrib and the 

smaller teeth of the fossil indicate that it is distinct 

from Trochodendraceae and Cercidiphyllaceae. The 

following characteristics of the fossil, however, 

indicate that it is assignable to Zizyphus: (1) palmate 

venation, (2) a steeply ascending pair of first lateral 

primaries and a weak pair of second lateral 

primaries, (3) closely spaced tertiary veins that 

connect the first lateral primaries to the midrib, (4)

medial secondary veins that are aparallel to the first 

lateral primaries that depart from the midrib more 

than half way to the apex, (5) camptodrome 

marginal tertiary veins and craspedodrome marginal quartary veins, (6) closely and regularly spaced 

and glandular teeth, (7) a broadly rounded base, and 

(8) a highly attenuated apex. In lacking aerodrome 

first lateral primary veins, Z. alaskana most closely 

resembles the leaves of the extant Z. fungi Merr. of 

Hainan; the latter species, however, has numerous 

intersecondary veins that are lacking in Z. alaskana 

and smaller and fewer teeth. Intercostal venation 

similar to that of the fossil is found in Z. inermis 

Merr. from the Philippine Islands, but this species 

has smaller teeth and aerodrome first lateral 

primaries. In size and spacing of the teeth, Z. 

alaskana is similar to Z. yunnanensis from the 

Paratropical Rain forest of southern China, but this 

living species has aerodrome first lateral primaries. 

Although all characters of the fossil can be found in 

leaves of extant members of Zizyphus, no living 

species has leaves that have all the characters of Z. 

alaskana. It should be noted that the modern leaves 

morphologically most similar to the fossil are found 

in species of the Paleotropical rain forest; the warm 

temperate and neotropical species of Zizyphus do not 

have leaves closely similar to Z. alaskana.

Holotype. USNM 43371A, B.

Occurrence. Kushtaka Formation, loc. 11158.

Family VITACEAE 

Genus Vitis Linnaeus

Vitis sp. 

Plate 15, figures 2, 4, 5

Discussion. This species is represented by abundant material, but I have not examined a sufficient 

amount of cleared material of extant members of 

Vitis to diagnose the species. The lack of lobes, 

narrow leaf shape (for Vitis), and shallowly cordate 

base, make this species readily recognizable. What 

appears to represent the same species is also known 

in an early Ravenian assemblage from the Puget 

Group.

Specimens. USNM 43383-43385.

Occurrence. Kushtaka Formation, loc. 11166; 

Kulthieth Formation, loc. 11170.

Order MALVALES

Genus Tilia Linnaeus

Tilia carbonensis Wolfe, new species

Plate 27, figures 1, 5

Diagnosis. Leaves simple; venation palmate; 

shape ovate; base broadly rounded, apex acute; first 

lateral pair of primary veins extending about one-
SYSTEMATICS 81

1 mm

FIGURE 34. Ultimate venation of Plafkeria sp. USNM 43388A, 

loc. 11167.

third distance to apex, craspedodrome, giving off 

about four or five craspedodrome lateral secondary 

veins; six to eight pairs of medial secondary veins, 

straight to slightly curving, craspedodrome, giving 

off no or up to two (greater basally) craspedodrome 

branches on basal sides; intercostal tertiary veins 

closely and uniformly spaced, perpendicular to 

secondaries, branching to form a quartary vein 

perpendicular to tertiary; quartary branch of tertiary 

extending midway to adjacent tertiary, bending 

sharply to parallel tertiary and extending to join 

secondary; teeth large, uniformly spaced, aristate 

shaped, separated by angular sinuses.

Discussion. The palmate venation, closely and 

uniformly spaced tertiary veins, and the aristate 

teeth indicate that the fossils represent Tilia. No 

extant species of Tilia that I have examined appears 

to be closely related to T. carbonensis. Most species 

have either a cordate base or more numerous teeth or 

both.

Holotype. USNM 43386A.

Paratype. USNM 43387.

Occurrence. Kushtaka Formation, loc. 11167.

Family TILIACEAE 

Genus Plafkeria Wolfe, new name 

Willisia Wolfe, 1968, U.S. Geol. Survey Prof. Paper 571, p. 24.

The name Willisia was originally proposed by 

Warming in 1901 for a genus of Podostemaceae. The 

name that I proposed in 1968 for a genus of tiliaceous 

leaves is thus clearly a junior homonym. This genus 

is here renamed Plafkeria, in recognition of George 

Plafker's extensive contributions to our knowledge 

of the geology of the Gulf of Alaska region.

Type species. Plafkeria rentonensis (Wolfe) 

Wolfe, new combination: Willisia rentonensis Wolfe, 

1968, U.S. Geol. Survey Prof. Paper 571, p. 24, pi. 7, 

figs. 3, 5; text figure 19A.

Plafkeria sp. 

Plate 27, figure 6; text figure 34

Discussion. The incompleteness of the specimen 

unfortunately does not allow it to become a namebearing specimen, although it has features that

1 cm

FIGURE 35. Marginal venation of Saurauia. A, S. sp., USNM 

43389, loc. 3846. B, S. reinwardtiana, USGS Ref. Colln. 505.

distinguish it from the other two known species of 

Plafkeria. The Kushtaka species differs from the 

genotype P. rentonensis in having broader, more 

widely spaced intercostal tertiary veins, a more 

strongly developed pair of second lateral primary 

veins, and areoles that are slightly larger. The 

Kushtaka species differs from P. obliquifolia in 

having a less broad base, the inframarginal loops 

closer to the margin, and the second pair of lateral 

primaries less strongly developed. The morphologic 

gradation indicates that these three species of 

Plafkeria may represent a lineage; no Kummerian 

representative of the lineage is yet known.

Specimen. USNM 43388A.

Occurrence. Kushtaka Formation, loc. 11167.

Order PARIETALES 

Family ACTINIDIACEAE 

Genus Saurauia Wildonow

Saurauia sp. 

Plate 18, figure 1; text figure 35

Discussion. Several features of the specimen 

indicate that it represents Saurauia: widely spaced, 

sharp, small teeth; numerous, closely spaced, 

branching intercostal tertiary veins that tend to be 

oriented perpendicular to the midrib; broadly 

curving secondary veins that curve sharply towards 

the apex as they approach the margin and form an 

extensive series of loops; and craspedodrome 

marginal tertiary veins. The Kushtaka leaf resembles most closely S. reinwardtiana Miq. of Indonesia. 

The only significant differences noted are in the 

slightly wider spacing of both teeth and intercostal 

tertiary veins in the Kushtaka leaf.

Specimen. USNM 43389.

Occurrence. Kushtaka Formation, loc. 3846.

Family THEACEAE
82 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Genus and species indetermined

Plate 25, figure 3

Discussion. Several leaves have (1) a short, thick 

petiole, (2) a margin that is entire in the basal part 

and that has a few, irregularly spaced, rounded teeth 

in the apical part, (3) secondary veins that are 

straight to slightly curving and that loop angularly 

near the margin, and (4) a coriaceous texture. 

Therefore, these leaves probably represent Theaceae, but the poor preservation of the venation does 

not allow a generic assignment.

Specimen. USNM 43390.

Occurrence. Kushtaka Formation, loc. 11165.

Family DIPTEROCARPACEAE

The published fossil record of dipterocarps is 

meager, particularly in regard to Paleogene occurrences. Rasky (1956) has recorded a flower of the 

family from the late Eocene of central Europe, but 

the family is notably lacking in the extremely 

diverse early Eocene London Clay flora. Because of 

the apparent scarcity of the family in the Paleogene, 

I concluded several years ago that the members of 

the family either had leaves that did not preserve 

well or that the family had not yet achieved a 

significant place in tropical forests; I had, therefore, 

not examined closely the extant foliage. Today, of 

course, the family is one of the dominant families of 

the Paleotropical rain forest and extends into the 

Paratropical Rain forest. Regarding the preservation factor, the leaves hold together extremely well in 

the clearing process, and it thus seems probable that 

the leaves should preserve well.

Leaves of most genera of Dipterocarpaceae are 

characterized by having tertiary veins that are 

considerably thinner than the secondary veins and 

that are closely spaced and undulatory or anastomosing. The secondary veins, moreover, either extend 

to, or almost to, the margin without looping; the 

secondaries thin conspicuously near the margin, and 

the loops are typically not conspicuous. Leaves of 

some members of Rhamnaceae particularly Berchernia, Karwinskia, and Rhamnidium are similar 

in venation, but they can be distinguished from 

Dipterocarpaceae of the type just discussed. Karwinskia and Rhamnidium both have pellucid dots, 

which can be discerned on well-preserved fossils. 

Even in poorly preserved material, however, 

dipterocarps can be distinguished from Rhamnaceae 

by the character of the tertiary venation that extends 

from the midrib to the secondary veins. In the 

rhamnaceous genera mentioned, the tertiary veins 

typically are perpendicular to the midrib and extend 

in an approximately straight line to intersect the

B D

FIGURE 36. Venation of Parashorea. A, B, P.pseudogoldiana, 

lectotype, USNM 38978, loc. 3847. C, P. pseudogoldiana, 

hypotype, USNM 222841, loc. 11195, D, P. malaanon, USGS 

Ref. Colln. 1663.

secondary veins obliquely. In Dipterocarpaceae the 

tertiary veins are also typically perpendicular to the 

midrib, but they curve basally and intersect the 

secondary veins at approximately a right angle. On 

this basis, the leaves described as Rhamnidium 

chaneyi Potbury (1935), Rhamnites cashmanensis 

Wolfe, and R. franklinensis Wolfe (1968) are 

members of Dipterocarpaceae. The preservation of 

the specimens on which these species are based is 

such that pellucid dots would have been preserved.

Genus Parashorea Kurz 

Parashorea pseudogoldiana (Hollick) Wolfe, new combination

Plate 16, figures 6-8; plate 19, figure 6, text figure 36A, B 

Rhamnuspseudogoldianus Hollick, 1936 (part), U.S. Geol. Survey

Prof. Paper 182, p. 139, pi. 79, figs. 1, 2 only. 

Rhamnus marginatus auct. non Lesquereux. Hollick, 1936, U.S.

Geol. Survey Prof. Paper 182, p. 140, pi. 78, figs. 1, 2. 

Cornus hyperborea auct. non Heer. Hollick, 1936, U.S. Geol. Survey Prof. Paper 182, p. 158, pi. 99, figs. 2, 3.

Discussion. The fossils display the marginal and 

intercostal venation of Dipterocarpaceae noted 

previously. In their broadly rounded to slightly 

cordate base and numerous steeply ascending 

secondary veins, the fossils are most similar to 

leaves of the extant Parashorea.

A fragmentary specimen from the Kulthieth 

Formation has well-preserved ultimate venation.
SYSTEMATICS 83

D

FIGURE 37. Marginal and intercostal venation of Barringtonia. 

A, C, B. sp., USNM 43392B, loc. 11170. B, D, B. acutangula, 

USGS Ref. Colln. 1986.

The pattern is strikingly similar to the extant Parashorea in having small, quadrangular areoles that 

lack freely ending veinlets and in having the next 

coarser order of venation forming a quadrangular 

mesh. The areoles of the fossil, however, are 

somewhat larger than the areoles of P. malaanonan 

collected from British North Borneo. The larger 

areoles probably indicate that the Alaskan climate 

was somewhat cooler than that of the Melanesian 

region today (p. 23).

Lectotype. USNM 38978, which is the specimen 

figured on Rollick's (1936) plate 79, figure 2, lower 

left leaf.

Syntype.-USNM 43391.

Hypo topes. USNM 39034, 39035, 222841.

Occurrence. Kushtaka Formation, loc. 3847; 

Kulthieth Formation, loc. 11195.

Order MYRTIFLORAE

Family BARRINGTONIACEAE

Genus Barringtonia Forster

Barringtonia sp. 

Plate 17, figures 3, 4; text figure 37

Discussion. The fossils superficially resemble 

some leaflets of Juglandaceae. In that family the 

fourth order intercostal venation forms a quadrangular or polygonal mesh, but in the fossils the fourth 

order intercostal veins are perpendicular to and 

directly connect the tertiary veins. This unusual 

fourth order venation, the closely serrate margin, 

and the two series of marginal loops (the marginal 

quartary veins are craspedodrome) represent a 

highly characteristic combination of characters that 

among extant plants I have seen only in leaves of 

Barringtonia. Except for the wider spacing and 

higher degree of percurrency of the intercostal 

tertiary veins of B. duktothensis, the fossils are very 

similar to B. speciosa Bl.

Specimen. USNM 43392A, B.

Occurrence. Kulthieth Formation, loc. 11170.

Family RHIZOPHORACEAE

Genus Kandelia Wight and Arn.

Kandelia wangi Wolfe, sp. nov.

Plate 19, figs. 1, 5a, b.

Diagnosis. Leaves pinnately veined, simple; 

oval apex broadly rounded, base decurrent along 

petiole; midrib thick; about six pairs of secondary 

veins; pronouncedly decurrent along the midrib in 

the basal half of the lamina, departing at angles of 

about 30° in the basal half and about 45° in the upper 

half, extending straight towards the margin, curving 

apically about three-fourths the distance to the 

margin, camptodrome, with the loops almost 

paralleling the margin; between two adjacent 

secondary veins one or two strong veins extend 

admedially, branch irregularly about one-half the 

distance to the margin; tertiary veins widely and 

irregularly spaced, branching irregularly to form a 

coarse, irregular mesh; fourth order veins extending 

into the mesh, thinning conspicuously, branching 

irregularly to form areoles; areoles intruded by 

irregularly and profusely branching veinlets; margin entire, without marginal vein; petiole thick and 

short.

Discussion. Leaves of several genera of Rhizophoraceae are closely similar in having secondary 

veins that extend straight towards the margin and 

loop approximately parallel to the margin and an 

irregular intercostal venation. In Rhizophora, 

Bruguiera, and Ceriops, however, there are typically 

two or more strong veins between and parallel to the 

secondary veins. In Carallia there are large, 

irregular loops between the secondary loops and the 

margin. In Cassipourea the margin has a few, 

irregularly spaced teeth. In Blepharistemma the 

secondary and intercostal venation is formed by 

conspicuously weak veins.

In Kandelia candel (L.) Druce, all significant 

features of the fossils can be matched although the 

secondary veins are more closely spaced in the 

fossils than in this extant species. Notable similarities, other than those common to the several genera 

of Rhizophoraceae previously mentioned, occur in 

the marginal and ultimate venation. The marginal 

venation has a single series of small, angular loops 

of irregular shape between the secondary loops and 

the margin. The areoles are composed of irregularly 

shaped fourth order veins and are intruded by thick, 

profusely branching veinlets.

Holotype. USNM 222843.

Paratype. USNM 222844.

Occurrence. Kulthieth Formation, loc. 11195.
84 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

1 cm

FIGURE 38. Marginal venation oiAlangium. A, A. bergensis, 

holotype, USNM 38926, loc. 3847. B, A. longiflorum, USGS 

Ref. Colln. 1931,

Family ALANGIACEAE

Genus Alangium Lamarck

Alangium bergensis Wolfe, new species

Plate 16, figure 2; text figure 38

Malpoenna magnified (Saporta) Hollick, 1936 (part, nontypic), 

U.S.. Geol. Survey Prof. Paper 182, p. 118, pi. 66, fig. 5, only.

Diagnosis. Leaves simple, venation palmate; 

shape slightly obovate; base asymmetric and 

rounded, apex acuminate with drip-tip; five primary 

veins, the first lateral pair extending about half 

distance to apex; second lateral primary veins weak; 

eight or nine lateral secondary veins per first lateral 

primary, extending almost straight and looping 

sharply close to margin; four or five pairs of medial 

secondary veins, gradually curving, looping sharply 

near margin and forming a series of small angular 

loops adjacent to margin; intercostal tertiary veins 

widely spaced, branching in a Y pattern; quartary 

venation forming an irregularly polygonal, large, 

mesh; margin entire.

Discussion. The fossil leaf is somewhat similar to 

leaves of certain Tiliaceae (Microcos, for example) 

and Sterculiaceae (Pterospermum). In Tiliaceae, 

however, the branches of the intercostal tertiary 

veins are approximately perpendicular to the 

tertiary veins. In Sterculiaceae, the intercostal 

tertiary veins are closely spaced and somewhat 

sinuous. In features of primary and intercostal 

venation, the fossil is similar to Alangium. Among 

extant species the fossil shows similarities to both A. 

chinense (Lour.) Rehd. and A. longiflorum Merr. The 

few medial secondary veins, extension of the first 

lateral primary veins about half way to the apex, and 

the narrow, slightly obovate shape, are features that

1 cm

FIGURE 39. Intercostal venation of Eugenia. 

A, E. sp., USNM 43395, loc. 11170. B, E. 

jambos, USGS Ref. Colln. 1709.

ally A. bergensis to A. longiflorum. The main feature 

that distinguishes leaves of these two species is that 

the looping of the secondary and tertiary veins is 

considerably closer to the margin in the fossil than in 

A. longiflorum; marginal looping similar to the fossil 

is found in A. chinense, which, however, has 

numerous medial secondary veins, first lateral 

primary veins extending only about one third the 

distance to the apex, and a broad, ovate shape. A. 

bergensis is considered to be most closely related to 

A. longiflorum.

Rollick's material from the Paleocene Hamilton 

Bay assemblage was also referred to Malpoenna 

magnifica; this material represents a species that 

has pinnate venation, weak basal secondary veins, 

and a pair of strong secondary veins departing from 

the midrib apically from the base, that is, the species 

has the diagnostic features of a member of 

Lauraceae.

Holotype. USNM 38926.

Occurrence. Kushtaka Formation, loc. 3847.

Alangium sp. 

Plate 16, figure 1

Discussion. The specimen has tertiary veins 

branching in a Y pattern and the irregular fourth 

order mesh of Alangium. A. bergensis cannot be 

represented, however, because of the highly asymmetric base, the presence of seven primary veins, and 

the more closely spaced intercostal tertiary veins.

Specimen. USNM 43393.

Occurrence. Kushtaka Formation, loc. 11166.

Family MYRTACEAE 

Genus Eugenia Linnaeus

Eugenia sp. 

Plate 16, figures 3-5; text figure 39A

Discussion. The continuous inframarginal vein 

(formed by looping of the secondary veins), the 

closely spaced and numerous secondary veins, and 

the numerous intersecondary veins paralleling the
SYSTEMATICS 85

secondaries indicate that these fossils are referable 

to Myrtaceae. Among extant species the fossils 

appear to be closest to members of Eugenia. In the 

highly linear shape, the fossils resemble the extant 

E. jambos L.; leaves of this species differ from the 

fossils in having the submarginal vein spaced 

farther from the margin, and the intervening space 

has an additional series of loops. The leaves of E. 

multiflora Hance are similar to the fossils in 

marginal venation, but the former are ovate.

Specimens. USNM 43394-43396.

Occurrence. Kulthieth Formation, loc. 11170. 

Eugenia? sp.

Plate 27, figure 7.

Discussion. The continuous inframarginal vein 

places this fragment in Myrtaceae. The anastomosing intercostal tertiary veins indicate an assignment 

to Eugenia, but considering the fragmentary nature 

of the specimen the determination is questioned. The 

leaf is wider and the intercostal areas more 

elongated than in the early Ravenian Eugenia that 

was previously discussed.

Specimen. USNM 43397.

Occurrence. Kushtaka Formation, loc. 11167.

Order UMBELLIFLORAE

Family CORNACEAE

Genus Mastixia Blume

Mastixia irregularis (Hollick) Wolfe, new combination

Plate 15, figure 6

Cornus irregularis Hollick, 1936, U.S. Geol. Survey Prof. Paper 

182, p. 158, pi. 99, fig. 1.

Discussion. The close spacing and anastamosing of the intercostal tertiary veins and the thickened 

margin (probably a marginal vein) indicate that this 

leaf is referable to Mastixia rather than to Cornus. 

Among extant species, the fossil resembles leaves of 

M. philippinensis Merr., but in that extant species 

the leaves have more closely spaced intercostal 

tertiary veins and the more basal secondary veins do 

not extend as far towards the apex.

Holotype. USNM 39033.

Occurrence. Kushtaka Formation, loc. 3846.

Order ERIC ALES

Family CLETHRACEAE

Genus Clethra Gronov

Clethra sp.

Plate 18, figure 2

Discussion. Although the fossils superficially 

resemble some leaflets of Juglandaceae, all the 

fossils are symmetric and have a petiole, indicating 

that they are simple leaves. The finely and uniformly

serrate margin, camptodrome secondary veins, 

craspedodrome tertiary veins, and intercostal tertiary veins that are numerous and regularly spaced 

and that are not perpendicular to the secondary 

veins are characters that indicate that the fossils are 

a species of Clethra. Among extant species of the 

genus, C. alaskana appears most similar to C. 

delavayi Franch. from Yunnan. The fossils differ 

chiefly in having the secondary veins extending 

closer to the margin before looping.

Specimen. USNM 43398.

Occurrence. Kulthieth Formation, loc. 11170.

Order TUBIFLORAE

Family VERBENACEAE

Genus Clerodendrum Linnaeus

Clerodendrum sp. 

Plate 7, figures 1-3, 6; plate 17, figure 1

Discussion. This species is represented by five 

incomplete specimens in proximity on one slab and 

its counterpart. The most complete specimen is the 

one figured on plate 7, figure 3 and its counterpart on 

plate 17. The following features of the fossils indicate 

an assignment to Clerodendrum: extremely thick 

and short petiole, extremely thick midrib that thins 

conspicuously in an apical direction, broadly 

curving and widely spaced secondary veins, that are 

camptodrome and form a series of almost quadrangular loops, intercostal tertiary veins that are thin 

and branch to form quartary veins that thin 

conspicuously, a thickened margin, and extremely 

small teeth. Whether the small teeth should be 

considered teeth is questionable; as in the leaves of 

some extant species of the genus, the "teeth" of the 

fossils appear to be only slight bumps. The fossils 

particularly resemble the extant C. nutans Wall, of 

Burma, differing primarily in the pronounced 

rounding of the base.

Specimens. USNM 43399, 43400, 43401 A, B, 

43402.

Occurrence. Kushtaka Formation, loc. 11158.

Genus Holmskioldia Retzius 

Holmskioldia speiri (Lesquereux) MacGinitie?

Plate 30, figure 4

Holmskioldia speirii (Lesquereux) MacGinitie, 1953, Carnegie 

Inst. Washington Pub. 599, p. 156, pi. 74, figs. 1, 2 (see 

synonymy).

Parana speirii Lesquereux, 1883, U.S. Geol. Survey Terr. Rept, 

v. 8, p. 172, pi. 28, fig. 15.

Discussion. This determination is queried because of the fragmentary nature of the specimen. In 

details of venation, however, the Alaskan specimen 

matches material from conterminous United States 

assigned to this species.
86 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

Specimen. USNM 43403.

Occurrence. Katalla Formation, Split Creek 

Sandstone Member, loc. 11168.

INCERTAE SEDIS

Genus Macclintockia Heer

Macclintockia pugetensis Wolfe

Plate 30, figures 5, 6, 8-10

Macclintockia pugetensis Wolfe, 1968, U.S. Geol. Survey Prof. 

Paper 571, p. 26, pi. 7, figs. 8, 9.

Discussion. Fragments of this characteristic 

species are present in almost every plant collection 

that has been made by geologists from marine rocks 

of Keasey and earliest Lincoln age in the Gulf of 

Alaska province. The leaves must have been very 

coriaceous to be so widely dispersed in the sea.

Hypotypes. USNM 43404-43408.

Occurrence. Kushtaka Formation, loc. 9891; 

Katalla Formation, Split Creek Sandstone Member, 

Iocs. 9552,9889,11168, Basin Creek beds Iocs. 11188?, 

GAS 29181; Kulthieth Formation, Iocs. 9551, 9553, 

9893, 9894, 11189.

Macclintockia sp. 

Plate 25, figure 8

Discussion. The basal parts of two leaves have 

the characteristic venation of Macclintockia. Both 

specimens are broader than M. pugetensis, and they 

have strong lateral veins that depart from the top of 

the petiole. Although possibly related to the 

Kummerian species, the middle Ravenian species 

appears to be a distinct entity.

Specimen. USNM 43409.

Occurrence. Kushtaka Formation, loc. 11164.

