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Graduate Student Theses, Dissertations, & Professional Papers Graduate School

1965

A survey of distribution patterns in the Montana alpine flora

Richard H. Pemble The University of Montana

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Recommended Citation Pemble, Richard H., "A survey of distribution patterns in the Montana alpine flora" (1965). Graduate Student Theses, Dissertations, & Professional Papers. 6680. https://scholarworks.umt.edu/etd/6680

This Thesis is brought to you for free and open access by the Graduate School at ScholarWorks at University of Montana. It has been accepted for inclusion in Graduate Student Theses, Dissertations, & Professional Papers by an authorized administrator of ScholarWorks at University of Montana. For more information, please contact [email protected]. A SURVEY OF DISTRIBUTION PATTERNS IN THE

MONTANA ALPINE FLORA

by

RICHARD HOPPE PEMBLE

B.A. Simpson College, 196j

Presented in partial fulfillment of the requirements for the degree of

Master of Arts

UNIVERSITY OF MONTANA

1965

Approved by:

Chairman, Bracrd of Examiners\

Dean, Graduate School

AUG ' *• Date

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ACKNOWLEDGEMENT

I wish to express my fullest appreciation to Dr. L.H. Harvey who

gave so freely of his time to help me in all phases of this study. I

should also like to acknowledge the help of Drs. R.S. Hoffmann, J. R«

Habeck, and S.J. Preece who served as members of my examining committee

and were kind enough to critically read this thesis prior to the writ­

ing of a final draft. My appreciation is also extended to Dr. W.E.

Booth who gave me unrestricted access to the herbarium of Montana State

University during my time spent there. And last, but not least, I

should like to acknowledge the help and encouragement of my wife, Helen,

who took over the many 'secretarial duties' involved with writing this

thesis.

I should also like to acknowledge the National Science Foundation

summer fellowship program for partial financial support during this

study.

•ii—

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. TABLE OF CONTENTS

PAGE

INTRODUCTION...... 1

Physiography of Montana...... 3

Past and Present Climate of Montana ...... 6

Description of the Alpine Zone ...... 10

T i m b e r l i n e ...... 12

Ecology of the alpine z o n e ...... 13

Definition of an Alpine Plant...... U 4

Floristic Elements ...... , 1^

PROCEDURE...... 18

Determination of Altitudinal Limits of Alpine Areas in Montana 18

Field Studies...... 18

Herbarium Material Studied ...... 19

Distribution Maps ...... 21

FLORISTIC ELEMENTS IN THE MONTANA ALPINE FLORA 2h

The Lowland Element ...... 2^

The Arc tic-alpine Element...... 26

Arctic-alpine species which are fully circumpolar or

nearly so ...... 26

Arctic-alpine species which are widely distributed in the

North American Arctic and that of eastern or western Eurasia, 35

Arctic-alpine species restricted to North American Arctic or

found, in addition, only on the Pacific Coast of Eurasia

and/or in Greenland......

-iii-

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PAGE

Cordllleran Element. hi

Species with a wide range in the western Cordilleras <> . . . » U8

Species restricted to Rocky Mountains...... 6l

Pacific Coast Element...... 68

Endemic s ...... o...... *..... ^ 2!

DISCUSSION ...... 80

SUMMARY...... 89

LITERATURE CITED ...... 91

APPENDIX I List of species constituting the lowland element

in the Montana alpine flora...... 96

APPENDIX II Distribution of species occurring in the

Montana alpine ...... 103

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LIST OF FIGURES

FIGURE PAGE

1. Maximum Glaciation in Montana During the Pleistocene. . . • . 11

2. Mountainous Montana ...... 23

3# Erigeron humilis Graham. The Montana Alpine Distribution of

a Gircumpolar Arctic-alpine Species Reaching its Southern

Limits in the Rocky Mountains in Northwest Montana. . . . 28

ii* Average July Temperatures in Montana...... 29

5. Gassiope tetragona (L.) D.Don var. saximontana (Small) C.L.

Hitchc, The Montana Alpine Distribution of a Circumpolar

Arctic-alpine Species Reaching its Southern Limits in

the Rocky Mountains in Extreme West-central Montana . • 31

6 . Saxifraga oppositifolia L, The Montana Alpine Distribution of

a Circumpolar Arctic-alpine Species Reaching its Southern

Limit in the Rocky Mountains in the Beartooth Plateau Area 33

7. Festuca brachyphylla Schulte. A Circumpolar Arctic-alpine

Species Reaching its Southern Limit in the Rocky Mountains

xn New Mexxc o...... 3^

8. Androsace lehmanniana Spreng. The Montana Alpine Distribution

of an Arctic-alpine Species Widely Distributed in the North

American Arctic and that of Western Eurasia and Which

Reaches its Southern Rocky Mountain Limit in North-

central Montana ...... o*...»*. 3^

-V-

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FIGURE PAGE

9» Pedicularis oederi Vahl, The Montana Alpine Distribution

of an Arctic-alpine Species Widely Distributed in the

North American Arctic and Western Eurasia Which Reaches

its Southern Limits in the Rocky Mountains in the

Beartooth Plateau Area...... 38

10. Eritrichium nanum (Vill.) Schrad, var. elongatum(Rydb.)Gronq.

The Montana Alpine Distribution of an Arctic-alpine Species

Widely Distributed in the North American Arctic and West­

ern Eurasia Which Reaches its Southern Limits in the

Rocky Mountains in New M e x i c o ...... liO

11. Agropyron latiglume (Scribn.&Smith)Rydb. The Montana Alpine

Distribution of an Arctic-alpine Species Restricted in the

Arctic to North America and Greenland ...... UU

12. Trisetum spicatum (L,) Richt. The Montana Alpine Distribu­

tion of an Arctic-alpine Species Which is Widely Distribu­

ted Including Stations in Both the Northern and Southern

Hemisphere as well as Alpine Areas in Europe and Asia . U7

13. General Distribution of the Cordilleran Element Widely

Distributed from Alaska Along the Rocky Mountains, in the

Cascades and Sierra Nevadas but not in the Coast Ranges

of California ...... h9

lii, Fhyllodoce empetriformis (Sw.) D.Don. The Montana Alpine

Distribution of the Cordilleran Element Widely Distribu­

ted from Alaska Along the Rocky Mountains, in the Cas­

cades and Sierra Nevadas but not in the Coast Ranges

of California ...... ^0

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1$, The General Distribution of the Cordilleran Element Widely

Distributed from Alaska Along the Rocky Mountains, Sierra

Nevadas, the Coast Ranges of California and Sometimes

Extending into the Cascades ...... ^2

16 . Cassiope mertensiana (Bong.) D.Don var. gracilis (Piper)

C.L.Hitchc, The Montana Alpine Distribution of the

Cordilleran Element Widely Distributed from Alaska

Along the Rocky Mountains, the Sierra Nevadas, the

Coast Ranges of California, and the Cascades. .... 53

17. The General Distribution of the Cordilleran Element Widely

Distributed from British Columbia and/or Alberta Along

the Rocky Mountains and Sierra Nevadas, sometimes ex­

tending into the Cascades but not the Coast Ranges

of California ...... 55

1 8 . Senecio fremontii T. & G. var, fremontii. The Montana

Alpine Distribution of the Cordilleran Element Widely

Distributed in the Rocky Mountains from British

Columbia and Alberta Along the Rocky Mountains and

Sierra Nevadas but not in the Coast Ranges of

California, ...... 56

1 9 . General Distribution of the Members of the Cordilleran

Element Which are Distributed in the Rocky Mountains

and the Sierras but not in Coast Ranges of California,

and Reach their Northern Limits in Southwestern and

South-central Montana...... 59

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FIGURE PAGE

20, Astragalus tegetarlus Wats. A Representative Montana DiS“

tribution of Members of the Cordilleran Element Which

are Distributed in the Rocky Mountains and the Sierras

but not in the Coast Ranges of California, and Reach

their Northern Limits in Southwestern and South-

central Montana...... 60

21, General Distribution of Cordilleran E]e ment Restricted to and

Widespread in the Rocky Mountains, 62

22. Erigeron simplex Greene. The Montana Alpine Distribution

of a Species Restricted to and Widespread in the Rocky

Mountains, a,,,,».»...... 63

23. General Distribution of Cordilleran Element Restricted to

the Rocky Mountains with Northern Limits in Southwestern

or West-central Montana...... 6^

2I4, Mertensia alpina (Torr.) G.Don. The Montana Alpine Distri­

bution of a Species Restricted to the Rocky Mountains

Which Reaches its Northern Limits in West-central Mon­

tana . 0 . 0 ...... **..* 66

25* General Distribution of Pacific Coast Element in Montana

Alpine Flora ...... o.*...... 6 ^

26» Eriogonum pyrolaefolium Hook. var. coryphaeum T. & G, The

Montana Alpine Distribution of a Species in the Pacific

Coast Element ^1

27 « General Distribution of Species Endemic to the Yellowstone

R e g i o n ...... ?li

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FIGURE PAGE

28, Erigeron rydbergii Gronq, The Montana Alpine Distribution

of a Species Endemic to the Yellowstone Region. » » . 75

29. General Distribution of Species Endemic to Western Montana

and Adjacent Areas, ...... 77

30, Stellaria americana (Porter) Standi, The Montana Alpine

Distribution of a Species Endemic to Western Montana

and Adjacent Areas...... 78

31. Aquilegia jonesii Parry, The Montana Alpine Distribution

of a Species Endemic to Western Montana and Adjacent

Areas and Further Restricted to Calcareous Areas. .... 79

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. INTRODUCTION

Biologists harve long ii'ecognized that the alpine and arctic timber-

lines form one of the outstanding borders between major vegetation types

and as a result these, as well as the region above timberline, the

arctic and alpine tundras, have been extensively studied. In the Rocky

Mountain region the alpine areas of Colorado, where in certain localities

there are hundreds of acres above timberline, have been intensively

studied (Marr, 19^8, 1959, 1961; Weber, 19^9, 1959, 1961). A second

Rocky Mountain alpine location which has been studied is the Beartooth

Plateau of northern Wyoming and southwestern Montana, an area which is

also large and easily accessible (Billings and Mooney, 1959; Bliss, 1956?

Johnson, 1962; Johnson and Billings, 1962). Among the earliest and most

comprehensive studies were those of Rydberg (1913, 19lii)o He included

in his series of publications on the phytogeography of the Rocky Moun­

tains three articles on the phytogeography of the Rocky Mountain alpine»

His observations which are relevant to the Montana alpine were based on

a limited number of collections primarily from the Yellowstone Park

region of Montana. An early study which dealt with the alpine flora

of Montana was that of Hawkins (1903), but his collections were limited

to south-central Montana.

Only three recent studies have been made which have been concerned

with alpine vegetation in Montsina, excluding those of the Beartooth

Plateau area. A longterm study undertaken by Drs. R.S. Hoffmann, R.D.

Taber, and T.J» Nimlos of the University of Montana and Samuel Bamberg

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of the University of Colorado and later of the University of California^

Davis (Bamberg; 1961, 1963; NimloS; 1962; Taber^ Hoffmann^ Nimlos, and

Bamberg; 1961), included seven study areas widely distributed through

west-central Montana, principally east of the Continental Divide*. Their

study was designed to determine and describe the principal plant commu­

nities, the associated vertebrate animal communities, and the soil types

of Montana alpine areas, Bamberg, in his report on the plant communities,

included a list of the species encountered in his study and made some ob­

servations concerning the floristic elements present in the Montana al»

pine flora. The second recent alpine vegetation study in Montana was

that of Choate (1963) which resulted in an ordination of the alpine veg­

etation of Logan Pass, Glacier National Park, The third study was a

floristic one of the Logan Pass area in Glacier National Park by Sammons,

It was not completed because of her death. The collections are deposited

in the herbarium of the Ikiiversity of Montana,

No work has been done which was specifically aimed at determining

the distribution of the species present in the alpine flora of the

state. The only recent works indicating distribution of Montana alpine

species, Hitchcock et al, (1955; 1959, 1961, 196U), describe species'

ranges in the Pacific Northwest, but for the most part these are of such

a general nature that they do not reveal true distributions. Booth

(1959) gives species' ranges based on collections in the herbarium at

Montana State University, Bozeman, which indicate occurrences by coun­

ties only.

Within Montana a wide variety of alpine environments are available

and for this reason the distribution of species within the alpine areas

of the state may be expected to be quite complex. The literature

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. available indicates the lack of information in this area and a need for

a study of the type which has been undertaken*

The purpose of my study has been twofold* First, I have tried to

determine as completely as possible the range of the alpine species

present in the flora of Montana as shown by collections in the following

herbaria: Ifriiversity of Montana (Missoula, Montana)j Montana State l&ii-

versity (Bozeman, Montana); Region One, United States Forest Service;

and Glacier National Park and by personal collecting in selected areas*

Second, using distribution maps b^sed on these collections and, in a

few instances, published reports I have tried to determine the floristic

elements in the Montana alpine flora, to show their total distribution

pattern, and to analyze their distribution in Montana*

Physiography of Montana

In order to fully appreciate the distribution of alpine species in

Montana an understanding of the present physiography as well as the

past history of the mountainous areas of the state is important* The

following discussion of the physiography of Montana is adapted from

Alden (19^3) and Perry (I962) who should be consulted if further infor­

mation on the physiography and geology of the state is desired*

The Rocky Mountains of today have undergone at least two successive

periods of mountain making* Earlier mountains developed in the Precam-

brian time have been eroded and destroyed and are not of much signifi­

cance in the structure of the present Rocky Mountains in Montana*

Beginning in the late Precambrian, layers of sand, mud, and limy

secretions were deposited which later turned into sandstone, shale, and

limestone* These sediments came from ancient seas which once spread

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entirely across this region. The former were deposited on the Pre­

cambrian complex and are known as the Belt Series, Sedimentation con­

tinued intermittently throughout most of the Paleozoic and until the

late Cretaceous of the Mesozoic With little folding or uplift occurring.

At the end of the Cretaceous compressive forces wrinkled the strata

into great folds miles in length and width. This folding process con­

tinued into the Cenozoic Era but it reached its climax at the end of the

Cretaceous. As a result of later developmental changes the exact posi­

tion of the first mountain ridges and valleys is not known. However^

before the first developmental change took place late in the Cretaceous^

volcanic activity broke forth in a long narrow belt from near Yellow­

stone Park almost to Augusta, Lewis and Clark County, a distance of

about 150 miles. Lava poured onto the horizontal sediments of the late

Cretaceous, These later became folded and faulted by the Rocky Mountain

deformation. The uplifted areas were quickly eroded, nearly as fast as

they were uplifted.

The forces of compression slackened as the Cenozoic Era progressed.

In the early Tertiary, Cretaceous conditions still were prevalent in

eastern Montana and the results were basin deposits of plant material

covered by sand and mud.

After the large folds of the Cretaceous occurred, deep-seated

igneous activity resulted in the intrusion of liquid rock which later

cooled and solidified, forming batholiths. Nearly all batholiths are

granitic in composition (Perry, 1962), The Bitterroot Range southwest

of Missoula is part of the large granitic Idaho batholith. Following

the faulting which formed this range, a long period of erosion of the

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. higher sediments resulted in the exposure of the granite« According to

Perry (1962), nearly ninety percent or more of the batholitic intrusions

lie in southwestern Montana^ south of a line from Missoula to Helena^

and west of Bozeman,

The present-day Rockies in western Montana were the result of block

faulting which was caused by forces acting vertically rather than hori™

zontally# In certain areas large sections of the fault block were

shifted hundreds of feet higher than adjacent segments. While some

block-fault movement may have occurred during the Pleistocene5 most is

believed to have taken place during the Tertiary (Perry^ 1962). This

process resulted in the main pattern of our present-day mountains#

The Rocky Mountains^ occupying about one-third of the area of the

state, sprawl across the western portion in long, generally parallel,

northwest-southeast trending chains. These consist of forty or more

individual mountain ridges which are generally termed ranges. Often the

term range is misleading because definite borders between mountain groups

are lacking# In this paper and on the distribution maps many smaller

•ranges • are indicated since their names are locally used and often

found on herbarium labels# Their use gives a more definite location in

an otherwise much more encompassing mountain range.

Half of the state has an elevation in excess of 5000 feet, however,

the mean altitude of 3,^00 feet is the lowest for the Rocky Mountain

States (Montana Almanac, 1958)# Height above sea level of Montana’s

mountains averages from 6OOO feet in northwestern Montana to about 10,000

feet on the Beartooth Plateau in southwestern Montana, Granite Mountain,

which lies near the Beartooth Plateau, is the highest peak in Montana

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. with an elevation of 12,799 feet»

The isolated island-like mountains of central Montana, somewhat

younger than the folded mountains of the Rockies, are of four different

types: (1 ) vertical uplifts, (2 ) intrusions of igneous rock, (3 ) piles

of lava, and (U) erosion remnants of an elevated area (Perry, 1962)»

Mountains caused by vertical uplift include the Big Horn (Pryor), the

Big Snowys, Little Belt, and the Beartooth Plateau» Those mountains of

central Montana caused by intrusions of igneous rock are the Judith,

Moccasin, Little Rockies, Crazy, and, in part, the Bearpaw Mountains»

These lavas bear no relationship, as to the origin, to the late Cretaceous

and Tertiary lavas in the mountains of southwestern Montana (Perry, 1962)*

Fast and Present Climate of Montana

"Climate is the most far-reaching of the natural elements control­

ling plant life" (Polunin, I960)» Many alpine species illustrate this

in that they require the cooler temperatures afforded by high altitudes

in order to persist. When these species are transplanted to areas of

warmer temperatures, even where there is no competition, they only sur­

vive for a short time.

In the geological past climatic changes have occurred and these

have been significant in their influence on the present distribution of

vegetation. Species are limited in their migrational ability by their

capacity to disperse themselves either vegetatively or by seeds»

Climate affects migrational ability to the extent that propagules can­

not move through areas where they cannot become established.

