FEB 28 1947
- 49.5 245"
SARGENTIA

A CONTINUATION OF THE
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY

VI

THE BOTANY OF SOUTHWESTERN
MACKENZIE

BY
HUGH M. RAUP

WITH THIRTY-SEVEN PLATES AND SIXTEEN
TEXT-FIGURES

PUBLISHED BY
THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY
JAMAICA PLAIN, MASS., U. S. A.
1947

H pl.i-37

f

SARGENTIA

A CONTINUATION OF THE
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY

A publication issued at irregular intervals by the Arnold Arbore-
tum of Harvard University. Issues can be obtained from the Arnold
Arboretum, Jamaica Plain, Mass. U. S. A. All correspondence
pertaining to Sargentia should be addressed to the Librarian.

No. I. Fijian Piant Sruptes, IJ]. Boranicat REsutts oF THE
1940-41 Cruise or THE “CHENG Ho.” By A. C. Smith (and collabo-
rators). Pp. 1-148, with five text-figures. July 20, 1942. $2.50.

No. II. THe Arattacear or Cuina. By Hui-Lin Li. Pp, 1-
134, with fourteen text-figures. Oct. 26, 1942. $2.25.

No. III. A Revision or tHE Genus Sapra CoLeBRooKe. By
Luetta Chen. With nine text-figures. THe Cuinesr anv INnpo-
CHINESE SPECIES OF Ormosta. By E. D. Merrill and Luetta Chen.
Pp. 1-120. Jan. 30, 1943. $2.00.

No. IV. Marerrats ror A FLora oF THE CONTINENTAL NortH-
wesT TERRITORIES OF Canapa. By A. E. Porsild. Tue Wittows
oF THE Hupson Bay Recion AND THE LABRADOR PENINSULA. By
Hugh M. Raup. With four plates. Pp. 1-135. Sept. 25, 1943.

$2.50.

No. V. FRaGMENTA PapuaNna [OBSERVATIONS OF A NATURALIST
IN NETHERLANDS NEw Guinea]. By H. J. Lam. Pp. 1-196, with
two maps and thirty-two text-figures. Translated from the Dutch by
Lily M. Perry. Feb. 6, 1945. $3.00.

No. VI. Tue Botany or SourHwesTerRn Mackenziz. By Hugh
M. Raup. Pp. 1-275, with thirty-seven plates and sixteen text-figures.
Feb. 28, 1947. $5.00.

SARGENTIA

A CONTINUATION OF THE
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY

VI

THE BOTANY OF SOUTHWESTERN
MACKENZIE

BY
HUGH M. RAUP

WITH THIRTY-SEVEN PLATES AND SIXTEEN
TEXT-FIGURES

PUBLISHED BY
THE ARNOLD ARBORETUM OF HARVARD UNIVERSITY
JAMAICA PLAIN, MASS., U. S. A.
1947

SARGENTIA

A CONTINUATION OF THE
CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
oF Harvarp UNIVERSITY

No. VI, pp. 1-275, with thirty-seven plates and sixteen text-figures
; Issued February 28, 1947

PRINTED IN UNITED STATES OF AMERICA BY
LANCASTER PRESS, INC., LANCASTER, PENNA.

THE BOTANY OF SOUTHWESTERN MACKENZIE
Hucu M. Raup

with thirty-seven plates and sixteen text-figures

TABLE OF CONTENTS

Page

DETERMINE IONE, AGKHOWIGUOINENES |... 6... sess + os oe cb edals cre ReREE ie Gate Rnan semana stale ietalat, = 2
Topography of southwemern Mackenzie ....... 0.4.5 save cubes os Meegubeetweaad dees: 6
PEigtOtY OE SEDIOCIBIOE caret kc es ie cece nce sone 0 Us Am Obs meee ts de> e5 10
Geology - and. O00)S Fuccim ete, fing 856i noe cece okay pol ee enC sale hae eis & yh 3% 17
Pre-Glactal Mela sides bsicccs once ccae cece whet OF om ene oS en Ries #5 <a 17
Region east of the Mackenzie Mountains ......,. <4 «0eeee es nN an Naa keeg's Waens 17
Mackenzie MGtntaitise eaciescccc 0.0 ce. ve ccc ce es oles 9 OpMmnmRERetSs tmp MenannEpT 213 Mois 70!/e 18
Glacial weoboity (2.4% wchiratela dds 0-0 sihaiesrals's 0 avid 6 45 05.8 eee OE EE Ge 482 ok 19
Glaciation’ east: of the Mackenzie : Moumtaintsi 2 fonts: is. Sates neo ihe sriclele divigi'e sis cues 20
Recent sedimentation sim southwestern’ Mackenziguwaa.s myc 6. oan few se se cle sh 000 + s,s 22
Glaciation: in the. Mackenzie: Mountains vacant + Sere os «ie tails ose os la ov oe 24
Geological observations af Brintnell Lake +l. Giga cue as <> conan sss ness cn cescace sees 25
Summary of topographic and geologic observations ...........00eeeee cece cree eens 28
Climate and. agriculture. % 1644 ><> <>< ++ << 6G sea EDD pia win’ dee eM elesia bed ese scee ce 29
Climate at Simppodt: 4 cccgodins 6540+ ++ as + 6cr ae acclaim RE <a ies 0 § 30
Seasonal phenomena and agriculture at Simpson ..............secceccccceesceesees 30
Comparison of the climates of Simpson and Brintnell Lake ...............-+.-+-0- 32
Plant: communitiesof southwestern Mackenzie) samen, sic. sete swe es ere 34
Introductory Qin oaadecbebrsleele l= sls e'o/e «(2 sia Deen tare slr felel ote coli EMRAO CeIn As ne les «3 4% 34
Plant communities in the“vicinity of Fort: Simpsonins......0-.---tencesctectcrscces 34
Flood plain spruce 20rGate .. 1.0.5: 55 peepee UNE An « C05 ose MER EME wes +s 0s 4 49 a5
Upland forests of white spruce and jack pine ........... ccc eee e tener eee eeers 35
Prairies’ il See cc eee dike! c a 5 so koe ra ERED se steele oa. 2) aM enIN eel. :0)¢ 090: 4 36

- Plants of sand’ bars and river banks -~. 2.2.0 GMEL Gs o's. ie gdp GR eee ee oe idee ee 37
Plants: of fields tandscléanines:.......:<:-./ aceite «= scolpaicle 5410 SIRiP ei e + Sia tie io! ot oherg 38
Vegetation of the Horn and Franklin Mountains .......... 0... ee ee eee e eee eee eeeeee 39
Plant comm«anities: of thes Btintnell . Lalketaneapetrs. ees 6. nis a >< seers arene ies 5 o's oehein 40
POrestse tien ook lee eee oat a sea ee cs ip tus ace) s aKeienNmeres 3rtte (ois (alles 4 41
Subalpine scrub communities... . ..)oin Beene tenes ve tawen dvdematek ee cy es sees ee 47
Alpine vewetation « .: vrescs aos +0.s0.04isth Seen ee sae ARMIES OMENS Hs ease essere 49
Damp alpine meadows ...%... «+++ «p55 eee EMBO GIRE RL eee sis eee s tens 51

Dry alpine meadows «1... .....+- +0000 Oks Sam NEE ONES ese 40s cee ees Se
Plants of loose slide rock and turf-banked terraces ..............-.sceeeeeeees 54

The vegetation of stone rings: ..i'ss 6d didip aonb ace ep V¥ew lea tWecawssegnenes 52
Ledge and crevice vegetation ......ccccesseeeececcrccecsssswsreseessenecssers 55
Plants of ravines and gorges below timber line ...............cceecceceerceceees 57
Flood plain communities .......... +. :speenaess EMMIS SWE sies 6% 6p == 5505's Biff
Plant communities along the Canol Roadiiiemmmiien eee. os cr ale es ens nee sain 59
Geographic affinities of the vegetation of southwestern Mackenzie .........-...-+-++45 60
Introductory chic cece ben beecees o00cee ek EE 5s se sceestonsberesenes 60
Geographic relationships of the forests of southwestern Mackenzie ............-..+: 61
Geographic relationships of species in the Brintnell Lake region .............-..0045 62
Wide-ranging forest: species ......... ca peEEMEMMMERE ERIN: 2c ccc nccsesccceesepe 64
Wide-ranging arctic-alpine plants . 0... 6c. ce eece ee cc cece nsec teres ce eesscecnes 66
Plants of the timber line region .. ....: ig ppee PEA eb les siete sens raeewens 68
‘Alaskan-cordilleran species .......+.2s cee ERE bn cess ce tacesereesceeees 69
Cordilleran species .........00:00so08s eee RB ewes ccevearcrersessueces 70

EERIOTS cc a's dca ccccsctsceess oan Gann Eo. , 5 ae od teu ee amcheeae 71

2 SARGENTIA [6

ee NI IIE AI 6 6K Kk vos SEs sb wget a MEL, vcs akon 72
Application of Hultén’s hypotheses to the Brintnell Lae HOCK 55 Ahi ss oisOes cc. 74
Ie NE PRM y 5 5500s Ud CLAN c's s ly Mo heade onde 76
Peoria CIO CAM MME 6s oo o's 5c 1SYs FC Dis odes ce MR LL. ors 76
eNO PRPIETICRM, CORN FRINGED 86.5 60. ions 055s vans be kb wen seis coe ek 79
Continental western American radiants ..............ccecseceevcceccecccce le, 80
NN SN 0 Sis oh Suds oj ss bn ss c3s chor vv 82
UEORL RISCUPMDOMRT DIME 665 56.0 e ccc ccccssevcccccacsctenpecad PPP WAL Mies ave 84
PN ie hit SMG WS lS 6 di o's was 4:0 5'5.d CREE SO oT wkend ce 86
Origins of the flora of southwestern Mackenzie ..............000-0--0000 000 90
Catalogue of the vascular flora of southwestern PARCKONSIE: «vais ss adh ies chen ke 95
TSI Pc eh eh CAV RL S00 is wads ss bn sc knne usec sade cdbeen ots mutate cas 257

INTRODUCTION AND ACKNOWLEDGMENTS

THE BROAD outlines of the botany of boreal America began to take form in
the first half of the nineteenth century, particularly in the works of Sir John
Richardson and William J. Hooker. Not since that time, with the exception
of John Macoun’s “Catalogue of Canadian Plants,” have any attempts been made
to prepare general descriptions either of the flora or of the botanical landscapes.
The necessity for more intensive studies and the difficulties of travel have re-
quired rather that the work should be carried on in parts, looking forward to
a more comprehensive revision of earlier concepts when the whole region should
have been examined in some detail. The consummation of such a revision has
been rendered increasingly complex in recent years by the development of new
techniques for the study of the plant cover, and new viewpoints in the fields of
biogeography and taxonomy. These new techniques and viewpoints have re-
quired that the whole structure be re-examined constantly in order to bring
modern interpretations of taxonomic and geographic phenomena into line with
progress in related fields.

It was to be expected that more intensive study should be subdivided on a
geographic basis, examples of which, as they apply to the Mackenzie drainage
basin, will be noted below. The present paper deals with one of these geo-
graphic areas, in the southwestern part of the District of Mackenzie (Northwest
Territories), and may be divided roughly into four parts. The first is a general
description of the region—its topography, geology, soils, and climates so far as
they are known, together with a review of exploratory history. Second is an
account of the plant communities that together form the botanical aspects of the
landscapes. Third is a study of the geographic affinities of some species and
communities, with an attempt to apply to the problem the recent hypotheses of
Fernald and Hultén concerning the development of boreal vegetation in post-
Pleistocene time. Finally there is an annotated catalogue of the vascular plants
known to occur in southwestern Mackenzie.

The area treated in this paper is bounded on the west and south by the borders
between the District of Mackenzie, Yukon, British Columbia, and Alberta.
On the east it extends to the Slave River, Great Slave Lake, and Marian River.
Its northern boundary is approximately at lat. 64° 30’, making the northern
limit on the Mackenzie River a short distance above Norman (see Fig. 1).
These boundaries originally were selected not because they had great botanical
significance but because they enclose an area not covered by any previous or
current investigations. Nevertheless, as will be shown subsequently, the western

Scale of miles
J 2s 50 7S

100

> %
q

winter L-
t isabella eS
&%
Seguir L Snare L lr rzzle be
fee a) or kney &-
Takatu L : v2 2 ep
BS gael Poy

Fru

a

ALS crn
h3

Navire,
(South ahan ni

e Kakisa “ ae

4
x ae,
i a ratinasy’

Aiard > “a> Nosy
aac <r ——a

ile : =

Pelitot PR ' Bistcho 2

Fic. 1. Map of southwestern Mackenzie and adjacent regions.

o

Louise Falls
Ae anne
Q Fa \5 auf talo
Leke 2

-_—_-
—_

woo?

ene |

12

—

[Zr61

TIZNANIVW NYALSAMHLAOS AO ANVLOG AHL “dAVY

4 SARGENTIA [6

and southwestern boundaries prove to be of considerable geographic importance
from a botanical standpoint. .

The Great Bear Lake and lower Mackenzie regions will be treated by A. E.
Porsild in his forthcoming Flora of the Northwest Territories, preliminary notes
for which have already been published (Porsild, 1937, 1943). Botanical inves-
tigations in areas to the south and southeast have been reported upon in some of
my own papers (1930, ’33, ’34, 35, °36, 42, ’45C), while the region of the Alaska
Highway in northern British Columbia will be the subject of papers growing
out of the field work of 1943 and 1944 (Raup, 1945A, B). The flora of Yukon
is being studied and published upon at the present time by Dr. Eric Hultén
(1941-45). The flora of southeastern Yukon, however, immediately adjacent
to the area discussed in the present paper was almost completely unknown until
1944, when A. E, Porsild made extensive collections along the Canol Road be-
tween Johnson Crossing and Norman Wells. Mr. Porsild’s main report, now in
preparation, will make a highly significant contribution to our knowledge of
the flora and plant geography of northwestern America. However, he has
already published (1945) an account of his collections from the eastern slopes
of the Mackenzie Mountains.

The material basis for the present studies is principally in three collections.
The first was made by Dr. James H. Soper and myself at Fort Simpson and at
Brintnell Lake in the Mackenzie Mountains during the summer of 1939 (see
Raup, 1939). The second is a collection made by A. E. Porsild on the eastern
slopes of the Mackenzie Mountains along the Canol Road in the autumn of
1944, and the third was made by Dr. V. C. Wynne-Edwards of McGill Uni-
versity in the same area during the summer of 1944 (see A. E. Porsild, 1945).
I have also had access to a few small collections, notably those of E. A. Preble
and Merritt Cary along the Mackenzie and on the slopes of the Nahanni Moun-
tains, of C. L. Crickmay and F. S. Nowosad along the Mackenzie and lower
Liard rivers, of Onion, Kennicott, and Hardisty at Great Slave L. and along the
Mackenzie, and of Brooke, Hume, Howe, and Bedford at Great Slave Lake.
Also I have examined numerous duplicates from the Richardson collections made
at the time of the first and second Franklin Expeditions and now to be found in
the herbaria at the New York Botanical Garden, Harvard University, and the
National Museum of Canada.

The list of the vascular plants cannot be regarded as in any sense complete.
A glance at the map will show huge areas that are yet unexplored botanically.
This applies not only to most of the Mackenzie Mountains, but also to the out-
lying Cretaceous plateaus that form the Eagle and Horn Mountains, from which
no plants have ever been collected. As in previously published lists of plants
from the northwestern wilderness, my chief aim has been to bring up to date
what is known, so as to form a sounder basis for further work and a clearer
idea of the lacunae in our knowledge.

The range maps published in this paper ? are based principally upon specimens
that I have seen in the herbaria at Harvard, the New York Botanical Garden,
the U. S. National Museum, the National Museum of Canada, and the Philadel-
phia Academy of Sciences. The ranges have in many cases, however, been filled
in from published records. The extent to which such records should be used
is always a problem; and it is common practice either to allow the maps to

1 The base map used for these ranges is by J. Paul Goode and is copyrighted and pub-
lished by the University of Chicago Press.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE be

stand upon specimens alone, or to include unverified data designated by suitable
symbols. I have preferred in the present instance to show the stations as of
uniform validity, trusting to care in the selection of published records for the
elimination of error. The rules used in this selection are few and simple. In
cases of some rare species, and of species whose determination is difficult and
controversial, no unverified records were admitted. Records far outside what
appear to be the “range patterns” of species were scrutinized carefully even if
specimens were available, and if the records were unsupported the stations were
put in with question marks. Recent monographic treatments usually have been
accepted at face value, though with reservations in cases of unusually contro-
versial groups. The floristic works most heavily relied upon are those done in
recent years by students who have had access to large collections and libraries,
though data on well-defined, long-established species have been accepted freely
from less authentic sources. The following are the papers most frequently con-
sulted in the preparation of the maps.

Aleutian Islands, Alaska, and Yukon: Hultén, 1937A; B; Hultén, 1941-45; Porsild, 1939;
Scamman, 1940; Anderson, J. P., 1943-45; Britton and Rydberg, 1901.

British Columbia, Western Alberta, Washington, and Montana: Henry, 1915; Rydberg,
1922; Macoun, John, 1877; Raup, 1934, 1942; Dawson, 1881, 1888A; Jones, 1936, 1938;
Piper, 1906; St. John, 1937; Standley, 1921; Farr, 1907.

Mackenzie, Keewatin, Alberta, Saskatchewan, and Manitoba: Porsild, A. E., 1937, 1943;
Macoun & Holm, 1921; Raup, 1934, 1935, 1936, 1942; Grgntved, 1936; Giissow, 1936;
Stormer, 1933; Rydberg, 1932; Hooker, 1829-1840; Preble, 1908; Macoun, J. M., 1896,
1911; Tyrrell, J. W., 1902; Tyrrell, J. B., 1896, 1897.

Labrador and the eastern Arctic: Polunin, 1940; Potter, 1934; Abbe, 1936; Woodworth,
1927; Wetmore, 1923; Bishop, 1930; Marie-Victorin, 1935; Raup, 1943; Simmons, 1906,
1913.

Greenland: Porsild, M. P., 1932; Bocher, 1933A, B; Gelting, 1934.

General: Macoun, J., 1883-1890; Fernald, 1925.

The field work of 1939 upon which much of this report is based, and without
which it could not have been prepared, was supported by generous grants from
several institutions and persons, to all of whom I am deeply grateful. The cost
of the expedition was met by grants from the Milton Fund at Harvard, the
General Science Fund of the American Academy of Arts and Scienaes, and the
Alexander Dallas Bache Fund of the National Academy of Science. Valuable
field equipment was kindly loaned by the National Museum of Canada. Per-
sonal contributions were made by an anonymous donor and by Mrs. Mary G.
Henry, whose interest in northwestern botany has been unflagging, and whose
pioneering field work in northern British Columbia is well known.

In the study of the material I have had much assistance and inspiration from
Mr. A. E. Porsild, Chief Botanist at the National Museum of Canada. He has
not only placed at my disposal a large amount of data on the flora of the eastern
slopes of the Mackenzie Mountains, but has also examined much of our material
in the light of his extensive knowledge of the whole flora of western arctic Canada
and Alaska. Furthermore his accumulated data on the ranges of arctic plants
have been made freely accessible to me for the filling in of gaps on the maps.

Several students of special groups have been kind enough to examine speci-
mens representing their particular fields. Dr. C. L. Hitchcock has looked over
our specimens of Draba; Dr. Bassett Maguire, Arnica; Dr. I. M. Johnston,
Boraginaceae; Mr. C. A. Weatherby, the ferns and fern-allies; Dr. Bernard
Boivin, Thalictrum; and Dr. Edgar T. Wherry, Polemonium. Professor M. L.

6 SARGENTIA [6

Fernald has, as usual, been a constant source of ideas and assistance in the study
of the flora. Dr. E. C. Abbe of the University of Minnesota has very kindly
permitted me to make use of his unpublished data from the east coast of Hudson
Bay for filling in the maps. I am indebted also to the authorities of the New
York Botanical Garden and the United States National Museum for access to
their material. Dr. William R. Maxon of the latter institution was especially
helpful in getting out a list of the Preble and Cary collections. Professor Kirk
Bryan of Harvard has very kindly read and criticized the geological parts of
the paper.

It is impossible to mention individually all the people who helped to forward
the field work. Mr. Harry Snyder of Montreal placed at our disposal indis-
pensable information about the Brintnell Lake country, with copies of the manu-
script map of the lake and its environs made during his expedition of 1937.
Officials of the Hudson’s Bay Company and the Mackenzie Air Service were,
as usual, most helpful in organizing our supplies and transportation. It is im-
possible to express too great appreciation of the kindness and hospitality of our
many friends at Fort Simpson. They made our stay there thoroughly comfort-
able and a continuous delight. Special thanks are due to Archdeacon and Mrs.
Harry G. Cook who took us into their home, and to Capt. George Isbister who
was in charge of the Government Radio Station. Infinite patience on the part
of Capt. Isbister made it possible for us to maintain radio contact with Simpson
throughout the summer.

Finally it is a pleasure to express my obligation to my wife and to Dr. James
H. Soper, field companions of the expedition of 1939. Without their unfailing
patience, enthusiasm, and good will, the measure of success achieved would have
been impossible.

THE TOPOGRAPHY OF SOUTHWESTERN MACKENZIE

The most prominent topographic feature of southwestern Mackenzie is formed
by the Mackenzie Mountains, which lie along its western border. These moun-
tains appear to be a part of the Rocky Mountain system, but their central axis
is displace eastward from that of the latter. The Rocky Mountains of northern
British Columbia recede to low elevations in the plateau country of the upper
Liard River. This plateau “extends eastward, right through to the Great Plains
region, between the extreme north of the Rocky mountains and the southern end
of the Mackenzie mountains. These two groups of mountains do not lie in
tandem; they lie in the same northwesterly strike, but Mackenzie mountains are
offset by a gap that centres about the mouth of Fort Nelson river and opens
out to the southeast” (Camsell, 1936).

The Mackenzie Mountain system begins at the Liard River and extends in
a northwesterly direction in a great arc some 600 miles long. According to
Camsell (1. c.) it covers an area of 75,000 to 100,000 square miles and is re-
garded as the largest mountain group in Canada. Keele’s description (1910),
made as he crossed the range from west to east by way of the Ross and Gravel
Rivers, will serve to characterize the topography.

The mountains as seen from the west present a massive and continuous front,
unbroken by lowland areas. The crestline is uneven in profile, however, and
the course of the ranges varies, with subdued types of mountains among the more
rugged ones. The structure is characterized by folding on a broad scale, with

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 7

the folds sometimes close-set or overthrust. The western slopes are trenched
by wide valleys that cut well back to the divide. Both hard and soft strata
appear, and to some extent have determined the major topographic features,
though the main drainage channels cut through both hard and soft rocks. The
higher, more rugged mountains are granite stocks which have resisted weather-
ing much better than the intervening sedimentary rocks. Glacial erosion and
deposit have greatly modified the surface features, smoothing the bedrock and
strewing the valley floors and lower slopes with drift and outwash. In this
western part of the range the mountains are commonly over 7000 feet high, and
a few reach 9000 feet.

“About forty miles east of the divide the topography changes in a marked
degree, and a more compact and rugged mountain region is entered. The
drainage channels of this region are confined in narrow valleys, with steep,
barren slopes of rock and talus, the rivers in the bottom flowing in a very con-
tracted bed, which at rare intervals opens out into a narrow alluvial flat.

“The structure of these mountains differs from that of the ranges to the west,
being apparently due to fracturing, buckling and faulting of the strata, and the
residual masses present the appearance of a series of faulted and tilted blocks.
The principal lines of fracture are in a northwest-southeast direction, and the
beds have a prevailing southwesterly dip.

“Escarpments produced by tilted strata, overlooking fault valleys, are the
most prominent features, but they do not appear to persist in alignment for any
great distance.

“The highest peaks are roughly pyramid-shaped masses, carved from the
harder of the stratified rocks of which the mountains are built. They vary in
elevation from 6,500 to 7,500 feet, with a height above Gravel river of 3,500 to
6,000 feet. The denudation of these mountains has not reached such an ad-
vanced stage as that exhibited by those to the westward; the valleys are narrow
and steep-sided, and the grade of the drainage channels is much steeper.

“The Mackenzie mountains, as a whole, have a maximum width of about 300
miles; there is no well-defined crest line, but they appear to be rather a complex
of irregular mountain masses, which are the result of deformation and uplift.
The topography of the western portion bears evidence of long-continued differ-
ential erosion, while the eastern portion has the appearance of being a more
youthful topographic stage. Both in geology and structure the eastern portion
of these mountains is closely related to the Rocky mountains in southern Canada.”

“A portion of the drainage of the western slope of the Mackenzie mountains
falls into the Frances river, and thence by the Liard and Mackenzie rivers into
Beaufort sea, but the greater part is taken by tributaries of the Yukon river to
Bering sea. All the drainage of the eastern slope falls into the Mackenzie river.

“There is a great disparity both in river development and stream grades
between the two sides of the divide.

“On the western slope the water flows from near the divide, for a long dis-
tance, through valleys of mature erosion with an easy grade before reaching the
master stream, the Yukon; while the streams on the eastern slope fall rapidly
for a comparatively short distance and reach a much lower level at their junction
with the Mackenzie. For example, the Ross and Pelly rivers have a combined
length of 450 miles, and enter the Yukon at an elevation of about 1,500 feet
above sea-level, while the Gravel river, 255 miles long, enters the Mackenzie
at a height of about 200 feet above sea-level.

8 SARGENTIA [6

“Owing to the great difference in precipitation the streams from the west
side of the divide carry down to the Yukon more than twice as much water as
the streams over an equal area on the eastern side. Thus the Gravel river and
its branches do the work on the eastern side of the divide, while the combined
efforts of the Hess, Macmillan, and Ross rivers are required to carry the water
from the western slope; the Gravel river being about equal in volume to the
Macmillan” (Keele, 1910).

Keele’s delineation of two types of topography in the Mackenzie Mountains
is borne out by more recent observations on the route of the Canol Road and
farther south in the South Nahanni River country. In fact the South Nahanni
appears to separate the two types, at least in its upper course. Flying west from
Simpson, one comes rather suddenly to the front range, and then passes over
a mountain mass whose summits, seen on the skyline, make a fairly even plain.
The tops commonly show a rolling surface with rounded contours, broken by
sharp cliffs and steep slopes into the deep valleys. This is the topography
described by Keele along the Gravel River, though west of Simpson it is not so
high. Porsild (1945) saw it at the “Plains of Abraham” about 80 miles west
of Norman Wells. The high granite peaks of the western elements of the range
appear west of the upper South Nahanni (see below), and apparently corre-
spond to those seen by Keele in the Christie Pass area.

The maximum height attained by the Mackenzie Mountains is as yet unknown.
A granite peak at Brintnell Lake has been estimated with reasonable accuracy
at about 8500 feet, while still higher elevations have been reported between there
and the Gravel River, as well as northward in the Bonnet Plume River region.

Some further discussion of the mountain topography will be found in the
description of vegetation in the Brintnell Lake region.

East of the Mackenzie Mountains lies a northern portion of the central con-
tinental plain. It is composed of two principal elements in our region: the
Alberta Plateau and the Mackenzie Lowland. These are bounded on the east
by the Laurentian Plateau (see Camsell & Malcolm, 1921). The Alberta
Plateau is represented only in the southern part. Its northern boundary extends
“from near Fort Smith on Slave river northwesterly along the south side of
Great Slave lake and thence at an equal distance along the Mackenzie river to
the Liard in longitude 122 degrees. Here the escarpment is lost, although from
McConnell’s observations it may swing southwest along the valley of Liard
river.” The northern margin of the plateau is a well-defined fall-line for
streams draining in that direction. Its surface is gently rolling, and marked by
glacial deposits of various kinds with an abundance of lakes. The drainage is
immature, with sluggish streams and many muskegs. “The top of the escarp-
ment is about 400 feet above the level of the adjacent lowland, and the surface
then rises gradually southwest and west to the foothills of the Cordillera”
(Camsell & Malcolm, 1921). Southwest of Great Slave Lake the plateau sur-
face is about 1100 feet above sea-level.

A plateau of similar nature borders the Mackenzie Mountains at the northeast.
It has been called the Peel Plateau by Camsell (1906; see also Camsell & Mal-
colm, 1921), and occupies the upper valley of the Peel River between the
Richardson and Mackenzie Mountains.

The Mackenzie Lowland, through which the Mackenzie River flows northward
to the Arctic, is bounded on the south and west by the Alberta Plateau escarp-
ment and the Mackenzie Mountains, while to the east it abuts against the

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 9

Laurentian Plateau. In general it is a broad plain of low relief, with a gradual
slope (about 8 inches to the mile, according to Camsell and Malcolm) northward
to the Arctic coast. “The surface as a rule does not stand much above the
main water-courses or the great lakes and the banks of these are rarely more
than 200 feet in height.” There are of course long extensions of the lowland
up the valleys of the larger rivers such as the Slave and Liard. Its surface in
general is poorly drained, with most of its streams leading toward the Mackenzie.
Muskegs and lakes abound.

Three other topographic features of southwestern Mackenzie should be noted.
Two of these are low plateaus that rise above the general level of the central
plain. The Horn Mountain Plateau lies about 20 miles north of the Mackenzie
between Providence and Simpson. Its surface is about 2500 feet above sea-level
and is said to be covered with muskegs and ponds (Whittaker, 1922, 1923).
The Eagle Mountain Plateau is southwest of Great Slave Lake and west of the
Hay River. It also reaches an elevation of about 2500 feet (see Cameron, 1922).
Horn Mountain and presumably also Eagle Mountain are capped by Cretaceous
rocks and correspond to the Caribou, Birch, and Watt Mountain Plateaus
farther south.

A third elevation above the central plain is the range known as the Franklin
Mountains. It more or less parallels the Mackenzie from a point near the
mouth of Willow Lake River northward beyond Great Bear River. Between
Willow Lake River and Wrigley the range ‘consists of sharp, but open, southerly
plunging folds, the axes of which control the trend of the river. Just north of
Willow Lake river, two ranges, about 10 miles apart, appear to represent the
east and west limbs of an eroded anticline, although it has not been established
that there are no intervening folds. The easterly ranges die out 4 or 5 miles
north of the latitude of Fish Lake river, the westerly range where traversed by
this river containing an open anticlinal axis, with its eastern limb truncated.
Farther north, due to the rising axis, older formations are exposed and the
mountains are high and more complex, and cover a broader belt.

“At Smith creek the western range has split into two, the eastern branch
being subdivided into several irregular ridges, extending in different directions.
Opposite Wrigley, the Roche-qui-trempe-a-l’Eau forms the westernmost ridge
of the Franklin system. Four miles to the east the mountains rise as a some-
what dissected mass, which may be grouped into two main ranges, with another
which forms the easternmost part of the system, dying out to the north . . . Cap
Mountain [northeast of Wrigley], . .. consists of a quartzite ridge dipping
steeply to the west and rising in elevation at its northern end where it forms
the highest mountain of the Wrigley region—its height being estimated at more
than 5000 feet above sea-level” (Williams, 1922).

“The western border of Franklin mountains, from the vicinity of Wrigley to
Saline river, is from 6 to 8 miles east of Mackenzie river. Exceptions occur in
the case of Roche-qui-trempe-a-l’eau; a small mountain ridge bordering the
Mackenzie 114 miles below Wrigley; and of low mountains situated 3 miles
east of the Mackenzie in the vicinity of Blackwater river. For 40 miles south
of Great Bear River, Franklin mountains are represented by a chain of about
six isolated knobs, none of which appears to rise more than 1000 feet above
Mackenzie river. Mount Charles, situated on the north bank of Great Bear
river, is a continuation of this chain, and rises about 1440 feet above Great Bear
river and probably 1500 feet above Mackenzie river” (Williams, 1923).

10 SARGENTIA [6

Great Bear River, which connects Great Bear Lake with the Mackenzie, has
cut its valley directly through the Franklin Mountains. Several other streams,
such as the Blackwater, rise in a lake country east of the range and have cut
deep valleys through it almost at right angles.

A minor elevation above the Mackenzie lowland is about 10 miles back from
the river and some 35 miles north of Simpson. It is plateau-like and is called
the Ebbutt Hills, about 1500 feet above the river level. Its top is said to be
covered with muskeg (Williams, 1922).

This brief discussion of the topography of the area will furnish a setting for
the following descriptions of exploratory history, geological formations, and
soils. It must be remembered that vast stretches of country away from the main
rivers and lakes remain to be explored, so that detailed topographic accounts are
impossible except in a few places.

HISTORY OF EXPLORATION

The first white men to visit southwestern Mackenzie were those in Alexander
Mackenzie’s exploring party of 1789. The narrative of this expedition gives
the course of the Slave and Mackenzie Rivers, outlines parts of the northwest
shore of Great Slave Lake, and gives the approximate position of the Caribou,
Horn, Eagle, Mackenzie, and Richardson Mountains. It is filled with observa-
tions on the plant and animal life of the region, as well as on the life of the
aborigines. Mackenzie’s voyage was designed to open the country for the fur
trade, and the great river soon became the principal highway in this trade and
the scene of its permanent establishments. No doubt a certain amount of ex-
ploration away from the main route was carried on by the traders, but no account
of it is available. Even the explorations of Sir John Franklin and his parties
(Franklin, 1823, 1828), which made such outstanding contributions to the geog-
raphy of the northwest, added little to the knowledge of our region. They
passed through it only as Mackenzie did, by way of the main river.

Attempts to use the Liard River as a trade route into the Cassiar region and
the upper Yukon proved futile, due to the difficulties of navigation. Part of
the upper Liard country was visited by the Hudson’s Bay Company trader
~ Robert Campbell, who traveled down the Dease and up the Liard and. Frances
Rivers to Frances Lake in 1840. Campbell established a route from Frances
Lake into the upper Pelly River by which he reached the Yukon. He is known
to have made a journey from the Yukon to Fort Simpson by this route and the
Liard River in 1852 (see Dawson, 1889, for a discussion of early travels on the
Yukon and upper Liard Rivers). The expense and risk of navigating the Liard
through the Rocky Mountains, however, forced the traders to give it up, and the
posts of the upper river were then supplied from the Pacific Coast.

It is unnecessary to enumerate all the travelers who have contributed to the
description of the upper Mackenzie River country, though a few outstanding ones
will be mentioned. Richardson’s account of his Franklin search expedition of
1848 (Richardson, 1851) is of particular value because of its general notes on
all of the Mackenzie basin. His collections made at the time of the Franklin
expeditions were described by W. J. Hooker (1840), while his general observa-
tions on natural history are to be found not only in his appendices to the Franklin
narratives and in the narrative of his journey of 1848, but also in other publica-
tions (1825, 1861). A few plants were collected in this period also by W. J. S.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 11

Pullen along the upper Mackenzie. Pullen was a British naval officer engaged
in the Franklin search. He spent two winters at Fort Simpson and traveled up
to Great Slave Lake in 1852 (Pullen, 1852). His specimens were enumerated
by Seemann in 1852-57.

Some valuable data on soil frost phenomena, as well as miscellaneous obser-
vations on the vegetation, were made by J. H. Lefroy in 1843 and 1844. He was
commissioned by the Royal Society to gather magnetic data throughout the
Mackenzie region, spending the winter of 1843-44 at Chipewyan and traveling
northward as far as Fort Good Hope in the following summer (Lefroy, 1857,
1883, 1886).

In 1859-60 Robert Kennicott, a naturalist working under the auspices of the
Smithsonian Institution and the Chicago Academy of Sciences, made a collecting
trip through the Athabaska—Mackenzie country. He spent the autumn of 1859
at Fort Simpson, making a trip of three weeks up to Fort Liard. In the fol-
lowing winter and spring, after another journey to Fort Liard, he visited Forts
Rae and Resolution on Great Slave Lake, and spent the summer of 1860 at the
latter place. He then descended the Mackenzie and crossed into the Yukon by
way of the Peel, Rat, and Porcupine Rivers. Kennicott made large zoological
and botanical collections and was extraordinarily successful at inspiring the local
traders to continue gathering material. A set of his plants is now in the
herbarium of the New York Botanical Garden (see Kennicott, 1869).

Emile Petitot, a Catholic missionary, spent many years in the Athabaska—
Mackenzie country, beginning about 1862. He published a series of papers
between 1875 and 1893 devoted primarily to the geographic features of the
country and its people. Few if any persons have ever traveled so widely in this
region or written so voluminously about it. His personal observations appear
to be thoroughly reliable, but they are so interspersed with hearsay and legend
that sorting them out is often difficult.

More accurate descriptions of the surface features and geology of our region
began to take form for the first time in the reports published by the Geological
Survey and the Department of Interior of Canada in 1888, 1890, and 1891.
These reports were based upon a series of remarkable survey journeys made in
1887 and 1888 by G. M. Dawson (1888), R. G. McConnell (1891) and William
Ogilvie (1890). Dawson and McConnell went to Dease Lake in Northern
British Columbia by way of the Stikine River and Telegraph Creek. From
Dease Lake they descended the Dease River to the Liard, where McConnell
left the main party and went down the latter stream. Dawson went up the
Liard and Frances Rivers to Frances Lake, then into the Pelly River by way
of the old trade route used by Campbell in the 1840’s. He then descended the
Pelly to the Lewes and returned to the coast by the latter stream and the Chilcoot
Pass. Dawson’s description of the western component of the Mackenzie Moun-
tains in the vicinity of Frances Lake is the first authentic account of the Macken-
zie range that is available.

McConnell surveyed the Liard to its junction with the Mackenzie in the
summer of 1887, and then spent the autumn of that year in the country along
the Slave, Salt, and Hay Rivers. He wintered at Fort Providence, making
journeys to Fort Rae in the north arm of Great Slave Lake. In the summer of
1888 he descended the Mackenzie and made the traverse to the Yukon by way of
the Peel and Porcupine. He returned to the coast by the Yukon and the Lewes.

Ogilvie spent the winter of 1887-88 on the Yukon, and then ascended the

12 SARGENTIA [6

Tat-on-duk River to its head and crossed a height of land to the headwaters of
the Porcupine. Then he went down the Porcupine and up the Bell to the Trout
River, a western tributary of the Peel. He reached the Mackenzie by way of
the Peel, and returned southward through the Mackenzie basin.

As a result of these three expeditions the Mackenzie River was more accu-
rately mapped, the Liard was surveyed for the first time up as far as the Frances,
and the configuration of the Mackenzie Mountain mass began to appear in its
true form. Also the broad outlines of the geology of the region were formu-
lated. All of the reports are filled with pertinent notes on the vegetation and its
distribution—particularly that of Dawson, who was one of the most discerning
naturalists ever to visit the Northwest. As appendices to his report he pub-
lished a classic essay on the distribution of trees and shrubs in Yukon and
adjacent British Columbia (see also Dawson, 1888B), a list of plants collected
on the journey, as well as notes on Indian tribes, zoological collections, rocks,
and astronomical and meteorological observations. For many years the plants
collected by him were the only specimens available from the vast inland region
of northern British Columbia and southern Yukon. Most of them are now to
be found in the National Herbarium of Canada.

The period of gold discovery in the Cassiar Mountains (in the 1870's) and
the later one in the Yukon region (1898) occasioned a great deal of travel and
exploration along the Liard and Peel Rivers. Many gold-seekers tried to reach
the Yukon by these routes, some successfully and some disastrously. (See the
reports by Dawson and McConnell cited above; also Graham, 1935 and Keele,
1910.) According to Keele, one of the parties crossed the Mackenzie Moun-
tains by way of the Gravel and Hess Rivers. Very few of these travelers ever
published adequate descriptions of the country they traversed so laboriously.
Another crossing of the Mackenzie Mountains in 1898 is reported by Keele.
This was made by a trader with a party of Indians from Fort Good Hope, who
started a post on the western side of the divide at the Stewart River.

In 1892 Miss Elizabeth Taylor traveled down the Mackenzie by the usual
route, making collections of natural history specimens. No published narrative
of her journey has come to light, but her collections are well known and have
been published upon by several students. Most of the plants are to be found
in the herbaria at the National Museum of Canada, the New York Botanical
Garden, and at Harvard. They bear excellent data as to locality and date, but
unfortunately most of the collection was numbered serially by specimens, with
some duplication and omission of numbers, so that duplicates cannot now be
recognized by the numbers alone.

A zoological collector, Frank Russell of the University of Iowa, worked at
Great Slave Lake and along the Mackenzie in 1893 and 1894. His report,
published in 1898, contains many valuable notes on forests and other types of
vegetation. He is one of the few travelers who have described any part of the
semi-open prairie country between Providence and Rae.

The map of the western and northern arms of Great Slave Lake was greatly
improved by Ogilvie (1890) and McConnell (1891), and pioneer geological
reconnaissance was done by McConnell especially around the western arm. The
first comprehensive geological reconnaissance at Great Slave Lake, however,
was made in 1899 by Robert Bell and his assistant J. M. Bell of the Canadian
Geological Survey (R. Bell, 1900; see also J. M. Bell, 1929). The significance
of ancient shore lines around the lake, pointed out by McConnell, was discussed

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 13

at somewhat greater length by Bell, who saw them in the eastern as well as the
western arms.

The first comprehensive biological investigation in the upper Mackenzie River
region was done by E. A. Preble and his associates of the United States Biologi-
cal Survey in 1903-04 (Preble, 1908). After an earlier trip in 1901 to Great
Slave Lake, Preble returned to Resolution in the spring of 1903, accompanied
by his brother, A. E. Preble, and Merritt Cary. Cary and A. E, Preble spent
the summer of 1903 collecting along the south shore of Great Slave Lake, and
along the Mackenzie down as far as Wrigley. Their chief bases were Hay
River, Providence, Simpson, the mouth of the North Nahanni River, and
Wrigley. While at the Nahanni they climbed Nahanni Mountain. They re-
turned to the United States in the autumn. Meanwhile E. A. Preble went up
the north arm of Great Slave Lake and through a chain of lakes to Great Bear
Lake. He descended Great Bear River to Norman, and went up the Mackenzie
to Simpson where he spent the winter of 1903-04. In the following summer
he continued his work down the Mackenzie to the Peel and Fort McPherson,
and then returned southward by the usual river route. Preble’s report, though
concerned largely with the animals of the region, is a veritable mine of biological
information on the whole Mackenzie basin. It also contains excellent general
descriptions of physiographic features, climate, and vegetation, with exhaustive
accounts of exploratory history. Some of the principal biogeographic boundaries
in the region had already been pointed out by Richardson, but it remained for
Preble to define them more accurately and to map them according to the Merriam
life-zone concept. Plants collected by the Preble expeditions are to be found in
the United States National Herbarium at Washington. An annotated list of
trees and shrubs was published in the report of 1908, but the herbaceous species
have never been listed except in the accession records of the Nationa! Herbarium.

A crossing of the Mackenzie mountains was first described by Joseph Keele,
of the Canadian Geological Survey (1910). Some of the western approaches
to the mountains were surveyed by McConnell in 1902 on the Macmillan River
(McConnell, 1906) and by Keele in 1904 on the upper Stewart River (Keele,
1906). In the summer of 1907 Keele made a reconnaissance survey along the
Pelly up to about 140 miles above its junction with the Ross River. He then
ascended the Ross to its headwaters, where he found a mountain pass to the
head of the Gravel River, which he descended to the Mackenzie. The journey
through the mountains required the entire winter and spring of 1907-08. Keele’s
report is a mine of first-hand information on the topography, climate, flora, fauna,
Indians, and geological structure of the Mackenzie Mountains. Until very re-
cently it was the only source of data on this little-known wilderness. I have
drawn upon it frequently and at length during our field work of 1939 and in the
preparation of the present paper.

The Ogilvie Mountains were crossed by Camsell in 1905 by way of the Stewart
River, Braine Creek, Nash Creek, and the Wind River. His report (1906), like
that of Keele, contains a wealth of data on the natural history of the country.
The physiographic province known as the Peel Plateau was first adequately
described as a result of this survey.

More intensive geological survey work around the western arm of Great Slave
Lake and along the upper Mackenzie was begun in 1916 by parties sent out from
the Canadian Geological Survey. A. E. Cameron was in charge of work around

*
‘

14 SARGENTIA [6

the lake and in the valleys of the Hay and Buffalo Rivers during the summers of
1916 and 1917 (see Cameron, 1917, 1918, 1922B; also Hume, 1921). His paper
on post-Glacial Lakes in the Mackenzie basin (1922A) was based in part upon
observations made during these years.

The years between 1920 and 1923 saw a great deal of geological survey
activity along the upper Mackenzie, largely inspired by the discovery of oil in
the Norman area. A report on the district between Great Slave Lake and
Simpson was published by E. J. Whittaker (1922); between Simpson and
Wrigley by M. Y. Williams (1922); on the North Nahanni and Root River
areas by G. S. Hume (1922) ; between Providence and Simpson by Whittaker
(1923) ; on the northern part of the Franklin Mountains by Williams (1923) :
the Redstone—Dahadinni Rivers area by Hume (1924); and on a part of the
lower Liard River by Hume (1923). The geology of the Norman area was
described by Hume in 1923, and by E. M. Kindle and T. O. Bosworth in 1921.
A general account of the geological structure of the whole Mackenzie River region
was published by D. B. Dowling in 1922.

A large amount of purely geographic information resulted from these surveys,
particularly for the eastern approaches to the Mackenzie Mountains and for the
Franklin Range east of the river. It was in this period also that the Survey
began to use a route to the upper Mackenzie by way of Fort St. John, B. C.,
and the Sikanni River. Their parties went to Peace River Crossing by rail
and then up to Fort St. John by boat. Here they took pack horses and traveled
northward to the Sikanni River, where they built scows. With the scows they
floated down the Sikanni and Nelson Rivers to the Liard and then to Simpson.
The advantage of this route was that the parties and their equipment could be
on the scene of the season’s work about a month before the opening of navigation
on Great Slave Lake, thus lengthening the possible field season by approximately
a third.

Although many useful notes on the occurrence of forest types are to be found
in the above reports, only a few plant collections were made. For the most part
they were gathered by such members of the field parties as had the requisite
interest. Brooke, Howe, and Bedford brought specimens from the southwest
and north shores of Great Slave Lake, and Bedford collected a few in the country
north of the north arm. One of the best collections was made by C. L. Crickmay,
an assistant to M. Y. Williams in 1922 along the lower Liard and upper Macken-
zie. Later, in 1945, Crickmay made another small collection in the Franklin
Mountains. In 1944 Mr. F. S. Nowosad of the Canadian Department of Agri-
culture brought a small but interesting collection from the lower Liard and upper
Mackenzie. All of this material is to be found in the National Herbarium of
Canada.

Three short papers by E. M. Kindle (1918, 1920, 1921), derived from obser-
vations made during his geological work on the Mackenzie, are of particular
interest from a phytogeographic standpoint. They deal with recent sedimenta-
tion along the main rivers of the Mackenzie basin, and with the arrival and
departure of winter conditions.

New maps of Great Slave Lake and of the Mackenzie and Slave Rivers were
made by the Canadian Topographical Survey during the 1920’s._ G. H. Blanchet
was in charge of the parties operating on Great Slave Lake, and he wrote an

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 15

excellent description of the lake and its environs (Blanchet, 1926). Most of
the region has now been photographed from the air, with new maps printed or
in preparation.

My own field work in the Athabaska—Great Slave Lake region has touched the
area treated in this paper only on two occasions. In 1927, en route to Great
Slave Lake, I made a few collections and notes along the lower Slave River, and
in 1929 our field party crossed from the mouth of Salt River to the Little Buffalo
by wagon, and made a canoe journey up the latter stream to Conibear (Moose)
Lake. Such collections as were available from the northern part of the Wood
Buffalo Park (north of lat. 60°) were incorporated in my botanical paper on
the Park (1935), while some descriptions of its vegetation types and the affinities
of its flora are to be found in two earlier papers (1930, 1933). The floristic
and vegetational features of the whole region around Athabaska and Great
Slave Lake were presented in two later papers (1936, 1946). Some further
contributions to our knowledge of the vegetation in the northern part of the
Wood Buffalo Park are to be found in an excellent paper by J. D. Soper on the
History, Range, and Home Life of the Northern Bison (1941). Mr. Soper
spent two years in the Park area (1932-34), traveling extensively and making
a general study of the biota.

Since the exploratory activity of the Geological Survey of Canada along the
Mackenzie and lower Liard in 1920-22, the most important work in our region
has been done in the Mackenzie Mountains. F. A. Kerr, a mining geologist,
published a short note on the topography and geology of the South Nahanni—
Frances River district in 1934; and the South Nahanni up as far as Virginia
Falls was examined by A. E. ‘Cameron in 1935 (see Cameron, 1936, 1938).
Both of these men refer to maps of the South Nahanni made by Fenley Hunter
in 1928, and of the Flat River country by R. M. Paterson about the same time,
but I have been unable to find reproductions of them.

Two papers, published in 1936 and 1937, are of particular interest to the
phytogeographer. Dr. Charles Camsell (1936) made a journey by plane from
Prince Rupert to Wrangell, and thence to Dease Lake. From here he flew
eastward over the plateau region of the upper Liard, and down to the Mackenzie.
His brief paper describing the flight contains some excellent photographs and
an account of the geography of this unexplored wilderness as seen from the air.
He points out, for the first time, the continuity and wide extent of the great
forested watershed of the upper Liard basin. It is this forest that appears to
isolate the alpine vegetation of the Mackenzie Mountains from that of the
Rockies. R. M. Anderson (1937) at once saw the zoogeographic significance
of the “Liard gap,” and emphasized its importance as a connecting route between
the “Canadian zone” fauna of the Mackenzie basin and that of the Yukon.

In 1937 Mr. Harry Snyder of Montreal took a survey party to Brintnell Lake
(shown on some maps as Glacier Lake), a small body of water near the head
of the South Nahanni River. The South Nahanni is exceedingly swift and
difficult of navigation, especially in its lower course. It flows in a deep canyon
for a considerable distance, and at about 150 miles above the Liard it forms
Virginia Falls, estimated to be about 300 feet-high. The country on either side
of the river is rugged and mountainous, making overland travel also difficult
and time-consuming. Only a few hardy trappers and prospectors have ever
penetrated the region by the ordinary means. Snyder had supplies and heavy

16 SARGENTIA [6

equipment taken up the river in barges as far as the falls, and then ferried both
party and supplies to Brintnell Lake by air. Accompanying Mr. Snyder were
Dr. George Goodwin, a zoologist from the American Museum of Natural His-
tory, and Mr. Fred Lambart, a surveyor. Triangulation points were set up on
the lake shores, and a topographic map was made of the lake and of the moun-
tains immediately surrounding it. This map was used to good advantage by
our party in 1939, and is in part reproduced here as Fig. 2.

Aerial photographs of the South Nahanni from the Liard to Virginia Falls,
of the valley of the Flat River, and of the Macmillan Lake country have been
made in recent years, and maps have been drawn from them. The discovery
of gold on the western tributaries of the South Nahanni has greatly increased
interest in the region, and further exploration will no doubt go forward rapidly.

The Arnold Arboretum expedition of 1939 reached Fort Simpson on June 8,
traveling by the usual steamer route from Waterways to Fitzgerald, and by air
from Fort Smith to Simpson. Supplies had been shipped to Simpson in the
preceding year. After about a week of collecting at Simpson, the party, equip-
ment, and supplies were flown to Brintnell Lake in a chartered Mackenzie Air
Service plane. We remained at the lake until the 20th of August, making col-
lections of the flora and records of its local distribution and the structure of
plant communities. Three weeks were then spent at Simpson, making further
collections. The return to Waterways followed the usual steamer route. Some
15,000 herbarium specimens were gathered during the season, about two-thirds
of which are of vascular plants and the remainder of lichens, mosses, and fungi.
Duplicates from this material will be found in various herbaria (see Raup, 1939).

Exploration of the central part of the Mackenzie range has been enormously
facilitated during the war by the construction of the Canol pipeline from Johnson
Crossing at the north end of Teslin Lake to Norman Wells on the Mackenzie,
A biological field party under the joint leadership of A. E. Porsild and A. L.
Rand, of the National Museum of Canada, traversed the pipeline road in the
season of 1944, making large collections which are now being worked up. Most
of Mr. Porsild’s botanical material was gathered on the western slopes of the
mountains, in the Territory of Yukon, but early in September he made a rapid
reconnaissance between Macmillan Pass and the Mackenzie, gathering such
specimens as were still available so late in the season.

Also in 1944, Dr. V. C. Wynne-Edwards, of McGill University, was on the
Mackenzie in connection with a study of fisheries. In late July he went up the
Canol Road to the Bolstead Creek Pump Station and made a collection of plants
from that vicinity. Earlier in the season he made another collection on the
upper slopes of Nahanni and Lone Mountains, west of the Mackenzie River near
the confluence of the North Nahanni; and he also gathered miscellaneous speci-
mens along the Mackenzie itself at various times during the season. Mr. Porsild
(1945) has published a list of his own and Wynne-Edwards’ collections from
the eastern slopes of the mountains, with descriptions of vegetation seen along
the Canol Road in this sector. His records are included in the present paper,
and I have drawn freely from his descriptive notes.

I am indebted to Dr. C. O. Hage of the Geological Survey of Canada for a
small collection of plants made at Trout Lake on Aug, 13, 1944. So far as I
am aware these are the only specimens available from the vast area beween the
Mackenzie and Liard, on the north, and the 60th parallel on the south.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 17

GEOLOGY AND SOILS
PRE-GLACIAL GEOLOGY

The bedrock that immediately underlies most of the area of southwestern
Mackenzie treated in this paper is of Paleozoic or more recent age. Such granitic
rocks as appear are Paleozoic or Cretaceous intrusives. The surface exposure of
pre-Cambrian granite, gneiss, and metamorphosed sedimentary rocks of the
Laurentian Plateau marks the eastern border of our region, and these rocks
undoubtedly underlie the Paleozoic sediments in at least a part of the country
east of the Mackenzie River.

Outcrops of rock occur at a few places along the Mackenzie and Liard Rivers,
but the principal sources of data on the content and sequence of formations are
in the Mackenzie and Franklin Mountains, where folding and faulting have
brought the deeper beds to the surface, or at the escarpments of the Alberta
Plateau and of the higher plateaus of the Horn and Eagle Mountains. It is
unnecessary for present purposes to describe these formations in detail. Ac-
counts of them are to be found in the reports of the Canadian Geological Survey
from the time of Dawson to the present, especially in the records of surveys made
between 1916 and 1923 (see discussion of exploratory history). Of greatest
significance from a phytogeographic standpoint are the content and surface con-
figuration of the beds that lie directly beneath the recent glacial or alluvial
deposits. It is from these beds that much of the material in the recent deposits
has come, and their topography has in large measure determined the disposition
of glacial deposits and the course of modern streams.