Genus Dicotylophyllum Saporta

Dicotylophyllum carbonensis Wolfe, new species

Plate 27, figures 8, 10

Diagnosis. Leaves simple; venation pinnate; 

shape linear, oval; base acute, apex acute; 7-10 pairs 

of secondary veins straight to typically broadly 

curving, craspedodrome or camptodrome; if camptodrome, branching to send more basal branch into 

tooth centrally and more apical branch to parallel 

margin and join next more apical secondary vein; 

intercostal tertiary veins approximately perpendicular to secondaries, forking, widely spaced; margin 

with widely and uniformly spaced almost spinose 

teeth separated by arcuate sinuses.

Discussion. I am unable to suggest what extant 

group these fossils might represent. They are similar 

to Dicotylophyllum flexuosa (Newb.) Wolfe, a 

widespread Paleocene species. This older species, 

however, has more closely spaced teeth and the 

secondary veins are typically camptodrome.

Holotype. USNM 43410.

Paratype. USNM 43411.

Occurrence. Kushtaka Formation, loc. 11167.

Dicotylophyllum kummerensis (Wolfe) Wolfe, 

new combination

Plate 25, figure 7; plate 30, figure 11

Artocarpoides kummerensis Wolfe, 1968, U.S. Geol. Survey Prof. 

Paper 571, p. 19, pi. 4, figs. 1, 7.

Discussion. I agree with MacGinitie (in press) 

that the leaves he described as Laurophyllum 

litseafolia and that I transferred to Artocarpoides 

represent neither Lauraceae nor Moraceae, but what 

family these leaves represent is unknown. The close 

relationship between MacGinitie's species and the 

species described from the upper part of the Puget 

Group is apparent, and I thus transfer A. kummerensis to Dicotylophyllum.

Specimens. USNM 43412, 43413.

Occurrence. Kushtaka Formation, Iocs. 9891, 

11164.

Dicotylophyllum spp.

Plate 17, figure 2. The specimen is an entire 

margined leaf that has steeply ascending secondary 

veins and numerous intersecondaries. It may 

represent Lauraceae, but is is too fragmentary for 

determination. USNM 43414, locality 11170, Kulthieth Formation.

Plate 18, figure 3. This is the holotype of 

Rhododendron crassum Hollick. Almost nothing 

remains of the secondary venation pattern, it is 

obscured by the extremely coriaceous texture. I 

therefore propose that Rhododendron crassum 

Hollick, 1936, p. 160, pi. 99, fig. 4, be declared a nomen 

nudum because of insufficient characters. USNM 

39036, locality 3847, Kushtaka Formation.

Plate 18, figure 5. The margin has widely spaced, 

small, and sharp teeth. The character of the margin 

and the widely spaced, branching intercostal 

tertiary veins are features also found in Pterostyrax; 

leaves of this genus, however, have straight 

secondary veins. USNM 43415, locality 11159, 

Kushtaka Formation.

Plate 18, figure 6. This leaf may be an aberrant 

leaf of Alnus scotti; the intercostal venation is 

particularly similar to that alder. None of the 

specimens attributed to A. scotti, however, have the 

pronouncedly angular sinuses. USNM 43416, locality 11158, Kushtaka Formation.

Plate 24, figure 2. The difference in angle of 

departure of the secondary veins on one half the 

lamina as compared to the other indicate that this 

may represent a leaflet. The secondary veins fork 

before entering the teeth, and one branch enters a 

tooth along the apical side. The specimen may 

represent Sapindaceae. USNM 43417, locality 11165, 

Kushtaka Formation.
SYSTEMATIC REVISIONS OF OTHER PALEOGENE SPECIES 87

1 cm

FIGURE 40. Venation of Salix and Lithocarpus. A, D, S. coatsi, paratype, UCMP 

8910. B, E, L. densiflora, USGS Ref. Colln. 45. C, F, L. glabra, USGS Ref. Colln. 

192.

Plate 24, figure 4. The specimen has an entire 

margin and a mucronate apex. A pair of weak basal 

secondary veins appears to be present; the specimen 

could represent a species of Lauraceae. USNM 43418, 

locality 11165, Kushtaka Formation.

Plate 24, figure 6. The linear shape, coriaceous 

texture, and entire margin occur in a wide range of 

plants. The venation is too poorly preserved for 

determination. USNM 43419, locality 9389, Kushtaka Formation.

Plate 24, figure 5. The margin appears to have 

irregularly spaced, rounded teeth. In combination 

with the straight, sharply looping secondary veins, 

this could indicate an affinity to Rutaceae. USNM

43420. locality 11165, Kushtaka Formation.

Plate 24, figure 8. The falcate shape indicates 

that the specimen represents a leaflet. A sapindaceous alliance is indicated by the broadly curving 

secondary veins and the few teeth that appear to be 

entered by veins along the apical sides. USNM

43421. locality 11165, Kushtaka Formation.

Plate 26, figure 8. The specimen has camptodrome secondary veins, percurrent tertiary veins, 

and widely spaced, small teeth. It may represent a 

rosaceous genus, such as Prunus, but characters for 

generic assignment are too few. USNM 43430, loc. 

11167, Kushtaka Formation.

Plate 27, figure 9. Two fragments of a lamina 

have an entire margin and broadly curving 

secondary veins. The material is too fragmentary to 

even suggest possible affinities. USNM 43422, 

locality 11167, Kushtaka Formation.

Plate 30, figure L The secondary venation, which 

is poorly preserved, appears to have been pinnate 

and camptodrome. The entire margin, rotund shape, 

and thick petiole separate this leaf from other 

members of the Alaskan Kummerian flora. USNM 

43423, locality 11168, Katalla Formation, Split Creek 

Sandstone Member.

Plate 30, figure 2. The asymmetry indicates that 

the specimen is a leaflet. The margin is nonentire, 

but the preservation is too poor to determine the 

manner of entry of the veins into the teeth. USNM 

43424, locality 11169, Kushtaka Formation.

Plate 30, figure 3. This poorly preserved leaf was 

apparently coriaceous. The teeth are few and widely 

spaced; the sinuses appear to have been arcuate. The 

secondary veins fork and send a branch into the 

teeth. USNM 43425, locality 11168, Katalla Formation, Split Creek Sandstone Member.

SYSTEMATIC REVISIONS OF OTHER PALEOGENE 

SPECIES

The late Roland Brown of the U.S. Geological 

Survey performed a service to Tertiary paleobotany 

in publishing, from time to time, short papers in 

which he described new species and revised 

previously published species. Any paleobotanist 

who deals with an entire assemblage of Tertiary 

plants has an inherent difficulty in attempting to be 

familiar with a wide array of foliar types; some of us 

are more familiar with certain taxonomic groups 

than with others, and hence errors of specific, 

generic, and even familial assignments are made. 

Some of the previously published determinations 

that I consider to be invalid are thus revised here; 

particular attention has been directed to systematic 

revisions that document some of the distributions 

mentioned in previous sections of this report. 

Salix coatsi (Axelrod) Wolfe, new combination

Text figure 40

Lithocarpus coatsi Axelrod, 1966b (part, typic), California Univ. 

Pubs. Geol. Sci., v. 59, 67, pi. 10, figs, 12-14; pi. 12, figs. 1-8 

only.

Discussion. Leaves of Lithocarpus (including the 

Asian species that many authors refer to Pasania) 

have a characteristic venation: the areoles are small 

(less than 0.5 mm in diameter) and typically lack
88 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

freely ending veinlets, the secondary veins (on 

toothed leaves) are typically craspedodrome or fork 

once to send a strong branch into the tooth, and the 

loops are approximately square. The teeth, in 

addition, lack glands. The specimens of Lithocarpus 

coatsi, however, have large (over 1 mm in diameter) 

areoles that are intruded by once- or twice-branching 

veinlets, and the secondary veins (on toothed leaves) 

are camptodrome, typically fork well within the 

margin, and the fork in turn gives off a weak branch 

into a glandular tooth; the marginal loops are 

irregular in shape and elongated perpendicular to 

the midrib. The fossils do not, therefore, represent 

Lithocarpus. A much closer comparison can be made 

with leaves of some willows, such as Salix exigua 

Nutt., in venation and the presence of glandular 

teeth.

Pterocarya orientalis (MacGinitie) Wolfe, 

new combination

Juglans orientalis MacGinitie, 1937, Carnegie Inst. Washington 

Pub. 465, p. 135, pi. 3, fig. 6; pi. 4.

Discussion. The Weaverville specimens and 

material from late Oligocene assemblages in Oregon 

represent Pterocarya rather than Juglans. Members 

of the section Cardiocaryon, to which MacGinitie 

(1937) likened this species, have leaflets that have 

rounded, broad, abmedially pointing teeth. In 

members of the section Platyptera of Pterocarya, the 

leaflets have teeth that are sharp and apically 

pointing, and the secondary veins are camptodrome 

and form only a single series of marginal loops; on 

this basis MacGinitie's species is transferred to 

Pterocarya.

Fagopsis groenlandica (Heer) Wolfe, new combination

Quercus groenlandica Heer, 1968 (part), Flora Fossilis Arctica, 

v. 1, pt. 1, p. 108, pi. 8, fig. 8; pi. 10, fig. 1 only; pi. 11, fig. 4; 

pi. 47, fig. 1.

1880, Flora Fossilis Arctica, v. 6, no. 2, p. 10, pi. 4, fig. 5. 

1883 (part), Flora Fossilis Arctica, v. 7, no. 2 p. 89, pi. 69, fig. 4. 

Hollick, 1936, U.S. Geol. Survey Prof. Paper 182, p. 99, pi. 57,

fig. 6. 

Brown, 1962 (part), U.S. Geol. Survey Prof. Paper 375, p. 58,

pi. 19, figs. 3, 6, 12; pi. 40, figs. 1, 2.

Quercophyllum groenlandicus (Heer) Koch, 1963, Medd. om 

Gr^nland, v. 172, no. 5, p. 34, pi. 5, figs. 1-4; pi. 6, figs. 1-3.

Discussion. Koch (1963, p. 36) noted the similarity between Quercophyllum groenlandicus and the 

Oligocene Fagopsis longifolia (Lesq.) Roll. The 

similarities between these two species are in the 

typically asymmetric base, uniformly spaced teeth 

that are one per secondary vein, overall shape of the 

teeth, and numerous, uniformly spaced secondary 

veins. The Paleocene Q. groenlandicus thus falls 

within the limits of Fagopsis.

Fagopsis groenlandica typically has more numerous secondary veins and rounder teeth than F. 

longifolia. An undescribed species of the genus from 

the Eocene Republic assemblage of Washington 

has the round teeth ofF. groenlandica and the fewer 

secondary veins of F. longifolia, indicating that the 

three species may constitute a phylad.

Fagopsis was transferred to Betulaceae by 

MacGinitie (1953), largely based on his interpretation of the associated and attached aments; any 

interpretation based on such impressions, however, 

is difficult. Leaves of all species of Fagopsis have a 

distinct marginal vein; this vein eliminates them 

from Betulaceae and allies the genus to Fagaceae. If 

Fagopsis is Fagaceae, it is clearly not extant; the 

marginal venation is that of Fagus (see following 

discussion of F. oregona), but the aments (some are 

also associated with leaves of Fagopsis near 

Republic) clearly indicate that the two genera can 

only be distantly related. It is possible that Fagopsis 

represents an extinct family of Fagales in which the 

aments are betulaceous and the leaves fagaceous. E. 

B. Leopold (oral commun., February 1969) reports 

that pollen extracted from the aments from 

Florissant is tricolpate; this kind of pollen is a 

feature of Fagaceae but not of Betulaceae.

Fagopsis groenlandica is one of the most 

characteristic and widespread species in the Paleocene. The species is known from many localities in 

conterminous United States, Alaska, and Greenland.

Fagus oregona (Chaney and Sanborn) Wolfe, 

new combination

Tetracera oregona Chaney and Sanborn, 1933, Carnegie Inst. 

Washington Pub. 439, p. 87, pi. 31, figs. 5-7; pi. 32, fig. 1.

Discussion. Leaves of Tetracera have irregularly 

polygonal areoles intruded by profusely branching 

veinlets that thin conspicuously; leaves of Fagus, as 

well as the fossils under discussion, have almost 

quadrangular areoles intruded by simple or oncebranching veinlets that do not thin conspicuously. 

That the fossils are not some other fagaceous genus 

is indicated by the character of the marginal 

venation: in other Fagaceae, the most abmedial 

tertiary vein that departs apically from a secondary 

vein is parallel to the margin and aparallel to the 

intercostal tertiary veins. In both extant Fagus and 

the fossils the most abmedial tertiary vein typically 

departs from the secondary vein in a course parallel 

to the intercostal tertiary veins. Sanborn's (1947) 

citation of Tetracera oregona is based on leaves that 

have widely spaced intercostal tertiary veins and a 

doubly serrate margin; they are, therefore, not 

conspecific with F. oregona.

Fagus oregona resembles F. pacifica Chan.,
SYSTEMATIC REVISIONS OF OTHER PALEOGENE SPECIES 89

which is known in the late Oligocene of Oregon. 

Similarities between these two species are in the 

sharp teeth, consistently craspedodrome secondary 

veins, and closely spaced intercostal tertiary veins. 

Dissimilarities include the more numerous secondary veins and oval shape of F. oregona.

The occurrence of Fagus in a subtropical 

assemblage such as the Goshen should not be 

surprising. Wang (1961, p. 145) notes that Fagus is 

abundant in the subtropical evergreen oak forest of 

China.

Calkinsia dilleri (Knowlton) Wolfe, new combination

Cinnamomum dilleri Knowlton, 1900, U.S. Geol. Survey 20th Ann.

Kept., pt. 3, p. 47, pi. 4, fig. 1. 

Knowlton, 1902, U.S. Geol. Survey Bull. 204, p. 59. 

Sanborn, 1935, Carnegie Inst. Washington Pub. 465, p. 18, pi.

4, figs. 1, 3, 4, 6-8.

Hyperbaena dilleri (Knowlton) Wolfe, 1968 (part), U.S. Geol. Survey Prof. Paper 571, p. 21, text fig. 15A only.

Discussion. As noted previously (p. 00), this 

species represents the genus Calkinsia, In the 

illustration of this species and modern comparative 

material (Wolfe, 1968, text fig. 15), the captions for 

15B and 15C were inadvertently reversed, that is 15B 

is Cinnamomum trichophylla and 15C is Hyperbaena mexicana.

Calkinsia franklinensis Wolfe

Calkinsia franklinensis Wolfe, 1968, U.S. Geol. Survey Prof. Paper

571, p. 20, pi. 4, figs. 4, 8. 

Hyperbaena dilleri (Knowlton) Wolfe, 1968 (part, nontypic), U.S.

Geol. Survey Prof. Paper 571, p. 21, references to Iocs. 1027,

9678.

Discussion. The early Fultonian specimens referred to Hyperbaena dilleri are merely slightly 

narrower leaves of Calkinsia franklinensis (see also 

p. 63).

Cinnamomophyllum bendirei (Knowlton) Wolfe, 

new combination

Cinnamomum bendirei Knowlton, 1902, U.S. Geol. Survey Bull.

204, p. 57, pi. 10, fig. 4. 

Philadelphus bendirei (Knowlton) Chaney, 1927 (part), Carnegie

Inst. Washington Pub. 346, p. 118, pi. 17, figs. 1, 2 only. 

Sassafras bendirei (Knowlton) Brown, 1940, Washington Acad.

Sci. Jour., v. 30, p. 350. 

Sassafras columbiana Chaney and Axelrod (nontypic), 1959,

Carnegie Inst. Washington Pub. 617, p. 179, pi. 34, fig. 2

only. 

Persea pseudocarolinensis auct. non Lesquereux. Chaney and

Axelrod, 1959 (part), Carnegie Inst. Washington Pub. 617,

p. 177, pi. 34, fig. 3 only.

Discussion. The generic determination of this 

species has almost come full circle. The validity of 

the reference of the species to Lauraceae was 

demonstrated by Brown (1940), but he considered 

that the species represented Sassafras. Leaves of 

Cinnamomum and Sassafras are, however, readily

separable, even if "unlobed" leaves of Sassafras are 

compared with leaves of Cinnamomum. In Cinnamomum, as in the fossils cited in the synonymy, a 

distinct marginal vein is present and the areoles are 

small, quadrangular, and typically lack freely 

ending veinlets. Sassafras, in contrast, lacks a 

marginal vein and the areoles are large, irregularly 

polygonal, and intruded by profusely branching 

veinlets. The fossils may represent Cinnamomum, 

but the leaves of genera such as Neolitsea are equally 

similar.

Cinnamomophyllum eocernua (Chaney and Sanborn) Wolfe, 

new combination

Octoea eocernua Chaney and Sanborn, 1933, Carnegie Inst.

Washington Pub. 439, p. 74, pi. 19, figs. 1-3, 5. 

Potbury, 1935, Carnegie Inst. Washington Pub. 465, p. 68,

pi. 6, fig. 1.

Discussion. These fossils have small, quadrangular areoles, most of which lack freely ending 

veinlets. If veinlets are present, they are unbranching. The areoles of Octoea are considerably larger, 

irregularly polygonal, and are intruded by branching veinlets. The fossils are, therefore, referable to 

Cinnamomophyllum.

Euptelea orientalis (Sanborn) Wolfe, new combination

Diospyros orientalis Sanborn, 1935, Carnegie Inst. Washington 

Pub. 465, p. 26, pi. 10, fig. 3.

Aralia taurinensis auct. non Celastrus taurinensis Ward. Sanborn, 1935, Carnegie Inst. Washington Pub. 465, p. 27, pi. 10, 

figs. 1, 2, 4.

Discussion. The sharp teeth separated by arcuate sinuses, closely spaced and branching intercostal tertiary veins that are perpendicular to the 

secondary veins, and the mixed condition of 

camptodrome or craspedodrome secondary and 

marginal tertiary veins are features that indicate 

that these leaves represent Euptelea. E. orientalis 

resembles most closely the extant E. pleiosperma 

Hook. f. and Thorns. Scott and Barghoorn (1955) 

have recorded the wood of this genus from the Clarno 

nut bed assemblage.

Fortunearia weedi (Knowlton) Wolfe, new combination

Quercus weedii Knowlton, 1899, U.S. Geol. Survey Mon. 32, p. 705,

pi. 87, fig. 4. 

Corylus macquarryi auct. non Heer. Knowlton, 1899, U.S. Geol.

Survey Mon. 32, pi. 86, fig. 3.

Discussion. These leaves, which are from the 

Yellowstone National Park Eocene, represent Fortunearia. This generic assignment is substantiated 

by the: basal pair of secondary veins, widely spaced 

and branching intercostal tertiary veins, craspedodrome (typically) or camptodrome secondary veins, 

and widely spaced and spinose teeth. The Yellowstone leaves are the first North American record of
90 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

this extant monotypic genus of the Mixed Mesophytic forest; the genus is not known to be represented in 

the Neogene Mixed Mesophytic forest of western 

North America. Note that the Eocene occurrence is in 

subtropical forest; the extant species is also found in 

the evergreen oak-laurel forest of China (Wang, 1961, 

p. 143) although the species is more common in the 

Mixed Mesophytic forest.

Platanus macginitiei Wolfe, new species

Platanus aceroides auct. non Goeppert. Chaney and Sanborn, 

1933, Carnegie Inst. Washington Pub. 439, p. 78, pi. 25, fig. 6. 

Sanborn, 1937, Oregon State Mon., Studies in Geology, no. 4, 

p. 22, pi. 7, fig. 1.

Diagnosis. Leaves simple; venation pseudopalmate, that is, veins equal in strength to midrib depart 

above base of lamina; shape oval to ovate, three 

(typically) or five "lobes" that are separated by 

shallow sinuses; 8-12 pairs of secondary veins, 

typically straight, craspedodrome; numerous craspedodrome marginal tertiary veins; intercostal tertiary 

veins branching, slightly arcuate, numerous, areoles 

formed by fifth order veins, about 1 mm in diameter, 

approximately square, intruded by once- to thricebranching veinlets; margin conspicuously toothed; 

teeth large (those entered by secondary veins) or 

small (those entered by tertiary veins), separated by 

arcuate sinuses; small teeth typically at least five per 

large tooth at widest part of leaf.

Discussion. This species of Platanus is one of the 

most characteristic in the North American Tertiary 

because of the many small teeth between the large 

teeth. The late Oligocene and earlier Miocene P. 

bendirei (Lesq.) Wolfe is probably related; this 

younger species typically has one or two (rarely as 

many as four or five) small teeth per large tooth. 

Thus far P. macginitiei is known only from 

assemblages of middle Oligocene age: Scio (including the Bilyeu Creek assemblage), Sweet Home, 

Goshen, Ohanapecosh, and the pre-Angoonian 

assemblages from the Kootznahoo Formation.

This species is named for Dr. H. D. MacGinitie, in 

recognition of his many contributions to the Tertiary 

paleobotany of North America.

Holotype.-USNM 245723 (Sweet Home, Oregon, 

loc. 9107).

Paratypes. UCMP 203; Oregon State College 

Paleobotanical collection No. 500.

Crataegus cuprovallis Axelrod

Crataegus cuprovallis Axelrod, 1966b, California Univ. Pubs. 

Geol. Sci., v. 59, p. 71, pi. 16, figs. 1-5.

Holodiscus harneyensis auct. non (Arnold) Axelrod, 1950. Axelrod, 1966b, California Univ. Pubs. Geol. Sci., v. 59, p. 72, 

pi. 16, fig. 10.

Discussion. I can see no morphologic basis for

separating the specimen relegated to Holodiscus 

from those of Crataegus cuprovallis. Extant species 

of Holodiscus have very thin veins forming the 

areoles, whereas leaves of Crataegus have thick 

veins forming the areoles. In this respect, the Copper 

Basin Holodiscus should be attributed to Crataegus.

Heteromeles cuprovallis (Axelrod) Wolfe, new combination

Text figure 41

Salix cuprovallis Axelrod, 1966b, California Univ. Pubs. Geol. 

Sci., v. 59, p. 65, pi. 9, figs. 1-3.

Discussion. Axelrod (1966b) compared the fossils 

to extant willows such as Salix barclayiAnders. and 

S. bebbiana Sarg. As the accompanying drawings 

show (see fig. 41), these modern willows have leaves 

that have numerous, uniformly spaced intercostal 

tertiary veins, and smoothly curving secondary 

veins that form an extensive series of loops. S. 

cuprovallis, on the other hand, has irregularly and 

widely spaced intercostal tertiaries that are typically 

oblique to the secondary veins, and almost straight 

secondary veins that fork conspicuously and do not 

form an extensive series of loops. The fossils are 

similar, however, to the leaves of the extant 

Heteromeles arbutifolia M. Roem.

Dracontomelon sanbornae Wolfe, new species

Allophylus wilsoni auct. non Chaney and Sanborn. Sanborn, 

1935, Carnegie Inst. Wash. Pub. 465, p. 25, pi. 9, fig. 5.

Diagnosis. Leaf compound; venation of leaflet 

pinnate; shape ovate; base asymmetric; apex 

acuminate with drip-tip; 14 or 15 pairs of secondary 

veins, uniformly spaced, broadly curving, curving 

sharply at margin, forming extensive series of small 

loops; intercostal tertiary veins approximately 

perpendicular to secondaries, closely spaced, typically branching midway between secondaries, one 

branch extending admedially and branching profusely; margin entire.

Discussion. The entire margin of the leaflet 

removes it from Allophylus wilsoni. In the intercostal tertiary venation particularly in the profusely branching veins that extend admedially the 

fossil is similar to the leaflets of Dracontomelon. The 

fossil particularly resembles D. mangiferum Bl., but 

leaflets of that extant species typically have fewer 

secondary veins that extend farther along the 

margin, slightly more closely spaced intercostal 

tertiary veins, and an oval shape. Scott (in Chandler, 

1964) has recorded fruits of this genus from the 

Clarno nut-bed assemblage, which is approximately 

coeval with the Comstock.

This species is named for the late Dr. Ethel I. 

Sanborn, in recognition of her contributions to the 

paleobotany of Oregon.
SYSTEMATIC REVISIONS OF OTHER PALEOGENE SPECIES 91

1 cm

FIGURE 41. Venation of Heteromeles and Salix. A, H. cuprovallis, holotype, UCMP 8963. B, H. arbutifolia, 

USGS Ref. Colln. 785. C, S. barclayi, USGS Ref. Colln. 904.

Holotype. UCMP 760.

Miquelia californica Wolfe, new species

Ficus densifolia auct. non Knowlton. MacGinitie, 1941, Carnegie 

Inst. Washington Pub. 543, p. 110, pi. 16, figs. 3-5.