The following discussion of the climate of Montana is adapted from

Climate and Man (19hl) and the Montana Almanac (1958), Montana's large

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area and great differences in elevation have given it diverse climates.

Factors which determine the nature of these climates include the barrier

effect of the mountainous areas in the westp altitude^ latitude^ the

interior position of the state on the continent^ and the direction of

movement of air masses and storms. West of the Continental Divide the

climate approaches that of the North Pacific Coast^ having milder win=

tersj cooler summers; more frequent^ abundant^ and evenly-distributed

precipitation; less wind movement; a greater degree of cloudiness; and a

shorter growing season than the eastern part. East of the Divide the

continental type of climate prevails.

The mountains are responsible for the protection of western Montana

from the cold waves coming out of the interior of Canada which are there­

fore usually confined to the eastern and north-central parts of the state,

The mountains are also responsible for creating a rain shadow in most of

the eastern districts while forcing the release of moisture as precipita­

tion from the rising air on the windward slopes in the west. Local var­

iation in climate may be observed which is due to altitudinal differ­

ences. This variation includes lower atmospheric pressures^, cooler

temperatures^ stronger winds^ and greater precipitation at higher eleva­

tions.

Montana is traversed by, or is in close proximity to,„ the principal

storm paths of the Northwest. During the winter months most of the

principal air masses which affect Montana originate over the Pacific^

although an occasional cold^ dry air mass builds up over central Canada

which then may dominate eastern Montana and infrequently the entire

state.

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During the summer a semi-permanent thermal low pressure area in

the southwestern United States influences Montana’s weather by causing a

flow of unstable air into the state and thus creating numerous thunder­

storms. Since Montana is in the normal path of frontal storms during

the fall, winter, and spring, it experiences a rapidly changing sequence

of weather conditions during those seasons.

Glaciation has produced striking changes in the land features of

Montana as well as in the vegetation of the state. The most recent

glaciation occurred in the Pleistocene epoch when repeated glaciation

was the outstanding event, with glaciers occupying three times the area

they occupy today (Flint, 1957)» The last great spread of glaciers dates

from less than 20,000 years ago.

During the Pleistocene epoch of the Quaternary Period, which began

about one million years ago, the climate of North America became cold

enough for thousands of feet of glacial ice to accumulate over most of

Canada, Due to its weight and mobility, the ice spread itself outward

from centers of accumulation. Much of the ice moved southward into the

northern Ikiited States. During this epoch the climate of North America

fluctuated from time to time, and as a result the ice advanced southward

into the United States and then melted back into Canada repeatedly.

There were four major advances of Pleistocene glacial ice of which the

Nebraskan was first, followed by the Kansan, Illinoian, and the Wisconsin.

The ice is believed to have melted far northward or to have disappeared

between each of these stages.

Fossil evidence indicates that evolution in plants, invertebrate

animals, and vertebrate animals occurred within the Pleistocene epoch and

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. that new species and genera evolved and that others became extinct

(Flint, 1957). Also evidenced by the fossil record is the fact that

shifts in populations occurred during this epoch. These population

shifts came about in response to temperature and moisture changes as

well as to the direct response to the presence of the glaciers them­

selves.

The most recent glaciation of North America, the Wisconsin, consis­

ted, at the maximum, of ice which was continuous across northern North

America from the Atlantic to the Pacific. The glaciers of that time are

placed in two groups according to their character and place of origin

(Flint, 1957). The Cordilleran complex consisted of the network of gla­

ciers which already occupied the mountains of western North America,

The Laurentide Ice Sheet consisted of one more or less continuous sheet

which spread over North America from Newfoundland to the Rocky Mountains,

The name Cordilleran Ice Sheet has been applied to the most con­

spicuous single element in the Cordilleran complex as it existed during

the last glacial maximum (Flint, 1957). This was a continuous mass of

interconnecting valley and piedmont glaciers and it covered most of the

mountainous country from the Coast and Cascade Ranges, in the west, to

the Rocky Mountains in the east. This mass of ice was centered in Bri­

tish Columbia, On the eastern slope of the Rocky Mountains the Cordil­

leran glaciers met and coalesced with the Laurentide Ice Sheet, Both of

these entered Montana (Fig, 1),

The Cordilleran Ice advanced southward for sixty or seventy miles

within the state. Its course followed the Rocky Mountain Trench and it

overrode most of the mountains, except for the highest ridges, as it

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moved south» Old Glacial Lake Missoula was formed when a lobe of itg

which moved southward across Idaho, dammed the Clark Fork valley and the

Kootenai River valley near the Montana-Idaho state line. This lake re­

mained long enough for many feet of sediments to accumulate on its floor.

It is probable that two separate developments of the lake occurred, one

in Illinoian and one in Wisconsin time (Perry, 1962).

South of the limit of continuous ice in western ttaited States were

at least seventy-five separate areas of glaciers (Flint, 19$7)« Nearly

every high mountain range in western Montana had an ice cap at some time

during the Pleistocene. The distribution of these areas is indicated

(Fig. 1) and the maximum extent of the glaciation is shown regardless of

stage at which it occurred, thus at different times the relative sizes

of the mountain glaciers might vary from that indicated on the map. To­

day several small glaciers are present in the mountains of Glacier

National Park and on the high mountains northeast of Yellowstone National

Park.

During the Pleistocene a continuous ice sheet is also believed to

have covered the high plateau northeast of Yellowstone Park in the south­

western part of the state (Flint, 1957)»

Description of the Alpine Zone

A third topic which must be considered before the distribution of

alpine species can be fully appreciated is that of the alpine zone itself,

its ecology, and the factors which determine its limits.

When one speaks of the alpine region, at least in the popular sense,

he may be making reference to any mountainous area, an opening within a

forested region of a mountain, exposed ridges, or a high meadow above

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■gston

Lake Missoula

Cordilleran Ice

Laurentide Ice

Loca5 M:)iintai.n Glaciers

Figiire 1. Maximum glaciation in Montana during the Pleisto* cene (after Alden, 19^3; American Geographical Society, 19^5).

- 11 -

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timberline. The lower limits of the alpine zone are usually considered

to be timberline, Timberline is^ however, not a well-defined boundary

and difficulties arise in using it as a criterion for the lower limits

of the alpine zone,

Timberline. Timberline is a broad zone which begins with the contin­

uous forest of the lower mountain slopes and gradually thins out to the

last krummholz, usually with no sharp line between these two regions, A

fact often overlooked is that the vegetation of this zone is a complex

of alpine and subalpine vegetation.

There are several theories regarding the relative importance of

various factors in determining the limitations of tree growth or the

timberline. These have been reviewed by Daubenmire (195U)a but essen­

tially they are: (1 ) excessive wind, (2 ) carbon dioxide deficiency, (3 )

snow depth, (U) desiccation during temperature inversions in the winter,

(5) heat deficiency, and (6 ) light deficiency. Because of a limited

amount of meteorlogic data several of these theories have not as yet

been tested, but it is likely that more than one will be found to be sig­

nificant in limiting tree growth (Daubenmire, 19$h)« Within western Mon­

tana one can observe the late snow-belt at Logan Pass and its effects in

depressing the altitude of the timberline. On Divide Mountain, in Gla­

cier Park, one can observe how exposure to the wind can depress timber-

line, According to Daubenmire (195^), however, "these influences never

have enough significance to disrupt the general conformity between

timberline and isotherms," He indicates that the heat deficiency hypoth­

esis may provide the explanation for altitudinal limitation of tree

growth. Support for this might well be obtained from the evident linear

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relationship between altitude of timberline and latitude, the altitude

of timberline decreasing as one goes north from the equator.

Ecology of the alpine zone. The study of the ecology of the alpine

has been approached in more than one way. In Colorado, where much of

the recent work has been done, the ecosystem approach has been followed.

According to Marr (I96I) "an ecosystem is an ecological unit or sub­

division of the landscape or geographic area that is relatively homo­

genous and reasonably distinct from adjacent areas. It involves organisms,

environmental factors, and ecological processes," Stands are concrete

vegetation units having homogenity and distinctness of features which

makes them stand out from the vegetation of surrounding areas, while

stand-types are units which combine the characteristics of many stands.

In following this approach, Bamberg (I96I) divides his alpine stand-types

for Montana into groups according to environmental relations, namely

whether the parent rock was hard-rock of igneous and metamorphic origin,

or whether it was the soft-rock type, primarily limestone, of sedimen­

tary origin, A further subdivision was made on the basis of winter

snow-free areas and winter snow-covered areas. The final arrangement

was on the basis of increasing moisture, Bamberg established sixteen

stand-types to characterize the vegetation on the eight study areas he

worked. He indicated that three stand-types are essentially not vege­

tated and these are the snow accumulation areas, the rock outcrops, and

the scree or talus types.

He points out the difficulty of sampling due to much microenviron­

mental and subsequent vegetational variation occurring in alpine areas.

He noted the importance of soil moisture in determining stand-types and

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found it effective to organize the discussion of these units in a se­

quence that followed their distribution along a moisture gradient*

Topography, relation to snowbanks, soil-frost action and soil drainage

influence soil moisture so that the gradient between stand-types may be

gentle or abrupt (Bamberg, I96I).

In Glacier National Park, Choate (1963), utilizing the technique

of ordination, demonstrated a continuum in the alpine area at Logan Pass.

She also observed the importance of soil moisture and her ordination

showed a gradation from wet meadow communities to the dry, rocky commun­

ities occurring on ridges and ledges.

Other environmental factors of importance in the alpine zone are

wind, depth, and duration of snow cover; frost phenomena or cryopedo-

genic processes; and, in certain areas, activity of small mammals.

Wind operates directly on the plants by increasing transpiration and by

causing mechanical breakage or scouring. It also affects soil moisture,

snow cover, and temperature. In any given area local topography may

greatly modify the direction and speed of the wind. Intense frost action

can exert a major control on vegetation in alpine environments (Johnson

and Billings, 1962) through repeated modifications of patterned ground.

These factors may act individually or together on the alpine vegetation.

The dynamics of the alpine tundra are not well-known even though it is

apparent that wind, moisture, and snow cover are important in determining

the character of a given stand (Bamberg, I96I).

Definition of an Alpine Plant

Many plants can exist in the climate above timberline although this

alone is not a good criterion to use in defining an alpine plant. The

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latter shoijld be able to reproduce in this environment. These

criteria delimit a group of species which would include some not con­

sidered to be alpine plants. However, to include only plants which

do not occur below timberline would be to disregard the majority of the

species occurring in the alpine. In this study the definition of an

alpine plant has been broadened to include those species which are prin­

cipally distributed near or above timberline or are very common in al­

pine areas, even if they are also fairly common in certain habitats at

lower elevations* In addition, certain species which are present in the

arctic and the alpine as well as below timberline have been discussed

under the category of alpine plants if they are widely distributed

throughout the alpine as well as the arctic,

Floristic Elements

A flora is a product of complex processes of evolution and it is

necessary to analyze it in terms of its particular origins. Several

methods of approach might be attempted in order to d etermine the history

of the alpine flora. One of the best would be to study the distribution

of Pleistocene and pre-Pleistocene fossil material in an attempt to use

the past as a guide to the present. This approach has received more

attention in Europe than in North America, It is limited by the absence

of a complete fossil record. However, the little data available has

proved valuable in understanding the origin of Pleistocene and post-

Pleistocene floras. Another method would be to select a species or a

group of species, study their ecological requirements and from this

deduce their past distribution. According to Dahl (1959), this method's

validity depends considerably on which species are included in the group.

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A third approach, and the one which has been followed in part in this

stndy, is to treat the flora as a whole and divide its component taxa

into groups according to their distribution patterns, ecological re­

quirements, adaptations to dispersal and other characteristics and try

to conclude from this something about the history of the groups and of

the flora as a whole* Such groups are called floristic elements*

The climatic changes which are responsible for plant migration are

also responsible for bringing about selection and segregation of species

and permit the intermingling of previously unassociated plants. Plant

distribution is controlled secondarily by the distribution of edaphic

factors. In order to analyze the floristic elements present in the

alpine flora of today, it is necessary to take these facts into account.

The evolution of floras is dependent upon plant migration, the

evolution of species, and the selective influences of climatic change

acting upon the varying tolerances of the species common to the flora.

The groupings of species within any given flora may differ widely ac­

cording to the principle upon which this grouping is based.

The division of a flora into floristic elements should be based on

five principles (Mason, 1959). First, the species should be grouped

according to their geographical distribution. Second, they should be

grouped according to their region of origin, thus reflecting the genesis

of the different parts of a given flora. Third, they should be grouped

according to the given routes by which they migrated into the floral re­

gion, As a species may penetrate into the domain of a given flora by

several routes, these latter elements may be difficult to establish.

Fourth, the species should be grouped according to the time they became

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part of the given flora, and fifth, they should be grouped according to

habitat preferences# According to Mason the establishment of such ele­

ments is of great significance in determining the history of a flora

and of those climatic changes to which it has been subjected#

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. PROCEDURE

This study has been restricted to the political boundaries of

Montana principally because of limited time and of limited access to

collections from other areas» The area covered by the study is large^

and thus the amount of field study possible was not great and the

majority of the data has come from herbarium studies.

Determination of Altitudinal Limits of Alpine Areas in Montana

Certain topographic features are of great importance in determining

the altitude of timberline in any given area so that at a given latitude

in the state, the altitude of timberline is not necessarily uniform,

Bamberg (1961) lists the lower altitudinal limits of the alpine areas

he visited in the state and these show the general linear relationship

of treeline to latitude which I also observed. I have arbitrarily

chosen to record as alpine anything collected above 9500 feet in the

south-central area of the state down to 6700 feet at Logan Pass in Gla­

cier National Park near the northern boundary of Montana. In setting

these limits as criteria I may be including some collections made below

timberline, I believe, however, that this will allow me to cover any

possible alpine situation which might otherwise be missed by setting

higher altitudinal limits as criteria in regions with which I am not

familiar.

Field Studies

During the summer of 196U thirteen trips were made to alpine areas

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located in extreme western ard in northwestern Montana* The areas were

chosen because of limited alpine collections from them. They wert in­

tensively collected when visited although the number of collections

from a particular area varied d'oe to the time of the S'lmmer at which it

was visited and number of visits made. Most of these areas^ therefore,

should still be studied throughout the growing season.

The following is a list of collected areas and the dates they were

visited:

Divide Mountain, Glacier National Park, Juiy 26, 196U Glacier County August 9, I96U August II45 I96U

Logan Pass, Glacier National Park, July 196U Glacier County August '', 1961+

Mount Henry, Glacier National Park, August 15, 196L Glacier County

Siyeh Pass, Glacier National Park, July I8, 19ÔU Glacier County August 8, I96L

Lunch Creek Cirque, Glacier National Park, A>igust it, I96U Glacier County

Trapper Peak, Bitterroot Mountains, July 1966 Ravalli County

Lolo Peak, Bitterroot Mountains, July 31, 196U Missoula County

Piquett Mountain, Bitterroot Mountains, August 25« I96U Ravalli County

St* Mary's Peak, Bitterroot Mountains, Auguir. -7, lybu Ravalli County

Herbarium Material Studied

All collections which would be considered alpine, according to the

definition previously given, which were in the herbarium of the

University of Montana, Missoula^ that of Montana State Uni^ersioy,

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Bozeman; as well as that of Region One, Ihlted States Forest Service,

Missoula; and selected families from the reference herbarium at Glacier

National Park were studied* The University of Montana herbarium has a

large amount of material from west of the Continental Divide and that of

Montana State University has numerous collections from the southwest cor­

ner of the state east of the Divide*

At the first, a complete survey of all collections in the herbarium

of the University of Montana was made in order to determine all the

vascular plants which were collected above timberline in the state* In

many instances there are several collections of the same species from a

particular area and these have all been recorded* While the survey of

the herbarium at Montana State University was believed to be complete for

alpine species, only stations which were not obtained from the herbarium

at the University of Montana were recorded* Duplicates of collections

on file at each institution have not, in general, been recorded* A

similar procedure was followed for the survey of the herbarium of the

Region One, Ikiited States Forest Service*

In making the herbarium survey, as well as in identifying my own

collections, the species concepts of Hitchcock, et al. (1955, 1959,

1961, I96U) have been followed for the Dicotyledoneae, Hitchcock and Chase

(1950) for the Gramineae, and Mackenzie (1931=1935) and F.J* Hermann

(I96U) for Garex. Determination of the Festuca ovina L. complex follows

Holmen (I96U). The rest of the Monocotyledoneae, Equisetaceae, Fclypod-

iaceae, Selaginellaceae, and Lycopodiaceae were determined according to

the concepts of either Davis (1952) or Moss (1959). Polunin (1959) is

the authority for the occurrence of species in the arctic* Munz (1959)

is the authority for California distributions*

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The distribution patterns of the species have been plotted on maps

which show the mountainous portion of Montana. On these maps the stip­

pled areas indicate the mountainous regions and the clear areasç, the

intermontane valleys (after Alden^ 1953)* The numbers on the stippled

areas refer to the names of the mountain masses (Alden, 1953.j Perry<1

1962) as indicated on the legend of figure 2» The maximum extents of

the Gordilleran Ice and the Laurentide Ice Sheet as they were reached

in Montana are also indicated. The major drainages have been included

and indicated by circled numbers.