Region east of the Mackenzie Mountains

The bedrock nearest the surface in most of the region east of the Mackenzie
Mountains appears to be of Devonian shale and limestone. These are the rocks
that outcrop around most of the western arm of Great Slave Lake and along the
upper Mackenzie and lower Liard Rivers. They also appear in the Alberta
Plateau escarpment along the Hay and Buffalo Rivers. They are generally flat-
lying except in the Franklin Mountains, where they are folded and fractured.
The principal variations in the surface formations, other than the Franklin Range,
are three in number. On the western shore of the north arm of Great Slave
Lake limestone and dolomite of Silurian age appear, but they are covered by the
Devonian rocks not far from the lake shore. Higher plateaus rising above the
Alberta Plateau and the Mackenzie Lowland are composed of Cretaceous sand-
stone and shale. The Horn and Eagle Mountains are formed of these deposits,
and there is a long extension of them into the plateau of the upper Liard River
west of the Mackenzie Mountains. Cretaceous beds of similar nature also ap-
pear on the banks of the Mackenzie between Norman and the Blackwater River.
In the Norman area some beds of Tertiary sandstone and shale have been found
overlying the Cretaceous. They have been classed as Eocene,

In all the region east of the Mackenzie Mountains, therefore, the only obstruc-
tions to the free movement of the Keewatin ice mass from east to west were the
ridges of the Franklin Range and the comparatively low Cretaceous plateaus.
It is to be expected that lobes of the ice were at least deflected by these uplands,
and that subsequent morainic deposits should show a topographic relation to
them. At the same time the surface rocks were soft enough to furnish the

18 SARGENTIA [6

glacier with a heavy load of material to be mixed with detritus from the pre-
Cambrian rocks to the eastward and to be deposited as a heavy mantle of drift
when the ice receded.

Mackenzie Mountains

Most of the rocks exposed in the Mackenzie Mountains are sediments of
Paleozoic age. The eastern and western components of the range, however,
show notable differences in the composition of the beds. Along the Pelly and
lower Ross Rivers, Keele described crystalline schists which are widespread in
the Yukon region. They are succeeded by various beds of quartzite, argillite,
slate, chert, limestone, sandstone, conglomerate, and volcanic tuff, often ar-
ranged in closely folded strata. Argillite, chert, and dark-colored limestone
were also described on the upper Macmillan River by McConnell. Keele states
that the mountains at the head of the Ross River “are formed of alternating beds
of dark compact quartzite and grey shale and slate,” and: “About ten miles east
of the divide some yellowish crystalline limestones occur in low isolated cliffs
along the embryo Gravel river, but the principal rocks are dark sandy shales,
striped grey slates and micaceous sandstone or quartzose schists. Rocks of this
character extend eastward to Mount Sekwi, about fifty miles from the divide,
and then end abruptly. . . . A radically different geological province begins at
Mount Sekwi, and limestones, dolomites, sandstones, and conglomerates, etc., of
various bright colors, replace the sombre rocks to the westward.” These are the
most prominent rocks of the eastern component of the mountains, through which
the Gravel and other rivers have cut their valleys. It is these strata that have
been block-faulted to form the characteristic topography already described for
the eastern area (PLATE II). Near the Mackenzie, Cretaceous sandstone and
conglomerate are found folded into the front ranges.

The western part of the range has been intruded by masses of granite that
form the cores of the higher and more rugged mountains (PLATE I). These are
well described by Keele on the upper Ross, and they have been seen also on the
upper Macmillan and Stewart Rivers. Farther south they form the so-called
Snyder Range, west of the upper South Nahanni. Being much harder than the
surrounding sediments they have weathered into serrated ridges which stand out
conspicuously among the other mountain summits of the region. Keele says
that the only igneous rock seen along the Gravel River, on the other hand, was
a sill of diabase between beds of sandstone below the mouth of the Twitya River.

The structural history of the Mackenzie basin has been discussed by D. B.
Dowling in a paper published by the Geological Survey of Canada in 1922.
Dowling’s, conclusions were based not only upon the earlier work of Dawson,
McConnell, and Keele, but he also had access to later reconnaissance in the upper
Yukon region and to the extensive survey work done on the upper Mackenzie
after 1916. The Mackenzie Mountains appear to have been formed by the com-
pression and crumpling of somewhat thinner beds than those in the Rockies
farther south. The crustal movement took place before the deposit of the Ter-
tiary sediments at Norman, which are regarded on fossil evidence to be of Eocene
age. Dowling states that “the northern mountains were formed early in the
period of crustal compression which was inaugurated towards the close of the
Cretaceous. They, therefore, antedate the outer ranges of Alberta which were
formed at the close of the Eocene.” The effects of these later movements can

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 19

be seen in the tilting of the Tertiary beds at Norman and in the folding of
Tertiary deposits of the Wind River plateau to the west.

Dowling points out further that “the order of formation within the two groups
of mountains seems to have been opposite in direction. The Rocky mountains,
originating in a series of large fault blocks followed by the formation of over-
thrust fault blocks that formed mountain ridges to the east, show a progression
from west to east. The Mackenzie mountains, with which may be grouped the
Franklin mountains, originating with crustal movements followed by the regular
Rocky mountain type of thrust fault blocks, show progressive development from
east to west.” The batholithic intrusions in the western part of the Mackenzie
range are tentatively considered as Cretaceous in age. “They are intimately
associated with the earlier intrusions of the coast range, but although very
similar, lithologically, they were apparently intruded somewhat later . . . ap-
parently, therefore, the country west of the Mackenzie was deformed and in-
vaded by granite between early Cretaceous and Eocene times. The mountains
between the Mackenzie and Yukon Rivers may have been outlined at this early
period.”

GLACIAL GEOLOGY

Most of the soils and minor topographic features of southwestern Mackenzie
are products of erosion and deposition during the Glacial Period, of lacustrine
deposition during the retreat of the ice, or of strictly post-Glacial alluvial sedi-
mentation. On rock outcrops, particularly in the mountains, there has been a
certain amount of residual soil formation since the Glacial Period, but it is of
minor importance in terms of the total surface area of the region. Most of the
soils are youthful in point of time, for they originated with the final retreat of
the last ice or after the withdrawal of glacially impounded lakes. They are also
immature so far as the development of profiles is concerned, due in part to their
youth and in part to the subarctic climate of the region. At the same time the
vegetation gives every evidence of youth and immaturity in the development of
community equilibria. In regions where soils have reached maturity and plant
communities have reached “climax” stages, strong positive correlations between
soil and vegetational boundaries cannot be expected ; but where both are youthful
and immature the relation of plant communities to the age and structure of soils
and surfaces is apt to be a close one. The patterns of development and present
distribution among the types of vegetation in our region, therefore, can be ex-
pected to show a correlation with the physiographic events of late Glacial and
post-Glacial time.

In studies of vegetation in the Athabaska-Great Slave Lake region (1930,
1935, 1946), considerable evidence has been accumulated to show that such a
correlation exists. It was possible to do this because sufficient knowledge of
vegetational and glacial features had been gathered to draw up the principal
geographic patterns involved. The field observations necessary to a similar
synthesis in the upper Mackenzie country have not yet been made, though enough
are at hand strongly to suggest that the correlation is possible. The operations
of the Canadian Geological Survey in this area have been concerned primarily
with stratigraphic problems, but nearly all of the reports contain notes on glacial
features and forests. These observations, supported by some of my own, form
the principal basis for the discussion which follows.

20 SARGENTIA [6

Glaciation east of the Mackenzie Mountains

The great lake region of the Mackenzie was affected by glacial ice only from
the east, while in the area under discussion here the Cordilleran glaciers also
had a part in forming the present topography. So far as is known the eastern
ice completely covered the country up to the base of the Mackenzie Mountains.
It was thick enough to override most if not all of the Franklin Mountains, fill
the valley of the Mackenzie between this range and the front of the Mackenzie
Mountains, and to push into the lower valleys of some streams draining the
latter. Keele (1910) found a gneissic boulder in the Gravel River valley 1800
feet above the Mackenzie, and Williams (1923) states that “Glacial erratics lie
on the very top of Cap mountain and within 400 feet of the top of mount Clark.
The boulders are Precambrian and are evidently from the east.” Hume (1924)
estimates that the ice in the Mount Clark area, opposite the mouth of the Gravel
River, must have been at least 4300 feet thick. Deposits from this ice along the
Mackenzie River have a thickness of at least 200 to 250 feet (Alcock, 1937).

It has been shown that during the retreat of the ice from southwest to north-
east across the Athabaska—Great Slave Lake region a series of lobes occupied
the valleys of the Athabaska, Peace, Wabiskaw, and Hay and Buffalo Rivers.
These lobes were separated from one another chiefly by the Cretaceous plateau
remnants known as the Birch, Caribou, and Eagle Mountains. During re-
treatal stages lakes were impounded in the valleys, with bottoms at levels which
are now about 1600 feet, 1100 feet, 800 feet, and 700 feet above sea-level
(Cameron, 1922A). Sandy outwash materials and lacustrine clays were laid
down in the bottoms of the lakes, and in the most recent expansions of Athabaska
and Great Slave Lakes great alluvial deposits have been formed by the Athabaska,
Peace, and Slave Rivers. The topography of the ancient lake bottoms is re-
lieved by lobate morainic systems representing temporary halts in the retreat
of the ice.

In the country west of the Slave and Athabaska Rivers the most recent alluvial
soils have developed a flood plain forest of white spruce whose earlier succes-
sional stages involve willows and balsam poplars. Light soils on the older and
higher surfaces (800-ft., 1100-ft., 1600-ft.) have a somewhat richer forest of
white spruce which appears to have developed from one of jack pine. Heavier,
clayey soils on the uplands, however, show a semi-open prairie vegetation,
wherein the grassland is interspersed with groves of aspen and white spruce.
The largest of the prairies are on the 800-foot level, but they are also wide-
spread on the bottom of the 1100-foot lake. Undrained depressions on all sur-
faces develop a typical muskeg forest of black spruce and tamarack.

It is thus evident that, whatever the actual causes, there is in this area a geo-
graphic correlation between surface age and origin on one hand and the distri-
bution of certain types of vegetation on the other. It is of particular interest
that so far as is now known the presence of semi-open prairies in this region is an
indicator for the presence of glacio-lacustrine deposits of clayey soils.

Semi-open grasslands have been known for many years along the lower Liard
River (Lefroy, 1857; Petitot, 1893). They have also been described northwest
of Great Slave Lake along the trail between Providence and Rae, and in 1939 our
party visited a prairie area on the upland west of Fort Simpson (see below for
description). Grassy openings were frequently observed during our flight from
Smith to Simpson. They appear on both sides of the upper Mackenzie, and

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 21

usually they can be distinguished from marsh or muskeg by their surface con-
figuration (Raup, 1935). Further evidence for the wide extent of prairie in
this region is the fact that the wood bison formerly ranged over a large part of it
__as far west as the Liard and Horn Rivers and as far north as Lac la Martre
(Raup, 1933; Soper, 1941). The summer feed of the bison was in the upland
prairies.

In spite of this evidence for their existence, actual descriptions of grasslands
are extremely rare in the exploratory literature. This is probably due to the
fact that summer travel away from the main streams has nearly all been by canoe
through rivers and lakes, or in winter time over the uplands. The prairies are
not usually seen immediately along the streams, while in winter, under a cover
of snow, they are not easily distinguished from muskegs or marshes.

According to Cameron (1922A), a lobe of the ice lay in the upper valley of
the Mackenzie during the lake stages of 1600 feet and 1100 feet. It was con-
fined on the north by the Horn Mountain Plateau and on the south by the Eagle
Mountains (Cameron Hills). Its westward extent is uncertain, but Whittaker
(1923) described “sandy drumlins” along the Jean-Marie River 20-50 miles
above its mouth, and another moraine about 80 miles above the mouth. One
of these hills is said to be about 125 feet high. The upper moraine appears to
be the dam which holds back McGill Lake. The Jean-Marie River makes an
S-curve between its head and the point where it joins the Mackenzie. When
the moraines are located on the map it is clear that they are not far apart and
are probably portions of a single system whose trend is roughly north and south.
Although Cameron makes no mention of them, they are approximately at the
hypothetical margin of the glacial lobe at the 1600-foot lake stage. At the same
time the lobe that occupied the Hay and Buffalo River valleys reached south-
westward beyond the 60th parallel. Its terminus is marked by moraines ex-
tending across the Hay River valley between the Caribou and Eagle Mountains.

When the lake in the Hay River valley stood at about 1100 feet the upper
Mackenzie lobe had retreated eastward. Cameron draws its border in such a
way as to cover the Mackenzie valley to a point just below Mills Lake. He
does not describe terminal moraines in this area, but notes ridges of drift on
the lower Kakisa River which flows into the head of the Mackenzie from the
south. These ridges apparently are tied into the very prominent moraine which
more or less parallels the southwest shore of Great Slave Lake and holds back
the water of Buffalo Lake. This moraine probably was formed when the Hay
and Buffalo River lobe halted for the formation of the 1100-foot lake. To the
southeast it connects with the Nini-sheth Hills in the northern part of the Wood
Buffalo Park (Raup, 1935).

Glacial striae are in agreement with this general concept of the glacial lobes.
Those in the Hay and Buffalo River valleys point in a southwesterly direction
up the valleys, while those seen along the upper Mackenzie also have a strong
southwesterly trend (Whittaker, 1922, 1923; Cameron, 1922B). The upper
valley of Jean-Marie River, which trends in a northwesterly direction, was
thought by Whittaker to have been widened and deepened by ice.

Events corresponding to those just described should be found farther north,
between Simpson and Norman, but they are as yet undescribed. It is presumed
that one or more lobes extended westward on the north side of the Horn Moun-
tains, having their retreatal stages east of the Wrigley area and in the basin
of Lac la Martre. Williams (1922) noted glacial grooves on Cap Mountain

22 SARGENTIA [6

near Wrigley pointing 12-17 degrees north of west, and morainic ridges approxi-
mately at right angles to this near Smith Creek on the east side of the Mackenzie
a short distance south of Wrigley. On Mount Charles, north of the Bear River,
Kindle recorded deep glacial grooves trending east and west (see Camsell &
Malcolm, 1921). Williams (1923) saw striae on the west side of Mount Clark
running west 20° north. It is not known whether this ice front in the Mackenzie
valley was earlier or contemporaneous with the front placed by Cameron in the
valley above Simpson.

Lacustrine sediments were reported by Whittaker (1922) along the Trout
River beginning about 35 miles above the Mackenzie and extending for about
22 miles. This places them on the Alberta Plateau, and probably between the
moraines which marked the 1600-foot and 1100-foot lake stages. If the existence
and timing of the upper Mackenzie lobe are correctly correlated with the lobes in
the Hay and Buffalo River valleys and in the Peace River valley, the Trout River
lake should correspond to the one whose deposits are at about 1100 feet in the
Wood Buffalo Park. Vegetational evidence noted above indicates that this or
an equivalent lake was widespread along the upper Mackenzie.

Prairies northeast of Providence described by McConnell (1890) and by
Russell (1898) are probably on lacustrine clays deposited during a lower lake
stage. Judging by Cameron’s map (1922A, p. 351) they could have been laid
down in the bottom of the 800-foot lake, when the ice had vacated all of Great
Slave Lake except McLeod and Christie Bays.

McConnell (1890) saw low-lying stratified sands along the Liard for a distance
of some 25 miles below the mouth of the South Nahanni. He thought that they
were of lacustrine origin and that they might be correlated with simjlar deposits
seen along the Mackenzie near the mountains about 60 miles below Simpson.
This suggests a post-glacial lake extending northward along the base of the
Mackenzie Mountains. I can find no further discussion of such a lake by geolo-
gists who have investigated this region, and I saw no evidence of it on the flight
west from Simpson. With the exception of small areas on the upland imme-
diately west of the Mackenzie, semi-open prairies were not seen north of the
Liard River between Simpson and the mountains.

The timing of the events noted above is conjectural. In a previous paper
(1946) I have assembled some evidence which suggests that the morainic sys-
tems of our region represent retreatal stages of the Late Wisconsin (W,) ice.
Jt is suggested further that the last of this ice did not disappear from Great Slave
Lake until comparatively recent post-Glacial time, perhaps near the beginning
of the post-Glacial climatic optimum which is thought to have had its greatest
development between 3000 and 9000 years ago,

Recent sedimentation in southwestern Mackenzie

The surface of Great Slave Lake now stands at about 500 feet above sea-level.
Immediately following the disappearance of the ice from its eastern end, how-
ever, it is thought to have been about 100 feet higher and to have been in the
form of a great cross whose southern arm extended southward in the valley of
what is now the Slave River to about the latitude of Fort Smith (Cameron,
1922A, p. 352). The southern arm was partly filled with lacustrine deposits,
which now appear as cut banks of sand which reach their greatest height (about
125 feet) above the river at Fort Smith. The present level of Great Slave Lake

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 23

is thought to have been attained in large measure by a differential uplift which
was greatest toward the eastern end and tended to “spill” the water out by way
of the Mackenzie. During this period the Slave River cut its broad modern
valley in the lake deposits, and began laying down the huge masses of alluvium
that are gradually filling the southwestern part of Great Slave Lake.

Great Slave Lake has thus become a huge settling basin for material carried
by the Slave River. A few other streams from the uplands southwest of the
Lake (Hay, Buffalo) make their contributions, but the Slave brings in by far
the most material. Streams entering from the north carry only small quantities
of debris, for they come from areas where fine-textured rock materials are not
so abundant, and where timber and other vegetation are thin. The Slave River,
on the other hand, carries all the water from the Athabaska and Peace Rivers,
which in turn drain most of the southern and southwestern parts of the Mackenzie
basin. A large portion of the load carried by the Athabaska is left in Lake
Athabaska, and at certain parts of the flood season some of that from the Peace
is left there also. But most of the debris from the Peace goes on down the
Slave, and on occasions of very high water in the Athabaska there is a visible
procession of drift timber through the western end of Lake Athabaska and into
the Slave. All of this makes up a phenomenal amount of material brought into
the south side of Great Slave Lake. Kindle (1918) may well be correct in
saying that “There is probably no lake in North America which receives any-
thing like the amount of driftwood which is poured into Great Slave Lake,
chiefly through the Slave River.”

As a result of drainage and sedimentation, therefore, the entire southern arm
of Great Slave Lake has been eliminated, with the production of thick alluvial
deposits around the southwest shores. Immediately above the recent alluvial
deposits are broad lacustrine plains of clay and sand. Vegetational development
on the two types of terrain is sharply differentiated.

Kindle (1. c.) has pointed out the contrast between the Slave and upper
Mackenzie Rivers. Very little of the material carried in by the Slave ever
leaves Great Slave Lake. The water of the Mackenzie between Simpson and the
lake is clear. The Slave meanders in a muddy flood plain, with banks of local
silt deposits or undercut alluvium. The upper Mackenzie flows through glacial
deposits of boulder clay, from which the boulders have been sorted out to form
broad pavements on either side of the river. These pavements, so even in many
places that they look as though they had been laid by hand, have had a tendency
to act as retaining walls for the river, preventing the formation of meanders and
maintaining the stream in straight courses for many miles.

Below the mouth of the Liard the Mackenzie again becomes turbid, but the
Liard water remains on the west side of the Mackenzie for a long distance.
According to Kindle’s observations the water on the east side of the Mackenzie
remains clear for about 160 miles below the mouth of the Liard.

Fort Simpson is situated on an island near the west bank of the Mackenzie
just below the mouth of the Liard (Prater II). The island is about two miles
long and half a mile wide, separated from the west bank by a narrow “sny”
which is nearly dry at times of low water. It appears to be covered principally
by alluvium from the Liard, with the boulder clay of the region at its base.
Devonian shale appears beneath the glacial deposits along the Mackenzie near-
by, but none was observed at Simpson.

The village is near the upper end of the island, on a bluff about 50 feet above

24 SARGENTIA [6

the river. In cross-section the island is highest on the Mackenzie side, sloping
off by a series of low steps or escarpments to the southwest. All of the lower
parts of the island are marked by what appear to be old longitudinal channels
formed and abandoned by the river as it cut its present course.

Further description of the Simpson area will be found in the discussion of
vegetation.

Glaciation in the Mackenzie Mountains

Glacial ice in the Mackenzie Mountains was thought by Keele (1910) to have
been confluent with the Cordilleran ice mass of the Rocky Mountains, and to
have had its gathering ground on the western slopes of the range. He estimated
its thickness in the Ross River region at about 3000 feet, which would mean that
it did not cover the higher peaks. He states that “the movement of the ice
during its maximum development was controlled to a great extent by the main
drainage valleys, and flowed down those almost, but not quite, to the Yukon
River. When the ice became thick enough on the western slopes . . . it began
to pour through the gaps and passes of the divide and to send streams down the
valleys of the Gravel river and its branches.” It is of further note that Keele
thought the ice divide shifted from west to east. He found granite from the
western slopes far down the eastern side of the range, and considered that during
the period of shrinkage the divide shifted to the present watershed because the
drift which he found in the pass between the Ross and Gravel Rivers was
deposited in a zone of stagnation where there was no movement of the ice.

The exact relationship between the two glacial systems at the eastern front
of the Mackenzie Mountains is yet to be worked out. Keele found Cordilleran
glacial deposits overlying those from the east in the lower valley of the Gravel
River, indicating that at least at one stage the eastern ice preceded the Cordilleran.
One of the most striking topographic features seen from the air while flying west
from Simpson is the complex system of moraines at the eastern base of the
mountains. They have a well-defined lobate form, convex eastward, and were
obviously formed during retreatal stages of valley glaciers. The unmodified
form and continuity of their outlines indicate that they have not been disturbed
by any later ice from the east. They must therefore be of the same age or
younger than near-by deposits of material from the east, but whether the latter
underlie them is of course unknown. The Cordilleran moraines extend a short
distance away from the base of the mountains, perhaps a few miles, but what the
extreme eastern limits of the ice were is yet to be determined. Striae east of the
mountains are prevailingly westward in direction, but Whittaker (1922, 1923)
reports two sets along the Trout and Redknife Rivers on the Alberta Plateau
south of Mackenzie between Providence and Simpson. The apparently older
set bear S. 50° E. and S. 45° E., respectively, suggesting a western origin for the
ice in this area at at least one period.

Valley glaciation similar to that described by Keele was ‘noted by Camsell
(1906) in the Ogilvie Mountains at the head of the Wind River. He states,
“All data collected with regard to the glaciation of the mountain section of the
Wind river point to the conclusion, that the region was not completely covered
by a great ice sheet during the glacial period; but that the valleys alone were
occupied by glaciers.” He proposed a glacier about 1000 feet thick flowing north-
ward down the valley of the Wind, producing a U-shaped profile and leaving
heavy deposits of gravel and clay on the valley floor. Camsell crossed the moun-

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 23

tains by way of Braine and Nash Creeks, in a pass estimated to be about 3500
feet above sea-level. He gives the elevation of the highest peaks there as about
6800 feet. He says that the region ‘‘was not covered by a large continental ice
sheet, but the valleys alone were filled to a depth of 1000 to 1500 feet with valley
glaciers. . . . Evidence of glaciation can be traced to a height of 4500 feet above
sea level, so that about 2000 to 2500 feet of the highest peaks protruded through
the ice.”

GEOLOGICAL OBSERVATIONS AT BRINTNELL LAKE

Brintnell Lake, near the head of the South Nahanni River, was, as previously
stated, the scene of about two months’ botanical exploration by the Arnold
Arboretum expedition of 1939. Because the following discussion of the flora
and its distribution involves repeated reference to this area, some observations
concerning its topography and geological history are necessary.

Brintnell Lake is about 31/4 miles long and about half a mile wide at its widest
point. The trend of its main axis is approximately W. 20° N., while the position
of its western end is about in latitude 62° 5’ N. and longitude 127° 35’ W. Its
altitude is about 2600 feet above sea-level (see Fig. 2; Pirates I, III, IV, XI).
The bottom has not yet been found, but it is evidently quite deep. Mr. Harry
Snyder, who was there with a survey party in 1937, informed me that he put
down about 1000 feet of cable without reaching the bottom. The lake is held
to its present level by a rock barrier at the eastern end, through which the out-
flowing stream pours in a narrow, unnavigable gorge. This stream joins the
South Nahanni about 5 miles down the valley and perhaps 800 or 1000 feet
below the lake.

The mountains rise steeply from the north and south shores to heights of
6000 to 7000 feet above sea-level. About two miles west of the lake there is
a higher peak that attains an elevation of approximately 8500 feet. Several
streams feed the lake from the neighboring slopes, but the main intake is from
a creek which flows through a great U-shaped valley to the west (PLATE IV).
This creek is fed by glaciers in the high mountains at the head of the valley about
7 miles above the lake. The valley floor west of the lake is made up of gravel
plains and broad marshy flats of light-colored glacial clay. When our party
arrived on June 16 the lake water was nearly clear, but within a few days it
became “milky” from glacial melt water and remained so throughout the summer.

The mountain rocks immediately surrounding the lake are Paleozoic shales
and slates. The higher peaks to the west and north, however, are composed of
light-colored granites which produce jagged, serrate profiles. The contact be-
tween granites and sediments is clearly seen where it angles up the side of a
mountain just west of the lake (Pate III). The presence of granite peaks in
the upper South Nahanni country was not known until quite recently. In terms
of Keele’s division of the Mackenzie range into eastern and western phases,
this region should be placed with the western section, where intrusive granites
form the cores of the higher mountains.

The slates and shales dip sharply to the southwest, forming high cliffs and
talus slopes on the north sides of the mountains (PLaTe X). The south-facing
slopes are somewhat less steep, but due to the sharp angle of the strata nearly
all of the slopes are precipitous. The valley that contains the lake, therefore,
is similar to the fault block valleys described by Keele along the Gravel River,
and it shows a similar orientation of dip and strike.

26 SARGENTIA [6

The contrast between the topography and soils of the shale and granite areas
is striking. A small stream, Frost Creek, entering the north side of the lake
near the western end, issues from a hanging valley in the shale mountains
(Pirate III). The lower end, or rather the rim, of this valley is about 1400
feet above the lake. From various points in the valley and on its lateral slopes
there are good views of the higher topography formed from this kind of rock.
In spite of the steep dip of the strata, many of the upper slopes have become

AW Scale of mites
XS oa 0 { Z

& Glaciers

ZX Fling &
rab Ue =e <4 Terrace
\< sl \- .
i: ; a

ey

ACRE

(27°45
127°35'

62°

Fic. 2. Map of Brintnell Lake and vicinity, redrawn from a manuscript map made by
Mr. Fred Lambart in 1937.

rounded and have produced through weathering a fair amount of soil (PLATE
X). Below the summits, however, there are large areas of slide rock, much
of which is still in motion, forming great “rivers” of rock down the slopes and
in the higher gullies. The degree of stability in this slide rock determines, and
is in turn conditioned by, the amount of vegetation growing on it. North slopes
are of course cliff-like and at the higher elevations are almost completely devoid
of vegetation. ‘The talus here is so active that plants can hardly get a footing.
By contrast the country a little farther west, where the granite peaks rise out

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 27

of the shales, shows very little soil produced in situ. The granite cores are
exfoliating in enormous blocks and slabs to form ponderous and almost sterile
talus slopes. The granite peaks show almost no green plant life.

The break-up of the mountain rocks and the formation of talus is going on
continuously and rapidly. While we were camped at the lake we frequently
heard the rumbling sound of landslides, and occasionally we saw clouds of dust
rising from the high slopes or caught sight of huge blocks of stone as they
tumbled off the cliffs and bounced to a standstill on the talus. Conspicuous
features of the lake shores are points formed by steep fans of rock debris (PLATE
IV). These are most highly developed on the south shore at the foot of the
precipitous north slope of Colonel Mountain.

The shales contain many stringers of calcite. These are most abundant on
the mountain south of the lake, a fact which appears to be reflected in the vege-
tation, for there seem to be more calcicolous plants there than on Red and
Terrace Mountains on the north side.

The glacier at the head of the main stream is obviously the remnant of a larger
one which once flowed down the lake valley. The U-shaped form of the valley,
with its sharply truncated transverse ridges, is clear evidence of this (PLate IV).
The truncated slopes have a gradient of about 34 degrees from the horizontal
and descend into the lake with scarcely any shore formation (PLaTE IV). All
of the fans mentioned above postdate the retreat of the ice.

Glacial deposits were found up to an altitude of approximately 4000 feet in
the main valley and 5000 feet in the valley of Frost Creek. In the main valley
they are of two kinds. On the south-facing slopes north of the lake are ridges
of gravel and boulder clay festooned diagonally down the steep slopes in a south-
easterly direction. The channels of streams cutting through the ridges are filled
with boulders from the till. These deposits are rather loose and uncompacted,
with little or no evidence of cementing. On the opposite side of the lake, masses
of till were found packed into the great gullies on the north slope of Colonel
Mountain, indicating that the gullies and intervening hogbacks were formed
before the last advance of the glacier. Presumably the till completely filled the
gullies to a height of about 4000 feet, at which level the remains of it form
shoulders on the truncated fronts of the hogbacks (PLate IV). The fact that
these shoulders have truncated surfaces which are confluent with those of the
underlying rocks of the hogbacks indicates that this till antedates the last advance
of the ice and must have been deposited during an older advance and retreat.

In some places tributary gullies have been formed on the lateral slopes of the
hogbacks in such a way as to excavate the back sides of the old till deposits.
In one such place a section of the till about 200 feet thick was examined (PLATE
V). The lower part is of fine sandy or clayey material interspersed with beds
of coarser gravels. Above this comes a bed of unsorted till containing coarse
material, and then the whole section is capped by a mass of heavy boulders set
in a matrix of finer materials. The entire section shows a large amount of com-
pacting and cementing, some of it almost resembling tillite. It is breaking up in
great columnar blocks which are often capped by large boulders that give them
a castellated appearance.

These two tills differ not only in age, position, and degree of cementation,
but also in the composition of their included rocks. Gravel and boulders washed
out of the younger deposits are mostly of granite and shale, while in the old

28 SARGENTIA [6

compacted till there are massive blocks of conglomerate, cherty stones, and
blocks of what appear to be a reddish quartzite.

North of the lake, in the hanging valley mentioned above, are tills which
form terraces on the valley slopes. The highest of these terraces is at about
5000 feet, near the rim of the valley, and is composed of unsorted drift in which
the boulders are principally of granite (PLates III, VI). Large erratic bould-
ers are frequent on the surface. There are at least two sets of the terraces,
representing two periods of stream dissection through the drift. The lower is
of very loose gravelly till, also made up principally of granite stones.

There is therefore unmistakable evidence of two advances and retreats of a
glacier in the main lake valley, with possible reflections of both of them in the
terraces of the hanging valley. The character of the rocks in the older till of
the main valley suggests that they came from the western slopes of the Mackenzie
range. If this is true it is consistent with Keele’s findings in the Ross and
Gravel River region. Also the prevalence of granites in the younger tills sug-
gests that the center of spread for the main ice mass may have shifted eastward,
as Keele thought it did. The absence of western rocks in the drift of the hang-
ing valley might well be due to its local origin as a tributary glacier among the
granite mountains immediately north of the lake.

On the rock barrier at the eastern end of the lake are some gravelly and sandy
ridges that look like outward deposits formed during the recession of the ice.
Similar deposits form gravel plains on the valley floor west of the lake.

The influence of frost upon the formation and movement of soils is everywhere
to be seen. Well-defined stone rings were found in a springy area on the till
in the upper valley of Frost Creek (PLate VI). The rings range in diameter
from a few inches to six feet or more. Many of the upper mountain slopes are
literally covered with turf-banked terraces in varying stages of stabilization
(Pirate IX). The rounded contours of many of the higher slopes on the shale
mountains appear to be due to long-continued soil formation and _solifluction
caused by frost (PLate X). In these higher altitudes frost action presumably
has been continuous throughout the Glacial Period, as it is very probable that
summits above 5000 feet were not covered by glaciers. Similar topography is
to be seen on many mountains in the South Nahanni country (PLATE II), as
well as along the Canol Road (see above).

SUMMARY OF TOPOGRAPHIC AND GEOLOGIC OBSERVATIONS

From a phytogeographic standpoint, the following topographic and geologic
features of southwestern Mackenzie are of outstanding interest. Most of the
soils, though immature, are deep and derived in large measure from relatively
soft rocks. The principal exceptions to this are on the higher slopes of the
mountains and in areas near the margin of the Laurentian Plateau. The pres-
ence of beds of limestone and calcareous seams in the slate has made possible
the growth of calcicolous plants throughout most of the region. The oldest land
surfaces are on the unglaciated higher slopes of the Mackenzie Mountains. Next
in age are the tops of the Horn and Eagle Mountains which separated the re-
treating lobes of the Keewatin ice sheet. The retreat of the Cordilleran and
Keewatin ice opened next the Alberta Plateau and the Mackenzie Lowland, as
well as the lower valleys of the Mountains. There appears to be a sequence

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 29

in age on the surfaces east of the mountains, progressively younger from west
to east; so that the most recently exposed lands are old lake bottoms imme-
diately surrounding Great Slave Lake and newly formed flood plain and delta
deposits on the shores of the lake itself. In the mountains the most youthful
surfaces are near the heads of the interior valleys, where considerable areas have
been uncovered by the recent retreat of the glaciers.

The Mackenzie Mountains have certain features that are of primary sig-
nificance to the plant geographer. First, they are sufficiently high so that their
summits and upper slopes are far above timber line, making possible the devel-
opment of an arctic alpine tundra vegetation. Second, they are isolated from
neighboring ranges by comparatively low, timber-covered plateaus and lowlands,
so that their alpine areas are not connected with those on the neighboring ranges
or with the arctic tundra. Third, they have a great variety of rocks, among
which there is a demarkation between eastern and western groups (just east of
the divide in the latitude of the upper Gravel River). East of this boundary
massive limestone, dolomite, and shale are very prominent, while west of it
quartzite, argillite, cherty rock, and sandstone are present in profusion along
with limestone. Furthermore the western group is intruded by great masses
of batholithic granite. Calcicolous floras can be expected therefore to develop
more extensively over the limestone of the eastern section, while at the same
time a greater instability of soils can be expected there due to the ease with
which limestone and shale are broken up by frost action. Botanical effects of
the difference in rock types are not simple, however, for, as Keele has pointed
out so clearly, the climatic and topographic differences between the east and
west slopes of the range are sharp and striking. The west slopes receive heavy
precipitation, which is carried off in broad, more nearly graded valleys, while
the eastern slopes have comparatively light rain- and snowfall and the valleys
are steep and canyon-like.

CLIMATE AND AGRICULTURE

The difficulties involved in a phytogeographically significant description of
the climate o7 southwestern Mackenzie are well-nigh insurmountable with our
present knowledge. It is presumed that when data gathered during the war
are released the situation will be much improved, but until that time we must be
content with the scanty records and interpretations now available.

In previous papers (1934, 1935, 1946) I have already discussed the climatic
phenomena of the Peace River region and of the Athabaska—Great Slave Lake
district. So far as is known, that of the upper Mackenzie valley is not far differ-
ent from that of the northern part of the Wood Buffalo Park and the western
Great Slave Lake country. Consequently there is no need for a repetition of
the discussion here. I shall limit this treatment, therefore, to a brief and general
characterization of the climate of the region of Fort Simpson; and I shall attempt
to draw up some contrasts between the climate of Simpson and that of the
Brintnell Lake district in the Mackenzie Mountains. On the occasion of our
1939 expedition we kept records of maximum and minimum temperatures at
Brintnell Lake, and notes on local weather. These can be compared profitably
with simultaneous records made at the government weather station in Simpson,
and with the daily weather maps for the same period.

30 SARGENTIA [6

CLIMATE AT SIMPSON

I have included in Taste 1 equivalent measurements for Fort Norman and
Fort Smith, chiefly in order to relate the Simpson area to its surroundings and
to my earlier discussions for the Athabaska—Great Slave Lake country.

It will be seen at once that there is a close similarity in the climatic features
of the upper Mackenzie valley and the Slave River country. The climate as a
whole is continental in character, with short, relatively warm summers and long
cold winters. Even though high temperatures are sometimes reached at Simpson
(the highest recorded is 94° F.), the growing season is short and hazardous.
July minima of 31° have been recorded at the weather station, so that a com-
pletely frostless season cannot always be depended upon. There is an average

TABLE1.!
SUMMARY OF METEOROLOGICAL DATA FROM Forts SMITH, Stimpson, AND NORMAN
Total Rainfall | Mean temperature | Mean daily range
Total June,
annual :
ides rainfall | July,
or: Aug. Jan. July Jan. July
Fort Smith 12.67 8.55 4.05 —15.1 59.9 16.1 26.7
1913-1934
Fort Simpson 13.45 7.67 4.77 —19.2 61.6 16.4 23.3
1897-1927
Fort Norman 10.70 7.02 5.05 — 18.6 59.2 14.6 24.5
1908-1928

* Data extracted from publications by Connor (1937, 1938).

season free of frost, however, of between 70 and 75 days. Between 40 and 50
percent of the annual precipitation falls in the months of May to August, inclu-
sive, and the mean annual snowfall is between 40 and 60 inches. The wettest
month at Smith and Simpson is July, but farther north, at Norman, it is August.

An outstanding feature of the climate from a botanical standpoint is the low
yearly precipitation. This is especially notable when the three months of the
most active growing season are considered (June, July, and August). Such
low precipitation in a southern climate would be a concomitant of desert condi-
tions, but in the Mackenzie region there is no apparent lack of available moisture
in the soil. It is presumed that the source of this moisture is in the permanently
frozen subsoil, and that the availability of the mositure is dependent upon sum-
mer temperatures sufficiently high to release it. Climatic values that might be
critical in the delimitation of vegetation, therefore, probably should be looked
for in temperature rather than in precipitation.

SEASONAL PHENOMENA AND AGRICULTURE AT SIMPSON

Detailed consecutive observations of seasonal phenomena are not available
for most of the Mackenzie settlements, although a few outstanding events such
as the opening and closing of navigation have been recorded faithfully. We are
dependent upon random notes made by various travelers for piecing together the

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 31

progression of changes that occurs during the year. The principal sources of
published information for the vicinity of Simpson are in papers by McConnell
(1891), Preble (1908), Kindle (1920), and Albright (1933A, 1937). These
will be drawn upon freely in the following notes.

Warm weather begins in spring during late April or early May. In 1888
McConnell reported that the temperature rose above freezing for the first time
on April 20, and after May 1 the snow rapidly disappeared. Preble states that
in 1904 the spring was somewhat earlier. In that year the average daily maxi-
mum temperatures rose above freezing during the first week of April, but the
average daily minima did not rise above 32° until the fourth week. They again
fell below it in the first week of May, and rose above it for the season in the
second week of May.

Preble noted that the buds and catkins of willows and alders “imparted a
brown tinge to the hillsides” on March 30. The snow had nearly disappeared
from the open fields by April 18, and the sap of the white birches began to flow
freely on April 20. Mosquitoes also first appeared on April 20. A little snow
fell on May 6, and the leaves of Ribes oxyacanthoides began to appear May 8.
Leaves of aspens and white birches were half an inch in length on May 14, and
violets bloomed at about the same time. Viburnum edule and Populus Tacama-
hacca were in leaf by May 18, and Vaccinium Vitis-Idaea was in flower. Ribes
oxyacanthoides and Calypso bulbosa were in flower on May 22. ‘There were
several inches of snow again on May 29. The leaves of the larch were first seen
on June 2 a few miles below Simpson.

The ice in the Liard river breaks up in spring before that of the Mackenzie.
In 1904 this occurred on April 29. The Liard ice drove a channel across the
Mackenzie and piled up high on the north bank. The breakup of the Mackenzie
ice below the Liard did not occur until May 2. According to Kindle this was
a little earlier than usual, for he states that the average dates of opening at
Simpson are between May 10 and 15.

The Mackenzie ice does not open between Great Slave Lake and Simpson until
two and a half or three weeks after the ice has gone out below the latter place.
Great Slave Lake itself usually opens between June 16 and July 2.

In the autumn the larger bays of Great Slave Lake become covered with ice
in October, early or late, depending upon the season. In the wider parts of
the lake the water may remain open until mid-November. The upper Mackenzie
usually closes in the latter half of November, though it is sometimes closed to
canoe navigation in mid-October. In terms of vegetation growth, the season
ends during the latter half of August or early in September. Most of the native
flora has set its fruit by the last week in August.

At the tirne of our arrival in Simpson on June 8, 1939, the spring was far
advanced. About fifty species of plants were found in flower there during the
week between June 9 and June 16. We experienced the first frost at Simpson
on August 24, soon after our return there from Brintnell Lake. Plant collecting
continued to be profitable until about mid-September, though most of the vascular
flora had already matured and lost its seed. The first killing frost has sometimes
been reported quite late. Albright states that it did not occur in 1930 until Sep-
tember 24, and Kindle recorded the first notable freeze in 1920 on September 25.
Kindle noted two to three inches of snow at Providence on October 1.

Successful gardens have been grown at Simpson for many years. Root crops,
such as potatoes, carrots, and rutabagas, are particularly successful, and some oats

32 SARGENTIA [6

and barley are raised. Squaw corn will not mature, but will produce roasting
ears. Vegetable marrow is raised from seed, and tomatoes will occasionally
ripen out of doors. Most tomatoes are taken in to ripen, however, for it is esti-
mated that they will ripen on the vine in only about two years in ten.

Approximately fifty acres are under cultivation, mostly on the island, but in .
part on a low flood plain on the west bank of the river. No cultivation has been
attempted except on the alluvial soils along the main river. Domestic livestock
is wintered successfully, supplementing the feed with wild hay cut in the neigh-
borhood. Some of this hay comes from the semi-open prairies on the upland
west of the river. Of crops raised in excess of local needs, potatoes and ruta-
bagas are most important. They are shipped down the river to missions where
gardening is not so profitable.

COMPARISON OF THE CLIMATES OF SIMPSON AND BRINTNELL LAKE

Our party spent 61 days at Brintnell Lake between June 16 and August 20.
During this period 60 maximum and minimum temperatures were recorded with
a Sixes type thermometer placed in a protected situation on the shore of the lake
about four feet above the ground. The altitude of Simpson is approximately
420 feet above sea-level, and that of the lake about 2600 feet. Taster 2 sum-

TABLE 2.

SUMMARY OF METEOROLOGICAL OBSERVATIONS AT SIMPSON AND BRINTNELL LAKE,
JUNE 16-AuGusr 20, 1939

Simpson Brintnell L.

Average maximum temperature 72.8 65.7
Absolute maximum temperature 84.0 83.5
Average minimum temperature 48.6 43.6
Absolute minimum temperature 36.0 32.0
Maxima

No. of days 80° or above 14 1

No. of days 72° or above 37 13

No. of days 65° or below 9 25

No. of days 60° or below 4 13
Minima

No. of days 50° or below 39 55

No. of days 40° or below 2 15

No. of days 32° or below 0 1
No. of days on which Brintnell L. maxima were higher than those

at Simpson (56 days compared) 10
No. of days on which Brintnell L. minima were higher than those

at Simpson 8
No. of days on which rain fell 19 29
No. of clear or partly cloudy days eye 54 41

marizes the observations taken at the lake, and offers a comparison with those
derived from meteorological records at Simpson for the same period of days.
Temperatures are in °F. and precipitation in inches.

The mean and absolute maximum or minimum temperatures do not contrast
the two places so well as the distribution of the maxima and minima. It is about
6 degrees colder at the lake as shown by the averages, but from the standpoint of
growing conditions it is more significant that the maxima rose to 72° or above
on nearly 3 times as many days at Simpson as they did at Brintnell Lake, and

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 33

that they remained at or below 65° on 25 days (about 42% of the total) at the
lake as against 9 days at Simpson. Likewise it is notable that the minima fell
below 50° on 55 nights at the lake, and below 40° on 15 nights, against 39 and
2 nights, respectively, at Simpson. ‘Reversals of the usually lower temperatures
in the mountains were noted on 10 days for maxima and 8 nights for minima.
The daily ranges of temperature do not differ greatly. At Simpson the greatest
was 37° anc the least 6° (mean 24.6°), while the greatest at the lake was 36°
and the least 4° (mean 22°).

Periods of storminess were somewhat more numerous at the lake. The ac-
companying chart (Fig. 3) shows the approximate dates of the stormy periods.

Although no rainfall measurements were made at the lake, the amount for the
60 days can safely be estimated as at least two or three times as great as that
at Simpson. In the same period only 1.29 inches were recorded at the latter
place. On 19 of the days for which rain was recorded at the lake it was noted
as light rain or mist, but on the other 10 days there were heavy downpours
lasting from one to several hours.

Dates 4 5 10 15 20 25 30
Simpson
June
Brintnell Lake YO, EZ
Simpson \LZ3. Yt; Win BZ Yt
July
Brintnelt Lake ZZ ULL, Cae “ZA UAE
Simpson WLLL
Auqust
Brintnell inte WWE EEE YJ

Fic. 3. Chart showing a comparison of periods of storminess at Simpson and Brintnell
Lake between June 16 and August 20, 1939.

It will be noted that in the stormy periods of June and most of July there
were coincidences between the two areas, but in August the correlation is not
so clear. Examination of the daily weather maps for the period in question
shows a relationship between the occurrence of storminess in the Brintnell Lake—
Simpson region on the one hand and the appearance of pronounced high pressure
systems moving in from the Pacific over southeastern Alaska and northern
British Columbia, in contact with low pressures moving eastward and southeast-
ward over central Alaska and Yukon. These lows appear to follow two courses.
They may come over the lower Mackenzie, apparently moving eastward around
the northern part of the Mackenzie Mountain mass, and thence southeastward up
the Mackenzie valley. Or they may move eastward over the upper Yukon and
upper Liard valleys, reaching the upper Mackenzie region by way of the Liard
gap. The stormy periods that appeared at Simpson and Brintnell Lake at about
the same time appear to be most closely related to the former of these two storm
tracks, although the latter sometimes affects Simpson also. It will be noted that

34 SARGENTIA [6

in four of the periods the Brintnell Lake region was affected a day or two earlier
than Simpson.

The storm of July 21-25 at Simpson seems to have been at the contact between
a pronounced low over Keewatin and a pronounced high coming in from the
north Pacific. This complex did not affect the Brintnell Lake area until the
23rd, when a low appeared from central Yukon, approaching on the west side
of the Mackenzie Mountains.

PLANT COMMUNITIES OF SOUTHWESTERN MACKENZIE
INTRODUCTORY

It has already been made clear that the botanical exploration of southwestern
Mackenzie is scarcely more than begun. This applies not only to purely floristic
aspects, but also to knowledge of vegetational structure. Descriptions of plant
communities drawn in anything approaching detail have been made in only three
areas: Fort Simpson, Brintnell Lake, and the vicinity of the Canol Road be-
tween Norman Wells and Macmillan Pass. <A fourth point of departure is to
be found in my earlier studies (1935, 1946) on the Athabaska—Great Slave Lake

_region, which overlaps the southeastern part of the area now under consideration,

The vegetation east of the mountains is considerably better known than that
in the mountains themselves. From notes made at Simpson, along the upper
Mackenzie, and in the Wood Buffalo Park, together with casual observations
made by various travelers in the past, it is possible to form a fair picture of the
types of vegetation in this area. There are, however, large lacunae to be ex-
plored, especially on the outlying Cretaceous Plateaus (the Horn and Eagle
Mountains) and on the Franklin Range. Likewise the semi-open prairies along
the Mackenzie and lower Liard and north of Providence remain to be studied.
Our examination of the grassland openings on the upland west of Simpson hardly
does more than point to this problem. For the Mackenzie Mountains it is
impossible to do more than describe the isolated areas that have been seen and
discuss the affinities of their flora, with full appreciation of the fact that only a
beginning has been made.

In the ensuing discussions I shall divide the region as suggested above, first
putting together what is known of plant communities outside the Mackenzie
Mountains, and then proceeding to an account of those at Brintnell Lake. Com-
munities along the Canol Road have already been published upon (Porsild,
1945), and will be mentioned only in outline or by way of comparison with the
Brintnell Lake area.

PLANT COMMUNITIES IN THE VICINITY OF Fort SIMPSON

Notes on the vegetation around Fort Simpson were made during the second
week of June and over a period of about three weeks in late August and early
September, 1939. The following descriptions of communities are formed from
these notes and from collections made by our party in the same intervals (see
Piate II). They are therefore somewhat incomplete due to the limited time
available, and also because of their seasonal restrictions.

For purposes of discussion the following plant communities or groups of com-
munities can be defined for this area: Flood plain white spruce forest; upland
forest of white spruce, jack pine, aspen and poplar; prairies; river sand bar

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 35

and river bluff associations; and old fields. A few local muskegs occur in the
vicinity, but they are of small extent and not different from muskegs described
elsewhere in the central Mackenzie basin.

Flood Plain Spruce Forests

As noted earlier in this paper, Fort Simpson is on a small island in the
Mackenzie River just below the mouth of the Liard. It is nearest the south-
west bank of the Mackenzie and is connected to the latter at stages of low water
by sand and mud bars. The whole island is part of a low flood plain, portions
of which also appear on the adjacent main river bank. The plain is quite lim-
ited, however, and lies at the foot of a high bluff which here borders the modern
valley of the river (PLATE II). The island is composed largely of silt, most of
which appears to have come from the Liard River. It is highest on the north-
east shore, where an eroding bluff descends to the river. From the top of this
bluff the surface drops off by a series of low terraces to the sand bars above
mentioned. In the lower areas there are many old channels, some of which still
contain sloughs. The channels are roughly parallel to the river and to the long
axis of the island.

It is probable that the entire island and other parts of the flood plain were
covered with forest prior to the founding of the settlement during the first decade
of the 19th century. Extensive clearings have been made for pasture and culti-
vation both on the island and on the adjacent mainland, but remnants of the
forest are stil! abundant.

Primary spp.: Picea glauca var. albertiana, Viburnum edule, Rosa acicularis, wood-
land mosses.

SECONDARY spp.: Equisetum pratense, Equisetum scirpoides, Carex bonangensis, Carex
media, Calypso bulbosa, Habenaria obtusata, Populus Tacamahacca, Salix arbusculoides,
Salix Bebbicna, Alnus tenuifolia, Betula papyrifera var. humilis, Geocaulon lividum,
Mitella nuda, Ribes hudsonianum, Ribes lacustre, Ribes oxyacanthoides, Fragaria vesca
var. americana, Rubus acaulis, Rubus pubescens, Viola renifolia var. Brainerdii, Shepherdia
canadensis, Cornus canadensis, Cornus stolonifera, Pyrola grandiflora var. canadensis,
Castilleja Raupii, Boschniakia rossica, Lonicera glaucescens.

Like most of the flood plain forests in our region, it is difficult to describe this
community except in terms of developmental stages which are now clearly to be
seen on the sand bars. Whether the older forest at higher levels on the island
had the same origin is of course conjectural, because we have no data on the
age of the deposits or on their relation to the advent of forests in the region.
At present the successional stages are nearly identical with many that I have
described (1935) along the lower Peace and Slave Rivers. A forest of balsam
poplars precedes the spruce, and this is in turn preceded by willows and alders
(see below). Intermediate stages are common. At Simpson the whole se-
quence can be seen simply by walking across the island, where the mature spruce
forest is on the older, higher terraces, while the stages are progressively younger
toward the sand bars of the back channel or “sny.” In the poplar woods alders
and willows are the primary shrubs, and mosses have only begun to cover the
ground. Young white spruces come up under the poplars and gradually
replace them.

Upland forests of white spruce and jack pine

Detailed notes on the upland forests west of the river opposite Fort Simpson
could not be made, but the following will be sufficient to characterize them.

36 SARGENTIA [6

They are composed principally of Picea glauca vars. albertiana and Porsildii,
Pinus Banksiana, Populus tremuloides, and occasional mixtures of Populus
Tacamahacca. The commonest shrubs are Rosa acicularis, Salix Bebbiana, and
Viburnum edule. Arctostaphylos Uva-ursi is abundant, as are also Carex
siccata, Bromus Pumpellianus, Aster Lindleyanus, and Hieracium canadense.

Except for scattered muskegs the surface of the upland is sandy and well-
drained, with low sand ridges here and there. The pine forms nearly pure
stands on these sandy hills. Evidence of fire is everywhere to be seen, and the
present composition of the forest is no doubt conditioned by burning. In fact
it strongly resembles a fire-induced type that is seen on most of the uplands of
the central Mackenzie basin (Raup, 1935, 1946). If this is the case the pri-
mary forest on the upland is probably of white spruce. Fires of varying intensity
and over soils of varying dryness have given rise to every possible mixture of
spruce with aspen, poplar, and pine, or to nearly pure stands of pine. Every
indication is that the spruce is finally rehabilitated if left undisturbed for perhaps
75 to 100 years.

Prairies

An outstanding feature of the vegetation in the Simpson area is the occurrence
of small prairie openings in the upland forest west of the river (Prates II, XI).
Some of the openings have been utilized by the people of the trading post as a
source of wild hay, and stock has sometimes been pastured in them. They vary
in size from patches only a few yards across to expanses of 30 acres or more.
The soils are rather loose and sandy in texture, but very dark brown in color.
The surfaces usually have little or no “saucer-form,” though some of the open-
ings have shallow wet depressions or stream channels in them.

The most widely spread community found in the prairies is as follows:

PriMARY spp.: Calamagrostis neglecta, Poa pratensis, Juncus Vaseyi (alternating).

SECONDARY spp.: Botrychium multifidum, Agrostis scabra, Poa palustris, Carex Bebbii,
Salix Bebbiana, Arabis hirsuta var. pyenocarpa, Geum macrophyllum var. perincisum,
Rosa acicularis, Rubus strigosus, Astragalus frigidus var. americanus, Vicia americana,
Epilobium angustifolium, Arctostaphylos Uva-ursi, Stachys palustris var. pilosa, Galium
boreale, Solidago decumbens var. oreophila, Aster Lindleyanus, Antennaria nitida.,

This community makes a thick turf in which the Calamagrostis is easily the
most abundant species, though in a few places it is superseded by Poa pratensis
or Juncus Vaseyi. Its principal variations are a dry phase, in which Arcto-
staphylos is abundant and Equisetum prealtum becomes very common, and a
wet meadow phase. The wet meadow is ‘not uniform, but is composed of
patches of nearly pure stands of Carex rostrata or of dense mats of Carex aurea.
Other characteristic species of the wetter ground are Deschampsia cespitosa,
Phalaris arundinacea, Carex leptalea, Habenaria hyperborea, Polygonum natans
f. Hartwrightit, Ranunculus Macounii, Potentilla norvegica, Mentha canadensis
var. glabrata, and Galium trifidum.

The margins of the prairies are apparently being invaded by the neighboring
woods, though the rate at which this is occurring could not be estimated. Young
jack pines, aspens, and balsam poplars are pushing in from the borders, the
aspens most vigorously.

These prairies have all the outward aspects of those in the Wood Buffalo Park
(Raup, 1935). Superficial examination suggests that they occur on more

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 37

porous soils, however, than most of the Park openings. Also their flora is much
smaller, and their primary species are somewhat different. Poa pratensis prai-
ries were seen at the eastern base of the Caribou Mountains, but Calamagrostis
neglecta was not found there. This may be due in part to seasonal aspects.
Juncus Vaseyi is common enough in the Park area, but I have not seen it occu-
pying any other than a secondary position in the communities there.

There is no evidence that the prairies are not “natural.” Continued grazing
and repeated burning can sometimes cause grassy openings in a forested country
(Raup, 1945) ; but those I have seen were obviously related to long-established
Indian camp sites, none of which were seen in this neighborhood. Neither were
there any old stumps or disturbances of the soil surface due to the wind throw
of trees.

There are no doubt other prairie areas in the vicinity which would throw
further light upon the composition and possibly upon the history of this type of
vegetation. Openings north of Providence have long been known, and prairie-
like vegetation has been seen by travelers south of the Mackenzie and west of
Great Slave Lake. Until further observations have been made in this little-
known region it is futile to attempt any generalizations with regard to the flora
and community structure of the grasslands.