Diagnosis. Leaves simple; venation pinnate; 

shape asymmetric, oval; base rounded, apex acute; 

five to seven pairs of secondary veins, departing at 

angles of 45°-80° (greater apically), looping sharply; 

intercostal tertiary mesh large, veins forking at right 

angles and thinning conspicuously into profusely 

branching fourth order mesh; margin entire.

Discussion. The characteristic third and fourth 

order mesh of the fossils allies them to Miquelia, a 

member of the tribe Phytocreneae of the Icacinaceae. 

M. californica is particularly similar to M. reticulata 

Merr. from the Philippine Islands. The most 

conspicuous difference between the two species is 

that all the secondary veins of this extant species 

depart at about an80° angle and form an intercostal 

area that is approximately trapezoidal.

Holotype. UCMP 2180.

Paratypes. UCMP 2179, 2181.

Sageretia laurinea (Lesquereux) Wolfe, new combination

Juglans laurinea Lesquereux, 1878, Harvard Museum Comp.

Zoology Mem., v. 6, no. 2, p. 35, pi. 9, fig. 11. 

Phyllites laurinea (Lesquereux) MacGinitie, 1941, Carnegie Inst.

Washington Pub. 543, p. 162, pi. 45, fig. 8.

Discussion. MacGinitie (1941) noted that the 

basal secondaries of these leaves were "pseudoprimaries," that is, they arose from the base of the

leaf. This feature together with the finely serrate 

margin and the intercostal tertiary veins being 

oriented obliquely to the secondary veins indicate 

that the fossils represent Sageretia. Comparisons 

can be made to species such as S. elegans (H.B.K.) 

Brongn.

Plafkeria obliquifolia (Chaney) Wolfe, new combination

Text figure 42 

Celtis obliquifolia Chaney, 1925, Carnegie Inst. Washington Pub.

349, p. 52, pi. 1, figs. 1, 3, 5. 

Viburnum thomae Chaney and Sanborn, 1933, Carnegie Inst.

Washington Pub. 439, p. 98, pi. 40, figs. 5, 6. 

Tilia williamsi Sanborn, 1947, Oregon State Monographs, Studies

in Geology, no. 4, p. 23, pi. 6, figs. 1-3. 

Alangium thomae (Chaney and Sanborn) Lakhanpal, 1958,

California Univ. Pubs. Geol. Sci., v. 35, p. 34, pi. 10, figs. 1,2. 

Discussion. The ultimate venation of these 

specimens indicates that they are not Alangium, 

because that genus has large areoles intruded by 

profusely branching veinlets. The leaves resemble 

Byttneriophyllum tiliaefolium (A. Br.) Knobl. and 

Kvac., particularly in shape and intercostal tertiary 

venation. The fourth order venation of Byttneria, 

however, is highly characteristic: the tertiary veins 

typically branch about halfway between the 

secondaries and send a fourth order branch almost 

perpendicularly but the branch curves rapidly to 

parallel the tertiary vein. This feature is well 

illustrated by Knobloch and Kvacek (1964, pi. 3, fig. 

2). In Plafkeria, the fourth order veins are numerous, 

branching, and extend between two adjacent
92 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

1 mm

FIGURE 42. Venation of Plafkeria, Byttneria, and Alangium, A, D, W. obliquifolia, hypotype, USNM 43431. B, 

E, B. aspera, USGS Ref. Colln. 560. C, F, A. chinense, 

USGS Ref. Colln. 1225.

secondaries (fig. 42). Sanborn's (1947) reference of 

her material to Tiliaceae, was, therefore, justified.

Camelia oregona (Chaney and Sanborn) Wolfe, 

new combination

Ilex oregona Chaney and Sanborn, 1933, Carnegie Inst. Washington Pub. 439, p. 80, pi. 22, figs. 2, 5, 6.

Discussion. Dr. Copeland McClintock of Yale 

University first pointed out to me that the Goshen 

material assigned to Ilex represented Camellia. This 

assignment is indicated by the irregularly shaped 

marginal areoles that are intruded admedially by 

profusely branching veinlets and by the rounded 

teeth that are separated by angular sinuses.

Doona? chaneyi (Potbury) Wolfe, new combination 

Text figure 43

Rhamnidium chaneyi Potbury, 1935 (part), Carnegie Inst. Washington Pub, 465, p. 75, pi. 11, figs. 1, 4, 5 only.

Discussion. These leaves have the intercostal 

venation typical of Dipterocarpaceae (p. 82). The 

similarity of Doona is in the almost ladder-like fifth 

order venation and the closely spaced intercostal

FIGURE 43. Venation of Dipterocarpaceae and Rhamnaceae. A, Parashorea 

cashmanensis, paratype, USNM 42130. 

B, P. malaanon, USGS Ref. Colln. 1663. 

C, Doona? chaneyi, syntype, UCMP 

933. D, D. ovalifolia, USGS Ref. Colln. 

1656. E, Rhamnidium elaeocarpa, USGS 

Ref. Colln. 1528.

tertiary veins (fig. 43). The reference to Doona is 

questioned because this genus is generally thought 

to have evolved on Ceylon and to have never reached 

mainland Asia. If so, D.? chaneyi presumably 

evolved from a common ancestor. One of Potbury's 

types is excluded from this species because it has 

more widely spaced intercostal tertiary veins and the 

areoles are almost twice as large as those of the three 

remaining elements.

Lectotype. UCMP 935.

Syntypes. UCMP 933, 934.

Parashorea chasmanensis (Wolfe) Wolfe, new combination

Text figure 43

Rhamnites cashmanensis Wolfe, 1968, U.S. Geol. Survey Prof. 

Paper 571, p. 23, pi. 6, fig. 2; text fig. 18.

Discussion. As noted previously (p. 82), this 

species has venation features characteristic of 

Dipterocarpaceae. In spacing of the intercostal 

tertiary veins, the fossils are most similar to 

Parashorea. Note (in fig. 43) that the ultimate 

venation pattern is clearly not rhamnaceous but is 

dipterocarpaceous. When I first described this 

species, I stated (Wolfe, 1968, p. 23): "I am not 

absolutely convinced that Rhamnites cashmanensis 

is a member of Rhamnaceae, but this familial 

reference is the best one that I can make at this time."
LOCALITY DATA 93

Alangium oregonensis Wolfe, new species

Celtis obliquifolia Chaney (non typic), 1927, Carnegie Inst. 

Washington Pub. 346, p. 115, pi. 13, fig. 12.

Diagnosis. Leaves simple; venation palmate; 

shape asymmetrical, ovate; base oblique and 

broadly rounded, apex acuminate; six primary veins, 

first lateral pair extending about one third to about 

half way towards apex, camptodrome; five to six 

pairs of medial secondary veins, looping sharply 

near margin; intercostal tertiary veins; intercostal 

tertiary veins spaced about 0.4 cm apart, perpendicular to secondary veins.

Discussion. These leaves have the ultimate 

venation of Alangium (see discussion of Willisia 

obliquifolia). Among extant species, the fossils most 

closely resemble in shape the leaves of A. chinense 

(Lour.) Rehd., but leaves of that species typically 

have fewer primary veins and more widely spaced 

intercostal tertiary veins.

Holotype.-UCMP 73.

LOCALITY DATA

3842. Grade Trail cabin, opening along west contact of coal with 

shale. Kushtaka Formation. Collector: Maddren, 1905. 

Middle Ravenian.

3846. From creek flowing into head of Canyon Creek from Mt. 

Chezum at elevation of 2,000 feet. Kushtaka Formation. 

Collector: Maddren, 1905. Lower Ravenian. (Reference to 

the topographic map published by Martin, 1908, indicates 

that the 2,000-foot contour on his map is well below the 

conspicuous break in slope, and this collection is thus 

from beds stratigraphically below localities 11163 and 

11164.

3847. From gulch on creek emptying into Berg Lake where Happy 

Hollow trail passes around shore. Kushtaka Formation. 

Collectors: Martin, Paige, and Maddren, 1905. Lower 

Ravenian.

3879. Yakutat Bay, west shore at Dalton's coal outcrop on Esker 

Stream. Kulthieth Formation. Collector: Tarr, 1905. 

Lower Ravenian.

9389. East slope Charlotte Ridge, 0.4 mile NE of NE cor. sec. 10, 

T. 17 S., R. 7 E., at approximate altitude of 1,000 feet. 

About 200 feet below base of Tokun Formation. Kushtaka 

Formation. Collector: F. F. Barnes, 1955. Middle Ravenian.

9551. Small stream 3.0 mile N. 45° E. of north end of Hanna Lake, 

near edge of moraine. Kulthieth Formation, probably 

upper part. Collector: Miller, 46AMr 43. Kummerian?

9552. Near top of Split Creek Sandstone Member, 6.11 miles S. 

761/2° W. from the point of intersection of the two principal 

tributaries of Burls Creek; lat 60°13'38" N., long 144°17' 

14" W. Katalla Formation. Collector: Rossman, 44ARm 1. 

Kummerian.

9553. Near USGS Cenozoic loc. 17831 (see Miller, 1961b); lat 60° 

18.5' N., long 142°47.0' W. Mollusks from this locality 

include Volsella aff. V. eugenensis (Clark), Tellina cf. 

lincolnensis Weaver, and Solena cf. clarki (Weaver and 

Palmer), as determined by H. E. Yokes (written communication, 1948). Yokes believes the mollusks to be of middle

9889.

9891.

9893.

9894.

9895.

1157.

11158.

11159.

11160.

11162.

11163.

11164.

11165.

11166.

11167.

11168.

11169.

11170.

11183.

11184.

11185.

Oligocene, i.e., Lincoln age. Upper part Kulthieth Formation. Collector: Miller, 47AMr 60. Kummerian. 

Sec. 10, T. 19 S., R. 6 E., lower part Katalla Formation. Collector: Shell Oil Co., 62H4 30. Kummerian. 

Sec. 10, T. 19 S., R. 6 E., lower part Katalla Formation. Collector: Shell Oil Co., 62H4 35. Kummerian. 

Sec. 30, T. 16 S., R. 9 E., in saddle just north of Doughton 

Peak. Kushtaka Formation. Collector: Shell Oil Co., 62H 

603 200. Kummerian. 

Sec. 19, T. 18 S., R. 16 E. Upper part Kulthieth Formation.

Collector: Shell Oil Co., 62H 1609 40. Kummerian. 

Sec. 19, T. 18 S., R. 16 E. Upper part Kulthieth Formation.

Collector: Shell Oil Co., 62H 1609 367. Kummerian. 

Sec. 25, T. 20 S., R. 18 E., Poul Creek Formation. Collector:

Shell Oil Co., 62J 266. Kummerian.

Lat 60°24.8' N., long 143°51.8' W. From bed of creek. Kushtaka Formation. Collector: Wolfe, 1968. Lower Ravenian. 

Lat 60°25.2' N., long 143°48.8' N. From west wall of 

canyon. Kushtaka Formation. Collectors: Wolfe and 

Larson, 1968. Lower Ravenian.

Lat 60°25.2' N., long 143°48.8' N. From west wall of 

canyon, about 200 feet north of loc. 11158. Kushtaka 

Formation. Collector: Gunn, 1968. Lower Ravenian. 

Lat 60°24.0' N., long 143°57.5' W. From slump area just 

south of creek. Kushtaka Formation. Collector: Wolfe, 

1968. Lower Ravenian.

Lat 60°23.9' N., long 143°57' W. From sandstone bed just 

beneath crest of ridge and about 800 feet higher than loc. 

11160. Kushtaka Formation. Collector: Wolfe, 1968. 

Lower Ravenian.

Lat 60°26.,5' N., long 143°57.4' W. From west side of small 

gully. Kushtaka Formation. Collectors: Wolfe, Larson, 

and Gunn, 1968. Middle Ravenian. 

Lat 60°26.7' N., long 143°57.5' W. Along east side of small 

gully. Kushtaka Formation. Collectors: Wolfe, Larson, 

and Gunn, 1968. Middle Ravenian.

Lat 60°25.5' N., long 144°7.7' W. Cliff face about 300 feet 

below contact with Tokun Formation. Kushtaka Formation. Collectors: Wolfe, Larson, and Gunn, August, 1968. 

Middle Ravenian.

Lat 60°28.5' N., long 143°37 W. At base of prominent cliff. 

Kushtaka Formation. Collectors: Wolfe, Larson, and 

Gunn, 1968. Lower Ravenian.

Lat 60°27.1' N., long 143°51' W. Just above prominent 

break in slope, from concretionary bed. Kushtaka 

Formation. Collectors: Wolfe and Larson, 1968. Upper 

Ravenian.

Lat 60°16.7' No., long 143°51,8' W. From southeast side of 

northeast tip of Nichiwak Mountain, about same altitude as top of Bering Glacier. Split Creek Sandstone 

member of the Katalla Formation. Collectors: Wolfe, 

Larson, and Gunn, 1968. Kummerian. 

Lat 60°27.5' N., long 143°48.8' N. On south side Carbon 

Ridge, about 50 feet below crest. Kushtaka Formation 

Collectors: Wolfe, Larson, and Gunn, 1968. Kummerian. 

Lat 60° 19.5' N., long 142°29.9' W. On south side of valley of 

creek. Kulthieth Formation. Collectors: Wolfe, Larson, 

and Gunn, 1968. Lower Ravenian.

Lat 60°6.1' N., long 139°11.5' W. North side of Haenke 

Glacier. Yakataga? Formation Collector: Plafker, 1968. 

Seldovian. 

Lat 60°9.3' N., long 140°20.5'W. Yakataga Formation.

Collector: Phillips Petroleum Co., 1967, Homerian. 

Lat 60°8.8' N., long 140°8.25' W. Poul Creek? Formation 

Collector: British Petroleum Co., GB 1215. Angoonian.
94 PALEOGENE FLORAS FROM THE GULF OF ALASKA REGION

11186. Southwest shore of Cenotaph Island in Lituya Bay; along 

strike of beds from a point 0.85 mile N. 84° W. of easternmost cape on the island to a point 0.68 mile S. 70° W. of 

the same cape. Unnamed beds of late Miocene age. 

Collector: Miller, 58AMr 333. Seldovian or Homerian.

11188. Unknown position in the Basin Creek Member of the 

Katalla Formation; 1.60 miles N. 11° W. from the point of 

intersection of the two principal tributaries of Burls 

Creek, lat 60°13'38" N., long 144°17'14" W. Collector: 

Rossman, 44ARm 27. Kummerian.

11189. The map furnished by British Petroleum has an area west 

of Hope Creek of about 10 mi2 circled and the notation 

that their locality GA3024 is "in this area." This area 

only has rocks that belong well up in the Katalla Formation. Because this locality produced both Macclintockia 

pugetensis and Platanus comstocki, the locality data are 

probably erroneous.

CAS 29181. Headwaters of Redwood Creek, 2,000 feet north of 

junction with east fork. Lower part Katalla Formation. 

Collector: Hanna, 1938. Kummerian.

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INDEX

[Italic page numbers indicate both major references and descriptions]

Page 

Acacia ............ .. ... ..... ......... .... 37

Acer.. ............. ... ... ....26,18,20,45,46,70

arcticum .... ......................................................... 16

glabroides . .... .. ... ....... ..... ... 14

(rubrum type) . . .......... ... .. ... 45

trilobatum productum ... ..... . . ...... 16

Aceraceae .. .............. .... . 14, 19, 43, 47

aceroides, Platanus . ... ... ............. ...... 71, 90

Acila .. .............. . . .... ............. . . ...... .... 6

decisa ... ..

shumardi 

Actinidia .. . . 

Actinidiaceae .

.... .. 6

6

.. 45 

. 27, 81

acutissima, Phytocrone ............... 9, 11, 27, 29, 30, 76;

pis. 12, 13 

Addicott, W. O., cited .................... .......................... 3, 5

Admiralty Island .......... .. . ................... . 14

post-Kummerian assemblages.. ................... .. 36

Ailanthus ... . ..... .. ... ..... ............ ........ . .. 45

Akebia ..... ........ . .... .. ..................... ..... 45

Alangiaceae .......... ............................ 27, 50, 84

Alangium.......... ...... 41, 44, 45, 46, 50, 84, 91, 93

bergensis ................. 9, 11, 27, 29, 30, 52, 84; pi. 16

chinense ... .................... ...... ...................84, 93

longiflorum ....... .. .................. ..... ...... ..... 84

oregonensis................... . . ......................... 93

thomae ...... .. ........................ ..... .... ............... 91

sp ......................................................... 11, 27, 84; pi. 16

Alaska, Fagopsis groenlandica .......................... 88

Paleocene rocks.................... ............................ 57

Alaska Peninsula ........ ................... ... ..... ............. 14

marine lithologic unit.... ......... ...... ............... 39

mollusks, marine .. ........... ............... ........ ... 40

Alaska Range ............. ................... ................... 14, 42

Angoonian rocks.. ...... ....... ...................... 46, 56

uplift .......... .. .................. ........ .................... 42

Alaska Range section .................... .. ..... ............ 20

Alaskan floras, ages ...... .. .......................... ....... . 1

alaskana, Clethra........................................................... 85

Dryopteris ..................... ......... . 10, 26, 30, 53; pi. 3

Euodia .................................... 11, 27, 29, 30, 73; pi. 9

Firmiana ................................................................. 14

Goweria ................................. 9, 11, 27, 30, 75; pi. 13

Melanorrhoea...................... 11, 27, 30, 52, 74; pi. 10

Salix ........................................................ ................. 18

Semecarpus......................................................... 52, 74

alaskanum, Artocarpidium .....................................52, 59

Asplenium.............................................................. 51, 53

Dryopteris .................................................................. 51

alaskanus, Stemonurus ................ 11, 27, 29, 78; pi. 13

Zizyphus .................................... ..9, 11, 27, 80; pi. 14

Alder ..................................................................... 57, 59, 86

Asian...................... ................................................... 58

leaves ........................................................................... 57

Allantodiopsis .............................................................. 12, 53

pugetensis .......... 9, 10, 12, 26, 30, 31, 53; pis. 2, 20

Allophylus............................................................................ 78

duktothensis........................ 11, 27, 29, 30, 78; pi. 14

peduncularis................................................................ 78

wilsoni .................................. 11, 34, 35, 78, 90; pi. 29

sp .......................................................... 11, 31, 79; pi. 24

Alnus................................l3, 16, 18, 34, 41, 44, 46, 57, 58

barnesi ..................................................... 16, 17, 20, 59

cappsi ..................................................................... 13, 20

(cappsi type)................................................................ 45

carpinoides .................................................................. 14

. ..... . .. ....... . ...... 58

10, 12, 32, 33, 34, 57, 58; pi. 26

......... . ......... 13, 14

....... . . .... ........... ......... 59

. ......... . .. ........ .. . .......... 59

Alnus Continued 

crispa ..... .. ..

cuprovallis . . ...

evidens . . .....

fauriei ... .

healyensis . .

incana..... .. .... . .... . .... .. . .. 20,21,58

japonica . . .... . .. ..... . ...... . . ... .. 58, 59

kefersteini . ..... . .. . ... ...... .. . . .... ..... .. . .... .. 16

martini ......... . ........ 10, 27, 30, 34, 58, 59; pis. 5, 21

nitida..... ...... .. ... . . .... ..... ..... . ....... ... .. . .............. 59

scotti .............. . ...............10, 27, 30, 52, 59, 86; pi. 4

sinuata . ....... .............. ... ... ....................... ...... ....... 59

sp ................. . .......... ........... 10, 32, 34, 59; pis. 26, 28

Altingia ....................... ..... .. ............... ......... . ... ......... 50

amblyrnhynca, Populus ........... . . .. ........ .. 17

Amelanchier ...................... ...... ................ ............... 71

Ampelopsis .... . . ........ ............. .. . .... . .... ..... .. 45

Amphi-transpacific distribution ....... . ...... ............. 51

Amphi-transpacific genera .. .................. ..... ........ 49, 50

Anacardiaceae..... ............ . ....... ............. .... 27, 31, 50, 74

genus ......................... .......................... ............. 11

genus and species indetermined ............ . 74; pi. 25

Anamirta ............... . ................................. 28,50,62

cocculus ....... . .. .... . .... ................ ... . ..... .. . . ... .. 62

milleri ........ . .. . .... . ............ 10, 27, 30, 62; pi. 6

Anemia. . . .... . .... .................. ....... ... ... .. .. ...... ... 52

elongata ............................ .. .... ......... ........................ 52

eocenica ... ... ...... ................ ............................ .. .... 9, 52

gunni... .......... .......... ........................9, 10, 26, 52; pi. 2

occidentalis ............ ...... ................................. ........... 52

sp .............................. ....... .................. 10, 34, 52; pi. 28

Angiospermae ... . .. . ....... .. ...................14, 26, 31, 32, 34

Angiosperms . .. . ......... ............ . . . .... ......... ....... ..... 43

trans-Pacific ...... ...... ...... . ................................... . 49

Angoonian assemblage, type ..... ....... ..... .............. ..... 54

Angoonian assemblages, Alaska ....... .. ... ...... .... ... 57

Oregon .... .. ... ... .... .............. ............................... ..... 46

Pacific Northwest. ................. .................................. 57

Angoonian Stage ................................ ........ ...... ...8, 13, 16

age ..................... ......... .. ... ......... .............. ............. 14

age equivalents .... .... ... ... ......................... ............... 8

Alaska ................ .... .... ................................. ............. 46

climate .............. ...... ....................... .. ........... ............. 36

definition ............ .... .......................... ........... ... ......... 14

flora...................... .. . .......................... ........... ............. 14

floral assemblage . . ...................... ........... ....... 14, 20

Oregon ................... . ...................................... ............. 46

temperate, cool.. . .. .......................... .. ........ ....... 36, 46

type section.... ...... . ............... . .. ....... ........ ........... . 14

Annonaceae ................... ................ ........... .. 27, 35, 50, 66

antipofi, Fagus ............. ...................................... ............. 16

apoensis, Engelhardtia... ................................... ............. 55

aquaticiformis, Planera ........................................... 51, 56

Aquifoliaceae ................. ...................................... ....... 33, 74

aquilina, Salix ...... ............................................... ............. 54

aragoensis, Whitneyella sinuata.................................... 4

Aralia taurinensis .............................................. ............. 89

arbutifolia, Heteromeles ........ ............................ ............. 90

Arbutus ............................................................... .. ....... 47, 48

arctica, Nyssa .................................................................... 18

Populus... ......................................................... ....... 17, 61

Trochodendroides ..................................... .. ............. 61

arcticum, Acer ............ ...... ............................... ...... ............ 16

Cercidiphyllum ............................................ ....... 17, 18

Arcto-Cretaceo-Tertiary Geoflora .................. ............. 43

Arctostaphylos .................................................................. 47

Arcto-tertiare Element .................................................... 39

Page 

Arcto-Tertiary concept, history .... . ................... 37

Arcto-Tertiary Geoflora ................ . ... ........ 43, 48

Arcto-Tertiary Geoflora concept . 1, 37, 38, 41, 43, 50

Chaney's ..... ......... . . .......................... 40

ariculaecordatus, Grewiopsis . . . ........... ..... .... 16

Artocarpidium .. .. .... ..... ...................... 41

alaskanum . ........ .......... ........ .... .... 52, 59

Artocarpoides......... ........................ ......... ... 86

kummerensis ........ ...................... ..... 31, 34, 86

Asia, Doona ......................................... .................... 92

Meliosma simplicifolia ...... ........................ . 79

southeast, Celastrus .............. . ..... ................. 75

aspera, Tilia . ..... ... .............. ............. .............. 14

Aspidaceae ..... ...... .............................. .... 26, 31, 53

Asplenium alaskanum ..... ....................... .......... 51, 53

magnum................................................................ 53

Astoria Formation, age .......................... ............. ... 19

Astoria, Oregon, marine lithologic unit................ 39

Atwood, W. W., quoted .... .... ...................................... 40

atwoodi, Vitis................................................................. 14

auriculata, Cocculus ............................. ...................... 63

Axelrod, D. I., quoted ... ........................................ 58

axelrodi, Prunus .. ................... 12, 32, 33, 71; pi. 26

B

balsamoides, Populus ......... ....................................52, 75

barclayi, Salix ........................... .................................. 90

barnesi, Alnus ........................................... 16, 17, 20, 59

Barringtonia..................................................... 28, 50, 83

duktothensis............................................................. 83

South Pacific .......................................................... 30

speciosa ......................... ........................................ 83

sp ................. ............................ 11, 27, 30, 83; pi. 17

Barringtoniaceae ....................................... 27, 28, 50, 83

Basin Creek Member, age ............................................. 13

early Blakeley ............................................................ 6

Katalla Formation ..................................................... 6

leaves ....................................................................... 6

molluscan fauna ........................................................ 6

plant assemblage .................................................... 13

Bear Lake Formation, 

Unga Conglomerate Member.......................... .... 20

bebbiana, Salix .................................................................. 90

bendirei, Cinnamomophyllum....................... .............. 89

Philadelphus .............................................................. 89

Platanus........................................................................ 90

Sassafras..................................................................... 89

(bendirei type), Carya ................................................ 44, 45

Platanus...................................................................... 44

Benzoin dilleri ................................................................... 76

Berchemia ...................................................................... 48, 82

Berg Lake .............................................................................. 4

fossil plant assemblages .................................... 4, 41

bergensis, Alangium .... 9, 11, 27, 29, 30, 52, 84; pi. 16

Bering River coal field .................................................... 40

Beringia.......................................................................... 49, 50

Betula ........................................... 17, 18, 35, 44, 46, 57, 60

(papyrifera type)........................................................ 45

stevensoni.................................................................. 57

sublutea ........................................................................ 17

sp ..........................................................10, 44, 60; pi. 28

Betulaceae........ 14, 19, 24, 27, 31, 32, 34, 43, 47, 57, 88

Bignoniaceae ..................................................................... 51

Bilyeu Creek assemblage............................................... 90

binervatum, Lygodium .................................................... 52

Biostratigraphy.................................................................... 7

blancoi, Phytocrene.......................................................... 77

99
100 INDEX

Blechnaceae .........................................................