A detailed state distribution map based on field collections^ her^

barium collections, and/or reported occurrences was made for each spe­

cies. Space prohibits the documentation of the stations indicated on

the distribution maps. Altitude of these stations has not been shown

on the maps, but this information has been deposited in the herbarium

of the University of Montana. The maps which have been included in this

thesis are representative of a particular pattern of distribution with­

in the state, or, in a few instances, they illustrate the distribution

of a particular species discussed in the text. For certain of the

categories a map of the general distribution pattern based on ranges

reported from outside of the state has been made. These are, however,

only to be taken as indicating a general pattern for the group.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. figure 2. MOUNTAINOUS MONTANA

MOUNTAIN SYSTEMS RIVERS

1. Purcell Mts. 18. Anaconda Range Q Kootenad. River (Pintlar Area) 2, Selish Mts. Marias River 19. Elkhom Mts. 3* Whitefish Mts. South Fork of 20. Bridger Range Flathead River U. Lewis Range (Glacier Park Rg.) 21. Beaverhead Range Teton River

Cabinet Mts. 22. Pioneer Mts. Missouri River

6, Mission Range 23. Ruby Range Clark Fork of Columbia River 7, Swan Range 2ii. Snowcrest Range Swan River 8, Flathead Range 25. Gravelly Range Blackfoot River 9* Goeur d'Alene Mts. 26. Tobacco Root Mts. Bitterroot River 10. Bitterroot Mts. 27. Centennial Mts. Beaverhead River 11. Garnet Range 28. Madison Range Madison River 12. Helena Mts. 29. Gallatin Range Gallatin River 13. Big Belt Mts. 30. Absaroka Range (Snowy Mts,) Yellowstone River lU. Little Belt Mts. 31. Beartooth Plateau 15• Castle Mts. Area

16. Sapphire Range 32. Crazy Mts.

17. Flint Creek Range 33. Highwood Mts.

3U. Big Snowys

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Figure 2, . Mountainous Montana (in part after Alden, 1953)*

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. FLORISTIC ELEMENTS IN THE MONTANA ALPINE FLORA

The Montana alpine flora appears to consist of five major

floristic elements which can be subdivided as followss

1. Lowland element (about 225 species)

2. Arctic-alpine element (about 110 species)

A. Species fully circumpolar or nearly so

Bo Species widely distributed in the North American Arctic and that of eastern or western Eurasia

C, Species restricted to North American Arctic or found^ in ad­ dition, only on the Pacific Coast of Eurasia and/or in Green­ land

3. Gordilleran element (about 100 species)

A. Species with a wide range in the western cordilleras

a* Widely distributed from Alaska along the Rocky Mountains and Sierra Nevadas, sometimes extending into the Cascades but not in the Coast Ranges of California

b. Widely distributed from Alaska along the Rocky Mountains, Sierra Nevadas, and in the Coast Ranges of California, sometimes extending into the Cascades

c. Widely distributed from British Columbia and Alberta along the Rocky Mountains and Sierra Nevadas, sometimes extending into the Cascades but not in the Coast Ranges of California

do Distributed in the Rocky Mountains and Sierra Nevadas but not in the Coast Ranges of California, and reach their north­ ern limits in southwestern or south-central Montana

Bo Species restricted to Rocky Mountains

ao Widespread in Rocky Mountains

bo With northern limits in southwestern or south-central Mon­ tana

Co Disjunct

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5* Endemics (about 20 species)

1« The Lowland Element

Many plants in the Montana alpine are more widely distributed

below timberline. These are here called the lowland element. Some of

these have only occasional alpine occurrences while others have an ap­

parent wide environmental tolerance and they occur commonly above as

well as below timberline, but show no apparent morphological segregation

between alpine populations and lowland populations. Other species which

may occur above timberline^ as well as below^ show marked dwarfness in

their alpine environments, which transplant experiments have shown is

sometimes genetically controlled. These individuals constitute alpine

ecotypes (Clausen, Keck, and Hiesey, 19^0, 19U7^ 19U8j Turesson, 1922,

1925, 1931) and retain their dwarfness when they are transplanted to

lowland environments. It is questionable whether any understanding of

alpine distribution patterns can be derived from the alpine distribu­

tions of those species which do not have alpine ecotypes because their

alpine distribution is only an incidental aspect of their total distri­

bution. These species have not been discussed in detail nor have their

overall distributions been classified although their presence in alpine

stations has been indicated in a list in the appendix. Another list of

these species only is included there. The maps of their alpine distri­

bution are deposited in the herbarium of the University of Montana.

The distribution of the alpine ecotypes of the other lowland species

which occur in the alpine has been discussed with those of other species

which have similar distribution patterns. In addition, some of the

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species which have been discussed in detail and included as “alpine

plants’ may better be classified as representative of the lowland

element, or at least of subalpine species* In the tabular distribution

of all species included in the appendix these have been indicated with

a question mark following their name.

2. The Arctic-alpine Element

A. Arctic-alpine species which are fully circumpolar or nearly so.

Arctic-alpine species in the Montana alpine which are fully or nearly

circumpolar includes Arenaria rubella (Wahlenb.) J.E.Smith, Arnica

alpina L., Campanula uniflora L., Garex bipartita All., Carex misandra

R.Br., Cassiope tetragona (L.) D.Don, Cystopteris fragilis (L.) Bernh,,

Deschampsia caespitosa (L.) Beauv., Draba fladnizensis Wuifen., Draba

glabella Pursh., Epilobium latifolium L,, Erigeron humilis Greene,

Festuca brachyphylla Schultes, Juncus biglumis L., Juneus castaneus Sm.,

Koenigia islandica L., Lycopodium selago L., Oxyria digyna (L.) Hill,

Phippsia algida (Phipps) R.Br., Poa alpina L., Poa arctica R.Br., Poa

glauca Vahl,, Potentilla nivea L., Ranunculus pygmaeus Wahlenb., Salix

arctica Pall,, Saxifraga caespitosa L., Saxifraga cernua L., Saxifraga

flagellaris Willd., Saxifraga oppositlfolia L., Sedum roseum (L.) Scop.,

Tofieldia pusilla (Michx) Pers., and Trisetum spicatum (L.) Richt.

In addition, Juncus albescens (Lange) Fern, forms a circumpolar series

with Juncus triglumis L., (Polunin, 1959) while Draba lonchocarpa Rydb,

is a derivative of the nearly circumpolar Draba nivalis Liljebl. (Hitch­

cock, et al., I96U).

A conclusion that appears obvious with regard to these species is

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that most of them formerly belonged to an arctic flora but were driven

south during the Pleistocene glaciation» After the ice retreated and

the snowline moved back up the mountain slopes3 some of them moved back

up the mountains and as a result now occur in these alpine areas» It is

also probable that certain species which were strictly alpine prior to

the Pleistocene migrated north with the retreat of the glaciers to become

a part of the arctic flora. Therefore, the area of origin of particular

arctic-alpine species is frequently difficult to determine.

The fact that the Continental Ice Sheet reached its southern limits

within Montana would seem to indicate that arctic species migrating south

in front of the ice might now, in consequence, have their southern limits

within the state. These would not include species able to disperse over

the intervening distances today or those whose environmental tolerance

doesn't limit them to tundra or glacial gravels in close proximity to

the glacial borders. Whether tundra preceded the glaciers into Montana

is not known (Flint, 1957). Several of these reach their southern

limits in the Rocky Mountains in the northwest corner of Montana, mostly

in Glacier National Park (Fig. 3). These limits are covered by the

extension of the ice during the Pleistocene. They are further restricted

within the area delimited by the average July temperature of 6l degrees

(Fig, L). Although this isotherm is derived from data collected at

stations which are well below the alpine zone it is believed that the

temperatures of these isotherms reflect the differences between the

temperatures in various alpine areas. The implication would seem to be,

therefore, that temperature has been a limiting factor in the distribu­

tion of these species. They include Juncus biglumis L,<, Carex bipartita

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Figure 3. Erigeron humilis Graham. The Montana alpine distribu­ tion of a circumpolar arctic-alpine species reaching its southern limits in the Rocky Mountains in northwest Montana,

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8

61

Figure Uo Average July temperatures in Montana (U.S. Dept, Agriculture, 19^1).

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All,, Erigeron humilis Graham, and Tofieldia pusilla (Michx.) Pers,,

all of which were collected in wet areas or mossy turf. These areas,

it should be pointed out, are characteristically cooler during the sum^

mer growing season than other alpine habitats,

A second group of circumpolar arctic-alpine species includes

those which reach their southern limits in the Rocky Mountains in ex“

treme west-central Montana (Fig, 5)» These would also appear to have

been driven south by the Pleistocene glaciation but evidently were able

to persist beyond the maximum southern edge of the continuous ice.

They either would not have been restricted to the glacial margins or

were able to migrate into mountains south of there as they now have

stations well south of drift borders. The species are Cassiope

tetragona (L,) D.Don var* saximontana (Small) C.D.Hitchc., and Arnica

alpina L. var, tomentosa (Macoun) Cronq. The collection of Cassiope

tetragona made on St, Mary's Peak in the Bitterroot Mountains is an ex­

tension of its known range south from Glacier National Park, I have

seen no collections or published records of this species or of Arnica

alpina whose southern station is in the Pioneer Mountains, between

these areas and Gfacier National Park,

A third group of circumpolar arctic-alpine species are those which

are widely distributed throughout the mountainous portion of the state

and have as their southern limits in the Rocky Mountains the Beartooth

Plateau area of southwestern Montana and northwestern Wyoming (Fig. 6 ).

The Beartooth Plateau is a high plateau believed to have been exten­

sively glaciated during the Pleistocene which, even today, has several

small glaciers (Perry, 1962; Flint, 195?), The upper krummholz limits

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Figure Cassiope tetragona (L,) D.Don var. saximontana (Small) G.L.Hitchc. The Montana alpine distribution of a cir­ cumpolar arctic-alpine species reaching its southern limits in the Rocky Mountains in extreme west-central Montana.

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on this plateau occur at about 9,800 feet and the total area above

timberline is quite extensive. According to Bamberg (1961) it is the

only area which he studied where all sixteen of his stand=types may be

found, thus, indicating a wide diversity of available habitats. The

species in this category which reach their southern limits here ares

Draba glabella Pursh, Juncus albescens (lange)Fern., Koenigia islandica

L., and Saxifraga oppositifolia L. In addition, several arctic species

which are not circumpolar have their southern boundary here. These will

be discussed later in the thesis.

A fourth group of circumpolar arctic=alpine species which have

their southern limits in the Rocky Mountains in the high mountains of

Colorado or New Mexico include the following speciess Campanula uniflora

L,, Carex misandra R, Br., Juncus castaneus Sm,, Lycopodium selago L.,

Phippsia algida (Phipps)R. Br,, Poa alpina L., Poa arctica R« Br.,

Poa glauca Vahl, Potentilla nivea L., Ranunculus pygmaeus Wahlenb.,

Salix arctica Pall, var. petraea Anderss., Saxifraga caespitosa L.

var. minima Blank., Saxifraga cernua L., and Saxifraga flagellaris WiHd,

All of the species just discussed are absent from the Sierra

Nevada and Coast Ranges of California though some extend into the

Cascades of Oregon and Washington. A final group of species which are

circumpolar arctic-alpine are present in both the Rocky Mountains and

the Sierra Nevadas. These include Arenaria rubella (Wahlenb.)J,E.

Smith, Cystopteris fragilis (L.)Bernh., Deschampsia caespitosa (L.)

Beauv., Draba fladnizensis Wuifen., Epilobium latifolium L,,

Festuca brachyphylla Schultes, Oxyria digyna (L,)Hill, and Sedum roseum

(L.)Scop. These apparently migrated south all along the broad front

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m vingston t

Figure 6, Saxifraga oppositifolia L, The Montana alpine distribution of a circumpolar arctic-alpine species reaching its southern limit in the Rocky Mountains in the Beartooth Pla­ teau area.

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DMTA

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Fig-ure 7. Festuca brachyphylla Schulte. A circumpolar arctic- alpine species reaching its southern limit in the Rocky Mountains in New Mexico,

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of the Gordilleran sheet unlike the previous groups which apparently

were restricted in their migration to the Rocky Mountains or else were

able to migrate along the front of the sheet if they were restricted to

narrow corridors in their southern migration,

B. Arctic-alpine species which are widely distributed in the North

American Arctic and that of eastern or western Eurasia. A second large

category of arctic-alpine species present in the Montana alpine flora

includes those species which are not, according to Polunin (1939), fully

or nearly circumpolar. These species are reported by him to be widely

distributed in the North American Arctic and that of eastern or western

Eurasia, Polunin indicates that his determination of circumpolar ranges

for the species so indicated in his manual are based on his field obser­

vations and herbarium study. It is possible that some of these species

may be shown to be circumpolar later. The arctic-alpine species which

are widely distributed in the North American Arctic and that of eastern

or western Eurasia have been divided into groups in a similar manner as

was done for the completely circumpolar ones.

Six of these species are believed to reach their southern limits

in the Rocky Mountains in north-central to northwestern Montana (Fig, 8 ),

These are Androsace lehmanniana Spreng., Carex nardina Boot, Dryas

integrifolia Vahl, Festuca vivlpara (L,) Small, Hedysarum alpinum L.,

and Lycopodium alpinum L, I have seen Montana collections of Carex

nardina, Lycopodium alpinum, and Festuca vivipara only from Glacier

National Park. Bamberg (1961) reported Hedysarum alpinum from there

and Hitchcock et al. (I961) indicates the range of the latter to be from

Alberta and British Columbia south to north-central Montana. Dryas

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Figure 8, Androsace lehmanniana Spreng, The Montana alpine distribution of an arctic-alpine species widely distributed in the North American Arctic and that of western Eurasia and which reaches its southern Rocky Mountain limit in north-central Montana,

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integrifolia has not been reported by Hitchcock for the state but

Bamberg (1963) reports it for the Big Snowys in central Montana and a

hybrid between it and Dryas octopetala L» for Glacier National Park.

The species ranges east from the Canadian Rockies along the inter­

national border to New Hampshire and north to Alaska and Greenland,

Androsace lehmanniana was previously known in Montana only from the

Big Snowys, but a collection of it was made on Divide Mountain in

Glacier National Park which is a marked northwestern extension of its

ranges in the state, Montana is the apparent western limit for these

six species which range east in Canada and sometimes into eastern

United States. An arctic or boreal origin is suspected for this group

on the basis of their fairly wide distribution across North America

east of the Rocky Mountains. They probably entered the Montana alpine

from the northeast,migrating in front of the Laurentide Ice Sheet

rather than along the Rocky Mountains,

A second group in this category includes species having the

Beartooth Plateau area as their southern limit in the Rocky Mountains

(Fig, 9)0 These are the following: Aster sibiricus L., Carex

podocarpa R,Br. (including C« tolmei Boott), Eriophorum vaginatum L»

(including E. callitrix Cham,), Parnassia kotzebuei Cham, var, kotzebuei,

Pedicularis oederi Vahl, Senecio resedifolius Less., sind Stellaria

calycantha (Ledeb.) Bong. Stellaria calycantha is a wide-ranging

species occurring as far west as the Sierra Nevada of California and as

far east as the New England states, A distribution pattern as extensive

as this would indicate a northern origin for the species. Pedicularis

oederi is widespread in the arctic from eastern Europe across Asia to

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ngston

Figure 9. Pedicularis oederi Vahl, The Montana alpine distribution of an arctic-alpine species widely distributed in the North American Arctic and western Eurasia which reaches its southern limits in the Rocky Mountains in the Beartooth Plateau area.

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Alaska-Yukon, This would indicate a likely arctic origin, Carex

podocarpa, Parnassia kotzebuei, Aster sibiricus, and Senecio resedifolius

extend south to Washington and/or Oregon as well as to the Beartooth

region in the Rocky Mountains, A possible alpine origin is proposed for

them on the basis of their limited arctic distribution which covers the

area from eastern Asia through the North American arctic, Senecio

resedifolius ranges in the arctic from eastern Europe across Asia and

Alaska-Yukon to western Canada indicating a probable arctic or boreal

origin.

The largest number of species in this category which reach their

southern limits in the high mountains of Colorado or New Mexico (Fig,

10) are as follows : Aster alpinus L,, Aster aboriginum Richards,

Astragalus alpinus L,, Carex atrata L,, Carex capillaris L., Carex

capitata L,, Carex hepburnii Boott, Carex scirpoidea Michx,, Dryas

octopetala L,, Eritrichium nanum (Vill,) Schrad, (including E, areti-

oides (Cham. & Schlecht) D.C.), Gentiana algida Pall,, Geum rossii

(R.Br.) Ser,, Kobresia bellardii (All.) Degl., Lloydia serotina (L.)

Reichb,, Saxifraga bronchialis L,, Smelowskia calycina (Steph,) C,A,

Mey., and Taraxacum lyratum (Ledeb,) DC., all of which are absent from

the Sierra Nevada Range and the Coast Ranges of California. Also inclu­

ded in this group are Achillea millefolium L., Androsace septentrionalis

L,, Antennaria alpina (L.) Gaertn,, Arenaria rossii R.Br,, Asplénium

viride Huds., Athyrium alpestre (Hoppe) Rylands, Cerastium beeringianum

Cham. & Schlech., Crepis nana Rich,, Draba crassifolia R. Grab,,

Epilobium alpinum L. agg,, Erigeron acris L., Luzula parviflora (Ehrh.)

Desv., Luzula spicata (L,) DC,, Salix glauca L,, Sibbaldia procumbens L.,

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•^IDAi

Figure 10. Eritrichium nanum (Vill.) Schrad, var, elongatum (Rydb,) Gronq, The Montana alpine distribution of an arctic- alpine species widely distributed in the North American Arctic and western Eurasia which reaches its southern Rocky Mountain limit in New Mexico,

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Silene acaulis L « , Solidago inultlradiata Ait. , and Taraxacum ceratophorum

(Ledeb.) DC. which occur in the Sierra Nevadas and/or Coast Ranges of

California as well as in the -K^ocky Mountains.