Plants of sand bars and river banks

From a phytogeographic standpoint the river banks and sand bars at Simpson
hold perhaps the most interesting part of the local flora. This is due not only
to the variety and geographic affinities of the plants, but also to the fact that
these situations contain some of the developmental stages leading to the flood
plain forests.

All of the shores present unstable and hazardous habitats for plants. All but
the higher parts of the bluffs are subject to flooding in spring, while at lower
levels there is a progressively shorter growing season determined by the gradual
ebb of the river during the summer. All the surfaces are subject to erosion by
the river, either in the season of high water or at the time of ice flow in the
spring. Late in the season the higher sand bars dry out and are subject to
wind erosion, even forming into small dunes. To withstand these rigors plants
must, in addition to withstanding the subarctic climate, be perennials with root
or rhizome systems of great flexibility, or annuals whose disseminules are
extremely hardy.

The plant associations here are nearly all open. The individual plants are
often widely scattered, with most of the ground surface unoccupied. It is im-
possible to designate well-defined communities except on the older sand bars,
although certain species are more abundant than others in specific habitats. A
general list of species noted on the sand bars is as follows:

Equisetum arvense,-Equisetum fluviatile, Agropyron latiglume var. pilosiglume, Agro-
pyron trachycaulum var. typicum, Beckmannia Sysigachne, Cinna latifolia, Deschampsia
cespitosa, Eleocharis palustris, Juncus alpinus var. rariflorus, Juncus balticus, Juncus no-
dosus, Salix interior var. pedicellata, Salix lasiandra var. recomponens, Salix macken-
sieana, Polygonum aviculare, Polygonum lapathifolium var. salicifolium, Rumex maritimus
var. fueginus, Corispermum hyssopifolium, Rosa Woodsii, Astragalus yukonis, Hedysarum
alpinum var. americanum, Sium suave, Plantago major, Solidago lepida var. elongata,
Aster falcatus, Aster junceus, Aster sibiricus, Erigeron philadelphicus, Artemisia biennis,
Artemisia Tilesii.

38 SARGENTIA [6

Damp sand at the borders of lagoons and sluggish channels has conspicuous
stands of Juncus nodosus, Juncus alpinus var., Juncus balticus, and Equisetum
arvense. Rumex maritimus var. fueginus, Beckmannia Syzigachne, Polygonum
lapathifolium var. salicifolium, and Sium suave also appear to reach greatest
abundance in these wetter areas.

The higher sand ridges and terraces are either colonized directly by willows,
Salix interior var. pedicellata, or have an intermediate association composed
principally of Equisetum fluviatile with a mixture of Eleocharis palustris. The
legumes, asters, and roses are found chiefly among the willows. Of the latter
the sand bar willow, Salix interior var., is the pioneer species. On older bars
it has an admixture of Salix mackenzieana and Salix lasiandra var. It is in
these willow thickets that Populus Tacamahacca and Alnus tenuifolia first ap-
pear as forerunners of the flood plain forests.

The most abundant species on windblown sand is Corispermum hyssopifolium.
In many places it is the only plant that inhabits this unstable habitat.

In an earlier part of the paper I have already described one of the more re-
markable physiographic features of the upper Mackenzie—the boulder pave-
ments which form the banks of the river for many miles. The rocks are of
varying size, from cobblestones to huge boulders, most of them with glacial
facets, but all much rounded and waterworn. They lie in a scanty matrix of
sand and silt which is usually damp at lower levels but may become quite dry
in late summer. As in the vegetation of the sand bars, plants are scattered
loosely, usually without semblance of arrangement into communities. In any
one locality the number of species is exceedingly small, but when taken together
from a mile or so of riverbank they show a surprising variety.

Triglochin palustre, Agropyron trachycaulum var. typicum, Calamagrostis inexpansa
var. brevior, Deschampsia cespitosa, Poa glauca, Eleocharis palustris, Juncus balticus,
Juncus nodosus, Allium Schoenoprasum var. sibiricum, Sisyrinchium angustifolium, Salix
glauca, Rumex maritimus var. fueginus, Chenopodium album, Corydalis aurea, Rosa
W oodsii, Astragalus alpinus, Astragalus tenellus, Astragalus yukonis, Hedysarum Macken-
sti, Linum Lewisii, Epilobium glandulosum var. adenocaulon, Apocynum sibiricum, Mentha
canadensis var. glabrata, Stachys palustris var. pilosa, Rhinanthus groenlandicus, V eronica
peregrina var. xalapensis, Solidago graminifolia var. major, Solidago lepida var. elongata,
Aster angustus, Aster falcatus, Aster junceus, Helenium autumnale, Achillea Millefolium,
Achillea sibirica.

This assemblage resembles somewhat that of the sand bars, but many of its
plants are drawn from more mesophytic habitats. The prevalence of heavy
boulders undoubtedly renders it far more reliable as a plant habitat than the
open sand. The above list can only be a partial one, for it is based upon one
set of observations in a limited area. Further looking along the river banks
would no doubt greatly increase it.

Steep river bluffs, composed of till, loose shale, or undercut banks of silt,
have an exceedingly scanty flora in the neighborhood of Simpson. A few
plants, however, were found only in these places: Braya humilis, Astragalus
aboriginum, Elaeagnus commutata, Crepis elegans. The only willow noted on
the bluffs was Salix arbusculoides. Acer negundo is growing on the steep bank
of the river at the village of Simpson, probably planted there by the early traders.

Plants of fields and clearings

An attempt was made to record the species growing as “weeds” in and around
Simpson. There are several pastures and cultivated areas in the vicinity, in

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 39

various conditions of tendance, and many of the roadways and dooryards are
grazed and fertilized by horses and cattle. A great variety of garden vegetables
and flowers have been grown or attempted through the years, so that there have
been abundant opportunities for the advent of weeds and escapes. Stock feed
brought down by boat has added to these sources. The following species were
noted in clearings and old fields:

Hierochloé odorata, Hordeum jubatum, Poa glauca, Poa pratensis, Carex aenea, Carex
praticola, Juncus bufonius, Sisyrinchium angustifolium, Polygonum aviculare, Rumex
mexicanus, Chenopodium capitatum, *Amaranthus retroflexus, Silene Mensiesii, Stellaria
calycantha, Stellaria longifolia, Stellaria longipes, *Stellaria media, Anemone canadensis,
Anemone multifida var. hudsoniana, Thalictrum occidentale, Arabis divaricarpa, Arabis
hirsuta var. pycnocarpa, *Brassica Kaber var. pinnatifida, *Capsella Bursa-pastoris, Draba
nemorosa, *Neslia paniculata, Fragaria glauca, Potentilla Anserina, Potentilla multifida,
Potentilla norvegica, Potentilla pennsylvanica, Viola adunca, Androsace septentrionalts,
Collomia linearis, Lappula Redowskii var. occidentalis, Stachys palustris var. pilosa, Galium
boreale, Aster junceus, Erigeron philadelphicus, Helianthus annuus, Artemisia Diennis,
Petasites sagittatus, *Sonchus oleraceus, Lactuca pulchella.

Perhaps the most striking feature of this group is the small number of intro-
duced European weeds (*)—only six species, or about 13.6%. None of the
six, with the possible exception of Capsella, are abundant or become pests in
the gardens. As in other parts of the central Mackenzie basin, the most numer-
ous dooryard weeds are Hordeum jubatum, Polygonum aviculare, Chenopodium
capitatum, Capsella Bursa-pastoris, Potentilla Anserina, and Potentilla multi-
fida. None of the 44 plants listed above have become serious garden pests.

VEGETATION OF THE HoRN AND FRANKLIN MOUNTAINS

So far as I am aware no botanist has ever visited the Horn Mountain plateau
or those parts of the Franklin Mountains that fall within our area. Botanical
knowledge of the Eagle Mountain Plateau is practically non-existent.

I can do no better than to quote directly the descriptions of the Horn Moun-
tains published by the geologist E. J. Whittaker in 1922 and 1923. In 1921
Whittaker and his party ascended the Horn River and climbed to the summit
of the plateau at its southeastern end. He writes, “These hills rise gradually
to a height of about 1,200 feet above the plain in a distance of about 6 miles.
At this height the slope is truncated by a level strip averaging 1144 miles wide,
and then follows an abrupt rise of 400 to 500 feet to the top of the range, which
is a plateau . . . , almost treeless, covered with caribou moss, and dotted with
small ponds. One of these ponds is worthy of being called a lake, being about
114 miles long by 34 mile wide.

“Only the southeastern end of the range was explored. It averages 8 miles
in width at this end, but apparently widens out farther to the northwest.

“The trees on the bare plateau of the range show plainly the effect of the
prevailing winds. The limbs all point to the southeast, and the northwest sides
of the trees are scarred and limbless. Mink, Birch, and Fawn lakes have willowy
and marshy shores on the northwest sides, but the southeast sides are covered
with boulders and an occasional sand or gravel beach. This is evidence of heavy
wave action on the southeast sides forming a beach in each case” (1922, p. 47B;

also Plate IA).

40 SARGENTIA [6

Whittaker again ascended the plateau in 1922, this time by way of the Rabbit-
skin River which enters the Mackenzie about 22 miles above Simpson. “Horn
mountains in this area rise rather abruptly from this level plain to a height of
about 1500 feet. The ascent is made in a gentle slope of about 300 feet in half
a mile and an abrupt rise of 1200 feet in about 300 yards. From the crest the
land rises very slowly for about a mile, then merges into the plateau which is
covered with stunted spruce and, more rarely, poplar” (1923, p. 95B).

The botany of the Franklin Range is almost completely unknown. It is pre-
sumed to be largely timbered, but I have been unable to find any description of
the amount or kind of forest. Judging by elevations given by geologists, there
must be some areas of alpine vegetation.

The Ebbutt Hills, according to M. Y. Williams (1922, p. 57B), are plateau-
like on top, and “covered with muskeg, black spruce, etc.”

Some of the plants collected by C. L. Crickmay may have come from the
Franklin Range. Many of them are alpine species that could hardly have been
found in the Mackenzie lowland. Crickmay’s notes read, however, “Mackenzie
River, Wrigley to Blackwater R.,” and “Blackwater R. to Norman.”

PLANT COMMUNITIES IN THE BRINTNELL LAKE AREA

The most prominent plant communities in the vicinity of Brintnell Lake in
the Mackenzie Mountains are the forests of the lower mountain slopes, a sub-
alpine scrub zone made up principally of willows and dwarf birch on the middle
slopes, and a varied group of tundra communities at higher levels. Timber line
is between 3800 and 5000 feet above sea-level; or 1200 to 2400 feet above the lake.
The subalpine scrub varies greatly in vertical width, but in places covers nearly
1000 feet. Timber lines on north- and south-facing slopes in some places differ
but little in elevation, although the character of the forests is quite different.

As stated elsewhere, most of the shores of the lake are precipitous, so that
space for the development of communities other than those just mentioned is
limited. A few small muskegs have developed, chiefly at the eastern and western
ends of the lake, while in the valley of the glacial stream that enters from the
west are deposits of mud upon which a marsh type of vegetation appears. The
- lowland along this stream also contains some gravel and sand bars that have
produced characteristic communities. Large fans composed of stones and gravel
occur at several places around the lake, to produce the nearest approach to
beaches that the area affords.

The descriptions presented below are derived from the amalgamation of two
sets of field notes. First are the records of collected specimens upon which
rather copious habitat and locality data were kept. Since every effort was made
to collect the entire flora of the area, these data when collated give a fairly com-
prehensive record of the existing habitats. Second, brief descriptions of the
more important communities were prepared in the field, with lists of the species
observed in given areas and photographs to illustrate typical sites. However,
in spite of this attempt to form a well-rounded description of the plant life, I
make no pretense whatever to completeness. The structure of the vegetation in
subarctic America is notoriously fickle, and that of the Brintnell Lake region is
no exception. New types of communities, or curious variations of the old ones,
are sure to turn up with further examination of the country.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 41

The following is an outline of the types of vegetation at Brintnell Lake.

Forests.
Lower slopes and lake shores—Picea glauca vars.
Higher south-facing slopes—Picea glauca vars.
North-facing slopes—Picea mariana, Picea glauca var.
Muskegs——Picea mariana, Larix laricina var.
Communities induced by fire.
SUBALPINE SCRUB.
ALPINE COMMUNITIES.
Damp meadows.
Stabilized scree slopes.
Slide-rock.
Ledges and crevices.
VEGETATION OF ALLUVIAL MUD AND GRAVEL DEPOSITS.
VEGETATION OF STONE AND GRAVEL LAKE BEACHES.

Forests

The most mesophytic forest at Brintnell Lake is situated on the lower slopes
of the mountains, extending only a few hundred feet above the level of the lake
(Pirate VII). Its structure and composition are as follows:

PrrMary spp.: Trees: Picea glauca var. albertiana. Shrubs: Betula papyrifera var.
commutata, Betula papyrifera var. humilis, Alnus crispa, Ribes triste, Rosa acicularis,
Shepherdia canadensis, Viburnum edule. Ground: Hypnum spp., Peltigera aphthosa,
Vaccinium Vitis-Idaea var. minus.

SECONDARY sSPp.: Cystopteris fragilis, Dryopteris fragrans, Dryopteris Linnaeana, Lyco-
podium annotinum var. pungens, Lycopodium complanatum, Corallorrhiza trifida, Cypri-
pedium passerinum, Habenaria hyperborea, Salix arbusculoides, Salix Scouleriana var.
coetanea, Geocaulon lividum, Actaea rubra, Ribes glandulosum, Ribes hudsonianum, Poten-
tilla fruticosa, Rubus acaulis, Lupinus arcticus, Cornus canadensis, Moneses unifiora, Pyrola
asarifolia var. incarnata, Pyrola grandiflora var. canadensis, Pyrola secunda, Pyrola virens,
Arctostaphylos rubra, Linnaea borealis var. americana, Solidago multiradiata, Arnica al-
pina, Senecio lugens.

The spruces commonly attain breast-height diameters of 2-3 feet and heights
of 60-100 feet. The mat of mosses and duff in the oldest forest observed was
at least 15 inches thick. The mineral substratum is usually very rough, com-
posed principally of the larger talus rocks that have reached the lower slopes
and the coarser materials of the upper parts of the alluvial fans. On sites that
have not been altered by fire the heavy mat of mosses completely covers all but
the largest boulders. In spite of their excellent diameter and height growth,
most of the trees on the slopes have much twisted grain and a large amount of
compression wood on their down-hill sides near the base. Nearly all are bent
to their vertical positions from near the base, indicating that they have suffered
from land-slip, or snow-slides, or both. The shrub layer is not dense, and
offers but little hindrance to travel through the woods. ‘The principal difficulties
of travel are the rough, bouldery substratum and the tangle of fallen logs that
are everywhere in the way.

There is but little variation in this rich forest around the lake. It is modified,
however, on drier sites and on the flood plain of the glacial stream west of the
lake. Picea glauca var. albertiana remains a primary species throughout, how-
ever. More xerophytic phases of the white spruce forest are found on a se-
ries of rock and gravel ridges east of the lake, on the warm south-facing slopes

42 SARGENTIA [6

below timber line north of the lake, and on the more unstable portions of the
gravel fans.

East of the lake the stream that drains it soon enters a steep rapid which lies
at the bottom of a picturesque winding gorge. The gorge is cut in shaly rocks
whose dip is here nearly vertical. On the north side of the stream is a series
of rocky and gravelly ridges separated by gently sloping plains covered with dry
open woods. Occasional depressions are filled with muskegs. The open woods
are composed of the following association.

Primary spp.: Picea glauca var. albertiana, Betula papyrifera var. commutata, Hypnum
spp., Cladonia spp., Cetraria spp.

SECONDARY spp.: Lycopodium complanatum, Festuca altaica, Betula glandulosa, Betula
occidentalis, Rosa acicularis, Lupinus arcticus. .

This forest is rather park-like, with the trees standing in a dense mat of
fruticose lichens and woodland mosses. Although the list of secondary species
given above is probably not complete, the total flora is quite scanty and scattered.
As will be noted elsewhere in this paper, the white birches are extremely difficult
to define specifically. Many of them combine the morphological characters of
varieties commutata and humilis.

The higher south-facing mountain slopes north of the lake, as well as gravel
fans on that side, are notable for the addition of another variety of the white
spruce, P. glauca var. Porsildii, described in this paper. The smooth, balsam-
like bark of this species and its more broadly pyramidal form make it easily
distinguishable in stands otherwise composed of var. albertiana. It was rarely
seen on the south side of the lake.

As one ascends the steep mountain slopes north of the lake the forest gradually
becomes more open, the trees smaller, and the ground cover composed of grasses
and other herbs in place of the heavy moss mat of the lower slopes (PLATE
VIIT). At the same time the shrub layer changes in character. The sub-
stratum remains exceedingly rough and rocky, with its surface variously broken
by gullies and ridges of glacial till. Most of the slope is blanketed with bouldery
till which in places is arranged in ridges whose summits slope downward to the
east. These appear to be lateral moraines deposited by the glacier which last
filled the valley. Streams such as Frost Creek have cut gorges in the till, paving
their beds with huge boulders that have been washed out of it. The channels
of these streams evidently have shifted frequently, as shown by the abandoned
gullies on either side.

The forests at 1000-1500 feet above the lake may be characterized as follows:

Primary spp.: Picea glauca var. albertiana, Picea glauca var. Porsildii, Juniperus com-
munis var. montana, Arctostaphylos Uva-ursi, Festuca altaica.

SECONDARY spp.: Lycopodium clavatum var. monostachyon, Lycopodium complanatum,
Agropyron trachycaulum var. novae-angliae, Agrostis scabra, Calamagrostis purpurascens,
Festuca brachyphylla, Poa Buckleyana, Poa nemoralis, Trisetum Spicatum var. Maidenii,
Carex atrata, Smilacina stellata, Salix arbusculoides, Salix Bebbiana, Salix glauca var.
berstipula, Salix Scouleriana var. coetanea, Alnus crispa, Betula occidentalis, Betula glan-
dulosa, Betula papyrifera var. commutata, Arenaria lateriflora, Arenaria verna var. pubes-
cens, Aconitum delphinifolium, Anemone Richardsonii, Aquilegia brevistyla, Arabis Drum-
mondti, Arabis Holboellii, Saxifraga tricuspidata, Amelanchier florida, Dryas Drummondit,
Potentilla fruticosa, Rubus strigosus, Lupinus arcticus, Shepherdia canadensis, Epilobium
angustifolium, Cornus canadensis, Vaccinium uliginosum, Vaccinium Vitis-Idaea_ var.
minus, Mertensia paniculata, Pedicularis labradorica, Linnaea borealis var. americana,
Viburnum edule, Solidago mutltiradiata, Antennaria isolepis, Antennaria sp., Artemisia
arctica, lichens (Peltigera, Cladonia).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 43

This forest is park-like, with the spruces branched from near the base.
Where the substratum is fairly well-stabilized the openings among the trees are
grassed over with the large tufts of Festuca altaica, interspersed with clumps
of Juniperus and mats of Arctostaphylos. Steeper slopes of gravel or loose
shale, such as occur on the faces of the morainic ridges, lack most of the grasses
and have only scattered Juniperus and Arctostaphylos with a few herbaceous
species that manage to gain a precarious foothold. It will be seen from the list
of secondary species that this high slope forest combines elements of the floras
of the rich lowland woods and the subalpine scrub.

Still higher on the mountainside, nearer timber line, the flora as a whole re-
mains the same, but a somewhat different arrangement of primary species ap-
pears. Thickets of Betula glandulosa, Alnus crispa, and Salix glauca var. per-
stipula begin to dominate the scene among the more scattered spruces, while on
the loose soil of very steep slopes Saxifraga tricuspidata, Vaccinium V itis-Idaea
var. minus, and Vaccinium uliginosum form low, patchy thickets among the
juniper bushes.

There is abundant evidence of fire on the mountainside in the neighborhood
of Frost Creek. Evidence of at least two fires could be discerned, the later one
probably less than 50 years ago. This later fire appears to have started near
the western end of the lake and to have swept eastward and upward so as to
consume all of the forest to timber line. Its eastern limit was not determined
with accuracy, although from the appearance of the forest as seen from the lake
it extended only a mile or so in that direction. In much older spruce forest
near the shore about a mile east of Frost Creek some carbonized remains were
found, indicating a much earlier fire. Other old forest in the vicinity gave no
evidence of ever having been burned. It is difficult to see how these fires were
started, unless they were ignited by lightning. Evidence of the presence of
trappers or prospectors is all very recent, probably dating within the last 25
years, and nowhere around the lake is there any sign of Indian habitation. No
old camp sites were found, and no trails other than occasional game trails.

The principal vegetational effects of the fires have been the expansion of the
white birches, aspen, and balsam poplar. In many places these have become
primary species in the burn forests (PLATE VII). At the same time the larger
willows have become primary in the shrub layer, notably Salix S couleriana var.
coetanea and S. arbusculoides. These often form single-stemmed trees as much
as 6 inches in diameter. On the ground there is a tangle of partly burned and
slowly decaying logs. The rehabilitation of the spruce forest from this inter-
mediate stage appears to be direct. There is little or no evidence that the fire
lowered the tree line, but the size of the stumps of fire-killed trees near timber
line indicates that they were much larger than those now growing there. The
present trees reach a diameter of about a foot, while the stumps are often
twice that.

All of the great quantity of down timber, however, cannot be attributed to
the fire, for there appears to be a normally high mortality rate among the trees
on the steep slopes. This appears to be due both to land-slip and snow-slides,
the latter probably by far the more important. Views of the steep south-facing
slopes from the mountains across the lake show what look like gullies, some of
them extending all the way from timber line to lake level (PLare III). There
is a complex interwoven system of these “gullies,” some of which are narrow,
while others are a hundred feet or more wide. Examination on the ground

44 SARGENTIA [6

shows that they are not gullies in the ordinary sense, but great furrows in the
forest, now in part overgrown with a dense tangle of willows and other shrubs
of the bordering woods together with young spruce, aspen, and balsam poplar.
At the foot of each furrow is an immense “jackstraw” tangle of dead logs.
Neither the debris at the bottom of the slope or in the furrows themselves involves
much mineral material, indicating that the damage has been done principally by
sliding snow and ice.

All the trees and shrubs in the furrows are bent downward from the base and
then curve upward again, indicating the repeated effects of sliding snow. The
prevalence of this form on all the steeper slopes suggests that the damage done
by snow is general, though not always concentrated in channels. It probably
accounts for a large part of the fallen timber.

At first glance the relative openness of the snow-slide furrows suggests that
they would be good routes for climbing the mountain. Our party tried one but
was very quickly disillusioned. The mass of debris on the ground and the
matted tangle of downward-pointing branches on the distorted shrubs make
climbing far more exhausting than it is in the neighboring forest.

The snowslide channels are seen only on very steep south-facing slopes. It
is presumed that they are caused by the early loosening of snow during spring
thaws at and above timber line. Insolation would no doubt be more rapid here
than under cover of forests at lower levels.

The great fans of gravel and stones that occur around the lake (Piate IV)
apparently are formed by torrential floods. The size of trees growing in the
more active channels near the shore indicates that such floods are not frequent.
None occurred during the season of 1939 that caused any noticeable displace-
ment of materials. In what part of the open season the floods come is not
known, but it is assumed that they come in late spring or early summer. Our
field party of 1943 experienced such a flood in the Summit Pass area along the
Alaska Highway. It came early in July, following a hard rain which lasted for
about 36 hours. Boulders several feet in diameter were moved by this flood,
and immense quantities of rock debris were shifted in the mountain gullies and
alluvial fans. Here again there was good evidence that at least a quarter of a
century had elapsed since a similar flood had occurred.

The surfaces of the fans are covered with gravel, stones, and boulders, the
finer of which are nearest the lake. The general contours are of varying steep-
ness depending upon the size of the materials. The slope of the one at the mouth
of Frost Creek is rather gentle, about 200 feet in 1600; but some of those on
the south shore, composed of large rocks from the steep north slopes of Colonel
Mountain, have a much larger gradient, about 200 feet in 1000. The contours
are broken by interlocking abandoned channels, some of which will no doubt
become active again with the next flood.

The vegetation on the fans is an open forest of spruce, balsam poplar and
large willows, interspersed with areas of nearly barren gravel or dense thickets
of shrubs and young trees (PLare XII). The flatter parts which receive suf-
ficient drainage, usually at the sides of the main channels, sometimes harbor
muskegs. The drier parts proved to be of considerable floristic interest because
of the heterogeneous mixture of species growing on them. Their flora is drawn
not only from the woods and shores around the lake, but also from the alpine
vegetation far above. The following list contains species found in the open
woods and thickets, and is made up from observations and collections on several

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 45

different fans. For a more complete account of the flora of these sites the list
should be supplemented with one given below in the discussion of alpine plants
at lake level.

Primary spp.: Picea glauca var. albertiana, Picea glauca var. Porsildii, Betula papy-
rifera var. humilis, Betula papyrifera var. commutata, Alnus crispa, Salix planifolia, Salix
alaxensis var. longistylis, Viburnum Gi. Shepherdia canadensis, Rosa acicularis.

SECONDARY SPP.: Agropyron latiglume, Agropyron trachycaulum var. novae-angliae,
Arctagrostis arundinacea, Calamagrostis canadensis var. robusta, Festuca brachyphylla,
Poa alpina, Poa arctica, Poa Brintnellii, Poa glauca, Carex atrata, Salix Bebbiana, Salix
commutata, Salix Scouleriana var. coetanea, Stellaria calycantha, Anemone parviflora,
Arabis Holboellii, Cardamine bellidifolia, Draba McCallae, Saxifraga radiata, Saxifraga
tricuspidata, Geum macrophyllum var. perincisum, Rubus strigosus, Sibbaldia procumbens,
Epilobium angustifolium, Epilobium lactiflorum, Pedicularis sudetica, Campanula lasio-
carpa, Erigeron angulosus var. kamtschaticus, Artemisia Tilesii, Taraxacum alaskanum,
Taraxacum lacerum, Taraxacum ceratophorum,

On more undisturbed parts of the fans this open forest merges imperceptibly
with the rich spruce woods already described. A moss mat appears on the
ground, anc woodland species of shrubs predominate to the exclusion of others.
As previously noted, the smooth-barked spruce, P. glauca var. Porsildii, is a
primary species only on the north side of the lake. Its occurrence on the south
side appeared to be limited to the lowermost parts of some of the fans.

Still another type of white spruce forest has developed on the mud, sand, and
gravel flats west of Brintnell Lake. It is rather open and park-like, with a mat
of mosses that is thin and much interrupted.

Primary spp.: Picea glauca var. albertiana, Picea glauca var. Porsildti, Populus Taca-
mahacca, Alnus tenuifolia, Equisetum palustre, Hypnum spp.

SECONDARY SPP.: Arctagrostis arundinacea, Calamagrostis canadensis var. robusta, Poa
pratensis, Cerastium Beeringianum, Aconitum delphinifolium, Parnassia palustris var.
neogaea, Ribes lacustre, Ribes oxyacanthoides, Rosa acicularis, Hedysarum alpinum var.
americanum, Oxytropis hyperborea, Oxytropis foliolosa, Moneses uniflora, Pyrola asari-
folia var. incarnata, Pyrola secunda, Arctostaphylos rubra, Mertensia paniculata, Solidago
multiradiata.

This forest resembles flood plain types in other parts of the Mackenzie basin.
It is of interest that Equisetum palustre here forms dense green swards on the
forest floor, much as FE. pratense does along the Slave, Peace, and Athabaska
Rivers. Possible developmental sequences leading’ to the spruce forests west
of the lake will be discussed in another place.

Most of the forests on the north-facing slopes fronting Brintnell Lake are
radically different from those just described. White spruce timber similar to
the more mesophytic type already noted occurs south of the lake, but is confined
to the more stable portions of the alluvial fans, reaching only about 200 feet
above the shore. With one exception, which occurs near timber line, the forests
of the higher slopes are of black spruce, Picea mariana.

The northerly slope of Colonel Mountain consists of a series of great gullies
separated by sharp “hogbacks” which extend nearly to the summit (about 7300
feet above sea level) (PLATE IV). The hogbacks are the products of an erosion
interval that extended through at least one interglacial period. Their lower
portions were removed to an elevation of about 4300 feet by the last advance
of the ice, so that they are now terminated by great triangular facets that reach
about 1700 feet above the lake. The facets have a gradient of about 34°, and are
clothed with forest to about 4000 feet, or about 1400 feet above the lake.

46 SARGENTIA [6

The steep slopes are rocky, but they are covered for the most part by a thick
mat of fruticose lichens and woodland mosses. This mat is often a foot or more
in depth and is nearly always damp. It thins out gradually toward timber line
so that the underlying rocks begin to be exposed. The forest is an open one
even at lake level, but it becomes still more so at higher elevations ( PLATE
VIII). The general aspect of the north-fagng mountainsides is strikingly dif-
ferent from that of the opposite slopes, due tothe peculiar greens of the fruticose
lichens. These greens are particularly noticeable, of course, in wet weather,
and contrast sharply with the higher grassy slopes north of the lake.

There is a certain amount of change in vegetation from bottom to top of the
forested zone, but except for a narrow zone at the top the whole is sufficiently
uniform to justify treating it as a unit. The following list is a general one,
therefore, and will be followed by notes on the principal variations.

PRIMARY spp.: Trees: Picea mariana. Shrubs: Salix glauca var. perstipula, Alnus
crispa, Ledum groenlandicum, Vaccinium uliginosum. Ground: Cladonia, Cetraria, HA yp-
num spp., Arctostaphylos rubra, Equisetum scirpoides, Andromeda Polifolia.

SECONDARY spp.: Arctagrostis arundinacea, Festuca altaica, Carex scirpoidea, Carex
vaginata, Tofieldia palustris, Corallorrhisa trifida, Goodyera repens var. ophioides, Habe-
naria obtusata, Salix alaxensis, Salix reticulata, Betula papyrifera var. humilis, Geocaulon
lividum, Oxyria digyna, Silene acaulis var. exscapa, Stellaria longipes var. Edwardsii,
Saxifraga tricuspidata, Dryas integrifolia, Rosa acicularis var. cucurbiformis, Astragalus
frigidus var. littoralis, Hedysarum alpinum var. americanum, Lupinus arcticus, Epilobium
latifolium, Pyrola secunda, Cassiope tetragona, Vaccinium microcarpum, Vaccinium Vitis-
Idaea var. minus, Mertensia paniculata, Pedicularis capitata, Pedicularis labradorica,
Boschniakia rossica, Pinguicula villosa.

The primary species just noted are characteristic up to 500-600 feet above
the lake. Above this general level Salix Richardsonii and Betula glandulosa are
added to those of the shrub layer, and Dryas integrifolia becomes an important
component of the ground mat. The white birches disappear almost entirely from
the association. Otherwise the aspect is much the same except that the black
spruces are more dwarfed and more widely spaced.

The upper 200-300 feet of the forest has a superficial resemblance to that
below, but closer examination shows a striking change. The black spruce is
replaced by Picea glauca var. albertiana, while the shrub and ground floras show
a transition to the subalpine and alpine vegetation.

PRIMARY spp.: Trees: Picea glauca var. albertiana. Shrubs: Salix glauca var. per
stipula, Salix Richardsonii, Betula glandulosa. Ground: Cladonia, Cetraria, Hypnum:
spp., Cassiope tetragona, Vaccinium uliginosum, Dryas integrifolia, Carex Scirpoidea.

_Ledum groenlandicum drops to a secondary position in the community, while
a few species such as Equisetum scirpoides, Corallorrhiza trifida, Goodyera
repens var. ophioides, Habenaria obtusata, Rosa acicularis var. cucurbiformis,
Pyrola secunda, Vaccinium microcarpum, and Pinguicula villosa become very
rare or drop out altogether. At the same time some additions are made: Shep-
herdia canadensis, Oxytropis ixodes, Empetrum nigrum, and Abies lasiocarpa.
This is the timber line community on the north-facing slopes. It gradually gives
way to a subalpine scrub or to some form of.alpine tundra, depending upon
slope, exposure, or substratum. It is broken here and there by exposed cliffs
of shale on which there is a ledge and crevice flora (see below), or by gullies
in which the willows and alders make dense tangles of shrubbery.

We saw no evidence of fire in any part of the communities on the north-facing
slopes. Consequently I am unable to suggest what its effects would be.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 47

Muskegs were observed on the low ground at both ends of Brintnell Lake,
and on various local areas of small size on the mountain slopes and lake shores.
The following is a general account of their structure and flora drawn from all
these sources:

Primary spp.: Picea mariana, Larix laricina var. alaskana, Ledum groenlandicum,
Sphagnum sp.

SECONDARY SPP.: Equisetum scirpoides, Equisetum sylvaticum var. pauciramosum, Equi-
setum variegatum, Selaginella selaginoides, Arctagrostis arundinacea, Calamagrostis cana-
densis var. robusta, Poa pratensis, Eriophorum brachyantherum, Carex aurea, Carex
capillaris, Carex concinna, Carex gynocrates, Carex leptalea, Carex loliacea, Carex media,
Carex membranacea, Carex scirpoidea, Carex vaginata, Juncus castaneus, Tofieldia palus-
tris, Corallorrhiza trifida, Cypripedium passerinum, Habenaria hyperborea, Habenaria
obtusata, Listera borealis, Salix Barclayi, Salix commutata, Salix glauca var. perstipula,
Salix myrtillifolia, Salix reticulata, Salix Richardsonii, Alnus crispa, Alnus tenuifolia,
Betula glandulosa, Arenaria lateriflora, Anemone parviflora, Ranunculus hyperboreus,
Ranunculus lapponicus, Cardamine pratensis, Rorippa islandica var. microcarpa, Chryso-
splenium tetrandrum, Parnassia Kotzebuei, Parnassia palustris var. neogaea, Ribes hud-
sonianum, Geum macrophyllum var. perincisum, Potentilla fruticosa, Rubus Chamaemorus,
Rubus acaulis, Hedysarum alpinum var. americanum, Oxytropis ixodes, Empetrum nigrum,
Epilobium davuricum, Epilobium latifolium, Andromeda Polifolia, Arctostaphylos rubra,
Cassiope tetragona, Vaccinium microcarpum, Vaccinium uliginosum, Gentiana propinqua,
Pedicularis capitata, Pedicularis sudetica, Boschniakia rossica, Pinguicula villosa, Valeriana
septentrionalis, Aster sibiricus, Erigeron elatus, Antennaria monocephala, Senecio inde-
corus, Senecio lugens, Senecio pauperculus, Taraxacum ceratophorum, Taraxacum lacerum,
Taraxacum lapponicum.

Pools and shores in the muskegs have a few floating or emergent aquatics
such as Hippuris montana, Ranunculus Gmelini var. Purshii, Carex aquatilis,
Carex physocarpa, Carex diandra, and Juncus balticus. Aquatic communities
are, however, very poorly represented in or around Brintnell Lake. No species
of Potamogeton, Nuphar, Myriophyllum, Ceratophyllum, or S parganium were
seen.

A cursory examination of the above list will show at once that the bulk of the
muskeg flora is typical of this habitat throughout most of the northwest. A few
species, such as Carex membranacea, Juncus castaneus, Salix reticulata, Salix
Richardsonii, Parnassia Kotzebuei, Cassiope tetragona, and Gentiana propinqua,
have a strong tundra affinity ; but most are of wide range in the subarctic forests
of the continent. No raised bogs were seen, but on gentle, well-watered slopes
the muskegs have a tendency, as in so many parts of the north, to spread upward.
Some modifications of water levels and muskeg communities were seen along a
sluggish tributary stream in the flats west of the lake, but no detailed studies of
them were made. The changes were caused by beaver dams,

Subalpine scrub communities

A shrub zone, or subalpine scrub, is not well defined on all the mountain
slopes at Brintnell Lake. It is best seen on Red and Terrace Mountains north
of the lake, though even here it varies greatly in prominence and vertical width
(Pate III). Willows and dwarf birch make up most of it. On the steep
north slopes of the glaciated facets of Colonel Mt. and on the hogbacks above
them, it can scarcely be discerned at all, but on the flanks of the hogbacks there
is usually a narrow band of shrubs which widens out somewhat in the inter-
vening valleys. Here it occupies a strip probably less than 100 feet wide in
vertical dimension.

48 SARGENTIA [6

The lip of the hanging valley from which Frost Creek falls into the main
valley of the lake is at about 3800 feet above sea-level, or about 1200 feet above
the lake. The upper valley of Frost Creek is U-shaped below 5000 feet and
evidently contained a tributary glacier at one time. A well-defined terrace of
till appears on its flanks at about 5000 feet. Timber line on the south-facing
slopes seems to be modified by the presence of the hanging valley. On Red and
Terrace Mountains east and west of the Frost Creek valley it rises to nearly
5000 feet, but within the hanging valley itself there are only a few scattered and
stunted trees above 4000 feet. Above timber on the south slopes there is only
a narrow zone of scrub, which disappears completely on the steeper areas. This
zone is greatly expanded in the hanging valley. It covers most of the valley
bottom and the lateral slopes up to the tops of the morainic terraces, or to about
5000 feet. It thus achieves a vertical width of about 1000 feet.

Floristically the subalpine scrub is a transition, drawing most of its species
from above and below. The following is a composite list made up from notes
gathered on both sides of the lake.

Primary spp.: Betula glandulosa, Salix glauca var. berstipula, Empetrum nigrum, Phyl-
lodoce glanduliflora, Cassiope tetragona, Festuca.altaica.

SECONDARY spp.: Lycopodium alpinum, Lycopodium complanatum, Abies lasiocarpa,
Picea glauca var. albertiana, Juniperus communis var. montana, Agrostis scabra, Calama-
grostis canadensis var. robusta, Poa arctica, Poa alpina, Trisetum spicatum var. Maidenii,
Carex atrata ssp. atrosquama, Carex phaeocephala, Carex Soperi, Carex podocarpa, Carex
pyrenaica, Carex scirpoidea, Carex vaginata, Luzula Wahlenbergti, Lloydia serotina,
Zygadenus elegans, Cypripedium passerinum, Salix alaxensis, Salix arctica, Salix Bar-
rattiana, Salix Barrattiana var. marcescens, Salix commutata, Salix planifolia, Salix
reticulata, Salix Richardsonii, Oxyria digyna, Polygonum viviparum, Silene acaulis var.
exscapa, Aconitum delphinifolium, Anemone narcissiflora, Anemone Richardsonii, Del-
phinium glaucum, Arabis Drummondii, Arabis lyrata var. kamchatica, Sedum roseum
var. integrifolium, Saxifraga Lyallii, Saxifraga punctata, Saxifraga tricuspidata, Dryas
integrifolia, Astragalus frigidus var. littoralis, Hedysarum alpinum var. americanum,
Oxytropis hyperborea, Viola epipsila, Shepherdia canadensis, Cornus canadensis f. pur-
purascens, Vaccinium Vitis-Idaea var. minus, Polemonium acutiflorum, Pedicularis capi-
tata, Pedicularis labradorica, Veronica alpina var. unalaschcensis, Linnaea borealis var.
americana, Erigeron unalaschkensis, Antennaria isolepis, Artemisia arctica, Petasites
frigidus, Arnica alpina, Senecio lugens, Senecio triangularis, Hieracium gracile.

A few species were found at Brintnell Lake only in the subalpine scrub zone:
Carex Soperi, Saxifraga Lyallii, Viola epipsila, Cornus canadensis f. purpu-
rascens, Petasites frigidus, and Senecio triangularis. Some of these were quite
rare, and none were common except locally.

Another small group, though found elsewhere on the mountains, appeared to
be especially abundant in the scrub. These are: Lycopodium alpinum, Abies
lasiocarpa, Lusula Wahlenbergii, Salix Barrattiana and var. marcescens, Del-
phinium glaucum, Aconitum delphinifolium, Polemonium acutiflorum, and Pe-
dicularis labradorica.

A few alpine species achieve their lowest limits in the scrub: Lycopodium
alpinum, Carex phaeocephala, Carex podocarpa, Lloydia serotina, Salix arctica,
Salix Barrattiana and var. marcescens, Anemone narcissiflora, Delphinium glau-
cum, Sedum roseum var. integrifolium, Saxifraga punctata, Phyllodoce glanduli-
flora, Veronica alpina var. unalaschcensis, Erigeron unalaschkensis, and Hiera-
cium gracile. Lower slope species that here reach the upper limits are as
follows: Abies lasiocarpa, Picea glauca var. albertiana, Agrostis scabra, Calama-
grostis canadensis var. robusta, Carex vaginata, Luzula Wahlenbergii, Cypri-

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 49

pedium passerinum, Salix commutata, Salix planifolia, Arabis Drummondi,
and Antennaria isolepis.

Thus in spite of its discontinuous distribution and variable width, the sub-
alpine scrub is seen to have a certain amount of floristic individuality, and at
the same time to constitute a meeting-ground for alpine and lower slope plants.
Of the 69 species listed for the whole zone, 15, or about 21.7%, may be regarded
as fairly characteristic. Twenty-five plants, or about 36.2%, reach their upper
or lower limits in the scrub. There is a small amount of overlap in the cate-
gories just mentioned, but the scrub as a habitat appears to have some particular
significance for a little over 50% of the species that grow there.

Some mention should be made of local geographic variations in the structure of
the scrub communities. At timber line in the valley of Frost Creek the principal
species are the willows, which form dense thickets 4-6 feet high. Along the
stream itself Salix alaxensis becomes a primary species in place of S. glauca
var. perstipula. At higher elevations the shrubs of all kinds become dwarfed,
and at the upper limits of the zone are often no more than a foot high. On the
south and southeast slopes of Red Mountain the willows outnumber the dwarf
birches; but on the east slope, as well as on the terraces of Terrace Mountain,
dwarf birch forms nearly pure stands. Cassiope and Phyllodoce are much more
prominent on the northeasterly slope of Red Mountain than elsewhere.

Throughout the scrub zone, but particularly toward its upper parts, the con-
tinuity of the thickets is much broken by grassy openings in which Festuca
altaica is a primary species. Most of the sedges and grasses as well as many
of the other herbaceous plants mentioned in the list are found in these openings.
A few, such as Delphinium, Aconitum, and Polemonium, seem to find their most
congenial habitat in the dense thickets. These thickets are difficult to climb
through because their interlaced branches are depressed downward like those
described in the snowslide areas.

Alpine vegetation

It is nearly impossible to organize the alpine vegetation into clear-cut com-
munities. In some places the plants form a fairly continuous turf in which
interrelationships among plants probably are developed, but over most of the
alpine surfaces interdependencies among species are difficult or impossible to
define. It is doubtful whether these loose aggregations of plants can be called
communities in the usual ecological sense. Their composition and distribution
are determined very largely by the physical nature of the substratum, and since
this is the case it seems best to organize them in terms of the substratum.

The nature of the alpine plant cover depends in large measure upon the
stability of the soil and the amount of water at or near the surface. Both of
these factors in turn are dependent for their expression upon the relative friability
of the underlying rock, the degree, of slope, the degree of consolidation of the
material in case it is a glacial till, and the length of time since the surface was
exposed by the glaciers. With regard to the last of these conditions it seems
reasonable to assume that most of the alpine surfaces around Brintnell Lake
are somewhat older than those now under forest and subalpine scrub. Evidence
to this effect is in the upper limit of obvious glacial erosion in the main valley
of the lake (about 4300 feet), and the upper limits of the younger till on the
slopes of Red and Terrace Mountains (about 5000 feet). The date of the last

50 SARGENTIA [6

advance of the ice is not known, nor is the length of the interval between this
and the preceding one.

It is possible to make a rough division of the alpine vegetation on the basis
of the stability of the surface, with subdivisions related to the size of the mate-
rials and their drainage conditions.

Soils essentially stable, though still subject to slow creep.
Alpine “meadows,” in which the soil is mostly covered by a turf.
1. “Damp meadows” along streams and in snow-flush areas.
2. “Dry meadows” on stabilized slide rock.
Soils in active motion at frequent intervals,
Steep slopes of slide rock.
3. Plants scattered in loose slide rock.
4. Plants in turfy banks or terraces.
Soil kept in motion by frost heaving on level or gently sloping surfaces.
5. Plants scattered in heaved soil or in turf at the margins of polygons.
Crevices and ledges.
6. Plants scattered or in turf.

It must be kept in mind that these subdivisions constitute only a working
scheme for the description of the alpine vegetation in the immediate vicinity of
Brintnell Lake. They do not account for types of alpine habitats such as lake
shores which were not observed there. Furthermore they are clearly defined
only in their extreme forms, and every conceivable intergradation among them
can be found. On the other hand the scheme takes cognizance of an environ-
mental influence which is probably of first importance in the determination of
the content and distribution of arctic-alpine vegetation. This is the action of
frost in the break-up of the parent rock and in the subsequent movement of the
fine materials to a state of rest. The influence of the vegetation in furthering
the latter process has been clearly recognized by such students as Antevs (1932)
and Polunin (1934-35), and Polunin (1. c.) has pointed out the historical sig-
nificance of the counter-influence of solifluction upon the development of the
vegetation.

Lack of success in defining alpine plant communities in the usual terms of
primary and secondary, or dominant and incidental species can be logically
accounted for by the long-continued frost action in alpine habitats. This factor
probably has been in effect during all of the Glacial and post-Glacial period, so
that the whole history of the occupance by plants of such alpine habitats as occur
in the Mackenzie Mountains is one of vegetational instability.

With these things in view it may well be advisable to give up trying to describe
a great deal of our alpine and arctic vegetation in terms of plant associations,
and to institute methods that will recognize the significance of a state of con-
tinuing instability. I have already discussed the difficulties that arise from too
close adherence to the prescribed ecological methods (1941; see also Griggs,
1934). In the ensuing descriptions it is to be understood that the term “pri-
mary species” is significant chiefly as a measure of frequency except in some
of the most stable meadow types.

Before beginning an account of the alpine vegetation some preliminary con-
siderations are necessary. It will be noted that all our data on this type of
habitat were gathered on the mountains of sedimentary rock. The granite peaks
to the westward of Brintnell Lake were inaccessible in the time available, and
probably are unscalable. Examined through field glasses they appeared to be
nearly sterile, with the cliffs exfoliating so rapidly as to make plant life nearly

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 51

impossible. The great talus at the foot of Mt. Harrison Smith, which appears
in Prats III, is practically devoid of plants in many places, even nearly down
to lake-level.

A curious phenomenon of local distribution may also be pointed out here. A
number of species were found to be absent or rare on one or the other of the
mountains north and south of the lake. The following are incomplete lists
which will illustrate this.

Common on the north slope of Colonel Mt., but absent or only rare or occa-
sional on Red and Terrace Mountains: Zygadenus elegans, Tofieldia palustris,
Salix Barraitiana, Salix Richardsonii, Oxyria digyna, Silene acaulis var. exscapa,
Stellaria longipes var. Edwardsii, Arenaria Rossii, Lychnis apetala, Saxifraga
oppositifolia, Saxifraga aizoides, Astragalus frigidus var. littoralis, Hedysarum
alpinum var. americanum, Oxytropis Maydelliana, Oxytropis pygmaea, Oxy-
tropis hyperborea, Oxytropis ixodes, Epilobium latifolium, Pedicularis capitata,
Senecio lugens, Crepis nana.

Common on the slopes of Red or Terrace Mountains, but rare or absent on
the north side of Colonel Mt.: Lycopodium alpinum, Lycopodium Selago, Hiero-
chloé alpina, Poa arctica, Carex albo-nigra, Luzula confusa, Luzula spicata,
Lugula arcuata, Corydalis pauciflora, Phyllodoce glanduliflora, Gentiana glauca,
Veronica alpina var. unalaschcensis, Erigeron eriocephalus, Erigeron jucundus,
Arnica mollis, Arnica alpina ssp. tomentosa.

Causes for these differences were not determined. They could be looked for
in local climatic differences due to north and south exposure, or to variations in
the chemical nature of the substratum. It is probable that there is more calcium
in the rocks south of the lake than north of it, due to an abundance of calcite
stringers in the shale. The flora of Colonel Mountain bears a closer resem-
blance to the extremely calcicolous communities described by Porsild (1945) on
the “Plains of Abraham” along the Canol Road than does that of Red and Ter-
race Mountains (see below).

Damp alpine meadows

Damp alpine meadows at Brintnell Lake are best developed along small brooks
where the latter flow over terraces of glacial till or other soil containing fines.
They also appear on moderately sloping surfaces where snow lies in depressions
well into the summer. Here the substratum may be of coarser debris because
of the continued source of water from the melting snow. These snow-flush areas
are commonly not so rich in species or individuals as brook valleys, for they are
apt to become quite dry in late summer. Damp meadows make up, together,
only a small part of the alpine vegetation, for surface water above timber line
in these mountains is not common. They are usually formed in a thick turf
which gives the impression of permanence and stability. It is common, how-
ever, to see large masses of this turf breaking up by transverse crevasses which
produce an appearance of draped festoons when seen at a distance. They are
thus constantly being dissected, later to form continuous mats again a little lower
down the slopes.

Primary spp.: Salix alaxensis, Salix glauca var. perstipula, Salix Richardsonu, Salix

Barrattiana, Betula glandulosa, Carex podocarpa, Festuca altaica.

SECONDARY SpP.: Equisetum arvense, Equisetum variegatum, Arctagrostis arundinacea,

Hierochloé alpina, Poa arctica, Carex atrata ssp. atrosquama, Carex deflexa, Carex neso-
phila, Luzula arcuata, Luzula spicata, Lloydia serotina, Salix arctica, Salix reticulata,

52 SARGENTIA [6

Oxyria digyna, Polygonum viviparum, Arenaria sajanensis, Aconitum delphinifolium,
Anemone narcissiflora, Anemone parviflora, Anemone Richardsonii, Delphinium glaucum,
Ranunculus Eschscholtsii, Ranunculus nivalis, Corydalis pauciflora, Cardamine bellidifolia,
Draba longipes, Draba McCallae, Sedum roseum var. integrifolium, Parnassia fimbriata,
Parnassia Kotzebuei, Saxifraga punctata, Savxifraga rivularis, Potentilla diversifolia var.
glaucophylla, Sibbaldia procumbens, Astragalus frigidus var. littoralis, Epilobium anagalli-
difolium, Pyrola minor, Cassiope tetragona, Gentiana glauca, Polemonium acutiflorum,
Mertensia paniculata, Myosotis alpestris ssp. asiatica, Pedicularis labradorica, Veronica
alpina var. unalaschcensis, Campanula lasiocarpa, Erigeron eriocephalus, Erigeron una-
laschkensis, Antennaria monocephala, Artemisia arctica, Arnica Lessingti, Hieracium
gracile, Taraxacum alaskanum.

The above list is of course a composite one, made up from various situations.
Nevertheless in some localities nearly all of the species may be found in close
proximity. The willows are mostly concentrated along the streams or ‘snow-
flush channels, while the grasses, sedges, and other herbaceous plants form the
turf on either side, with occasional willows and dwarf birches scattered here and
there. On many sites there is no small group of species that can be designated
as primary, for the community is a hodge podge mixture of many.

Dry alpine meadows

Dry alpine meadows as I have conceived them here include a great variety of
plant groupings. They occupy erosion and solifluction slopes that are fairly
well stabilized but which have no continuous source of surface water except in
the spring when most of the snow is melting. The mineral substratum is made
of angular fragments of slate and shale, in some places comminuted to form a
few inches of residual soil.

With the possible exception of slide rock, this type of vegetation covers more
area than any other in the Brintnell Lake region. The long, even slopes of the
gently rolling ancient surface seen from the air in the eastern parts of the
Mackenzie Mountains are covered with it. It is represented at Brintnell Lake
on the summits of Red and Terrace Mountains, and probably on the upper south-
facing slopes of Colonel Mt. It also appears on the lower parts of the great talus
accumulations between 4000 and 6000 feet.

The primary group of species in the dry meadows varies greatly with exposure
and the relative stability of the soil. Before noting these variations I shall give
a list of all the species observed in the dry meadows.

Woodsia ilvensis, Lycopodium alpinum, Lycopodium annotinum var. alpestre, Lyco-
podium complanatum, Lycopodium Selago, Picea glauca var. albertiana (occasional
dwarfs), Juniperus communis var. montana, Agropyron latiglume, Calamagrostis lap-
ponica, Calamagrostis purpurascens, Festuca altaica, Festuca brachyphylla, Hierochloé
alpina, Poa alpina, Poa arctica, Poa glauca, Trisetum Spicatum var. Maidenii, Carex bi-
partita, Carex nardina var. Hepburnii, Carex bhaeocephala, Carex podocarpa, Carex
pyrenaica, Carex scirpoidea, Lusula arcuata, Lusula confusa, Lusula spicata, Tofieldia
palustris, Zygadenus elegans, Salix arctica, Salix glauca var. acutifolia, Salix glauca var.
berstipula, Salix pulchra, Salix reticulata, Betula glandulosa, Oxyria digyna, Polygonum
viviparum, Arenaria sajanensis, Arenaria verna var. pubescens, Silene acaulis var. exscapa,
Stellaria longipes var. Edwardsii, Aconitum delphinifolium, Anemone narcissiflora, Anem-
one parviflora, Draba fladnizensis var. heterotricha, Draba glabella, Sedum roseum var.
integrifolium, Saxifraga oppositifolia, Saxifraga tricuspidata, Dryas integrifolia, Dryas
octopetala, Potentilla diversifolia var. glaucophylla, Potentilla emarginata, Potentilla fruti-
cosa, Potentilla nivea, Potentilla uniflora, Rosa acicularis, Astragalus frigidus var. lit-
toralis, Hedysarum alpinum var. americanum, Hedysarum Mackensii, Lupinus arcticus,
Oxytropis hyperborca, Oxytropis ixodes, Oxytropis Maydelliana, Oxytropis pygmaea,
Empetrum nigrum, Shepherdia canadensis, Epilobium angustifolium, Epilobium latifolium,

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 53

Pyrola asarifolia var. incarnata, Pyrola secunda, Arctostaphylos rubra, Cassiope tetra-
gona, Ledum groenlandicum, Phyllodoce glanduliflora, Vaccinium uliginosum var. alpinum,
Vaccinium Vitis-Idaea var. minus, Gentiana glauca, Gentiana propinqua, Polemonium acuti-
florum, Mertensia paniculata, Myosotis alpestris ssp. asiatica, Pedicularis capitata, Pedicu-
laris labradorica, Pedicularis lanata, Veronica alpina var. unalaschcensis, Linnaea borealis
var. americana, Campanula lasiocarpa, Solidago multiradiata, Antennaria media, Anten-
naria monocephala, Artemisia arctica, Arnica alpina, Arnica Snyderi, Senecio lugens,
Hieracium gracile.

This is by far the largest single assemblage of plants on the alpine areas, or
for that matter in any of the Brintnell Lake habitats. Over 60% of all the spe-
cies found above the subalpine scrub are here. In no one place, however, 1s
so large a proportion of them found together as is the case of the group in damp
meadows.

Variations in general aspect of the dry meadows can best be shown by field
notes made on primary species in various situations. The most complex aggre-
gations are on slopes of northern and eastern exposure. Here a nearly con-
tinuous turf is sometimes formed, in which the primary species are Dryas inte-
grifolia, Cassiope tetragona, Festuca altaica, Salix reticulata, Carex scir poidea,
Vaccinium uliginosum var: alpinum, and mats of dwarfed Betula glandulosa.
Phyllodoce glanduliflora and Empetrum nigrum are sometimes added to the
group. This complex was observed on the north slopes of Colonel Mt. and
on the north and east slopes of Red Mt.