Blepharistemma .............,...... ...... ...................

Boraginaceae .............................................

Boreoacala.............................. .,...,........., ..-.....-....

candoni oregonense ...,. . . 

Brachiodontes cowlitzensis ....... . ....

bracteata, Phytocrene ...... ...... ..

Brazil, Amazon lowland, leaf-margin

percentages . ....... ........ ... ..... - . ....

Bridge Creek flora, Pacific Northwest .. . .

radiometric age . .... ... ...... ....

British North Borneo, Parashorea

malaanonan............. - ..... .... , . 

British Petroleum Company, fossil plants 

Brown, Roland, cited... 

Brown, R. W., quoted ,.,..,.. 

browni, Eucommia .... ........ . ... .

Bruclarkia fulleri .. ...

Bruguiera .. ......

buchii, Cornus .... . ...

Burseraceae ....... -

Byttneria . .....

Byttneriophyllum tiliaefolium

Page 

. 34, 53 

....... 83

....... 51

......... 6

. . 6 

.. ..... 3

....... 77

,. 25 

... 54 

... 14

83

. 2

87

54

70

6

83

52

55

91

91

Caesalpinites .. .. . ...

sp

californianus, Pitar ... 

californica, Magnolia

Miquelia .

Zizyphus . 

Calkinsia ..... .

dilleri .

franklinensis

plafkeri .

pugetensis . .

ravenensis 

Calycacarpum 

Camelia oregona 

Camellia . .. 

Canadian shield area 

Cananga . ...

odorata

sp

Canarium ... 

Canavalia eocenica 

candel, Kandelia

................ .... 72

...... 11, 27, 30, 72; pis. 9, 25

... .... .. . 3

.... ... 68

... . . .... .. .... . . 91

. ..... ......... 61

... ... . 12, 32, 62, 89

... .... 12, 63, 89

....... . . 63, 89

10, 12, 27, 31, 62; pis. 6, 23 

...... ..12

, ... .... 12

... .44, 45, 46, 47, 49

... .. . . .... 92

... . ............ 92

..... ... 42

.... 50, 66

.. .. .. . .... 66

10, 27, 52, 66; pi. 8

55

.... . . 51

. . ....83; pi. 19

Canyon Creek, fossil plant assemblages .. ... 4

plant localities .... . . ..... 4

Capay stage... . ... . ...... . 8

cappsi, Alnus ... ... .... .... 13, 20

(cappsi type), Alnus . ..... 45

Carallia ... .... ... .. . 83

Carbon Mountain . ... ....... 4,6

carbonensis, Dicotylophyllum ... 11, 33, 34, 86; pi, 27 

Ilex . ..... .... . .... ... 11, 33, 74; pi. 26

Litseaphyllum .... 10, 34, 68; pi. 28

Salix .. ..... .... 10, 32, 55. pi. 26

Sorbus . .. 11, 32, 34, 71; pi. 27 

Tilia .. ... .. 11, 33, SO; pi. 27

Cardiocaryon ... .. .. ... 56, 88

(Cardiocaryon). Juglans . .. ..45,88

Carey's postulated orocline .... 36

carpinoides, Alnus ... .. ........ ... 14

Carpinus .... .... ..... 45, 57, 58

cobbi ... . . .. 13, 20

Carpolithes elytraeformis , . .. 52, 76 

Carya ... ... ... 44, 55

(bendirei type) . . .. . 44, 45

cashmanensis .. . 10, 12, 31, 55; pi. 20

Cascade Range, floras, Miocene .. 24

Cashman assemblage , -... - . - 9

cashmanensis, Carya .. 10, 12, 31, 55; pi. 20

Rhamnites.. ....... .... ,82,92

Cassipourea .. ....... . .... 83

Castanea ... .. .. .... ....... 47

Castanopsis ....... 60

Catalpa . ... . ...... ..... 45

Page 

Ceanothus .. ................ .......................................... 47, 48

idahoensis ...................... ........................................ 61

Cedrela ......... ...................... ............... .... 45, 46, 48

Celastraceae.................. .................... ........ 27, 31, 75

Celastrus ............... ... ................. ....... .. 41, 44, 45, 75

comparabilis. ... ...........9, 11, 27, 28, 30, 51, 52, 75;

pis. 11, 18 

conditionalis . ... .. ...... .............. ... .... .30

monospermoides ... ......... .... ..... .. .. 75

taurinensis ... ............... ....... 89

Celtis , ....... ........ .......... ............... . . 45

obliquifolia .... ..... .. .. ... .. .. ...... 91,93

Cercidiphyllaceae........... .... ...... .. .... 80

Cercidiphyllum .. ... 15, 17, 18, 43, 44, 45, 46, 48, 61

arcticum ............. .. .... .... ..... .... 17, 18

crenatum ...... .. ......... .13, 14, 15, 17, 61

elongatum , .......... ... , . ... .......... ... 61

piperoides ........... ... .... .. ... . . 61

Cercocarpus ... . ....... . ....... .... 47, 48, 71

Ceriops.. . ..... ........... . , .. .... . . ... 83

Ceylon, Doona ... ...... .... .. ..... .. ........ . . 92

lowland, leaf-margin percentages ... .. ....... 25

upland, leaf-margin percentages..... .... . . 25

Chalk Bluffs assemblage, California ... ............ 80

Chamaebatiaria . . .............. . ......... - -- - 71

Chancy, R. W., cited ...... ............ . .... - - 2

quoted .... . ,. . ...... ... - ... 16, 40, 54

chaneyi, Doona .... ...,. ..... .. .... 92

Prunus ..... ..... . .... ... .71

Rhamnidium .. .... . ....... . . 82,92

Chaney's proposed floral correlations . . 16 

Chaney's temperate floras ... ... ... ... 2

Charlotte Ridge, fossil plant assemblage . ......... 6, 8

mollusk assemblage ... ... .. 5

Charlotte Ridge locality .. 53 

chasmanensis, Parashorea .. ... 92

Chickaloon Formation, Matanuska Valley . 51 

China, forest, mixed mesophytic, leaf-margin 

percentages .... - - .25

Ilex micrococca ... .. . ... 75

Zizyphus yunnanensis . 80 

chinense, Alangium ... . 84, 93

chuitensis, Corylus ..... 58

Clamgulchian assemblages ... .20

cinnamomeum, Cinnamomum .... . . .. 52. 67

cinnamomeus, Phyllites . .. 52, 67 

Cinnamomum .. ... 2, 41, 45, 46, 48, 67, 89

cinnamomeum .... .... . 52, 67

dilleri . ... ... . 89

trtchophylla - ... . . 89

Cinnamomophyllum 15, 33, 41, 67, 89 

bendirei ...... , 89

eocernua . - - ... 15, 89 

kushtakensis .. ..... .... 10, 31, 67; pi. 23

latum ...... .... 10, 27, 30, 52, 67: pis. 8, 9

Cladrastis (japonica type) . ... .. . 44

Claiborne flora . . ... 57, 70

Clamgulchian assemblage .. .... .... 42, 53

Clarno assemblage, Oregon, upper, . 63 

Clarno Formation, John Day Gulch 

assemblage.. ... ... .. 33

Puget Group ..... .. ... . 33

West Branch Creek locality ... ... ... 56

Clarno nut bed ... ... 61, 63, 89, 90

Clarno nut bed assemblage, Oregon 35, 57 

Clerodendrum .... .. .. .. . 48, 85

nutans ...... . ... 85

sp ...... 11, 27, 30, 85; pis. 7, 17

Clethra . ... 44, 45, 85

alaskana . ... ... . .. 85

delavayi . ... .. 85

sp ... .. ..... 11, 27, 30, 85; pi. 18

Clethraceae . .. 27, 28, 31, 85 

Climate, Alaska . . .... 32

middle Ravenian . .... . , 32

paratropical, North America 50 

temperate .. . . 58

cool .... . 46

tropical, definition . ..... 24

Page 

Climate, tropical Continued

North America.... .. .. ....... ..... ....... 50

Climates, Paleogene, Alaskan ...... ................. 36

Climatic change, Oligocene ... ... ..... . .......... .... 50

Climatic summary .... ......... . ........ ...... .35

Climatic zonation.. ....... ............. ..... 41, 50

Climbing plants, relationship with climate ... 23 

Coal Bay, Alaska Peninsula ... ... .... 19

plant assemblages ..... . .. .... 20

coatsi, Lithocarpus ... ... .... .. - 87,88

Salix . ...... .... .87

cobbi, Carpinus . . . . . 13, 20

Cocculus.. . .. ... ... ... . 20, 44, 45, 48, 49, 63

Anamirta. ..... .. . ..... 62

auriculata . . . . ..... 63

flabella . . ........ 17, 44, 61, 63

trilobus .. ... ........ 63

sp ... .. 10, 27, 30, 32, 63; pis. 6, 26

Collawash, Oregon, Miocene . .... . . 29, 71

Collawash flora, Oregon . . 28, 46, 62 

Colubrina .. ..... . . . .. 47, 48

columbiana, Gari ....... 5

Sassafras . ........ .... .. 89

Turcicula ... .. .. . .......... 6

comparabilis, Celastrus ................ 9, 11, 27, 28, 30,

51, 52, 75; pis. 11, 18 

Comptonia ...... .... ............ ..... ................................. 45

Comstock flora, Oregon ... 12, 35, 63, 90 

comstocki, Mallotus . ... 71

Platanus . 11, 12, 34, 71; pi. 29 

concavum. Piper . . 51 

conditionalis, Celastrus .. .. .... . .. .. 30

Hampea . . .... .... - - ..51,65

Paratinomiscium ... . 10, 27, 30, 51, 65; pi. 7

condoni oregonense, Boreoscala .. ... .. ... .. 6

Coniferales 14, 26, 31, 32, 53 

Continental drift .. .... .... .. 36, 49, 50

Cook Inlet basin . ................ ... . .. 1, 14, 16, 46

latitude . . ... ............................. ...... 19

Cook Inlet Region, Paleocene assemblages .... . 18

coosensis, Whitneyella .... .................... ........ ........ .5

Copper Basin assemblage............. ...............19, 33, 90

Nevada. . ........... .................................. 19

Copper Basin floras, radiometric age ... ............. 12

Copper River basin .. .... .................. .................... 14

Cordova, Alaska, climate ,. .............. . ............. 36

Cornaceae .................................... . ..... .,27, 50, 85

Cornus .... .................... ............. .... .41, 45, 85

buchii .. ...... .................. ................ 52

hyperborea ..... ........................................ 52, 82

irregularis ... ......... . ... ........ ............ 51, 52, 85

Corylites.. . ... ........ ...... ...... ..................... 57, 58

Corylopsiphyllum .... .......... ...................... .. .. 70

Corylopsis ....... .... ..... ....... ... ... 33, 44, 45, 47, 70

sp .. ... .......... .......... . ..... ..11, 32, 70; pi. 26

Corylus . . .......... ....................17, 57, 58

chuitensis ... .,......... ............................. ... 58

harrimani . . .....,................................... 58

macquarryi.......... .......... .... .......................... 89

cowlitzensis, Brachiodontes .......................... ...............3

Ficopsis ........ ..... ........................................... 5, 1

crassifolia, Euodia ................. ... .................. ........ 73

Luvunga .. . ............, ......... .. ... ... .. 74

crassum. Rhododendron.......... ...... ............ .29, 51, 86

Crataegus . ... .... ... ...................................... . 71, 90

cuprovallis. .. ............ .... .. .. ................ 90

crenata, Grewia . ........ ........,.... . .... 61

crenatum, Cercidiphyllum . ..... 13, 14, 15, 17, 61

crispa, Alnus .......... ............................................ .... 58

Cryptocarya ............ ......... ......... ............ ......34, 67, 68

infectoria .......... .......... ........... ............................. 69

presamarensis ........ .... ..... .. ....... 51, 68

cuprovallis, Alnus ... 10, 12, 32, 33, 34, 57, 58; pi. 26 

Crataegus ....... .................... ........................... 90

Heteromeles .... ....... .......... .... 90

Salix . ..... ..... ... ...........- ...... ....... -- - 90

Cyathea . ... ........... .. .. ...... ......... ... . 53

inequilateralis ... ..8, 10, 26, 30, 51, 53; pi. 3

pinnata .. .. ......... . ...................... . 53

Cyatheaceae . ... .... . ...... ... .... ................... 26, 53

Cycads ...,...... ... .... .. ..................... ................... 41
INDEX 101

Page

D

Daphnogene kanii ... ........................................... ... 52

Kanii ....... ..................................................... 38

daphnogenoides, Populus ......................... ............... 61

Dasymaschalon ................................................... . .. 66

trichophorum.......... .. ..................................... .. 67

sp ..... ... .. .............. . ............ 10, 27, 66; pi. 8

decheni, Rhamnus ............... ...... ......... .......... . 51

decisa, Acila . ....... .... . ...... ..................... . .... .....6

deflexiloba, Quercus ............................................... 19

delavayi, Clethra .. ................................................ .. 85

densifolia, Ficus .................................................... .. 91

Diatoms, marine.. .....,.................................. ... 20

Dicotyledonae .... ............................................ 14, 26

Dicotyledones.. ........ .. ..................... 31, 32, 34, 37

Dicotyledonous genera ...................................... .... 44

Dicotyledonous species, Alaskan, 

Lower Ravenian ............. .......................... ..... 28

drip-tips ... ........... ..... ..................... ..... .. 29

Dicotyhphyllum . .......................... ...33, 44, 86

carbonensis . .. .... ........ 11, 33, 34, 56; pi. 27

flexuosa .. . ................. ............... 18, 44, 51, 86

kummerensis ......... . ....... 11, 12, 86; pis. 25, 30

richardsoni. ...... ............ .... ... 16, 17, 44, 61

spp .. .... ........ . .. 86; pis. 17, 18, 24, 27, 30

dilleri. Benzoin . .. .. ....................................... 76

Calkmsia...... .. .................. ... ........12, 63, 89

Cinnamomum ..... ... ....................................... ... 89

Goweria ... .... .... .. 8, 9, U, 27, 30, 52, 76; pi. 11

Hyperbaena ... .................... ............... ..... 62, 89

dimorpha, Populus .. .. ................. .. .................... 54

Diospyros. ...... .... ... .... ............ ............. 45,48

orientalis ..... .. .......... ........................ .... .... 89

Diploclisia . ........... ...... ........................... .. 50, 63

sp - - . . . ..... ..... . ....... . 10, 27, 30, 63; pi. 6

Dipterocarpaceae ......, ........27, 31, 33, 35, 50, 82, 92

Dipterocarps, Paleogene occurrences . ................. 82

Dipteronia ... ... ... ...... . ............... ..... .. 46, 47

Disco Island, Greenland ..... ......................... ...... 39

disputabilis, Piper .............................. ............. . ..... 51

dissecta, Platanus ......... .......... ....... ................... 19

(dissecta type), Platanus ........ . ........... .... 46

Domengine stage ....... .......... ........ ..8

Doona ..... . . ..... ... . . . .. ......... 92

chaneyi. . . ...... . . ........ 92

Dorofeyev, P. I., cited ..... ..... ...... 43

Dracontomelon .. .... .. . ... ...... 50, 90

mangiferum . . .......... .... . .... 90

sanbornae . . ... ......... 90

Dryophyllum ...... ... ... ..... ..... 60

pugetensis .... ...... 10, 12, 31, 60; pi. 21

stanleyanum . . .... .... ...... 52, 74

tennesseensis. .... ... ... .... ...... ... .. 60

Dryopteris .. . ...... . .... ... , 41, 53

alaskana . ........... . . 10, 26, 30, 53; pi. 3

alaskanum .. .. . ........ .... ..... 51

gibbsi ... .... . .... ... ... . .53

Duchesnea. . .. .... .. ....... .... ....... 46

Duktoth River ... ... .. . . 6,9

Duktoth River assemblage, age .. ..... .... .. . 9

duktothensis, Allophylus ........ 11, 27, 29, 30, 78; pi. 14

Barringtonia...... . .............................................. 83

Meliosma . .. ............. ........ . 11, 27, 30, 79; pi. 14

E

East Indies, leaf-margin percentages .. .... ... 25

Ehretia ... . .. . ..... 45

elegans, Sageretia . ... ...... ,. . .. 91

elongata. Anemia .... .. . .. ........... ... . 52

elongatum, Cercidiphyllum ... ...... ... ... 61

elytraeformis, Carpolithes... . . ,. .... 52, 76

Paleophytocrene .... . 11, 27, 30, 52, 76; pi, 13

Endiandra . .. . . ................ 67

Enemtensky assemblage . ..... ..... ... 42

Enemtensky Formation, Kamchatka .... ..., . 42

Engelhardtia .... .. . ......... ....... 55

apoensis ... .... .... ...... ... ....... .. 55

(Engelhardtia) . . ..... .... ... 55, 56

(Oreamunoa). .... - ... ... .... ...... .55

(Engelhardtia), Engelhardtia ............................... 55, 56

Engelhardtia sp ........... .....10, 31, 32, 46, 47, 55; pi. 20

Engler, Adolph, quoted................................................ 39

Eocene-Oligocene boundary, Europe............................8

foraminiferal .. .........................................................8

mammalian ........... ............ .................................8

marine molluscan.,.... ......... ...... .........................8

eocenica, Anemia ......... . ............. . ......... .......... 9, 52

Canavalia . .... . ........ . ............................... 51

eocernua, Cinnamomophyllum ..... ... ........ . 15, 89

Octoea. ............ ...... ..... .. ...... ..................... 89

Eohypserpa. ...... .... ... ........................ 50,63

seeds ,. ..... ..... . . .......... .... .. ............ 63

Eometrix martini ... .... . .......... ........ ... ... ......,. 4

Ericaceae ... .... ....... ......... ........ 43, 46, 47, 49

Ericales .... .... ..... ....... .....................27, 85

Erythropalum .... ... .... ... . .... .... ..... .... 50

Esker Stream, coal-bearing rocks.. ......... ..........7

plant species..... ........... ...... ..... . .. ... . .9

Eucommia ... ...... .. .... ....... ......... 32, 44, 47, 70

seeds ... ..... . .......... ......... ..... ... 70

browni... .. .. ... ...... ... ..... ...... 70

montana ...... ......... .... .. ..... .. . ... .... 70

sp ... ............ .......... .... . 11, 31, 70; pi. 24

Eucommiaceae.... .. .... ... .... ........ 31, 70

Eugenia .... ..... . .... .. ........ .... ..... ... 84

jambos ... . ......... ........ .. ...... 85

multiflora . ..... .. .... ..... .... 85

sp .............. ............ 11, 27, 30, 33, 34, 84; pis. 16, 27

Euodia .............................................. 28, 44, 45, 48, 50, 73

alaskana ................................ 11, 27, 29, 30, 73; pi. 9

crassifolia ...... .. .... .. . ....... ..... 73

Euonymus nevadensis .. ... ...., .. ........... .. .... 71

Euphorbiaceae . ........ .. ... ..... .. 71

Euphoria ..... ... ... ........... .... .. .. 23

Euptelea ... . .. ... . ....... ... 43, 44, 45, 46, 47, 89

orientalis .... .... .. .......... . .. 89

pleiosperma .... ,,.... .. .. .. . .... . ... 89

Eurasia, flora, Eocene,... ... ..... ..... .. . 28

europaeus, Glyptostrobus .. . ...... ... ... . 18, 52

Eurya . .. ... ......... .... ... ... 45

evidens, Alnus .. .. ... ... .... .......... 13, 14

Exbucklandia.. .. . . ........ ... 45,46,48

exigua, Salix .. ....... . .... .... ... . 88

Fagaceae ................................ 14,31, 32, 34, 43, 46, 60, 88

pollen ..... ....,.............................,.....,...,..._........,.,.......,. 88

Fagales.. . . ..... . ... .... 14, 27, 31, 32, 34, 57, 88

Fagopsis........ ....... ....... ............................. 44, 46, 88

leaves .. ...... ...... ... . .............................. 88

groenlandica..- ........... . . ................... ........ 44, 88

longifolia ...... . ... ..... ................................... 88

Fagus ... . .... .... 13, 15, 20, 35, 44, 45, 47, 60, 88, 89

antipofi. ... .. .... ... . ........ .... ............... 16

oregona ... .... ............ ........... ... ..... . 15, 88

pacifica ..... ............ .. .. ........... ............. 88, 89

sp .................................. ... 10, 14, 34, 44, 60; pi. 28

fauriei, Alnus..... .. .... . . ..... .... ... ...... 59

Ferns, tree . .............. ....................... 31

Ficopsis cowlitzensis.... ................ ................ 5, 7

Ficus densifolia .. ......... ............ .... .... ... 91

sordida .... ..... ............ ...................................... 77

Filicales ... . ...... .... .. .., .. .. 26, 31, 34, 52

Filkineae... .... ....... . .... ... 26,31,34

Firmiana alaskana ........... .......... ...... .... .14

flabella, Cocculus . ... ..... ................... 17, 44, 61, 63

flexuosa, Dicotylophyllum. ......... . - 18, 44, 51, 86

Flora, Chickaloon..................... ............. ............... 41

Claiborne ....... ............ ... .... ...... - ....... ... 57

Fort Union . .......... .... ............. .... ..... 39, 40

Kazakhstan .... .... - ... .... . ........................ .. 43

Kenai,.... ..................... ......................... 41

Kobuk River ...... . . ... . ..... .... ........ 41

Madro-Tertiary .. ...... ........ ........ ,. - 41

Sezanne ...... ... ..... ........... ... ............ .... 38

Simonova , ....... ................... ...... ... 41

Thanet, Gelinden ... ... ........... ..... 38

Upper Dui, Sakhalin ..... .... ........... ...... ....... 41

Floral lineages, tropical, distribution ........................ 49

tropical, migration.................................................... 49

Floral migration route, Southern Hemisphere ..... 50

Floral migration routes, tropical, Paleogene .......... 49

Floral specimens, Hollick's............................................ 51

Floras, Alaska Peninsula .............................................. 40

Alaskan, Eocene ...................................................... 39

Kummerian.......................................................... 34

Oligocene....................... .................................. 46

Paleogene ........................................................... 41

Paleotropical ..................................................... 28

Ravenian, middle............................................. 31

upper........................................................... 32, 33

Tertiary .............................................. 39, 40, 41

Antrim ...................................................................... 38

Arctic, Greenland..................................................... 40

Miocene ............ ..........................................39, 40

Spitzbergen..................................................... 40

Tertiary ......................................................40, 41

Atanekerdluk ........................ ............................38, 39

Berg Lake ................ .................... .................2, 40, 41

Bridge Creek..................................................... 40, 41

Eocene ................................................................. 1, 41

Europe .............................................................. 38

Gulf of Alaska.................................................... 41

Gulf of Alaska Coastal Province ....................... 51

herbarium specimens ........................................ 51

Icelandic assemblages ........................................ 39

Indomalaysian region........................................... 32

Latah, Washington ....... ............................... 46, 80

latitude, high ...........,..................-...... .-.--..-.-.. 41

leaf..................... .... .......................................... 51

Mull ............. ........................................................... 38