Achillea millefolium ssp. lanulosa is a wide-ranging species of

which variety alpicola (Rydb.) Garrett, is the alpine and subalpine

ecotype (Hitchcock et al., 1955)• Only the alpine occurrences of this

species have been plotted for the state. Epilobium alpinum, according

to Hitchcock et al.(l96l), includes at least four other taxa that are

nearly always accorded specific rank. These include E. clavatum Trel.,

E. hornemannii Reichb., E, lactiflorum Hausskn., and E« oregonense Hausskn.

According to Hitchcock et al. (1961), "these names appear to distinguish

little more than conspicuous phases of almost continuous variation in

which neither genetic nor geographic barriers have developed to delimit

the different variants except in a mechanical way." I have seen Montana

alpine collections of variety alpinum, variety clavatum, variety lacti­

florum, and variety nutans (Hornem.) Hook, and geographical isolation,

at least in Montana, of these variants is non-existent. Erigeron acris

is divided into three varieties (Hitchcock et al., 1955) to which most

of the Montana alpine material is referred as variety debilis Gray,

According to Hitchcock et al, (1959), Eritrichium nanum consists of

three rather weakly defined varieties, with the plants of the United

States referred to as variety elongatum (Rydb.) Cronq. Geum rossii

consists of three weakly distinguishable races of which two occur in the

Montana alpine (Hitchcock et al., I96I). The variety rossii reaches its

southern Rocky Mountain limit in the Mission Mountains of Montana. The

variety turbinatum (Rydb.) G.L,Hitchcock is apparently restricted in

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. the Rocky Mountain region from central and western Montana south to

New Mexico and west to the Wallowa Mountains of northeast Oregon

(Hitchcock et al,, I96I), Saxifraga bronchialis consists of at least

three varieties outside of the arctic (Hitchcock et al., 1961) to which

the Montana material I have seen is referred to as variety austromontana

(Wieg.) G,N,Jones, Silene acaulis is represented in Montana by two

clearly marked races (Hitchcock et al., I96U), The variety subacaules-

cens (Williams) Fern, & St.John is supposed to range in the Rocky

Mountains from southern Montana to New Mexico and Arizona, The variety

exscapa (Allioni) DC, is supposed to be more restricted to northern

Montana, Idaho, Oregon, and north in the Rocky Mountains to the arctic.

While the characters separating the two varieties appear fairly well

marked, the latter do not seem to show any geographical isolation with­

in the state. Smelowskia calycina is, according to Hitchcock et al,

(I96I1) represented in Montana by the variety americana (Regel & Herd,)

Drury & Rollins,

Achillea millefolium, Androsace septentrionalis, Antennaria alpina ,

Astragalus aboriginum. Astragalus alpinus, Carex atrata, Garex hepburnii,

Gerastium beeringianum, Crepis nana, Draba crassifolia, Dryas octopetala,

Epilobium alpinum, Eritrichium nanum, Geum rossii, Lloydia serotina,

Luzula spicata, Sibbaldia procumbens, Silene acaulis, Smelowskia calycina,

and Solidago multiradiata are widespread in the Montana alpine. Only

limited collections of Aster alpinus, Athyrium alpestre, Arenaria rossii.

Asplénium viride, Garex scirpoldea, Gentiana algida, Kobresia bellardia,

Luzula parviflora, Salix glauca, and Taraxacum certophorum have been

seen, but on the basis of their overall distribution they are expected

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to be more widespread in the Montana alpine then these collections would

indicate.

C. Arctic-alpine species restricted to North American Arctic or

founda in addition, only on the Pacific Coast of Eurasia and/or in

Greenland. The final category of incompletely circumpolar arctic-

alpine species in Montana (Fig, 11) includes those which are restricted

to the arctic areas of North America and/or adjacent Greenland and

easternmost Asia. An alpine origin for some of these species is quite

possible on the basis of their restricted arctic distribution. The

species in this category, which are absent from the Sierra Nevadas and

Coast Ranges, include : Agropyron latiglume (Scribn. & Smith), Anemone

drummondii Wats., Anemone multifida Poir., Arenaria obtusiloba (Rydb.)

Fern., Bupleurum americanum Coult. & Rose, Carex nigricans C.A. Meyer,

Carex practicola Rydb,, Carex spectabilis Dewey, Draba lanceolata Royle,

Erigeron caespitosus Nutt., Erigeron compositus Pursh, Festuca

baffinensis N.Polun., Oxytropis viscida Nutt., Pedicularis groenlandica

Retz,, Ranunculus gelidus Kar. & Kir., Salix vestita Pursh, Selaginella

rupestris (L.) Spring, Senecio fuscatus Hayek., Senecio lugens Rich.,

Senecio pauciflorus Pursh, Silene repens Pers., and Veronica wormskjoldii

Roem, & Schult.

Agropyron latiglume, Bupleurum americanum, Draba lanceolata,

Erigeron caespitosus, Festuca baffinensis, Salix vestita, Selaginella

rupestris, Senecio fuscatus, Senecio lugens, and Silene repens are ap­

parently absent from the Sierra Nevadas. Festuca baffinensis, Senecio

fuscatus, and Silene repens range from the arctic south in the Rocky

Mountains apparently to the Beartooth Plateau region, Salix vestita

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Figure 11, Agropyron latiglume (Scribn, & Smith)Rydb. The Montana alpine distribution of an arctic-alpine species restricted in the afctic to North America and Greenland,

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reaches the arctic in eastern Canada. This species is reported by-

Hitchcock et al. (I96I4.) for the Big Snowys but I have only seen col­

lections from Glacier National Park, the Flathead Range, the Swan Range

and the Mission Mountains in north-central Montana. It is, however,

reported to reach the Wallowa Mountains of Oregon and may be expected

in more alpine areas throughout this state. Only limited numbers of

collections of Draba lanceolata. Ranunculus gelidus, Selaginella

rupestris, and Senecio lugens have been seen from Montana but are ex­

pected to be more widespread in the state on the basis of their general

distribution patterns. Collections of Agropyron latiglume, Bupleurum

americanum, and Erigeron caespitosus indicate that they are widespread

in the alpine flora of the state.

The species in this category which occur in the Sierra Nevadas and/

or Coast Ranges as well as in the Rocky Mountains includes Anemone

drummondii, Arenaria obtusiloba, Carex practicola, Erigeron compositus,

Pedicularis groenlandica, Senecio pauciflorus, and Veronica wormskjoldii.

Only two collections of Carex practicola were seen but this species is

also to be expected from more Montana alpine localities. The phase of

Erigeron compositus which is generally confined to the higher elevations

is the variety discoideus Gray. Certain collections I have seen of this

species from Montana alpine areas are better referred to the variety

glabratus Macoun which is the more common phase accurring generally at

moderate elevations in the mountains (Hitchcock et al., 1955)o I have

not seen a collection of Senecio pauciflorus from Montana, but it is

reported by Bamberg from the Flint Creek Mountains and would be expec­

ted in the state. Anemone drummondii, Arenaria obtusiloba, Pedicularis

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groenlandica, and Veronica wormskjoldii are widespread in alpine areas

of the state.

There are a few arc tic-alpine plants in Montana (Fig, 12) which

have a wide distribution including stations in both the northern and

southern hemispheres as well as in alpine localities in Europe and Asia,

These species includes Gentiana prostrata Haenke., Lycopodium clavatum

L., Phleum alpinum L., and Trisetum spicatum (L.) Richt, According to

Polunin, only Trisetum spicatum is fully circumpolar, Phleum alpinum

ranges in the arctic from eastern Europe, eastern Canada, west and east

Greenland, and probably Alaska-Yukon (Polunin, 19$9), Lycopodium

clavatum occurs in the arctic of west Greenland and eastern Asia, and

Gentiana prostrata reaches the eastern Asian arctic and that of Alaska-

Yukon (Polunin, 1959)* All of these species probably migrated into the

southern hemisphere during the Pleistocene when conditions were favor­

able for migration. After the Pleistocene the warmer temperature and

drier conditions resulted in the disappearance of these species in the

central part of their ranges and the formation of a bipolar distribution.

As these species are also present in Eurasia it is most probable that

their origin was in the arctic or at least boreal regions, Trisetum

spicatum was probably circumpolar and Phleum alpinum nearly so prior to

the Pleistocene,

3. Cordilleran Element

The third major element present in the alpine flora of Montana

consists of those species which do not reach the arctic but which are

distributed in the cordillera of western North America, The major

assumption underlying this category is that these species did not

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Figure 12. Trisetum spicatum (L,) Richt, The Montana alpine distribution of an arctic-alpine species which is widely distributed including stations in both the northern and southern hemisphere as well as alpine areas in Europe and Asia,

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originate in the arctic but in alpine areas of the mountains of western

North America» This category can be subdivided into several more or

less natural groups based on the general distribution patterns of the

included species,

A* Species with a wide range in the western Cordilleras, a* The

first group consists of species which are widely distributed from Alaska

along the Rocky Mountains and Sierra Nevadasj, sometimes extending into

the Cascades but not in the Coast Ranges of California, It includes:

Agropyron subsecundum (Link) Hitchc., Anemone multifida Poir., Anemone

occidentalis Wats., Antennaria umbrinella Rydb., Aster foliaceus Lindl,,

Draba oligosperma Hook., Hieracium gracile Hook., Juncus drummondii E,

Meyers, Juncus mertensianus Bong,, Oxytropis viscida Nutt., Fhyllodoce

empetriformis (Sw,) D.Don, Potentilla diversifolia Lehm., Ranunculus

eschscholtzii Schlecht, Romanzoffia sitchensis Bong,, and Senecio

cymbalaroides Nutt.

Agropyron subsecundum ranges east across Canada to Newfoundland and

south to the mountains of Maryland. It is represented by an alpine form,

variety andium (Scribn. & Smith) Hitchc,, occurring from Montana to

Washington south to Colorado and Nevada, Anemone multifida ranges from

the valleys and foothills to the higher peaks. The variety tetonensis

(Porter) C.L.Hitchc. is more characteristic of the higher peaks and is

the variety to which all of the Montana alpine material has been refer­

red, According to Hitchcock et al, (19SS) the alpine and subalpine

varieties of Aster foliaceus are variety apricus Gray, which ranges

from southern British Columbia to northern California and east to

Colorado and Montana, and variety foliaceus, which ranges northward, at

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Figure 13* General distribution of the cordilleran element widely distributed from Alaska along the Rocky Mountains, in the Cascades and Sierra Nevadas but not in the Coast Ranges of California.

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Figure lU, Fhyllodoce empetriformis (Sw.) D.Don. The Montana alpine distribution of the cordilleran element widely distributed from Alaska along the Rocky Mountains, in the Cascades and Sierra Nevadas, but not in the Coast Ranges of California,

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. decreasing elevations from Glacier National Park^, where it occurs in the

high mountains » The common Rocky Mountain variety of Draba o~1 i gosperma

is the variety oligosperma to which all the alpine material 1 have seen

is referred. According to Hitchcock et al. (1961) Potentilla diversi­

folia is an extremely variable species. The alpine material from Mon­

tana is referable to variety diversifolia, the most widespread^ and

var. perdissecta (Rydb.) C.L.Hitchc. The variety multisecta Wats, is

supposed to occur less commonly in Montana, but I have not seen any al­

pine material which could definitely be referred to this variety although

two collections from the Bridger Mountains appear to approach it, even

though they were considered closer to variety perdissecta. Ranunculus

eschscholtzii is a widespread polymorphic species differentiated into

several races which, in their extreme forms, are strikingly distinctive

and often considered as a separate taxa (Hitchcock et al,, 196It). All

of the alpine collections from Montana that I have seen appear to be

best referred to variety suksdorfii (Gray) Benson. Anemone occidentalis

and Romanzoffia sitchensis have been included in this group on the basis

of their overall distribution patterns, although these species both

reach their southern limits in the Rocky Mountains in northwest Montana

(Hitchcock et al., 1959, I96I4). Two other species, Hieracium gracile

and Anemone multifida have the same North American distribution pattern

as this group but are also known from South America. They are widespread

in the Montana alpine.

b. The second group consists of species which are widely distributed

from Alaska along the Rocky Mountains, Sierra Nevadas, and in the Coast

Ranges of California, sometimes extending into the Cascades (Fig. 15)»

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J

Fig-ure 15« The general distribution of the cordilleran element widely distributed from Alaska along the Rocky Mountains, Sierra Nevadas, and in the Coast Ranges of California, sometimes extending into the Cascades.

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Figure 16, Gassiope mertensiana (Bong,) D,Don var. gracilis (Piper) C,L,Hitchc, The Montana alpine distribution of the cordilleran element widely distributed from Alaska along the Roaky Mountains, the Sierra Nevadas, the Coast Ranges of California, and the Cascades,

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They are; Agrostis scabra Willd.^ Agrostis variabilis Rydb,, Arnica

latifolia Bong., Garex nigricans C.A.Meyer, Carex spectabilis Dewey,

Cassiope mertensiana (Bong,) G.Don, Danthonia intermedia Vasey.

Most of the alpine material of Agrostis scabra is referrable to the

variety geminata (Trin.) Swallen which is more restricted in distribution

than the typical, wide-ranging variety scabra. The high altitude form

of Arnica latifolia is the variety gracilis (Rydb.) Cronq. Cassiope

mertensiana includes several distinctive variants (Hitchcock et al.,

1959) of which only the variety gracilis (Piper) C.L.Hitchc, is known

from Montana. Danthonia intermedia is a wide-ranging species extending

east in Canada to Newfoundland and Quebec and south to northern Michigan.

Agrostis variabilis is more restricted in its range than this, occurring

from British Columbia and Alberta south to Colorado and California.

c. The third group consists of species which are widely distributed

from British Columbia and Alberta along the Rocky Mountains and Sierra

Nevadas, sometimes extending into the Cascades but not in the Coast

Ranges of California. (Fig. 17). Species in this group are; Arabis

lemmonii Wats., Arabis lyallii Wats., Arnica mollis Hook,,Arnica

rydbergii Greene, Carex albo-nigra Mack., Carex illota Bailey, Carex

nubicola Mack,, (includes G. haydenniana Olney), Carex phaeocephala

Piper, Draba paysonii Macbr., Eriogonum ovalifolium Nutt., Gaultheria

humifusa (Grah.) Rydb., Gentiana calycosa Griseb., Lewisia pygmaea (Gray)

Robins., Luzula glabrata (Hoppe) Desf., Mimulus tilingii Regel,

Pedicularis contorta Benth., Phacelia hastata Dougl,, Poa epilis Scribn.,

Poa rupicola Nash., Ribes montigenum McClatchie, Senecio fremontii T.&G.,

and Senecio werneriaefolius Gray, many of which are widely distributed

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Figure 17. The general distribution of the cordilleran element widely distributed from British Columbia and/or Alberta along the Rocky Mountains and Sierra Nevadas, sometimes extending into the Cascades but not the Coast Ranges of California.

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Figure 18, Senecio fremontii T, & G, var, fremontii. The Fontana alpine distribution of the cordilleran element widely distributed from British Columbia and ^Iberta along the Rocky Fountains and Sierra Nevadas but not in the Coast Ranges of California.

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in the Montana alpine.

Arabis lemmonii is divided into three infraspecific taxa (Hitchcock

et al., I96I4), two of which are present in the Montana alpine. These

are variety drepanoloba (Greene) Rollins and variety lemmonii. Draba

paysonii is divided into two infraspecific taxa, variety paysonii and

variety treleasii (Schultz) C.L.Hitchc. Although the two varieties are

supposed to have distinct ranges (Hitchcock et al,, I96U), the Montana

material indicates that they occur together here. Eriogonum ovalifolium

is a very wide-ranging species and one that is influenced considerably

by local ecological conditions (Hitchcock et al., I96U). According to

Hitchcock et al. (196U), several variants are recognizable, of which all

the Montana alpine material I have seen is referrable to the variety

depressum Blank. Gentiana calycosa is a wide-ranging species showing

much variation (Hitchcock et al,, 1959). The variety asepala (Maquire)

C.L.Hitchc. occurs in Montana (Hitchcock et al,, 1959) although all the

material I have seen appears to best fit variety obtusiloba (Rydb.)

G.L.Hitchc. Lewisia pygmaea is divided into two varieties (Hitchcock

et al., I96U) of which the variety pygmaea is the common form in the

Rocky Mountains. Mimulus tilingii is separated into two varieties but

only variety tilingii seems to occur in Montana. Four varieties of

Phacelia hastata are recognized (Hitchcock et al., 1959), of which va­

riety alpina (Rydb.) Cronq, occurs in alpine areas in south-central and

southwest Montana and the variety leptosepala (Rydb.) Cronq. in similar

areas of northwest Montana,. Pedicularis contorta is divided into two

similar but well-marked varieties (Hitchcock et al., I96I); the variety

ctenophora (Rydb.) Nels. & Macbr. and the variety contorta. The former

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is occasionally found as far west as Lima, in Montana, but is more com­

mon in the Big Horn Mountains of Wyoming. Hitchcock et al. (I96I) re­

port an isolated population in Idaho County, Idaho which has pink

flowers as in the variety ctenophora but is glabrous as in the variety

contorta. The proper taxonomic status of these has not yet been deter­

mined. A collection fitting this description was made on Trapper Peak

in the Bitterroot Mountains of Ravalli County, Montana where the variety

contorta was also collected. This would seem to indicate that the first

population is not geographically isolated. Although the variety

ctenophora was not collected here, it is quite probable that these two

populations represent hybrids formed by the two varieties and that the

latter are not, therefore, as well separated taxanomically as claimed,

Salix nivalis is questionably divided into two phases (Hitchcock et al.,

I96L) of which all of the material I have examined seemed best placed

in variety nivalis. The variety saximontana (Rydb.) Schneid. shares a

nearly co-extensive range with the variety nivalis and would be expected

to occur in the Montana alpine, Senecio fremontii has three infraspeci­

fic races of which only the variety fremontii occurs in Montana,

d. The fourth group consists of species which are distributed in

the Rocky Mountains and Sierra Nevadas but not in the Coast Ranges of

California, and reach their northern limits in southwestern or south-

central Montana (Fig, 19). They include; Agropyron scribneri Vasey,

Astragalus tegetarius Wats., Carex scopulorum Holm, Collomia debilis

(Wats.) Greene, Haplopappus suffruticosus (Nutt.) Gray, Hulsea algida

Gray, Ivesia gordonii (Hook.) T. & G,, and Synthyris pinnatifida Wats,

of which the material I have seen is referable to the variety canescens

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Figure 19. General distribution of the members of the cordilleran element which are distributed in the Rocky Mountains and the Sierras but not in the Coast Ranges of California and reach their northern limits in southwestern and south-central Montana.