The alpine surface of Red Mountain shows further variation as follows. At
about 4800 feet on the south-facing slope is a rather open association of Vac-
cinium uliginosum var. alpinum, Festuca altaica, and a partial cover of lichens.
Among the latter the xerophytic Stereocaulon is predominant. Betula glandu-
losa and Salix glauca var. acutifolia are common in widely spaced, low mats.
At about 5900 feet, on the rounded slopes of the summit, the primary species are
Dryas octopetala, Vaccinium uliginosum var. alpinum, and a lichen mat made
up of Cladonia and Cetraria. Most of the ground is covered, but here and there
are open areas of loose shale or finer soils kept barren by frost action. In de-
pressions where snow had lain for a large part of the summer Phyllodoce glandu-
liflora is abundant.

The highest point of Red Mt. is a narrow ridge on which a thin soil has
weathered out of the shale. The most abundant plants are Festuca brachyphylla,
Poa arctica, and a loose mat of Cladonia and Cetraria. In a few places there
are mats of Vaccinium uliginosum var. alpinum and Cassiope tetragona. The
whole association is open, with much ledge and loose shale exposed.

Just east of the summit, at the beginning of a long slope of fairly well stabilized
talus, the aspect changes at once. Here the primary species become Festuca
altaica, Cassiope tetragona, Dryas octopetala, Vaccinium uliginosum var. al-
pinum, and masses of fruticose lichens. At 5000-5500 feet on the easterly and
northeasterly slopes this group is enlarged by the addition of Empetrum nigrum,
and becomes the turf community described above.

Some contrasts can be seen among these complexes even from a distance. On
some south-facing slopes Festuca altaica is very prominent, due to the small
development of other species with it. It is a bunch grass that grows in large
tufts often a foot or two thick and well separated from one another. When
growing thus it can be distinguished from far away, but it disappears when
mixed with many other species. :

54 SARGENTIA [6

Plants of loose slide rock and turf-banked terraces

Slide rock of loose, angular shale fragments is of wide occurrence above timber
line at Brintnell Lake. In some places whole mountainsides are composed of
continuous “rivers” of this shale (PLates IX, X). The parent rock does not
always break up into small pieces, however, so that here and there are masses
of coarser material, forming steep slopes of large angular boulders. The shale
is highly mobile, but the boulder fields are more or less stabilized and possess a
flora similar to that of crevices and ledges. Only plants whose water require-
ments are low, and whose root and rhizome systems are adaptable in the ex-
treme can live in the loose shale. The following is a list of those observed or
collected :

Woodsia ilvensis, Carex albo-nigra, Carex nardina var. Hepburnii, Carex nesophila,
Carex phaeocephala, Carex podocarpa, Lusula arcuata, Luzula confusa, Lusula spicata,
Zygadenus elegans, Salix arctica, Salix glauca var. acutifolia, Salix glauca var. perstipula,
Salix pulchra, Arenaria Rossii, Arenaria sajanensis, Arenaria verna var. pubescens, Lych-
nis apetala, Stellaria longipes var. Edwardsii, Aconitum delphinifolium, Arabis lyrata var.
kamchatica, Draba cinerea, Draba lanceolata, Draba nivalis var. elongata, Draba prae-
alta, Saxifraga aizoides, Saxifraga tricuspidata, Dryas Drummondii, Dryas integrifolia,
Dryas octopetala, Hedysarum Mackensii, Oxytropis Maydelliana, Oxytropis pygmaea,
Campanula lasiocarpa, Solidago multiradiata, Aster sibiricus, Erigeron jucundus, Arnica
mollis, Arnica Lessingii, Arnica alpina ssp. tomentosa, Arnica Snyderi, Crepis nana.

Except in certain instances to be described below, this assemblage of plants
cannot be called a community in any sense. The shale is scarcely more than
at an angle of rest, so that a touch is all that is needed to set it in motion. Most
of the plants are widely scattered, and there are large areas completely devoid of
vegetation of any kind. Roots and rhizomes are usually just beneath the sur-
face and nearly always extend up the slopes from the aerial parts of the plants.
From a tiny, dwarfed herbaceous perennial, all alone in the loose and sliding
shale and giving every appearance of being a chance seedling, one often can
excavate an elaborate system of rhizomes and roots extending many feet up the
slope and giving evidence of great age. Some plants of tufted habit seem to
slide with the shale without serious damage.

There is, of course, every kind of transition between the completely open
aggregations and the more or less stable communities described above as dry
alpine meadows. Only one of these stages need be given special notice. Some
of the more or less woody species which are most efficient in their mat-forming
proclivities, especially Dryas integrifolia, Saxifraga tricuspidata, Cassiope tetra-
gona, Salix glauca var. acutifolia, and Salix arctica, may form mats of inter-
laced stems and roots that can cling together in spite of the movement of the
shale. On the mats thus formed other plants can find at least temporary secur-
ity, and sometimes small areas of turf are formed, involving mosses and fruticose
lichens.

Many shale slopes have the appearance of being festooned with bands of
vegetation, or streaked with them up and down (PLate IX). These bands are
made up of the mats just described. On broad slopes whose surfaces are more
or less convex, and on which the movement of materials downward is fairly
uniform laterally, the mats are arranged transversely; but where the shale
“flows” down gully-like depressions, with lines of greater or less mobility ar-
ranged down slope, the mats are on the areas of greatest stability and follow
the lines of “flow.” The mats are never anchored with equal success at every

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 35

point, so that they commonly move en masse under weights of shale dammed up
above them, thus assuming the characteristic draped appearance. For the same
reason the mats arranged parallel to the lines of movement are often discon-
tinuous.

Causes for the movement of the shale other than gravity could not be accu-
rately evaluated. Torrential rains in spring and summer could start some of
it, as could also large amounts of meltwater at the time of the spring thaw. Or
it might be due to alternate freezing and thawing of water among the stones
early in the spring. The last is undoubtedly the principal cause on more nearly
stabilized slopes where turf-banked terraces are also sometimes formed. It is
of note that those seen on shale in the Brintnell Lake area were mostly on north-
facing slopes. South- and west-facing shale slides on Red and Terrace Moun-
tains were nearly barren, suggesting that the shale is there too mobile for the
mat-forming plants to get a foothold. This suggests that local climatic factors
are significant, resulting in more movement of snow in the spring, or more
intense frost action after the snow has left.

The vegetation of stone rings

Well-defined stone rings were observed at Brintnell Lake in only one place.
This was on a deposit of glacial till in the upper valley of Frost Creek, at an
altitude of about 4500 feet (PLate VI). The area where they were found is
wet and springy, with small pools of water here and there. The rings vary in
size from a Jew inches to about 6 feet, the larger ones bordered by till boulders
as much as 3 feet in diameter.

The movement of materials by frost is so active that the stone rings are nearly
devoid of vegetation except for crustose lichens on the larger boulders. A single
species of vascular plant was growing sporadically in the stirred soil of the
rings, Saxifraga Lyallii. Curiously enough, this species was found nowhere else
in the region. The area of the rings is well within the subalpine scrub zone,
as it is developed in the valley of Frost Creek, and the characteristic plants of
this zone encroach upon the outer rings and push into the more stable portions
around the large boulders.

The production of stone rings and stripes appears to be much more active on
the limestone mountains described by Porsild along the Canol Road (1945,
p. 5). He states, “Everywhere on sloping ground solifluction and live stone
creeps can be observed and on level ground polygon formation is active. Out-
crops of rock everywhere show evidence of rapid weathering by frost action.”
Porsild gives no detailed studies of the relation of plant growth to these phe-
nomena, although some of his notes on local habitats can be so interpreted (see
below ).

Ledge and crevice vegetation

Outcrops of creviced ledges occur on all the mountain slopes. Composed as
they are largely of shaly rocks, they are breaking up very rapidly under the
influence of subaerial erosion and frost cleavage. As stated previously, most
of the sedimentary rocks of the region dip sharply to the southward and south-
westward. Consequently the largest areas of shale slide-rock are on the south-
and west-facing slopes, while the most extensive cliff development is on the
upper north-facing ones (PLaTtE X). The north slope of Red Mt., for instance,

56 SARGENTIA [6

is a series of precipitous cliffs some 200 feet high, at the foot of which is a talus
accumulation that reaches down another 300 feet or so to a small stream. Still
higher cliffs appear on the north side of Colonel Mountain toward the summit,
Hogback ridges are everywhere broken by steep outcrops of bedrock.

Most of the flora of the ledges is scattered in crevices or in small accumula-
tions of residual soil that have formed on cliffs. Occasionally one finds a high
ledge that is fairly level, so that its residual soil stays in place and will retain
some water. Here a turf develops, carpeted by meadow types of community
like those already described. Such places are rare, however, and the association
of plants is usually open. As on slide rock, it is nearly impossible to designate
primary species. It might be proper to say that crustose lichens play this role,
for they cover most of the bare rock. Among vascular plants, Dryas integri-
folia, Saxifraga tricuspidata, Salix glauca var. acutifolia, and Trisetum Spicatum
var. Maidenii are probably the most common species in crevices and on dry
ledges. The following is a list of species found on these sites:

Cystopteris fragilis, Woodsia ilvensis, Juniperus communis var. montana, Calamagrostis
purpurascens, Poa alpina, Poa glauca, Poa paucispicula, Trisetum spicatum var. Maidenii,
Carex nardina var. Hepburnii, Lusula confusa, Lusula spicata, Salix glauca var. acuti-
folia, Arenaria Rossii, Arenaria sajanensis, Arenaria verna var. pubescens, Silene acaulis
var, exscapa, Arabis lyrata var. kamchatica, Cardamine bellidifolia, Draba cinerea, Draba
lanceolata, Draba nivalis var. elongata, Draba praealta, Saxifraga aizoides, Saxifraga
nivalis, Saxifraga oppositifolia, Saxifraga tricuspidata, Dryas integrifolia, Potentilla emar-
ginata, Potentilla nivea, Potentilla uniflora, Astragalus alpinus, Hedysarum Mackenszii,
Oxytropis ixodes, Oxytropis Maydelliana, Oxytropis pygmaea, Gentiana propinqua, Aster
sibiricus, Erigeron jucundus, Arnica Lessingii, Arnica Snyderi.

It will be noted that this list has much in common with that for shale slide
rock. The cliffs constitute an extremely xerophytic and physically hazardous
habitat, perhaps a little more stable than some of the slides. As previously
noted, talus made up of larger rock has an extremely poor flora which closely
resembles that of ledges.

A phase of the Mackenzie Mountain alpine vegetation not seen in the Brintnell
Lake area is described by Porsild (1945) on the “Plains of Abraham” along the
Canol Road. This locality is between 70 and 80 miles west of Norman Wells,
at an altitude of about 5700 feet. It is underlain by massive limestones which
give a strongly calcicolous aspect to the flora. Porsild designates four more or
less distinct habitats, “each conditioned largely by exposure to the prevailing
wind, snowcover and soil.”

The most stable of these habitats is on “level, sheltered flats, on gentle slopes
and at the foot of slopes where small terraces form. ...’ Here are listed
Arctagrostis latifolia, Puccinellia Vahliana, Carex atrofusca, Carex membra-
nacea, Kobresia simpliciuscula, Juncus triglumis, Juncus biglumis, Lloydia sero-
tina, Tofieldia palustris, Arenaria humifusa, Arenaria Rossii, Thalictrum al-
pinum, Papaver Keelei, Braya humilis, Draba alpina var. nana, Saxifraga
aizoides.

“On coarse, unsorted, broken limestone rock of the windswept ridges and
rocky summit,” the following species are noted: Carex misandra, Luzula nivalis,
Salix arctica, Salix phlebophylla, Arenaria arctica, Papaver Keelei, Saxifraga
oppositifolia, Dryas integrifolia.

In masses of Cassiope tetragona on “small islands” on gentle slopes were
found Thalictrum alpinum, Eutrema Edwardsii, Parrya nudicaulis, Potentilla

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 57

biflora, Oxytropis Maydelliana, Oxytropis terrae-novae, Pedicularis capitata,
Senecio atropurpureus.

Very unstable limestone talus has scattered plants of Carex nardina var. Hep-
burnii, Carex rupestris, Melanidion borealis, Potentilla uniflora.

From Porsild’s habitat notes on his collections it is possible to draw up a list
of species for still another type. This corresponds to our damp or wet alpine
meadows at Brintnell Lake, and is as follows: Equisetum variegatum, Arctagrostis
latifolia, Puccinellia Vahliana, Carex atrofusca, Carex membranacea, Carex
scirpoidea, Kobresia simpliciuscula, Juncus biglumis, Juncus ?triglumis, Lloydia
serotina, Tofieldia palustris, Salix arctica, Polygonum viviparum, Arenaria humi-
fusa, Arenaria sajanensis, Thalictrum alpinum, Anemone parviflora, Braya
humilis, Saxifraga Hirculus, Gentiana propinqua, Pedicularis sudetica, Erig-
eron unalaschkensis, Senecio resedifolius.

Plants of ravines and gorges below timber line

Rocky ravines and gorges near lake level have a flora drawn principally from
the more xerophytic elements of the slope forests. It is unnecessary to list it
in full. Nevertheless these situations harbor a few unique species which justify
their special notice. .

The rocky gorge of the stream which drains eastward from the lake has al-
ready been mentioned. The cliff-faces here have a rich cover of crustose lichens,
while on the ledges are mats of moss and fruticose lichens in which are dwarf
junipers and a few small white spruces and white birches. Calamagrostis pur-
purascens and Poa glauca are also common there.

Creeks that drain the north slope of Colonel Mt. have cut deep into the bed-
rock between the hogbacks, so that by climbing over the recently formed stony
fans we could get into narrow gorges with precipitous and often overhanging
cliffs on either side. Two of the new species described in this paper, Poa
Brintnellii and Lychnis brachycalyx, were found in one of these gorges, as well
as a species of saxifrage (Saxifraga sibirica) which proves to be an addition to
the flora of the continent. These places are probably among the most difficult
plant habitats which the region affords. They are subject to torrential floods,
falling rock debris, rapid disintegration of the rock itself, and a growing season
reduced to lowest terms. In many spots there is never any direct sunlight.

Ravines on south-facing slopes are in sharp contrast. Their streams are
lined with tall willows, and the forest flora pushes into every crevice and soil
pocket. On the other hand no plants of unusual geographic or taxonomic
interest could be found in them.

Flood plain communities

A striking vegetational feature of the Brintnell Lake area is on the flood plain
of the glacial stream at the western end of the lake. The youngest of the stream
deposits are mud flats formed of the light gray rock flour brought down from
the glacier. This material is in the form of a small delta at the mouth of the
creek. A short distance above the lake coarser materials, sand and gravel,
occur as low plains and ridges. Most of the sand and gravel appears to be
coming from small tributary streams which come off the nearby mountainsides
at steep gradients and spread their loads on the plain. Occasional torrential
floods in the main creek probably are effective in reworking the sand and gravel.

58 SARGENTIA [6

I have already described the principal forest types on the flood plain, a white
spruce type and a muskeg forest. One of the few places at Brintnell Lake in
which developmental sequences in vegetation can be postulated reasonably is
on this flood plain, and stages leading to the forest types can be defined with
fair precision. The following notes will be presented, therefore, in the order
of a proposed scheme for the successional development of the forests.

The most recently formed mud flats at the mouth of the creek have a dense
stand of Eriophorum Scheuchzeri and Juncus castaneus (PLate XI). The two
species are completely mixed, but are nearly alone in forming the rather close
cover. When the Eriophorum is in fruit, its cottony heads turn the flats white
and make a conspicuous feature of the landscape. Small pools cut off from the
main lake in the formation of the delta are devoid of aquatic plants, but are
usually surrounded by pure stands of Equisetum palustre. The latter species
also appears to form a transition community between the Eriophorum—Juncus
association and one of grasses.

On surfaces that are a little older, the principal species are Calamagrostis
canadensis var. Langsdorfi and Arctagrostis arundinacea. Arctagrostis attains
its greatest abundance in the transition areas from Equisetum palustre, while
the Calamagrostis is thickest on the higher parts of the marsh and on low natural
levees along the creek. Equisetum palustre remains common throughout the
Calamagrostis—Arctagrostis stage. Other plants in this community are Equi-
setum fluviatile, Poa pratensis, Stellaria borealis, and occasional willows, Salix
commutata,.

On fairly well-drained sites the Calamagrostis gives way directly to a shrub
border of species derived from the neighboring flood plain forests: Alnus crispa,
Rosa acicularis, Viburnum edule, and Ribes triste. At the transition from
Calamagrostis to shrubs are several species which are not found elsewhere in
the area or have their greatest development here: Heracleum lanatum, Geum
macrophyllum var. perincisum, and Artemisia Tilesii. Other common plants
are Agropyron trachycaulum vars. unilaterale and novae-angliae. The shrubs
pass directly into the white spruce woods.

Much of the shore at the western end of the lake, away from the new mud
flats, is marsh. The principal species nearly everywhere is Carex aquatilis,
though Calamagrostis neglecta and C. canadensis var. robusta are also common.

On poorly drained mud flats back of the shore, either the Eriophorum—Juncus
or the Carex aquatilis shore community may develop into a distinctive associa-
tion which seems to be a forerunner of the muskeg. This association occupies
wide areas, and has the most complex flora to be found on the muds.

PRIMARY spp.: Equisetum variegatum, mosses.

SECONDARY spp.: Selaginella selaginoides, Agropyron trachycaulum var. Langsdorfi,
Agrostis scabra, Festuca altaica, Carex microglochin, Tofieldia palustris, Cypripedium
passerinum, Habenaria hyperborea, Salix commutata, Salix reticulata, Alnus tenuifolia,
Betula glandulosa, Anemone parviflora, Parnassia palustris var. neogaea, Dryas Drum-
mondii, Dryas integrifolia, Hedysarum alpinum var. americanum, Pyrola asarifolia var.
incarnata, Arctostaphylos rubra, Antennaria sp.

The moss and the Equisetum form a close mat over most of the ground. The
latter is so abundant that as one walks through it there is a continuous crackling
sound from the breaking siliceous stems. The willows, alders, and dwarf
birches are only occasional except at the landward margins of the community
where they become primary and pass into the muskeg. Dryas is commonest

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 59

on the drier sites, especially where there is a gravel or sand plain nearby. Oc-
casional small white spruces and balsam poplars also appear on dry sites.

The most xerophytic habitats in the valley west of the lake are the gravel and
sand plains. For the most part they have open plant associations, with the
following species involved :

Agropyron latiglume, Festuca brachyphylla, Zygadenus elegans, Populus Tacama-
hacca, Arenaria dawsonensis, Draba praealta, Dryas Drummondii, Fragaria glauca, Gen-
tiana propinqua, Campanula lasiocarpa, Aster sibiricus, Antennaria isolepis, Taraxacum
alaskanum, Taraxacum lapponicum.

Much of the sand and gravel is barren, with the above plants scattered here
and there. Over large areas, however, Dryas Drummondii forms tight mats to
the exclusion of nearly everything else. Next to the Dryas, Aster sibiricus,
which also tends to form mats, is probably the commonest species. The stabili-
zation of the soil by these mats, with the accumulation of some humus, appears
to set the stage for Populus Tacamahacca, which comes up to form a scrubby
bush at the borders of the plains and finally grows into an open woods in which
young white spruces come up. All stages in the transition to spruce through a
stand of balsam poplar can be seen on the gravel plains and on the gravel fans
from which they came. Similar transition woods are to be seen in reduced form
on nearly all the gravel fans around the lake,

TABLE 3 gives a tentative outline for the course of these flood plain successions.

TABLE 3.

PROVISIONAL SCHEME OF PLANT SUCCESSIONS ON FLOOD PLAIN DEPOSITS AT
WEST END OF BRINTNELL LAKE

Picea glauca Picea mariana—Ledum— Picea glauca
Sphagnum (muskeg)
Shrubs
Shrubs Populus Tacamahacca
Calamagrostis
canadensis var.
Equisetum variegatum Dryas Drummondit
Equisetum palustre
Carex aquatilis Open herbaceous
Pa association
Eriophorum—Juncus
~v i al —————
Newly formed Marshy shores Sand and gravel
mud flats plains and bars

PLANT CoMMUNITIES ALONG THE CANOL RoaAp

Mr. A. E. Porsild has published (1945) excellent descriptions of vegetation
types which he saw along the Canol Road between Macmillan Pass and Norman
Wells. It is unnecessary to transcribe his notes here, but a brief guide to them

60 SARGENTIA [6

will facilitate comparison with those made by our party. A few references have
already been made to them. Page citations refer to his paper:

Mackenzie valley forests were described in the area between Norman Wells
and the lower Carcajou River (pp. 3-4), while muskegs and lower slope muskeg
forests were noted near the crossing of Little Keele River about 50 miles west
of Norman Wells (pp. 4-5). Higher slope and timber line forests and those of
gravel plains along the streams are noted briefly along the upper Carcajou River
and along Bolstead Creek (pp. 7-8), and again in the valley of the Twitya
River (p. 8).

Alpine communities are described in some detail on the “Plains of Abraham,”
between 70 and 80 miles west of Norman Wells and at an altitude of about 5700
feet (pp. 5-7).

Further notes on alpine vegetation are given for Macmillan Pass (pp. 10-12),
along with a few comments on the geographic affinities of the flora in this pass.
Porsild points out that Macmillan Pass appears to mark a well-defined floristic
boundary between the eastern and western slopes of the Mackenzie Mountains.

Frost action phenomena such as stone rings are described on the Plains of
Abraham (pp. 5-7), and peculiar frost mounds in peat, known as “pingoes,”
were seen at Macmillan Pass (pp. 10-11).

GEOGRAPHIC AFFINITIES OF THE VEGETATION
OF SOUTHWESTERN MACKENZIE

INTRODUCTORY

Thus far I have attempted only to achieve botanical descriptions of some
landscapes in southwestern Mackenzie. The geographic breadth of these de-
scriptions and the amount of detail in them are extremely limited; but there are
enough, I believe, to bring out the more important phytogeographic patterns
in the area as a whole. An intriguing aspect of the vegetation is that of its
developmental history, which, if it can be learned, will throw a great deal of
light upon present trends of change and the causes for present distribution.
Some evidence with which to reconstruct this history is to be found in the
existing distribution and floristic content of plant communities, and in the geo-
graphic affinities both of the species and of the communities.

A complete analysis of geographic relationships would require a far more
extensive knowledge of the plant life of all of southwestern Mackenzie than
now exists. Lacking such extensive detailed knowledge, it seems best to select
for study the areas for which we have the most complete information. The
region of Brintnell Lake is outstanding in this respect, though it is small and
not thoroughly representative of the Mackenzie Mountains. For the plateaus
and lowlands east of the mountains our floristic knowledge is far from complete,
but enough is known about their forest types to justify proposing the general
geographic relationships of these types to other forests of the northwest.

Such conclusions as I have reached regarding the origin and history of the
vegetation of this region, therefore, are based largely upon interpretations of
the flora of the Brintnell Lake area, and of the plant communities both at Brint-
nell Lake and in the lowlands east of the mountains. Before setting down these
interpretations it is necessary to present some analyses of the geographic pat-
terns that are available.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 61

GEOGRAPHIC RELATIONSHIPS OF THE FORESTS OF SOUTHWESTERN MACKENZIE

The upland forest of white spruce, jack pine, aspen, and balsam poplar, with
its accompanying muskegs, is probably the most widespread timber type in
southwestern Mackenzie east of the mountains. It was noted by Whittaker
(1923) between the Mackenzie and the Horn Mountains, by Williams (1922)
between Simpson and Wrigley, by Whittaker in the region south of the upper
Mackenzie, and by McConnell (1891) along the lower Liard. It is character-
istic of the uplands of the Wood Buffalo Park (Raup, 1935). Porsild (1945)
has some brief notes on the region between Norman Wells and the Carcajou
River which suggest that aspen, poplar, and pine are not so abundant there or
have dropped out, and that white spruce and white birch are the principal forest
trees on the uplands. A similar transition must occur on the eastern slopes of
the Mackenzie Mountains west of Simpson and in the South Nahanni River
valley. The northwestern limit of the jack pine in the Mackenzie valley is
probably somewhere between Simpson and Norman. Preble (1908) placed it
at about lat. 64° 30’.

Upland forests closely resembling those along the upper Mackenzie are wide-
spread in the upper Liard basin (Raup, 1945), but here the lodgepole pine re-
places the jack pine and there is very little balsam poplar in the burn succes-
sions. The transition from lodgepole to jack pine presumably is in the Liard
gap and on the slopes of the northern Rocky Mountains (McConnell, 1891), and
it is supposed that the two species overlap there (Halliday and Brown, 1943).

All of the forests along the upper Mackenzie and lower Liard Rivers are
placed by Halliday (1937) in his “Mackenzie Lowland Section.” In his dis-
cussion of this section, however, he makes no subdivision of it into flood plain
and upland types such as appears to be necessary in this region. He sets a
northern boundary for his “Mixedwood Section” approximately at the northern
margin of the Alberta Plateau southwest of Great Slave Lake and south of the
Mackenzie and Liard Rivers. The northwestern phase of the Mixedwood as
Halliday describes it scarcely can be distinguished from some phases of his
Mackenzie Lowland forest. It now seems more logical to redefine the Macken-
zie Lowland Section so that it includes only flood plain types, composed prin-
cipally of white spruce and balsam poplar. Upland types in the upper Mackenzie
and lower Liard country could then be considered representatives of the far-
northwestern expression of Halliday’s Mixedwood, in which white spruce, aspen,
jack pine, and balsam poplar are the most prominent trees, and in which there
is some admixture of white birch.

This proposal necessitates greatly narrowing the area of the Mackenzie Low-
land type, confining it to recent flood plain deposits along the rivers. The
Mixedwood should then be extended northward in the Mackenzie valley, possibly
to a limit that could be marked by the extent of the jack pine as an important
tree. The eastward range of the Mixedwood, between the upper Mackenzie
and the Laurentian Plateau, is as yet unknown.

At Brintnell Lake the lower slope forests of white spruce and white birch
without pine bear a resemblance to those described by Porsild (1945) in the
Mackenzie valley between Norman Wells and the Carcajou River. These ap-
pear to be beyond the northern limit of the spruce-pine-aspen-poplar type of
the upper Mackenzie, and it has already been suggested that a similar change in
composition must occur on the lower eastern slopes of the mountains. The

62 SARGENTIA [6

white spruce—-white birch forest has no designation in Halliday’s classification
for Mackenzie basin types. The nearest approach to it is his “Yukon Section”
as it was described in 1937 (p. 23). He extends this section eastward in the
upper Porcupine River region approximately to the height of land west of
McPherson, but it is possible that it should reach also into the lower Mackenzie
country.

Whether or not Halliday’s “Upper Liard Section” extends eastward to the
South Nahanni River valley is yet unconfirmed. It has been so designated on
a map of the Alaska Highway area (Raup, 1945). The Upper Liard and
Mixedwood Sections are in reality quite similar in general appearance, structure,
and flora. The principal difference by which they can be separated is the pres-
ence of lodgepole pine in the former and jack pine in the latter. On this cri-
terion the eastern extent of the Upper Liard Section in our region is to be
determined by the limits of the lodgepole pine in the Mackenzie Mountains, and
the two sections probably overlap in the Liard Gap. Pines are known to be
abundant in the region of Clark, McMillan, and Landing Lakes, in the Mackenzie
Mountains southwest of Brintnell Lake. They were described to me by Mr.
William Clark of Fort Simpson, and they appear in various places on the aerial
photographs. It is presumed that they are lodgepole pines, although this is
yet to be verified.

The forests of black spruce on the steep north-facing slopes at Brintnell Lake
are likewise difficult to place in Halliday’s classification. A somewhat similar
type is of wide occurrence in the Rocky Mountains of northern British Co-
lumbia ; but there it usually contains lodgepole pine and some white spruce and
has been placed tentatively in a phase of Halliday’s “Foothills Section” (Raup,
1945).

GEOGRAPHIC RELATIONSHIPS OF SPECIES IN THE BRINTNELL LAKE REGION

Of the 283 species and varieties of vascular plants collected at Brintnell Lake,
I have mapped 271. When the maps of range limits are superposed it becomes
possible to designate a number of rather well-defined patterns. The actual num-
ber of these patterns depends of course upon the degree of refinement that
enters into the selection of criteria. Needless to say, the scanty existing knowl-
edge of geographic factors in boreal America makes it impossible to carry the
refinement very far. Comparison of the maps soon brings out, however, two
phytogeographic transition zones» which appear to overshadow any others that
appear. These are the arctic timber line, and the coastal mountains of south-
eastern Alaska, British Columbia, and western Washington. Some indication
of the significance of these “boundaries” may be had from the following figures.
Of the species considered in the maps 199 approach or cross the arctic timber
line, and 164 of them are either stopped by it or cross it to a limited distance.
Nearly all (283) of the Brintnell Lake plants extend to the western coastal
mountains or forests, but about 196 of them avoid all or part of these habitats.
In view of the obvious phytogeographic importance of the coast range and the
arctic timber line, I have drawn up a tentative classification of range patterns
based in large part upon the plants’ behavior with relation to them. A third
geographic boundary is not so well defined in all cases, but is sufficiently clear
to necessitate recognition. This is marked by the eastern ranges of the Cor-
dillera. About 25% of the Brintnell Lake flora does not extend east of the

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 63

Rocky Mountains, or has limited extensions into the northern great plain of
the continent.

About 4.2% of the flora at Brintnell Lake (12 spp. and vars.) is composed of
plants described as new in this paper, recently described from the lower Macken-
zie district, or known in North America from this single area. Tentative ranges
can be drawn for a few of these, but so little has been discovered about them that
I have preferred to keep them in a separate category, and I have not included
maps of them. They are as follows:

Picea glauca var. Porsildii, Poa Brintnellii, Carex Soperi, Salix Barrattiana var. mar-
cescens, Salix Bebbiana var. depilis, Salix glauca var. perstipula, Lychnis brachycalyx,
Saxifraga sibirica, Rosa acicularis var. cucurbiformis, Arnica alpina, Arnica Snyderi,
Antennaria sp.

At the other extreme in range extension are three cosmopolitan species:
Cystopteris fragilis, Equisetum arvense, and Carex aquatilis s. 1, These are
among the most wide-ranging plants in boreal America, and the first two are
cosmopolitan throughout much of the world. In Canada and Alaska they thrive
in nearly all parts of the arctic, alpine, and forest regions, without regard to
boundaries that are limiting to most other plants. Of the three, Equisetum
arvense seems to have the most varied stations. Cystopteris fragilis for some
reason has not been collected on the arctic coast of Alaska, the southwest coast
of Hudson Bay, the Richmond Gulf area on the east coast of Hudson Bay, or
in the Hamilton Inlet region of Labrador. Carex aquatilis is known on the
British Columbia coast only on Vancouver Island, and should perhaps be in-
cluded with reservation among the cosmopolitan species.

The remaining 268 species and varieties can be arranged in five general cate-
gories: (1) species whose principal areas are in the region of the Canadian
coniferous forest, and whose ranges are wide in this region, extending for the
most part all the way across the continent; (2) arctic or arctic-alpine wide-
ranging species with ranges concentrated north of the arctic timber line, or in
both the arctic lowland and in the cordilleran alpine areas; (3) an intermediate
group of wide-ranging subarctic species which cannot be placed with either of
the above, but which appear to have their greatest concentration on or near the
arctic timber line; (4) a group of Alaskan-cordilleran species which have only
limited extensions east of the Rocky and Mackenzie Mountains, and most of
which are arctic-alpine in general affinities; (5) a cordilleran group, most of
which are alpine but not arctic, and whose ranges are concentrated in the Rocky
Mountains and coast ranges. Further subdivisions of these five categories can
be made by noting first the actual relation of the wider ranges to the arctic
timber line, and second the relation of the ranges to the western coastal mountains
or the Pacific slope forests. The following discussion will follow the above
outline.

In order to compare the range maps in PLates XIII—-XXXVII more easily
I have drawn lines connecting marginal stations in such a way as to outline the
limits to which the various species are now known to extend. For common,
much collected plants this can be done without difficulty, but in cases of rare or
poorly collected species the outlines of geographic limits must be regarded as
tentative. The most difficult cases are those with stations separated by hun-
dreds of miles. Here it is often impossible to decide whether to consider the
ranges as continuous or disjunct. Of the ranges mapped in this study I have
designated only nine as being discontinuous, although it is possible that a few

64 SARGENTIA [6

others might be included. In defense of my tendency to reduce the number of
disjunct ranges as much as possible I can only say that one of the results of
the collecting and exploration in arctic and subarctic America during the past
twenty years has been to fill in and make continuous many ranges that were
formerly thought to have separate eastern and western components.

Wide-ranging forest species

The most extensive plant ranges in Canada and Alaska, with the exception of
the three cosmopolitans already mentioned, are exhibited by a small group of
about 19 species. These plants do not stop at the arctic timber line, but reach
far out into the tundra, particularly in the eastern arctic. Most of them are
spread throughout the timbered parts of Alaska. Eight of them (Fig. 4-a)
have the widest ranges because they also enter the forests of the Pacific slope
from southern Alaska to western Washington:

Equisetum scirpoides, Poa pratensis s. 1., Corallorrhiza trifida, Alnus crispa s. 1., Rubus
Chamaemorus, Epilobium angustifolium s. 1., Pyrola secunda, Ledum groenlandicum.

Fic. 4. Wide-ranging forest species extending north of the arctic timber line.

The other 11 species either avoid the coastal forests altogether, or enter them
only in southern Alaska or Washington. Thus they are more strictly con-
tinental in the far west (Fig. 4-b) :

Calamagrostis neglecta, Carex vaginata, Habenaria obtusata, Salix planifolia, Betula
glandulosa, Ranunculus Gmelini var. Purshii, Ranunculus lapponicus, Draba lanceolata,
Potentilla fruticosa, Hedysarum alpinum var. americanum, Vaccinium Vitis-Idaea var.
minus.

The remainder of the widespread forest species are more rigorously limited
by the timber line, though some of them occur far north on the Labrador coasts,
southern Baffin Island, or in Greenland. There are 80 of them, divisible into
four groups on the basis of their geographic relation to the Pacific slope forests.
The first of these groups (Fig. 5-a) contains 30 species that appear in the coastal
forests both in southern Alaska and in British Columbia and western Wash-
ington. Several of them, perhaps half a dozen, do not appear to reach the
western coast of Alaska.

Dryopteris Linnaeana, Equisetum fluviatile, Equisetum palustre, Lycopodium anno-
tinum s. 1., Selaginella selaginoides, Juniperus communis var. montana, Agrostis scabra,
Poa nemoralis, Carex leptalea, Carex canescens, Carex Garberi s. 1., Juncus balticus s. De
Smilacina stellata, Habenaria hyperborea, Salix Bebbiana, Arenaria lateriflora, Stellaria
calycantha s. 1., Actaea rubra, Ribes lacustre, Ribes triste, Shepherdia canadensis, Cornus
canadensis, Cornus stolonifera, Moneses uniflora, Pyrola asarifolia var. incarnata, Pyrola

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 65

minor, Heracleum lanatum, Arctostaphylos Uva-ursi (incl. var. coactilis), Linnaea bo-
realis s. 1., Viburnum edule.

The seconc group, of 13 species (Fig. 5-b), occurs in some of the coastal
forests of southeastern Alaska, but becomes more continental in British Columbia
and Washington. A few of them, probably not more than four, fail to reach the
western Alaskan coast.

hed \

N CBG ew
= — 7 g
¢ iAS ve é:

AY
2 IN Ss si
d NY See
SN ARN ATS
se WANS

(pe

SS
SEDO
SS
CPT x
PSO OK i, TA
eS ee,

Fic. 5. Wide-ranging forest species whose northern limits are at or near the arctic
timber line.

Equisetum sylvaticum s. 1., Lycopodium clavatum var. monostachyon, Lycopodium com-
planatum s. 1, Carex gynocrates, Eriophorum brachyantherum, Cypripedium passerinum,
Populus Tacamahacca, Populus tremuloides, Geocaulon lividum, Parnassia palustris var.
neogaea, Ribes hudsonianum, Dryas Drummondii, Rosa acicularis s. 1.

A third group of 14 species (Fig. 5-c) is still more continental in range,
reaching the coast timber only in southeastern British Columbia and western
Washington. They avoid the wet slope forests of southern Alaska, and none
of them extends to the western coasts of Alaska. Four of these plants seem to
have disjunct ranges: Arctostaphylos Uva-ursi var. adenotricha, ‘Betula papy-
rifera var. commutata, Arabis Holboellii, Antennaria subvicosa. I have in-
cluded them here because a hypothetical joining of their parts would result in
ranges similar to the others in Fig. 5-c.

Carex aurea, Betula papyrifera var. commutata, Arenaria dawsonensis, Arabis Drum-
mondii s. 1., Arabis Holboellii, Geum macrophyllum var. perincisum, Potentilla norvegica,
Pyrola virens, Arctostaphylos Uva-ursi var. adenotricha, Erigeron angulosus var. kamt-
schaticus, Erigeron elatus, Antennaria subviscosa, Senecio indecorus, Senecio pauperculus. ©

The fourth, and almost strictly continental, group contains 23 species (Fig.
5-d). With one or two possible exceptions these plants avoid the Pacific slope
forests entirely, and all but five of them fail to reach the west coasts of Alaska.
A few are even more restricted and do not extend west of the northern Rocky
Mountains.

66 SARGENTIA [6

Picea glauca s. 1., Picea mariana, Agropyron trachycaulum var. novae-angliae, Agro-
byron trachycaulum var. unilaterale, Calamagrostis canadensis var. robusta, Carex con-
cinna, Carex diandra, Carex media, Goodyera repens s. 1., Listera borealis, Salix myrtilli-
folia, Alnus tenuifolia, Betula occidentalis, Aquilegia brevistyla, Rorippa islandica var.
microcarpa, Ribes glandulosum, Ribes oxyacanthoides, Amelanchier florida (incl. A. hu-
milis), Fragaria glauca, Rubus acaulis, Rubus strigosus, Mertensia paniculata, Valeriana
septentrionalis.

Wide-ranging arctic-alpine plants

There are 66 wide-ranging arctic-alpine plants in the Brintnell Lake flora.
As among the forest species, one can first divide them into two groups accord-
ing to their actual relation to the arctic timber line. In spite of the fact that the
principal ranges of nearly all of them extend throughout much of the arctic
archipelago, 26 of them are commonly found far south of the arctic limit of
trees (Fig. 6-a, b, c). In so doing they achieve the most extensive ranges of
all our arctic-alpine plants and, as did the forest species that crossed the timber
line, they approach a cosmopolitan distribution in Canada and Alaska.

Fic. 6. Wide-ranging arctic and arctic-alpine species which extend far south of the
arctic timber line.

The most extensive ranges are those of 7 species (Fig. 6-a) which occur not
only on the Rocky and Mackenzie Mountains but also on the coastal ranges all
the way from the Alaska Peninsula to Washington. They extend southward in
the central plains at least as far as Lake Athabaska.

Equisetum variegatum, Lycopodium Selago, Festuca brachyphylla, Poa alpina s. 1.,

Carex scirpoidea, Salix arctica s. 1., Empetrum nigrum.

Another group, of 8 species, is predominantly continental south of the moun-
tains in southeastern Alaska (Fig. 6-b). They occur on the coast and Alaska
ranges of southern Yukon and Alaska, as well as on the mountains around the
Lynn Canal, but in British Columbia and Alberta they are largely confined to
the Rockies and Selkirks.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 67

Poa glauca, Trisetum spicatum s. 1., Carex capillaris s. 1., Juncus castaneus, Polygonum
viviparum, Saxifraga tricuspidata, Dryas integrifolia, Astragalus alpinus.

A third group of these extremely wide-ranging arctic-alpine plants contains
11 species (Fig. 6-c) and is almost completely continental in the west. South-
east of the Prince William Sound region of Alaska these plants are confined to
the more northerly ranges of the coastal mountains, and to the Rockies and
Selkirks. Only two of them come out to the coast in northwestern Washington.

Dryopteris fragrans s. str., Calamagrostis purpurascens, Arenaria verna var. pubescens,

Ranunculus hyperboreus, Cardamine pratensis s. 1., Draba cinerea, Chrysosplenium tetran-

drum, Saxifraga aizoides, Potentilla nivea s. 1. Pyrola grandiflora s. 1., Taraxacum

lacerum.

For the remaining 40 arctic-alpine plants the arctic timber line appears to be
a more formidable barrier. They either stop at the timber line or come only
a short distance south of it. Like the preceding group they can be arranged
in three divisions depending upon whether they extend to the mountains of
the Pacific coast.

First are 12 species of wide range on the coastal mountains (Fig. 7-a). One
of these, Dryas octopetala, has a broken range. It is widespread in the Cor-
dillera and reappears in northwestern Greenland.

Agropyror, latiglume, Poa arctica, Luzula spicata, Oxyria digyna, Silene acaulis var.

exscapa, Cardamine bellidifolia, Saxifraga oppositifolia, Saxifraga rivularis, Dryas octo-
petala, Epilobium latifolium, Epilobium anagallidifolium, Erigeron unalaschkensts.

4» KS

* ZN oe EN
KYA Y GET Lire
RE See

Re Fe “Se = <a 3

S

7 eee eee ~~

Fic. 7. Wide-ranging arctic and arctic-alpine species whose southern limits are at or
near the arctic timber line.

Another group of 10 species ranges the coastal mountains only in southern
and southeastern Alaska (Fig. 7-b). In British Columbia and Alberta they
reach their southern limits in the Rockies and Selkirks, and are strictly con-
tinental.

Hierochloé alpina, Carex bipartita, Eriophorum Scheuchzeri, Lusula Wahlenbergii,

Salix reticulata, Salix Richardsonii, Oxytropis foliolosa, Cassiope tetragona, Pedicularis
capitata, Pedicularis sudetica.

68 SARGENTIA [6

Finally, a group of 18 arctic-alpine plants, with one exception in western
Washington, avoid the coastal mountains entirely (Fig. 7-c).

Calamagrostis lapponica s. 1., Carex membranacea, Lusula confusa, Arenaria humifusa,
Arenaria Rossii, Lychnis apetala, Stellaria longipes var. Edwardsii, Ranunculus nivalis,
Draba fladnizensis var. heterotricha, Draba glabella, Saxifraga nivalis, Potentilla emargi-
nata, Oxytropis Maydelliana, Epilobium davuricum, Vaccinium uliginosum var. alpinum,
Pedicularis lanata, Erigeron eriocephalus, Crepis nana.

Two features of the arctic-alpine plant ranges should be noted especially.
Nearly all of them extend to the western Alaskan coasts, in contrast to the
forest plants among which 39 do not accomplish this. Second, there are a few
of the arctic plants, probably not more than 8, which do not occur in the Rocky
Mountains at all, but find their southern limits in the Mackenzie Mountains and
in southern Yukon or Alaska.

Plants of the timber line region

There are 31 species which constitute something of an enigma in that, al-
though they have wide subarctic ranges across most of the continent, they can-
not properly be classed either with forest or arctic-alpine categories. They
extend into the tundra, but avoid most of the high arctic regions. At the same
time they are common to abundant in the northern part of the forest. They
occupy a wide range of habitats, although none of them are characteristic of
rich woods, and about two-thirds of them are perhaps most abundant in muskegs
or wet tundra. Most of them appear in the northern Rockies, but two or three
do not come south of the 60th parallel. All but two or three reach the western
Alaskan coasts.

Sa 3 s

Fic. 8. “Timber-line species.”

A glance at Fig. 8-a and -b will show that most of these “timber line” species
are continental in the more southern parts of their ranges. I have arranged
them in two groups, carrying out the plan used in other categories. The first
group contains 10 species which occur in the coastal strip as far south as south-
eastern Alaska, while beyond that area all but 3 retreat inland (Fig. 8-a). One
species of disjunct range is included here, Lycopodium alpinum.

Woodsia ilvensis, Lycopodium alpinum, Calamagrostis canadensis var. Langsdorfi,
Cerastium Beeringianum, Anemone parviflora, Anemone Richardsonii, Sibbaldia procum-
bens, Andromeda Polifolia, Vaccinium microcarpum, Vaccinium uliginosum.

A second group, of 21 species, avoids the coastal strip except in southwestern
British Columbia, where four of them come out to Vancouver Island (Fig. 8-b).

\

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 69

Three species with discontinuous ranges are in this group: Sagina Linnaei,
Epilobium lactiflorum, Oxytropis txodes.
Carex deflexa, Carex microglochin, Tofieldia palustris, Salix glauca s. 1., Arenaria
sajanensis, Sagina Linnaei, Parnassia Kotsebuei, Hedysarum Mackenszii, Oxytropis hud-
* sonica, Oxytropis ixodes, Epilobium lactiflorum, Cornus canadensis f. purpurascens, Arcto-
staphylos rubra, Gentiana propinqua, Pedicularis labradorica, Veronica alpina var. una-
laschcensis, Pinguicula villosa, Solidago multiradiata, Antennaria isolepis, Taraxacum
ceratophorum, Taraxacum lapponicum.

Alaskan-cordilleran species

An arrangement of Alaskan and cordilleran components of the Brintnell Lake
flora on the basis of their relation to timber line proves to be impracticable.
Of the 72 species counted in these categories, only ten can be regarded as forest
plants, and mest of these ten are likely to be found far above or beyond the
timber line. All the others are plants commonly found in the alpine or western
arctic tundra. However, as I have noted elsewhere in this paper, a great many
alpine plants of the Mackenzie and northern Rocky Mountains are commonly
found in the timbered areas at low levels. About half of the western alpine
plants were found below timber line at Brintnell Lake, and no doubt further
search would yield more of them. It is not possible, therefore, safely to classify
them in relation to timber line.

mm.

Fic. 9. Alaskan-cordilleran species.

I have arranged the 72 species in two main groups, the first of which ranges
throughout central and western Alaska, many of them north to the arctic coast.
Most of them come into the mountains farther south, so that I have called them
the Alaskan-cordilleran group. There are 45 of these, 16 of which (Fig. 9-a
and -c) range southward into the western United States. The whole group can
be separated on the basis of its coastal extensions. First is a wide-ranging series
of 11 species, all of which occur on the coastal mountains of southeastern Alaska,
and many of them on the coast ranges farther south (Fig. 9-a).

70 SARGENTIA [6

Carex physocarpa, Lusula arcuata, Lloydia serotina, Zygadenus elegans, Arabis lyrata
var. kamchatica, Sedum roseum var. integrifolium, Viola epipsila, Gentiana glauca, Aster
sibiricus, Artemisia arctica s. 1., Petasites frigidus.

Allied to these, but with their southern extensions terminating in Yukon or
the Alberta and British Columbia Rockies, are 15 species, all of which are
coastal in southern or southeastern Alaska (Fig. 9-b).

Cryptogramma crispa var. sitchensis, Arctagrostis arundinacea, Festuca altaica, Poa
paucispicula, Carex loliacea, Salix alaxensis s. 1, Salix arbusculoides, Salix pulchra,
Betula papyrifera var. humilis, Aconitum delphinifolium, Anemone narcissiflora, Saxi-
fraga punctata s. 1., Boschniakia rossica, Campanula lasiocarpa, Arnica Lessingii.

Two groups are more strictly continental in the region southeast of Prince
William Sound, although a couple of species appear at the coast in the Puget
Sound area. Five species range southward into the United States (Fig. 9-c).

Delphinium glaucum, Potentilla uniflora, Myosotis alpestris s. 1., Artemisia Tilesti s. iF
Senecio lugens.

The remainder, 14 species, extend from Alaska to the northern Rocky Moun-
tains, and avoid the coastal mountains except west of Prince William Sound

(Fig. 9-d).

Larix laricina var. alaskensis, Carex nesophila, Carex podocarpa, Corydalis pauciflora
Draba longipes, Saxifraga radiata, Astragalus frigidus var. littoralis, Lupinus arcticus,
Oxytropis hyperborea, Oxytropis pygmaea, Gentiana arctophila, Polemonium acutiflorum,
Antennaria monocephala, Taraxacum alaskanunm.

,

Cordilleran species

The last category of ranges to be considered involves 27 plants that are more
strictly cordilleran than any of the above. They do not reach the arctic coasts
proper, either in Alaska or Mackenzie, and their eastward extensions into the
northern plains are less numerous and less pronounced. All of them range
southward into the western United States. They fall rather clearly into two
groups: one with a strong coastal relationship and the other just as strongly
continental.

= S J
. sN
IY way.
\ ~ v
ae >
tae Hs
|
TPS
~ ii x
a *
A 4 } ¥
Ny ;
cute RY J
hehe aah S|
’ 1 =e ~

Fic, 10. Cordilleran species.

Sixteen of the 27 follow the coastal mountains from the Alaska Peninsula to
western Washington, although a couple of them retreat inland a short distance
in the latter region. They extend throughout the northern Rocky Mountains, but
most of them are confined to the southern ranges of Alaska. None of them
reach the lower Yukon valley (Fig. 10-a), although two have been found at the
Mackenzie River delta. Our Mackenzie Mountain collections have constituted
northeastern range extensions for most of these species.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 71

Abies lasiocarpa, Carex nardina var. Hepburnii, Carex phaeocephala, Carex pyrenaica,
Juncus Drummondt, Salix Barclayi, Salix commutata, Salix Scouleriana s. 1., Ranunculus
Eschscholtzti, Draba nivalis var. elongata, Parnassia fimbriata, Saxifraga Lyallii, Hip-
puris montana, Phyllodoce glanduliflora, Senecio triangularis, Hieracium gracile.

The other 11 cordilleran plants avoid the coastal strip except in the south,
where four of them appear in the Puget Sound region. A few of them (4)
reach central Alaska, and two are found at the Mackenzie delta (Fig. 10-b),
This group contains the very few Rocky Mountain plants which have their known
northern limits in the Mackenzie Mountains.

Poa Buckleyana, Carex albo-nigra, Carex atrata ssp. atrosquama, Salix Barrattiana,

Draba McCallae, Draba praealta, Potentilla diversifolia s. 1., Erigeron jucundus, Anten-
naria media, Arnica mollis, Arnica alpina ssp. tomentosa.

Discussion

TABLE 4 gives a summary of the range patterns just described. Some gen-
eralizations from these figures will serve to bring out further the regional char-
acteristics and affinities of the Brintnell Lake flora. Fully 70%, or about 199
of the species, have wide ranges across the continent, most of them extending

TABLE 4.
No. of
spp. and
vars. Per cent

Endemics and plants of limited known range 12 4,2
Cosmopolitan in Canada and Alaska 3 1.0
Wide-ranging forest species

Reaching coasts of Alaska, B. C., and western Washington 38 13.4

Reaching coasts of southeastern Alaska 13 4.6

Avoiding coasts, southeastern Alaska to Washington 48 17.0
Wide-ranging arctic or arctic-alpine species

Reaching coastal mountains, Alaska to Washington 19 6.7

Reaching coastal mountains of southeastern Alaska 18 6.4

Avoiding coastal mountains, S. E. Alaska to Washington 29 10.2
Wide-ranging ‘‘timber-line”’ species

Reaching coast of S. E. Alaska 10 Pes)

Avoiding coasts, S. E. Alaska to Washington 21 7.4
Alaskan-cordilleran species

Reaching coasts, Alaska to Washington 11 3.9

Reaching coasts of southern and southeastern Alaska 15 3.3

Avoiding coasts, S. E. Alaska to Washington 19 6.7
Cordilleran species

Reaching coasts, southern Alaska to Washington 16 5.6

Avoiding coas*s, southern Alaska to Washington 11 3.9

Totals 283 99.8

from Alaska to the Gulf of St. Lawrence. Approximately 25.4% are Alaskan
or cordilleran, while the remaining 4.2% are endemic or local.

The wide-ranging plants fall into two main groups, arctic-alpine and forest
species. In the first there are 66 species, while in the second there are 99,
A third group of 31 species (10.9%) are intermediate between the first two,

72 SARGENTIA [6

and three (1%) are cosmopolitan. This is not a complete statement of the
proportions of arctic-alpine and forest elements, however, because approximately
62 of the 72 Alaskan and cordilleran plants are of prevailingly alpine or arctic
affinity and should be added to the wide-ranging ones. Thus the total of arctic,
arctic-alpine, and alpine plants is about 128, or about 45% of the total vascular
flora, and that of forest species is 109, or about 38% of the whole.

Although the flora as a whole is strongly continental in character, a great
many of its species are able to live in the damp forests of the north Pacific
slopes or on the coastal mountains. The range maps give a rough index to the
degree of continentality which is achieved by the Brintnell Lake flora, a fact
worthy of note. There are about 140 species (49.4%) which avoid the coastal
forests and mountains completely or nearly so, and are the most rigorously
continental plants if our flora so far as the west is concerned. Another 56
(19.8%) avoid the British Columbia and Washington coasts, but occur in south-
eastern Alaska. The remaining 87 (30.7%) inhabit, in addition to the con-
tinental ranges, the coastal strip from Alaska to Washington.

Another figure of some interest for evaluating the floristic affinities of our
area is the number of plants that do not reach the western coasts of Alaska.
There are approximately 86 of these, or about 30% of the flora.

TABLE 5 will summarize the above generalizations.

TABLE 5.
No. of
spp. and
vars. Per cent
Endemics and plants of limited known range 12 4.2
Cosmopolitan species 3 1.0
Plants of wide range across the continent 199 70.3
Plants with ranges mainly in Alaska and the Cordillera 72 25.4
Plants of the forested country 109 38.0
Arctic, arctic-alpine, or alpine species 128 45.0
““Timber-line”’ ranges 31 10.9
Strictly continental with regard to S. E. Alaska, B. C., and
Washington 140 49.4
Plants that avoid the coasts of B. C. and Washington, but
occur in S, E, Alaska 56 19.8
Plants whose ranges extend into the coastal strip of S. E.
Alaska, B. C., and Washington 87 30.7
Plants that do not reach the western coasts of Alaska 86 30.0

ALPINE PLANTS BELOW TIMBER LINE

Among the 283 species and varieties of vascular plants found in the Brintnell
Lake area it is possible to make the following subdivisions based upon their
relation to timber lines.

Lower altitudinal limits in subalpine scrub 62 21.9%
Confined to the subalpine scrub 6 2.1%
Upper altitudinal limits in subalpine scrub 131 46.3%

Widespread above and below timber line 84 29.7%

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 73

Of the plants confined to areas above timber line nearly all prove to have
definitely arctic or alpine ranges in the boreal part of the continent. Most of
them are but rarely found in the northern fringes of the forest. Of the 131
species not found above the subalpine scrub, however, about 20 also belong in
the category of arctic-alpine species. These plants probably should be in the
fourth group given above, and were not seen above timber line at Brintnell Lake
because suitable habitats were not available in the areas observed by our party.
Many of them are plants of wet mossy tundra which is rare or nonexistent on
the dry alpine slopes around the lake. Two other species in the lower slope
group also should not be considered characteristic. These are Carex aquatilis
and Carex capillaris, both of which are ‘“‘wides,” almost universally distributed
throughout the forest and the tundra. The total number of species that can
be considerec. characteristic of the slopes and bottoms below timber line is
therefore reduced to 109.