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

other Alaskan, correlation ................................ 16

Pacific Northwest, Eocene .... ......................... 44

Oligocene............ ........................................ 44

Paleocene................. ............ .. ..................... . 44

pollen, Cretaceous, Late....... ................ ........... 44

Ravenian, late ........................... ......................... 45

Ruby . ............ . ........ -- ................................. 71

seed ................................................................................ 51

Siberia .......................................................................... 41

subtropical .................................................................. 41

temperate, Neogene ........... ................................. 24

tropical, America ... . ........ - ... .............. ......... 50

Asia, southeastern ............ ....... .................. 50

United States, western conterminous .......... . 28

Florida Keys, leaf-margin percentages................... 25

Florissant assemblage, Colorado ................. 46, 71, 88

Floristic analysis ..... ..... ......... - -........... 20, 26, 31, 32

Floristic zonation ......... ..... ................................... 41

Foliar physiognomy .... . ................... ... ......... 23

relationship with climate............................ 23, 24

Foraminifera .....................,.......... .......... ... ...... - 20

Eocene ....... ................ ....................... 16

Mclntosh Formation ........ ........ .................... 8

Forest, Boreal. . . .... .... ................................. 34

Complex Notophyll Vine .... ........ .......... ..... 29

conifer-sclerophyllous, broad-leaved,

Nevada................................................................ 21

coniferous . ...................... 19, 21, 26, 34

boreal .... .......... .. .......................... ... 21

coastal .................... . ............................. 40

Paleogene ... ... ........................ ................ 50

Cordova . .... ... ..................... .. ....... .... 40

deciduous, broad-leaved ........ 19, 20, 21, 43, 45, 46

broad-leaved, eastern Asia ....................... 33

Pacific Northwest .......................... ... ... 34

Paleogene ............... .......................... ...... 50

temperate,.... ............. ........ ..... ..... 42

temperate, warm .............. . ............. 26

dipterocarp, culled, Philippines ......... .......... 29

virgin, Philippines ... .................... .... ... 29

evergreen, broad-leaved ................. 26, 36, 37, 42

broad-leaved, China ... ..... .. .... 56, 89, 90

notophyllous..... ........ ... ..... ......... 32

hardwood, mixed northern, Alaskan ............... 34

Hong Kong ............ ........... . ..... .. 24

lowland... . ........ . ..... .................. .. . 40
102 INDEX

Page

Forest Continued

midmountain, Philippines.................................... 29

Mixed Mesophytic .... 23, 28, 33, 46, 48, 56, 63, 90

Asia...................................................... 34, 47, 48

east central.................................................. 26

eastern ......................................................... 48

Eurasia, central.............................. .. ....... 43, 49

Miocene ........................... ..... .................. 47

North America..... .......... . .. .... ........45, 48

eastern .............................................. 48, 49

western............... ........... ....... 48, 49, 58

origins................... . ...... . .......... . . 43

Pacific Northwest.................. ........... ...... 47, 56

theory ......................... ... .... ........ ......... 43

montane.......... ....................... ............. 25

monsoon................... ...... . ..... ............ ........ 29

mossy, Philippines ............ .. ..... ...... ..... . 29

notophyllous...... ....... ....... . 32

oak-laurel.......... .. ... ..... . . .... 25

China ..... ... ........ ......... .. ... 26

eastern Asia..... ...... ...... ..... ... .. .. 32

Taiwan ..... ............... .. ....... .......... ... 26

Pacific Northwest........................ ........ ... 40

Paleocene........................................ ...... .... . ...... 44

Platanaceae, Cretaceous ........ ...... ....... .... . 42

semi-deciduous....................... ........... ...... ... .... 25

Siberia ................................ .. .................... 42

Simple Mesophyll Vine ...... .. . ........... . .... 29

subtropical ................. .....25, 26, 28, 35, 44, 45, 60

eastern Asia......... .... .. ........ .............. 32

Eocene ........................ . ....... ...... .... 33

Kummerian Stage, Alaskan ...... . 34

lowland................ .. ..... .... .... .. . 71

Paleogene .... .... . . ...... .. 47, 48

Taiwan......... .... ............... .............. ... 24

temperate.... ........ ..... ...... ..... 23, 35, 42, 61

mesic..... ........... .... . ........... 33

Nevada..................... ... ..... ... .... . . 19

upland ...... . . .......... . 71

warm........ ..... ............ 25, 26, 45, 49, 60, 61

Mixed Mesophytic ... ..................... .28

tropical...... ... ............ ......... . . . ....... ....... 50

Hawaii ... .......... .... .... ... ..... ... 25

monsoonal region............. ...... ........ 25

North America, eastern ..... ....... 49

Pacific Northwest....... . .... .... ..... . .. 45

Paleogene ............. ... .............. 45, 47, 48, 49

Eurasia.... ....... .... ... . .................... ... 49

North America, eastern . .... ...... ... .49

North America, western . .. ................. 49

Queensland.. . ... ........... ...... 25

upland ... .. . . ...... .............. ..... ..... 61

warm temperate, Pacific Northwest ............. . 19

Forest formations, Australia, eastern ..... .... . 28

Webb's subdivisions ..... ... ... ...... ............. 28

Fort Union formation........... .. ...................... 40, 61

Fort Union Group ............... ...... ................ 39

Fortunearia ..... ...... ... ......... 45, 47, 70, 89

weedi......... ...................... . . . ........... ....... 89

Fossil, Collawash species .... .. ....... .... . 62

Goshen species .......... ............... ... . 62

Katalla species . ......... . ... ...... 62

Fossil plants, Katalla district ......... ...... . . . 2, 16

Malaspina district .... .... ....... ...... ......... ... 2

Fossil record, Neogene ....... . .................. ... ... 48

Fossils, Chalk bluffs .. ........ . .... .... .. 61

leaves ........... ............................................. 51

preservation .............. ......... . ... 51

Fothergilla . ....... .... ... .... ..... ..... . . 44, 70

sp .. . . ......... ........ ..... . . ..... .... 14

foveolata, Paleophytocrene........... .......... .... 76

Fragraria .......... ... . . .... ... ...... . 46

franklinensis, Calkinsia. ........... . .......... 63, 89

Rhamnites. ..... ...................... ....... .... 82

Franklinian Stage, age equivalent. ...... ..... 8

Fraxinus ............... ......... ... . ..77,18,41

Port Moller specimen ..... .. 17, 18

herendeenensis . . .. ........ ...... . . ...... 17

lateralis .... .... .... .... ... ...... ............. 52, 56

Page

Frederika Formation................................ .................. 42

Fruit assemblage, Siberia .................... ........... ...... . 42

fulleri, Bruclarkia......................................... .......... ..... 6

Fultonian Stage, age equivalent . ... ..................... ....8

fungi, Zizyphus ......................................... .................. 80

furuhjelmi, Quercus ................................................ 16

G

Gardner, J. S., quoted...... .... ...... ........... ..... ....... 38

Gari columbiana ... ............. ... .. ..... ..... .... .5

Garrya ..... ... . ..... .... ....... .... .......... .. 47, 48

Gem-Harris horizon.... ...... ................ ..... ... ..... .....9

geniculata, Ilex ............ ........... .... ...... ..... ..... 75

Geoflora, concept.... ........... .. ..... ..... .... .. . .... 43

Geraniales.. ........... ... ..... ..... . 27,72

gibbsi, Dryopteris .. ..... .......... ..... ........... .... 53

Ginkgo . ... ... ....... .... . ......... ..... .... 18, 41, 48

Cook Inlet Region..... ...... ... .. . ... 18

Port Moller assemblages ........................................ 18

Seldovian assemblages............................................ 18

sp . ... ............ ..... ........ ............. ....... . 14

Ginkgoaceae ..... ...... .. ........... ..................... ... 14

Ginkgoales ....... ......... ..... .... ........... .... ......... 14

Girroniera ........... ...... ...... ......... ...... 60

sp ..... ................................. .....10, 27, 29, 60; pi. 5

glabroides, Acer .. ... ..... .... ...... ..... ... ... 14

glandulifera, Populus ... .... ..... .... ..... ... ... ..... 61

glauca, Knema... ...... .... ........ .... ........... 66

glyptostroboides, Metasequoia . 13, 14, 18 

Glyptostrobus... . ................. ......... ..... 18, 33, 53

Chignik Bay ................ ............ ................... 18

Cook Inlet region .......... ...... . ..... ...... . 18

Port Moller region ..... ....... .... ..... .... .. 18

Seldovian species ............. . ............... . ....... 18

europaeus.. .. ...... . ...... ........... ..18,52

nordenskioldi.... ............... ........ ..... ...... 18, 53

sp ............ 10, 13, 26, 30, 31, 32, 52, 53; pi. 3, 20, 21

Good, Ronald, quoted ...... .......... ... 37, 38

Gordonia . ........... . ............................. 45, 48, 49

Goshen flora. ...... ..... ..... .... ..... 15, 16, 78, 89, 90, 92

Goweria ... .. ......... . ...... ..... ...... 75

alaskana ..... ....... ............ 9, 11, 27, 30, 75; pi. 13

dilleri. . .... ....... 8, 9, 11, 27, 30, 52, 76; pi. 11

linearis .... .. . ......... ..................... .. ........... 75

Gray, Asa, quoted ... ... .... ... ...... .......... ... .. 38

Great Britain, Paleocene rocks .... ...... ... 57

Green River Canyon section, Puget Group ......... ....9

Greenland, Fagopsis groenlandica .... ... ....... . 88

Paleocene rocks. ... ....... ................ ...... 57

palms, Tertiary .. . ................ .......... 50

Grewia crenata ...... .. ....... .......... ..... 61

Grewiopsis. ..... ... ................... . . 44

ariculaecordatus .... . . ..................... 16

groenlandica, Fagopsis ............. . ...... 44, 88

Quercus ....... .................... ...... ... ..... 88

groenlandicus, Quercophyllum ....... ........ ............ 88

Gulf of Alaska . .......................... ... .... .....1

Eocene rocks. ... ... ............... ....... .37

floras. .. .............. ..... .............. .. .... .1

Eocene .......... ................ . . ... 36, 41

fossil plants .. .............................. ... ...1,2

latitude ...... ........... ........ ...... .... 19

Paleogene floras .. ... ...... ...............1,2,13

Gulf of Alaska Coastal Province, floras .. ...... 51, 86

Gulf of Alaska region, Hollick's name-bearing 

specimens .... ............ .... . ....... ... ........ 51

Kummerian assemblages..... ..... .. .... 12, 26

Ravenian assemblage ........ ..... . ... 26

Gulf of Alaska section, leaves, Neogene age .... 13

marine mollusks ... ... ... ...... ..... .. 13

post-Kummerian age . .. ........... .13

Gunn, Donald, cited . . .. ..... ... .2

gunni, Anemia ......... ... 9, 10, 26, 52; pi. 2

Gymnospermae .. .... .... .... . 14, 26, 31, 32

H

Hainan, lowland, leaf-margin percentages.............. 25

Pyrenacantha volubilis .......................................... 78

Page

Hainan Continued

upland, leaf-margin percentages.......................... 25

Zizyphus fungi.... ...................... ............................ 80

Halesia.. ................... ................................................. 45

Hamamelidaceae ..... 14, 31, 32, 43, 45, 46, 47, 50, 70

Hamamelis ...... ...... ..... ............ ....................... 44, 70

Hamamelites ..... .................................................. 70

Hamilton Bay assemblage .. . . ......... ..................... 84

Hampea .... .......... ........... .. ............................. 65

conditionalis...... .. .... ......... ...... ............... 51, 65

Hanna Lake .............. ... ..... ..... ............................ 7

harneyensis, Holodiscus .. .. ........... ......... ..... 90

Prunus ................. ............ ... ............... 71

harpyiarum, Tetrapteris.... .. . . ... .. .... .. ...... 51

harrimani, Corylus ............. ......... . ................... 58

Hawaii, lowland, leaf-margin percentages .......... 25

upland, leaf-margin percentages......... .... ...... 25

healyensis, Alnus ........ ............................................... 59

Hemitelia... ................ .................................. ........... 31

pinnata.... ......... ............... ............. .......... 8, 53

Herendeen Bay, Alaska Peninsula ... .. ............. 20

Herendeen Bay assemblage .............. .. ......... . 20

herendeenensis, Fraxinus ......................................... 17

hesperia, Salix......................................... . ............... 19

Heteromeles ................. ......... ...... ........... ... ..... 47

arbutifolia ........ ...... .... ..... ............................ 90

cuprovallis............... .. .......... .......................... 90

Hollick, Arthur, quoted.......... ................... .. 2, 16, 40

Holmskioldia .. ... ......... .............. ................. 34, 85

speiri ................................................... 11, 34, 85; pi. 30

Holodiscus. ..... . . ..... .......... .......... ...... 47,71,90

harneyensis . .... .. ........ ... ..... ........... ...... 90

Homerian assemblages . ... .............. ..... .. . .... 20

Homerian Stage, age .... ........... ... ...... ................ 20

type section. ................... ...................... .............. 20

Homotaxis .................... ..... . .......... ... .. 1, 38, 39, 40

Hong Kong, leaf-margin percentages ........ ............. 25

Humble Oil Company.................... .............. .... ..... 2

Huxley, Thomas, quoted ...................... .................. 38

Hydrangea .... ............. .......... ............... .. ........... 44

Hyperbaena . .... ............... . ................ .................. 63

leaves ........ ....... ................................................. 63

dilleri. ........................................ ..... ........... 62, 89

mexicana ... .. ............... ................................. 89

hyperborea, Cornus ..... .... .................... ......... 52, 82

Hypserpa .... ...... .... .. ...................... 34, 50, 63, 64

nitida .. .............................. ......... .................... 63

sp ............ . ...... ... . ............. 10, 34, 35, 64

I

Icacinaceae ... ...... 19, 27, 31, 32, 35, 50, 56, 71, 75, 91

leaves ..................... . ......................... .. . ......... 9

idahoensis, Ceanothus .................................................... 61

Idesia .............................. . ....... ............................. 48

Ilex......... .. ........................ ...... ......... 45, 46, 74, 92

carbonensis .................................. 11, 33, 74; pi. 26

geniculata ..... ................... .......... ............. 75

micrococca. ...... ....... . ...................................... 75

oregona...... ......... . ..................... . . ............ 92

Illiciaceae ........ ..... ................. ....................27, 62

Illicium.. ......................... .. .................. .................. 9, 62

lanceolatum ................. ... .......................... ... 62

sp ................. ...... ................ 10, 27, 62; pi. 15

inaequilateralis, Pteris ... ..... ............. ....... 51, 53

incana, Alnus......................... ......... ...... ... 20, 21, 58

Indonesia, Stemonurus scorpiodes ........... ............. 78

inequilateralis, Cyathea... .8, 10, 26, 30, 51, 53; pi. 3

inermis, Zizyphus............................ ............... ........... 80

infectoria, Cryptocarya .............................................. 69

inopinum, Mohrodendron . ... ................................... 51

lodes ................................................................................ 50

Iltis, Hugh H., cited.......................................... ............... 3

Incertae sedis.................................................................. 86

Indonesia, Saurauia reinwardtiana .......................... 81

Introduction .......... ......................................... ...1, 7, 21

irregularis, Cornus..... ................................... 51, 52, 85

Mastixia . . ................ ... 11, 27, 52, 85; pi. 15
INDEX 103

Page

jambos, Eugenia ...... ....................................................... 85

Japan, Ilex geniculata ..................... ............................. 75

Japan Current.............................................................. 42

japonica, Alnus........ ................................................. 58, 59

(japonica type), Cladrastis ...............'.......................... 44

John Day Gulch assemblage ................... .. 33, 35, 71

Oregon ................................................................... 57

Joseph Creek, British Columbia, upper 

Ravenian assemblage ............................. .......... 35

Juglandaceae.... 14, 26, 31, 32, 43, 46, 47, 55, 83, 85

Juglandales .. .............. .................14, 26, 31, 32, 55

Juglans..... ... . ................ 7, 18, 33, 41, 44, 56, 88

laurinea ...................................................................... 91

magnified .................................................................. 14

nigella.... ..................................................................... 18

orientalis ..................................................................... 88

thermalis .. ....... ..... ..... ...... ... 52, 76

(Cardiocaryon) ..... ... .... .... ..... .. 45, 88

sp ..... ..... ....... . ... ............... .. . ..... 32

sp . .... .... ..................... ... .... .... 10, 56; pi. 26

K

kachemakensis, Salix ........ ........... 18

Kamchatka ... .... . .. ......... ................. ..... 20

Kandelia .......... ............. ... ...... . . .................... 83

Asia, southeast ........................................................ 30

candel....................... .......................................83; pi. 19

China ......................................................................... 30

wangi ............................................... 11, 27, S3; pi. 19

kanii, Daphnogene ........................................................... 52

Kanii, Daphnogene ..... ........... ...................... ..... 38

Karwinskia .......... ................ ........... ..............47, 48, 82

Katalla district .......................................... ....................3

orocline..................................................... ..... 36

Tertiary rocks ................................... ................ 36

Katalla Formation.............. ................................ ..6

Basin Creek Member................... ............. .... 6, 86

fossil assemblages .................................................... 2

Katalla district ................... ......................... ..3

Kummerian assemblages............ ... ............... 12

Nichiwak Mountain................................ ..... ... 6, 12

plant assemblages ............... ............................. 19

Split Creek Sandstone Member................. ....6, 12,

60, 62, 64, 68, 69, 78, 79, 86, 87 

Katalla specimens, Cinnamomophyllum 

kushtakensis ........ ........................................... .67

katallaensis, Litseaphyllum..... ......... 10, 34, 68; pi. 29

kaulfussi, Lygodium ............................. ..................... 52

Kazakhstan .... .............................. ........................... 42

Keasey Formation .............................................. 12

marine rocks...... ................................................. 86

Keasey stage, age equivalent .... . .................. 12

kefersteini, Alnus ......... .......... .. . . ................... 16

Kenai flora, type.......................... ...... ........ ...........2

Kenai floras, Chaney's correlation ........................ 18

Kenai Formation ........................................................1

age....................................... .... ......... ................ 40

Cook Inlet region ... ....................... ....... . 13, 14

floras.......... ...... ............................ ..............1,2,14

plant assemblage .................................. 13, 14, 19

Kenai group, mollusks ............. ......... .. .... ...... 40

Kenai Group, Angoonian Stage . ......... ........ 13, 16

Homerian Stage ............... . .... .............. ... 13

plant-bearing beds.... ...... .. ... ...... .39

Seldovian Stage . ..................................... 13

Keteleeria.... .................. ..... ... ........ .... ..... 48

Knema ......... .................... .... ... ... 28, 41, 50, 65

glauca .......... ..................... . ... ........ ........ 66

korthalsii.... .. ........... .......... .. .................. 66

sp ............ ..... . ....................... 10, 27, 66; pi. 8

Knowlton, F. H., quoted .... ....... .. .... . . ... 2, 39

Kootznahoo Formation, Admiralty Island ......... . 14

Kanalku Bay ....................................... ..... ....... 14

Long Island ................................ ....... . .......... 14

plant assemblages ............ ....... . . ..... 16

pre-Angoonian assemblages .... ..... .. 90

type section ..... ........... ... . . .. .... .. 14

Page

Kootznahoo section .......................................................... 20

korthalsii, Knema.............................................................. 66

Kuiu Island.......................................................................... 14

Kulthieth Formation........................ 6, 52, 53, 54, 56, 59,

60, 62, 64, 65, 66, 68, 69, 71, 74, 75,

76, 77, 78, 79, 80, 82, 83, 85, 86

age................. ............. ......................................... 9, 13

Duktoth River .............. ..............................................9

Esker Stream .................................................. .........7

fossils, abundance ......................................................3

Hanna Lake area ............................. .........................7

Kulthieth River ....................................................... 6, 7

Kummerian assemblages............. ...................... 12

Lincoln stage........................................................ ...6

Marvine Glacier ..................................... . ........ 13

mollusks, diagnostic ..... .......................................7

Eocene, Tejon age........................ ....................7

marine ........... ................................... ......... 6, 13

Oligocene......... ....... ..................... ..................7

Paleogene plants................................ .....................3

plant assemblages ...................................... ..6, 7, 19

plant locality ............ ..................................... 6, 7

plant megafossils .....................................................6

Poul Creek boundary...................................................7

Samovar Hills ..................................................... 7, 9

unknown unit..............................................................7

Yakataga district......................................................9

Kulthieth River............................................................ 6, 7

Kummer sandstone bed ...................................................9

kummerensis, Artocarpoides ..... ...................31, 34, 86

Dicotylophyllum ..................... 11, 12, 86; pis. 25, 30

Kummerian assemblages, Alaskan, 

strand element... ..................................................... 35

Kummerian Stage .............................. ............12, 16, 34

age equivalent.................................................... . 12

Alaskan, forest, subtropical............................... 34

Alaskan assemblage .....................29, 34, 35, 36, 87

climate ............................................ ............... . 36

floral assemblages ............................................ 14, 19

floristic analysis... .............................................. 34

Pacific Northwest assemblage............................ 35

subtropical ........................................................... 36

vegetational analysis ..................... .................. 35

Kushtaka Formation............................ 3, 52, 53, 54, 55,

56, 57, 58, 59, 60, 61, 62, 63, 65, 66, 67, 68, 69,

70, 71, 72, 73, 74, 75, 76, 77, 78, 79, 80, 81

82, 83, 84, 85, 86, 87

age ....... .................. ................. ......................... 2, 9

age equivalent....... ............................................... 6

Berg Lake ...................................................................... 9

Berg Lake flora............................................................ 2

Canyon Creek ... .. ........... .. .......................... ..4

Carbon Mountain........................... ... ................6

Carbon Ridge....................................................... 12

Charlotte Ridge. ........................ ....... .....5,6

coal beds, flora .... ............................................. ...2

coal measures... ...................................... . ... 2, 3

floras.................................. .......................................8

fossil localities .......... . ..... . ....................... 5, 6

Katalla district ...... ................... . ... .... ........ .3

Keasey age ............................. .. .................. ...6

Kummerian assemblages.................................... 12

middle Ravenian assemblages ... ............... 9, 12

mollusks. ........................... ....... ................. .. . 2, 6

plant assemblages .................. ........................ 19

Tokun contact ......... ................. ............. .... .. 6

kushtakensis, Cinnamomophyllum .. 10, 31, 67; pi. 23 

Litseaphyllum ... ....... ............... ....... ....... 31

Meliosma ................ .... ....... 11, 27, 30, 79; pi. 14

Ulmus...... ......... . ...... ... ..... . . .. ...... 31

lanceolatum, Illicium ................... .. ...... ....... .... 62

Land-bridge dispersals. ............................................. 51

Land Bridges, trans-Pacific .................................... 49

tropical, latitude .... ............................ ........... 50

langsdorfii, Sequoia ..... .... ..... .................... ...... .. 18

LaPorte assemblage, northern California............ 35

Page

Larson, Willard, cited ........................................................ 2

larsoni, Lygodium ................................... 10, 31, 52; pi. 20

/ate, Neolitsea .................................................................... 67

Latah flora, Washington .......................................... 46, 80

lateralis, Fraxinus ...................................................... 52, 56

latifolia, Luvunga.............................................................. 74

latum, CinnamomophyllumlO, 27, 30, 52, 67; pis. 8, 9 

Lauraceae .................................. 19, 27, 31, 32, 33, 34, 35,

..............................41, 46, 47, 48, 49, 50, 67, 73, 84, 86, 87

fruits ............................................................................. 67

leaves ............................................................................ 67

venation................................................................ 67

Oregon ......................................................................... 46

organ............................................................................ 67

Laurels, Asian.................................................................... 68

laurinea, Juglans ............... .............................................. 91

Phyllites.............................. ........................................ 91

Sageretia ................................................................. 91

Laurophyllum ... ............................................................. 68

litseafolia..................................................................... 86

presamarensis ................... .................................. 52, 68

raminervum ...... ......................................................... 68

Laurophyllus ........................... ........................................ 68

Lauras ......... .................................................................67, 69

similis .. ....................................................................... 69

Leaf margins, entire ....................................................... 24

entire, dicotyledonous species ............................. 28

habitat ........................ ............................................... 24