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Figure 20, Astragalus tegetarius Wats. A representative Montana distribution of members of the cordilleran element which are distributed in the Rocky Mountains and Sierra Nevadas but not the Coast Ranges of California, and reach their northern limits in southwestern and south- central Montana.

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(Pennell) Cronq. All of these species appear to be south of the drift

border in Montana with the exception of Garex scopulorum which Bamberg

(1961 ) reports from Glacier National Park. This station is several

hundred miles north of the rest of the apparent range of the species and,

although the report may be valid, I doubt it on the basis of our present

knowledge. Four varieties of Collomia debilis may be recognized (Hitch­

cock et al., 1959), of these both variety debilis and variety camporum

Pays, reach Montana, but the alpine material I have seen seems best

referred to the variety debilis.

B. Species restricted to Rocky Mountains, a. The first group con­

sists of species which are widespread in the Rocky Mountains (Fig.21).

They are: Agrostis humilis Vasey, Antennaria lanata (Hook.) Greene,

Artemisia michauxiana Bess., Garex nova L.H,Bailey, Garex pyrenaica

Wahlenb., Gastilleja rhexifolia Rydb., Glaytonia megarhiza (Gray) Parry,

Dodecatheon pauciflorum (Durand) Green, Draba incerta Pays., Erigeron

simplex Greene, Haplopappus lyallii Gray, Juncus parryi Engelm., Luzula

wahlenbergii Rupr., Fhyllodoce glanduliflora (Hook.) Goville, Fhyllodoce

X intermedia (Hook.) Gamp., Potentilla ovina Macoun., Potentilla

quinquefolia Rydb., Salix cascadensis Cock., Saxifraga adscendens L.,

Saxifraga lyallii Engl., Senecio porteri Greene, Telesonix jamesii (Torr.)

Raf., and Townsendia parryi Eat.

Glaytonia megarhiza is widely distributed but disjunct over its

total range (Hitchcock et al,, 196ii) although variety megarhiza , which

occurs in Montana, is fairly common here. Dodecatheon pauciflorum is a

very wide-ranging species which extends from Alaska south to Mexico and

east to Pennsylvania, The reduced form of alpine situations in the

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Figure 21. General distribution of cordilleran element restricted to and widespread in the Rocky Mountains,

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Figure 22. Erigeron simplex Greene. The Montana alpine distribu­ tion of a species restricted to and widespread in the Rocky Mountains.

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Rocky Moimtalns may, according to Hitchcock et al. (1959), be awarded

recognition as an infraspecific variant, variety watsonii (Tidestr.).

Saxifraga lyallii is the only member of this group which has its

southern limits in Montana,extending south in western Montana to

Beaverhead County (Hitchcock et al., 1961), although I have seen no

alpine material from any farther south in Montana than the Mission Moun=

tains. Senecio porteri is reported by Hitchcock et al, (1955) only

from the Wallowa Mountains of northeastern Oregon and from Colorado.

Bamberg reports this species for the Beartooth Plateau which, if correct,

would give it a distribution pattern similar to the rest of the group.

All of the Montana collections of Telesonix jamesii are referred to the

variety heucherformis (Rydb.) Bacigalupi. The species is usually but

not always found on limestone (Hitchcock et al., I96I). Two species in

the group have a wider distribution or are closely related to species

occurring elsewhere. Antennaria lanata is related to Antennaria carpath-

ica (Wahl.) R.Br. of the Old World (Hitchcock et al., 1955) and both

are probably derived from a once more wide-ranging species, probably of

northern origin. Saxifraga adscendens is included here although its

North American variety, oregonensis (Raf.) Breit, occurs as far north in

the Rockies as Alberta and British 'Columbia, It also has an European

variety.

b. The second group consists of species with northern limits in

southwestern or south-central Montana (Fig. 23). They are : Artemisia

scopulorum Gray, Galtha leptosepala DC., Garex chimphila Holm,

Gastilleja pulchella Rydb., Ghaenactis alpina (Gray) Jones, Erigeron

ursinus D.C. Eat., Haplopappus pygmaea (T, & G.) Gray, Hymenoxys

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Figure 23. General distribution of cordilleran element restricted to the Rocky Mountains with northern limits in southwestern or west- central Montana,

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Figure 2h. Mertensia alpina (Torr.) G.Don. The Montana alpine distribution of a species restricted to the Rocky Mountains which reaches its northern limits in west-central Montana,

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grandiflora (T. & G.) Parker, Kobresia macrocarpa Glokey, Mertensia

alpina (Torr.) G.Don, Mertensia perplexa Rydb., Penstenion montants

Greene, Primula parryi Gray, Townsendia montana Jones, Trifolium

dasyphyllum T, & G., Trifolium nan van Torr,, and Trifolium parryi Gray.

These species apparently do not occur in the Bitterroot Mountains north

of Trapper Peak, Ravalli County.

According to Hitchcock et al, (196U), Galtha leptosepala probably

extends north from Montana to Alaska, although he saw no material from

north of the Canadian border, A local variant of the species, variety

sulfurea C.L, Hitchc., occurs in the Lost River Mountains of Custer

County, Idaho.

c. The third group consists of species which are disjunct. They

are Castilleja occidentalis Torr,, Draba apiculata C.L,Hitchc., and

Erigeron lanatus Hook.

The first species is apparently related to the boreal Castilleja

pallida (L.) Spreng. from which it may be derivative. Draba apiculata

is composed of two races and has a range which is very disjunct

(Hitchcock et al., 196I4.). The variety apiculata is reported from

northeast Park County, Montana, the Grand Tetons of Wyoming, and in the

Unita and Wasatch Mountains of Utah, The variety daviesiae C, L, Hitchc,

is known only from the Bitterroot Mountains of Montana, According to

Hitchcock et al,, it should be found in areas between the latter and

that of variety apiculata, Erigeron lanatus is common in the high

mountains of northwest Montana, southern Alberta, presumably southern

British Columbia, and was recently collected in Colorado (Hitchcock et

al,, 1961 ).

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U* Pacific Coast Element

The fact that the Pacific Coast flora contributes in certain

localities to the Rocky Mountain flora has been known for a long time

(Kirkwood, 1927). The effect of the Pacific Coast flora is most no­

ticeable in the latitudes of the westerlies, where there are strongly

developed storm tracts. Here, steady wind following the storm tracts

blows directly inland from the coastal strip. The climate and its

characteristic flora are extended far inland as a peninsula with all

zones within this climatic peninsula showing the effects of the coastal

climatic influence in their floras. They retain their typical dominants

but add a few species which have distribution centers lying along the

coast (Daubenmire, 19U3).

The fourth major floristic element in the Montana alpine flora is

thePacific Coast element (Fig, 25). The species included in this cate­

gory are; Aster alpigenus (T. & G.) Gray, Campanula parryi Gray,

Campanula scabre11a Engelm., Cryptantha nubigena (Green) Pays,, Eriogonum

pyrolaefolium Hook., Saxifraga tolmei T. & G., Senecio subnudus DC,, and

Veronica cusickii Gray,

Aster alpigenus, Campanula parryi, Senecio subnudus, and Veronica

cusickii are referred here with some hesitancy, as they are actually

absent from the Coast Ranges although Aster alpigenus and Senecio

subnudus occur in adjacent mountains to the east. Three fairly well-

defined geographic races of Aster alpigenus may be distinguished

(Hitchcock et al,, 1955), and of these, the variety haydenii (Porter)

Cronq. occurs in Montana. Campanula parryi reaches its western limits

in the Wenatchee Mountains of Washington and extends east to central

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Figure 25* General distribution of Pacific Coast element in the Montana alpine flora.

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and north-central Idaho and adjacent Montana (Hitchcock et al., 1959).

The alpine material of Campanula parryi which I have seen is referred

to variety idahoensis McVaugh. Veronica cusickii extends from the

Wallowas and higher Blue Mountains of northeastern Oregon across north

and central Idaho to western Montana (Hitchcock et al., 1959). The lat­

ter two species might better be included with the cordilleran element

although their ranges occur only within the weather peninsula in Idaho

and Montana, with the Pacific Coast element. Eriogonum pyrolaefolium

consists of two varieties (Hitchcock et al., 196U). The Montana

collections represent the variety coryphaeum T. & G. Only the variety

ledifolia (Greene) Engl. & Irmsch. of Saxifraga tolmiei was seen in the

Montana alpine material I have examined. Although it is reported by

Hitchcock et al* (1961) only from St. Mary's Peak in the Bitterroot

Mountains, it was collected on Trapper's Peak and Lolo Peak and would

therefore appear to be more widespread in the Bitterroots than previously

believed. In addition, Bamberg (I963) reports this species from the

Highland Mountains east of the Bitterroots.

Campanula parryi, Campanula scabre11a, Eriogonum pyrolaefolium,

Saxifraga tolmiei, and Veronica cusickii appear to be confined to that

area of western Montana where the influence of the Pacific Coast cli­

mate is most apparent. This includes the area east of the Continental

Divide from the Bitterroot Mountains in the south to Glacier National

Park in the north. The other three species included in the Pacific

Coast element are apparently more widely distributed in western Montana

extending as far east as the Beartooth Plateau area.

Bamberg (1963) includes several additional species as representative

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Figure 26. Eriogonum pyrolaefolium Hook. var. coryphaeum T, & G, The Montana alpine distribution of a species in the Pacific Coast element.

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of this element in the alpine flora of the state. These are the follow­

ing: Gollomia debilis (Wats,) Greene, Garex leporinella Mack,,

Galyptridinm umbellatnm Greene, Hulsea algida Gray, Pedicularis contorta

Benth, Penstemon confertus Dougl., and Sednm donglasii Hook. Garex

leporinella was not known from Montana (Mackenzie, 1931-193$)^ however,

Bamberg (1963) reports the species from Glacier National Park and, if

the identification is correct, the species may be considered part of the

Pacific Goast element in the alpine flora, I have included Gollomia

debilis and Hulsea algida with the species occurring in the southern

Cordillera of the Sierra Nevadas and/or the southern G as cades and the

southern Rocky Mountains. These are species which reach their northern

limit in the Rocky Mountains in southwestern and west-central Montana.

On the basis of its overall distribution I feel that Pedicularis

contorta is better referred to the category of central cordilleran spe­

cies which occur in both the Sierra Nevada and Rocky Mountains, I

have considered Penstemon confertus, Sedum stenopetalum Pursh, (includes

S, douglasii), and Spraguea umbellatum Torr. (includes Calyptridium

umbellatum) to be more representative of the lower montane flora than

of the alpine, although they can be referred to the Pacific Goast ele­

ment in the montane flora.

5. Endemics

The last major category of species making up the Montana alpine

flora are those which are classified as endemics. These are species

with a distribution pattern confined to a single region or locality

although, as Polunin (i960) points out, the term is largely relative

and therefore certain of these species might better be placed in a

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category which implies wider distribution.

The first major group of endemics included in this category are

those which are restricted to the Yellowstone region (Fig. 27). General­

ly the boundaries of this region are from the southwestern corner of

Montana to the northwestern corner of Wyoming, although they may be

extended to central Idaho in the case of certain species. These species

are: Castilleja nivea Pennell & Ownbey, Ghinophila tweedy! (Canby &

Rose) Henderson, Circium polyphyllum (Rydb.) Petrak., Erigeron rydbergii

Cronq., Pedicularis cystopteridiafolia Rydb., Pedicularis pulchella

Pennell, Salix dodgeana Rydb., Trifolium haydennii Porter.

Pedicularis cystopteridiafolia is related to the circumboreal

species, Pedicularis sudetica Willd, which is not known to occur in

Montana but is reported from Colorado, the latter population approaching

the former (Hitchcock et al., 1959)» If it arose from an isolated frag­

ment of Pedicularis sudetica then it probably has evolved since the

Pleistocene from a population which became isolated during the retreat

of the glaciers. Another species which may have evolved in a similar

manner is Salix dodgeana. It is related to the nearly circumboreal

Salix rotundifolia Trautv,

A second group of endemics is one whose principal distribution is

in western Montana but which may occur also from southern Alberta and

British Columbia to northwestern Wyoming (Fig. 29)® These species ares

Aquilegia jonesii Parry, Cardamine rupicola (Rydb.) C.L,Hitchcock,

Car ex plectocarpa F.J,Hermann, Papaver pygmaeum Rydb., Penstemon

ellipticus Coult.& Fish., Phacelia lyalli (Gray)Rydb,, Senecio

megacephalus Nutt., Stellaria americana (Porter) Standi., Synthris canbyi

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7-

Figure 27. General distribution of species endemic to the Yellowstone region.

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vlngston

Figure 28, Erigeron rydbergii Cronq, The Montana alpine distribu­ tion of a species endemic to the Yellowstone region.

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Pennell, and Townsendia spathulata Nutt.

Cardamine rupicola is known from the Swan and Mission Mountains of

Montana. Garex plectocarpa is known only from Logan Pass, Glacier Na­

tional Park, although a collection from Sperry Glacier, Glacier National

Park, approaches this species. Papaver pygmaeum only occurs in the

higher mountains of northwestern Montana and southeastern Alberta. Ac­

cording to Hitchcock et al, (I96I) , a somewhat similar species, Papaver

nudicaule L., which is present in the arctic, extends south in the

Rocky Mountains to Alberta and reoccurs in east-central Utah and Colo­

rado, It is conceivable that Papaver pygmaeum arose from an isolated

population of Papaver nudicaule. Synthyris canbyi is known only from

the Mission Range in Montana. It is apparently related to Synthris

laciniata (Gray) Rydb. of Utah (Hitchcock et al., 1959) and may have de­

veloped from this species during its northern migration under the influ­

ence of isolation.

Aquilegia jonesii ranges from southern Alberta to northwestern

Wyoming (Fig. 31). The distribution fo this species is notably disjunct

as the plant occurs only in limestone areas (Hitchcock et al,, 196U).

It has been proposed that the main soil factor involved in the distribu­

tion of alpine plants appears to be the degree to which the humus col­

loids are saturated with cations, predominantly calcium ions (Rune,

1953)» There is no indication that the local occurrence or the general

distribution of any alpine species is determined by more specific fac­

tors, such as amount of potassium or phosphorus or micronutrient salts

available, except for a limited number of species associated with mag­

nesium rich soils or with copper ores (Rune, 1953)»

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Figure 29. General distribution of species endemic to western Montana and adjacent areas.

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f

vingston

Figure 30, Stellaria americana (Porter) Standi, The Montana alpine distribution of a species endemic to western Montana and adjacent areas.

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*V

Xk n

{20 vingston

m

Figure 31. Aquilegia jonesii Parry, The Montana alpine distribu­ tion of a species endemic to western Montana and adjacent areas and further restricted to calcareous areas.

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Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. DISCUSSION

"Only a very few plants with a world—wide range show no taxonomic

variation" (Hulten, 19^9). Such plants are presumably very old forms

that have had their biotypes reduced by the selection pressures of the

different climatic conditions under which they have grown. Most plants

are more or less clearly separable into geographical races which, in

turn, can be broken down into local populations. The races, whether

they are large or small, are frequently described as distinct species

without mention of the earlier known species with which they form world­

wide taxa (Hult^, 1959). As these species or their close relatives may

be concealed under a variety of names, each of which refer to a small

geographical race or to isolated populations in Europe, Asia, or America,

the difficulties in trying to map the total area of the world-wide taxon

are obvious. The latter should always be interpreted in its widest sense

and its subdivisions, where known, should be indicated. This also applies

to taxa whose distribution cover smaller areas if they show geographic

variation patterns.

When the distribution of many circumpolar arctic-alpine species is

mapped in some detail they are found to have ranges which consist of a

more or less continuous ring around the North Pole and another concen­

tric, but broken ring along their southern boundary (Hulten, 1959).

Hulten indicates that when the ice retreated the arctic species

were able to follow it north to a certain extent but that the increasingly

warmer and drier conditions which arose, combined with the rapidly

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spreading forests, soon exterminated them in the lowlands. They could

survive to the present day, however, in the mountains where they be­

came isolated on each due to the disjunctness of the mountain groups*

They lost most of their bio types due to the changing conditions and

their populations in the different mountain groups evolved into forms

which were not uniform. This would explain why the arctic-alpine spe­

cies often have several closely related but different counterparts

along the southern rim of their present range (Hult^, 1959), According

to Hulten the northern ring probably developed after the glacial period

through a fusion of populations which had survived the glaciation in

different réfugia after the glaciation had disrupted the parent species*

pre-glacial circumpolar distribution*

There are two possible origins for species which are arctic-alpine

but not circumpolar or nearly so* The first is that these species

originated in the arctic and became part of the alpine flora through

migration. The second is that these species originated in the alpine

areas to the south and migrated to the arctic probably during or just

after the Pleistocene*

An arctic-alpine species which originated in the arctic would not

necessarily have a circumpolar distribution pattern. It would seem

likely, however, that if it was one of those species which range from

the European arctic across that of Asia and into that of North America

and/or Greenland that it probably had an arctic origin prior to the

Pleistocene and migrated south to become part of the alpine flora. This

assumption follows from their fairly extensive distribution in the arc­

tic* However, a particularly aggressive alpine species which migrated

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to the arctic during or just after the Pleistocene might also be expec­

ted to develop an extensive range in the arctic in the period of time

available following its introduction there. It would seem that those

arctic-alpine species which are restricted to the North American arctic

or have reached only to far eastern Asia or possibly western and/or

eastern Greenland would be the more likely ones to have had an alpine

origin in North America,

The origin of those species in the group whose distribution, or

the distribution of whose closely related species, is in the alpine

areas in the temperate latitudes must be considered. These species,

if present only in the alpine areas of the western hemisphere and in the

arctic, could conceivably have originated in the alpine location and

migrated to the arctic, A group of closely related species in which

none of the individuals are circumpolar, but in which the group is,

whose members occur in the alpine areas of Europe and North America

probably had an arctic origin. One of the group could have had alpine

origin if it had originated in one of alpine areas and migrated north

during one part of the Pleistocene and south into the alpine of another

continent during a later part of this period.