It is of interest to examine further the ranges of the 84 species that appear
to cross the timber line with impunity. Two of them are obviously wides:
Cystopteris fragilis and Equisetum arvense. Another 20 can be classed as be-
longing to the boreal forest regions. They appear to reach their greatest con-
centration within the forest zone either in the west or extending across the
continent. Thus they can be looked upon as invaders of the alpine areas from
below. Needless to say, most of them are not found far above the subalpine
scrub. They may be listed as follows: Lycopodium complanatum, Juniperus
communis vat. montana, Zygadenus elegans, Betula glandulosa, Delphinium
glaucum, Arabis lyrata var. kamchatica, Draba lanceolata, Dryas Drummondu,
Potentilla fruticosa, Rosa acicularis, Hedysarum alpinum var. americanum,
Shepherdia canadensis, Epilobium angustifolium, Pyrola asarifolia var. incar-
nata, Pyrola secunda, Ledum groenlandicum, Vaccinium Vitis-Idaea var. minus,
Mertensia paniculata, Linnaea borealis var. americana, Aster sibiricus.

Fifty species appear to be derived from the arctic and alpine elements of the
flora: Dryopteris fragrans, Equisetum variegatum, Agropyron latiglume, Arcta-
grostis arundinacea, Calamagrostis purpurascens, Festuca altaica, Festuca brachy-
phylla, Poa glauca, Poa alpina, Poa arctica, Poa paucispicula, Trisetum spicatum
var. Maidenii, Carex atrata ssp. atrosquama, Carex pyrenaica, Carex scirpoidea,
Luzula arcuata, Salix alaxensis, Salix glauca var. perstipula, Salix reticulata,
Salix Richardsonii, Oxyria digyna, Polygonum viviparum, Arenarta verna var.
pubescens, Silene acaulis var. exscapa, Stellaria longipes var. Edwardsu, Aconi-
tum delphinifolium, Cardamine bellidifolia, Draba longipes, Draba McCallae,
Draba praealta, Saxifraga aizoides, Saxifraga tricuspidata, Dryas integrifolia,
Astragalus alpinus, Astragalus frigidus var. littoralis, Lupinus arcticus, Oxy-
tropis hyperborea, Empetrum nigrum, Epilobium latifolium, Cassiope tetragona,
Polemonium acutiflorum, Pedicularis capitata, Campanula lasiocarpa, Anten-
naria monocephala, Artemisia arctica, Arnica alpina, Arnica Snyderi, Senecio
lugens, Crepis nana, Taraxacum alaskanum.,

Finally there are 12 species from the group that I have designated as “timber
line species”: Carex deflexa, Tofieldia palustris, Anemone parviflora, Anemone
Richardsonii, Parnassia Kotzebuei, Sibbaldia procumbens, Oxytropis ixodes,
Hedysarum Mackensii, Arctostaphylos rubra, Gentiana propinqua, Pedicularis
labradorica, Solidago multiradiata.

Five of the 6 plants confined to the scrub appear to be more nearly related
to the arctic and alpine elements than to those of the forests. The sixth, Cornus

74 SARGENTIA [6

canadensis f. purpurascens, is merely a color form of the common bunchberry of
the woods. '

In the preceding analysis of floristic affinities, a division is made of all the
flora of the Brintnell Lake area into three groups exclusive of local plants and
“wides”: plants of boreal forests, plants of arctic-alpine affinities, and a group
of “timber line species.” The total numbers in these categories are 109, 128,
and 31, respectively. Of all the species of boreal forest affinities, therefore, 20,
or about 18.3%, are able to extend their local ranges above timber line; and as
previously noted most of them do not reach far above the scrub. On the other
hand 50 species of arctic-alpine affinity, or about 39% of the total, are found
both above and below timber line. Nearly all of them are found at lake level.
If the 20 alpine forms found around the lake but not above timber were to be
added to this group the percentage would be still greater, about 54.7%. As
would be expected, the percentage among the ‘‘timber line” species is also rela-
tively high: 38.7%.

It appears, therefore, that the timber line is a far more significant boundary
for boreal forest plants than for arctic-alpine ones. There is the suggestion,
further, that the forest is intrusive at Brintnell Lake, having invaded an alpine
tundra whose remnants are still to be found at all levels in the vicinity.

Most of the alpine species at lake level are found immediately on the shores
of the lake, either on the gravel beaches of the fans or in local muskeggy spots
at or just above the level of the lake. These are the most rigorous, unstable
habitats at the lower altitudes, due to stream erosion and deposition, wave action,
ice push, and frost heaving. They are the areas which the slope and lowland
forests have not yet been able to occupy, and they could well serve as relic areas
for a tundra flora.

APPLICATION OF HULTEN’s HypoTrHEsEes TO THE BRINTNELL LAKE FLORA

The flora of the Mackenzie Mountains, so far as it is at present known, should
be examined and studied in the light of Dr. Eric Hultén’s thesis (1937) con-
cerning the origin and dispersal of all arctic and boreal floras. The geographic
position of the Mackenzie Mountains in northwestern America, their recent
botanical exploration, the apparent partial isolation of their alpine flora during
post-Glacial time, and the simplicity and apparently incipient endemism in their
flora all suggest that this flora might be used as a test of Hultén’s thesis. In
the following discussions it must be borne constantly in mind that I am treating
only one phase of the Mackenzie Mountain flora, that in the vicinity of Brintnell
Lake. Mr. Porsild’s collections of 1944 on the westerly slopes of the range
show a much larger assemblage of plants, whose geographic affinities will have
to be studied before a balance can be struck for the whole region.

It is unnecessary to review in detail all of the reasoning and implications of
the theories advanced by Hultén, for this has already been done by several stu-
dents (see Stebbins, 1942; Raup, 1941; Halliday & Brown, 1943; Cain, 1944),
The following is a brief outline of the broader geographic aspects, with a few
critical notes that have grown out of the present investigation.

In connection with his floristic studies of Kamtchatka and the Aleutian Islands
Hultén mapped, sometimes in detail and sometimes by limits, the ranges of
hundreds of arctic and boreal species. He organized this factual material on
the basis of what he called “equiformal areas.” That is, when large numbers of

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 75

ranges are superposed they fall into a group of patterns which are more or less
clearly defined geographically, and “equiformal” within themselves. Each equi-
formal area shows a region of concentration in number of species, which is
called its “centre.” Since it is assumed that each species has acquired its
present range by dispersal from a point of origin or survival, the regions of con-
centration within the equiformal areas are regarded as fundamental centers of
origin for the various major elements in the flora; and the “equiformal areas”
become “equiformal progressive areas” that are thought to indicate the general
patterns of dispersal among the major elements. The species are termed “‘ra-
diants” from the various “centra.”

The geographic arrangement of centra as worked out by Hultén is as follows:
They are “. . . in North-Eastern Siberia and in the Amur-Manchurian region.
Another occurs in the Altai-Sajan region, sending out radiants towards the
Arctic shore. A third centre is northern Japan, whence numerous plants radiate
to the north and to the coast of the Asiatic Continent. A centre of great im-
portance is the region around the northern part of the Bering Sea. It sends
out progressive radiants reaching symmetrically as well to the west into arctic
Asia and Europe as to the east to Eastern America, and also often extends arms
along both the Asiatic and American Pacific coast. In America radiants pro-
ceed from the Yukon valley along the Arctic American coast, others centre
around the Arctic Archipelago, and others again have the centre of their pro-
gressive figures in the State of Washington and radiate along the American
coast or along the Rocky Mts. to Alaska. Of the plants discussed . .. no
groups could be formed having their centres in northern Europe or western
Siberia, or in North-Eastern America or in the country between Yukon Valley
and the Great Lakes” (Hultén, 1937, p. 25).

Hultén immediately draws a correlation between the distribution of his centra
and the distribution of ice during the Pleistocene. No centra could be found
in areas that were covered by ice during the maximum advance of the glaciers ;
and the existing floras have all been derived from areas of refuge close to the
ice. It is maintained, on genetical grounds, that the ability of the species to
disperse themselves from their refugia has not been uniform. Those confined to
small refugia, under difficult climatic conditions and in small populations, are
considered to have been so depauperated of biotypes that they have been exceed-
ingly slow to spread. Others had large areas and populations, either within the
generally glaciated regions or south of the ice, so that they retained their inherent
variability and aggressiveness and could quickly invade lands freed of ice.

It is thought that the “Linnaean species” of the present boreal flora originated
in the last great interglacial or earlier, and that large numbers of them achieved
wide dispersals during that time. Their present areas are looked upon as reduc-
tions from these wide ranges, with post-Glacial re-expansions that have been
conditioned by the amount of depauperation suffered during the maximum ice
advance.

If the general thesis outlined by Hultén is tenable, then the flora of the
Mackenzie Mountains should show affinities with his various centra which,
within the general limits of accessibility, are directly proportional to the size of
the relic populations, and presumably inversely proportional to their degree of
biotype depauperation.

Before proceeding with such an analysis of the Brintnell Lake flora it will be
well to examine critically the disposition made by Hultén of certain species, and

76 SARGENTIA [6

to make some additions to his lists. It is impossible in the present study to
make a complete critical revision of the equiformal progressive areas, for to do
so would require the remapping of all the species’ ranges. Only such notes will
be presented as have appeared in the investigation of the Brintnell Lake plants.

Hultén’s arrangement of equiformal areas is open to modification or criticism
along three lines. First, it is possible to make additions to his lists, at least
in the American boreal flora. The Brintnell Lake region is found to have ap-
proximately 45 species not discussed by him at all. When these are sorted into
patterns they have a distribution among the equiformal areas, however, that
makes no serious modification in the latter. Second, there are apparent gaps in
the ranges of boreal American plants which are gaps in exploration rather than
in the actual ranges. Lacunae in our knowledge of the more uniform floras of
eastern glaciated regions are not so troublesome ; but the gaps in northern British
Columbia and Yukon are more serious, for the distribution of species into equi-
formal areas sometimes depends upon their behavior in this region. Some of
Hultén’s dispositions of species are therefore open to modification due to range
extensions discovered since his study was made. Third, Hultén’s original sort-
ing of species among his equiformal areas can be questioned in many cases. This
is particularly notable in his treatment of wide-ranging forms.

Although suggestions along these three lines will be made in the discussions
that follow, it will be noted that only a few actual changes in Hultén’s lists have
been made, other’than by the addition of species not treated by him. Such
changes as may be made are in many cases matters of judgment, and even when
all are taken.together they probably will make no serious modification of the
structure. For the purposes of the present paper therefore, and until the gen-
eral level of definitiveness in the whole structure is raised, it has seemed best
to leave many of the doubtful ranges where Hultén placed them.

Southern Beringia radiants

Two groups of equiformal areas are centered in the Beringian region. First
to be considered is in southern Beringia. It extends both east and west from
the Aleutian chain—to Kamtchatka and the Kuriles on one hand and southward
along the Pacific coast of America to the United States on the other. Hultén
lists 77 species and varieties here, some of which are of wide range in the area,
but many of which show discontinuous and widely separated stations. They
are thought to have occupied small, isolated refugia during the Pleistocene, and
to have been badly depauperated of biotypes. Furthermore they are in general
oceanic and coast-bound, though a few penetrate inland in Alaska and eastern
Siberia. The Mackenzie Mountains are isolated from them not only because
of the suspected inability of the plants to migrate, but also because of their
obvious preference for coastal habitats. Only one of the 77 species was found
at Brintnell Lake, Carex nesophila; but in view of its recent discovery also in
the Richardson Mountains and in the central Yukon region it probably belongs
among the northern Beringian groups rather than among the southern, more
oceanic ones.

“ Northern Beringia radiants

Plants whose equiformal areas are thought to center in Northern Beringia
are discussed by Hultén in four groups: species with ranges principally (1)

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 77

American, (2) Asiatic, (3) bilateral across Northern Beringia, and (4) having
extensions southward to and along the north Pacific coasts. We are concerned
here only with those that appear in America.

Approximately 114 species are noted in the first of these four categories, and
31 of them have been listed at Brintnell Lake. They are treated by Hultén in
three groups: first a “rigid” group which is predominantly arctic; second, a
“plastic” group that is more southern ; and finally a very plastic group with wide
ranges in the subarctic. Hultén believes that these three groups had somewhat
different histories during and after the Pleistocene, depending upon their orig-
inal habitat preferences. The first lived in the more northern, arctic parts of
Beringia where they were greatly reduced by the maximum ice. They include

some of our “most arctic” species. The second are thought to be “. . . less
arctic and not montane. . . . They do not occur in the Arctic Archipelago and,
except Beringia, they occupy chiefly glaciated areas. . . . They thus possess a

”

much more plastic area than the former group... .” The third are thought
to have “. . . covered so extensive an area on the D-interglacial that they had
reached the southern boundary-line of the maximal ice before the maximal
glaciation.” With such large undepauperated populations to draw from they
have been among the most successful plants in filling the gaps made by the
glaciers.

The equiformal figures made from the ranges of the rigid and plastic groups
appear to be fairly consistent, with a few exceptions. We find among the rigid
species Saxifraga tricuspidata and Dryas integrifolta, both of.which are widely
distributed in the northern Rocky Mountains as well as in the Arctic. Hulten
notes that they appear to have been especially successful at spreading in spite
of their supposed depauperation. It seems more reasonable to place them
among his Arctic-Montane group (see below). Among his plastic species, so
long as they are confined to the far northwest, the figures appear reasonable ;
but when species with wider ranges are considered many are found to be nearly
identical with his Continental Western American radiants. Such plants are
Betula papyrifera var. humilis, Salix arbusculoides, Carex physocarpa, Parnassia
Kotzebuei, Cypripedium passerinum, and Solidago multiradiata. Finally it is
difficult to see why three of the primary species of the Canadian forest, Picea
glauca, P. mariana, and Larix laricina s. 1., along with several of the more char-
acteristic species of large sections of this forest region * should be placed among
the Northern Beringian radiants. Their ranges are nearly or quite identical
with many other forest species that are classified among the Continental Western
American radiants, and there is no clear reason to think they have had a dif-
ferent history. If there is evidence that a species had access to the larger
refugia during the Pleistocene, it seems unnecessary and somewhat misleading to
attach it to an equiformal area centering on one of the lesser refugia.

With these things in mind I have altered Hultén’s lists by reclassifying the
wide-ranging plants above mentioned. The resulting total list of Northern
Beringian radiants is thereby reduced to approximately 90, of which 15 have
been found at Brintnell Lake. The 15 species are as follows:

2 Mertensia paniculata, Geum macrophyllum var. perincisum, Aquilegia brevistyla, Ribes
hudsonianum, Agropyron dasystachyon, Festuca saximontana, Lusula saltuensis, Ribes glan-
dulosum, Impatiens biflora, Cornus stolonifera, Orchis rotundifolia, Primula mistassinica,
Corydalis sempervirens.

78 SARGENTIA [6

Rigid group:

Taraxacum alaskanum Salix Richardsonii
Saxifraga radiata Salix alaxensis
Astragalus frigidus var. littoralis Oxytropis Maydelliana
Oxytropis pygmaea . Carex membranacea
Lupinus arcticus Taraxacum lacerum
Plastic group:
Carex podocarpa Poa paucispicula
Arctagrostis arundinacea Polemonium acutiflorum

Larix laricina ssp. alaskensis

The second main group of Northern Beringia radiants with which we are
concerned are those called by Hultén the Bilateral Radiants of Northern Beringia.
There are 45 of these, including one (Erigeron eriocephalus) that I have added.
Five of the 45 are found at Brintnell Lake. Reference to Hultén’s map (PI.
13) and discussion (pp. 57-60) indicates that this group is analogous to his
American and Asiatic radiants from Northern Beringia, and differs from them
chiefly in extending on both sides of Bering Strait. The most widespread of
them become the high arctic circumpolar species. Many of them now show gaps
in their ranges, but Hultén thinks they were probably all completely circumpolar
in the D-interglacial. With them he groups the so-called “amphiAtlantic” and
“west-arctic” species, such as Saxifraga aizoides and Arenaria humifusa, whose
Bering Straits-eastern Asiatic components are thought to have been destroyed
by the last ice. The question naturally arises as to whether this is justifiable if
the Beringian area is to be looked upon as the most important refuge available
to such plants. Recent collections, however, particularly in the Rocky Moun-
tains, suggest that it may not be necessary, at least in America, to relate them
to Beringia. Furthermore, Arenaria humifusa is now reported from there.

The five species found in the Mackenzie Mountains are Arenaria humifusa,
Saxifraga aizoides, Potentilla emarginata, Pyrola grandiflora, and Erigeron
eriocephalus. A glance at their ranges shows that all but the last have now
been found in the northern Rocky Mountains, so that it probably is more logical
to consider all but the Erigeron with the Arctic-Montane radiants (see below)
rather than with the Northern Beringia radiants. This group of plants brings
out the significance of the gap in the botanical exploration of northern British
Columbia.

The third group of Northern Beringia radiants represented in America is
called “Arctic—Pacific.” It is characterized by Hultén as “. . . closely related
to that one [arctic circumpolar plants] and with exactly the same history, the
only difference being that it includes those arctic circumpolar plants that have
also reached the southern shore of Beringia. . . . (It should be remembered
that all arctic plants are to be regarded as more or less oceanic, as they are con-
fined to a strip along the shore of an ocean).” He lists 41 species here, of
which 11 have been found at Brintnell Lake. The latter are as follows: Poa
glauca, Luzula Wahlenbergii, Salix arctica s. 1., Salix pulchra, Aconitum del-
phinifolium, Anemone Richardsonii, Ranunculus nivalis, Draba cinerea, Pingui-
cula villosa, Antennaria monocephala, Arnica Lessingii.

I find it difficult to place all of these 11 species in any single category of
ranges, and many of them do not seem to fit the conditions set up by Hultén
for his Arctic—Pacific group. Antennaria monocephala and Aconitum delphini-
folium are strongly represented in southern Beringia; but they are scarcely

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 79

arctic in Alaska, and in the eastern part of their ranges they are decidedly con-
tinental. The same can be said of Anemone Richardsonii except that it does
have a somewhat more arctic extension. It is represented in the Arctic Archi-
pelago, however, only in the western part. Poa glauca is a decidedly continental
species except in the Arctic, and it appears to be only slightly represented on
the north Pacific coasts. All four of these could as well be placed with the
Continental western American radiants, or the last two with the Arctic-Montane
group. Poa glauca, a highly polymorphic species, is actually one of our most
widely dispersed boreal grasses. Except for the North Pacific coast, it could
be grouped, so far as boreal America is concerned, with such wides as Cystop-
teris fragilis and Equisetum arvense.

Lack of clarity in the taxonomy of arctic Draba makes it impossible for me
to know what Hultén had in mind for D. cinerea, but as the species is mapped
here it can hardly be related to his Arctic—Pacific group. It may belong with
the Arctic-Montane plants.

The realignments noted above reduce Hultén’s original list to 36 species, and
they reduce the Brintnell Lake representatives to 6. Two of the 6 remain some-
what doubtful. The northern and eastern ranges of Luzula Wahlenbergu are
probably not complete because it has not always been clearly distinguished from
related species, but as mapped here it is not a conspicuously arctic plant.
Pinguicula villosa is likewise not notably arctic, but is rather a “timber line
species.”

The “Atlantic-Pacific” species of Hultén’s classification are for the most part
strongly oceanic in habitat preference, and are not represented at Brintnell Lake
at all.

Western American coast radiants

Hultén lists 263 species as having a strong Pacific coast affinity in America.
I have added 4 more, making a total of 267. Of these, 23 have been found at
Brintnell Lake.

Hultén’s list is divided into four groups which are mapped separately (Pls.
16-19). The first two contain plants that are distinctly western American in
the sense that they do not reach the eastern coasts, while the last two comprise
species that have continuous ranges across the continent or have separate eastern
and western components. The selection of species for these last two groups
appears to be somewhat arbitrary, for it involves drawing plants from the great
mass of subarctic wides, as it did in the case of the Northern Beringia Radiants.
One could place most of these species just as well in the Western Continental
Radiants or group them with the Boreal Circumpolar wides.

The first two groups, however, can be related to the Pacific coasts. They are
divided into those that spread widely from Southern Beringia southeastward
into the United States, and those that do not reach Bering Sea at all, but have
an extensive range in the north Pacific area. Of the former group Hultén
thinks “. . . that they occupied the southern coast of Beringia in the early part
of the D-interglacial and radiated from there towards California, some few also
towards Yezo. They had reached the southern boundary of the ice before the
maximum glaciation, which they survived on the one hand in Beringia and on
the other hand south of the ice.” Concerning the latter group he states that it
“|. is naturally very large, as it comprises those plants that lived south of
the ice during the maximum glaciation and then progressed along the coast

80 SARGENTIA [6

towards Alaska when the ice-sheet retreated. They can have migrated parallel
to the southern edge of the ice during the glacial. Probably the coast was among
the first areas to be free from ice and as furthermore there must have existed a
dry continental shelf, although a narrow one, the plants south of the ice must
have had a good opportunity, without competition from other plants, of advanc-
ing northward along the coast.”

Hultén’s figures for the two groups just mentioned have been modified not
only by the addition of 2 species, but also by changing 2 species from the second
to the first. Draba nivalis var. elongata (D. lonchocarpa) and Phyllodoce glan-
duliflora have now been found on the Beringian coasts. The two species added
are Carex pyrenaica and Cryptogramma crispa var. sitchensis. The first was
taken from Hultén’s Arctic-Montane Radiants.

The plants represented in Hultén’s Plates 18 and 19, with their extensive
ranges across America, are thought to have survived the last ice “. . . in an
unbroken zone along the southern margin of the ice,” and to illustrate the great
spreading capacity of plants not greatly depauperated of biotypes.

The 23 species of West American Coast radiants represented at Brintnell
Lake are as follows:

Ranunculus Eschscholtsii Salix Scouleriana
Salix Barclayi Salix commutata
Juncus Drummondit Senecio triangularis
Saxifraga Lyallii Parnassia fimbriata
Hieractwm gracile Carex phaeocephala
Hippuris montana Carex leptalea
Draba nivalis var. elongata Heracleum lanatum
Arabis lyrata var. kamchatica Cornus canadensis
Cryptogramma crispa var. sitchensis Agrostis scabra
Phyllodoce glanduliflora Habenaria hyperborea
Carex pyrenaica Alnus crispa

Viburnum edule

Continental western American radiants

This group is looked upon as more or less parallel, historically, to the West
Coast Radiants, differing chiefly in being strongly continental. It is broken up
into two main groups, “. . . those that do not reach the Bering Sea but stop
in the Yukon valley, and those that reach Beringia. The former group appar-
ently survived the maximum glaciation in the upper Yukon valley... or in
the mountains south of it, the latter group also possessed stations in Beringia
during that period.”

Hultén lists 158 species as Continental West American Radiants. The ranges
of 138 of them are presented in his Plates 20, 21, and 22, while the other 20
are given similar histories although they are now thought to show gaps in their
ranges. On the basis of Brintnell Lake studies I have added 19 species not
discussed by Hultén. It is now possible to alter Hultén’s subdivisions of the
group to a certain extent. Salix myrtillifolia, Salix Barrattiana, Salix Bebbiana,
Actaea rubra, and Carex concinna all extend farther west in Alaska than he
indicates; and Populus tremuloides, P. Tacamahacca, and Betula glandulosa
(incl. B. sibirica) probably reach Bering Sea. The following are listed as not
reaching the eastern American coasts, although they all do so: Salix myrtillifolia,
Listera borealis, Salix Bebbiana, Actaea rubra, Agropyron trachycaulum var.
unilaterale, Carex concinna.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 81

I have also added 13 species by reclassifying plants listed by Hultén in other
categories. One of these, Arabis Drummondii, is taken from his Pacific Coast
radiants, but the remainder are from his Northern Beringian elements (see

above). The latter are as follows:

Betula papyrifera var. humilis
Salix arbusculoides
Cypripedium passerinum
Solidago multiradiata
Parnassia Kotzebuei
Aquilegia brevistyla

Mertensia paniculata
Ribes hudsonianum
Ribes glandulosum
Cornus stolonifera
Picea glauca

Picea mariana

Including the additions noted above there are 190 species in Hultén’s Conti-
nental West American group. Of these, 73 have been found at Brintnell Lake.
No attempt will be made to subdivide them according to Hultén’s scheme, since
this would involve the reclassification of a number of his ranges. In the fol-
lowing list, however, the 73 species are arranged, in general, according to his

sequence.

Salix myrtillifolia

Salix Barrattiana

Delphinium glaucum

Listera borealis

Arctostaphylos Uva-ursi var. adeno-
tricha

Arnica mollis

Abies lasiocarpa

Alnus tenutfolia

Actaea rubra

Agropyron trachycaulum var. unilaterale

Geum macrophyllum var. perincisum

Carex concinna

Fragaria glauca

Carex albo-nigra

Carex atrata ssp. atrosquama

Antennaria media

Draba praealta

Potentilla diversifolia s. 1.

Draba McCallae

Erigeron jucundus

Poa Buckleyana

Ribes hudsonianum

Aquilegia brevistyla

Betula papyrifera var. humilis

Populus tremuloides

Shepherdia canadensis

Ribes lacustre

Populus Tacamahacca

Carex aurea

Carex Garberi var. bifaria

Betula papyrifera var. commutata

Carex deflexa

Geocaulon lividum

Ledum groenlandicum

Betula glandulosa

Smuilacina stellata

Erigeron elatus

Senecio pauperculus

Amelanchier florida

Senecio indecorus

Agropyron trachycaulum var, novae-
angliae

Dryas Drummondii

Arenaria dawsonensis

Arabis Holboellii

Ribes oxyacanthoides

Arnica alpina ssp. tomentosa

Cornus stolonifera

Ribes glandulosum

Arabis Drummondii

Senecio lugens

Gentiana propinqua

Zygadenus elegans

Rubus acaulis

Habenaria obtusata

Ranunculus Gmelini var. Purshti

Hedysarum alpinum var. americanum

Salix Bebbiana

Ribes triste

Carex gynocrates

Pyrola asarifolia var. incarnata

Viola epipsila

Carex physocarpa

Draba longipes

Picea glauca

Picea mariana

Mertensia paniculata

Parnassia Kotzebuei

Solidago multiradiata

Cypripedium passerinum

Salix arbusculoides

Gentiana glauca

Campanula lasiocarpa

Boschniakia rossica

Hultén’s selection of plants for this list can be criticized at several points.
Many of the more wide-ranging species could with equal propriety be placed

82 SARGENTIA [6

with his Arctic-Montane group, particularly with those that do not possess an
arctic branch. Others are not strictly continental in the west, and could be
classified with the Pacific Coast Radiants. The following might be placed in
the latter category: Actaea rubra, Ribes lacustre, Ledum groenlandicum, Pyrola
asarifolia var. incarnata, Viola epipsila, Carex physocarpa, Campanula lasio-
carpa, Boschniakia rossica.

Arctic-montane plants

Of all the groups set up by Hultén, that of Arctic-Montane plants is probably
the most complex. The maps given in his Plates 27-30 cover the species rep-
resented in America, and involve 144 plants. An additional 56 species are
considered to have had their ranges derived from similar patterns, but they are
not included in the maps.

The group as a whole is perhaps best characterized in Hultén’s own words:
“The species belonging to this large and interesting group differ distinctly in
one specific property from those previously treated. ... They possess the
capacity for living not only on lowlands of arctic or subarctic character but also
on mountains in far more southerly districts. . . . They are thus either mountain
plants or arctic plants. The peculiar point about the group . . . is that it com-
bines both these specialties and that its plants are distributed as well along the
Arctic Shore as on the mountains. Some of them can even stand the condi-
tions along the northern Pacific shore and protrude considerably even there.”
It is thought that the broad range of habitats used by these plants enabled them
to migrate during the Pleistocene, and that consequently they are an arctic-
alpine group that was not so much depauperated of biotypes as were other
groups whose habitat selection was not so wide.

Hultén subdivides the American Arctic—Montane plants into four groups,
as follows: (1) those thought to have had their montane element eliminated by
the Pleistocene, (2) those with a montane and western arctic development in
America but with their largest areas in Eurasia, (3) those having both arctic
and montane branches but not extending to the Gulf of St. Lawrence region,
and (4) a group similar to the last but reaching the Gulf of St. Lawrence. The
disposition of plants among these four groups is open to question. For instance,
47 species are listed in group (1), of which 14 were found at Brintnell Lake.
Of the latter, two (Pedicularis labradorica, Arctostaphylos rubra) are scarcely
arctic. The first appears in the Archipelago only in southern Baffin Island, and
does not go north of Disco Bay in Greenland. The second has not been found
either in the Archipelago or Greenland. All of the 14 occur in the Rocky
Mountains, and 8 of them south at least as far as the Jasper Park district. None
of the 14 should be in Hultén’s group (1), therefore. In group (3), Poa
arctica reaches the Gulf of St. Lawrence, and Luzula arcuata has no arctic
branch at all. In group (4) Epilobium davuricum and Eriophorum Scheuch-
geri do not show montane branches in the British Columbia and Alberta Rockies.
It should be pointed out further that a number of the plants listed in group (4)
are not conspicuously arctic, especially in the east. These are as follows:
Betula occidentalis, Taraxacum ceratophorum, Veronica alpina var. unalasch-
censis, Carex media, Salix planifolia, Valeriana septentrionalis, Oxytropis folio-
losa. Some of these are definitely limited to the forested regions: Betula
occidentalis, Carex media, Valeriana septentrionalis.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 83

A notable source of error in Hultén’s selections may be discussed here, though
it also appears in other parts of his paper. He includes in his group } plants
thought to have had their montane elements destroyed during the Late Wisconsin
period. Such plants, however, lacking montane extensions, become eligible to
classification as northern Beringian Radiants. If this reasoning were carried
to its logical conclusion most of the more plastic northern Beringian plants,
those with wide arctic ranges, could be taken out of that category and placed
with the arctic-montane group. The only real criteria upon which we can base
judgments in these complicated problems are the actual existing positions and
sizes of the ranges of the plants, and their degree of specific segregation. From
these we can draw inferences as to the geographic preferences and spreading
capacities of the plants. If an arctic plant fails to appear in the Rocky Moun-
tains it seems to me more logical to assume either that it is unable to migrate
there or has not yet had time to, than to class it with other Arctic-Montane plants
which have achieved the wider range. This is particularly true since the
Arctic-Montane group as a whole is thought to have suffered much less de-
pauperation during Late Wisconsin time than the Beringian groups.

In the course of this study I have made several changes in Hultén’s lists of
Arctic-Montane plants. Twelve species and varieties found in the Mackenzie
Mountains were not discussed by him, but seem most nearly related to this
series. Twenty other species have been taken from four of his other equiformal
groups and added here because their areas appear more nearly to fit the Arctic—
Montane pattern. These are as follows: from Northern Beringia Radiants—
Dryas integrifolia, Saxifraga tricuspidata; from Bilateral Radiants of Northern
Beringia—Arenaria humifusa, Saxifraga aizoides, Potentilla emarginata, Pyrola
grandiflora, Taraxacum lapponicum, Arenaria sajanensis, Sagina Linnaei; from
Arctic—Pacific Radiants—Draba cinerea, Antennaria monocephala, Anemone
Richardsonii, Poa glauca, Aconitum delphinifolium; from Boreal—Circumpolar
plants—Petasites frigidus, Aster. sibiricus, Chrysosplenium tetrandrum, Carex
microglochin, Calamagrostis canadensis var. Langsdorfi, Lycopodium Selago,
Cardamine pratensis, Calamagrostis lapponica.

The total amended list of Arctic-Montane plants now contains 235 species,
of which 100 have been found at Brintnell Lake. The latter are as follows.
Although they are not broken down into the 4 subdivisions made by Hultén, the
sequence is approximately that proposed by him.

Festuca altaica : Saxifraga punctata
Sedum roseum var. integrifolium Artemisia arctica

Salix reticulata Myosotis alpestris
Pedicularis labradorica Corydalis pauciflora
Vaccinium uliginosum var. alpinum Lloydia serotina
Dryopteris fragrans Anemone narcissiflora
Hierochloé alpina Aconitum delphinifolium
Arctostaphylos rubra Antennaria monocephala
Salix glauca Artemisia Tilesti s. 1.
Lycopodium alpinum Petasites frigidus
Arenaria verna var. pubescens Aster sibiricus
Saxifraga nivalis Pedicularis capitata
Pedicularis sudetica Pedicularis lanata
Cerastium Beeringianum Cassiope tetragona
Cardamine pratensis Agropyron latiglume
Calamagrostis lapponica s, 1. Saxifraga tricuspidata

Potentilla uniflora Poa arctica

84

SARGENTIA

Lychnis apetala

Lusula arcuata

Dryas octopetala

Oxytropis hyperborea
Arenaria Rossii

Gentiana arctophila

Crepis nana

Oxytropis hudsonica
Oxytropis ixodes

Antennaria isolepis

Cornus canadensis f. purpurascens
Draba cinerea

Antennaria subviscosa
Anemone Richardsonii

Betula occidentalis
Chrysosplenium tetrandrum
Hedysarum Mackensti
Epilobium davuricum
Calamagrostis purpurascens
Stellaria longipes var. Edwardsii
Carex scirpoidea

Festuca brachyphylla
Epilobium latifolium
Cardamine_ bellidifolia
Erigeron unalaschkensis
Carex nardina var. Hepburnii
Saxifraga rivularis
Ranunculus hyperboreus
Astragalus alpinus

Taraxacum ceratophorum

Silene acaulis var. exscapa
Lusula spicata

Veronica alpina var. unalaschcensis
Carex media

Carex bipartita

Poa alpina

Polygonum viviparum
Oxyria digyna

Epilobium anagallidifolium
Carex capillaris

Juncus castaneus

Trisetum spicatum var. Maidenti
Saxifraga oppositifolia
Eriophorum Scheuchzeri
Salix planifolia

Sibbaldia procumbens
Anemone parviflora
Valeriana septentrionalis
Lusula confusa

Draba glabella

Oxytropis foliolosa
Sagina Linnaei

Arenaria sajanensis
Taraxacum lapponicum
Pyrola grandiflora
Potentilla emarginata
Saxifraga aizoides
Arenaria humifusa

Poa glauca

Dryas integrifolia

Potentilla nivea Carex microglochin
Draba fladnizensis var. heterotricha Calamagrostis canadensis var. Langs-
Tofieldia palustris dor fi

Lycopodium Selago

A few of the maps included in the present paper are at variance with Hultén’s
due to difference in species concepts. Salix glauca, for instance, is considered
by Hultén in a very broad sense, to include S. cordifolia. I have mapped it in
a more restricted sense, and have included only forms distinguished as S. glauca
by Schneider. Cassiope tetragona is mapped in its broad sense, to include
subsp. saximontana. In Draba fladnizensis, apparently considered by Hultén
in a broad sense, I have mapped only var. heterotricha.

Boreal circumpolar plants

This is the last, and one of the larger groups of plants set up by Hultén.
He has mapped 214 species, and discusses 41 others, or 255 in all. He calls it
the “boreal group” and distinguishes it as follows: “Two groups of circumpolar
plants have now been discussed, viz., the arctic and arctic-montane ones... .
It now remains to discuss the boreal group. . . . The arctic group survived the
maximum glaciation chiefly north of the maximum ice, the boreal plants were
so sensitive to the cold that they were able to survive chiefly on the refugia
south of the ice.” It is thought that one of the principal results of this is that
these plants maintained relatively large populations during the later episodes of
the Pleistocene, and were not much depauperated of biotypes. The most wide-
spread plants in the boreal world are in this group.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 85

Six plates (36-41) and a text-figure (fig. 14) are devoted by Hultén to
maps of such plants of the boreal group as occur in America. His subdivisions
of the group are based largely upon the degree of continuity of the ranges.
First are those that show no appreciable gaps in the whole circumboreal area
(Pl. 36), and then comes a series of ranges showing various kinds of gaps. No
attempt will be made here to review his disposition of species through the series ;
but it should be said that numerous changes might be made, especially in Ameri-
can ranges, due to recent exploration and the filling in of supposed gaps. For
instance 38 species are mapped in Pl. 37 to illustrate species with gaps in the
central portion of boreal America. Seven of these were found in the Mackenzie
Mountains, but none of the seven now shows an appreciable discontinuity. A
number of others in the list have been filled in in the same manner.

I have added only 7 species to Hultén’s lists, although some have been taken
out and reclassified elsewhere. The latter are Chrysosplenium tetrandrum,
Carex microglochin, Calamagrostis canadensis var. Langsdorfi, Lycopodium
Selago, Petasites frigidus, Aster sibiricus, all of which appear to be more prop-
erly placed in the Arctic-Montane group. Two of the additions have been
taken from the Arctic-Montane group and placed here because of their wide
boreal rather than arctic ranges: Epilobinm lactiflorum, Stellaria calycantha.
The other 5 additions were found in the Mackenzie Mountains but are not dis-
cussed by Hultén. One of these is the polymorphous Poa pratensis, which I
have mapped in its broadest sense.

The total amended list of Boreal Circumpolar Species is therefore of 262
plants, of which 52 have been found in the Mackenzie Mountains. The latter
are as follows:

Lycopodium clavatum var. mono- Carex diandra

stachyon Goodyera repens var. ophioides
Cystopteris fragilis Lycopodium complanatum
Equisetum arvense Poa nemoralis
Epilobium angustifolium Juniperus communis var. montana
Dryopieris Linnaeana Equisetum sylvaticum var. pauciramo-

Woodsia ilvensis

Poa pratensis s. 1.
Calamagrostis neglecta

Carex aquatilis s, 1.

Carex canescens

Corallorrhiza trifida

Parnassia palustris var. neogaea
Equisetum palustre

Equisetum fluviatile

Vaccinium microcarpum

Rubus Chamaemorus

Linnaea borealis

Potentilla norvegica

Moneses uniflora

Roripfa islandica var. microcarpa
Rubus strigosus

Vaccinium uliginosum
Empetrum nigrum

Vaccinium Vitis-Idaea var. minus
Pyrola secunda

Equisetum scirpoides

sum

Carex loliacea

Eriophorum brachyantherum
Potentilla fruticosa
Andromeda Polifolia

Rosa acicularis

Arenaria lateriflora
Calamagrostis canadensis var. robusta
Equisetum variegatum
Lycopodium annotinum s. 1.
Juncus balticus s. 1.

Pyrola minor
Arctostaphylos Uva-ursi
Pyrola virens

Epilobium lactiflorum
Stellaria calycantha
Selaginella selaginoides
Ranunculus lapponicus
Carex vaginata

Draba lanceolata

_ Erigeron angulosus var. kamtschaticus

86 SARGENTIA [6

It will be noted that in several cases I have mapped only American varieties
of the circumpolar species, but in most of them the range patterns are essen-
tially the same as those of the aggregate species.

Discussion

Before proceeding to a discussion of the origin and development of the vege-
tation of southwestern Mackenzie it will be well to compare the results of my
own survey of the geographic affinities of the Brintnell Lake flora with those
obtained from an application of Hultén’s hypotheses. In order to do this I
have examined Hultén’s disposition of species, as emended in the present paper,
in the light of their actual ranges as indicated on the maps. The accompanying
Tastes 6 and 7 show first the percentages of the Brintnell Lake plants, ar-
ranged according to Hultén, which are drawn from the various groups of actual
ranges (Figs. 4-10), and second a summary of these percentages as they are
related to the larger categories of ranges.

It is clear that Hultén’s Boreal Circumpolar group, as represented at Brint-
nell Lake (52 spp.), is prevailingly of wide continental ranges (98%). It is
derived mainly from forest species (78.8%), and if half the “timber-line” and
cosmopolitan elements are added, its forest element can be raised to a slightly
higher proportion (86.4%). About 50% is of species which extend to the
coastal strip in the west.

The Arctic-Montane plants (100 spp.) are also derived from wide-ranging
groups, but to a somewhat lesser extent than the Boreal Circumpolar plants
(82%). As would be expected, they come principally from our arctic and
alpine elements (72%), and if half the “‘timber-line” element is added this can
be raised to 82.5%. In contrast to the Boreal Circumpolar plants, about 51%
of the Arctic-Montane group is more or less strictly continental with regard to
the western coastal strip.

Among Hultén’s Continental West American plants (73 spp.) the proportion
of species with wide continental ranges falls to 68.5, leaving 31.5% to come from
Alaskan and cordilleran range patterns. They are prevailingly of forest affini-
ties (71.2%, or 73.9% if half the “timber-line” element is added). They are
more rigorously continental in the west than the Arctic~Montane group, with
64.4% failing to reach the Pacific coast.

Western American Coast Radiants represented at Brintnell Lake (23 spp.)
come principally (69.6%) from our Alaskan and cordilleran groups. Most of
them (65.2%) are species with alpine or arctic-alpine range patterns, and
nearly all of them (95.7%) occupy the coastal strip from southern Alaska to
western Washington.

Beringian Radiants (23 spp. at Brintnell L.) are mostly of western affinity
(60.9% ), and strongly arctic-alpine (91.3%, or 93.4% if half the “timber-line”
percentage is added). They are strictly continental in the west to the extent
of 65.2%, and only 1, or 4.3%, extends throughout the coastal country to
western Washington.

So far as the above comparisons go the correlation between Hultén’s arrange-
ment and my own is fairly good. They do not present a complete picture,
however, as a glance at the tables will show. We find, for instance, that while
Hultén has derived 78.8% of the Boreal Circumpolar species of the Brintnell
Lake flora from strictly forest groups, he has also derived 7 % of his Arctic—

1947]. RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 87
TABLE 6.
Conti-
iy as W. nental Arctic- Boreal
Beringia American W. Mcasanes Circum-
Radiants Coast Mdiicican |. Plast polar
Radiants i ik Plants
Cosmopolitan in
Canada and
Alaska (3)! 5.7
Reaching coasts
of Alaska, B. C.
and W. Wash. (7) 30.4 | (10) 13.7 (21) 40.4
Wide-
ranging Reaching coasts
forest of S. E. Alaska (7) 9.6 (6) 11.5
species
Avoiding coasts,
S. E. Alaska to
Washington (29) 39.7 | (5) 5.0} (14) 26.9
Reaching coastal
mts., Alaska to .
Wide- Washington (1) 4.3 (16) 16.0] (2) 3.8
ranging
arctic Reaching coastal
or mts. of S. E.
arctic- Alaska (2) 8.7 (16) 16.0
alpine
species Avoiding coastal
mts., S. E. Alaska
to Wash. (5) 21.7 (24) 24.0
: Reaching coast
Wide- | of S. E. Alaska (6) 6.0] (4) 7.7
ranging
ae Avoiding coasts
gai S. E. Alaska to
species —_| Washington (1) 4.3 (4) 5.4] (15) 15.0| (1) 1.9
Reaching coasts
Alaska to
Washington (1) 4.3] (4) 54] (6) 6.0
Alaskan- Reaching coasts
cordilleran | of S. and S. E.
species Alaska (5) 21.7 (1) 43] (4) 54] (4) 40) (1) 1.9
Avoiding coasts
S. E. Alaska to
Washington (9) 39.1 (3) 4.1] (7) 7.0
Reaching coasts,
S. Alaska to
a er Washington (14) 60.8] (1) 1.4] (1) 1.0
1s ares Avoiding coasts,
S. Alaska to
Washington (11) 15.0

1 Figures in parentheses indicate the numbers of species.

88 SARGENTIA | [6
Montane plants, 71.2% of his Continental West American plants, 34.8% of
his West American Coast Radiants, and 4.3% of his Beringian Radiants from
the same source. Most of the species in this source are wide-ranging across
the continent.

TABLE 7.
W. Conti- : Boreal
Beringia | American nental Arctic- Circum-
Radiants Coast W. Montane polar
. American Plants
Radiants Plants
Plants
Plants of wide range across the
continent (9) 39.1 (7) 30.4 | (50) 68.5 | (82) 82.0 | (51) 98.0
Ranges mainly in Alaska and the
Cordillera (14) 60.9 | (16) 69.6 | (23) 31.5 | (18) 18.0] (1) 2.0
Plants of the forested country (1) 4.3} (8) 34.8 | (52) 71.2 | (7) 7.0 | (41) 78.8
Arctic, arctic-alpine, or alpine
species (21). 91.3 | (15) 65.2 1 (17)}228.3.4 toa) 720-113) 59
““Timber-line”’ species (1) 4.3 (4) 5.4] (21) 210] (5) 9.6
Cosmopolitan ; (3K OOF
Continental with regard to S. E.
Alaska, B. C.,and Washington | (15) 65.2 (47) 64.4 | (51) 51.0 | (15) 28.8
Avoiding coasts of B. C. and
Washington, but occurring
in S, E. Alaska (7) 30.4] (1) 4.3 | (11) 15.0 | (26) 26.0 | (11) 21.1
Extending into coastal strip, S. E.
Alaska to Washington (1) 4.3 | (22) 95.7 | (15) 20.5 | (23) 23.0 | (26) 50.0

This can be illustrated also with the Arctic-Montane group, of which 82.5%
(including half the “timber-line” element) are derived from my arctic-alpine
plants. Hultén derives approximately 10.7% of his Boreal Circumpolar group,
26% of his Continental West American plants, 65.2% of his West American
Coast Radiants, and 93.4% of his Beringian Radiants from the same source.

As previously stated, Hultén, in building his “equiformal progressive areas”
outward from their centra, had to draw increasing numbers of species from the
great groups of wide-ranging arctic-alpine and forest plants. The above tables
and comparisons give a rough index of the extent to which he did this. Corre-
lations with actual range patterns, made within his major groups, are fairly good.
As would be expected a large percentage of the Boreal Circumpolar plants come
from the wide-ranging forest species. Likewise it is logical to find that most
of the Beringian and Arctic-Montane groups are among the widespread arctic
and alpine ranges. Since most of the Alaskan—Cordilleran element in the Brint-
nell Lake flora consists of arctic-alpine species, it is to be expected that Hultén’s
West Coast Radiants should show a large proportion from this source. It is
also reasonable that his West Coast Radiants should all be present in the coastal
strip, and that his Continental West American species should be prevailingly
continental in the west. At the same time it is clear that the principal range
patterns as I have designated them are more or less heterogeneous with regard
to the lesser representations of Hultén’s categories.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 89

Among the Beringian Radiants 39.1% (9 spp.) are plants of wide range
across the continent, nearly all of them arctic. Judging by their actual range
patterns these species could as well be placed among the Arctic-Montane group,
and their attachment to a Beringian center of origin is difficult to justify. They
indicate the submergence of the Beringian equiformal progressive areas into the
heterogeneous assemblage of wide-ranging arctic-alpine species.

The same analysis may be made of the West American Coast Radiants. No
less than 30.4% of them are plants of wide continental range, about two-thirds
of them from the arctic-alpine ranges and about one-third from the forested
country. These 30.4% (7 spp.) could as well be classified as Arctic-Montane
or Boreal Circumpolar. Among the Continental West American plants repre-
sented at Brintnell Lake 68.5% (50 spp.) are of wide range across the con-
tinent, most of them in the forests. An obvious reason for the selection of these
species from the forest wides is their relative continentality in the west, but in
Hultén’s arrangement even this breaks down to the extent of about 35% of the
cases.

The above analysis suggests that the identity of Hultén’s “equiformal pro-
gressive areas,” in terms of the species that compose them, can be maintained
only in or near their “centra” where groups of plants having limited ranges are
concentrated. It is possible to find, in some cases, concentric ranges that justify
the enlargement of the centra in certain directions, indicating trends of geo-
graphic development. Hultén has done this admirably, and has pointed out
not only the probable locations of the centra, but also the probable routes by
which boreal America has acquired its post-Glacial flora. Most of the arctic
and subarctic flora of North America, however, is composed of “pools” of wide-
ranging species, into which Hultén’s equiformal concentric ranges disappear.
The great arctic-alpine, or Arctic-Montane pool absorbs on one hand the North-
ern Beringia Radiants, and on the other the alpine elements of the Continental
Western American, and parts of the Western American Coast Radiants. Other
parts of the last two, as well as some Beringian elements as they were originally
constituted by Hultén, become lost in the wides of the forested regions.

The sorting of the wides, therefore, into groups representing their centers of
origin seems impracticable except within certain limits. Arctic-alpine species
can be separated reasonably well from those of the forests. Indications of con-
tinentality can be found in the behavior of plants with relation to the western
coastal strip, and in the extent of their extreme northwestern limits. With our
present knowledge of post-Glacial climatic and geological history, and with our
present limited data on existing environmental conditions as well as on the
genetic constitution of the species, it seems unwise to attempt further sub-
divisions.

It is remarkable, however, as will be shown below, that in spite of the obvious
discrepancies between Hultén’s arrangement of the species and the evidence from
their actual. ranges, the correlations as a whole are positive enough to justify
about the satne conclusions from his methods with regard to the origin of the
Brintnell Lake flora as are reached by the methods I have used. This I believe
can be put down to Hultén’s perspicacity in handling the somewhat imperfect
data he had on North American ranges.

90 SARGENTIA [6

ORIGINS OF THE FLORA OF SOUTHWESTERN MACKENZIE

Whether or not there were ice-free land surfaces in southwestern Mackenzie
during the Late Wisconsin (W,) glacial episode is still uncertain. I have
shown in a recent paper (1946) that a large triangular area in Alberta and
southern Saskatchewan was in all probability uncovered during this period.
The location of the northern apex of the triangle, formed by the junction of
Keewatin and mountain glaciers, is unknown, but it is not likely that it reached
north to the 60th parallel. For the Brintnell Lake region I have presented evi-
dence that summits and upper slopes above approximately 5000 feet were not
covered.

If there were ice-free summits in the Mackenzie Mountains, their flora must
have been composed of the arctic-alpine remnants of a wide pre-Late Wisconsin
dispersal of these plants. A few species found at Brintnell Lake suggest this.
First are those of wide range in Eurasia but not known elsewhere in North
America: Saxifraga sibirica, Calamagrostis lapponica s. str., Arnica alpina s. str. ;
second are the suspected endemics, of which there are eight. Most of the
endemics, however, appear to be “incipient” rather than well-defined morpho-
logical segregates. Considering the small number of these unique plants, the
paucity of the Brintnell Lake flora as a whole, and the present state of botanical
exploration in vast areas of the northwest, it is hardly justifiable to set up the
Mackenzie Mountain region, at least so far as the eastern slopes are concerned,
as a refuge equivalent to those of Beringia, the north Pacific coasts, or the
Yukon Plateau. It seems more reasonable to assume that although a few alpine
plants may have persisted at Brintnell Lake during Late Wisconsin time, most
of the flora has arrived during and since the retreat of the last valley glaciers.
The findings of Mr. Porsild on the western slopes of the mountains may require
modification of this view.

The length of time involved since the final retreat of the ice is highly con-
jectural. On the basis of Antevs’ conclusions (1931, 1933) I have suggested
(1. c.) that the ice did not disappear entirely from Great Slave Lake until near
the advent of the post-Glacial climatic optimum. This period is commonly
thought to have occurred between 3000 and 9000 years ago. It is not impos-
sible that the latest moraines at Brintnell Lake are of approximately equivalent
age. Porsild (1945) believes that he has found some evidence in Macmillan
Pass of a climate milder than now exists there.

If Hultén’s theories of equiformal progressive areas and varying biotype de-
pauperation are tenable, then it should be possible to draw up a sequence show-
ing the relative availability of plants for the colonization of the Brintnell Lake
area during the retreat of the glaciers. It is presumed that for a time after the
disappearance of the ice the mountains and adjacent valleys and plains had free
access to the alpine and tundra floras of the Rockies, the Cassiars, the Yukon
Plateau, the Richardson Mountains, and the arctic lowlands. With the ameliora-
tion of the climate coniferous forests eventually covered the valleys of the Mack-
enzie, upper Liard, Peel, and Wind Rivers, and finally the divides between these
streams and the Yukon system. In so doing they effected the isolation of the
alpine and arctic elements of the Mackenzie Mountain flora.

Here again the time intervals are conjectural, but probably can be brought
within reasonably definite limits. There is fairly good evidence that forests
did not appear in the central Mackenzie basin until after the final retreat of the

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 91

ice from Great Slave Lake and until after the last of the post-Glacial lakes in
the Athabaska—Great Slave Lake region had been drained. Using the reason-
ing referred to above, therefore, forests did not enter southwestern Mackenzie
until the period of the climatic optimum, probably as late as 7000 years ago.
They can hardly have reached the higher, interior valleys and divides until
somewhat later. The prairies of the upper Mackenzie River country are thought
to have developed directly from some form of tundra on the heavier lacustrine
soils.

According to Hultén’s theory the first aggressive floristic elements to invade
the glaciated lands were the Arctic-Montane plants, and such parts of the
Beringian and other groups as are supposed to have reached refugia south of
the ice before the maximum advance. In both cases these plants were able to
maintain large populations during the last great ice advance, and to maintain
their genetic plasticity. The most plastic of all Hultén’s groups, and certainly
the most wide-ranging species, are among the Boreal Circumpolar plants, but
since a very large proportion of this group is of forest species, it cannot be ex-
pected to show so high a representation in the Mackenzie Mountains as the
arctic-alpine group.

Next to the Arctic-Montane plants in a decreasing scale of availability would
come the Continental West American Radiants. These are thought to have
survived in the Yukon valley and the northern Rocky Mountains, many of them
below the ice boundary. Since they contain a large arctic-alpine element they
can be expected to be well represented in the Mackenzie Mountains. Further-
more a part of their refuge areas are directly adjacent to the Mackenzie Moun-
tains.

The various Beringian Radiants of Hultén should be rather poorly represented,
not only because many of them are thought to have lost their spreading capacity,
but also because they, like other alpine elements, were eventually stopped by the
advance of the forests. Another small part of the Mackenzie Mountain flora
should be derived from the coastal radiants, for they could not be expected to
spread far outside the habitats of the relatively warm and wet north Pacific
shores. It is true that Hultén allows many of these coastal species to spread
widely across the continent, but they are more southern types that would not be
expected at high latitudes.

In Taste 8 I have compared Hultén’s floristic elements of the Brintnell Lake
area in two ways. First the percentages of the total known flora which are

TABLE 8.
% of % of
Hultén’s Found at total Hultén’s
totals Brintnell Brintnell amended
(amended). Lake L. plants totals
Local plants, or plants of limited
known range 12 4.2
Beringia Radiants 248 23 8.1 oS
Atlantic-Pacific plants 16 0 0 0
W. American Coast Radiants 267 23 8.1 8.6
Continental W. American Radiants 190 73 25.8 38.4
Arctic-Montane plants 235 100 o0.3 42.6
Boreal Circumpolar plants 262 52 18.3 19.9

Totals 1218 283

a SARGENTIA [6

derived from the various sources are compared, and second, the percentages of
the total numbers of radiants (as revised in this paper) from the various sources
that are represented in the mountains. Approximately the same results are
reached by both methods. The Arctic-Montane group is present in largest
numbers. Over 42% of all the plants listed in the group have been found at
Brintnell Lake, and they make up about 35% of the local flora there. Next are
the Continental Western American plants, and third, the Boreal Circumpolar
group. The Beringian groups, taken together, and the western American Coast
Radiants have each supplied only 8.1% of the flora. The most strictly coastal
group set up by Hultén, of Atlantic—Pacific plants, is not represented at all.