Lower Ravenian assemblages, analysis ......... 28

Middle Ravenian assemblages, analysis.......... 32

nonentire ................................................................... 24

upper Ravenian................................................ 33

percentages............................................ 25, 26, 35, 41

Alaska ......................... ..................................... 46

Kazakhstan ...................................................... 43

Oregon, coastal................................................. 46

Miocene ........................................................... 46

relation to climate ................................................... 24

Upper Ravenian assemblages, analysis .......... 33

Leaf-size classes, Raunkiaer's system.................. 28, 29

Leaves, alder ...................................................................... 57

Cinnamomum ........................................................... 89

Fagus .... ................................................................... 88

Greenland ................................................................ 70

Gulf of Alaska section .......................................... 13

Neogene age.............................................................. 13

Neolitsea ........ ...... .................................................... 89

Paleocene............................ ..................................... 70

Sassafras ................ ........ ...................................... 89

texture.... . ................................................................ 23

liliaceous ........................ .......................................... 81

venation................................... ................................ 51

density ............................ ................................. 23

Legumes....................................................................... 72

Leguminosae ........................................... 27, 31, 49, 72

Leguminosites sp .................... ................................. 31

Leitneria . ...................................................................... 43

Lecythidaceae ................................ ................................. 50

Lianes.............................................................. 31, 32, 35

Alaskan, lower Ravenian .. ................................. 30

leaves ................................................................... .. 23

Licaria ............................... ........................................... 67

Limacia ...... ....... .......................................... 28, 50, 64

oblonga.......... .......................... ............................ 64, 65

scandens ............................................................ 64

stenophylla ... .................10, 27, 29, 30, 64; pis. 6, 7

velutina .................................................................... 64

Lincoln stage, Molopophorus stephensoni zone ...... 8

Lindera ....................................................... 34, 48, 67, 69

linearis, Goweria ................................ ............ ............. 75

Liquidambar... ............................. 20, 32, 44, 45, 46, 70

(pachyphylla type)............................................. 44

sp ......................................................11, 31, 70; pi. 24

Liriodendron.............................................................. 44, 45

Lithocarpus. .................................................... 60, 87, 88

coatsi................ .................. ............................. 87, 88

Litsea ............... ............... .... .... ............ 34, 48, 67, 69

litseafolia. Laurophyllum ...... ........ ....................... 86

Litseaphyllum .............................................................. 67, 68

carbonensis.............. .......................... 10, 34, 68; pi. 28
104 INDEX

Page 

Litseaphyllum Continued

katallaensis .................................... 10, 34, 68; pi. 29

kushtakensis ................................... ....................... 31

praelingue ......................................... 10, 68, 69; pi. 22

presamarensis ...... 9, 10, 27, 30, 31, 34, 35, 68, 69;

pis. 9, 22, 28 

presanguinea ................................... 10, 34, 68; pi. 29

raminervum ........................................................... 34

schorni ............................................... 10, 31, 68; pi. 25

similis ........................ 9, 10, 27, 30, 69; pis. 9, 23, 29

venation................................................................ 69

spp ......................................................................... 69

sp .............. 10, 27, 31, 34, 69, 70; pis. 9, 22, 28, 29

LituyaBay... ....................... .... .... ............. 13

London Clay flora ......... 21, 22, 27, 28, 50, 56, 70, 82

longiflorum, Alangium... .......................................... 84

longifolia. Fagopsis .......... .................................... 88

luvunga... ................... ........ ................. 28, 41, 50, 73

crassifolia ............. ....... .. . .......................... 74

latifolia.... ......... .. .......................... ..................... 74

spatiosa ........................11, 27, 29, 30, 52, 73; pi. 10

Lygodium ................. ............ ... .......................... 52

binervatum ........................ ........ ..................... 52

kaulfussi .......... ............ ..... .............................. 52

larsoni......... ..... ............................ 10, 31, 52; pi. 20

Lyonothamnus ............................................ 21, 47, 48, 71

M

Macaranga ....... ......................................................... 50

McClintock, Copeland, cited............................ ......... 92

MacClintockia trinervis ............................................. 38

Macclintockia. ................................................... 35, 44, 86

Paleocene species ....... .......................................... 35

pugetensis ............................ 11, 12, 34, 52, 86; pi. 30

age range................................... .................... 12

sp ........................................................11, 31, 86; pi. 25

MacGinitie, H. D., cited ........................................... . 3

quoted ........... ................................................. 24, 41

macginitiei, Platanus .......................................... 15, 90

Machilus ....................................................................67, 69

macilenta, Rimella............................................ ............. 3

M'clntosh Formation, age ................................ .......... 8

foraminifera ................ .......................................... 8

plant assemblage ............................ ..................... 8

Washington ................................................ ............. 8

MacNeil, F. S., cited ........... .................. ..... ......... 4, 6

quoted ....................................................... ... ........... 41

macquarryi, Corylus .................. . ................ .......... 89

Madagascar, Celastrus...................................... ............ 75

Maddren, A. G., quoted .................................................... 2

magnified, Juglans ...................................................... 14

Malpoenna ....................................................... 52, 84

Magnolia ............................... 2, 41, 43, 44, 45, 48, 49, 62

californica .......................................... ..................... 68

avails.... ................................................................ 52, 66

reticulata .......................................... 10, 34, 62; pi. 30

wormskioldi ................................. ....................52, 66

Magnoliaceae....................................................... 34, 47, 62

magnum, Asplenium ........................................................ 53

Mahonia.. ........................... ............................................... 48

malaanonan, Parashorea .................................83; pi. 19

Malaspina district .................................................6, 13, 14

fossil collections ............................................................3

unnamed marine siltstone and 

sandstone sequence ..................................................6

Malaya, Celastrus monospermoides .......................... 75

leaf-margin percentages......................................... 25

Mallotus................................................................................ 71

comstocki.................... .................................................. 71

Malpoenna ...................................................................... 2, 41

magnifica ...... ....................................................... 52, 84

Malvales ..................................................................14, 33, 80

mangiferum, Dracontomelon ........................................ 90

Mangroves, Asia, southeast.......................................... 30

China ............................................................................ 30

distribution.................................................................. 22

Manila, leaf-margin percentages ................................ 25

marginatus, Rhamnus ........................................52, 66, 82

Page

Marine megafossrl geochronology, Pacific Coast.. ..8

Martin, G. C., quoted........... .. . . ........... .. 2

martini, Alnus ... . ...... 10, 27, 30, 34, 58, 59; pis. 5, 21

Eometrix ............................................................. ..... 4

Marvine Glacier, plant assemblage.......... ........... .... 13

plant-bearing beds ........... ..... ................. ........ 13

Mason, H. L., quoted................................... ....20, 37, 41

Mastixia........................................... ....................... 50, 85

irregularis ........ ......... ..... 11, 27, 52, 85; pi. 15

philippinensis .................. .. .. ... .... ........... 85

Matanuska Valley . ......... ... ........... ....... 14

Mazatlan, Mexico. ........ .... ... .......... ....... ..54

Megafossil assemblages, Atlantic coast.................. 48

Melanesia, Celastrus .................................................... 75

Parashorea malaanonan...... ...... .. .. ............ 83

scandent vines....................................... ............... 74

Melanolepis.. ................... ... .................... .. ..... .. 50

Melanorrhoea.. .......................... .................. . 41, 74

alaskana . ...........................11, 27, 30, 52, 74; pi. 10

Melia.................... .................. ........... ................. 46

Meliaceae........................................................ .......... 27, 72

Meliosma .. .. ......... ........................................ . . 50, 79

duktothensis ........ .......... . 11, 27, 30, 79; pi. 14

kushtakensis .... ... ..... ........11, 27, 30, 79; pi. 14

simplicifolia .................................. ...................... 79

Menispermaceae ................19, 27, 31, 32, 34, 35, 41, 47,

50, 56, 62, 75 

Mesophyllous tropical plants, texture...................... 23

Metasequoia ....... ......................... .......... 14, 18, 48, 53

glyptostroboides ........ ............. ...............13, 14, 18

occidentalis .. ....... ............................ .............. 18

Metroxylon ......................... ............................................ 54

mexicana, Hyperbaena...... ............................................ 89

micrococca, Ilex..... .......................................... ................ 75

Microcos..... ........................................................................ 84

Microfossil assemblages, Atlantic coast .................. 48

Microphyllous plants, texture ...................................... 24

microphyllus, Zizyphus .................... .......................... 61

Miller, D. J., cited...................... ......................................2

quoted...........................................................................7

milleri, Anamirta ................... ............10, 27, 30, 62; pi. 6

minuta, Salix ................................................................... 18

Miquelia . ...... ....................................................... 50, 91

californica............. ....... ...... ................... ........ 91

Miquelia reticulata.... .......... ............................. ...... 91

Mixed Mesophytic forest. 23, 28, 33, 46, 48, 56, 63, 90 

Alaska ...................................................................... 56

Asia, eastern ............................................................. 48

depauperate .......................... ............................. 48

Japan ........................................................................... 58

lineages ................................ ....................................... 49

Miocene ....................................................................... 47

Neogene .................................................. 45, 56, 71, 90

North America.................................................... 45, 48

eastern ........................................................... 48, 49

western ......................................................48, 49, 58

Oregon ........................................................................ 71

origins........................................................................... 43

Pacific Northwest........................................ 47, 56

Tertiary ........................................................................ 49

theory ........................................................................ 43

tropical disjunctions ............................................... 51

Mixed Mesophytic genera .............................................. 45

Atlantic coast.............................................................. 48

Cretaceous..............................................................43, 44

Eocene ......................................................................... 44

extinctions................................................................... 46

organs, diverse .......................................................... 44

mixta, Rhus ...............................................11, 27, 74; pi. 19

Mohrodendron ................................................................ 2, 41

inopinum ...................................................................... 51

Mollusks, Eocene ......................................................3, 4, 40

Eocene, Tejon age-.............-............-..-............5, 6, 7, 8

localities, stratigraphic relationships ....................1

marine ....................... 4, 7, 15, 16, 20, 39, 40, 41, 42

localities...................................................................7

Molopophorus gabbi zone ............................ 6, 12, 13, 15

Molopophorus stephensoni zone.............................. 6, 13

Lincoln stage .......................................................... 8, 12

Page

Molopophorus tejonensis... . ... ..... ........ . ..5

Monocotyledonae ... ............................ ................. 26

Monocotyledones ...... ................. ..... ..... 31

monospermoides, Celastrus . . .................... ...... 75

montana, Eucommia... .................. .... ......... ...... 70

Moraceae ...... ...... ... ............... ............ ............... 86

multiflora, Eugenia . .. .......................... ................... 85

Myrica . ....... ... ... ........... ............. ................. 45

Myristica ................. .. . ................... 28, 41, 50, 66

sp ............................. ..... 10, 27, 30, 52, 66; pis. 5, 8

Myristicaceae ... ....... ...... . ....27, 28, 31, 50, 65

Myrtaceae ...............................................27, 31, 33, 50, 84

Myrtiflorae .... ...... .. .... ............................27, 33, 83

N

Nectandra ............................... ......... ... ........ 67, 68

presanguinea . .......... .... ........ ....... ... ....... .. 68

Nenana Gravel, deposition........... ............ ........... 42

Neogene assemblages.................. ................................ 42

North America .......... ..... ... . .. ... ................ . 46

Siberian .... .............................................................. 41

Neolitsea ...................... ........... ........................ ...... 48, 67

lata......................................... ................................. 67

nevadensis, Euonymus................................... .............. 71

Prunus ........ .... ......................................................... 11

New Guinea, Celastrus monospermoides ................ 75

Newberry, J. S., quoted ......... ................................... 39

newberryi, Ulmus.......................................................... 60

(newberryi type), Ulmus........... ........................... . 44

Nichiwak Mountain..... ... ................................ 6, 12

nigella, Juglans.......................... .................................... 18

Nipa.... ........................................................................ 21, 56

distribution........................................................ 22

Taiwan.................................... ................................ 21

nitida, Alnus ................................................................ 59

Hypserpa............................ ........................ ..... 63

nordenskioldi, Glyptostrobus... ............................. 18, 53

Norton Sound assemblage ........................................... 41

nutans, Clerodendrum .................................................. 85

Nyssa ............................... ............................. 18, 44, 48, 49

Port Graham ...... .................................................. 18

Port Moller region .............. .................................... 18

arctica.. ....................................................................... 18

o

Oaks, lobed ............................................................43, 45, 46

obliquifolia, Celtis ......................... ........................... 91, 93

Plafkeria ........................... ............................14, 81, 91

Willisia........................................................................ 93

oblonga, Limacia ........................................................ 64, 65

occidentalis, Anemia........... ........................................... 52

Metasequoia ............................................................ 18

Ocotea..................-...-......-.........--.-.---..............67, 89

eocernua........ ................................................................ 89

odorata, Cammga.............................................................. 66

Ohanapecosh assemblage............................................ 90

Olacaceae ......................................................................... 50

Olax.. .................................................................................. 50

Oligocene-Miocene boundary, Europe...........................8

Olympic Peninsula, Washington ............................... 25

Oreamunoa....................... ......................................48, 55, 56

(Oreamunoa), Engelhardtia .......................................... 55

oregona, Camelia ..... ..................................................... 92

Fagus ..................................................................... 15, 88

Ilex......................... ...................................................... 92

Tetracera ...................................................................... 88

oregonense, Boreoscala condoni ......................................6

oregonensis, Alangium................................................... 93

Oreopanax....... ..................................................................... 48

orientalis, Diospyros ..... .................................................. 89

Euptelea....................................................................... 89

Juglans......................................................................... 88

Pterocoryo........................................ ..................... 14, 88

Orocline ............................................................................... 36

Osmunda sp ...................................................................... 13

Ostrya..............................................................................57, 58

Ostryopsis............................................... ........................... 58

ovalis, Magnolia.......................................................... 52, 66
INDEX 105

(pachyphylla type), Liquidambar.... ................ 44

Pachysandra ........ . . .............. . ... ............... ....... 45

Pacific Coast marine megafossil chronology ........... 8

Pacific Northwest, lowlands................. ................ 19

uplands ......... ................. ..................... 19

pacifica, Fagus .. ............ .................. .. ... 88, 89

Paleocene assemblages, subtropical .... ........ ....... 33

temperate, warm, high latitude.. . ................. 33

Paleogene assemblages, Alaskan, latitude ..... ....... 36

Eurasia......... ....... .. .............. ............. 44

European .............................. .............................. 39

North America ............ ....................... .......... 44

Paleogene climates, Alaskan .. ....... . .................. 36

Paleogene floras, Gulf of Alaska .. ......... ............... 1

Paleogene forest, Eurasia ....... ............... ............... 49

Paleogene formations, Katalla district............. ....... 3

Paleogene migration routes, tropical.......... ... ....... 49

Paleogene plant assemblages, Gulf of 

Alaska region..... ....................................................... 3

Paleogene plant localities, age ................................. 7

stratigraphic relationships ................................. 7

Paleogene species, systematic revisions.................. 87

Paleogene subtropical assemblages........................ 24

Paleogene tropical assemblages .... ............. .... 24

Paleogene tropical forest, North America, 

eastern ...................................................................... 49

Paleogene forest, North America, western ............ 49

Paleophytocrene ........................... .................. 46, 75, 76

elytraeformis .................... 11, 27, 30, 52, 76; pi. 13

foveolata .......... ............... ...... ............................ 76

Paliurus ............... ............................................................ 45

Palmacites .. .................................................................... 54

genus and species indetermined ......................... 54

sp ........... . ................................ ........ 10, 54; pi. 4

Palmae ........................................................ 26, 31, 50, 53

gen. indet..................................................................... 10

Palms .... ....................................................... 31, 41, 42

fan ........................ ............................................. 54

Alaska ...... ...................................................... 50

central...................... .......................... 42, 50

Paleocene rocks, Matanuska Valley .......... 54

feather ...... ............................................................... 53

Greenland, Tertiary............................................... 50

leaves .........................................,............................. 54

sago.............................................................................. 54

(papyri/era type), Betula ............................................. 45

Pantropical distribution............... ................................. 51

Pantropical families ....................................................... 50

Parashorea ...............................................28, 41, 50, 82, 92

chasmanensis .......................................................... 92

malaanonan ....................................................S&\ pi. 19

pseudogoldiana .................. 9, 11, 27, 29, 30, 52, 82;

pis. 16, 19 

Paratinomiscium .............................................................. 65

conditionalis.......................... 10, 27, 30, 51, 65; pi. 7

Paratropical, concept ..................................................... 26

Parietales........ ..................................... 27, 31, 81

Pasania ............................................................................. 87

peduncularis, Allophylus................................................ 78

Peraphyllum......................................................... 47, 48, 71

Perse washingtonensis...................................................... 5

Pcrsea ............................................................2, 41, 48, 67, 69

praelingue.................................................................... 68

pseudocarolinensis.............................................. 69, 89

spatiosa .................................................................. 52, 73

Philadelphia bendirei..................................................... 89

Philippine Islands, Allophylus peduncularis.......... 78

Celastrus monospermoides ................................... 75

Cryptocarya infectoria ............................................ 69

Euodia crassifolia..................................................... 73

leaf-margin percentages ........................................ 25

Miquelia reticulata.................................................... 91

Zizyphus inermis ...................................................... 80

philippinensis, Mastixia ................................................ 85

Phoebe ................................................................................. 67

Phoenicites .......................................................................... 53

sp . .................. ....................... 10, 31, 53; pi. 20

Phyllites cinnamomeus ........................................... 52, 67

Phyllites Continued age 

laurinea ........................................................................ 91

Physical stratigraphy....................................... ............. 3

Phytocrene ....................... ........................ 28, 50, 75, 76

acutissima.............. 9, 11, 27, 29, 30, 76; pis. 12, 13

blancoi ........ ................................................ .... 77

bracteata .................... ........................................ 77

sordida ................... ......9, 11, 27, 29, 30, 77; pi. 13

sp ...... ..................... ................ ........................... 29

Phytocreneae ...... ..... ........................................ 30, 91

Pinaceae.............. ..................................................... 47

Pinnacle Hills .............................................................. 13

plant assemblages ............................................. 7, 20

Yakataga Formation.. ............................7, 13, 20

pinnata, Cyathea ............................................................ 53

Hemitelea ............................................................... 8, 53

Piper ........................................................ ........................... 2

concavum .................................................................... 51

disputabilis................................................................. 51

piperoides, Cercidiphyllum ............................................ 61

Tetracentron................ 10, 12, 27, 31, 34, 44, 61, 80;

pis. 5, 22, 28 

Zizyphus ............. ..................................................... 61

(piperoides type), Tetracentron ................................... 45

Pistacia ................... ... ................................................. 48

Pitar californianus...... ................................................. 3

Plafker, George, cited ........................................................ 2

plafkeri, Calkinsia ...........10, 12, 27, 31, 62; pis. 6, 23

Plafkeria ........................................... ........15, 33, 34, 81, 91

obliquifolia .................................................... 14, 81, 91

rentonensis ................................................................. 81

thomae ...................................................................... 15

sp ................... . ...I1 - 33, 81; pi. 27

Planera.................................................................................. 41

aquaticiformis ...................................................... 51, 56

Plant assemblages, Alaskan, temperate, warm .... 46

Arctic .......................................................................... 39

Tertiary .............................................................. 39

Bridge Creek..................... ................................... 46

fossil, Alaska....................................................... 40, 43

Siberian ................................................................ 43

lowland, Oligocene................................................. 46

Paleogene, European....... ...................... .............. 39

Willamette.................................................................. 46

Plant fossils, unknown unit......................................... 7

Plant stage epochs.................- ................................... 8

Plant tolerances, changed....................................... 21, 22

Plant venation, relationship with climate................ 23

Plants, dispersal............................................................... 47

evergreen................................................................... 37

megafossil assemblages, fluviatile. .................. 24

lacustrine.. ................................................... 24

tropical, dormancy......... ..................................... 37

Platanaceae ............................................................... 34, 71

Platanus...--.--.-.-.-.-..-................................ 15, 44, 71, 90

aceroides .............................................................. 71, 90

bendirei ..................................................................... 90

(bendirei type) .......... ..... .. .. 44

comstocki.................................... 11, 12, 34, 71; pi. 29

dissecta......................................................................... 19

(dissecta type) .....................- -.......- . . ..- - 46

macginitiei ........................................................... 15, 90

n. sp... -. ..... .. .. 12

Plate, Pacific, northeastern ..................................-. 36

thrust, upper. ..............................-.. .....- ...-.- 36

Platycarya....-.-.-.-........-- 21, 32, 41, 44, 45, 46, 47, 56

China .. ........ ....-..........---..- ---.. .--- 21

Taiwan..........- ....- . . ... .. -- - - 21

pseudobrauni .............. 9, 10, 26, 30, 31, 51, 52, 56

pis. 4, 5, 21 

strobilacea.................................................................... 56

Platyptera............................................................................ 88

pleiosperma, Euptelea...................................................... 89

Podostemaceae ......... ................... .......................... 81

Pole, Chukchi Sea ............................................................ 37

Poles, Cretaceous ........ ................................................... 37

magnetic, past..... .................................................... 37

North America ....... ................................. 37

Paleogene .................................................................... 37

Page 

Poles Continued

rotational........... ........... .... .................. 37

Pollen, fossil, Quaternary ............................ ................ 31

ulmaceous ........................... ..................... ............. 60

Pollen assemblage, Bering Sea ......... ...... .......... 20

Pollen floras, Miocene.... ............................................. 13

Queen Charlotte Islands .............. ......... .......... 13

Populus.............................................. 18, 41, 45, 47, 54, 55

amblyrnhynca............................................................ 17

arctica..... .............................................................. 17, 61

balsamoides .................... ................................. 52, 75

daphnogenoides ..................................................... 61

dimorpha.................................................................. 54

glandulifera ............................................................. 61

leaves ............................................................................ 33

Port Moller assemblages ........................................ 18

richardsoni............................................................ 17, 61

rotundifolia................................................................ 61

smilacifolia................................................................ 61

speciosa ................................................................. 61

xantholithensis ........................................................ 61

zaddachi..................................................................... 61

sp ........................... ......... .10, 32, 54; pi. 26

Parana speirii .................................................................... 85

Port Graham, Seldovian assemblage .. ...... 20

Port Graham assemblage, age............... ..... 19, 41

Port Moller assemblages, Acer .................................... 16

Betula........................................................................... 17

Cercidiphyllum ......................................................... 17

Salix .............................................................................. 18

Port Moller floral assemblages ............................. 16, 18

Port Moller floral localities, age .................................. 16

Port Moller region, Alaska Peninsula. ................ 16

Post-Kummerian rocks ....... ................... 13

Potentilla............................................................................ 71

Poul Creek, mollusks ....... ........................................n

Poul Creek Formation. - ............... . 7, 60

fossils, lack ....... . ..........................3

Juglans............................................................................n

nonmarine equivalent.... .................. 13, 20

plant megafossils ................................................... 6, 7

Poul Creek........... .................................................... 7

Poupartia.............................................................................. 50

praelingue, Litseaphyllum ................... 10, 68, 69; pi. 22

Persea........................................................................... 68

Precipitation, Alaska, summer .................................... 48

presamarensis, Cryptocarya.................................... 51, 68

Laurophyllum ...................................................... 52. 68

Litseaphyllum ...... 9, 10, 27, 30, 31, 34, 35, 68, 69;

pis. 9, 22, 28 

presanguinea, Litseaphyllum.............. 10, 34, 68; pi. 29

Nectandra.................................................................... 68

Principes .......................................... . ....26, 31, 53

productum, Acer trilobatum.......................................... 16

Prosopis................................................................................ 37

Prunus ............................................................................ 71, 87

axelrodi ....................................... 12, 32, 33, 71; pi. 26

chaneyi.......................................................................... 71

hameyensis ............................................................... 71

nevadensis .................................................................. 11

serotina ........................................................................ 71

pseudobrauni, Platycarya..9, 10, 26, 30, 31, 51, 52, 56;

pis. 4, 5, 21 

Ulmus...................................................................... 51, 56

ps(.udocarolinensis, Persea ...................................... 69, 89

pseudogoldiana, Parashorea .............. 9, 11, 27, 29, 30,

......................................................................52, &2;pls. 16, 19

pseudogoldianus, Rhamnus ........................... 52, 82

Pseudolarix.......................................................................... 48

Pteris inaequilateralis................................................ 51, 53

Pterocarya.......................................... 18, 20, 32, 44, 56, 88

Port Moller assemblages ........................................ 18

orientalis................................................................ 14, 88

pugetensis ........ 10,12, 31, 32, 33, 34, 45, 56; pi. 21

Pterospermites.............................................................. 38, 44

Pterospermum.................................................................... 84

Pterostyrax.......................................................................... 86

Ptycoraphis.......................................................................... 54

Puget Group..........................9, 12, 28, 52, 55, 61, 63, 67,

68, 70, 75, 86
106 INDEX

Page 

Puget Group Continued

Cashman assemblage............................................... 9

Clarno Formation.................................................... 33

early Ravenian assemblages ............... 8, 9, 75, 80

floras, middle Ravenian..................... ............ 32, 69

Franklinian Stage ................................ .................8

Fultonian Stage ....................................... 8, 9, 75, 76

Green River Canyon section....................................9

John Day Gulch assemblage .......................... . 33

Mixed Mesophytic genera ...................... ....... 32

nonmarine rocks..................... ....... ................ 35

stages .......... ......................................... ..... .... 8

Steel's Crossing assemblage..............................9

substages........... ...... ..................... ...... .... ......8

Tukwila Formation .......... ......... ............ ..... ... .8

Washington, plant assemblages.......... ....... 7,8

pugetensis, Allantodiopsis ............9, 10, 12, 26, 30, 31,

53; pis. 2, 20 

Calkinsia .................................................................... 12

Dryophyllum ............................ 10, 12, 31, 60; pi. 21

Macclintockia....... ............. 11, 12, 34, 52, 86; pi. 30

Pterocarya ........10, 12, 31, 32, 33, 34, 45, 56; pi. 21

Pycnarrhena.................... ............................. ..... 28, 50, 65

sp .....................................................10, 27, 30, 65; pi. 6

Pyrenacantha ..... ......... .......... ....... ...... 28, 50, 77

volubilis .................. ..... ....... ................. 78

sp ................................................ 11, 27, 30, 77; pi. 11

Pyrtaceae, genus.......................... ... .11

Pyrus ........................ ............. . . .. ..... 71

Q

Queen Charlotte Islands, pollen floras .......... .... 13

Quercophyllum groenlandicus ... ......... .88

Quercus ... ..... .... .... . ..... .... 20, 44, 45, 60

deflexiloba. .... ...... . . ... .... .. .. .. .. . 19

furuhjelmi ... ... . . .......... .. ... .. 16

groenlandica..... ...... .. . ........... ....... .. 88

weedii . .. ....... ................ .. .......... 89

R

Raging River Formation

Rain forest, Alaskan, lower Ravenian . Australian .. ............ ... . ... ... ...

coniferous .... ..........

extratropical. .........