The rarest of the arctic-alpine species in the alpine are those

with disjunct alpine colonies. These are separated from the main dis­

tribution area of the species by a distance greater than their present-

day dispersal capacity. They seem to consist of those species which

are limited to the most mesic habitats (Weber, 1959), Johnson (1962)

proposes that in the Beartooth Plateau region the bog environments,

which are capable of dissipating the excessive summer heat, have per­

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mitted such species to persist there. Weber indicates that the most

productive of the alpine habitats, in reference to the number of arctic

disjuncts they harbor, are situated wherever high ranges tend to run in

an east—west direction and lie between or connect the principal north-

south -orienta ted mountains. These ranges are all rather similar in that

they are more constantly mesic throughout the year than contrasting

north-south-orientated ones* Slower run-off, late remaining snow beds,

development of some small bogs and many small lakes at the higher ele­

vations are the result of the less direct sunlight in these areas in the

summer months. In these regions the disjunct species occur on the north

slopes or at quite low elevations at the base of these slopes (Weber, 1961).

The Bitterroot Mountains in Ravallii County, Montana would seem to

fit these requirements and the collections of Gassiope tetragona from

St. Mary’s Peak in these mountains is that of such a disjunct species.

This species was collected on the north slope of an east-west ridge in

a pocket of snow accumulation where it would appear that the temperature

and more or less persistent snow would provide the necessary mesic hab­

itat for it to persist after the Pleistocene. As this particular area

of the Bitterroot Range (from Lolo Peak in Missoula County south to

Trapper Peak in Ravalli County) does seem to fulfill the criteria set

forth by Weber for localities having disjuncts, it is suspected that

further investigation of the Bitterroot Mountains will yield more of them,

The southern limits of different arctic-alpine species occur at dif­

ferent latitudes. This would seem to indicate that their limits are con­

trolled in part by temperature, and Dahl (1955) thinks that in the alpine

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the average yearly maxim-um summer temperature is the critical one.

The distribution patterns of the cordilleran element can be ex=

plained mainly by age, presumed differences in migration abilities, and,

in some groups, by the influence of the Pleistocene glaciation. As has

been previously stated, the major assumption underlying this group is

that of an alpine origin. The first group of the cordilleran element

are those widespread species which have extensive ranges extending

south from Alaska into the Rocky Mountains, the Sierra Nevadas and/or

the Coast Ranges of California probably originated prior to the

Pleistocene and developed their present ranges under its influence. Be­

cause they are present in both the Sierra Nevadas and the Rocky Mountains,

it is believed that they originated in more northern alpine areas, pos­

sibly even in high boreal ones. Another group of widespread cordilleran

species with an extensive range does not extend as far north as Alaska,

Their ranges are extensive enough to indicate a relatively old, prob­

ably pre-Pleistocene, origin but apparently they were unable to migrate

any farther north than British Columbia or Alberta, Those species which

are present in both the southern Rocky Mountains and the Sierra Nevadas

have also been grouped under the wide-ranging species. Their origin was

almost certainly in the central or southern alpine areas of North Amer­

ica, They were not able to migrate to the north or, if they did, could

not persist in those northern alpine localities.

The second major group of the cordilleran element are those alpine

species restricted to the Rocky Mountains or which, in a few instances,

are also in adjacent Cascades in Washington, Those distribution pat­

terns in this group which were discussed were|first, those of the species

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fairly widespread in the Rocky Mountains; second, those of the species

more restricted to the central Rocky Mountains; and last, those of the

species restricted to the southern Rocky Mountains, It seems logical to

assume that the Rocky Mountains were the site for the origin of these

species, and that their migration was influenced by the factors previous­

ly mentioned, A noteworthy observation concerning these species which

are restricted to the southern Rocky Mountains or to the central Rocky

Mountains is that they apparently have little, if any, infraspecific

variation. Two possible explanations might be given for this. The first

is that there is actually no variation in these species and the second

is that these species have not been carefully enough studied to determine

what variation is present, A second observation regarding those species

which are primarily confined to the southern Rocky Mountains, but which

reach their northern limit in southwestern Montana, is that the latter

show no apparent correlation with any present day edaphic, climatic, or

physiographic factor. This might indicate that their isolation as a

group has been due to historical factors.

The endemic species in the alpine flora may be assigned to two pos­

sible origins. They may be very old species whose range was once far

more extensive that it is today or they may be a species of fairly re­

cent origin which has been unable to spread due to the unfavorable con­

ditions for migration since their origin. These species also exhibit the

apparent lack of subspecific variation that is shown by the groups of the

cordilleran species which were just discussed. Either very old species

depleted of biotypes and with a very restricted range, or young species

which have not migrated out of a limited area would be expected to show

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this lack of variation»

The present distribution patterns of species making up the alpine

flora of Montana have been most recently influenced by the Pleistocene

and its effects are most obvious» For this reason the patterns I have

discussed have been considered in light of their immediate pre-Pleisto­

cene form and their subsequent Pleistocene development. Neither the

origin of the pre-Pleistocene distributions nor that of alpine vegeta­

tion has been or will be extensively covered, however, some theories

regarding the origin of alpine and arctic vegetation will be briefly

noted.

Tundras or tundra-like landscapes have probably been in existence

since land vegetation itself. They may not have always existed in

present-day arctic areas. In fact, during the late Eocene and early

Oligocene, worldwide warmth permitted the existence of tropical and tem­

perate plants far north of their present ranges (Arnold, 1959)» Arctic

conditions at sea level were probably not in existence at this time and

tundra-like conditions existed only at high altitudes (Good, 1953).

These may have occurred only as small strips which were wide enough to

maintain localized tundra floras» During the progressive cooling of the

middle and late Tertiary, which culminated in the Pleistocene, these

tundra strips increased and, with their localized floras, moved down the

mountains and eventually stretched into the boreal areas abandoned by

the former Arcto-Tertiary flora which had migrated south (Arnold, 1959).

According to him, in the boreal area the tundra spread rapidly onto the

arctic plains and ultimately developed into a circumpolar tundra belt.

Some of the species in the retreating boreal vegetation may have become

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adapted to the changing conditions in this area and were incorporated

into the arctic flora.

By the end of the Pliocene modern tundra floras had become well

established in the Northern Hemisphere (Arnold^ 1959). If this occur­

red, then it may be said that the alpine flora antedates that of the

arctic, and that the alpine afforded a place of development for the

present arctic flora. According to Good (1953), however, during

the Cretaceous and earlier Tertiary physiographic conditions may have

been such that there were no elevations high enough to permit the exist­

ence of tundra environments. Such environments may not have developed

until after the orogeny which formed our present mountains in the later

Tertiary. The time lapse, however, between the orogeny of the Miocene

and Pliocene and the Pleistocene glaciation is long enough to have

allowed the development of tundra conditions in the mountains and they

therefore, probably existed before those in the arctic. Thus, the pre-

Pleistocene arctic flora probably originated, at least in part, from

that of the alpine (Good, 1953).

If the former did occur then closely related taxa with disjunct

areas in the alpine and arctic may well have developed from pre-

Pleistocene ancestors which grew in alpine areas of the temperate

latitudes. While it is probable that their distribution patterns were

greatly modified by the Pleistocene, it is possible that some of them

had developed ranges extending from alpine to arctic areas prior to the

Pleistocene, The result Pleistocene on these species may have been only

range expansions which were later disrupted by post-PleistoCene climatic

changes.

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Ideas regarding the origin and distribution of the arctic and

boreal flora have been stated by Hooker (1862), Fernald (192$, 1926),

Simmons (1913), Holm (1922), Marie-Victorin (1938), Wynne-Edwards (1937,

1939), Hulten (1937), and others. All these ideas have been reviewed

by Raup (19^1) and more recently, in part, by Flint (1957) with emphasis

on geological, particularly glacial data. They point up the fact that

there is still considerable difference of opinion in this area.

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A major problem encountered in this study was the determination of

which plants to include as 'alpine plants'. While the alpine zone is

generally agreed to be that region above timberline^ even though the

latter is not well-defined, definitions of what an alpine plant is vary

considerably. The majority of the plants which are found above timber-

line also extend, to varying degrees, below timberline. Some are found

only as low as the krummholz, some occur at moderate elevations in the

mountains, and others as low as inter-montarle valleys. Very few plants

have all their stations above timberline.

Our knowledge of the alpine flora and its distribution in Montana

is limited by the lack of collections from many of the alpine areas.

Areas from which alpine collections are noticeably lacking are the

Mission, the Swan, and the Flathead Ranges of northwestern Montana,

Numerous collections are available from those areas which are easily

accessible, such as the Logan Pass area in Glacier National Park, and

the Beartooth Plateau region in south-central Montana, Collections from

alpine areas which are less easily accessible are very limited.

Nevertheless, utilizing available herbarium collections, some field

collections and some published reports, the composition of the alpine

flora of Montana and the distribution of its species throughout the

state has been worked out in fair detail. This flora has been divided

into categories on the basis of the overall distribution patterns of its

species and their distribution in Montana, These patterns are usually

explainable on the basis of age, place of origin, and dispersal ability

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of the included species. Other major factors determining these patterns

are the edaphic, physiographic, and climatic ones. The most important

edaphic factors appear to be the pH of the soil and the amount of

available soil moisture. The most important of the physiographic fac­

tors are altitude, directional trends and degree of isolation of the

mountain systems. The temperature fluctuations and the resulting glaci­

ation of the Pleistocene have been two of the most influential of the

climatic factors in that they have caused the major migrations of alpine

and arctic species.

Field collections resulted in significant range extensions of two

arctic-alpine species, Androsace lehmanniana Spreng, and Gassiope

tetragona (L.) D.Don var, saximontana (Small) C.L.Hitchc, The collec-

I tion of the former from Divide Mountain, Glacier National Park, is a

western extension of the range of the first species and the collection

of the latter on St. Mary's Peak in the Bitterroot Mountains is a

southern extension of the range of the second species. There were no

new state records for any of the species collected and I have not at­

tempted to determine county records which have not been listed by Booth

(1959).

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. LITERATURE CITED

Alden, W.C. 1953. Physiography and glacial geology of western Montana and adjacent areas. U.S. Geol, Survey. Prof, Paper 231.

American Geographical Society of New York. 19ii5, Glacial map of North America. Special Papers^ Number 60 (Part 1) Western Half.

Arnold, Chester A. 1959. Some paleobotanical aspects of tundra development. Ecology UO; IU6 -IU8

Bamberg, Samuel A. 1961. Plant ecology of the alpine tundra areas in Montana and adj. Wyoming. M.A,Thesis. University of Colorado, Dept, of Biology. 163 p.

1963. Ecology of the vegetation and soils associ­ ated with calcareous parent material in the alpine region of Montana. Ph.D. Thesis. University of California, Davis. 91p.

Billings, W.D. and H,A, Mooney, 1959. An apparent frost hummocksorted polygon cycle in the alpine tundra of Wyoming, Ecology UO(l)s 16-20.

Bliss, L.C, 1956. A comparison of plant development in microenviron­ ments of arctic and alpine tundras. Ecol. Monogr. 26(U)s 303-337*

Booth, W.E. and J.C. Wright, 1959. Flora of Montana. Part II, Dicotyledons. Montana State College, Bozeman. 280 p.

Choate, Charlu M. 1963* Ordination of the alpine plant communities at Logan Pass, Glacier National Park, Montana. M.S. Thesis, Montana State University, Missoula, Montana, 113 P*

Clausen, J,, D.D. Keck, and W.M. Hiesey, 19U0. Experimental studies on the nature of species. I. Effects of varied environments on western North American plants. Carnegie Inst, Washington Publ., 520: 1-U52.

______, 19U?« Heredity of geograph- ically isolated races. A, N*, 81: 1114.-133*

o I9U8 , Experimental studies on the nature of species. III. Environmental responses of climatic races of Achillea. Carnegie Inst. Washington Publ,, 58ls 1-129.

Dahl, Eilif. 1955» Biogeographic and geologic indications of un­ glaciated areas in Scandinavia during the glacial ages. Bull. Geological Society of America. 66: 1^99-1520,

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Dahl, Eilif, 1959* Pleistocene history of the flora of the North Atlantic region with special reference to Scandinavia, Recent Advances in Botany, Ninth Inter, Botan, Congr. Is 919-928.

Daubenmire, R.F, 19U3, Vegetational zonation in the Rocky Mountains, Botanical Review 9(6); 325-393*

195U. Alpine timberlines in America and their interpretation, Butler Univ. Bot. Stud. 11; 119-136,

Davis, R,J. 1952. Flora of Idaho, W,C, Brown and Co., Dubuque, Iowa, 828 p .

Femald, M.L, 1925* Persistence of plants in unglaciated areas of boreal America. (Mem, Gray Herb, 2); Mem. Am. Acad. 15(3),

. 1926. The antiquity and dispersal of vascular plants. Quart, Rev, Biol. 1: 212-2h5<

Flint, R.F, 1961, Glacial and Pleistocene geology. John Wiley and Sons, Inc., New York and London, 553 P,

Good, Ronald. 1953, The geography of the flowering plants, Longmans, Green and Co., London, New York, and Toronto. 1152 p.

Hawkins, P.H, 1903, The alpine flora of Montana. M.S. Thesis, Montana State College, Bozeman. 2h p.

Hermann, F.J. I96U, A new Carex from Glacier National Park, Montana. Leaflets West. Bot. 10; 65-68.

Hitchcock, G.L., Arthur Cronquist, Marion Ownbey, and J.W. Thompson, 1955. Vascular plants of the Pacific Northwest. Part 5* Gompositae* Ünivèrsityicf Washington, Seattle!. 3U3p«

1959. Vascular plants of the Pacific Northwest. Part U, Ericaceae through Campanulaceae, University of Washington, Seattle. 510 p.

" 1961. Vascular plants of the Pacific Northwest, Part 3* Saxifragaceae to Ericaceae. University of Washington, Seattle, 613 p.

I96U. Vascular plants of the Pacific Northwest, Part 2. Salicaceae to Saxifragaceae, University of Washington, Seattle. 597 p.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. “ 9 3 “

Holm, T* 1922. Contributions to the morphology, synonymy, and geograph­ ical distribution of arctic plants. Rep, Canad, Arctic Exped,, 1913-18, 5(B).

Hooker, J.D. 1862. Outlines of the distribution of arctic plants, Trans. Linn. Soc. London 23: 201-3U8.

Hulten, E, 1937. Outline of the history of arctic and boreal biota during the Quartenary Period,

o 1939. Alpigenous and arctogenous plants; ranges of cir- cumpolar and arctic-montane plants. Ninth Inter, Bot, Congr, 1: 907-911.

Johnson, F.J. 1962, The occurrences of new arctic-alpine species in the Beartooth Mts. of Wyo.-Mont, Madrono, l6; 229-231.

_. and W.D, Billings, 1962, The alpine vegetation of the Beartooth Plateau in relation to cryopedogenic processes and pat­ terns. Ecol, Mono, 3 2 (2 )Î 103-133.

Kirkwood, J.E. 1927. The Pacific element in the Rocky Mountain flora. Northwest Science 1: lU.

Mackenzie, K.K. 1931-33» North American flora. Volume I8 , Gyperaceae- Cariceae, New York Botanical Garden, L?8 p,

Marr, J.W, 1938. Lee slope stands in the upper part of the forest- tundra ecotone on Niwot Ridge, Boulder County, Colorado, Jour, Colo-Wyo, Acad, Sci, L(10): Ll,.

_ , 1939. Forms of tree islands in alpine tundra, (Abstract). Jour, Colo.-Wyo. Acad, Sci, L(ll): 3h

_ , 1961, Ecosystems of the east slope of the Front Range in Colorado, Univ. of Colo. Studies Ser, D(8); I3U p.

Mason, H.S, 1939. Principles of geo, distribution. Madrono, 3:l8l- 190.

Montana Almanac. 1938. Montana State University Press, Missoula, Montana. U69 p.

Moss, E.H. 1939, Flora of Alberta, University of Toronto Press. Toronto, Canada, 3li6 p,

Munz, P.A. and David D, Keck. 1939. A California flora. University of California Press, Berkeley and Los Angeles, I68I p,

Nimlos, T,J. and R.C. McConnell. 1962. The morphology of alpine soils in Montana, Northwest Science, 36(U)s 99-112.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. — 9U=*

Perry, E.S* 1962. Montana in the geologic past. State of Montana, Bureau of Mines and Geology, Bulletin 26, ?8 p,

Polunin, Nicholas, 19^9. Gircumpolar arctic flora. Glaredon Press, Oxford. London, England, Slh p.

i960. Introduction to plant geography, McGraw-Hill Book Go., Inc., New York, Toronto, and London. 6U0 p,

Raup, H.M, I9UI, Botanical problems in boreal America. The Bot. Rev, 7(3 & U): lU?-2li7 .