It will be seen at once that these results correlate very well the series ex-
pected on the basis of Hultén’s theory. They lend considerable support to his
general concept. It should be noted that the results from similar comparisons
made with Hultén’s unrevised figures are not far different from the above. The
positions of the Boreal Circumpolar and Continental West American groups are
reversed, but they are not far apart in any case. Some interesting figures appear
when the Beringian Radiants are broken down into the various elements outlined
by Hulten (Taste 9).

TABLE 9,
i Hultén’s
totals % of % of
(amended) Found at total Hultén
for the Brintnell Brintnell radiants
centra Lake L. plants (amended )
S. Beringia Radiants 77 er 0.35 1.29
N. Beringia Radiants 90 15 533 16.66
Bilateral Radiants of N. Beringia 45 1 0.35 222
Arctic-Pacific plants 36 6 Zi13 16.66
Totals 248 WX 8.16 9.27

Although the actual numbers of these species that have reached the Brintnell
Lake region is small, the proportions of the total Northern Beringian and Arctic
Pacific Radiants are 16.6%. In spite of their relative depauperation, therefore,
these plants are nearly as well represented as the Boreal Circumpolar group.
It is possible to interpret them as evidence that forests have isolated the alpine
flora of the Mackenzie Mountains rather recently, allowing time for Beringian
plants to reach the mountains in considerable numbers.

Using data derived from maps presented in this paper, it is possible to arrange
a similar sequence of floristic influences that have affected the Brintnell Lake
region. The size and character of existing ranges can be used as indicators of
the success with which plants have migrated from the Pleistocene refugia.

Considering first the alpine flora, by far the largest single group (66 spp.) is
derived from wide-ranging arctic-alpine plants—51.5% of the alpines, or about
23% of the total flora. These are the species that are thought to have main-
tained comparatively large populations during the Pleistocene, and to have been
able to migrate during that time. Second in extent of range are the species that
have occupied wide areas in Alaska or the northern Cordillera or both, some of
them extending eastward toward Hudson Bay (maps, Fig. 9-a, b, c; Fig. 10).
There are 49 of these—about 38.2% of the alpines. Finally there are 13 spe-

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 93

cies of far northwestern range (map, Fig. 9-d), only a few of which extend
south of the 60th parallel. They comprise about 10.1% of the alpine plants at
Brintnell Lake.

The plants of wide range on the mountains of Alaska, Yukon, British Columbia,
and Alberta may well have had access to the large refuge of the Yukon Plateau
and to possible refugia in the northern Rocky Mountains and eastern foothills,
as well as to those of Beringia. The smallest ranges are those of the Alaskan
species which were probably limited to Beringian refugia.

Nearly all of the forest plants at Brintnell Lake are derived from wide-ranging
species in boreal America. I have already noted that only ten of the western
plants represented are typically forest types. The Brintnell Lake forest flora is
clearly derived from lands to the southeastward and has undoubtedly come up
from the Liard valley. That of the central Mackenzie region, in turn, as I
have suggested in an earlier paper, has probably developed from the amalgama-
tion of populations that persisted through Late Wisconsin time in the East and
in the valleys and foothills of the Rocky Mountains. All of these species could
have maintained fairly large ranges during that time, and have no doubt increased
their spreading capacity by subsequent fusion of populations.

It is impossible to assess accurately the influences of eastern and western
forest elements in the Brintnell Lake flora. I have pointed out elsewhere (1946)
that morphological characters in tree populations from eastern and cordilleran
refugia can be found intermingled in the forests of the central part of the
Mackenzie basin. Eastern characters apparently are lacking in the Brintnell
Lake forests. Eastern forms of the white spruce, white birch, aspen, and larch
were not found there, nor was the jack pine. Whether this applies also to
herbaceous and shrubby elements cannot now be determined.

Some further observations on the occurrence of eastern and western geo-
graphic varieties may be of interest here. When the whole forest flora of the
Mackenzie basin is considered, it is found that these eastern and cordilleran
components within species complexes are limited almost entirely to trees and
certain shrubs. Wide-ranging herbs of the Canadian forest show a remarkable
uniformity throughout, or a few have varietal segregates that are not readily
related to either eastern or western areas.

Among shrubs, on the other hand, it is possible to find a few such segregates.
Alnus rugosa in the east has a cordilleran counterpart, A tenwifolia, while in
Alnus crispa there is almost complete uniformity. Viburnum edule, Myrica
Gale, Potentilla fruticosa, and Betula glandulosa are essentially uniform through-
out. Rosa acicularis has several varietal segregates based upon the form of the
ripe fruit; but for the most part they are not readily set off geographically,
although R. acicularis var. lacorum may eventually prove to be the most com-
mon in the northern Cordillera. Ericaceous shrubs are exceedingly uniform
with the possible exception of Kalmia polifolia, whose var. microphylla is cor-
dilleran. Among the willows there is a varying amount of specific segregation.
In Salix § Cordatae, the cordata-lutea complex of the east has a close western
relative in S. Mackenzieana. Likewise S. serissima and S. lasiandra are re-
garded as seperate species, though they merge in the Athabaska country. The
sand bar willow, Salix interior, appears to merge, somewhere in the northern
Rockies, with S. exigua. On the other hand, one of the commonest willows in
the north, S. Bebbiana, is remarkably uniform throughout. The same is true
of S. planifolia, S. candida, and S. myrtillifohia.

94 SARGENTIA [6

All of the trees, with the possible exception of black spruce and balsam poplar,
have definable eastern and western varietal or specific segregates. I have
already pointed out that the degree of segregation in the tree species is roughly
proportional to their mesophytism, ie., the farther they are from the arctic
timber line the more clearly the varieties or species can be defined. Species
complexes that have not actually merged their eastern and western parts, such
as the firs, hemlocks, and arbor-vitaes, are still looked upon as having separate
species in the east and west. This idea can also be applied to shrubs and herbs,
with the result that the farther south it is carried the better the segregation
becomes. Most of the willows mentioned above as having eastern and western
species pairs are usually regarded as separate entities, though with many inter-
mediate forms, while there are other pairs still farther south, such as Salix sub-
coerulea and S. pellita, which are usually kept separate without question. Among
non-boreal species, both herbaceous and shrubby, eastern and western identities
or pairs have long been recognized. They exhibit some of the remarkably dis-
rupted ranges which have been so thoroughly studied by Fernald (1925).

From the above notes the general pattern of specific and varietal segregation
in the Canadian forest becomes clear. The farther north one goes the poorer
it becomes. There is, however, the anomaly of a widespread community in
which the segregation of geographic varieties is more distinct among trees than
among herbs, with shrubs in an intermediate position. At present it is possible
only to advance a theory to explain this. If our premise is correct that the
widespread boreal forest of the northern interior plains has been formed by the
amalgamation of eastern and western components since the retreat of the Late
Wisconsin ice, then we can visualize the process as being one of hybridization
and the intermingling of biotypes. This process would be most rapid in species
with the shortest life cycles, which would be the herbs. The shrubs would ac-
complish the fusion somewhat more slowly, and the trees most slowly of all.

There appears some corroborative evidence for such a gradation when the
shrubs alone are considered. The species complexes of upland woods, muskegs,
and slough margins show the most complete uniformity, whereas shrubs of river
banks have maintained better segregation. This is particularly true of the
willows and alders. Adherence to specific limited habitats may thus have con-
stituted a barrier to free interbreeding. On the other hand Cornus stolonifera,
also a plant of river banks and flood plain woods, shows poor segregation of
geographic varieties. It is easily dispersed by birds, however, and may thus
have overcome the handicap of its habitat requirements.

In the light of this reasoning, the varietal segregates which appear at Brintnell
Lake suggest that most of the forests came there from the foothills of the northern
Cordillera rather than from the central Mackenzie basin. Nevertheless eastern
elements may be there among herbaceous and shrubby species, submerged in
uniform wide-ranging populations that have grown out of the mergence of
eastern and western glacial components.

I have not considered the intermediate “timber-line” group, but whether it
is placed with forest or arctic elements, it can be thought to have shared the
wide ranges of either, and to have maintained great spreading capacity.

By way of summary, therefore, we may picture the Mackenzie Mountains as
having been one of the latest montane areas to lose its glaciers (remnants of
them still exist). While the glaciers were retreating there was a period of time
during which the lower slopes were tundra-covered, with the tundra more or

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 95

less continuous across the neighboring valleys and to neighboring mountain
ranges. The length of the period is problematical, for it is impossible with
present data to measure it accurately. There is some evidence, however, that
forests did not come into the lower valleys until about 7000 years ago, and that
they did not cover the divides until somewhat later. In any case the tundra
period must have been one in which the arctic and alpine plants of all the north-
western refugia had open routes of migration to the Mackenzie Mountains.
There is evidence that they availed themselves of these routes in differing pro-
portions, depending first upon their broad habitat preferences, and second upon
their inherent abilities to migrate. Their success at colonizing the mountains
was conditioned also by the remoteness of the various refugia. Plants of con-
tinental habitat preferences were of course most successful; and of these the
wide-ranging Arctic-Montane group were outstanding because of both accessi-
bility and lack of biotype depauperation. Survivors in the Yukon valley and
the northern Rockies were second in numbers, for they were only partially
depauperated and were near at hand. Northern Beringian and Coastal Radiants
were next in importance, probably held in check both by distance and lack of
plasticity.

The process of alpine colonization appears to have been checked, at least for
the Brintnell Lake region, before it had reached anything like completion.
Evidence for this is to be found in the general paucity of species in the alpine
plant cover. The “partial” nature of the flora is to be seen in all the common
arctic families and genera. Such groups as Potentilla, Senecio, Arnica, and
Pedicularis, all represented in surrounding regions by half a dozen or so species,
have only 2 to 4 species each in the Brintnell Lake district. The colonization
appears to have been stopped by the advance of forests into the surrounding
valleys. These forests brought a new element to the mountain flora, derived
from refugia south of the ice or on the slopes of the more southern mountains.
Whether any of it came from the Yukon valley is uncertain. It was derived
from Hultén’s Boreal Circumpolar, Continental Western American, and possibly
in part from his western coastal groups. In terms of range patterns outlined
in this paper it was drawn chiefly from wide-ranging forest species and in part
from the “timber-line” group. By nature it is aggressive, but probably shows
so small a portion of the total Brintnell Lake flora as it does only because of
the short time available for its invasion, and because of the subarctic situation
of the Mackenzie Mountains.

CATALOGUE OF THE VASCULAR FLORA OF
SOUTHWESTERN MACKENZIE

The ensuing catalogue contains 725 species and varieties of vascular plants.
They are distributed among the major groups as follows: Pteridophyta—35 ;
Gymnospermae—9 ; Monocotyledoneae—201 ; Dicotyledoneae—480. The list
cannot be looked upon as anything more than an attempt to bring existing
knowledge of the flora of southwestern Mackenzie together in one place so that
it can be studied and enlarged more intelligently.

It is clear, from an examination of range patterns and the floras of surround-
ing regions, that the catalogue almost certainly will be enlarged very considerably
with further collecting. Using Porsild’s work on the lower Mackenzie and the
western Arctic (1943) together with my own studies in the Athabaska—Great

96 SARGENTIA [6

Slave Lake area (1936) it is possible to draw up a list of about 240 species and
varieties known to occur in one or the other of the two areas, but not yet
collected in southwestern Mackenzie. A great many of these, particularly the
more southern element which occurs in the Wood Buffalo ‘Park, are to be ex-
pected in our region on the Alberta Plateau and in the Mackenzie Lowland
physiographic provinces. To have included them in the present catalogue,
however, would have necessitated the assumption of range extensions for which
there is no actual evidence.

I have admitted certain species to the catalogue although no specimens of
them have yet been collected in our region. There are 47 of them, selected from
the surrounding floras because they have been found in the Mackenzie basin both
north and south of our area. A smaller number, 22, are included on the basis
of early records made by Richardson at the time of the Franklin Expeditions.
Some of these are admittedly doubtful, but I am inclined to honor them because
of the surprising number of supposedly erroneous Richardson records that have
been verified in recent years.

In citing literature and synonyms I have included only material that bears
directly upon the flora of this region. The symbols indicating the herbaria from
which I have cited specimens are as follows:

G.—Gray Herbarium

Can.—National Herbarium of Canada

N.—Herbarium of the New York Botanical Garden

US.—United States National Herbarium

Unless otherwise indicated, the specimens cited are those collected by Dr.
J. H. Soper and myself in the season of 1939. The first set of these collections,
including types of novelties, will be found in the Gray Herbarium of Harvard
University, and the second set in the herbarium of the National Museum of
Canada at Ottawa.

OPHIOGLOSSACEAE

Botrychium Lunaria (L.) Sw. in Jour. Bot. Schrad. 18002: 110 (1801); Hultén, FI.
Alaska & Yuk. Pt. 1: 48 (1941); Porsild in Nat. Mus. Can. Bull. 101: 14, 29
(1945).

Osmunda Lunaria L. Sp. Pl. 1064 (1753).
Mackenzie R. between Wrigley and Blackwater R., Crickmay 124 (Can.); Nahanni Mt.,

Wynne-Edwards 8406 (Can.) ; Bolstead Cr., Canol Rd. mile 111 E., Wynne-Edwards 8230

(Can.).

Botrychium Lunaria (L.) Sw. var. minganense (Victorin) Dole, Fl. Vermont, ed. 3:
1 (1937); Hultén, Fl. Alaska & Yuk. Pt. 1: 48 (1941); Porsild in Nat. Mus. Can.
Bull. 101: 29 (1945).

Botrychium minganense Victorin in Trans. Roy. Soc. Can. 21: 331 (1927).
Mossy slope near summit of Lone Mt., Wynne-Edwards 8407 (Can.).

Botrychium multifidum (Gmel.) Rupr. Beitr. Pfl. Russ. Reich. 11: 40 (1859).
Osmunda multifida Gmel. in Nov. Comm. Acad. Sci. Petrop. 12: 517 (1859).
Abundant in a small prairie opening on the upland west of Fort Simpson,

9917. Known in the Mackenzie basin from one other locality, Sand Pt., on
the north shore of Lake Athabaska (Raup & Abbe 4565, 4649, 4654).

Botrychium virginianum (L.) Sw. var. europaeum Angstr. in Bot. Notiser 1854: 68
(1854); Porsild in Nat. Mus. Can. Bull. 101: 29 (1945).
Osmunda virginiana L. Sp. Pl. 1064 (1753).
In damp hollow near summit of Lone Mt., Wynne-Edwards 8408 (Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 97

POLY PODIACEAE

Woodsia ilvensis (L.) R. Br. Prodr. Fl. Nov. Holl. 158 (1810); Hooker, Fl. Bor.-Am.
2: 259 (1840); Macoun, Cat. Can. Pl. Pt. 5: 283 (1890); Hultén, Fl. Alaska & Yuk.
Pt. 1: 14 (1941).

Acrostichum ilvense L. Sp. Pl. 1071 (1753).

Among boulders in coarse slide-rock, south slope of Red Mt., alt. 4500’, 9787; shale cliffs
on west slope of Terrace Mt., alt. 5000’, 9518; shale cliffs near summit of ridge on Terrace
Mt., northeast of camp, alt. 5000’, 96817.

Occasional to common locally. Noted in Fl. Bor.-Am., from “Canada...
to... Bear L.” ;

Woodsia glabella R. Br. apud Richardson, Bot. App. Frankl. Journ. 754 (1823);
Hooker, Fl. Bor.-Am. 2: 259 (1840); Macoun, Cat. Can. Pl. Pt. 5: 282 (1890);
Hultén, Fl. Alaska & Yuk. Pt. 1: 13 (1941); Porsild in Nat. Mus. Can. Bull. 101:
14, 29 (1945).

Liard R., Nahanni Butte to Simpson, Crickmay 1 (Can.); west ridge of Franklin Mts.
at “River-between-two-mountains,” alt. 1000’, Crickmay 8 (Can.); Smith Cr., 4 mi. about
mouth, Crickmay 25 (Can.) ; Lone Mt., Wynne-Edwards 8409 (Can.) ; mountain range west
of Bolstead Cr., about mi. 111 E., Wynne-Edwards 8229 (Can.).

Common on dolomitic rocks in the central and southern parts of the Mackenzie
basin. The type of the species is a Richardson specimen from Great Bear Lake.

Cystopteris fragilis (L.) Bernh. in Neu. Jour. Bot. Schrad. 12: 26 (1806); Raup in Jour.
Arn. Arb. 17: 188 (1936); Hultén, Fl. Alaska & Yuk. Pt. 1: 16 (1941); Porsild in
Nat. Mus. Can. Bull. 101: 29 (1945).
Polypodium fragile L. Sp. Pl. 1091 (1753).
Dryopteris austriaca sensu Porsild in Nat. Mus. Can. Bull. 101: 29 (1945), non Woynar.
Northwest shore of Great Slave L., Bedford (Can.); north of the height of land between
Great Slave and Great Bear Lakes, Bedford (Can.); Liard R. between Nahanni Butte and
Simpson, Crickmay 2, 3 (Can.); west ridge of Franklin Mts. at “River-between-two-moun-
tains,” alt. 1000’, Crickmay 7 (Can.); rocky spruce woods on southward-facing slope west
of Brintnell L., alt. app. 2800’, 9474; shale crevices and slide-rock, north slope of Colonel
Mt., alt. 5500’-6000’, 9644; crevices in coarse slide-rock, south slope of Red Mt., alt. 4500’,
9727; shale cliffs in gorge on north slope of Colonel Mt., alt. 3500’, 9835; north peak of
Nahanni Mt., Wynne-Edwards 8410, 8411 (Can.); Lone Mt., Wynne-Edwards 8412, 8414
(Can.).
Occasional throughout the area; the only fern collected on Colonel Mt. except
Dryopteris Linnaeana, which was at the foot of the slope. One of the few
plants able to live in the loose shale of steep slopes.

Cystopteris montana (Lam.) Bernh. in Neu. Jour. Bot. Schrad. 12: 26 (1806); Hultén,
Fl. Alaska & Yuk. Pt. 1: 18 (1941); Porsild in Nat. Mus. Can. Bull. 101: 29 (1945).

Polypodium montanum Lam. FI. Franc. 1: 23 (1778).

Liard R., between Nahanni Butte and Simpson, Crickway 4 (Can.); north peak of
Nahanni Mt., Wynne-Edwards 8412 (Can.).

Dryopteris Lirmaeana C. Chr. Ind. Fil. 275 (1905); Hultén, Fl. Alaska & Yuk. Pt. 1:
25 (1941).

Polypodium Dryopteris L. Sp. Pl. 1093 (1753); Hooker, Fl. Bor.-Am. 2: 259 (1840)
at least in part.

Phegopteris Dryopteris (L.) Fée; Macoun, Cat. Can. Pl. Pt. 5: 270 (1890).

Mossy spruce woods on south shore of Brintnell L., 9330.

Occasional in the above habitat, but not seen elsewhere in our region. Noted
in Fl. Bor.-Am., as occurring from “Canada... to... Bear Lake.”

98 SARGENTIA [6

Dryopteris Robertiana (Hoffm.) C. Chr. Ind. Fil. 289 (1905); Hultén, Fl. Alaska &
Yuk. Pt. 1: 29 (1941); Porsild in Nat. Mus. Can. Bull. 101: 29 (1945).
Polypodium Robertianum Hoftm. Deutsch. Fl. 2: 20 (1795).
ry 3 ca Wynne-Edwards 8416 (Can.) ; Willow Lake R., 44 mi. above mouth, Crickmay
WSU0Gan.).

Dryopteris fragrans (L.) Schott. Gen. Fil. pl. 9 (1834); Hultén, Fl. Alaska & Yuk. Pt.
1: 23 (1941); Porsild in Nat. Mus. Can. Bull, 101: 14 (1945).

Polypodium fragrans L. Sp. Pl. 1089 (1753).

Aspidium fragrans (L.) Sw.; Hooker, Fl. Bor.-Am. 2: 261 (1840) ; Macoun, Cat. Can.
Pl. Pt. 5: 276 (1890). J

Rocky spruce forest on south-facing slope (granite) west of Brintnell L., alt. app. 2800’,
9483; among boulders in coarse slide-rock, south slope of Red Mt., alt. 4500’, 9754; Little
Keele R., Canol Rd., mi. 55 E., Porsild 11783 (Can.) ; Trout Cr., Canol Rd., mi. 118 E.,
Porsild 11835 (Can.).

Occasional to common in the Brintnell Lake localities cited. Noted in FI.
Bor.-Am. as occurring from “Saskatchewan to the Arctic Sea-coast.”

Dryopteris fragrans (L.) Schott. var. remotiuscula Kom. See Brown, Index N. Am,
Ferns 68 (1938); Porsild in Nat. Mus. Can. Bull. 101: 29 (1945).

Thelypteris fragrans (L.) Nieuwl. var. Hookeriana Fern. in Rhod. 25: 3 (1923).
Lone Mt., Wynne-Edwards 8415 (Can.).
Dryopteris spinulosa (O. F. Miill.) Watt in Can. Nat. II. 13: 159, 403 (1867); Ktze.
Rev. Gen. Pl. 2: 813 (1891).
Polypodium spinulosa O. F. Miill. Fl. Fridr. 113, f. 2 (1767).
North of the height of land between Great Slave and Great Bear Lakes, Bedford (Can.).
Apparently rare in the northern part of the Mackenzie basin, and occasional
in the southern and southwestern parts (Raup, 1934, 1936).

Asplenium viride Huds. Fl. Angl. 385 (1762); Porsild in Nat. Mus. Can. Bull. 101:
29 (1945).

Lone Mt., Wynne-Edwards 8417 (Can.).
Previously known in the Mackenzie basin from an unverified record by John
Macoun in the mountains along the upper Peace R.

Pellaea glabella Mett. ex Kuhn in Linnaea 36: 87 (1869). See Am. Fern JOUrage
3-5, 77-87 (1917); and 11: 39-40 (1921).
Pteris atropurpurea sensu Hooker, Fl. Bor.-Am. 2: 264 (1840), non L.
West ridge of Franklin Mts., on limestone, Crickmay 9 (Can.).
Rare in the southern and central parts of the Mackenzie basin, where it is
confined to limestone or dolomitic rocks. Noted by Hooker (FI. Bor.-Am., as
Pteris atro-purpurea) from “Canada to Bear Lake. . . .”

Cryptogramma crispa (L.) R. Br. var. acrostichoides (R. Br.) C. B. Clarke in Trans.
Linn. Soc. II. 1: 460 (1880). See Rhod. 37: 238-247 (1935).

Osmunda crispa L. Sp. Pl. 1067 (1753).

Cryptogramma acrostichoides R. Br. apud Richardson, Bot. App. Frankl. Journ. 767
(1823) ; Hooker, Fl. Bor.-Am. 2: 264 (1840) ; Macoun, Cat. Can. Pl. Pt. 5: 261 (1890).

Cryptogramma crispa (L.) R. Br. ssp. acrostichoides (R. Br.) Hultén, Fl. Alaska &
Yuk. Pt. 1: 40 (1941).

North of the height of land between Great Slave and Great Bear Lakes, Bedford (Can.).

Variety acrostichoides is common in dry crevices, chiefly on pre-Cambrian
rocks, throughout the Athabaska—Great Slave Lake country. Though very little
collected in southwestern Mackenzie, it probably is widely spread there also.
Noted in FI. Bor.-Am., from “Cumberland House Fort to Bear Lake.”

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 99

Cryptogramma crispa (L.) R. Br. var. sitchensis (Rupr.) Hultén, Fl. Alaska & Yuk.
Pt. 1: 40 (1941).

Allosurus sitchensis Rupr. Beitr. z. Pfl. Russ. Reich. 3: 46 (1845).

Rocky eastern slope of Red Mt., alt. 5800’, just below summit, occasional, 9725.

Our plant is clearly the same form as that noted by Hultén as the commonest
one in the Aleutian Islands (FI. Aleut. Isl. 51-52, 1937). It has broadly del-
toid sterile fronds, tripinnately dissected, with the ultimate divisions small and
obovate. In the division of its fronds it closely resembles C. crispa (L.) R.
Br., as pointed out by Hultén, but its general form and striped scales relate it
more closely to the American C. acrostichoides. Fernald (in Rhod. 37: 238-
247, 1935) has maintained the latter as a variety of C. crispa, while Hultén (FI.
Alaska and Yuk. Pt. 1: 40, 1941) now gives it subspecific rank. Hultén ap-
parently considers var. sitchense as a segregate from ssp. acrostichoides, and
uses a quadrinomial to express the relationship.

Polypodium virginianum L. Sp. Pl. 1085 (1753). See Rhod. 24: 125 (1922).

North shore of Great Slave L., Bedford (Can.).

Common in the Wood Buffalo Park and the Athabaska-Great Slave Lake
district, but not yet found northwest of Great Slave Lake.

EQUISETACEAE

Equisetum arvense L. Sp. Pl. 1061 (1753); Macoun, Cat. Can. Pl. Pt. 2: 249 (1890);
Raup in Jour, Arn. Arb. 17: 190 (1936); Hultén, Fl. Alaska & Yuk. Pt. 1: 51
(1941); Porsild in Nat. Mus. Can. Bull. 101: 14 (1945).

Damp sand bar along Mackenzie R., west of Simpson, 9874; gravelly thickets along shore
of Brintnell Lake, 9309; shale slope near west end of lake, watered by mineral spring, 9570;
wet ground along small stream on north slope of Colonel Mt., alt. 4500’, 9404.

Common to abundant in the habitats noted above, but not observed elsewhere
about Brintnell Lake. Wet rock slides at the mouths of mountain springs are
made conspicuous at a distance by the bright green of this species, which is
almost the only vascular plant growing on them. Porsild has noted it as common
in Macmillan Pass and elsewhere along the Canol Road.

Equisetum pratense Ehrh. in Hanov. Mag. 9: 138 (1784); Hulten, Fl. Alaska & Yuk.
Pt. 1: 56 (1941).

Common in damp lowland woods near Simpson, 9095.

Not previously observed north of the Wood Buffalo Park (Raup, 1936, p. 190).

Equisetum sylvaticum L. var. pauciramosum Milde, Mon. Equiset. 292 (1865); Raup in
Jour. Arn, Arb. 17: 190 (1936). See Rhod. 20: 129-131 (1918).
Equisetum sylvaticum sensu Hooker, Fl. Bor.-Am. 2: 269 (1840), in part; Hultén, FI.
Alaska & Yuk. Pt. 1: 57 (1941), in part.

Resolution, Great Slave L., Kennicott (N); northwest shore of Great Slave L., Bedford
(Can.) ; mossy bank on north shore of Brintnell L., 9184.

Occasional to common in lowland woods around Brintnell L. Noted in FI.
Bor.-Am., from “Canada . . . to Fort Franklin on the Mackenzie River,” and
probably common throughout the upper Mackenzie valley forests. Collected at
Great Bear L. by R. Bell (Can.).

Equisetum palustre L. Sp. Pl. 1061 (1753); Hooker, Fl. Bor.-Am. 2: 269 (1840);
Macoun, Cat. Can. Pl. Pt. 2: 250 (1890); Hultén, Fl. Alaska & Yuk. Pt. 1: 55
(1941); Porsild in Sargentia 4: 8 (1943); Raup in Jour. Arn. Arb. 17: 190 (1936).

Lower Slave R., below Grande Detour, Raup 40; marshy mud flats at mouth of stream

at west end of Brintnell L., 9494; poplar-spruce woods near west end of Brintnell L., 9448.

100 SARGENTIA [6

This species is abundant on the newly formed mud flats at the mouth of the
glacial stream that comes into the western end of Brintnell L. It shares dom-
inance there with Eriophorum Scheuchzeri, Juncus castaneus, Carex aquatilis,
and Arctagrostis arundinacea. It is also abundant in the rather open woods
which develop on these flats, forming a nearly continuous green sward on the
forest floor. It is rare in other habitats about the lake. Reported in Fl. Bor.-
Am. as far north as the “shores of the Arctic Sea,” and by Porsild at Great
Bear L. and in the Mackenzie delta.

Equisetum fluviatile L. Sp.’Pl. 1062 (1753).

Equisetum limosum L. Sp. Pl. 1062 (1753); Hooker, Fl. Bor.-Am. 2: 269 (1840);
Hultén, Fl. Alaska & Yuk. Pt. 1: 54 (1941) ; Raup in Jour. Arn. Arb. 17: 191 (1936).
Lower Slave R., Raup 25, 26; northwest shore of Great Slave L., Bedford (Can.) ; wet
meadow along glacial stream near west end of Brintnell L., 9700.
Common on the higher parts of the wet mud flats at the west end of Brintnell
L., where it is associated with Poa, Calamagrostis, Stellaria, and Arctagrostis.
Noted in Fl. Bor.-Am. northward “. . . to the shores of the Arctic Sea.”

Equisetum prealtum Raf. Fl. Ludov. 13 (1817).
Dry sandy margin of an upland prairie opening west of Simpson, 9927.
Common in places, with spreading-ascending stems forming a loosely tangled

mat. Not previously reported in the Mackenzie basin north of the upper

Slave R. (Raup, 1936, p. 191).

Equisetum scirpoides Michx. Fl. Bor.-Am, 2: 281 (1803); Hultén, Fl. Alaska & Yuk.
Pt. 1: 57 (1941); Porsild in Nat. Mus. Can. Bull. 101: 14, 29 (1945).

Swampy woods near Simpson, 9090; mossy thickets on south shore of Brintnell L., 9339;
in damp hollow on summit of Lone Mt., Wynne-Edwards 8418 (Can.); north peak of
Nahanni Mt., Wynne-Edwards 8419 (Can.); along Canol Rd. at Sekwi R., mi. 174 E,,
Porsild 11843 (Can.).

Abundant in spruce forests throughout the region.
Equisetum variegatum Schleich. Cat. Pl. Helvet. 27 (1807); Hooker, Fl. Bor.-Am. 2:

270 (1840), in part; Hultén, Fl. Alaska & Yuk. Pt. 1: 59 (1941); Porsild in Nat.
Mus. Can. Bull. 101: 14 (1945).

Mud flats at west end of Brintnell L., 9809; mossy muskeg thickets at west end of lake,
9708; mossy thickets along south shore of lake, 9359; grassy alpine meadow on north slope
of Colonel Mt., alt. 4000’, 9607; along Canol Rd. at Sekwi R., mi. 174 E., Porsild 11842
(Can.) ; noted by Porsild also in Macmillan Pass, mi. 225 E.

Common to abundant below timber line, in the Brintnell L. area, but only
occasional above. It is a primary species over large areas of mud flat at the
west end of the lake and appears to form a definite stage in the vegetational
development between the marsh and the spruce-poplar woods.

LYCOPODIACEAE

Lycopodium Selago L. Sp. Pl. 1102 (1753); Hooker, Fl. Bor.-Am. 2: 266 (1840);
Macoun, Cat. Can. Pl. Pt. 2: 287 (1890); Hultén, Fl. Alaska & Yuk. Pt. 1:
71 (1941); Porsild in Nat. Mus. Can. Bull. 101: 14 (1945).

Northeast slope of Red Mt., Brintnell L. area, alt. 5000’, 9269; Bolstead Cr., Canol Rd.,
W ynne-Edwards 8231 (Can.).
Common on the turfy slopes above timber line north of Brintnell L., but not
seen on Colonel Mt.
Lycopodium annotinum L. var. alpestre Hartm. Skand. Fl. ed. 2, 294 (1832); Hultén,
Fl. Alaska & Yuk. Pt. 1: 63 (1941). See Rhod. 17: 123-125 (1915).
Northeast slope of Red Mt., Brintnell L. area, alt. 5000’, 9266.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 101

Abundant on turfy slopes above timber line. The leaves are much more
serrate than those of var. pungens, which was collected at lower levels. A
specimen with serrate leaves (W ynne-Edwards 8420, Can.) cited by Porsild,
l. c. 29, as var. pungens, probably belongs here, and possibly also his record for
L. annotinum in Macmillan Pass.

Lycopodium annotinum L. var. pungens (La Pylaie) Desv. in Mém. Soc. Linn. Paris
6: 182 (1827); Raup in Jour. Arn. Arb. 17: 192 (1936); Hultén, Fl. Alaska &
Yuk. Pt. 1: 63 (1941). See Rhod. 17: 123-125 (1915).

Northwest shore of Great Slave L., Bedford (Can.) ; north of the height of land between
Great Slave and Great Bear Lakes, Bedford (Can.); rich woods near shore at west end
of Brintnell L., 9490.

Common in a few places around Brintnell L., growing in a mat of woodland

mosses and lichens.

Lycopodium clavatum L. var. monostachyon Grev. & Hook. in Hook. Bot. Misc. 2:
375 (1831); Hultén, Fl. Alaska & Yuk. Pt. 1: 66 (1941). See Rhod. 12: 50 (1910).

Stony ridge on mountainside northeast of our camp at Brintnell L., alt. 3700’, 9680.

Common locally in rather dry, open spruce woods. Not previously observed
in the Mackenzie basin north of Lake Athabaska (Raup, 1936, p. 192).

Lycopodium obscurum L. var. dendroideum (Michx.) D. C. Eat. in Gray, Man. ed.
6, 696 (1890); Hultén, Fl. Alaska & Yuk. Pt. 1: 69 (1941). See Rhod. 23: 188
(1921).

Lycopodium dendroideum Michx. Fl. Bor.-Am. 2: 282 (1803).

Near Norman, Mackenzie R., Miss E. Taylor 155 (Can.).

Apparently rare in the northern part of the Mackenzie basin, though it is
locally abundant about Lake Athabaska and in the southwestern sections (Raup,

1934, 1936).

Lycopodium alpinum L. Sp. PI. 1104 (1753); Hultén, Fl. Alaska & Yuk. Pt. 1: 62
(1941).
East slope of Red Mt., Brintnell L. area, alt. 5000’, 9742; west slope of Terrace Mt., alt.
4800’, 9507.
Occasional to common at Brintnell L. on grassy slopes in the upper edge of
the birch scrub and above it.

Lycopodium complanatum L. var. canadense Victorin in Contr. Lab. Bot. Univ.
Montreal No. 3: 70 (1925); Hultén, Fl. Alaska & Yuk. Pt. 1: 68 (1941); Porsild
in Nat. Mus. Can. Bull. 101: 29 (1945).

Woods near Ft. Liard, Nowosad (Can.) ; thicket on hogback, north slope of Colonel Mt.,
alt. 4300’, 9629; in moss-lichen mat in open spruce woods on rocky knoll near east end of
Brintnell L. 9584; open woods along Frost Creek, north of camp, alt. 3500’, 9270; north
peak of Nahanni Mt., Wynne-Edwards 8421 (Can.).

Occasional to common in open woods and dry thickets. All of the specimens
collected in the Mackenzie Mts. have the short, fruiting peduncles of var. cana-
dense Victorin. Forms which should also be referred here were collected by
the writer on Lake Athabaska (1936, p. 192), and by A. E. Porsild in Alaska
(Rhod. 41: 173. 1939).

SELAGINELLACEAE

Selaginella selaginoides (L.) Link, Fil. Hort. Berol. 158 (1841); Hultén, Fl. Alaska
& Yuk. Pt. 1: 74 (1941); Porsild in Nat. Mus. Can. Bull. 101: 14, 30 (1945).

Lycopodium selaginoides L. Sp. Pl. 1101 (1753).

102 SARGENTIA [6

Mossy muskeg thickets near west end of Brintnell L., 9711; north peak of Nahanni Mt.,
Wynne-Edwards 8422 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., Porsild 11827 (Can.) ;
same, Wynne-Edwards 8232 (Can.); Little Keele R., alt. 2200’, mi. 51 E., Porsild 11781
(Can.) ; Macmillan Pass, mi. 225 E., Porsild 11182 (Can.).

ISOETACEAE

Isoétes Braunii Dur. in Bull. Soc. Bot. France 11: 101 (1864); Hultén, Fl. Alaska &
Yuk. Pt. 1: 76 (1941). See Ann. Mo. Bot. Gard. 9: 156-173 (1922).
This species has not yet been found in southwestern Mackenzie, but it is to
be expected there. It has been collected near Lake Athabaska (Raup & Abbe
4617) and at Great Bear L. (Porsild 3688).

PINACEAE

Picea glauca Voss var. albertiana (S. Brown) Sarg. in Bot. Gaz. 67: 208 (1916);
Porsild in Nat. Mus. Can. Bull. 101: 14 (1945).

Picea albertiana S. Brown in Torreya 7: 126 (1907).

Pinus alba sensu Hooker, Fl. Bor.-Am. 2: 163 (1839), in part, non Michx.

Picea canadensis sensu Preble in N. A. Fauna 27: 517 (1908), non B. S. P.

Picea glauca Voss, Hultén in Fl. Alaska & Yuk. Pt. 1: 80 (1941), in part.

Mackenzie R., Onion (N); rocky cliff along gorge east of Brintnell L., tree 10’ high, 2”
D. B. H., 9580; stony fan at camp, Brintnell L., tree 65’ high, 14” D. B. H., no. 9663;
Sekwi R., Canol Rd., mi. 174 E., Porsild 11972 (Can.) ; headwaters of Sekwi R., mi. 180 E.,
alt. 3900’, Porsild 11840 (Can.).

This is the commonest tree in the Brintnell Lake region. It forms rather
dense forests on the rocky talus slopes about the lake and grows abundantly on
the gravelly floor of the valley west of the lake. On some rocky and sandy
knolls about the eastern end it makes an open, park-like wood with a lichen-heath
mat on the ground. On higher mountain slopes it extends into the alpine zone
farther than any of the other trees, gradually becoming stunted and reduced to
the proportions of a depressed shrub. On the north slope of Colonel Mt. it
clothes the larger rocky fans with a thick timber 50 to 70 feet high, but gives
way to P. mariana on the steeper slopes between the fans. It reappears at about
3500’, however, and makes the upper fringe of the timber, along with occasional
patches of Abies. On the fans and south-facing slopes north of the lake it is
mixed with P. glauca var. Porsilditi on the drier soils, but forms nearly pure
stands on moister sites such as occur at the bases of the mountains. Although
tall and straight, most of the trees are spirally twisted, and many have eccentric
cores especially at the base. This is due to their situation on steep slopes which
are subject to land-slip and snow-slides.

Picea glauca var. albertiana is the characteristic tree over large portions of
the central Mackenzie basin. It is clearly recognizable in the country imme-
diately south of the timber line and on the foothills of the Rockies, but in the
central lowlands and plateaus it appears to merge with typical P. glauca (see
Raup, 1946, for further discussion of its distribution).

Picea glauca Voss var. Porsildii var. nov. Prate XII.

Arbor ad 15-17 m. alta habitu conica vel anguste ovata, ramis inferioribus
exceptis paene horizontalibus, trunco basim versus ad 30 cm. diametro; cortice
pallido-canescente fissuris paucis longitudinalibus basim arboris versus exceptis
sublevi, pustulis resinosis multis ornato; ramulis glabris_pallido-viridibus ;

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 103

squamis geramarum margine tenuibus laceratisque ; foliis linearibus 9-22 mm.
longis glauco-viridibus, sterigmatibus brevibus erectis vel satis divergentibus ;
conis 2.5-5 cm. longis, maturitate ad 3 cm. crassis; bracteis obovatis vel late
cuneatis, apice rotundatis vel truncatis, integerrimis; squamis tenuibus, 3-45
mm, longis, 2 mm. latis, rotundatis, obtusis vel subdeltoideis, apice interdum
emarginatis, margine laceratis.

Stony fan at Camp Point, Brintnell L., tree 35’ high, 9666 (Type) ; same, tree 55’ high,
9664; same, tree 40’ high, 9665; mud flat near west end of lake, tree 20’ high, 44%” D. B. H.,
9803; near shore of Great Slave L., Hay River, 9984, 9985.

This tree is most nearly related to P. glauca and its northern variety alber-
tiana, and since it appears to differ from typical P. glauca in no essential char-
acters of fruit, it seems best to designate it as a geographic variety. It differs
from P. glauca var. albertiana chiefly in its conical or narrowly ovate outline
and in its relatively smooth, balsam-like bark beset with resin blisters. Variety
albertiana is very narrowly pyramidal or even linear in form, spire-like, and its
bark is rough and flaky. At Brintnell L. the two are growing side by side on
the stony fans, particularly on the north shore, and are sharply contrasted (see
Pi. XII). In characters of cone, branchlet, and leaf, the differences are not so
apparent, although the cone-scales of var. Porsildti tend to be more rounded in
outline, and less wedge-shaped toward the base than those of var. albertiana;
also the bracts of the former are shorter and more angular toward the apex than
in var. albertiana. From typical P. glauca it differs principally in its smooth
bark and, again, in its short, angular scales (in P. glauca they are up to 5 mm.
long).

Variety Porsildii is abundant on the drier mountain slopes and stony fans
of southern exposure about Brintnell L. It occurs on the south shore, but only
on stony fans near the water’s edge. On dry talus slopes it extends up to about
4000’ north of the lake. In the neighborhood of Fort Simpson it was observed
on the bluffs and uplands west of the Mackenzie River; and it also appeared
to be common on the south shore of Great Slave L. at Hay River. There are
two specimens in the Herbarium of the Arnold Arboretum collected by J. G.
Jack in the Jasper Park district which match our northern material. Both have
pieces of the nearly smooth bark attached. They are as follows: Athabaska R.
bank, near Jasper Park Lodge, 35’ high, 5-6” diameter, “Blisters in bark,” Sept.
8, 1922, 2807; Maligne R. above gorge, 45’ high, 6” diameter, “Large resin
blisters on bark,” Sept. 4, 1922, 2719. A third specimen, collected by Prof.
Jack at Lake Beauvert, 2669, is described as having “bark nearly smooth or but
slightly scaling,” but the twigs are decidedly pubescent! It is from a tree 40’
high and 8” in diameter. No trees with pubescent twigs were found in the
Mackenzie region.

Abies balsamea was reported on the upper Mackenzie by Richardson (Arct.
Search. Exp. 2: 316), with the following statement: it “. . . was traced beyond
the 62nd parallel on the canoe route.” The writer has seen no specimens, how-
ever, from north of the Athabaska delta; and this is consistent with Preble’s
findings. It is not impossible that the Mackenzie record by Richardson was
based upon observations of P. glauca var. Porsildii, the bark of which so closely
resembles that of the balsam. It should be noted also that var. Porsildii has
proved to be common along the Alaska Highway in southern Yukon and northern
British Columbia.

104 SARGENTIA hte

Picea mariana (Mill.) B. S. P. Prel. Cat. N. Y. 71 (1888); Preble in N. Am. Fauna 27:
518 (1908); Hultén, Fl. Alaska & Yuk. Pt. 1: 82 (1941); Porsild in Nat. Mus.
Can. Bull. 101: 14 (1945).

Abies mariana Mill. Gard. Dict. ed. 8, no. 5 (1768).

Pinus nigra sensu Hooker, Fl. Bor.-Am. 2: 163 (1839), non Ait.

Damp mossy woods near shore of Brintnell L., tree 30’ high, 5’ D. B. H., 9565.

Common in muskegs and low woods about Brintnell L. and ascending to
about 3500’ on the steep north-facing mountain slopes just south of the lake.

Here it grows in an open park-like stand with a thick mat of mosses and fruti-

cose lichens on the ground, and a rather open shrub layer of willows and dwarf

birches, At the higher levels it becomes stunted and tends to reproduce by
layering, sending out numerous “branches” through the dense moss-lichen mat.

Noted by Preble (1908, p. 518) “. . . northward nearly to the limit of the

forest. . . ,” and in Fl. Bor.-Am. (2: 164, as Pinus nigra) “. . . throughout

Canada, to lat. 65°... .” Porsild makes the following comment on its dis-

tribution along the Canol Rd., “Noted as common on the eastern end of the

road, but becoming scarce in the mountains where no extensive, black spruce

forest was seen. Last trees were noted on Trout Creek about mile 120 E.”

Abies lasiocarpa (Hook.) Nutt. N. Am. Sylva 3: 38 (1849); Hultén, Fl. Alaska &
Yuk. Pt. 1: 87 (1941); Porsild in Nat. Mus. Can. Bull. 101: 14 (1945).

Pinus lasiocarpa Hooker, Fl. Bor.-Am. 2: 163 (1839).

Upper valley of Frost Cr., north of our camp at Brintnell L., alt. 4500’-4800’, small trees
up to 10’ high, 2’-3’’ D. B. H., 9527; north slope of Colonel Mt., alt. about 4000’, 9376;
hogback on north slope of Colonel Mt., alt. 4200’ (bark only), 9602; Macmillan Pass, Canol
Rd., mi. 225 E., Porsild 11184 (Can.).

The alpine fir as found at Brintnell Lake displays some variation from its
usual form, and with further study it may justifiably be segregated as a geo-
graphic variety. It appears to differ from typical A. lasiocarpa in having very
small cones (2.5 to 5 cm. long) with correspondingly small scales. The wings
of the seeds are usually smaller, and are broader in proportion to length than in
A, lasiocarpa (6% X 884 mm. vs. 7 X 12 mm.); also they are more rounded
at the base, often cordate at their outer insertion on the seed, while in the typical
form they are usually cuneate in outline. Leaves of the fruiting twigs are acute
to rounded, or even slightly emarginate, not sharp-pointed as in the species;
leaves of sterile twigs are strongly emarginate.

Common at timber line, where it is usually surrounded by the dwarf birch
scrub (Betula glandulosa) and grows in small clumps with much dead wood in
them. On high, wind-swept slopes the plants are distorted and dwarfed.

The northern limit of the range of Abies lasiocarpa is usually given as about
lat. 61° in Alaska and Yukon (Sargent, Silva 12: 113). There is a specimen
in the Arnold Arboretum Herbarium collected by G. M. Dawson in July, 1887,
and labeled, “Mts. above Finlayson.” This is presumably Finlayson Lake, on
the divide between the Francis and Pelly River valleys. It is in lat. 61° 40’.
Dr. Dawson states (Geol. Surv. Can. 3: 118 B, 1888), that “On the ridges west
of the head of Finlayson Lake Abies subalpina becomes moderately abundant.”
Our collections, therefore, do not extend the range far northward (62° 5’), but
they establish the species in the isolated Mackenzie Mountain system to the

eastward.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 105

Larix laricina (Du Roi) Koch var. alaskensis (Wight) comb. nov.

Larix alaskensis Wight in Smithson. Misc. Coll. 50: 174 (1908).

Larix laricina sensu Porsild in Nat. Mus. Can. Bull. 101: 14 (1945).

Muskeg at eastern end of Brintnell L., tree 20’ high, 4” D. B. H., 9575; Godlin R., Canol
Rd., mi. 142 E, alt. 3600’, Porsild 11839 (Can.) ; Twitya R., Canol Rd., mi. 124 E., Porsild
11837 (Can.).

Common in muskegs about the eastern end of Brintnell L., but not found
elsewhere in that vicinity.

Larix alaskensis was described by W. F. Wight (Smithson. Misc. Coll. 50:
174, pl. 17, 1907) from material collected at Tanana, Alaska, by A. J. Collier.
Recent students have considered it synonymous with L. laricina (Porsild, Contr.
Fl. Alaska, Rhod. 41: 174, 1939; Rehder, Man. Cult. Trees and Shrubs, ed. 2, p.
98, 1940), following Ostenfeld and Larsen (Kgl. Danske Videnskab. Selsk. Biol.
Meddel. 9: 85-95, 1930). The latter authors did not see the type of L. alasken-
sis, but from specimens collected at Ft. Gibbon on the Yukon R. they confirmed
the characterization given by Wight: “The cone scales are longer [than in L.
laricina] in proportion to their breadth; the bracts of the cone scales are ovate
and without a projecting mucronate point at the apex, while L. laricina has
bracts short-oblong to nearly orbicular in outline, and commonly emarginate or
lacerate on either side of a mucronate projection at the apex.” Ostenfeld and
Larsen, however, did not believe that the characters given were sufficient to
separate the northwestern plants from the typical species. They cited similar
material from Lake Superior and from Churchill, Hudson Bay, stating that the
alaskensis type would better be considered as a “form of growth.”

Although he has not given it separate status, Hultén (Fl. Alaska and Yuk. Pt.
1: 79-80) recognized the existence of an Alaskan form of the American larch,
and states that he has not seen any specimens of the common southern type in
Alaska. I have examined a large series of specimens from most of the range
of the species, as well as a part of the type of L. alaskensis, and have concluded
that the northwestern form is at least worthy of being maintained as a geographic
variety. To the differences noted by Wight should be added another: the outer
margins of the bracts in typical L. laricina are firm, with the purplish color ex-
tending nearly or quite to the edge, while in var. alaskensis they are papery or
chartaceous, and usually yellowish-hyaline, with a tendency to become inrolled
in the mature cones. It is probably true, as Ostenfeld and Larsen have pointed
out, that var. alaskensis will appear far eastward in the range of the species,
particularly in northern latitudes. Specimens collected by the writer at Tal-
theilei Narrows on Great Slave L. (no. 90, cited under L. laricina in Phytogeog.
Studies Atha».—Great Slave L. Reg. I. p. 194) belong to it, as well as material
collected on Lake Athabaska (no. 4663 1. c.) and in the Wood Buffalo Park
(no. 1517, Bot. Invest. Wood Buff. Pk. p. 98). Labrador specimens in the
Gray Herbarium show the same tendency, though apparently not so marked as
in those from the northwest. It is probable that far northwestern material re-
ferred to in Fl. Bor.-Am. (2: 164, under Pinus microcarpa), “From York
Factory to Point Lake, in lat. 65°. . . ,” also belongs here.

Pinus Banksiana Lamb. Pinus 1: pl. 3 (1803); Hooker, Fl. Bor.-Am,. 2: 161 (1839);
Macoun, Cat. Can. Pl. Pt. 1: 468 (1884).

Pinus divaricata Du M. de C.; Preble in N. Am. Fauna 27: 516 (1908).

Common in the central and southern parts of the Mackenzie basin (Raup,
1935, 1936, 1942), but records for the country below Great Slave L. are yet

106 SARGENTIA [6

based only upon observations. Noted in Fl. Bor.-Am. from “. . . Canada, to
lat. 64°, and even beyond that, on the sandy banks of the Mackenzie River.”
There is a Richardson specimen in Herb. Can. (no. 25131), but no locality is
given. The species was reported by Dawson (after McConnell) on the lower
Liard R. in 1887. Preble has the following notes on the northwestern exten-
sions of the jack pine: “It occurs commonly on the Simpson Islands and about
the Northern Arm of Great Slave Lake. On my route to the northward of
Fort Rae it was seen in many places on Grandin River, reaching a diameter of
18 inches, but not growing very tall. North of the height of land it was rare,
but I saw many trees on a portage a few miles north of the outlet of Lake
Hardisty.

“On the Mackenzie it is common north to about latitude 64° 30’. . . . On the
north slope of the Nahanni Mountains, 75 miles below Fort Simpson, the species
ascends to about 1,000 feet and then becomes a dwarfed shrub and disappears.
On the southern slopes of the same mountains it occurs as a fairly well-grown
tree to about 2,000 feet, near the summit.”

Pinus contorta Loudon var. latifolia Engelm. ex Wats. in King, Rep. U. S. Geol. Expl.
40th Par. 5: 331 (1871); Hultén, Fl. Alaska & Yuk. Pt. 1: 77 (1941).

Pinus Murrayana Balf. Rep. Bot. Oreg. Exped. 2 (1853).
Pinus contorta Loudon var. Murrayana (Balf.) Engelm. in Wats. Bot. Calif. 2: 126
(1880).

As noted elsewhere in this paper, the occurrence and distribution of the lodge-
pole pine in southwestern Mackenzie is still to be discovered. It is presumed
that the lodgepole and jack pines overlap somewhere in the lower Liard country
and on the foothills of the northern Rockies. Pines in the Liard valley above
the “Liard Gap” all appear to be lodgepole, and it is supposed that those in the
Mackenzie Mountains west of the South Nahanni R. are also of this species.
No pines of any kind were seen at Brintnell L., and Porsild makes no mention
of any along the Canol Road east of Macmillan Pass until he reaches the
Mackenzie valley, where Pinus Banksiana appears.

Juniperus communis L. var. montana Ait. Hort. Kew. 3: 414 (1788); Hultén, FI.
Alaska & Yuk. Pt. 1: 89 (1941); Porsild in Nat. Mus. Can. Bull. 101: 30 (1945),
in part; Raup in Jour. Arn. Arb. 17: 195 (1936).

Juniperus communis var. nana Hooker, Fl. Bor.-Am. 2: 165 (1839).

Juniperus communis L. sensu Porsild in Nat. Mus. Can. Bull. 101: 14 (1945), in part.

Juniperus nana sensu Preble in N. Am. Fauna 27: 519 (1908), non Willd.

Bear R., Wood Buffalo Park, Russell 38 (G); Resolution, Great Slave L., E. A. Preble
206 (US); Wrigley, Crickmay 148 (Can.) ; hogback on north slope of Colonel Mt., Brint-
nell L. area, alt. 4300’, 9603; north peak of Nahanni Mt., Wynne-Edwards 8423 (Can.) ;
Nahanni Mts., Preble & Cary 68 (US).

Common on dry rocky slopes around Brintnell L. usually above timber line
(between 3800’ and 4500’), where it often occurs as a primary species in the
rather open communities. Noted in Fl. Bor.-Am. from Lake Huron “to the
barren country of the Hudson Bay Co.’s territories.” Undoubtedly common
throughout southwestern Mackenzie, though collections of it are rare. Porsild
noted it as common in many places along the Canol Rd.

Juniperus horizontalis Moench. Meth. 699 (1794); Hultén, Fl. Alaska & Yuk. Pt. 1:
91 (1941); Porsild in Nat. Mus. Can. Bull. 101: 30 (1945).

Juniperus sabina sensu Preble in N. Am. Fauna 27: 519 (1908), non L.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 107

Wrigley, Crickmay 149 (Can.); north peak of Nahanni Mt., Wynne-Edwards 8424
(Can.) ; Nahanni Mt., Preble & Cary 73 (US).

Preble states that this species extends “throughout the forested belt,’ but the
above are the only specimens I have seen from below Great Slave L.

TYPHACEAE

Typha latifolia L. Sp. Pl. 971 (1753); Hooker, Fl. Bor.-Am. 2: 170 (1839); Macoun,
Cat. Can. Pl. Pt. 4: 69 (1888).

Although this species is noted in Fl. Bor.-Am,. as occurring “Throughout
Canada to Fort Franklin,” there is no other evidence of its presence on the upper
Mackenzie. It is common, however, in the Wood Buffalo Park and is to be
expected around the western arm of Great Slave L.

“er

SPARGANIACEAE

Sparganium eurycarpum Engelm. in Gray, Man. ed. 2, 430 (1856); Porsild in Sargentia
4: 9 (1943). See Rhod. 24: 26 (1922).
Common at the margins of ponds and slow streams in the Athabaska—Peace
delta region, and collected by Porsild along the Mackenzie R. near Norman. It
probably occurs at least sporadically throughout the upper Mackenzie lowland.

Sparganium multipedunculatum (Morong) Rydb. in Bull. Torr. Bot. Cl. 32: 598 (1905);
Raup in Jour. Arn. Arb. 17: 195 (1936). See Rhod. 27: 190 (1925).

Sparganium simplex var. multipedunculatum Morong in Bull. Torr. Bot. Cl. 15: 79
(1888).