.. . 31 

....... 28

.... 25

.... 25

montane........... ........... ..... .. .. 25,26,29

paleotropical. ... .. ...... ........... .... .... 82

paratropical ...... .. 21, 23, 25, 28, 29, 32, 35, 36,

42, 44, 45, 54, 56, 57, 58, 60, 61, 63, 82 

Alaskan .. ........... ....... .......... ...... 36, 49, 50

broad-leaved evergreen.......... ..... .......... 25

China .. ... .. .... ... .. ..... .... 73, 80

Hainan . ......... .. ..... ... 25

Hawaii . . . .28

Paleogene .... . .... .... 49, 50, 58, 71

Taiwan ... .... ..... 25, 28, 42

United States, conterminous . 33, 50 

warm..... ....... . . ......... 50

submontane ............... .......... 25, 26, 28, 29, 36

Philippine Islands ........ .... .... ...... .... .. 28

subtropical . . .... 25, 29, 42, 45, 56, 57, 58, 61

Paleogene . ........ 71

United States, conterminous .... ......... 33

temperate..... .... .. . . ....... ... 28

tropical....... . ..... 23, 24, 26, 28, 44, 50, 54

raminervum, Laurophyllum.. .... ........ . 68

Litseaphyllum ...... ... ....... ... ... 34

Ranales ........... . ... 27,31,32,34,61

ravenensis, Calkinsia .... .... . 12

Ravenian Stage. .... .. .. .. ... ....... ..... ........ 8

definition, original ... ............. 8

dicotyledonous species ... . .32

early, Alaskan assemblage . . 29, 35, 36, 50, 63 

climate ........... . . ..... . .. ..... ... 35

Eocene assemblages ........... ...... .. ...... 51

evergreen species ..... ...... .. . .......... . ..... 37

floras...... ...... . .. ..... ..... ..... .. . 19

light .. .. ... .... ..... ... .... 37

paratropical . . .... 36

Ravenian Stage, early Continued

time equivalents ............................................ 8

late, Alaskan assemblage ....................... 29, 36, 45

climate ......................................................... 36

temperate................................................... 36, 45

lower ............ ...................................................... 8, 26

flora, Alaskan ..... ............................. 26, 27, 28

Eocene . ................................ .............. 26

Washington ....................................... 28

floristic analysis ...... ......... . ... ...... ~ 26

time equivalent ......... ............ ........... 8

Transition stage .................... - ........ 8

vegetation .... ................... ..... .............. 36

middle .... . .......... ... ................ ...... .. 9, 31

Alaskan assemblage . ...... ................... 29, 36

climate . ...... ....... . ..... . ... .. .. ........ 36

floras.... . ........ ..... .. .... 19,31,32

floristic analysis ..... . ..... .... ~~~...... 31

subtropical .. . .. .. .. .. ....... .. .... .. 36

time equivalent. ..... . ............ . ....... .. 8

vegetational analysis ...... .. ...... ......... 32

upper............ . ... - ............... .... ... ...... 12

definition, original ............ . .. ... .. . 9

floristic analysis ...... ...... ... .... ... ....... 32

type section. ....... ...... ......... . .... .......... 9, 12

vegetational analysis ..... . ....... -........... 33

reinwardtiana, Saurauia . .................... ... .......... 81

rentonensis, Plafkeria... .... .... ............. .... .. ... 81

WiWsia . ... ....... ....... ..... .......... ....... .. 81

Republic assemblage .... . ..... ...... - ... 35, 61

Washington .. . . .... ..... ..... ...... . 57, 88

Republic Basin floras, radiometric age ...... ... 12

reticulata, Magnolia ... .............. .. .. 10, 34, 62; pi. 30

Miquelia ..... . .... .... ..... ..... ...... ......... 91

Rex Creek assemblage ...... ..... . .... ... . .... 54

Rhamnaceae. . .................. 27, 33, 43, 79, 82, 92

Rhamnales .... .................. .. ..... 14, 27, 33, 79

Rhamnidium . .... ... .......... . ... . ...... 82

chaneyi . . .. ...... ..... - ... - 82, 92

Rhamnites cashmanensis ..... . ..... ...... -.. 82, 92

franklinensis . ........... ...... .......... 82

Rhamnus ...... ... . .......... ..... .... - ..... - 41,45, 79

decheni........ .... .... . . ........ - ... - 51

marginatus ..... .... ..... ..... ..... .. - 52, 66, 82

pseudogoldianus ........ ................... 52, 82

sp .... .. .............. ............. 11, 33, 79; pi. 27

Rhizophora .. ..... .. .... . ... .. .. .. . .. 83

Rhizophoraceae ......... ...... .... ... 30, 83

Rhododendron... .... ... .. ... .. ..... 41,47

crassum .... . ........ . ..... . 29, 51, 86

Rhus .... . .... .... ..... . - 44, 45, 74

mixta ........ ... ..... .... 11, 27, 74; pi. 19

Richards, P. W., quoted . .... ....................... 23

richardsoni, Dicotylophyllum . ...... 16, 17, 44, 61

Populus. .. ... .... ....... ... . ..... .. 17, 61

Salix . - - . . . . -- -- - - - 55

Rimella macilenta .. .......... ..... ...... .... 3

Rocky Mountain, Paleocene assemblages .... 71

Rocky Mountain region, Maestrichtian rocks . 57 

Rocky mountains, ancestral .. ...... ................ 54

Rocky Mountains, Oligocene assemblages .. ....... 34

Rosaceae .... .... 19, 32, 33, 43, 46, 47, 49, 71

Rosa .. ... ... . .......... .. . ..... 71

Rosaceous genera, Alaskan, upper Ravenian .... 33

Rosaceous genus, British Eocene ... .... ....... 33

Resales ... ............ ....... .. 14, 27, 31, 32, 34, 70

rotundifolia, Populus... ..... .. ...... .. ... 61

Vitis . ... .... . ... .... . ............... 14

Rubiaceae ... .... ... . ... ... . . .... .... ..... . 51

(rubrum type), Acer .. .. ...... ... ......... . 45

Rubus ... .. ....... .... .. .... 33, 45, 46, 71

seeds .. ... . .......... ... ...... 71

Ruby assemblage, Montana .. .... .... .. ... 46

Rocky Mountains .. ..... ........... .. 34

Rutaceae . . .. .... 27, 31, 33, 50, 73, 87

Sabalites

Page

Sabalites Continued

sp ..... ............ ....................... -- .-- 26, 30, 31

Sabiaceae............. . .................................. 27, 31, 79

Sachalin, Arctic Miocene........ - ............. ..... ...... 39

Sageretia . ................ - ... - . ....... -44, 45, 50, 80, 91

elegans ............ . ....... .............. ....... 91

laurinea .... ... ...................................................... 91

sp .........................................................11, 27, 80; pi. 15

St. Lawrence Island............... ... .... ....................... 14

Sakhalin ........... ................ .... .............................. 20

Salicaceae .. . ....................... 19, 24, 32, 43, 46, 47, 54

Salicales . ... . ........... .................................... .... 32, 54

Salix ........... ... ............. ..75,20,33,45,47,54,55

alaskana ...... .. .... .... ....... .. ........ ........... 18

aquilina . ....... . .......... .. ..... ..... ...... ........ 54

barclayi ...... ........ .... ........ ........................ 90

bebbiana ..... ..... ..... ... .. .... .......... ......... 90

carbonensis.. . ...... . ... ...... 10, 32, 55; pi. 26

coatsi........... ... ...... .. .......... ..... .. ... 87

cuprouallis ..... ............ .... .. . ............ 90

exigua .. ...... ............... ... ................ . .......... 88

hesperia ........ ............... ............................. ...... 19

kachemakensis ............ .................................... 18

minuta . ..... .. ............. ....................... ............. 18

richardsoni. ............ ............................................ 55

wilsoni ........................ .............................. ...... 55

sp . .. ........... .......... ................ 10, 32, 55; pi. 26

Sambucus ............................ ...................... .. ...... 45

Samovar Hills, marine mollusks ..........................7

plant assemblage ........ .... .................... .9

sanbornae, Dracontomelon .................. .............. 90

Sapindaceae .........................23, 27, 31, 34, 50, 78, 87

Sapindales ........... ..................... 14, 27, 31, 33, 34, 74

Sapindus .. ..................... ........................ ......... 79

sp ...... .... ........... .. ........... 11,31, 79; pi. 24

Sapotaceae .. ....... ............................................. 49

Sassafras............ .......................... ............... 48,67,89

bendirei ......... ......... ..... ..... . ... .. ..... 89

columbiana .................... .. ............. . ...... 89

Saurauia . .......... ................ .... ..... .... .... 50, 81

reinwardtiana ....... . .. ... ............. ...... 81

sp....... .... ................ .. ..... 11, 27, 81; pi. 18

scandens, Limacia ...... ............. ............ 64

Schizaeaceae ........ ...... .... ........ .... 26, 31, 34, 52

Schoepfia..... .... ... .... ........... . ..... 48

Schorn, H. E., cited .... .... ... ......... ..... ...........3

schorni, Litseaphyllum ..... .... .... ....10, 31, 68; pi. 25

Scio flora .. .............. ... ...... .... 15, 90

Sclerophylls ............................ ... ...... .. ..... 26

scorpiodes, Stemonurus . ... ...... .... .... 78

scotti, Alnus ..... . .......... 10, 27, 30, 52, 59, 86; pi. 4

Seed assemblage, Siberia........ ..... . .... .... .. ... 42

Seldovian assemblages, Alaska ........... ... 18, 20, 54

Seldovian Stage .. . ......... ..... ....... ... 8, 13

floral assemblage .............. .... ......... ... 18, 20

type section....... ... .... ... ........... 72

Semecarpus. . ........ ... ..... .. ....... .2,41,74

alaskana ......... . . ......... ... .. 52, 74

Sequoia langsdorfii . .... ........... ..... ..... 18

serotina, Prunus ..... . ..... ......... .... . 71

serrulata, Trochodendroides ...... ... ........ ...... 61

Zizyphus . ..... ... .... ............ .61

Shell Oil Company, fossil plants . ............ . ... .2

Shockum Mountains .. ............. ... . ........4

mollusks locality . . . ............. ......... .....A

shumardi, Acila. ... .. ............. .... ..... ..6

similis, Laurus.. .... .. ... . .................. .. ...... 69

Litseaphyllum ... 9, 10, 27, 30, 69; pis. 9, 23, 29 

simplicifolia, Meliosma . . ... ...... . ..... 79

Sinomenium ... ... ... .... ...... ........ .43

Sinowilsonia .... . . .. .... .......... 44, 45, 47, 70

sinuata, Alnus ... ... . ... .... .... ..... ...... .. 59

aragoensis, Whitneyella .. . ................ ...... .... 4

Whitneyella......... ..... .................... ............. .. 5

Sitkinak Island, Angoonian assemblage. .. .... 20

marine beds .... .. .............. ... .... ... .. 14

nonmarine beds. .......... .. .... ..... . . . 14

plant-bearing beds ....... . ........ ... 13

smilacifolia, Populus... .......... ... ... ...... .. 61
INDEX 107

Smith, P. S., quoted..... 

Sorbus ..... ..... .... .....

carbonensis ..... ....

sordida, Ficus... . ....

Phytocrene . . ... 

spatiosa, Luvunga ... .

Persea . ........ ...

speciosa, Barringtonia

Populus . .

Page

.. .............. . .. ......... 2, 40

.. .............. ...... ......46, 71

...... -11,32,34, 71; pi. 27

........... .................... 77

.9,11,27,29,30, 77; pi. 13

.11,27,29,30,52, 73; pi. 10

.. ... .... ............ 52, 73

. .. .......... .... ..... .. 83

... ... .... ...... ....... ... .... 61

speiri. Holmskioldia .. ... ... .. ..... 11, 34, 85; pi. 30

speirii, Parana... ... ... . ... .... ... ....... .... ... 85

Spiraea . . .... ..... . . .. ... .... .... ..... .... 46

Spitzbergen, Arctic Miocene.... .... ............... . 39

Split Creek Sandstone Member, gastropod. . .. 6

Katalla Formation . ......6,12,60,62,64,68,69

. ... .... . . .... ... . 78, 79,86, 87

leaves ........ .... .... . .. .. .. ... ... .6

mollusks. ...... . .... 6

Nichiwak Mountain. . .... ..... ..... .... 6, 12

plant assemblage . ....... .... .. 6, 13

stanleyanum, Dryophyllum ....... . ..52,74

Steel's Crossing assemblage.. .... ..... .. 19, 63

Puget Group ... .... .... .... ...... 9

Steenis, van, C. G. C. J., quoted . . 49, 50

Stemonurus.

alaskanus

scorpiodes . ..........

stenophylla, Limacia..

.. . 28, 50, 78

11, 27, 29, 78; pi. 13

...... ..... ..... ..... . 78

10, 27, 29, 30, 64; pis. 6, 7

Sterculiaceae .. ..... ... ... .... .... .... ... . 14, 84

stevensoni, Betula .. ... ...... .57

Stillwater Formation . ..... ................ . ...3

Berg Lake section. .. . ... ..... ......... ....3

Carbon Mountain ... .... ... .. . .. ..4

coal beds ...... .... . .... ..... .. ..... .... ...3

Katalla district . .... ... ... .... ... 3

lateral equivalent .... . ..... ....3

mollusks .. ..... .... . ......... .. 3, 4

plant localities . ..... .... ...... .... ... 3, 4

Shockum Mountains.... .... ... . . ..... . . 4

type section. . ..... ... .... . .......... 3

strobilacea, Platycarya .. . .............. ............... 56

Styrax .... .............

sublutea. Betula .... .... ....

Subtropical, concept .... ..... . ... ...

definition . ......... . ...

sureni, Toona . ... ... .... ...

Sweet Home, Oregon... .....

Sweet Home flora ... . ... .... .

Sympetalae . ...... ......... ....

Symplocos... ... .... .... ....

Systematics. .... .. .........

Systematic revisions, Paleogene species

.. . 45 

..... 17 

.. 26 

.. 25 

72,73 

... 90 

. 15, 90 

.. 51 

45,49 

.. . 51 

..... 87

Taiwan, leaf-margin percentages . 

taurinensis, Aralia . .... ... .. . . .... .

Celastrus ...... .. . ..... ......

Taxodiaceae .. . .... ... .. . 14, 26,

Tejon stage ..... . . .....

Tejon stage of Weaver . ..... ...

tejonensis, Molopophorus .. .... ...... ....

tennesseensis, Dryophyllum .... ..... ..

Terminalia . ....... .. ... ... .. .

sp ... . ...... .... . .

Ternstroemites sp . .. ... . . ..... ...

Tetracentraceae .... ...

Tetracentron..

piperoides

32,35.43,44, 

10, 12, 27, 31,

. ........ . 25

...... ..... 89

.. .......... 89

31, 32, 47, 53

.......... . . 8

.... ... ... 5

. ..... .... 5

... . ...... 60

... 2, 41 

... .... . 52, 66

.. ... 31

27, 31, 34, 61 

45, 46, 47, 61 

34, 44, 61, 80;

(piperoides type) ... .... ....

Tetracera .... ..... ......

oregona . .. ..

Tetrapteris harpyiarum .. 

Tetrastigma ..... .

Theaceae . ... .. ... . ..... .... .

genus ... ... .... ....

genus and species indetermined . 

thermatis, Juglans

pis. 5, 22, 28 

...... ....... 45

.... ... . 51

..... 50

. 31, 32, 50, 81 

. ... .... 11

...... 8^; pi. 25

. . 52, 76

thomae, Alangium .. ....................................................... 91

Plafkeria .................. ............................................... 15

Viburnum ................................. .............................. 91

Tiger Mountain Formation, late Fultonian 

assemblages .................................................................. 8

Tilia..................... .... ..................... .......20,23,33,45,50

aspera ..... ...... ................................................. 14

carbonensis.................................. 11, 33, 80; pi. 27

williamsi ..... ..... ............................................... 91

Tiliaceae........ ............................. 14, 23, 33, 81, 84, 92

tiliaefolium, Byttneriophyllum ................................ 91

Tinomiscium. ...... .... ...... ....... .................. 50, 62, 65

Tinospora ..... .. ...................................................... 50, 62

Tivelina vaderensis ...................... ........................ 4

Tokun Formation ......... ............ ............................ 12

age range.. .... ........... ....... .... . ........ ............ 6

Charlotte Ridge ................... ..... ...................... 5

fossil assemblages .............. ......... ............... 2

Katalla district ............. .. ..... .... .............. 3

Keasey age .............. . ........... ................... 6

marine mollusks ... .... ... ................. .... 4, 6

Tolstoi Formation, age.. ........ . .................. 19

Coal Bay, Alaska Peninsula .................... ..... 19

foraminifera. .. ........ .. . ...... . ....... .. 19

fossil plants .. .... ..... ....... ...... ...... .... 19

marine mollusks . ......................... ............... 19

Toona .. ..................... ....... .. .... ... . 44, 72

sureni ........ .......... ....... . ..... . .... ..... ..72,73

sp ........ .... ........... ............. ....... .11, 27, 72; pi. 19

Toxicodendron. .. ..... .............................. 45,46,74

Transition stage, lower Revenian.............. ............... 8

trichophorum, Dasymaschalon ........................... 67

trichophylla, Cinnamomum .................. ................. 89

trilobatum productum, Acer.......................... ........ 16

trilobus, Cocculus ......... . ........... ......... . .... .... 63

trinervis, MacClintockia .. . ........ ........ ... .... ..... 38

Trinity Islands .. . ............ ..... ..... ............. ........... 14

Trochodendraceae ..... .. . .... ...... ........ .. . 47, 80

Trochodendron ...... . ...... ... ...... . 43, 45, 46, 47

Trochodendroides arctica .. .......... ........ .................. 61

serrulata .... ... . ........... . . .............................. 61

zaddachi........... .... . ... ................. .... .. ..... 61

Tsadaka assemblage... ................................. ...... 54

Tsadaka Formation, Cook Inlet region ........ ... 13, 14

plant assemblage ......... .. ... 13, 14, 19, 46

Tubiflorae . .................. . .. ... . . 27, 34, 85

Tukwila Formation, Puget Group .................... ....... 8

Turcicula columbiana.. ..... . ............ .6

Turritella uvasana ........ .... . ... .5, 7

u

Ulmaceae .............. .. . .... 27, 31, 43,

Ulmus.............. .... . ... ............20, 32,

kushtakensis .... ........ .... ..... . ...

newberryi......... ............. ....... .. ....

(newberryi type) . ........... ........... ...

pseudobraunii ..... ... ............

sp ........................ .. ......... ..... 10,

Umbelliflorae .............. .. .. .. .... .....

Umbellularia ............ ...... ......

Unga Conglomerate Member, 

Bear Lake Formation ..... ....... . .....

marine mollusks . ........ .. ............

Unga Island, marine beds....... ... .. .. .. .

mollusks........ ... ................. ..........

United States, conterminous, Fagopsis 

groenlandica ... ... .... . ...

eastern, Celastrus.......... ... ...

Unknown unit . .. .. ..... ...... ............. ..

Unnamed marine siltstone and 

sandstone sequence ..........

age .. ................. .. ......

marine megafossils .......... ....

Unnamed stage, Alaskan . ................... .......

Upper Dui assemblage . ..... . ...... ....

Ural Mountains . . ....... ........... ......... ...

Urticales ....... ............ . . .... .........

uvasana, Turritella ...... . . ..... ........... . .