Rune, Olof. 1953, Plant life on serpentines and related rocks in the north of Swedenî Acta phytogeographica Suecica, v. 37, 1-139,

Rydberg, P.A. 1913, Phytogeographical notes on the Rocky Mt. région. I, Alpine region. Bull, Torrey Bot. Glub, hOt 677-686,

, I9IU. Phytogeographical notes on the Rocky Mt. region. II, Origin of the alpine flora. Bull. Torrey Bot. Glub, Ulî 89-1 0 3 .

______• I9IU, Phytogeographical notes on the Rocky Mt. region. Ill, Formations in the alpine zone. Bull. Torrey Bot, Glub, Ulî U59-U82.

Simmons, H,G, 1913« A survey of the phytogeography of the arctic American archipelago, Lunds Univ. Arssk. N.F. Afd, 2, 9(19),

Taber, R.D., R.S, Hoffmann, T.J, Nimlos, and S, Bamberg, 1961, Alpine ecosystems of the northern Rocky Mountains, Bull, of the Ecological Soc. of Am., h2(ii)s liiO,

Turesson, G. 1922. The genotypical response of the plant species to the habitat, Heredetas. 3s 211-350,

, 1925, The plant species in relation to habitat and cli­ mate, Hereditas. 3 s 11:7-236,

______. 1930. The selective effect of climate upon the plant species. Hereditas, Iks 99-152.

United States Department of Agriculture. 19kl« Yearbook of Agriculture. Glimate and Man, U.S. Government Printing Office, Washington, I2U 7 p,

Victoria, Fr, Marie-. 1938, Phytogeographical problems of eastern Canada, Gontr, Lab, Bot. Iftiiv, Montreal, No. 30: Repr, from Am, Mid, Nat. 19: 189-558.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. - 9 5 -

Weber, W„A« 19U9o Preliminary observation on plant distribution pat­ terns in Colorado. I. The relict flora. Jour. Golo.-Wyo. Acad. Sci. k(l): U5-U6.

« 1959o Some features of the distribution of arctic relicts at their austral limits. Recent Advances in Botany. Ninth Inter. Botan, Congr. 1; 912-91^.

1961o Alpine floristic components of the southern Rocky Mountains. (Abstract), Bull. Ecol. Soc. Amer. U2(ii)s I6U,

Wynne-Edwards, V.G. 1937. Isolated arctic-alpine floras in eastern North America; a discussion of their glacial and recent history. Trans, Roy, Soc, Can. 111(5) 31: 1-26.

______, 1939. Some factors in the isolation of rare alpine plants, Trans. Roy. Soc. Gan. Ill 33(5): 35-U2.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. APPENDIX I

List of species constituting the lowland element in the Montana

alpine flora,

POLYPODIACEAE

Cryptogramma crispa (L,) R,Br, var, acrostichoides (R,Br,) Clarke. Dryopteris filix-mas (L,) Schott, D* phegopteris (L.) Ghr. Polystichum lonchitis (L.) Roth.

EQUISETAGEAE

Equisetum arvense L. E. variegatum Schleich.

LYCOPODIAGEAE

Lycopodium annotinujn L.

SELAGINELLAGEAE

Selaginella densa Rydb.

GRAMINEAE

Agropyron trachycaulum (Link.) Malte Agrostis alba L, A, exarata Trin. A. idahoensis Nash A. scabra Willd, var. scabra A. thurberiana Hitchc. Bromus pumpellianus Scribn, Galamagrostis canadensis (Michx.) Beauv. Co purpurascens R.Br. Deschampsia atropurpurea (Wahl.) Scheele Elymus cinereus Scribn. & Merr. Festuca idahoensis Elmer. F, saximontana Rydb. (see Moss) Hesperochloa kingii (S.Wats.) Rydb, Hordeum jubatum L. Oryzopsis exigua Thurb. Poa cusickii Vasey P. fendleriana (Steud.) Vasey P. gracillima Vasey P. interior Rydb, P. leptocoma Trin. P. longiligula S. & W.

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Poa reflexa Vasey & Scribn, P, secunda Presi, P. stenantha Trin, Trisetnm wolfii Vasey

GYPERACEAE

Carex atrosquama Mack, C, anrea Nutt, Pranklinii Boott, ? C, geyeri Boott. C, kelloggii Boott, C, limosa L, 0^. microptera Mack, C, pseudoscirnoidea Rydb, C, scoparia Schk. stenochlaena (Holm,) Mack, Eleocharis pauciflora (Lightf,) Link var, fernaldii Svenson Scirpus caespitosus L,

JUNGAGEAE

Luzula divericata S, Wats,

LILIAGEAE

Allium cernum Roth, A, schoenoprasum L, Galochortus apiculatus Baker nuttallii Torrey, Erythronium grandiflorum Pursh, Tofieldia glutinosa (Michx,) Pers, ssp, montana G,L,Hitchc, Stenanthium occidentale A, Gray Veratrum viride Att, Xerophyllum tenax (Pursh,) Nutt, Zygadenus elegans Pursh, Zm, venenosus S. Wats, var, gr amine us (Rydb,) Walsh,

ORGHIDAGEAE

Habenaria saccata Greene

SALIGAGEAE

Salix brachycarpa Nutt, var, brachycarpa 8, Candida Flues, S, commutata Bebb, drummondii Barratt, So myrtillifolia Anderss,

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BETULACEAE

Betula gland-ulosa Michx. var. glandulosa

POLYGONACEAE

Eriogonijm androsaceam Benth, E. caespitos’um Nutt, E. chrysops Rydb. Ê. flavum Nutt, ssp, flavum E, flavum Nutt, ssp. piperi (Greene) Stokes var, linguifoljum Gandg, !• flavum Nutt, ssp. piperi (Greene) Stokes var. piperi "E. umbellatum Torr, var. subalpinum (Greene) M.E.Jones Polygonum bistortoides Pursh. P. engelmannii Greene P. viviparum L. Rumex paucifolius Nutt.

PORTULACACEAE

Claytonia lanceolata Pursh, var. lanceolata Lewisia triphylla (Wats.) Robins, Spraguea umbellata Torr. var. umbellata

GARYOPHYLLAGEAE

Arenaria aculeata W ats. A, capillaris Poir, var. americana (Macquire) Davis A, congesta Nutt, var, congesta A. congesta Nutt. var. lithophila Rydb. A, macrophylla Hook. A, nuttallii Pax. var. nuttallii Gerastium arvense L. Silene douglasii Hook, var, douglasii S. parryi (Wats.) Hitchc, & Maquire Stellaria crassifolia Ehrh, longipes Goldie var. altocaulis (Hulten) C.L.Hitchc. S. umbellata Turex

RANUNGULAGEAE

Anemone parviflora Michjc. A. patens L, Aquilegia flavescans Wats, var, flavescens Delphinium bicolor Nutt, D. glaucescens Rydb. D. nuttallii Pritz. var. fulvum G.L.Hitchc. Ranunculus flammula L, var. filiformis (Michx.) Hook. Trollius laxus Salisb,

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CRUCIFERAE

Arabia drummondii Gray A* hir suta (L ,) Sc op, A. nuttallii Robins. Draba aurea Vahl, var. aurea D« praealta Greene D. stenoloba Ledeb. Physaria didymocarpa (Hook.) Gray var. didymocarpa Thlaspi fendleri Gray Sedum lanceolatum Torr. var. lanceolatum S. rhodanthum Gray stenopetalum Hook.

SAXIFRAGACEAE

Heuchera cylindrica Dougl. var, alpina Wats, Ho cylindrica Dougl. var. glabella (T. & G.) Wheelock Leptarrhena pyrolifolia (D,Don)~R.Br. Mite11a breweri Gray, M. pentandra Hook. Parnassia fimbriata Konig. var. fimbriata Saxifrage arguta D. Don S. debilis Engelm, ferruginea Grab, var, macounü Engl. & Irmsch. S. mertensiana Bong. S. oecidentalis Wats, S. rhomboidea Greene

GROSSULARIACEAE

Ribes cereum Dougl. R, lacustre (Pers.)Poir.

ROSAGEAE

Dryas drummondii Richards Potentilla arguta Pursh, var. convallaria (Rydb.)T. Wolf P. concinna Richards var, concinna P. fruiticosa L. P. glandulosa Lindl, var. intermedia (Rydb.)C.L.Hitchc. P. glandulosa Lindl. var, nevadensis Wats, P. glandulosa Lindl. var, pseudorupestris (Rydb.)Breit, P. gracilis Dougl, var, flabelliformis (Lehm.)Nutt, Spiraea densiflora Nutt, var. densiflora S, densiflora var. splendens (Baumann^G.L, Hitchc.

LEGUMINOSAE

Astragalus bourgovii Gray, A. vexilliflexus Sheld. var. vexilliflexus

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Hedysarvan bore ale Nutt, var, boreale H* sulphurescens Rydb, Lupinus argenteus Pursh, var, argenteus L, argenteus Pursh. var. depressus (Rydb,) G.L.Hitchc. L. lepidus Dougl. var. utahensis (Wats.) G.L.Hitchc. L. wyethii Wats. Oxytropis besseyi (Rydb.) Blank, var. argophylla (Rydb.) Barneby 0. campestris (L.) DG, var. cusickii (Greenin. ) Barneby 0. lagopus Nutt, var, atropurpurea (Rydb.) Barneby O, sericea Nutt, var, sericea 0. sericea Nutt, var, spicata (Hook.) Barneby Trifolium beckwithii Brew,

LINACEAE

Linum perenne L. var. lewisii (Pursh.) Eat. & Wright

HYPERIGAGEAE

Hypericum formosum H.B.K, ssp, scouleri (Hook.) G.L.Hitchc. var. nortoniae (M.E.Jones) G.L.Hitchc,

VIOLAGEAE

Viola adunca Sm. var, bellidifolia (Greene) Harrington

ONAGRAGEAE

Epilobium angustifolium L.

UMBELLIFERAE

Angelica dawsonii Wats. Gymopterus bipinnatus Wats. C, glaucus Nutt, Heracleum lanatum Michx. Lomatium ambiguutn (Nutt. ) Goult. & Rose La cous (Wats.) Goult. & Rose L. cusickii (S.Wats.) Goult. & Rose L« sandbergii Goult, & Rose Lo triternatum (Pursh.) Goult. & Rose ssp. triternatum Musenion vaginatum Rydb.

ERIGACEAE

Arctostaphylos uva-ursi (L.) Spreng, Kalmia polifolia Wang, var, microphylla (Hook.) Rehd, Menzlesia ferruginea Smith var. glabella (Gray.) Peck Pyrola minor L. P. uniflora L. Vaccinium globulare Rydb. V. myrtillus L, V. scoparium Leiberg

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PRIMULACEAE

Dodecatheon jefferyi Van Houtte Douglasia montana Gray

GENTIANACEAE

Frasera speciosa Dougl. Gentlana affinis Griseb. G. amarella L. G. dentosa Rottb.■ Swerta perennis L.

POLEMONIACEAE

Gilia spicata Nutt. Phlox albomarginata M.E» Jones P* diffusa Bentho var. longistylis (Wherry) Peck P. pulvinata (Wherry) Cronq. Polemonium pulcherrimum Hook, var, pulcherrimum

HÏDROPHYLLACEAE

Phacelia heterophylla Pursh. P* sericea (Grah.) Gray var. sericea

BORAGINAGEAE

Eritrichium howardii (A.Gray) Rydb. Hackelia floribunda (Lehm.) Johnst. H. jessicae (McGregor) Brand Mertensia ciliata (Torr.) G.Don var. ciliata M. oblongifolia (Nutt.) G.Don M. viridis A. Nels* Myosotis sylvatica Hoffm. var. alpestris (F.W.Schmidt) Koch

SGROPHULARIAGEAE

Besseya rubra (Dougl.) Rydb. Bo wyomingensis (A.Nels) Rydb. G as tille j a crista-galli Rydb. flava Wats. Mimulus lewisii Pursh. Pedicularis bracteosa Benth. var. bracteosa P. bracteosa Benth. var. canbyi (Gray) Cronq» P. bracteosa Benth. var. siifolia (Rydb.) Gronq. P. parryi Gray. var. purpurea Parry P. racemosa Dougl. var. alba (Pennell) Gronq. Petistemon attenuatus Dougl. var. attenuatus P. attenuatus Dougl. var. pseudoprocerus (Rydb.) Gronq. P. fructicosus (Pursh) Greene var. fructicosus P. humilis Nutt.

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Fenstemon procerus Dougl. var. procerus? (Hook.) Gronq. Veronica peregrina L. var. xalapensis (H.B.K.) St.John & Warren

LENTIBULARIACEAE

Pinguicula vulgaris L.

RUBIACEAE

Galium boreale L.

VALERIANACEAE

Valeriana edulis Nutt, V, sitchensis Bong,

GAMPANÜLAGEAE

Gampanula rotundifolia L,

GOMPOSITAE

Agoseris aurantica (Hook.) Greene var. aurantiaca A, glauca (Pursh.) Raf. var. dasyphylla (T. & G.) Jeps. Anaphalis margaritacea (L.) B, & H. Antennaria corymbosa E, Nels, A, dimorpha (Nutt.) T. & G. A, parviflora Nutt, A, rosea Greene Arnica cordifolia Hook, var, cordifolia A. cordifolia Hook, var. pumila (Rydb.) Maquire A, diversifolia Greene A, longifolia P.O. Artemisia campestris L. ssp, borealis (Pall.) Hall & Clem, var, borealis A, frigida Willd. Aster perelegans Nels, & Macbr, Grepis runcinata (James) T. & G. ssp, runcinata Erigeron ochroleucus Nutt, var, scribneri (Canby) Gronq, E, peregrimus (Pursh, )Greene ssp. callianthemus (Greene)Gronq, var. call- ianthemus E, peregrimus (Pursh. )Greene ssp, callianthemus (Greene )Gronq. var, scaposus (T,&G,)Gronq, Gaillardia aristata Pursh, Senecio amplectens Gray. var. holmii (Greene) Harrington So canus Hook. S, crassulus Gray S, integerrimus Nutt, var, exaltatus (Nutt.) Gronq. So triangularis Hook, Taraxacum officinale Weber.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. APPENDIX II

Distribution of species occurring in the Montana alpine. A plus

sign indicates a collection which I have seen in the herbarium or col­

lected in the field and a minus sign indicates a reported occurrence

which I have not verified. All reported occurrences are those of

Bamberg (I9 6 I, 1963) except for the following species which are re­

ported by Johnson (1962) from the Beartooth Plateau: Carex capitata,

Carex misandra, Draba glabella, Eriophorum callitrix, Festuca baffinensis,

Juncus albescens, Juncus castaneus, Kobresia macrocarpa, and Phippsia

algida. Bamberg did not recognize any taxa at the infraspecific level

and certain of his reported occurrences have not been included if, ac­

cording to Hitchcock et al. (195$, 1959, 1961, 196U), infraspecific

variation occurs in the species and if more than one of the variants

occurs in Montana. An asterisk following the species name indicates

its presence in the arctic according to Polunin (1959)» A question mark

following the species name indicates those species which have been dis­

cussed in the text and questionably considered alpine species but which

might better be classified with the lowland element. The species making

up the lowland element have not been separated in this table but a list

of them is given in Appendix I.

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O a g ; I Ü P I co +3 Jh t>s -aî O O a Ot (2 Ml

Lewis Range *■ + +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts,

Big Snowy Range

Highland Mts, -loU-

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Lewis Range + + 4-

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range + 4 -

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts,

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Mission Range

Swan Range

Flathead Range

Bitterroot Range +

Big Belt Mts, +

Little Belt Mts.

Flint Creek Range -

Anaconda Range +

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range +

Gallatin Range

Absaroka Range + + Beartooth Plateau *

Crazy Mts.

Big Snowy Range

Highland Mts,

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Mission Range

Swan Range

Flathead Range + + +

Bitterroot Range ♦ *■ ♦

Big Belt Mts. - +

Little Belt Mts. -

Flint Creek Range -

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range +

Gallatin Range +

Absaroka Range +

Beartooth Plateau *

Crazy Mts « -

Big Snowy Range

Highland Mts.

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Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts*

Flint Greek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts* ♦

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts*

Big Snowy Range

Highland Mts*

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Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts,

-1 0 9 .

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Mission Range

Swan Range

Flathead Range

Bitterroot Range +

Big Belt Mts.

Little Belt Mts.

Flint Greek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range ♦

Gallatin Range

Absaroka Range

Beartooth Plateau ♦

Crazy Mts.

Big Snowy Range

Highland Mts.

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Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big.Belt Mts.