Sparganium simplex sensu Hooker, Fl. Bor.-Am, 2: 169 (1839), in part; Macoun, Cat.
Can. Pl. Pt. 4: 70 (1888) in part; non Huds.

Fort Franklin, Mackenzie R., Richardson (N).

Common in the central and southern parts of the Mackenzie basin, and noted
in Fl. Bor.-Am. “Throughout Canada to Fort Franklin, abundant.” No mate-
rial is yet available, however, from Great Slave L. or the upper Mackenzie valley.

Sparganium angustifolium Michx. Fl. Bor.-Am. 2: 189 (1803); Raup in Jour. Arn. Arb.
17: 196 (1936); Hultén, Fl. Alaska & Yuk. Pt. 1: 92 (1941); Porsild in Sargentia
4: 8 (1943).

Locally abundant in ponds and slow streams in the Athabaska-Great Slave L.
region, and collected by Porsild at Great Bear L. and in the Eskimo Lakes basin.
Probably widespread in the upper Mackenzie valley, though no specimens are
available.

Sparganium minimum Fries, Summa Veg. Scand. 2: 560 (1849); Hultén, Fl. Alaska &
Yuk. Pt. 1: 94 (1941).

Mackenzie 8., Onton (N).

The above specimen constitutes the only record for the species north of Lake
Athabaska, where it is common. The actual locality of the Mackenzie collection
is unknown.

POTAMOGETONACEAE

Potamogeton filiformis Pers. var. borealis (Raf.) St. John in Rhod. 18: 134 (1916);
Raup in Jour. Arn. Arb. 17: 198 (1936); Hultén, FI. Alaska & Yuk. Pt. 1: 99
(1941).

Potamogeton borealis Raf. Med. Repos. 2: 354 (1808).
Northwest shore of Great Slave L., Bedford (Can.).

108 SARGENTIA [6

Potamogeton vaginatus Turcz. in Bull. Soc. Nat. Hist. Mosc. 11: 102 (1838); Hultén,
Fl, Alaska & Yuk. Pt. 1: 104 (1941). See Rhod. 18: 131 (1916) and 20: 191
(1918).

Though not yet collected in our region, this species is very common in the

Wood Buffalo Park area and has been collected at Great Bear L. (Porsild, G).

Potamogeton pectinatus L. Sp. Pl. 127 (1753); Hultén, Fl. Alaska & Yuk. Pt. 1: 101
(1941),
Shallow water at lower end of Bluefish Island, Mackenzie R., near Simpson 9958, 9960.
Abundant in this situation, and probably widespread in the Mackenzie valley.
Collected by Porsild in the Mackenzie delta (6913, 6915, 7244, Can.).

Potamogeton foliosus Raf. var. macellus Fernald in Mem. Gray Herb. 3: 46 (1932).
Collected in the Wood Buffalo Park (Raup 1562, 132, 1561) and at Great
Bear L. (A. E. & R. T. Porsild), but not yet found on the upper Mackenzie.

Potamogeton Friesii Rupr. Beitr. Russ. Reich. 4: 43 (1845); Fernald in Mem. Gray
Herb. 3: 52 (1932); Hultén, Fl. Alaska & Yuk. Pt. 1: 99 (1941).
Not yet collected along the upper Mackenzie, but known in the Wood Buffalo
Park (Raup 1564) and at Great Bear L. (A. E. & R. T. Porsild Can., G).

Potamogeton alpinus Balbis var. tenuifolius (Raf.) Ogden in Contr. Gray Herb. 147:
90 (1943).
Potamogeton tenuifolius Raf. Med. Repos. Lex. 3, 2: 409 (1811).
Potamogeton alpinus Balbis ssp. tenuifolius (Raf.) Hultén, Fl. Alaska & Yuk. Pt. 1: 98
(1941).

This species has been collected in the Athabaska-Great Slave L. region (Raup
6498a, 6618, 6621, 6622, 6741, 6742), at Great Bear L. (A. E. & R. T. Porsild),
and in the Eskimo Lakes basin (4. E. & R. T. Porsild), but it has not yet been
observed on the upper Mackeizie.

Potamogeton gramineus L. var. typicus Ogden in Contr. Gray Herb. 147: 143 (1943);
Hultén, Fl. Alaska & Yuk. Pt. 1: 100 (1941).
Potamogeton gramineus L. var. graminifolius Fries, Novit. Fl. Suecicae 36 (1828) ; Raup
in Jour. Arn. Arb. 17: 196 (1936).
Northwest shore of Great Slave L., Bedford (Can.) ; margin of lowland slough near west
bank of Mackenzie R. near Fort Simpson, 9894,
Sterile plants only, creeping on damp muck, collected Aug. 24. Common in
this situation, but not seen elsewhere in our region. Common in the Wood
Buffalo Park, and collected at Great Bear L. by A. E. & R. T. Porsild.

Potamogeton gramineus L. var. maximus Morong ex Bennett in Jour. Bot. 19: 241
(1881). See Ogden in Contr. Gray Herb. 147: 148 (1943).

Known in the Wood Buffalo Park (Raup 1546, 1548) and at Great Bear L.
(A. E.& R. T. Porsild), but not yet found on the upper Mackenzie.
Potamogeton praelongus Wulf. in Arch. Bot. Roem. 3: 331 (1805); Raup in Jour. Arn.

Arb, 17: 197 (1936); Hultén, Fl. Alaska & Yuk. Pt. 1: 103 (1941). See Ogden
in Contr. Gray Herb. 147: 159 (1943).

Common in the Athabaska—Great Slave L. region (Raup 1570, 1571, 1572,
133), and found in the Eskimo Lakes basin (A. E. & R. T. Porsild), but not
yet collected on the upper Mackenzie.

Potamogeton Richardsonii (Bennett) Rydb. in Bull. Torr. Bot. Cl. 32: 599 (1905);
Raup in Jour. Arn. Arb. 17: 197 (1936).
Potamogeton perfoliatus var. Richardson Bennett in Jour. Bot. 27: 25 (1889).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 109

Potamogeton perfoliatus ssp. Richardsonii (Bennett) Hultén, Fl. Alaska & Yuk. Pt. 1:
102 (1941). ‘
Northwest shore of Great Slave L., Bedford (Can.); Ft. Rae, Bedford (Can.); shallow
water at lower end of Bluefish Island, Mackenzie R. near Simpson, 9959; Liard R., between
Nahanni Butte and Simpson, Crickmay 5 (Can.).

Common at the Bluefish Island station. In fruit late in August.

SCHEUCHZERIACEAE

Triglochin maritimum L. Sp. Pl. 339 (1753); Hooker, Fl. Bor.-Am. 2: 168 (1839); Raup
in Jour. Arn. Arb. 17: 198 (1936); Hultén, Fl. Alaska & Yuk. Pt. 1: 105 (1941).
North of the height of land between Great Slave and Great Bear Lakes, Bedford (Can.).
Noted by Hooker from “Cumberland House Fort ... to the Mackenzie
River”; and there is a specimen in Herb. N collected by Onion which probably
came from the Mackenzie, though it is without locality data. Collected by
A. E. & R.T. Porsild at Great Bear L. (3261, 3671, 4664, 5089, 5165) and in

the Mackenzie delta (1899, 2301).

Triglochin palustre L. Sp. Pl. 338 (1753); Hultén, Fl. Alaska & Yuk. Pt. 1: 106 (1941);
Porsild in Nat. Mus. Can. Bull. 101: 33 (1945).
Rocky-sandy bank of Mackenzie R., opposite Simpson, 9951; shore of Mackenzie R. 10
mi. below Mills L., Wynne-Edwards 8545 (Can.).

Occasional; found fruiting late in August.

ALISMACEAE
Sagittaria cuneata Sheldon in Bull. Torr. Bot. Cl. 20: 283 (1893); Raup in Jour. Arn.
Arb. 17; 198 (1936).
Northwest shore of Great Slave L., Bedford (Can.).

GRAMINEAE
Bromus ciliatus L. Sp. Pl. 76 (1753); Raup in Jour. Arn. Arb, 17: 199 (1936); Hulten,
Fl. Alaska & Yuk. Pt. 2: 249 (1942).
Bromus purgans L. var. vulgaris Hooker, Fl. Bor.-Am. 2: 252 (1840), in part.
Resolution, Kennicott (N); Fort Norman, Richardson (G).

Bromus Pumpellianus Scribn. in Bull. Torr. Bot. Cl. 15: 9 (1888); Hultén, Fl. Alaska
& Yuk. Pt. 2: 251 (1942); Porsild in Nat. Mus. Can. Bull. 101: 34 (1945). See
Porsild in Rhod. 41: 182-183 (1939).

Upland woods west of the Mackenzie R. near Simpson, 9935; Saline R., above Norman,
W ynne-Edwards 8567 (Can.).

Common; spikelets maturing late in August. Some of the glumes on these
specimens are hairy, suggesting var. arcticus (Shear) Porsild (1. c.), but other
characters of this variety, such as very hairy sheaths and upper leaf-surfaces, are
absent.

Festuca brachyphylla Schultes, Mant. 3 (Add. 1): 646 (1827); Hultén, Fi. Alaska &
Yuk. Pt. 2: 241 (1942); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945). See
Fernald in Rhod. 37: 250-252 (1935).

Stony beach at Camp Pt., Brintnell L., 9176; gravelly thickets near lake shore at Camp
Pt., 9368; dry sandy plain near west end of Brintnell L., 9451; shaly south slope of Terrace
Mt., alt. 4000’-5000’, 9669, 9678; turfy ledges on summit of Red. Mt., alt. 5900’, 9771; shale
slopes on south side of Red Mt., alt. 5000’, 9783; shale cliffs in gorge on north slope of
Colonel Mt., alt. 3500’, 9822; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8237
rian,).

Occasional to common locally ; in anthesis late in June, and mature in August.
This species is orfe of the principal ones forming the open association on the very

110 SARGENTIA [6

summit of Red Mt. It was not seen on Colonel Mt., however, except in the
deep gorge on the lower north slope.

Festuca rubra L. Sp. Pl. 74 (1753); Raup in Jour. Arn. Arb. 17: 199 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 2: 244 (1942). See Rhod, 35: 132-135 (1933).
This species has not been collected on the upper Mackenzie, but it is known
from Lake Athabaska (Raup 6723A, 6905, 6941A) and at Great Bear L. (J. M.
Bell, Can.).

Festuca rubra L. var. arenaria (Osbeck) Fries, Fl. Hall. 28 (1817); Raup in Jour. Arn.
Arb. 17: 199 (1936). See Rhod. 35: 132-135 (1933).

Festuca arenaria Osbeck in Retz. Sup. Prodr. Fl. Scand. 1: 4 (1805).

Like the last, this variety is known at Lake Athabaska (Raup 171, 4432, 4573,
6713, 6723, 6900, 6941), and at Great Bear L. and the arctic coast (Richardson,
G), but has not yet been collected between.

Festuca saximontana Rydb. in Bull. Torr. Bot. Cl. 36: 536 (1909); Raup in Jour. Arn.
Arb. 17: 199 (1936). See Rhod. 37: 250-252 (1935).

Festuca ovina sensu Hooker, Fl. Bor.-Am. 2: 250 (1840), in part; Macoun, Cat. Can. Pl.
Pt. 4: 235 (1888), in part, non L.

Festuca brachyphylla Schultes ssp. saximontana (Rydb.) Hultén, Fl. Alaska & Yuk. Pt.
2: 242 (1942).

Common in the Athabaska—Great Slave Lake region and far northward, though
no material is yet available from the upper Mackenzie region. Noted in FI.
Bor.-Am. northward “. . . to Bear Lake,” and there are Richardson specimens
from the latter place in Herbs. G and N.

Festuca altaica Trin. in Ledeb. FI. Alt. 1: 109 (1892); Hultén, Fl. Alaska & Yuk. Pt.
2: 239 (1942); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).

Mackenzie R., Wrigley to Blackwater R., Crickmay, no. 127 (Can.); and from Black-
water to Norman, Crickmay, no. 154 (Can.); along small brook on east slope of Red Mt.,
alt. 4500’, 9278; alpine meadows and scrub on north slope of Colonel Mt., alt. 4000’-6000’,
9396, 9615; open spruce woods on rocky knoll near east end of Brintnell L., 9576; turfy
ledges on summit of Red Mt., alt. 5900’, 9792; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-
Edwards 8236 (Can.).

Abundant above timber line, and occasional at lower elevations; in anthesis
late in June. This is one of the most abundant grasses in the Brintnell L. dis-
trict. It is a primary species in most of the alpine meadows, where it is made
conspicuous, even at a distance, by its habit of forming large tussocks in which
there remain a great many dry dead leaves. Its tall culms and panicles are
often dark purplish red. The lemmas of our plants are very scabrous, almost
hairy, thus resembling those of Alaskan specimens rather than those of northern
Rocky Mt. material. Plants from the upper Peace R. Rockies (Raup & Abbe,
no, 3956) have nearly smooth lemmas. Porsild states that Festuca altaica is a
dominant species in Macmillan Pass.

Glyceria grandis Wats. in Gray, Man. ed. 6: 667 (1890).

Glyceria maxima (Hartm.) Holmb. ssp. grandis (Wats.) Hultén, Fl. Alaska & Yuk. Pt.
2: 229 (1942).

Resolution, Kennicott (N).

Glyceria pulchella (Nash) K. Schum. in Just, Bot. Jahresb. 29, Pt. 1: 513 (1903); Raup
in Nat. Mus. Can. Bull. 74: 103 (1935); Hultén, Fl. Alaska & Yuk. Pt.-2'-o82
(1942). i

Panicularia pulchella Nash in Bull. N. Y. Bot. Gard. 2: 157 (1901).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 111

Along the lower little Buffalo R., Russell 44 (G); damp slough in lowland woods near
Simpson, 9840; grassy swale near Ft. Liard, Nowosad (Can.).

Common in the Simpson habitat ; spikelets mature in the latter part of August.
Not previously noted in the Mackenzie basin north of the Wood Buffalo Park.

Puccinellia Nuttalliana (Schult.) Hitche. in Jepson, FI. Calif. 1: 162 (1912); Raup in
Jour. Arn. Arb. 17: 200 (1936).

Poa Nuttalliana Schult. Mant. 2: 303 (1824).
Poa airoides Nutt.; Hooker, Fl. Bor.-Am. 2: 245 (1840).

Great Slave L., Richardson (G); southwest and north shores of Great Slave L., Howe
91994 (Can.).

Puccinellia Vahliana (Liebm.) Scribn. & Merrill in Contr. U. S. Nat. Herb. 13: 78
(1910); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).

Poa Vahliana Liebm. in Fl]. Dan. tab. 2401 (1845).

Plains of Abraham, Canol Rd., mi. 82 E., on bald, wind swept summit of limestone moun-
tain, alt. 6000’, in wet clay and limestone rubble, Porsild 11787 (Can.).

Poa alpina L. Sp. Pl. 67 (1753); Hooker, Fl. Bor.-Am. 2: 246 (1840); Macoun, Cat.
Can. Pl. Pt. 4: 222 (1888); Raup in Jour. Arn. Arb. 17: 201 (1936); Hultén, Fl.
Alaska & Yuk. Pt. 2: 198 (1942); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).

Mackenzie R., between Blackwater R. and Norman, Crickmay 155 (Can.); stony lake
shores at Brintnell L. Camp, 9168, 9572; high shale slopes and summit of Red Mt., alt. 5000’
5900’, 9781; creviced shale cliffs, north slope of Colonel Mt., 9606; damp mossy slopes on
old slide-rock, gorge on north slope of Colonel Mt., alt. 3200’, 9824; open spruce woods on
a stony fan, south shore of Brintnell L., 9828; Bolstead Cr., Canol Rd., mi. 111 E., wet alpine
meadow, alt. 5500’, Wynne-Edwards 8239 (Can.); same loc., alpine slope, 8238 (Gan.).:
noted by Porsild as common in Macmillan Pass, mi. 225 E.

Common above timber line, and occasional on gravelly lake shores; found in
anthesis about mid-June, and occasionally as late as mid-August. Extremely
variable in habit, especially at the lower elevations. Noted in FI. Bor.-Am.,
“From the elevated grounds of the Saskatchewan .. . to Bear Lake... .”

Poa pratensis L. Sp. Pl. 67 (1753); Hooker, Fl. Bor.-Am. 2: 246 (1840); Raup in Jour.
Arn. Arb. 17: 201 (1936).
Poa alpigena (Fr.) Lindm., sensu Hultén, Fl. Alaska & Yuk. Pt. 2: 197 (1942), in part;
Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).

East shore of Trout L., C. O. Hage (G); top of river bluff at Simpson, 9/21; cleared
field at Simpson, 9141; damp thickets west of Simpson, 9839, 9849; upland prairie west of
Simpson, 9898; gravelly willow-alder thickets along shore at Brintnell L. Camp, 9465, 9467;
damp stony beaches at Brintnell L. Camp, 9166; mossy shore of old stream channel at west
end of Brintnell L., 9495; damp mud flats near mouth of glacial stream at west end of Brint-
nell L., 9689; noted by Porsild in a number of places along the Canol Road between Mac-
millan Pass and Norman.

Occasional to common in the habitats cited above; in anthesis about mid-June
at Simpson, but not until the second week in July at Brintnell L. Noted in Fi.
Bor.-Am. from “Saskatchewan to Bear Lake.”

I am here considering Poa pratensis in its broad sense, including the northern
forms that have been called P. alpigena or P. pratensis var. alpigena. The ma-
terial that I have seen from southwestern Mackenzie is extremely variable, and
I have been unable to break it down satisfactorily into more than one entity, or
successfully to separate any of it from the common Poa pratensis of the central
and southern parts of the Mackenzie basin. If such a separation were made it
would be on artificial grounds.

Hultén evidently regards nearly all pratensis-like Poa in Alaska and Yukon
as P. alpigena, but notes (1. c. 198) that wherever other types in this group occur

112 SARGENTIA [6

“there seem to exist all kinds of intermediate forms.” True Poa pratensis he
regards as an introduced plant along the coasts (1. c. 219).

Poa pratensis is common throughout the central and southern parts of the
Mackenzie basin, in a variety of forms that merge imperceptibly with those of
the northern Great Plains. It is hardly conceivable that it is an introduced plant
in the Athabaska—Great Slave L. region. It is frequent in several habitats, but
has its greatest development in the wild upland prairies of the Wood Buffalo
Park. To consider it as introduced into these prairies would be to single it out
artificially and unjustifiably from the otherwise entirely native flora. The mate-
rial from southwestern Mackenzie seems to me to contain only variants in this
highly complex native population of pratensis-like forms. If a Poa alpigena
exists in the far northwest I suspect that it is connected with Poa pratensis in
the upper Yukon and Liard R. regions in such a way as to make it inseparable
as anything more than a variety.

It is of interest that Polunin (Nat. Mus. Can. Bull. 92: 73, 1940) arrived at
a similar conclusion with regard to eastern arctic Poa pratensis. There also he
found himself unable to effect a separation of Poa alpigena ftom the mass of
material as a whole.

Number 9727 of our collections at Simpson is worthy of remark. It was col-
lected from a “turf” on the much-frequented river bluff at Simpson. Here it is
grazed repeatedly and has a depauperate habit, with culms only 4-10 inches tall,
and with short, spreading leaves mostly rising from the base.

Poa arctica R. Br. in Suppl. App. Parry’s Ist Voy. 288 (mispaged 188) (1824); Hooker,
Fl. Bor.-Am. 2: 246 (1840); Hultén, Fl. Alaska & Yuk. Pt. 2: 201 (1942); Porsild
in Nat. Mus. Can. Bull. 101: 15 (1945),

Poa flexuosa sensu Hooker, Fl. Bor.-Am. 2: 245 (1840), in part, non Wahl,

Poa cenisia sensu Macoun, Cat. Can. Pl. Pt. 4: 224 (1888), non All.

Gravelly thickets on shore of Brintnell L. at Camp Pt., 9177, 9178, 9469; dry shale slopes
on south side of Red Mt., alt. 5000’, 9747, 9782; in willow-birch scrub, upper valley of Frost
Cr., north of Brintnell L. Camp, alt. 4000’, 9532; turfy ledges on summit of Red Mt., alt.
5900’, 9796, 9797, 9798, 9799; dry gravelly watercourse on N. slope of Colonel Mt., alt. 5000’,
9621; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5500’, Wynne-Edwards 8240,
8241 (Can.) ; noted by Porsild as common in Macmillan Pass, Canol Rd., mi. 225 E.

Occasional to common in the habitats noted, except on the summit of Red Mt.,
where it is abundant as a primary species on some of the turfy ledges. Collected
in anthesis on the lake shore as early as June 20, and on the mountain summit as
late as Aug. 7. Noted in Fl. Bor.-Am. as occurring from the “Rocky Mountains
to the Arctic Sea-shores and Islands in every direction.” At least a part of the
material cited in Fl. Bor.-Am. under P. flexuosa belongs here.

Extremely variable in size and habit, but fairly consistent as a species through-
out our collections. All of our material appears to represent ssp. Williamsii
(Nash) Hultén, Fl. Alaska & Yuk. Pt. 2: 202 (1942). This subspecies, how-
ever, seems to me of doubtful significance when the great variability of P. arctica
in the distinguishing characters throughout its range is considered. For discus-
sions of this variability see Hultén, 1. c., and Polunin in Bot. Can. E. Arct. 71-72.

Poa Brintnellii sp. nov. Fic. 11.

Gramen perenne 5-8 dim. altum, rhizomatibus bene evolutis ; inflorescentia late
pyramidata 6-12 cm. longa, diffusa, ramulis inferioribus plerumque binis, ramis
apice vel apicem versus tantummodo spiculiferis; culmis et inflorescentiis glabris
flexuosis; laminis 1.5—2.5 mm. latis ad 25 cm. longis, planis, laxis, superioribus

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 113

brevibus (3 cm. longis), ligulis 1-2 mm. longis rotundatis, vaginis minute as-
peratis ; spiculis 6-7 mm. longis, 2.5-3 mm. latis, basi acutis, gluma prima 3 mm.
longa, secunda 4 mm. longa, lemmate 4.5 mm. longo ovato-oblongo acuto pur-
purascente scabro 5-nervio, nervis mediis et lateralibus lanatis, internervis glabris,
nervis intermediis satis conspicuis, lemmate basi arachnoideo; antheris 2 mm.
longis.

Open spruce woods on stony fan, south shore of Brintnell L., 9827 (Type) ; mossy banks
on an old shale slide in gorge, north slope of Colonel Mt., alt. 3300’, 9830. Common in the
habitats noted; spikelets maturing about mid-August.

= y: 2 “he ae at 2
et he Ke Le c d cecren eof :

Fic. 11. Poa Brintnellii Raup, sp. nov.

Judging by its well-developed rhizomes this plant is in the section Pratenses
and seems nearest to P. arctica, which has a spreading inflorescence with spike-
lets only at or near the tips of the branches. However, its glabrous internerves,
rather prominent intermediate nerves (especially in mature specimens), and its

114 SARGENTIA [6

tall sprangling habit distinguish it from P. arctica. Also the bases of spikelets
are not so much rounded as in P. arctica. In spikelet characters it somewhat
resembles P. paucispicula or P. leptocoma, but neither of these produces such
prominent rhizomes. Its inflorescence also resembles that of P. nervosa, but
the latter has no cobweb at the base of the lemma. Hultén has described a Poa
arctica ssp. longiculmis, in Fl. Alaska & Yuk. Pt. 2: 202 (1942), which has a
tall slender habit of growth. His plants have closely involute basal leaves, how-
ever, and the lemmas are somewhat hairy.

Poa palustris L. Syst. Nat. ed. 10, 2: 874 (1759); Hultén, Fl. Alaska & Yuk. Pt, 2: 218
(1942). See Rhod. 18: 235 (1916).

Common in an upland prairie west of Simpson, 9896; grassy swale near Ft. Liard, Nowo-

sad (Can.).

Poa nemoralis L. Sp. Pl. 69 (1753); Hultén, Fl. Alaska & Yuk. Pt. 2, 217 (1942).

Dry open woods on south-facing slope of Red Mt., above Brintnell L. Camp, alt. 3500’,
9676.

Common in places, but not widely distributed in the Brintnell L. area. Spike-
lets maturing early in August.

Poa leptocoma Trin. in Mém. Acad. St. Pétersb. VI. Math. Phys. Nat. 1: 374 (1830);
Hultén, Fl. Alaska & Yuk. Pt. 2: 215 (1942).

Mackenzie R., between Blackwater R. and Norman, Crickmay 153 (Can.); and between
Wrigley and Blackwater R., Crickmay 126 (Can.).

Poa paucispicula Scribn. & Merrill in Contr. U. S. Nat. Herb. 13: 69 (1910).

Poa leptocoma sensu Porsild in Nat. Mus. Can. Bull. 101: 15 (1945), non Trin.

Damp stony and sandy beaches near Brintnell L. Camp, 9171, 9674; shale cliffs and slide-
rock on the summit of Red Mt., alt. 5900’, 9768; Macmillan Pass, Canol Rd., mi. 225 E.,
Porsild 11194-A (Can.).

Occasional to common in the habitats noted, but not seen elsewhere ; in anthesis
late in June.

This species was originally described from Alaskan material collected by
Coville & Kearney at Yakutat Bay. Scribner and Merrill considered it to be
most closely related to P. cenisia All., in the section Pratenses, but Hitchcock
(Man. Gr. 120-121) has probably placed it more correctly in the section Palus-
tres, near P. leptocoma Trin. WHultén, in his Fl. Aleut. Isls. 87, has, in fact,
reduced it to P. leptocoma, considering, incorrectly I think, that it is only a
lower-growing alpine form of the latter (see also Hultén, Fl. Alaska & Yuk.
Pt. 2: 215-216, 1942). Poa leptocoma was described from material collected
at Sitka by Mertens, but its range, according to Hitchcock’s interpretation, is
much more southern than P. paucispicula, reaching to New Mexico and Cali-
fornia. Fragments of the type of P. leptocoma were seen by Scribner and
Merrill, and presumably by Hitchcock, at the U. S. National Herbarium and
at the Missouri Botanical Garden; and Scribner and Merrill related it to P.
reflexa. Alaskan and northern cordilleran specimens, as well as our own, have
the oblong, obtuse, purplish lemmas and rather broad spikelets of P. paucispicula.
In P. leptocoma (sensu Hitche.) the spikelets are narrower, with acuminate
lemmas; and in Gray Herbarium specimens the spikelets are greenish and not
so much confined to the ends of the inflorescence branches as in P. paucispicula.
With these things in view it seems best to maintain P. paucispicula as a distinct
species representing the group of Poa which has short rootstocks but lacks the
long creeping rhizomes of the Pratenses. In its inflorescence it shows the same

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE Lis

tendency toward reduction in the number of spikelets and their concentration
toward the ends of the branches that is seen in the P. arctica complex in the
section Pratenses.

Poa glauca Vahl in Fl. Dan. tab. 964 (1790); Raup in Jour. Arn. Arb. 17: 201 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 2: 208 (1942); Porsild in Nat. Mus. Can. Bull.
101: 15 (1945),

Poa laxa sensu Hooker, Fl. Bor.-Am. 2: 246 (1840), in part, non Haenke.

Top of river bluff at Simpson, 9120; dry southerly slope in abandoned field, Simpson,
9137, 9138, 9159; rocky-sandy shore of Mackenzie R., opposite Simpson, 9967, 9968; Macken-
zie R., between Blackwater R. and Norman, Crickmay 152 (Can.); Norman, Richardson
(G, N) ; damp stony shores and gravelly thickets at Brintnell L. Camp, 9166-A, 9470; muskeg
thicket at west end of Brintnell L., ? 9710 (fragment) ; ledges on shale cliffs in gorge east
of Brintnell L., 9582; shale slopes on south side of Red Mt., alt. 5000’-5500', 9775, 9784, 9794;
shale cliffs in gorge on north slope of Colonel Mt., alt. 3500’, 9833; Bolstead Cr., Canol Rd.,
mi. 111 E., alpine slope, alt. 5500’, Wynne-Edwards 8242 (Can.).

Occasional to common, usually in dry exposed places. It is abundant in
clearings anc on river banks around Simpson. In early anthesis about mid-June.
Material cited in Fl. Bor.-Am. as P. Java, at least in part, from “Hudson’s Bay
to the Arctic Islands,” belongs here.

Poa Buckleyana Nash in Bull. Torr. Bot. Cl. 22: 465 (1895).

East shore of Trout L., C. O. Hage (G); southerly slope of Red Mt., alt. 3500’—4000’,
9668, 9776; dry stony ridge on south slope of Terrace Mt., alt. 4000’, 9670.

Occasional to common in the upper edge of the timber on these warm south
slopes, but not seen elsewhere in the region; in anthesis early in August. The
Brintnell Lake plants have nearly smooth sheaths, and lemmas almost smooth
except for a thin pubescence near the base and on the nerves. The lower parts
of the plants as well as the spikelets are quite glaucous, so that they have a visible
“bloom” on them. Hitchcock (Man. Gr., 936) reduces P. Buckleyana to P.
scabrella (Thurb.) Benth., but the nearly smooth sheaths and lemmas of our
plants seem to justify maintaining it. Poa Buckleyana has not been previously
noted in the Mackenzie basin north of Lake Athabaska.

Schizachne purpurascens (Torr.) Swallen in Jour. Wash. Acad. Sci. 18: 204, fig. 1
(1928); Hultén, Fl. Alaska & Yuk. Pt. 2: 193 (1942).

Trisetum purpurascens Torr. Fl. N. and Mid. U. S. 1: 127 (1823).

Grassy swale near Ft. Liard, Nowosad (Can.).

This species has been collected only once in the upper Mackenzie region,
though it is common in the Wood Buffalo Park (Raup in Nat. Mus. Can. Bull.
74: 105, 1935) and has been found at Great Bear L. (Porsild 3643).

Agropyron trachycaulum (Link) Malte var. typicum Fernald in Rhod. 35: 169 (1933);
Raup in Jour. Arn. Arb. 17: 202 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 262
(1942). See Fernald in Rhod. 35: 161-182 (1933) and 36: 417 (1934) for dis-
cussion of this and the following forms of A. trachycaulum.

Triticum trachycaulum Link, Hort. Bot. Berol. 2: 189 (1833).

Sand bars along Mackenzie R. west of Simpson, 9869; thickets along wood road near
Simpson, 9872; rocky-sandy bank of Mackenzie R. opposite Simpson, 9969; Norman, Rich-
ardson (G).

Occasional to common in the Simpson district; spikelets maturing late in
August.

It is notable that A. trachycaulum, so far as is known at present, is represented
in the Mackenzie R. district east of the mountains only by the typical form.

116 SARGENTIA [6

In the Brintnell L. region of the Mackenzie Mts. it is represented only by vars.
novae-angliae and unilaterale, as follows.

Agropyron trachycaulum (Link) Malte var. novae-angliae (Scribn.) Fernald in Rhod.
35: 174 (1933).

Agropyron novae-angliae Scribn. in Brainerd, Jones & Eggleston, Fl. Vermont, 9, 103
(1900).

Saline R., above Norman, Wynne-Edwards 8569 (Can.) ; along dry rocky gully on south-
facing slope of granite mountain just west of Brintnell L., alt. about 3000’, 9486; thickets on
north shore of Brintnell L. near camp, 9599; muddy bank of creek entering west end of
Brintnell L., 9697; warm, shaly south slope of Red Mt., in upper part of timber, alt. 4000’,
9779.

Occasional to common, mostly at or near lake-level; in anthesis about mid-
July. Variety novae-angliae has not thus far been reported north of the upper
Peace R. region in the Rocky Mts., and has not been noted at all in the Macken-
zie basin east of the mountains.

Agropyron trachycaulum (Link) Malte var. unilaterale (Cassidy) Malte in Nat. Mus.
Can, Ann. Rept. 1930: 46 (1932).

Agropyron unilaterale Cassidy in Colo. State Agric. Coll. Exp. Sta. Bull. 12: 63 (1890).

Agropyron subsecundum (Link) Hitche.; Hultén, Fl. Alaska & Yuk. Pt. 2: 261 (1942).

Muddy bank of creek entering west end of Brintnell L., 9699.

Common here, but not seen elsewhere in the Brintnell L. region; in anthesis
early in August. Probably common also in the upper Mackenzie valley, though
it has not yet been collected there. Frequent in the Wood Buffalo Park.

Agropyron latiglume (Scribn. & Sm.) Rydb. in Bull. Torr. Bot. Cl. 36: 539 (1909) ;
Hultén, Fl. Alaska & Yuk. Pt. 2: 258 (1942); Porsild in Nat. Mus. Can. Bull.
101: 15 (1945). See Malte in Nat. Mus. Can. Ann. Rept. 1930: 27-48 (1932), and
Fernald in Rhod. 35: 169 (1933),

Agropyron violaceum (Hornem.) Lange var. latiglume Scribn. & Sm. in U. S. Dept.
Agric. Div. Agrost. Bull. 4: 30 (1897).

Bank of Mackenzie R. at Simpson, 9/19; Mackenzie R. between Blackwater R. and
Norman, Crickmay 150 (Can.) ; local summit on hogback, north slope of Colonel Mt., alt.
4800’, 9610; thickets at margin of timber on gravel fan, south shore of Brintnell L., 9690,
9696; dry shale slope, south side of Red Mt., alt. 5000’, 9773; dry gravel plain near west
end of Brintnell L., 9801; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11189 (Can.).

Occasional to common ; found in anthesis at Brintnell L. in late July and early
August ; the Simpson specimen was just coming into flower about mid-June.

Our specimens show great variation in the amount of pubescence on the
lemmas (those in no, 9610 are nearly glabrous), and in the shape of the glumes.
In the Simpson material the latter have broad scarious margins and are abruptly
contracted at the apex, while those from Brintnell L. have narrower margins
and more attenuate apices. Some of the Brintnell L. plants, notably those from
the lower levels (nos. 9690, 9696, 9801), suggest forms of A. trachycaulum in
general aspect and in the form of the glumes. The writer is inclined to agree,
however, with the late M. O. Malte (1. c.) in considering A. latiglume as a cir-
cumpolar arctic-alpine species complex in its own right, distinguished by its
broad, flattish glumes which tend to be oblanceolate or obovate and conspicuously
scarious-margined.

Agropyron latiglume (Scribn. & Sm.) Rydb. var. pilosighume Hultén, Fl. Alaska &
Yuk. Pt. 2: 259 (1942),

Sand bar along Mackenzie R. west of Simpson, 9888.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 117

Occasional in this habitat; spikelets maturing in late August. The strongly
pubescent spikelets of this curious grass at once suggest A. dasystaciyon, but it
possesses none of the prominent creeping rhizomes of that species. In general
habit it looks like typical A. trachycaulum, with its spikes somewhat elongated,
scarcely imbricated, and the lower spikelets often distant. The closest relation-
ship, however, is clearly with A. latiglume, indicated by the pubescent lemmas
and the rather flat, oblong to obovate or oblanceolate, broadly scarious-margined
empty glumes which are abruptly contracted toward the apex. It differs from
A. latiglume in having pubescent glumes that are considerably shorter than the
lemmas and much shorter than the spikelets, and in having an elongated, scarcely
imbricated inflorescence. I have not seen any material cited by Hultén as var.
pilosiglume, but venture to place the Simpson plants under this name at least
tentatively.

Arctophila fulva (Trin.) Anders. Gram. Scand. 49 (1852); Hultén, Fl. Alaska & Yuk.
Pt. 2: 224 (1942); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).
Poa fulva Trin. in Mém. Acad. Sci. St. Pétersb. VI. Math. 1: 378 (1831).
Macmillan Pass, Canol Rd., mi. 225 E., Porsild (Can.).

Elymus canadensis L. Sp. Pl. 83 (1753); Raup in Jour. Arn. Arb. 17: 203 (1936).
Great Slave L., Richardson (G).

Elymus innovatus Beal, Grasses N. Amer. 2: 650 (1896); Raup in Jour. Arn. Arb. 17:
204 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 273 (1942); Porsild in Nat. Mus.
Can. Bull. 101: 16, 30 (1945). '

Elymus mollis R. Br. in Richardson, Bot. App. Frankl. Journ. 732 (1823) ; Hooker, Fl.
Bor.-Am. 2: 255 (1840); Macoun, Cat. Can. Pl. Pt. 4: 246 (1888); non Trin.

Dry ridges at mouth of South Nahanni R., Nowosad (Can.); Mackenzie R., between
Wrigley and Blackwater R., Crickmay 125 (Can.) ; Lone Mt., Wynne-Edwards 8427 (Can.) ;
Bolstead Cr., Canol Rd., mi. 111 E., alt. 4420’, Porsild 11828 (Can.); same place, Wynne-
Edwards 8243 (Can.) ; Sekwi R., Canol Rd., mi, 174 E., Porsild 11845 (Can.).

Noted in Fl. Bor.-Am. from “Saskatchewan to Bear Lake. . . 2”. Common in
the Wood Buffalo Park, and collected on the lower Mackenzie and at Great
Bear L. by Porsild (Sargentia 4: 13, 1943). Porsild states that it is very com-
mon along the Canol Rd.

Elymus arenarius L. var. villosus E. Mey. Pl. Labrad. 20 (1830). See St. John in
Rhod. 17: 98-103 (1915).
Elymus arenarius L.; Hooker, Fl. Bor.-Am, 2: 255 (1840), in part.
Elymus arenarius L. ssp. mollis (Trin.) Hultén in Vet. Akad. Handl. III. 5: 153 (1927),
Fl. Alaska & Yuk. Pt. 2: 268 (1942).

Southwest and north shores of Great Slave L., Howe 91989 (Can.) ; northwest shore of
Great Slave L., Bedford (Can.).

Common on the shores of larger lakes in the central part of the Mackenzie
basin, and noted in Fl. Bor.-Am. from “Saskatchewan to Bear Lake and the
extreme Arctic shores and islands.” There is a Richardson specimen from
Bear L. at Herb. G, but none from the upper Mackenzie have yet been seen.

Hordeum jubatum L. Sp. Pl. 85 (1753); Hooker, Fl. Bor.-Am. 2: 256 (1840); Macoun,
Cat. Can. Pl. Pt. 4: 243 (1888); Raup in Jour. Arn. Arb. 17: 204 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 2: 266 (1942).
Resolution, Harper 90056 (Can.) ; shore of Mackenzie R., 10 mi. below Mills L., Wynne-
Edwards 8544 (Can.) ; dry terrace in open field at Simpson, 9853; Saline R., above Norman,
W ynne-Edwards 8570 (Can.).

118 SARGENTIA [6

Our specimens are badly infected with a fungus which has greatly reduced
and distorted the inflorescences. Noted in Fl. Bor.-Am. from the “Saskatche-
wan to the Mackenzie River.”

Trisetum spicatum (L.) Richter, Pl. Eur. 1: 59 (1890); Hultén, Fl. Alaska & Yuk. Pt.
2: 184 (1942); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).

Aira spicata L. Sp. Pl. 64 (1753).
Trisetum subspicatum (L.) Beauv.; Hooker, Fl. Bor.-Am. 2: 244 (1840), in part.

Liard R., Nahanni Butte to Simpson, Crickmay 6 (Can.); Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8234 (Can.) ; noted by Porsild as common in Macmillan Pass, Canol
Rd., mi. 225 E,

Trisetum spicatum (L.) Richter var. molle (Michx.) Beal, Grasses N. Amer. 2: 377
(1896); Raup in Jour. Arn. Arb. 17: 204 (1936); Porsild in Nat. Mus. Can. Bull,
101: 30 (1945). See Rhod. 18: 195-198 (1916).

Avena mollis Michx. Fl. Bor.-Am. 1: 72 (1803).

Trisetum subspicatum (L.) Beauv. var. molle Gray; Macoun, Cat. Can. Pl. Pt. 4: 212
(1888).

North peak of Nahanni Mt., Wynne-Edwards 8426 (Can.).

Variety molle is the common form of the species in the central part af the
Mackenzie basin. Judging by specimens I have seen from Great Bear L.
(Richardson, G, N; J. M. Bell, Can.) it is common in that region also. The
record in Fl. Bor.-Am., “Rocky Mountains ... to the Arctic Sea-shore and
Islands,” probably refers in large part to this variety.

Trisetum spicatum (L.) Richter var. Maidenii (Gand.) Fernald in Rhod. 18: 196 (1916).

Trisetum subspicatum (L.) Beauv. f. Maidenii Gand. in Bull. Soc. Bot. France 49: 182
(1902).

Stony shore of Brintnell L., 9172; shale cliffs on north slope of Colonel Mt., alt. 4800’—
6000’, 9392, 9612; dry ridges on north slope of Terrace Mt., alt. 4000’-5000', 9675, 9683;
turfy ledges on summit of Red Mt., alt. 5900’, 9770; dry shale slope on south side of Red
Mt., alt. 5000’, 9774.

Occasional to common, chiefly on ledges above timber line, but reappearing
on the lake shore; in anthesis in the latter part of June, and mature in August.
Our plants are extremely variable in the color and form of the inflorescence, but
most of them fall into var. Maidenii which has bronze to purplish spikelets and
the inflorescence somewhat elongated and interrupted toward the base. The
greenish to straw-colored var. molle is not represented, although, as previously
noted, it is quite common in the central part of the Mackenzie basin. Three of
the numbers cited (9172, 9392, 9683) approach the typical variety in having
dense ovate inflorescences on some of the culms. Other culms in the same
colony may have them more elongate and interrupted.

Deschampsia cespitosa (L.) Beauv. Ess. Agrost. 91, 149, 160, pl. 18, f. 3 (1812); Hooker,
Fl. Bor.-Am, 2: 242 (1840), in part; Macoun, Cat. Can. Pl. Pt. 4: 209 (1888), in
part; Hultén, Fl. Alaska & Yuk. Pt. 2: 175 (1942).

Aira cespitosa L. Sp. Pl. 64 (1753).

Sand bar at mouth of small creek entering Mackenzie R. opposite Simpson, 9957; rocky-
sandy bank of Mackenzie R. opposite Simpson, 9962; dry ridges at mouth of South Nahanni
R., Nowosad (Can.).

Common in this habitat; spikelets mature the last week in August. Noted in
Fl. Bor.-Am. from “Saskatchewan to Bear Lake.” Our plants are relatively tall
and robust, apparently representing the species rather than var. glauca (see
Fernald in Rhod. 28: 154). The panicles are often 2.5-3 dm. long. Number

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 119

9957 is worthy of special note since it is strongly viviparous. This is carried
to such an extreme in some individuals that the whorls of the inflorescence are
composed of dense fascicles of proliferating spikelets which are strongly tinged
with purple. The occurrence of vivipary in North American D. cespitosa seems
to be rather unusual. The species is common throughout those parts of the
Mackenzie basin visited by the writer, but no viviparous forms have been ob-
served previously. Hitchcock (Man. Grasses of U. S., p. 290) states that the
species is “rarely with proliferous spikelets,” and in the large series of American
specimens at the Gray Herbarium only one such form could be found (Fernald
& Long 19947, Digby Co., Nova Scotia, as D. cespitosa var. glauca). Accord-
ing to Ostenfeld (Fl. Arctica, 113) the proliferous form of the species is D.
alpina (L.) R. & S. Hultén failed to find this in the Aleutians (Fl. Aleut. Isls.,
80-82), or in Alaska (Fl. Alaska & Yuk., 173-180). Porsild (Rhod. 41: 179)
does not mention it among his collections from Alaska. Simmons makes no
note of vivipary in Deschampsia (Aira) cespitosa either in his flora of Elles-
mereland or of the American Arctic Archipelago. Abbe, however, collected
plants along the northern Labrador coast which have been deternuned as D.
alpina (L.) R. & S. (Rhod. 38: 140). Polunin (Fl. E. Arct. Can. 55-60)
places all eastern Arctic viviparous forms of the cespitosa group in D. alpina, and
records it only from southern Baffin and northern Labrador. He regards it as
identical with plants from south Greenland and almost identical with those of
northern Europe and Spitzbergen. Gelting states (Meddel. om Grgnland, Bd.
101, Nr. 2: 188) that in East Greenland vivipary does not occur in the group.
On the other hand viviparous forms are rather common in northern Europe.
None of the eastern American or European D. alpina is tall and robust like our
Simpson specimens, nor does it show the tendency to dense fasciation in the
inflorescence,

Deschampsia brevifolia R. Br. Suppl. App. Parry’s 1*' Voy. 291 (mispaged 191) (1824);
Porsild in Nat. Mus. Can. Bull. 101: 15 (1945). See Fernald in Rhod. 36: 90
(1934).

Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11197 (Can.).

Vahlodea atropurpurea (Wahl.) Fries ssp. paramushirensis (Kudo) Hultén, F. Kamtch.
1: 111 (1927); Hultén, Fl. Alaska & Yuk. Pt. 2: 180 (1942); Porsild in Nat. Mus.
Can. Bull. 101: 15 (1945).
Deschampsia atropurpurea var. paramushirensis Kudo in Jour. Coll. Agric. Hokkaido
Univ. Sapporo 11: 71 (1922).
Macmillan Pass, Canol Rd., mi. 225 E., just east of the Yukon-Mackenzie divide, Porsild
11194 (Can.).

Calamagrostis purpurascens R. Br. apud Richardson, Bot. App. Franklin’s Journ. 731
(1823); Hooker, Fl. Bor.-Am. 2: 240 (1839); Raup in Jour. Arn. Arb. 17: 206
(1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 170 (1942); Porsild in Nat. Mus. Can.
Bull. 101: 15, 30 (1945). See Fernald in Rhod. 35: 213 (1933).

Gravelly thickets on shore of Brintnell L. near camp, 9308; alpine rock crevices on north
slope of Colonel Mt., 9420; shale cliffs in gorge near east end of Brintnell L., 9588; hogback
on north slope of Colonel Mt., alt. 4800’, 9611; wooded south-facing slope of Red Mt., alt.
3500'—4000’, 9667, 9673, 9777; Lone Mt., Wynne-Edwards 8425 (Can.) ; Sekwi R., Canol Rd.,
mi. 174 E., alt. 3625’, Porsild 11844 (Can.).

Common; in anthesis in late June or early July. This is one of the com-
monest grasses on rocky alpine slopes, though it is nowhere a primary species
in the plant associations. On dry, warm slopes and crevices at lower elevations
it grows tall (8-10 dm.), with paler purple in the spikelets than the typical form.

120 SARGENTIA [6

Noted in Fl. Bor.-Am. as occurring on the “Rocky Mountains and Arctic sea-
shore.” Frequent throughout most of the Athabaska—Great Slave L. region
(Raup, 1935, 1936).

Calamagrostis lapponica (Wahl.) Hartm. Handb. Skand. Fl. ed. 1: 46 (1820); Hultén,
Fl. Alaska & Yuk. Pt. 2: 166 (1942).

Arundo lapponica Wahl. Fl. Lapp. 27 (1812).

Turfy ledge on summit of Red Mt., Brintnell L. region, alt. 5900’, 9790.

Common in this situation but not seen elsewhere. Found in early anthesis
on Aug. 7.

The existence of typical Calamagrostis lapponica on the American continent
has long been questioned. Scribner & Merrill (Contr. U. S. Nat. Herb. 13:
61, 1910) considered specimens seen by them from Dall R., Alaska (O. A. Piper)
to represent the typical form, Hultén (FI. Alaska & Yuk. Pt. 2: 166-167, 1942)
apparently supports this view, and cites several other Yukon and Alaska collec-
tions which he also regards as typical. Stebbins (Rhod. 32: 56, 1930) did not
recognize any typical material in America, but described a var. brevipilis from
the Straits of Belle Isle, in Quebec. Britton & Rydberg (Bull. N. Y. Bot.
Gard. 2: 154, 1901) considered specimens they had from the upper Yukon to
be typical. Porsild (Sargentia 4: 10-12, 1943) has described a var. nearctica
from the Mackenzie delta region which he distinguishes from the typical species
by its smaller stature and by its shorter and narrower panicles and glumes. He
regards this as probably the principal representative of C. lapponica in America,
with a distribution from western Alaska to the northwest coast of Hudson Bay,
and southward into the Canadian Rockies. He considers that many American
specimens determined as C. lapponica are referable to C. neglecta or C. hyper-
borea.

It is impossible for me to judge the relative merits of these views regarding
all the northwestern material, but our material from the Brintnell Lake area
matches very well the rather large series of Eurasian specimens in the Gray
Herbarium. The awn is attached somewhat higher on the lemma (% to %
way up from the base) than in some of these specimens, but the character is
sufficiently variable to make it of little diagnostic value. Stebbins’ var. brevipilis
(considered by Porsild a form of C. neglecta) is distinguished by its short
callus hairs (7 the length of the lemma) ; but in our plants the callus hairs are
fully as long as the lemma, closely resembling those of the typical form. Var.
brevipilis is also noted as having “innovations numerous,” but here also our
specimens match the Eurasian.

Porsild’s var. nearctica is described as having lax panicles which are narrow
to somewhat open. Our specimens have rather dense, upright panicles, the
whole aspect of the plants resembling that of C. purpurascens. Otherwise they
match Porsild’s description rather well. They have anthers 2 mm. long, lemmas
3.5-4 mm., paleas 4 the length of the lemmas, glumes 4-5 mm., awns about as
long as the lemmas, not twisted and scarcely bent.

As Stebbins has pointed out (1. c.), C. lapponica seems to combine some of
the principal characters which are commonly used to define the Sections Calama-
gris and Deyeuxia. It has the long callus hairs of the former and the thick
lemma of the latter. Its short palea (14-%4 as long as the lemma) places it in
the subsection Orthoatherae of Deyeuxia, while its bent and slightly geniculate
awn suggests the subsection Ancylatherae.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 121

Calamagrostis canadensis (Michx.) Nutt. Gen. 1: 46 (1818); Hooker, Fl. Bor.-Am. 2:
240 (1839), in part; Raup in Jour. Arn. Arb. 17: 206 (1936). See Stebbins in
Rhod. 32: 42 (1930).

Arundo canadensis Michx. Fl. Bor.-Am. 1: 73 (1803).

Deyeuxia canadensis (Michx.) Hooker; Macoun, Cat. Can. Pl. Pt. 4: 204 (1888).

Along lower Slave R., near Pt. Brulé, Raup 231, 232; mouth of Rocher R., Great Slave
L., Harper 90055 (Can.).

The typical form, with very short spikelets, is common in the Athabaska and
Slave R. lowlands, but appears to reach its northern limits at Great Slave L. or
on the upper Mackenzie.

Calamagrostis canadensis (Michx.) Nutt. var. robusta Vasey in Wheeler, Rept. Wess:
Surv. 100 Merid. 6: 285 (1878); Raup in Jour. Arn. Arb. 17: 207 (1936). See
Stebbins in Rhod. 32: 42 (1930).

Calamagrostis canadensis (Michx.) Nutt. ssp. Langsdorfi (Link) Hultén, Fl. Alaska &
Yuk. Pt. 2: 161 (1942), in part.

Northwest shore of Great Slave L., Bedford (Can.) ; east shore of Trout L., C. O. Hage
(G) ; grassy swale near Ft. Liard, Nowosad (Can.) ; gravelly thickets along shore of Brint-
nell L. at Camp Pt., 9466; mossy thickets on north shore of Brintnell L. near camp, 9600.

Common in the open association of herbs that borders the willow and alder
thickets at the shore of Brintnell L.; in anthesis about mid-July. Probably
common throughout southwestern Mackenzie, though the only specimen I have
seen that may have come from the upper Mackenzie valley is without locality
data (Onion, Kennicott & Hardisty, N). Collected at Good Hope by Miss E.
Taylor (Can.). This is the commonest form of the species in the Athabaska-
Great Slave L. region. Hultén, in Fl. Alaska & Yuk., reduces var. robusta to
his ssp. Langsdorfi, which hardly seems justifiable in view of the geographic
position of the former in the interior of the continent.

Calamagrostis canadensis (Michx.) Nutt. var. Langsdorfi (Link) Inman in Rhod. 24:
143 (1922); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945). See Stebbins in
Rhod. 32: 43 (1930).

Arundo Langsdorfi Link, Enum. Pl. Hort. Berol. 1: 74 (1821).

Calamagrostis canadensis (Michx.) Nutt. ssp. Langsdorfi (Link) Hultén, Fl. Alaska &
Yuk. Pt. 2: 161 (1942), in part.

Mackenzie R., between Blackwater R. and Norman, Crickmay 151 (Can.) ; damp mud
flat near mouth of creek at west end of Brintnell L., 9688; Macmillan Pass, Canol Rd., mi.
225 E., Porsild 11201 (Can.).

Common to abundant at the west end of Brintnell L. on the higher parts of
the newly formed mud-flats, near the first willows; a primary species, associated
in some places with Arctagrostis arundinacea; in anthesis early in August.
Hitchcock, in Am. Jour. Bot. 21: 135 (1934), identified most if not all Ameri-
can plants long regarded as var. Langsdorfi with var. scabra (Pres!) Hitche.,
and did not admit var. Langsdorfi to the American flora. I am inclined to agree
with Hultén (1. c.), however, that this large-flowered form of C. canadensis is
circumpolar.

Calamagrostis inexpansa Gray var. brevior (Vasey) Stebbins in Rhod. 32: 50 (1930);
Raup in Jour. Arn. Arb. 17: 207 (1936).
Calamagrostis stricta var. brevior Vasey in Wheeler Rep. 6: 285 (1878).
Calamagrostis inexpansa Gray; Porsild in Nat. Mus. Can. Bull. 101: 33, 34 (1945);
Hultén, Fl. Alaska & Yuk. Pt. 2: 164 (1942), in part.
Rocky-sandy bank of Mackenzie R., opposite Simpson, 9961; outwash gravel plain along
the Mackenzie near the mouth of the N. Nahanni R., Wynne-Edwards 8535 (Can.) ; Saline

122 SARGENTIA [6

R., above Norman, Wynne-Edwards 8568 (Can.) ; “Cumberland House to Bear L.,” Rich-
ardson (G),

Common at the Simpson locality ; witlr maturing spikelets in late August.

Calamagrostis neglecta (Ehrh.) Gaertn., Meyer & Scherb. Fl. Wetterau 94 (1799);
Hultén, Fl. Alaska & Yuk. Pt. 2: 167 (1942). See Stebbins in Rhod. 32: 53
(1930).

Arundo neglecta Ehrh. Beitr. 6: 84, 137 (1791).

Calamagrostis coarctata (Torr.) Hooker, Fl. Bor.-Am. 2: 240 (1839), at least in part.

Upland prairie west of Mackenzie R. near Simpson, 9905; wet meadow at west end of
Brintnell L., 9715.

Abundant at the Brintnell L. station, where it is a primary species along the
lake shore near muskeg thickets; in anthesis the first week in August. In the
prairie near Simpson it is a primary species in the drier parts, and was maturing
its spikelets late in August. The record for C. coarctata, “woody country,” in
Fl. Bor.-Am. probably refers to this species.

Agrostis scabra Willd. Sp. Pl. 1: 370 (1798); Raup in Jour. Arn. Arb. 17: 208 (1936);
- Hultén, Fl. Alaska & Yuk. Pt. 2: 155 (1942). See Fernald in Rhod, 35: 207-212
(1933).

Wet gravelly shore of Brintnell L. at Camp Pt., 9468; along dry rocky gullies and on
fans on southward-facing slope of granite mountain W. of Brintnell L., 9484; dry shale
slopes on south side of Terrace Mt., alt. 3500’-5000', 9679; damp mud holes and mossy
muskeg thickets at W. end of Brintnell L., 9714; “Cumberland House to Bear Lake,”
Richardson (G).