46, 47, 60

41, 45, 60

... . 31

.. ... 60

... ....... 44

.. 51,56 

60; pi. 23 

. 27, 85 

47, 48, 67

.. .. 20 

....13, 20 

.. 18, 20 

..... .. 17

75 

.. 7

...... 6

... ... 6

.... ...6

..... ..... 16

... . 42

...... 42

27, 31, 60 

. ... 5,1

Page

vaderensis, Tivelina .......................................................... 4

Vakhrameyev, V. A., quoted ........................................ 42

VauQuelinia ....................................................................... 71

Vegetation, Alaskan, lower Ravenian ...................... 31

Alaskan, Ravenian, latitude................................. 37

southern................................................................ 19

Australian types ....................................................... 29

Eocene ......................................................................... 50

fluviatile habitat........................................................ 24

lacustrine habitat.................................................... 24

late Ravenian ........................................................... 46

subtropical .......................................................... 42, 45

narrow-leaf.......................................................... 42

temperate...........................................42, 45, 46, 54, 58

Alaska ............................................................... 71

Oligocene............................................................ 46

Pacific Northwest............................................. 71

warm.................................................... 26, 43, 46

history ................... ..................................... 43

tropical..... .................................................. 42, 45, 70

Oligocene ........................................................ 46

zonation . .. .......... ........... ........................36,41

Vegetational analysis. .................... 20, 21, 23, 28, 32

Vegetational history, Neogene, latitudes, 

northern....... .......................................................... 42

velutina, Limacia ........... .......................................... 64

Verbenaceae............ ........ ..................27, 31, 34, 50, 85

Viburnum thomae ... ... ..... . ...... ........................... 91

Vinea... .. .. ... ......... .. .... .................................. 56

Vitaceae ... ............. ..... .. ............14, 27, 31, 50, 80

Vitis. . ... ............................44, 45, 48, 49, SO

atwoodi.................... ............................................... 14

rotundifolia . . ......... ..... ............ ......... ......... 14

sp ........... ..... ..................... 11, 27, 30, 80; pi. 15

Yokes, H. E., cited ............ .............. ......................... 6

volubilis, Pyrenacantha ..... ....................... ............ 78

w

wangi, Kandelia ... ..... .... . ..... ..11, 27, 83; pi. 19

washingtonensis, Perse ........... ............ .................. 5

washingtoniana, Whitneyella .......................... ...... 6

Weaverville assemblages, Oregon ........................ 88

Webb, L. J., quoted .......... .. .... .... ... .............. 29

weedi, Fortunearia .............. ... ........................ 89

weedii, Quercus . .. ........... . ............. .............. .. 89

West Indies, leaf-margin percentages ....... .... .. 25

Whitneyella coosensis. ............... ........... ............... 5

sinuata .... .......... ..... .................................. 5

aragoensis. . ... ..................... ............ 44

washingtoniana .... .. ..... .............. ..... .... 6

Willamette flora, radiometric age......... .... ..... 14, 15

williamsi, Tilia ...... ............... ....... ..... ....... .... 91

Willisia . ......... ............ ...................... 46, 81

obliquifolia .... . ...... .. ..... .................. - 93

rentonensis ........ ..... ............... ............ ... 81

wilsoni, Allophylus . ........... 11, 34, 35, 78, 90; pi. 29

Salix ... . .... ... .... ..... .... ....... ........ 55

Wind River Formation, Wyoming ... .. ................ 54

Wolfe, J. A., quoted .................. ......................... 41, 92

Woodwardia ...... . .................. ............... . ...... . 53

sp . ... . .... ... .. ....... ........10, 34, 53; pi. 28

wormskioldi, Magnolia.......... ............ .... - .... 52, 66

Wrangell Lava ..... ... .. .................... ..... 42

Wrangell Mountains ..... . .... . ...... . .. . . 42

X, Y

xantholithensis, Populus ... .... ....... . .............. 61

Xylonagra ........ .......... ............................ ........47,48

Yakataga district ................. ........ ........ .... 6

fossil collections . ........ ... .............. ..... - - 3

orocline. .... .. ....... .. .. . ..................... 36

Tertiary rocks ..... ........ . . ....... ...... .. 36

Yakataga Formation ...... ..... ........... . ........... 7, 13

leaves ... ............... ................. .... ...... 20

marine bed . . ........ ..... .... .......... ....... 7
108 INDEX

Page

Yakataga Formation Continued

marine megafossils .................................................. 20

Pinnacle Hills .................................................7, 13, 20

plant megafossils .............................................6, 7, 20

stratigraphic equivalent.......................................... 13

Yellowstone Park, Eocene beds..............................53, 89

Yukon River........................................................................ 54

Yunnan, Clethra delavayi ........................................... 85

yunnanensis, Zizypkus.... ............................................. 80

Page

z

zaddachi, Populus ............................................................ 61

Trochodendroides .................................................... 61

Zelkova.................................................................... 45, 48, 60

Zhilin, S. G., quoted......................................................... 43

Zizyphus ....................................................... 44, 45, 48, 80

alaskanus ....................................9, 11, 27, 80; pi. 14

californica ................................................................... 61

fungi ................................................ ...... ................. 80

inermis ........................................................................ 80

Page

Zizyph us Continued

microphyllus............................................................... 61

piperoides .................................................................... 61

serrulata........................................................................ 61

yunnanensis................................................................ 80

Zonation, altitudinal....................................................... 19

climatic.................................................................... 41, 50

floristic.................................................................... 19, 41

latitudinal.............................................................. 19, 41

vegetational ....................................................... 19, 41
PLATES 1 30

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

A, West side of Carbon Mountain; photograph taken from the lower slopes of Cunningham 

Ridge above locality 11162. Doughton Peak is to the left; locality 9891 is to the left of 

this peak. Most of the rocks shown are probably of middle and late Ravenian age.

B, Looking east along the crest of Carbon Mountain from locality 11169. Note the strong 

folding in the Kushtaka Formation.
GEOLOGICAL SURVFY PROFESSIONAL PAPER 997 PLATE 1

THE KUSHTAKA FORMATION ON CARBON MOUNTAIN
PLATE 2

[All figures natural size]

FIGURES 1, 2. Allantodiopsis pugetensis Wolfe. (p. 53.).

1. Hypotype, USNM 43251; loc. 11159.

2. Hypotype, USNM 38957; loc. 3879. 

3. Anemia gunni Wolfe. (p. 52).

Holotype, USNM 43246; loc. 11170.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 2

LOWER RAVENIAN FLORA
PLATE 3

[All figures natural size]

FIGURES 1, 4. Dryopteris alaskana (Holl.) Wolfe. (p. 53). 

1. Lectotype, USNM 38664; loc. 3879. 

4. Syntype, USNM 245724; loc. 3879.

2. Glyptostrobus sp. (p. 53). 

USNM 38724; loc. 3879.

3. Cyathea inequilateralis (Holl.) Wolfe. (p. 53). 

Holotype, USNM 38665; loc. 3846.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 3

LOWER RAVENIAN FLORA
PLATE 4

FIGURES 1-3. Alnus scotti Wolfe. (p. 59).

1. Paratype, USNM 43273; loc. 11170, X 1.

2. Paratype, USNM 43274; loc. 3847, X 1.

3. Holotype, USNM 38899; loc. 3847, X 1.

4. Platycarya pseudobrauni (Holl.) Wolfe. (p. 56). 

Hypotype, USNM 43266; loc. 11170, X 1.

5. Palmacites sp. (p. 54).

USNM 43258; loc. 11160, X V6.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 4 

*?

LOWER RAVENIAN FLORA
PLATE 5

[All figures natural size]

FIGURES 1, 2. Platycarya pseudobrauni (Holl.) Wolfe. (p. 56).

1. Hypotype, USNM 43267; loc. 11170.

2. Hypotype, USNM 38893; loc. 3847. 

3, 5. Alnus martini Wolfe. (p. 58).

1. Holotype, USNM 43271; loc. LI 158.

2. Paratype, USNM 43272; loc. 11170. 

4. Tetracentron piperoides (Lesq.) Wolfe. (p. 61). 

Hypotype, USNM 43288; loc. 11170.

6. Myristica sp. (p. 66).

USNM 39019; loc. 3847.

7. Girroniera sp. (p. 60).

USNM 43285; loc. 11158.

8. Girroniera sp. (p. 60).

USNM 43286; loc. 11170.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 5

LOWER RAVENIAN FLORA
PLATE 6

[All figures natural size]

FIGURES 1-3. Limacia stenophylla Wolfe. (p. 64).

1. Holotype, USNM 43303; loc. 11159. 

2, 3. Paratypes, USNM 43304, 43305; loc. 11159.

4. Pycnarrhena sp. (p. 65). 

USNM 43308; loc. 11166.

5. CocculuK sp. (p. 63).

USNM 43299; loc. 11166.

6. Anamirta milleri Wolfe (p. 62).

Holotype, USNM 43293; loc. 11170.

7. Diploclisia sp. (p. 63).

USNM 43301; loc. 11170.

8. Calkinsia plafkeri Wolfe. (p. 62).

Paratype, USNM 43294; loc. 11157.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 6

LOWER RAVENIAN FLORA
PLATE 7

(All figures natural size]

FIGURES 1-3, 6. Clcrodendrum sp. (p. 85).

USNM 43399, 43400, 43401 A, 43402; loc. 11158. 

4, 7. Limacia stenophylla Wolfe. (p. 64).

4. Paratype, USNM 43306; loc. 11159. 

7. Paratype, USNM 43307; loc. 11166. 

5. Paratinomiscium conditionalis (Holl.) Wolfe. (p. 65). 

Holotype, USNM 39117; loc. 3879.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 7

LOWER RAVENIAN FLORA
PLATE 8

[All figures natural size]

FIGURE 1. Myristica sp. (p. 66).

USNM 38667; loc. 3847. 

2. Cinnamomophyllum latum (MacG.) Wolfe. (p. 67).

Hypotype, USNM 38914; loc. 3847. 

3, 4. Knema sp. (p. 66).

USNM 38915, 43309; loc. 3847.

5. Dasymaschalortf sp. (p. 66). 

USNM 43310; loc. 11158.

6. Cananga sp. (p. 66).

USNM 38911; loc. 3879.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 8

LOWER RAVENIAN FLORA
PLATE 9

[All figures natural size]

FIGURES 1, 4. Litseaphyllum similis (Knowlt.) Wolfe. (p. 69). 

Hypotypes, USNM 43320, 43321; loc. 11157. 

2. Litseaphyllum presamarensis (Sanb.) Wolfe. (p. 68).

Hypotype, USNM 43325; loc. 3846.

3, 10. Cinnamomophyllum latum (MacG.) Wolfe. (p. 67). 

3. Hypotype, USNM 43314; loc. 11166. 

10. Hypotype, USNM 43315; loc. 11157. 

5, 8, 9. Euodia alaskana Wolfe. (p. 73).

5. Holotype, USNM 43349; loc. 11158. 

8, 9. Paratypes, USNM 43350, 43351; loc. 11158.

6. Caesalpinites sp. (p. 72). 

USNM 43347; loc. 3847.

7. Litseaphyllum sp. (p. 69). 

USNM 43335; loc. 11159.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 9
PLATE 10

[All figures natural size]

FIGURES 1, 4. Luvunga spatiosa (Holl.) Wolfe. (p. 73). 

1. Hypotype, USNM 43352; loc. 3847. 

4. Holotype, USNM 38920; loc. 3847. 

2, 3, 5. Melanorrhoca alaskana (Holl.) Wolfe. (p. 74).

2. Hypotype, USNM 38885; loc. 3846.

3. Hypotype, USNM 43353; loc. 11158. 

5. Holotype, USNM 39020; loc. 3847.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 10

LOWER RAVENIAN FLORA
PLATE 11

[All figures natural sizej

FIGURES 1, 4-6. Celastrus comparabilis Holl. (p. 75).

1. Hypotype, USNM 43356; loc. 11158.

4. Hypotype, USNM 43357; loc. 11170.

5. Hypotype, USNM 38744; loc. 3847.

6. Holotype, USNM 38956; loc. 3879.

2. Pyrenacantha sp. (p. 77). 

USNM 43366; loc. 11170.

3. Goweria dilleri (Knowlt.) Wolfe. (p. 76). 

Hypotype, USNM 38901; loc. 3847.
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LOWER RAVENIAN FLORA
PLATE 12

[All figures natural size]

FIGURES 1-6. Phytocrene acutissima Wolfe. (p. 76).

1-3, 5. Paratypes, USNM 43359-43361, 43363; loc. 11158. 

4, 6. Holotype, USNM 43362A, B; loc. 11158.
PROFESSIONAL PAPER 997 PLATE 12

LOWER RAVENIAN FLORA
PLATE 13

[All figures natural size]

FIGURE 1. Phytocrene acutissima Wolfe. (p. 76). 

Paratype, USNM 43364; loc. 11166.

2. Paleophytocrene elytraeformis (Holl.) Wolfe. (p. 76). 

Holotype, USNM 39127A; loc. 3879.

3. Goweria alaskana Wolfe. (p. 75).

Holotype, USNM 43358A; loc. 11158.

4. Phytocrene sordida (Lesq.) MacG. (p. 77).

Hypotype, USNM 43365; loc. 11170. 

5, 6. Stemonurun alaskanus Wolfe. (p. 78).

5. Paratype, USNM 43367; loc. 11158.

6. Holotype, USNM 43368; loc. 11158.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 13

LOWER RAVENIAN FLORA
PLATE 14

[All figures natural size]

FIGURES 1, 2. Meliosma kushtakensis Wolfe. (p. 79).

1. Paratype, USNM 43381; loc. 11158.

2. Holotype, USNM 43382; loc. 11166. 

3, 4. Allophylus duktothensis Wolfe. (p. 78).

3. Holotype, USNM 43372; loc. 11170.

4. Paratype, USNM 43373; loc. 11170. 

5, 9. Zizyphus alaskanus Wolfe. (p. 80).

Holotype, USNM 43371A, B; loc. 11158. 

6-8. Meliosma duktothensis Wolfe. (p. 79).

6. Holotype, USNM 43378; loc. 11170. 

7, 8. Paratypes, USNM 43379, 43380; loc. 11170.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 14

LOWER RAVENIAN FLORA
PLATE 15

| All figures natural size]

FIGURE 1. Sageretia sp. (p. 80).

USNM 43370; loc. 11166. 

2, 4, 5. Vitis sp (p. 80).

USNM 43383-43385; loc. 11166. 

3. Illicium sp. (p. 62).

USNM 43292; loc. 11159. 

6. Mastixia irregularis (Holl.) Wolfe. (p. 85). 

Holotype, USNM 39033; loc. 3846.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 15

LOWER RAVENIAN FLORA
PLATE 16

[All figures natural size]

FIGURE 1. Alangium sp. (p. 84).

USNM 43393; loc. 11166. 

2. Alangium bergensis Wolfe. (p. 84).

Holotype, USNM 38926; loc. 3847. 

3-5. Eugenia sp. (p. 84).

USNM 43394-43396; loc. 11170. 

6-8. Parashorea pseudogoldiana (Holl.) Wolfe. (p. 82).

6. Hypotype, USNM 39035; loc. 3847.

7. Lectotype, USNM 38978; loc. 3847.

8. Syntype, USNM 43391; loc. 3847.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 16
PLATE 17

[All figures natural size]

FIGURE 1. Clerodendrum sp. (p. 85).

USNM 43401B; loc. 11158. 

2. Dicotylophyllum sp. (p. 86). 

USNM 43414; loc. 11170. 

3, 4. Barringtonia sp. (p. 83).

USNM 43392A, B; loc. 11170.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 17

LOWER RAVENIAN FLORA
PLATE 18

[All figures natural size]

FIGURE 1. Saurauia sp. (p. 81).

USNM 43389; loc. 3846.

2. Clethra sp. (p. 85).

USNM 43398; loc. 11170.

3. Dicotylophyllum sp. (p. 86). 

USNM 39036; loc. 3847.

4. Celastrus comparabilis Holl. (p. 75). 

Hypotype, USNM 245725; loc. 11170.

5. Dicotylophyllum sp. (p. 86). 

USNM 43415; loc. 11159.

6. Dicotylophyllum sp. (p. 86). 

USNM 43416; loc. 11158.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 18

LOWER RAVENIAN FLORA
PLATE 19

[All figures natural size]

FIGURES 1, 5a,b. Kandelia wangi Wolfe. (p. 83).

1. Holotype, USNM 222843; loc. 11195, X 5. 

5a. Paratype, USNM 222844; loc. 11195, X 1. 

5b. Holotype, USNM 222843; loc. 11195, X 1.

2. Kandelia candel (L.) Druce. Extant species for comparison. 

USGS Ref. Colln. 1752. X 5.

3. Rhus cf. R. mixta (Lesq.) MacG. (p. 74). 

USNM 222840; loc. 11195, X 1.

4. Toona sp. (p. 72).

USNM 222842; loc. 11195, X 1.

6. Parashorea pseudogoldiana (Holl.) Wolfe. (p. 82). 

Hypotype, USNM 222841; loc. 11195, X 5.

7. Parashorea malaanonan (Blanco) Vid. Extant species for comparison. 

USGS Ref. Colln. 1663, X 5.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 19

LOWER RAVENIAN FLORA
PLATE 20

[All figures natural size]

FIGURES 1, 2. Lygodium larsoni Wolfe. (p. 52).

1. Holotype, USNM 43248; loc. 9389.

2. Paratype, USNM 43249; loc. 11165. 

3, 6. Glyptostrobus sp. (p. 53).

USNM 43254, 43255; loc, 11165.

4. Allantodiopsis pugetensis Wolfe. (p. 53). 

Hypotype, USNM 43252; loc. 11165.

5. Phoenicites sp. (p. 53).

USNM 43257; loc. 11165.

7. Engelhardtia sp. (p. 55). 

USNM 43264; loc. 11165.

8. Carya cashmanensis Wolfe. (p. 55). 

Hypotype, USNM 43263; loc. 11165.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 20

MIDDLE RAVENIAN FLORA
PLATE 21

[All figures natural size]

FIGURES 1, 5, 6. Dryophyllum pugetensis Wolfe. (p. 60).

1, 6. Hypotypes, USNM 43281, 43283; loc. 11165.

5. Hypotype, USNM 43282; loc. 3842. 

2a. Glyptostrobus sp. (p. 53).

USNM 43256; loc. 11165.

2b. Platycarya pseudobrauni (Holl.) Wolfe. (p. 56). 

Hypotype, USNM 245726; loc. 11165.

3. Alnus martini Wolfe. (p. 58).

Paratype, USNM 43279; loc. 11163.

4. Pterocarya pugetensis Wolfe. (p. 56). 

Hypotype, USNM 43268; loc. 11165.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 21

MIDDLE RAVENIAN FLORA
PLATE 22

(All figures natural size]

FIGURES 1, 2. Litseaphyllum presamarensis (Sanb.) Wolfe. (p. 68). 

Hypotypes, USNM 43326, 43327; loc. 3842.

3. Tetracentron piperoides (Lesq.) Wolfe. (p. 61). 

Hypotype, USNM 43289; loc. 11165.

4. Litseaphyllum sp. (p. 69). 

USNM 43338; loc. 11165.

5. Litseaphyllum sp. (p. 69). 

USNM 43337; loc. 11165.

6. Litseaphyllum praelingue (Sanb.) Wolfe. (p. 68). 

USNM 43324; loc. 11165.
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PLATE 23

[All figures natural size|

FIGURE 1. Ulmus sp (p. 60).

USNM 43287; loc. 11165. 

2. Litseaphyllum similis (Knowlt.) Wolfe. (p. 69).

Hypotype, USNM 43322; loc. 11165. 

3, 4, 6, 7b. Calkinsia plafkcri Wolfe. (p. 62).

3, 6, 7b. Paratypes, USNM 43295, 43297, 43298; loc. 11165.

4. Paratype, USNM 43312; loc. 11165. 

5, 7a. Cinnamomophyllum kushtakensis Wolfe. (p. 67).

5. Holotype, USNM 43311; loc. 9389. 

7a. Paratype, USNM 43312; loc. 11165.
PROFESSIONAL PAPER 997 PLATE 23

MIDDLE RAVENIAN FLORA
PLATE 24

[All figures natural size]

FIGURE 1. Liquidambar? sp. (p. 70). 

USNM 43342; loc. 11165.

2. Dicotylophyllum sp. (p. 86). 

USNM 43417; loc. 11165.

3. Allophylus sp. (p. 79).

USNM 43376; loc. 11165.

4. Dicotylophyllum sp. (p. 87). 

USNM 43418; loc. 11165.

5. Dicotylophyllum sp. (p. 87). 

USNM 43420; loc. 11165.

6. Dicotylophyllum sp. (p. 87). 

USNM 43419; loc. 9389.

7. Eucommia sp. (p. 70).

USNM 43343; loc. 11165.

8. Dicotylophyllum sp. (p. 87). 

USNM 43421; loc. 11165.

9. Sapindus? sp. (p. 79).

USNM 43377; loc. 11165.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 24

MIDDLE RAVENIAN FLORA
PLATE 25

[All figures natural size]

FIGURES 1, 2, 6. Litseaphyllum schorni Wolfe. (p. 68).

1, 6. Paratypes, USNM 43332, 43334; loc. 11165. 

2. Holotype, USNM 43333; loc. 11165.

3. Theaceae, gen. and sp. indetermined. (p. 82). 

USNM 43390; loc. 11165.

4. Anacardiaceae, gen. and sp. indetermined. (p. 74). 

USNM 43354; loc. 11165.

5. Caesalpinites sp. (p. 72). 

USNM 43348; loc. 11165.

7. Dicotylophyllum kummerensis (Wolfe) Wolfe. (p. 86). 

Hypotype, USNM 43412; loc. 11164.

8. Macclintockia sp. (p. 86). 

USNM 43409; loc. 11164.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 25

MIDDLE RAVENIAN FLORA
PLATE 26

[All figures natural size]

FIGURE 1. Populus sp. (p. 54).

USNM 43259; loc. 11167.

2. Salix sp. (p. 55).

USNM 43262; loc. 11167.

3. Juglans sp. (p. 56).

USNM 43265; loc. 11167. 

4, 10. Alnus sp. (p. 59).

USNM 43275; 43276a; loc. 11167.

5. Cocculus sp. (p. 63).

USNM 43300; loc. 11167.

6. Corylopsis sp. (p. 70).

USNM 43341A; loc. 11167.

7. Alnus cuprovallis Axelr. (p. 58).

Hypotype, USNM 43270; loc. 11167.

8. Dicotylophyllum sp. (p. 87). 

USNM 43430; loc. 11167.

9. Prunus axelrodi Wolfe. (p. 71).

Holotype, USNM 43345A; loc. 11167. 

11, 13. Salix carbonensis Wolfe. (p. 55).

11. Paratype, USNM 43261; loc. 11167. 

13. Holotype, USNM 43260; loc. 11167. 

12. Ilex carbonensis Wolfe. (p. 74).

Holotype, USNM 43355A; loc. 11167.
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UPPER RAVENIAN FLORA
PLATE 27

[All figures natural size]

FIGURES 1, 5. Tilia carbonensis Wolfe. (p. 80).

1. Holotype, USNM 43386A; loc. 11167. 

5. Paratype, USNM 43387; loc. 11167. 

2. Rhamnus sp. (p. 79).

USNM 43369; loc. 11167. 

3, 4. Sorbus carbonensis Wolfe. (p. 71).

Holotype, USNM 43346A, B; loc. 11167.

6. Plafkeria sp. (p. 81).

USNM 43388A; loc. 11167.

7. Eugenia? sp. (p. 85).

USNM 43397; loc. 11167.

8, 10. Dicotylophyllum carbonensis Wolfe. (p. 86). 

8. Holotype, USNM 43410; loc. 11167. 

10. Paratype, USNM 43411; loc. 11167. 

9. Dicotylophyllum sp. (p. 87). 

USNM 43422; loc. 11167.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 27

UPPER RAVENIAN FEORA
PLATE 28

[All figures natural size]

FIGURE 1. Woodwardia sp. (p. 53).

USNM 43253; loc. 9891.

2. Fagus sp. (p. 60).

USNM 43284; loc. 9552.

3. Alnus sp. (p. 59).

USNM 43278; loc. 9891.

4. Betula sp. (p. 60).

USNM 43280; loc. 11168.

5. Anemia? sp. (p. 52).

USNM 43247; loc. 9891.

6, 9. Litseaphyllum carbonensis Wolfe. (p. 68). 

6. Paratype, USNM 43316; loc. 11169. 

9. Holotype, USNM 43317; loc. 11169. 

7. Tetracentron piperoides (Lesq.) Wolfe. (p. 61).

Hypotype, USNM 43290; loc. 11169. 

8a. Litseaphyllum sp. (p. 69).

USNM 43339; loc. 11169.

8b, 10a,b. Litseaphyllum presamarensis (Sanb.) Wolfe. (p. 68). 

Hypotypes, USNM 43328-43330; loc. 11169.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 28

KUMMERIAN FLORA
PLATE 29

[All figures natural size]

FIGURES 1, 3. Litseaphyllum katallaensis Wolfe. (p. 68). 

1. Paratype, USNM 43318; loc. 9551.

3. Holotype, USNM 43319; loc. 11168. 

2. Litseaphyllum similis (Knowlt.) Wolfe. (p. 69).

Hypotype, USNM 43323; loc. 11169. 

4, 6. Allophylus wilsoni Chan. and Sanb. (p. 78).

4. Hypotype, USNM 43374A; loc. 11168. 

6. Hypotype, USNM 43375; loc. 9553. 

5. Litseaphyllum sp. (p. 69). 

USNM 43336; loc. 11168.

7. Platanus comstocki (Sanb.) Wolfe. (p. 71). 

Hypotype, USNM 43344; loc. 9891.

8. Litseaphyllum sp. (p. 70). 

USNM 43340; loc. 9891.

9. Litseaphyllum presanguinea (Chan. and Sanb.) Wolfe. (p. 68). 

Hypotype, USNM 43331; loc. 9891.
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KUMMERIAN FLORA
PLATE 30

[All figures natural size]

FIGURE 1. 'Dicotylophyllum sp. (p. 87). 

USNM 43423; loc. 11168.

2. Dicotylophyllum sp. (p. 87). 

USNM 43424; loc. 11169.

3. Dicotylophyllum sp. (p. 87). 

USNM 43425; loc. 11168.

4. Holmskioldia speiri (Lesq.) MacG.? (p. 85).

USNM 43403; loc. 11168.

5, 6, 8-10. Macclintockia pugetensis Wolfe. (p. 86). 

5. Hypotype, USNM 43404; loc. 9891. 

6, 10. Hypotypes, USNM 43405, 43408; loc. 11168. 

8, 9. Hypotypes, USNM 43406, 43407; loc. 11189. 

7. Magnolia reticulata Chan. and Sanb. (p. 62).

Hypotype, USNM 43291; loc. 11168. 

11. Dicotylophyllum kummerensis Wolfe?. (p. 86). 

USNM 43413; loc. 9891.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 997 PLATE 30

11

KUMMERIAN FLORA