Little Belt Mts. -

Flint Creek Range »

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts* -»

Madison Range

Gallatin Range

Absaroka Range 4-

Beartooth Plateau + *

Crazy Mts*

Big Snowy Range

Highland Mts*

-Ill-

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Lewis Range ■4- + ♦ + + + *■ +

Mission Range

Swan Range + ♦

Flathead Range ♦

Bitterroot Range +

Big Belt Mts. ■¥

Little Belt Mts. •¥

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range ♦

Absaroka Range +

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-112 -

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Q)

-P a r t to +5 I o to CO Oj % "H c 5 p - p I U •H ctf c g 0) cO -S Ü -o HO O g -p O g -p 0) .5 0 fH -p fH iH CO W HO c: 0) P CO S o tfl CO -P ra •H ttJ s 1—i to ctJ •H d '—1 P •H § rt g XI •H >> C I Xi •H rH Q* to JC 0) i o a> (0 p ft XJ o •H CO A o n) g 1 «H k HO I I I 'ôtf s O . k, « 3 o I CO E-t EH l I M CH Ml DC Lewis Range + +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-113-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CO

•H1 a nJ cd * CD 03 •H % cd cd I *H Ü F, (Q cd I •H -P C •P cd a -P «2 rt 03 cd 03 •H % U a X} -o P § % Ü -P •H Cd 0 03 Ü o bO :p (0 X3 03 X u 1 cd O O •H > Ü o Ü XI •è IÜ0 & Ï CO Co| tO| 031 W| 03| W| tÔ| o3f 031 o5| Ml Lewis Range ♦ + + + <#■

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts#

Little Belt Mts*

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts# +

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts#

Big Snowy Range

Highland Mts#

-Iiu-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ‘H 0> % A m to o o a CM o CO % 1 to *HI •ë CD f H T3 CO o CO 0) ■ H u O CO n j c CO ÇL, rH f t Cm o -p ft rH I—4 •H O tu 0) ■H o Cm u Cm T) aJ w c u CO rH (0 !>> rH O § 0 U cd ft cd (d CtO x: r—1to fH > u k 0 o Cm CO I I I o Cm a O > f t 0) • . . a pq 1 M l Ml M l M l Ml Ml Ml Lewis Range + +

Mission Range

Swan Range

Flathead Range 4-

Bitterroot Range +

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-115-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. %

% o I+) CO 0 M O N •-d •H CD cd cd -p •H r H o CD CD % CO •HO cd •H C «H cd EE ccJ I CO •H t—I N bO bO be "0 •H bO U O •H >> !>j CO •H q) JC B A A ■x> Â s •H O) à CD , A u cd x> '0 bO bo •H .3 a X CO CD Q) Ë f * M l s oil P # l ft! IO o I I »4I Lewis Range 4- 4-

Mission Range + +

Swan Range 4"

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range 4-

Gallatin Range 4-

Absaroka Range * + 4-

Beartooth Plateau * 4- 4-

Crazy Mts,

Big Snowy Range

Highland Mts,

- 116 -

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd ■p m % .Q 0> L, > 0) cd cd 'g rt p p Î * g iH cd m CO p 1—i cd n) OJ CD . o CO •r4 1—i cu •H # tuO u - p CO CD •H 3 m P ÜI u » g § § cd o 3 CO cd > Ü > Ü o e g u k cd k I i g k o . . •» CD M <*!! <1 «I *=«1 -*îi <*;i -*!| o Lewis Range +

Mission Range

Swan Range +

Flathead Range +

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range +

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range * +

Absaroka Range ♦

Beartooth Plateau ♦

Crazy Mts*

Big Snowy Range

Highland Mts.

-117-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. § cd "ü) •H § g •H •H 0> o •H 5 01 •H X3 % ctf * cd 0) "l-t (D cd % X> •H x; ja % 01 - p ■+J 1 I g Cm T3 cd g rO IO •H •H •H Ü 01 -P (D 01 CQ 1 I g cd o O rH cd cti g OJ T 3 f-t fH I f g r H Ü i o o CD ; o -P wl c/5i Ml Ml CO c o l COÎI CO;i

Lewis Range + + 4- +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range -

Anaconda Range 4-

Bridger Range *

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range ?

Gallatin Range

Absaroka Range

Beartooth Plateau ♦

Crazy Mts.

Big Snowy Range -

Highland Mts.

- 118 -

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. to CO c cd •H ttf <1> m cd 1— 1 •H I—I 1—1 G, •H e« Ü C cd •H m 0) cd G Ciû •â c** EQ OJ Ü G (U o 1 o > to 0> CO g cfl % Cl) 0» CO o 0 rH > o - p 'O - p g «J •r4 - p G 1: S3 1— I •H G 0> o •ë Ïh 0) ra «H ■H (D 1— 1 •H 1—I QJ 0) ♦ •H g bfl I O G, a •H > •r-j - p > a> 0 rH +:> • * c r « cd co COI -aSI I I <

Lewis Range + 4- ♦

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts*

Little Belt Mts*

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range ♦

Pioneer Range +

Gravelly Range

Tobacco Root Mts*

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts*

Big Snowy Range

Highland Mts*

-119-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. •H •H fSJ -P •H rH •H O CSI JC -P CO o rH I—I CO O “H o ■r-t X! T3 o CO Ü CJ CO c CO I C CO p % O 0) s I H P o ’V -P y *H S p o ■S 0} +5 Ü fH rH p •H 03 CD rH Cd 'H p B 1 >1 M P. rH o JQ O I nJ . • § I a Q oi « O i l «I *aS

Lewis Range + + ♦

Mission Range +

Swan Range +

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range +

Beartooth Plateau 4-

Crazy Mts.

Big Snowy Range

Highland Mts.

— 120—

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd Si cd o rp O $ :h Ü cd *H % p CQ % -2 § cd Q) 0 CL •H & ri ‘E3 m (D •H 0 cd U I r-fI I I c t3 § Ü ÜÛ -S o » # •a(! I -*;i <«!i o a| A I A l A l A l Lewis Range

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range - 4-

Anaconda Range +

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts, —

Madison Range +

Gallatin Range 4-

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

- 121 -

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd cd OJ A % A & c * ( h g •H cd cd cd o - p * o & o cd cd o n) to t f—1 A o o * g ■H Ft, o S) Cd TJ I0 t-i 0) cd c : •H cd X •H T3 <0 C Ü o o HP o *o •H o rH Im o cd iH T3 u x t o o cd ■à rH o a Ù0 Ito I ■H •iH £3 A cd § U (0 P rH > & to to A k I—I1 cd Cd I > 0) rH # a x : a 0 | «I 0 | A g Lewis Range

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts. ♦

Little Belt Mts, +

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts. ♦

Madison Range +

Gallatin Range 4>

Absaroka Range +

Beartooth Plateau

Crazy Mts,

Big Snowy Range

Highland Mts,

— 122—

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. •H fp «2 CO ë -HCO •H CO -p «J •H -g c oJ p 0) I—t 0 I o o. * "S CO o o CTÎ •H W)i (I) X! rt -p ■g 0) o -P s I 8 w § § a nJ (0 ë CO I ■H f, nJ CO ISl rH CO I rH I CO o a o -2 to - p 0) T5 I cx -P W col col col 3 s;i I CÛI 1 Lewis Range + —

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts,

—12 3—

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd § +3 G cd o -P * E cd CO O cd * (0 •H W) * CO 05 •F-j rH G Cd B td •H 1— 1 O cd -P CO >H CD 0 Cd Ï3 CO o «H (D G Î3 P *fH X3 •H 0 p •H * m •S cS r-j f CO cd x: cd ■H G P •H •H E 0 •H e F—i •H CD •H 0 Ü 0 Ü A G iH • CD 1— 1 t ) m n Cd c CO . G •H uo rH •H o o G n t cd çd j ü h k G 1 )h u s u CD CD rH CD ü o . rO > Ü > O T) 1 «P F—! 6 O o . . . . . to c0| C O | toi toi toi toi co| co| co| Lewis Range + + 4- + 4- + ♦ + Mission Range

Swan Range + 4-

Flathead Range 4- •4*

Bitterroot Range + ♦

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-12U-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. * •H to *Hta •fH CO g “H 0 •H cd •H O rH 0) •H Cm O «H •H rH CO ^4 cd •H O CD •H Q> -H H 3 Cm * « (h x: r H cd CO O ■H o cd 1 O r H p r H +3 •o •r-3 P 0) C cd ÜD Cd O •H CD (I) > 0 ) r H s •H O > O r H n I to 0) ■ H , , o 0) M l M l I Q l oi oi CO Lewis Range 4 - + +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts*

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range +

Beartooth Plateau 4-

Crazy Mts,

Big Snowy Range

Highland Mts,

-125-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CO •H CO •H Cd -P S "H cd to IM rH -P to Cd o (UO ■ p 0 ■H a (H o CD to cd J-. cd •H •H o cu rH to rH O iH CO CO CO I tfO ■H c P D. fH P i 1—1 U 1 •P o •H u X) ■P •H 5 O h cd -P > P Cd "r4 •H •H k u > CO Q) XJ TD 1 0) -P c5 > bo > Caû > O , CÎ3 PI o. PMl ful 0,1 0,1 0.1 Lewis Range

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts. *

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau +

Crazy Mts, +

Big Snowy Range

Highland Mts.

-126.

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. ç -,

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Mission Range

Swan Range + +

Flathead Range + 4 - +

Bitterroot Range ♦ +

Big Belt Mts. -

Little Belt Mts. + + ♦ +

Flint Creek Range — — -

Anaconda Range +

Bridger Range +

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts. +

Madison Range + ♦ 4-

Gallatin Range ? *

Absaroka Range ♦

Beartooth Plateau ♦

Crazy Mts. -

Big Snowy Range

Highland Mts. -

-127-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd * c6 0) 0) CO § CO cu Ü X3 Cj 0 ) Ml C Cl CO U i 1-4 is 0) § u > § CO § tC) I ^1 «-9I k4l g o| o| oi Lewis Range _

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range -

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range -

Highland Mts.

- 128-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. "H "H sz ccJ 4^ 0) •H o ccJ M to cd •H -P “r-t o F, cd Ü * ‘H 3 0) o (D 0) to g g CO • H XI •H £3 •rC o o . . - p a 0 C 13 <3 «mIll o } •H CO CO T3 CD 0) O Ü g •H •H § F, t— 1 M CO c: O. . O •H O (D O CO rH A 0 F, g O f g •H A H 4-> Cd !■ s CO m > t T3 Æ k i >> •H c 1 to t> K £3 O 0| 6^1 E-.I ■H 1 Lewis Range + + +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Greek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range *

Gallatin Range +

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-129-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd * u •H i o •H c rH ■H -P cp I * g •H CL cd •H CO fH 1— t > +3 o a iH cd cd *Ü % 0) o rH Cd T3 X> O rH o I c C c m (H o ■H a cd *H -H Ü I «aî CL S) -P •H (d O pH r ^ rH rH o 1— 1 i > Cd O •H CL & M Ml M l M l M l M l 3 Lewis Range +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range + ♦

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts,

-130-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. H CO CO 0 - p cd cO 0 C § 0 P •H •H cd p CO CO 0 C cd I 0 cd tUO •H 0 0 CO CD rH § •H •rf CD 'rf o P •H fH •HP P Ù0 Fi Ü CO CO hû cd •H Cd CD Cd 0 0 ‘H F, c F, > E Ff Fi Ü CD F. P W CD Ü n CD C §• CD 0 0) c o l •H T3 P O -p CU ■P Cd rH •H 0 | CO CU •H CU •H CO o rH Ü 4-^ o| 0 F, CO C o ■H O CO 0 o | Ü CO P CO CD -p CO > . CO o CO ICP cd o o| J I I ^"41 1 S o Lewis Range +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts. -

Little Belt Mts. -

Flint Greek Range +

Anaconda Range +

Bridger Range +

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts. -

Madison Range +

Gallatin Range ♦ ♦

Absaroka Range ♦

Beartooth Plateau ♦

Crazy Mts.

Big Snowy Range -

Highland Mts. »

.131'

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. to ‘H E Î-I (0 o * qj oj cO JH CÜ CtJ o rH d a O *H E O 0) E rH T3 'g -P ■H «H fn tu «H * O I 0) r—I to cd •H I cd hû P •H 0) rH g C rH •H to g ClJ Ü P 0) •H rH Q) •H o A -g a TJ ■+J E(H •H g O to ÎH •H-P a "H 0) a s •S •H 5 ■H •H rH 1—j .2 o tot> N rH bJO Ml o Ü k to o Eto qj 1 nJ x : , , O O QL. I A i l (Cl A.I t=»l >1 Lewis Range + + + H-

Mission Range +

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range + +

Gallatin Range + +

Absaroka Range

Beartooth Plateau

Orazy Mts,

Big Snowy Range

Highland Mts,

- 1 3 2 -

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. * nJ C'l § * ■H G 0> flj Cu 0 *H cu % Ü 0) s: •H tM 0 OJ -P -p m c\5 Ê-I 0 I s 03 a 0 ce g *r4 O 0 Ü !— I g g Cl 03 Q) E o 03 ■P T3 T3 I • H 0 i G C O « m «*! <*!l Q OI a 6 & c b Oi OI

Lewis Range +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts. 4-

Madison Range

Gallatin Range

Absaroka Range *

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-133-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. Oj OS CO U % -P •H U OS rH 0) CQ w CO c X i r H •iH •H CO *H -P a SiI fw •H01 fH •H 0) * 0 •H fH CIS -H X! CIS I X) •H-P g m CIS Ü CIS *-P X» CIS CO S a -P t CIS (IS TS m o o CIS a CIS 0 CO u & t s •HX) CO rH c a ci $:i -p 1 CIS A 1—1 P ♦H O CO CIS •H CO CIS Cm S -p O «H g CD k § OS u •-a 1Ü X) 1 X •H (X f—1 > •H o T3 > g • 1—1 r H tH I I—I g o H X o o OI 0 | O I I o O a. p.! A,| a* Lewis Range m m ♦

Mission Range

Swan Range

Flathead Range

Bitterroot Range ♦ +

Big Belt Mts.

Little Belt Mts.

Flint Greek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range + 4-

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-13U*

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. cd CD (O Ü •H •H CO cd G cd 1— 1 CD § to cd CO g o cd A t) ü -p CD Cd * -p •H CO cd cd CO t—4 -H -p c O rH u CO rt (0 •H c - P t>j > G cd o. Ck X! cd x : g cd CD £X •H cd cd fH •p (— 1 O •H CD Q) cd cd U rH O OI ■H P CD 1—1 -r4 so Ü rH CO P Cd G CQ g § ■H CD -p Ü > X i x ; iH CO S Cd •H I JZ • a I a, C L ,| I A.I 0 . 1 A . I a i I a Ml Lewis Range + +

Mission Range

Swan Range

Flathead Range

Bitterroot Range *

Big Belt Mts* -

Little Belt Mts* +

Flint Creek Range +

Anaconda Range +

Bridger Range +

Beaverhead Range

Pioneer Range

Gravelly Range +

Tobacco Root Mts*

Madison Range +

Gallatin Range +

Absaroka Range + +

Beartooth Plateau +

Crazy Mts « ♦

Big Snowy Range -

Highland Mts.

-135-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. •rf rH rt li T3 bû C % 1 ■H Ü to CtJ X> a, cd Cd -p rH to to "E! ci JCO •rH •H o *r4 o g CtJ 10 Q ci rH C ü nJ •H a* 0) nJ •H to ÜÛ CtJ Ü S -p Ü 0) 05 •H ■P Sj o rH CtJ -P O 0) •H Ü k -0 to to ■P u s . 0) to to O CtJ e œ K| S SI SI SI mi O o| Lewis Range

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Greek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

- 136 -

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. CtJ CO o CD ttJ p CtJ cd •H Ü CQ (0 CO 0) •iH •H CtJ O •H > 0) ■iH bO u CD u nJ to G p P o C (t* •H 5 •H g -p ‘fH O rt (tj -p <-t r—j •H r—I (0 CtJ CO a to cd iH o CtJ CD •tH ‘H •iH k o o g AI Ü 0) X) Æ •H ■H C CtJ s s rH *iH Ü X X! CO •H rH f-H Ü m ft rH G Ü 5 x: O ‘fH Ü u § O a. k C +> i •H > o to •H 6 T3 I nJ . » • jH CD o o| o| ol o a t Ok our eCl Lewis Range + -*• +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts,

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau +

Crazy Mts,

Big Snowy Range -

Highland Mts.

-137-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission. c- TO cd 0 cd •H TO k -P 1—1 0 0 G cd O ■P o o Sh o Cd o 0 -p cd cd 0 0 m % 0 G G o g • a 0 0 o 0 o o •ri •H p> X) TO TO G U a, cd -p 0 0 0 ta 0 0 cd 0 Ü TO •H -p -P u TO *H O 1 ra F Ü A 1 I—4 G. 0 c A P> Ü «H •H CÏ O JG o A "H P* o E •H -P TO i X) rH 0 u i CD rH U > Ü O I t TO > 0 XJ I «H c 0 0 . CM AI

Mission Range +

Swan Range 4-

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range ♦

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau +

Crazy Mts.

Big Snowy Range

Highland Mts.

-138-

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Mission Range

Swan Range

Flathead Range

Bitterroot Range + +

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Range

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

-139-

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Lewis Range + + +

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts. -lUo-

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Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts#

Little Belt Mts.

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts.

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Mission Range

Swan Range +

Flathead Range +

Bitterroot Range

Big Belt Mts.

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range +

Gallatin Range ♦

Absaroka Range ♦

Beartooth Plateau * +

Crazy Mts.

Big Snowy Range

Highland Mts.

-1U2-

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Lewis Range + -#■

Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts#

Little Belt Mts*

Flint Greek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts#

Madison Range

Gallatin Range

Absaroka Range +

Beartooth Plateau — +

Crazy Mts,

Big Snowy Range

Highland Mts.

-1U3-

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Mission Range 4

Swan Range

Flathead Range ♦ +

Bitterroot Range + 4-

Big Belt Mts. 4

Little Belt Mts. ♦ 4

Flint Creek Range 4

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts.

Madison Range 4

Gallatin Range 4

Absaroka Range ♦ 4

Beartooth Plateau ? 4 4 4

Crazy Mts.

Big Snowy Range

Highland Mts.

-1U14-

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Mission Range

Swan Range

Flathead Range

Bitterroot Range

Big Belt Mts.

Little Belt Mts.

Flint Greek Range -

Anaconda Range +

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range *■

Tobacco Root Mts. ♦

Madison Range ♦

Gallatin Range ♦

Absaroka Range

Beartooth Plateau

Crazy Mts.

Big Snowy Range

Highland Mts. -1L2-

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Mission Range

Swan Range 4-

Flathead Rai;ge -*

Bitterroot Range ♦

Big Belt Mts,

Little Belt Mts,

Flint Creek Range

Anaconda Range

Bridger Range

Beaverhead Range

Pioneer Range

Gravelly Range

Tobacco Root Mts,

Madison Range

Gallatin Range

Absaroka Range

Beartooth Plateau

Crazy Mts,

Big Snowy Range

Highland Mts,

-lL6-

Reproduced with permission of the copyright owner. Further reproduction prohibited without permission.