Occasional to common in the Brintnell LL. habitats noted, but not found on
cold north-facing slopes. In anthesis about mid-July. Two of the numbers
(those from lake-level, 9468 and 97 14) have some of the lemmas awned and
probably represent f. Tuckermani Fern. The awns are attached above the mid-
dle of the lemmas and scarcely extend beyond the tips of the glumes.

Cinna latifolia (Trev.) Griseb. in Ledeb. Fl. Ross. 4: 435 (1853); Hultén, Fl. Alaska &
Yuk. Pt. 2: 148 (1942).

Agrostis latifolia Trey. in Goepp. Beschr. Bot. Gard. Breslau, 82 (1832).

Sand bars along Mackenzie R., just west of Simpson, 9887.

Occasional ; found with maturing spikelets in the last week of August. Cinna
latifolia has not been reported previously in the Mackenzie basin north of Lake
Athabaska (Raup, 1936, p. 208). In its more southern range it is a charac-
teristic species of rich woods and thickets, but at Simpson it was seen only on
river sand bars.

Arctagrostis latifolia (R. Br.) Griseb. in Ledeb. Fl. Ross. 4: 434 (1853); Porsild in Nat.
Mus. Can. Bull. 101: 15 (1945).

Colpodium latifolium R. Br. Verm. Schr. 1: 414 (1825).

Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8235 (Can.).

Noted by Porsild as common also in wet places on the alpine “Plains of
Abraham,” along the Canol Rd. near mi. 81 E.

Arctagrostis arundinacea (Trin.) Beal, Grasses N. Amer. 2: 317 (1896); Raup in Jour.
Arn. Arb. 17: 208 (1936); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945).
Vilfa arundinacea Trin. De gram. unifl. et sesquifl. 127 (1829).
Colpodium arundinaceum (Trin.) Hooker, Fl. Bor.-Am. 2: 238 (1839).
Arctagrostis latifolia (R. Br.) Griseb. var. arundinacea (Trin.) Griseb. in Ledeb. F1.
Ross. 4: 435 (1853) ; Hultén, Fl. Aleut. Isls. 71 (1937), Fl. Alaska & Yuk. Pt. 2: 145
(1942).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 123

Along wood road on bluff above Mackenzie R. west of Simpson, 9907; gravelly willow-
alder thickets on shore of Brintnell L., 9464; mossy shore of an old stream channel near west
end of Brintnell L., 9496; along small brook on west slope of Terrace Mt., alt. 4800’, 9533;
muskeg thicket on north shore of Brintnell L., 9567, 9568; damp stony beach at mouth of
Frost Creek, 9672; wet meadow on mud flat near west end of Brintnell L., 9698, 9701;
gravelly lake shore at Camp Pt., 9800; open wood of black spruce on north slope of Colonel
Mt., alt. about 3500’, 9819; “Cumberland House to Bear Lake,” Richardson (G).

Common to abundant on the shores of Brintnell L., and a primary species on
some parts of the recently formed alluvial deposits at the western end; at higher
elevations it is only occasional. In anthesis about mid-July. Noted by Porsild
as common in Macmillan Pass along the Canol Rd.

Some students have considered the genus Arctagrostis to be monotypic (Grise-
bach in Ledebour’s Flora Rossica; Hultén in Fl. Aleut. Isls.; Holm in Bot. Can.
Arct. Exp. Pt. B. p. 4), with A. latifolia as the typical species. Others, such
as Scribner & Merrill in Grasses of Alaska, 55, and Porsild in Rhod. 41: 178,
have maintained A. arundinacea as a distinct species. More recently Hultén,
in Fl. Alaska & Yuk. Pt. 2: 148, recognized a second species, A. poaeoides Nash,
in Alaska, but still maintained A. arundinacea as a variety of A. latifolia. There
are at the Gray Herbarium numerous well prepared specimens of A. latifolia
from most parts of the American Arctic. They are all relatively short plants
with dense inflorescences in which there is very little tendency to the elongation
of branches or main axis. The spikelets are usually 4-5 mm. long and the
anthers 2-2.7 mm. long (usually 2.1-2.5 mm.). Arctagrostis arundinacea in
the Brintnell L. region, on the other hand, commonly grows to a height of 1.2
m., with inflorescences freely branched and as much as 3 dm. long. Some of
the lower branches are naked for more than half their length. The spikelets are
shorter than those of A. latifolia (3.54 mm.), and the anthers vary from 1.2
to 2.1 mm. long (usually 1.4-1.8mm.). Another character is that of the “collar”
which subtends the lowest whorl of the inflorescence. In A. arundinacea it is
much more obliquely cut than in A. Jatifolia. With these differences in mind it
seems best to maintain the two as separate species.

As it first comes into flower the inflorescence of A. arundinacea has a “monili-
form” appearance due to the elongation of the main axis without accompanying
development of the branches. The latter soon grow out, however, and while the
plant is in anthesis the whole inflorescence is expanded so that the rather stiff
branches come off almost at right angles. After anthesis the branches are con-
tracted toward the main axis, the whole inflorescence becoming strict and re-
maining so until maturity.

Alopecurus alpinus Sm. FI. Brit. 3: 1386 (1804); Hultén, Fl. Alaska & Yuk. Pt. 2: 140
(1942); Porsild in Nat. Mus. Can. Bull. 101: 15 (1945). See Rhod. 27: 196 (1925).

Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11191 (Can.).
Noted by Porsild as rare or occasional in Macmillan Pass.

Alopecurus aequalis Sobol. Fl. Petrop. 16 (1799); Raup in Jour. Arn. Arb. 17: 208
(1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 139 (1942).

Alopecurus aristulatus Michx.; Hooker, Fl. Bor.-Am, 2: 233 (1839).
Alopecurus geniculatus L. var. aristulatus Munro; Macoun, Cat. Can. Pl. Pt. 4: 188
(1888).
Damp slough in lowland woods near Simpson, 9841; grassy swale near Ft. Liard, Nowo-
sad (Can.).
Common; in fruit late in August. Noted in Fl. Bor.-Am. from “Canada to
Bear Lake.”

124 SARGENTIA [6

Phleum alpinum L. Sp. Pl. 59 (1753); Porsild in Nat. Mus. Can. Bull. 101: 14 (1945)
Phleum alpinum L. var. americanum Fourn., sensu Hultén, Fl. Alaska & Yuk. Pt. 2: 136
(1942).
Noted by Porsild at Macmillan Pass along the Canol Rd., mi. 225 E., alt.
about 5000’.
Hultén, |. c., considers the northwestern American plants distinct from those
of northern Scandinavia, but admits that they cannot always be told apart.

Muhlenbergia Richardsonis (Trin.) Rydb. in Bull. Torr. Bot. Cl. 32: 600 (1905).

Vilfa Richardsonis Trin. in Mém. Acad. St. Pétersb. VI. Sci. Nat. 41: 103 (1840).

This species is abundant on the Salt Plains west of the upper Slave R. (Raup,
Jour. Arn, Arb. 17: 209, 1936), and has been collected by Porsild along Bear
R. (Sargentia 4: 9, 1943). It has not yet been seen, however, along the upper
Mackenzie.

Beckmannia Syzigachne (Steud.) Fernald in Rhod. 30: 27 (1928); Raup in Jour. Arn.
Arb. 17: 209 (1936); Porsild in Sargentia 4: 9 (1943); Hultén, Fl. Alaska & Yuk.
Pt. 2: 188 (1942).

Beckmannia erucaeformis sensu Richardson, Bot. App. Franklin’s Journ. 731 (1823), non
R. & S,

Damp lowland sloughs and sand bars along the Mackenzie R. near Simpson, 9873; Simp-
son, Miss E. Taylor 90 (N, Can.) ; grassy swale near Ft. Liard, Nowosad (Can.).

Common on river flood plains northward to the Mackenzie delta; spikelets
mature at Simpson during the last week in August.

Spartina gracilis Trin. in Mém. Acad. St. Pétersb. VI. Sci. Nat. 41: 110 (1840); Raup
in Jour. Arn. Arb. 17: 209 (1936).

Junction of the Little Buffalo and Nyarling Rivers, Russell 62 (G).

The range of Spartina polystachya, which may have included both S. gracilis
and S. pectinata, is given in Fl. Bor.-Am. 2: 242 (1840) as “Saskatchewan to
Slave Lake,” but there is a specimen of S. gracilis in Herb. G marked “Cumber-
land House to Bear Lake.” This specimen was originally marked S. poly-
stachya.

Spartina pectinata Link in Jahrb. Gewachsk 13: 92 (1820); Raup in Jour. Arn. Arb.
17: 210 (1936).

Spartina polystachya sensu Hooker, Fl. Bor.-Am. 2: 242 (1840), in part, non Willd.

Resolution, Onion, Kennicott & Hardisty (N).

See note under S. gracilis.

Hierochloé alpina (Sw.) R. & S. Syst. Veg. 2: 514 (1817); Hultén, Fl. Alaska & Yuk.
Pt. 2: 132 (1942); Porsild in Nat. Mus. Can. Bull. 101: 14 (1945).

Holcus alpinus Sw. in Willd. Sp. Pl. 4, 2: 937 (1805).

Alpine meadows on south slope of Red Mt., alt. 4600’, 9232; along small brook on east
slope of Red Mt., alt. 4500’, 9267; shale rock slide on west slope of Terrace Mt., alt. 5500’,
9536; summit and higher eastern slopes of Red Mt., alt. 5800’-5900', 9791; Bolstead Cr.,
Canol Rd., mi. 111 E., Wynne-Edwards 8233 (Can.).

Common to abundant in high alpine meadows on the north side of Brintnell
L., but not seen on the N. slope of Colonel Mt.; in anthesis the last week of June.

Hierochloé odorata (L.) Wahl. Fl. Upsal. 32 (1820); Raup in Jour. Arn. Arb. 17: 210
(1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 133 (1942).

Holcus odoratus L. Sp. Pl. 1048 (1753).

Resolution, Kennicott (N); old fields, clearings and. roadsides at Simpson, 9135; in
meadows, Bell R., Mackenzie R., McConnell 29487 (Can.).

Abundant at Simpson; in anthesis about mid-June.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 1WA)

Phalaris arundinacea L. Sp. Pl. 55 (1753); Hooker, Fl. Bor.-Am. 2: 234 (1839); Macoun,
Cat. Can. Pl. Pt. 4: 185 (1888); Hultén, Fl. Alaska & Yuk. Pt. 2: 131 (1942).

Damp places in upland prairie west of the Mackenzie R. near Simpson, 9908; grassy swale
near Ft. Liard, Nowosad (Can.).

Common in this habitat ; spikelets maturing in the last week of August. Noted
in Fl. Bor.-Am. from “. . . Canada... to the Bear Lake.”

CYPERACEAE

Eriophorum Scheuchzeri Hoppe, Bot. Taschenb. 104, app. tab. 7 (1800); Raup in Jour.
Arn. Arb. 17: 210 (1937); Hultén, Fl. Alaska & Yuk. Pt. 2: 284 (1942); Porsild
in Nat. Mus. Can. Bull. 101: 16 (1945).

Eriophorum capitatum sensu Hooker, Fl. Bor.-Am, 2: 231 (1839), in part; Macoun, Cat.
Can. Pl. Pt. 4: 104 (1888), in part; not Host.

Simpson, coll. ?, May 19, 1860 (N) ; mud flat at west end of Brintnell L., 9265.

Abundant at Brintnell L.; in flower the last week of June. A primary species
on the most recently formed deposits of glacial mud, where it is associated with
Juncus castaneus. At least a part of the material cited in Fl. Bor.-Am. under
E. capitatum, “From Quebec ... to the Arctic coasts and islands,” belongs
here. Noted by Porsild at Macmillan Pass, on the Canol Road.

Eriophorum vaginatum L. Sp. Pl. 52 (1753); Raup in Jour. Arn. Arb. 17: 211 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 2: 286 (1942); Porsild in Sargentia 4: 14 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8245 (Can.).

Eriophorum Chamissonis C. A. Mey. forma albidum (F. Nyl.) Fernald in Rhod, 23:
131 (1921); Raup in Jour. Arn. Arb. 17: 210 (1936).

Eriophorum russeolum Fries var. albidum F. Nyl. in Acta Soc. Sc. Fenn. 3: (1852) and
in Anders. Bot. Notiser 1857: 58 (1857).

Eriophorum russeolum Fries var. leucothrix (Blomgr.) Hultén, Fl. Alaska & Yuk. Pt.
2: 283 (1942).
Resolution, Kennicott (N).
Occasional in the Athabaska—Great Slave L. region, and probably also through-
out the Mackenzie country.

Eriophorum brachyantherum Trautv. Fl. Ochot. Phaenog. in Middend., Reise i. d.
Aussersten N. v. O. Sibieriens 1, 2: 58 (1856); Hultén, Fl. Alaska & Yuk. Pt. 2:
279 (1942).

Eriophorum opacum (Bjornstr.) Fernald; Raup in Jour. Arn. Arb. 17: 210 (1936) ;
Porsild in Sargentia 4: 14 (1943) ; Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

Liard R., Nahanni Butte to Simpson, Crickmay 8 (Can.); muskeg thickets and mossy
banks around the western end of Brintnell L., 9245.

Abundant in the Brintnell L. area; in fruit late in June. Noted by Porsild on
a small tributary of the Little Keele R., along the Canol Rd.

Eriophorum spissum Fernald in Rhod. 27: 208 (1925); Raup in Jour. Arn Arb. 17:
211 (1936); Porsild in Sargentia 4: 14 (1943).
Eriophorum vaginatum L. ssp. spissum (Fernald) Hultén, Fl. Alaska & Yuk. Pt. 2: 286
(1942). ;
This species has not yet been collected in the upper Mackenzie valley, though
it is to be expected there. It is common in the Wood Buffalo Park, and has
been collected on the lower Mackenzie (Stringer, Can.).

Eriophorum angustifolium Roth, Tent. 1: 24 (1788); Hooker, Fl. Bor.-Am. 2: 231
(1839); Raup in Jour. Arn. Arb. 17: 211 (1936); Hultén, Fl. Alaska & Yuk. Pt.
2: 276 (1942); Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

*

126 SARGENTIA [6

Liard R., Nahanni Butte to Simpson, Crickmay 7 (Can.); Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8244 (Can.); Macmillan Pass, Canol Rd., mi. 225 E., Porsild
11217 (Can.).

Noted in Fl. Bor.-Am. as occurring from “Canada to Arctic America.” Por-
sild reports it along the Canol Road, and also at Little Keele R. and on the
“Plains of Abraham.”

Eriophorum gracile Koch in Roth, Catalect. 2: add. 259 (1800); Hooker, Fl. Bor.-Am.
2: 232 (1839); Macoun, Cat. Can. Pl. Pt. 4: 106 (1888); Hultén, Fl. Alaska &
Yuk. Pt. 2: 280 (1942).

Noted in Fl. Bor.-Am. as occurring from “Saskatchewan to Arctic America.”

Otherwise we have no Mackenzie basin records for the species except at Lesser

Slave Lake (Brinkman 4542, N).

Scirpus caespitosus L. var. callosus Bigel. Fl. Bost. ed. 2, 21 (1824). See Rhod. 23:
22 (1921). ,
Eleocharis caespitosa (L.) Link; Hooker, Fl. Bor.-Am. 2: 229 (1839), in part.
Noted in FI. Bor.-Am. as occurring “Throughout Canada to near the shores of
the Arctic Sea. . . ,” but I have seen no material from north of Great Slave L.

Scirpus validus Vahl. Enumer. Pl. 2: 268 (1806); Raup in Jour. Arn. Arb. 17: 212
(1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 295 (1942); Porsild in Sargentia 4:
14 (1943),

North of the height of land between Great Slave and Great Bear Lakes, Bedford (Can.) ;
Mackenzie R. near Norman, Porsild 5367 (Can.).

Scirpus microcarpus Pres!, Rel. Haenk. 1: 195 (1828); Raup in Jour. Arn. Arb. 17:
212 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 292 (1942).

Resolution, Kennicott (N).

Eleocharis pauciflora (Lightf.) Link, Hort. Berol. 1: 284 (1827); Porsild in Sargentia
4: 14 (1943).
Scirpus pauciflorus Lightf. Fl. Scot. 2: 1078 (1777) ; Hultén, Fl. Alaska & Yuk. Pt. 2:
294 (1942).
Known in the Athabaska—Great Slave L. district (Raup 6833, 374), and at
Great Bear L. (Porsild 1. c.), but not yet collected along the upper Mackenzie.

Eleocharis palustris (L.) R. & S. Syst. 2: 151 (1817); Hooker, Fl. Bor.-Am, 2: 228
(1839); Raup in Jour. Arn. Arb. 17: 212 (1936); Porsild in Sargentia 4: 14 (1943).

Scirpus palustris L. Sp. Pl. 47 (1753) ; Hultén, Fl. Alaska & Yuk. Pt. 2: 204 (1942).

Sand bars along Mackenzie R. near Simpson, 9903; sandy bank of Mackenzie R. opposite
Simpson, 9966; grassy swale near Ft. Liard, Nowosad (Can.).

Occasional to common locally; in fruit the last week in August. Noted in
Fl. Bor.-Am., “Throughout Canada to Bear Lake,” and collected by Porsild at
Bear Lake (4668, Can.) and along the Bear R. (3376, Can.).

The Simpson specimens are of low stature (1.5-2.5 dm.) and grow from
rather stout, long, horizontal rootstocks. The bristles on the achenes are a little
longer than usual, but otherwise the specimens can be placed in var. typica Rouy.

Eleocharis acicularis (L.) R. & S. Syst. 2: 154 (1817); Porsild in Sargentia 4: 14 (1943).
Scirpus acicularis L. Sp. Pl. 48 (1753) ; Hultén, Fl. Alaska & Yuk. Pt. 2: 289 (1942).
Bank of Mackenzie R. opposite Simpson, Crickmay 40 (Can.).

Common in the Wood Buffalo Park (Raup in Jour. Arn. Arb. 17: 213) and
collected frequently by Porsild (1. c.) at Great Bear L. and on the lower

Mackenzie.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 127

Kobresia simpliciuscula (Wahl.) Mackenzie in Bull. Torr. Bot. Cl. 50: 349 (1923);
Hultén, Fl. Alaska & Yuk. Pt. 2: 297 (1942); Porsild in Nat. Mus. Can. Bull.
101: 16 (1945).
Carex simfliciuscula Wahl. in Sv. Vet. Akad. Handl. (Stockh.) 24: 141 (1801).

Plains of Abraham, Canol Rd., mi. 82 E., on bald, wind-swept summit, alt. 6000’, Porsild
11788 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8246 (Can.) ; Sekwi R.,
Canol Rd., mi. 174 E., Porsild 11846 (Can.).

Porsild states that this species is common along the Canol Road.

Carex nardina Fries var. Hepburnii (Boott) Kiikenth. in Pflanzenr. 38 (IV. 20): 70
(1909); Hultén, Fl. Alaska & Yuk. Pt. 2: 299 (1942), at least in part; Porsild in
Sargentia 4: 15 (1943), in Nat. Mus. Can. Bull. 101: 16 (1945).

Carex Hepburnii Boott in Hooker, Fl. Bor.-Am. 2: 209, tab. 207 (1839); Mackenzie in
N. Am. FI. 18: 22 (1931), tab. 2 (1940).

Rock slides, crevices and turf on north slope of Colonel Mt., alt. 5000’—5500', 9394, 9604,
9605; Plains of Abraham, Canol Rd., mi. 82 E., alt. 6000’, Porsild 11789 (Can.) ; Bolstead
Cr., Canol Rd., mi. 111 E., Porstld 11829 (Can.).

Occasiona) to common on the north slope of Colonel Mt., but not seen else-
where in the Brintnell L. region; mature in August.

Carex capitata L. Syst. Nat. ed. 10, 1261 (1759); Mackenzie in N. Am. FI. 18: 22
(1931); Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

Sekwi R., Canol Rd., mi. 174 E., Porsild 11854 (Can.).

Carex pyrenaica Wahl. in Vet. Akad. Handl. (Stockh.) 24: 139 (1803); Hultén, FI.
Alaska & Yuk. Pt. 2: 308 (1942), in part; Mackenzie in N. Am. FI. 18: 26 (1931);
Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

Damp stony beach at Camp Pt., Brintnell L., 9165; along rocky stream in upper valley
of Frost Cr., north of Brintnell L. camp, alt. 4300’—4500’, 9298, 9535; Fort Norman, Richard-
son (G, N); Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11203 (Can.).

Common in the alpine site at Brintnell L., but rare at lake level; mature about
mid-July.

Carex incurva Lightf. Fl. Scot. 544, pl. 24, f. 1 (1777); Richardson, Arct. Search. Exp.
2: 345 (1851); Mackenzie in N. Am. FI. 18: 31 (1931).

This species was noted by Richardson (1. c.) as occurring in the “Valleys of
the Saskatchewan and Mackenzie,’ and there aré Richardson specimens from
Great Bear L. in Herb. G. Mackenzie stated that he had seen material from
“Mackenzie,” presumably the Richardson specimens just mentioned. Otherwise
we have no records for it in the Mackenzie basin.

Carex siccata Dewey in Am. Jour. Sci. 10: 278, pl. F, fe 18 (1826); Macksiie in N.
Am. FI. 18: 39 (1931).

Dry upland woods west of Mackenzie R. near Simpson, 9901.
Common; achenes mature in late August.

Carex chordorrhiza Ehrh. in L. f. Suppl. Pl. Syst. Veg. 414 (1781); Mackenzie in N.
Am. FI. 18: 41 (1931); Hultén, Fl. Alaska & Yuk. Pt. 2: 315 (1942); Porsild in
Sargentia 4: 17 (1943).

This species has not yet been collected in the upper Mackenzie valley, though
it has been found at L. Athabaska (Raup in Jour. Arn. Arb. 17: 213), and at

Great Bear L. and in the lower Mackenzie country (Porsild 1. c.).

Carex diandra Schrank, Cent. Bot. Anmerk. 49 (1781); Mackenzie in N. Am. FI. 18:

72 (1931); Hultén, Fl. Alaska & Yuk. Pt. 2: 316 (1942); Porsild in Sargentia 4:
17 (1943).

Boggy swamp at west end of Brintnell L., 9453.

128 SARGENTIA [6

Common, forming dense tussocks near the edge of the lake; fruit maturing
about mid-July. Common throughout the Athabaska—Great Slave L. region
(Raup in Jour, Arn. Arb. 17: 213), and collected by Porsild (1. c.) at Great
Bear L.

Carex disperma Dewey in Am. Jour. Sci. 8: 266 (1824); Hooker, Fl. Bor.-Am. 2: 211
(1839); Mackenzie in N. Am. Fl. 18: 85 (1931); Hultén, Fl. Alaska & Yuk. Pt.
2: 327 (1942); Porsild in Sargentia 4: 17 (1943).

Margin of swamp in woods near Simpson, 9112.

Occasional in the Simpson area; with flowers and young fruit in mid-June.
Noted in Fl. Bor.-Am. as “Common in British America.” Common in muskeg
thickets in the Athabaska—Great Slave L. region (Raup in Jour. Arn. Arb. 17:
214), and collected by Porsild (1. c.) at Aklavik and Great Bear L.

Carex tenuiflora Wahl. in Vet. Akad. Handl. (Stockh.) 24: 147 (1803); Mackenzie in
N. Am. Fl. 18: 86 (1931); Raup in Jour. Arn. Arb. 17: 214 (1936); Hultén, FI.
Alaska & Yuk. Pt. 2: 328 (1942); Porsild in Sargentia 4: 18 (1943),

Occasional in the Athabaska—Great Slave L. region, on the lower Mackenzie,
and at Great Bear L., but not yet collected along the upper Mackenzie.

Carex loliacea L. Sp. Pl. 974 (1753); Mackenzie in N. Am. Fl. 18: 87 (1931); Hultén,
Fl. Alaska & Yuk. Pt. 2: 328 (1942).

Wet muskeg thickets in the valley west of Brintnell L., 9249, 9487; damp stony shore of
Brintnell L. at Camp Pt., 9194.

Occasional to common at or near lake-level, but not seen at higher elevations ;
mature about mid-July.

Carex bipartita All. Fl. Ped. 2: 265, pl. 89, f. 5 (1785); Mackenzie in N. Am. FI. 18: 88
(1931). See Bull. Torr. Bot. Cl. 50: 348-349 (1923).

Carex Lachenalii Schk.; Hultén, Fl. Alaska & Yuk. Pt. 2: 320 (1942); Porsild in Nat.
Mus. Can. Bull. 101: 16 (1945).

Dry shale slope on south side of Red Mt., alt. 5000’, 9746; along small stream, east slope
of Red Mt., alt. 4500’, 9296.

Occasional in the Brintnell L. area; found in anthesis in late June and early
August. Noted by Porsild as common in Macmillan Pass along the Canol Rd.,
mi. 225 E.

Carex bonanzensis Britton in Bull. N. Y. Bot. Gard. 2: 160 (1901); Porsild in Sargentia
4: 18 (1943).

Wet woods near Simpson, 9097.

Carex bonanzensis was described from the upper Yukon region, and has been
found in the Mackenzie region at only two places, Simpson and in the Mackenzie
delta (Porsild 6879, Can.). It is closely related to C. canescens, but differs
from the latter in having brown scales with prominent green mid-ribs and in
producing somewhat smaller perigynia.

Carex canescens L. Sp. Pl. 974 (1753); Mackenzie in N. Am. Fl. 18: 95 (1931); Hultén,
Fl. Alaska & Yuk. Pt. 2: 324 (1942); Porsild in Nat. Mus. Can. Bull. 101: 16
(1945).

Carex curta sensu Boott in Hooker, Fl. Bor.-Am. 2: 214 (1839), in part, non Good.

Damp stony beach at Camp Pt., Brintnell L., 9196; Macmillan Pass, Canol Rd., mi. 225 E.,
Porsild 11216 (Can.).

Occasional at Brintnell L.; in anthesis late in June. The note in FI. Bor.-
Am., “Common in British America,” probably refers in part also to the following.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 129

Carex canescens L. var. subloliacea (Laestad.) Hartm. Handb. Skand. Fl. ed. 4, 299
(1843); Raup in Jour. Arn. Arb. 17: 214 (1936).

Carex canescens L. f. subloliacea Laestad in Nova Acta Soc. Sci. Upsal. 11: 282 (1839).
Carex curta sensu Boott in Hooker, Fl. Bor.-Am. 2: 214 (1839), in part, non Good.
Twelve mi. east of Moraine Pt., northwest shore of Great Slave L., Bedford (Can.).
Common in the Athabaska—Great Slave L. region.

Carex gynocrates Wormsk. in Drejer, Nat. Tidssk. 3: 434 (1841); Mackenzie in N. Am.
Fl. 18: 98 (1931); Raup in Jour. Arn. Arb. 17: 214 (1936); Hultén, Fl. Alaska &
Yuk. Pt. 2: 302 (1942): Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

Carex dioica sensu Boott in Hooker, Fl. Bor.-Am. 2: 208 (1839), non L.

Wet mossy muskeg near east end of Brintnell L., 9353, 9577, 9581; Plains of Abraham,
Canol Rd., alt. 6000’, Porsild 11795 (Can.).

Common; mature in late July. Porsild also noted it along a small tributary
to Little Keele R. along the Canol Rd., mi. 51 E. In FI. Bor.-Am. it is said to
occur from “Carleton House to the Arctic Sea.” Our 9587 has unusually short
perigynia, scarcely longer than the scales. They are somewhat more abruptly
beaked than the typical form, and the beaks are not so prominently ceflexed at
maturity.

Carex Macloviana d’Urv. in Mem. Soc. Linn. Paris 4: 599 (1826); Mackenzie in N. Am.
Fl. 18: 126 (1931); Porsild in Nat. Mus. Can. Bull. 101: 16 (1945).

Carex festiva Dewey; Boott in Hooker, Fl. Bor.-Am. 2: 215 (1839), in part.

Mackenzie R., between Blackwater R. and Norman, Crickmay 159 (Can.) ; Macmillan
Pass, Canol Rd., mi. 225 E., Porsild 11211 (Can.).

The record under C. festiva in Fl. Bor.-Am., “Cumberland House, Bear Lake,”
probably refers in part to this species. Hultén (FI. Alaska & Yuk. Ft. 22 33D)
apparently considers that northwestern American material of the C. acloviana
complex represents ssp. pachystachya (Cham.) Hultén (1. c.). Mackenzie,
however, maintains the two as separate species, the latter distinguished by its
plano-convex perigynia with beak inconspicuously white-hyaline-tipped. Ac-
cording to Mackenzie, C. Macloviana has the body of its perigynia concave ven-
trally, with the beak strongly white-hyaline-tipped.

Carex phaeocephala Piper in Contr. U. S. Nat. Herb. 11: 172 (1906); Hultén, FI.
Alaska & Yuk. Pt. 2: 319 (1942).

Shale rock-slides and rocky stream channels on summit and upper slopes of Red Mt., alt.
4300’'-5900', 92977, 9529, 9531, 9772, 9780, 9795.

Common above timber-line on Red Mt., but not seen elsewhere in the Brint-
nell L. district. In anthesis late in June. Our specimens have darker scales
(some of them almost black) than most of the material in the Gray Herbarium.
The scales, except the lowermost in the spikes, have margins slightly if at all
hyaline, and the beaks of the perigynia are only slightly hyaline at the tips.
Descriptions of both C. phaeocephala and the closely related C. platylepis (cf.
Mackenzie in N. Am. Fl. 18: 139, 142) state that these species have prominent,
hyaline margins on the scales; but in the material at the Gray Herbarium there
is considerable variability in this character. Our plants are therefore placed
here until further study proves whether or not they are worthy of segregation.

Carex Soperi sp. nov. Fic. 12.

Culmi 20-35 cm. alti, e rhizomatibus squamosis 2 mm, crassis singillatim vel
pauci-aggregatim adscendentes, graciles sed rigidi, acute 3-angulati, supra scabri ;
foliis e culmis basim versus plerumque adscendentibus, pallido-viridibus, 6-15
mm. longis, 2.5—3 mm. latis, planis, longo-acuminatis, 3 vel 4 per culmum fer-

130 SARGENTIA [6

tilem, vaginis ventro albo-hyalinis ; capitulis dense aggregatis ovatis; spicis circa
5, gynocandris ; perigyniis atro-rubro-fuscis, margine subnigris, late ovatis, alatis,
3.54 mm. longis, 1-1.5 mm. latis, ad rostrum circiter 0.6 mm. longum rotunda-
tum contractis, dorso plurinerviis, ventro leviter paucinerviis, parte superiore
alae et rostri subtiliter serrulata, rostro atro-fusco vel nigro apice obliquo;
squamis costa subviridi et margine angusto hyalino exceptis atro-rubro-fuscis vel
nigris, longitudine corpore perigynii subaequalibus latitudine angustioribus ;
acheniis lenticularibus, stigmatibus 2.

Damp grassy place in willow-birch scrub, upper valley of Frost Cr., north of Brintnell L.,
alt. 4000’, 9534 (Type).

Common in this situation but not seen elsewhere. In flower characters and
the general appearance of leaves and culms these plants closely resemble C.
festivella Mackenzie, but they differ from this as well as from most of the section
Ovales in being non-caespitose. The culms rise singly or few together at in-
tervals of several inches, and must be collected individually like those of C.
siccata.

Carex praticola Rydb. in Mem. N. Y. Bot. Gard. 1: 84 (1900); Mackenzie in N. Am.
Fl. 18: 140 (1931); Hultén, Fl. Alaska & Yuk. Pt. 2: 319 (1942).
Cultivated field at Simpson, 9140; Mackenzie R., between Blackwater R. and Norman,
Crickmay 157 (Can.).

Common at Simpson; in anthesis about mid-June.

Carex Crawfordii Fernald in Proc. Am. Acad. 37: 469, pl. 1, f. 12, 13 (1902); Mackenzie
in N. Am. Fl. 18: 145 (1931); Raup in Jour. Arn. Arb, 17: 215 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 2: 320 (1942).

“Carlton House to Bear Lake,” Richardson (G).

In Fl. Bor.-Am. this species was probably included in C. scoparia, which was
noted as “Common in British America.” It is common about L. Athabaska,
but the above records are the only evidence for its presence in southwestern
Mackenzie.

Carex Bebbii Olney (Caric. Bor.-Am. 3, name only, 1871; L. H. Bailey in Mem. Torr.
Bot. Cl. 1: 55, as synonym, 1889); Fernald in Proc. Am. Acad. 37: 478 (1902).
Muskeg along road west of the Mackenzie R. near Simpson, 9899; upland prairie west
of the Mackenzie R., near Simpson, 9889.

Occasional ; fruit mature in late August.

Carex aenea Fernald in Proc. Am. Acad. 37: 480, pl. 3, f. 65, 66 (1902); Mackenzie in
N. Am. FI. 18: 169 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 320 (1942).

Southerly slope in an old field at Simpson, 9136.
Occasional ; in anthesis about mid-June. This material is very young and is
placed here tentatively. Its inflorescences are rather stiff for C. aenea, though

this is a common condition in young plants. Number 9136 matches specimens
collected on L. Athabaska in 1935 (Raup in Jour. Arn. Arb. 17: 215).

Carex leptalea Wahl. in Vet. Akad. Handl. (Stockh.) 24: 139 (1803); Mackenzie in N.
Am. Fl. 18: 173 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 306 (1942),

Mackenzie R., Richardson (N); damp place in upland prairie west of Mackenzie R. near
Simpson, 9902.

Occasional in the Simpson area and observed once at Brintnell Lake; fruit
maturing in late August. Occasional also in the Athabaska—Great Slave L.
region (Raup in Jour. Arn. Arb. 17: 215), and collected at Great Bear L. by
Porsild (Sargentia 4: 18).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 131

s i

LECRER

Fic. 12. Carex Soperi Raup, sp. nov.

132 SARGENTIA [6

Carex obtusata Liljebl. in Vet. Akad. Handl. (Stockh.) 14: 69, ¢. 4 (1793); Mackenzie
in N. Am. FI. 18: 181 (1935); Hultén, FI. Alaska & Yuk. Pt. 2: 305 (1942); Por-
sild in Sargentia 4: 18 (1943).
This species has not yet been observed in the upper Mackenzie country, but
is common in parts of the Athabaska-Great Slave L. region (Raup in Jour. Arn.
Arb. 17: 215) and along the lower Mackenzie (Porsild, 1. c.).

Carex supina Willd. ssp. spaniocarpa (Steud.) Hultén, Fl. Alaska & Yuk. Pt. 2: 365
(1942).

Carex spaniocarpa Steud. Synop. Cyp. 225 (1855).
Carex supina Willd. ex Wahl.; Mackenzie in N. Am. Fl. 18: 182 (1935) ; Raup in Jour.
Arn, Arb, 17: 215 (1936).
Known in the Athabaska—Great Slave L. region (Raup, 1. c.) and at Great
Bear L. (Richardson, G, N), but not yet collected in the upper Mackenzie
country.

Carex deflexa Hornem. Dansk. Oecon. Pl. ed. 3, 1: 938 (1821); Mackenzie in N. Am.
Fl. 18: 199 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 263 (1942).

Pebbly beach on shore of Brintnell L., 9167; along small stream, east slope of Red Mt.,
alt. 4500’, 9284.

Common in the above situations but not seen elsewhere ; spikes well developed
in late June.

Carex Rossii Boott in Hooker, Fl. Bor.-Am. 2: 222 (1839); Mackenzie in N. Am. FI.
18: 201 (1935); Raup in Jour. Arn. Arb. 17: 216 (1936); Hultén, Fl. Alaska &
Yuk. Pt. 2: 364 (1942).

This species has been collected just at the southeastern corner of our region,

at Fort Smith (Raup 326).

Carex tonsa (Fernald) Bickn. in Bull. Torr. Bot. Cl. 35: 492 (1908); Mackenzie in N,
Am. FI. 18: 205 (1935). See Fernald in Rhod. 44: 288-290 (1942).

Carex umbellata sensu Boott in Hooker, Fl. Bor.-Am. 2: 222 (1839), in part; Richardson

in Arct. Search. Exp. 2: 344 (1851) ; non Schk.

Carex umbellata Schk. var. tonsa Fernald in Proc. Am. Acad. 37: 507 (1902); Raup in

Jour. Arn. Arb. 17: 216 (1936).

This plant was noted by Richardson (1. c.) at Methye Portage and Fort
Simpson, as C. umbellata. As such it was noted in Fl. Bor.-Am. as occurring
in “British America.” I have seen a Richardson specimen from Chipewyan
(Herb. G), and have collected the species on the south shore of L. Athabaska
(6724). Both of these specimens represent var. tonsa. The Simpson record
made by Richardson, presumably also the variety, is the only evidence we have
of the plant north of Athabaska L.

Carex scirpoidea Michx. Fl. Bor.-Am. 2: 171 (1803); Boott in Hooker, Fl. Bor.-Am., 2:
208 (1839); Macoun, Cat. Can. Pl. Pt. 4: 112 (1888); Mackenzie in N. Am. FI.
18: 207 (1935); Raup in Jour. Arn. Arb, 17: 216 (1936); Hultén, Fl. Alaska &
Yuk. Pt. 2: 303 (1942); Porsild in Nat. Mus. Can. Bull. 101: 16, 30 (1945).
Northwest shore of Great Slave L., Bedford (Can.); Mackenzie R., Richardson (N);
Mackenzie R., Wrigley to Blackwater R., Crickmay 129 (Can.); Blackwater R. to Norman,
Crickmay 156 (Can.); Norman, Richardson (N); mossy thickets along south shore of
Brintnell L., 9363, 9364; south slope of Red Mt., alt. 4600’, 9237; north slope of Colonel Mt.,
alt. 3500’-5500’, 9395, 9626; north peak, Nahanni Mt., Wynne-Edwards 8431 (Can.) ; Lone
Mt., Wynne-Edwards 8432 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards
8252, 8253, 8254 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11853 (Can.).

Occasional to common on the shores of Brintnell L., particularly on north ex-
posures. On the cold north slope of Colonel Mt. it is abundant, and above

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 133

timber line it becomes a primary species on turfy slopes. It is also a predominant
species on parts of the south slope of Red Mt. In anthesis in July.

Carex concinna R. Br. apud Richardson, Bot. App. Frankl. Journ. 751 (1823); Boott
in Hooker, Fl. Bor.-Am. 2: 223 (1839); Macoun, Cat. Can. Pl. Pt. 4: 158 (1888);
Mackenzie in N. Am. FI. 18: 212 (1935); Raup in Jour. Arn. Arb. 17: 216 (1936) ;
Hultén, Fl. Alaska & Yuk. Pt. 2: 365 (1942); Porsild in Nat. Mus. Can. Bull.
101: 30 (1945).

Mackenzie R., Richardson (N); muskeg thickets at west end of Brintnell L., 9250; north
peak, Lone Mt., Wynne-Edwards 8429, 8430 (Can.) ; dry dolomitic summit of Nahanni Mt.,
W ynne-Edwards 8428 (Can.).

Common in the Brintnell Lake habitat noted above, but not seen elsewhere in
the district ; fruit maturing in late June. Noted in Fl. Bor.-Anv. from “Cumber-
land House to Mackenzie River.” Porsild (Sargentia 4: 18) has collected it
at Great Bear L. and in the lower Mackenzie region.

Carex Richardsonii R. Br. apud Richardson, Bot. App. Frankl. Journ. 751 (1823);
Mackenzie in N. Am. FI. 18: 213 (1935); Raup in Jour. Arn. Arb. 17: 216 (1936).

Fort Smith, Slave R., Raup 303, 1885.

In the original description the locality for this species was given as “W,”
meaning “wooded country from latitude 54° to 64° north.” Boott, in Fl. Bor.-
Am., gave the locality as “Norway and Cumberland House.” The species oc-
curs rather commonly in the Wood Buffalo Park, and has been collected just
within the southeast corner of our region at Ft. Smith. It undoubtedly occurs
along the upper Mackenzie.

Carex rupestris All. Fl. Pedemont. 2: 264, pl. 92, f. 1 (1785); Boott in Hooker, Fl. Bor.-
Am. 2: 209 (1839); Macoun, Cat. Can. Pl. Pt. 4: 113 (1888); Mackenzie in N. Am.
Fl. 18: 220 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 304 (1942); Porsild in Nat.
Mus. Can. Bull. 101: 16 (1945).

Norman, Richardson (G) ; Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11791 (Can.).
Collected by Porsild (Sargentia 4: 18) also at Great Bear L. and in the lower
Mackenzie country.

Carex glacialis Mackenzie in Bull. Torr. Bot. Cl. 37: 244 (1910); Mackenzie in N. Am.
Fl. 18: 221 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 366 (1942); Porsild in Nat.
Mus. Can. Bull. 101: 16 (1945).

Bald limestone mountain on Plains of Abraham, Canol Rd., Porsild 11790 (Can.) ; Sekwi

R., Canol Rd., mi. 174 E., Porsild 11849 (Can.).

Carex eburnea Boott in Hooker, Fl. Bor.-Am. 2: 226, pl. 225 (1839); Macoun, Cat. Can.
Pl. Pt. 4: 157 (1888); Mackenzie in N. Am. FI. 18: 223 (1935); Raup in Jour. Arn.
Arb. 17: 216 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 366 (1942); Porsild in
Nat. Mus. Can. Bull. 101: 16, 30 (1945).

Norman, Richardson, June, 1826 (G, N) (apparently part of the type material) ; north
face of Lone Mt., Wynne-Edwards 8434 (Can.) ; mossy slopes on summit of Lone Mt.,
Wynne-Edwards 8435 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., alt. 3625’, Porsild 11947
CCani):

Widely distributed in the central and northern parts of the Mackenzie basin,
apparently confined to limestone or dolomitic rocks (Raup, 1. c.; Porsild in

Sargentia 4: 19).
Carex Garberi Fernald var. bifaria Fernald in Rhod. 37: 255 (1935); Hultén, Fl. Alaska
& Yuk. Pt. 2: 333 (1942).
Carex Hassei sensu Mackenzie in N. Am. Fl. 18: 233 (1935), at least in part, non Bailey.
Stony beaches near west end of Brintnell L., 9174,

134 SARGENTIA [6

Common in this habitat; perigynia beginning to mature in the last week of
June. Specimens representing the typical form have been collected at Great
Slave L. (Raup in Jour. Arn. Arb. 17: 217), but the northwestern extent of
these forms is yet uncertain. Porsild (Sargentia 4: 19) reports C. Garberi,
presumably in its typical form, at Great Bear L. and in the Mackenzie delta.

Carex aurea Nutt. Gen. 2: 205 (1818); Mackenzie in N. Am. FI, 18: 234 (1935); Raup
in Jour. Arn. Arb, 17: 217 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 332 (1942).

Fort Smith, Raup 295; Mackenzie R., Richardson (G) ; upland prairie west of Mackenzie
R., near Simpson, 9934; wet mossy muskeg thickets at west end of Brintnell dnp D7 OL

Common ; mature in August. The plants collected at Brintnell L. are char-
acterized by distinctly pear-shaped perigynia. This form is nearly all lost in
drying. The Simpson specimens, as well as others collected in the Mackenzie
basin earlier, all have well-rounded perigynia. Mackenzie (1. c.) describes the
species as having orbicular-obovoid perigynia and says nothing about a pyriform
tendency, although he cites C. pyriformis Schw. as a synonym. Schweinitz de-
scribed this as having “capsules pyriform” (Ann. Lye. N. Y. 1: 69, 1824), but
later he reduced it, himself, to C. aurea. Whether or not the form is worthy of
recognition can be decided only after more field notes on the shape of the peri-
gynia have accumulated, especially since the character is largely lost in drying.

Carex vaginata Tausch in Flora 4: 557 (1821); Macoun, Cat. Can. Pl. Pt. 4: 153 (1888);
Raup in Jour. Arn. Arb. 17: 217 (1936); Hultén, Fl. Alaska & Yuk, Pt. 2: 372
(1942); Porsild in Nat. Mus. Can. Bull. 101: 16, 30 (1945).

Carex saltuensis Bailey, Macoun, Cat. Can. Pl. Pt. 5: 282 (1890) ; Mackenzie in N. Am.
Fl. 18: 241 (1935).

Carex phaeostachya sensu Boott in Hooker, Fl. Bor.-Am. 2: 226 (1839), in part, non
* Smith.

Damp mossy thickets and gravel shores around Brintnell L., 9248, 9360; mossy banks in
willow-birch scrub near iron spring near west end of Brintnell L., 9528; open mossy woods
on north slope of Colonel Mt., 9419; north peak of Nahanni Mt., alt. 2500’, Wynne-Edwards
8436 (Can.) ; Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11794 (Can.) ; Bolstead Cr.,
Canol Rd., mi. 111 E., Wynne-Edwards 8255, 8256, 8257 (Can.).

Occasional to common around the shore of Brintnell L., and very common
on the north-facing slope of Colonel Mt. up to about 3500’: mature in the latter
part of July. Noted in Fl. Bor.-Am. at “Fort Norman, Mackenzie River.”

Carex capillaris L. Sp. Pl. 977 (1753); Boott in Hooker, Fl. Bor.-Am, 2: 227 (1839);
Macoun, Cat. Can. Pl. Pt. 4: 163 (1888); Mackenzie in N. Am. Fl, 18: 295 (1935);
Raup in Jour. Arn. Arb. 17: 217 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 374
(1942); Porsild in Nat. Mus. Can. Bull. 101: 16, 30 (1945).

Norman, Richardson (G, N) ; wet mossy banks and muskeg thickets around Brintnell es
9173, 9244; north peak of Nahanni Mt., Wynne-Edwards 8437 (Can.); Bolstead Cr., Canol
Rd., mi. 111 E., Wynne-Edwards 8260 (Can.) ; Sekwi R., Canol Rd., alt. 3625’, mi, 174 Big
Porsild 11852 (Can.) ; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11209 CCan:)is

Common to abundant in the Brintnell L. habitats: mature late in July.

Carex Oederi Retz. var. viridula (Michx.) Kiikenth. in Pflanzenr. 38 (IV. 20): 674
. (1909); Hultén, Fl. Alaska & Yuk, Pt. 2: 377 (1942).
Carex viridula Michx. Fl. Bor.-Am. 2: 170 (1803) ; Mackenzie in N. Am. Fl. 18: 303
(1935).
Carex Oederi Retz. var. pumila (Cosson & Germ.) Fernald in Rhod. 8: 201 (1906) ;
Raup in Jour. Arn. Arb. 17: 217 (1936) ; Porsild in Sargentia 4: 19 (1943).

This species has not yet been observed along the upper Mackenzie, but it has
been collected at Great Slave (Raup, |. c.) and Great Bear (Porsild, 1. c.) Lakes.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 135

Carex petricosa Dewey in Am. Jour. Sci. 29: 246, pl. W, f. 70 (1836); Porsild in Sar-
gentia 4: 19 (1943), in Nat. Mus. Can. Bull. 101: 16, 30 (1945).

Lone Mt., Wynne-Edwards 8438 (Can.) ; north peak, Nahanni Mt., Wynne-Edwards 8439
(Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alt. 4420’, Porsild 11830 (Can.) ; same place,
W ynne-Edwards 8251 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11848 (Can.).

Carex misandra R. Br. App. Parry’s Voy. 283 (1823); Mackenzie in N. Am. Fl. 18:
312 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 273 (1942); Porsild in Nat. Mus.
Can. Bull. 101: 16 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., alt. 6000’, Porsild 11792 (Can.) ; Bolstead Cr

Canol Rd., mi. 111 E., Wynne-Edwards 8247, 8248, 8249, 8250 (Can.).

Porsild (1. c.) noted the species also in Macmillan Pass at mi. 219 E.

Carex atrofusca Schk. Riedgr. 106, pl. Y, f. 82 (1801); Mackenzie in N. Am. FI. 18:
313 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 373 (1942); Porsild in Sargentia 4:
20 (1943), in Nat. Mus. Can, Bull. 101: 16 (1945).

Bald limestone mountain on Plains of Abraham, Canol Rd., alt. 6000’, mi. 82 E., Porsild
11793 (Can.).

Carex lasiocarpa Ehrh. var. americana Fernald in Rhod. 44: 304 (1942).
Carex lasiocarpa Ehrh. sensu Mackenzie in N. Am. Fl. 18: 330 (1935) ; Raup in Jour.
Arn. Arb. 17: 217 (1936).

Evidence for the occurrence of C. lasiocarpa in southwestern Mackenzie is
very scanty. A record in Fl. Bor.-Am. under the closely related C. lanuginosa
Michx. reads “Canada to Mackenzie River.” The latter species apparently
has not been collected anywhere in the Mackenzie basin, but C. lasiocarpa has
proved to be abundant on the south shore of Lake Athabaska (Raup, |. c.) and
probably will be found farther northward. It is possible that the early note
in Fl. Bor.-Am. refers to it.

Carex rariflora (Wahl.) Smith in Engl. Bot. pl. 2516 (1813); Boott in Hooker, Fl. Bor.-
Am. 2: 224 (1839); Macoun, Cat. Can. Pl. Pt. 4: 150 (1888); Mackenzie in N. Am.
Fl, 18: 348 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 367 (1942); Porsild in Nat.
Mus. Can. Bull. 101: 16 (1945).
“Mackenzie River ex Boott” (G); “Fort Franklin,” Richardson (N); arctic tundra at
east end of Macmillan Pass, Canol Rd., mi. 204 E., Porsild 11885 (Can.).

Noted in Fl. Bor.-Am. at ‘Fort Franklin, Mackenzie River.”

Carex limosa L. Sp. Pl. 977 (1753); Boott in Hooker, Fl. Bor.-Am. 2: 224 (1839);
Mackenzie in N. Am, FI. 18: 350 (1935); Raup in Jour. Arn. Arb. 17: 218 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 2: 369 (1942); Porsild in Sargentia 4: 20 (1943).

This species has not been collected in the upper Mackenzie country, though
it is to be expected there. It is occasional throughout much of the Athabaska—

Great Slave L. region (Raup, |. c.), and has been collected by Porsild (1. c.)

at Great Bear L. and in the Eskimo Lakes basin.

Carex paupercula Michx. var. irrigua (Wahl.) Fernald in Rhod. 8: 76 (1906); Raup in
Jour. Arn. Arb. 17: 218 (1935),

Carex limosa L. var. irrigua Wahl. Act. Holm. 162 (1803).
Carex magellanica sensu Hultén, Fl. Alaska & Yuk. Pt. 2: 370 (1942), non Lam.
Carex paupercula Michx. sensu Mackenzie in N. Am. Fl. 18:°351 (1935); Porsild in
Sargentia 4: 20 (1943).
This species has been collected in the Athabaska—Great Slave L. region (Raup,
I. c.) and at Great Bear L. (Porsild, 1. c.), but has not been seen along the
upper Mackenzie.

136 SARGENTIA [6

Carex media R. Br. apud Richardson, Bot. App. Frankl. Journ. 750 (1823); Porsild in

Sargentia 4: 20 (1943), in Nat. Mus. Can. Bull. 101: 16 (1945). See Rhod. 35: 220-
223 (1933), 41: 203-205 (1939), 44: 303-304 (1942).

Carex Vahlii sensu Boott in Hooker, Fl. Bor.-Am. 2: 216 (1839) ; Mackenzie in N. Am.
Fl. 18: 357 (1935) ; non Schk.

Carex Vahlii Schk. var. inferalpina (Wahl.) Fernald; Raup in Jour. Arn. Arb. 17: 218
(1936).

Carex norvegica Retz. ssp. inferalpina (Wahl.) Hultén, Fl. Alaska & Yuk. Pt. 2: 348
(1942).

Carex alpina sensu Macoun, Cat. Can. Pl. Pt. 4: 136 (1888), non Liljebl.

Damp roadway in lowland woods near Simpson, 9092; stony beaches and muskeg shores
of Brintnell L., 9170, 9175, 9197, 9362; Macmillan Pass, Canol. Rd., mi. 225 E., Porsild
11208 (Can.).

Common in muskeg thickets at Brintnell L., but rare or occasional elsewhere
in the vicinity; mature in July. In Fl. Bor.-Am. C. Vaslii is noted as “Common
in British America, to the Arctic Coast.” Porsild (Sargentia 4: 20) notes C.
media as common in the wooded country of the Mackenzie northward to the delta,
and at Great Bear L.

Carex nesophila Holm in Am. Jour. Sci. IV. 17: 315 (1904); Porsild in Nat. Mus. Can,
Bull. 101: 16 (1945).

Grassy meadow on north slope of Colonel Mt., alt. 5000’, 9613; loose shale on summit of
Red Mt., alt. 5900’, 9789; mountain slope south of Little Keele R., Canol Rd., mi. 55 E.,
Porsild 11784 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8258 (Can.).

Common locally on the mountains at Brintnell L.; found flowering in late July
and early August.

Carex podocarpa R. Br. apud Richardson, Bot. App. Frankl. Journ. 751 (1823); Boott
in Hooker, Fl. Bor.-Am. 2: 224 (1839); Macoun, Cat. Can. Pl. Pt. 4: 149 (1888);
Mackenzie in N. Am. Fl. 18: 362 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 362
(1942); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945). See Porsild in Rhod. 41:
205-206 (1939),

Shale slides and turfy slopes, south side of Red Mt., alt. 5000’-5800', 9752, 9793; north-
east and east slopes of Red Mt., alt. 4500’-5500’, 9283, 9788; west slope of Terrace Mt., alt.
4500’-5000’, 9538; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8259 (Can.).

Occasional to common on south- and west-facing alpine slopes at Brintnell L.,
but abundant on the north and east sides of Red Mt. Here it is one of the most
prominent species in the turf. In flower late in June; mature early in August.
Porsild (1. c.) notes it as common in Macmillan Pass and elsewhere along the

Canol Rd.

Carex atrata L. ssp. atrosquama (Mackenzie) Hultén, Fl. Alaska & Yuk. Pt, 2: 354
(1942).

Carex atrosquama Mackenzie in Proc. Biol. Soc. Wash. 25: 51 (1912), in N. Am. Fi.
18: 366 (1935); Porsild in Sargentia 4: 21 (1943), in Nat. Mus. Can. Bull. 101: 17
(1945).

Pebble beaches and stony thickets, shore of Brintnell L., 9/64; grassy meadow, north
slope of Colonel Mt., alt. 5000’, 9614; south slope of Red Mt., in damp thickets, 9233;
springy place in birch scrub, upper valley of Frost Cr., alt. 4500’, 9517; Macmillan Pass,
Canol Rd., mi. 225 E., Porsild 11215 (Can.).

Occasional to common in the Brintnell L. sites; collected with immature peri-
gynia late in July. Mackenzie described this species from northern Rocky Moun-
tain material, stating that it differs from C. atrata L. and others of this relationship

in having somewhat inflated perigynia (little flattened at maturity) and the peri-

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 137

gynia not much broader than the achenes. There is a large series of Eursian C.
atrata in the Gray Herbarium; and on these the mature achenes are often inflated
and scarcely margined by compression. In fact our material is a good match for
many of the old-world specimens. There seems to be a tendency to smaller
spikes and shorter bracts.on the American plants, but I am inclined to agree with
Hultén that they should be recognized at most as a subspecies.

Carex albo-nigra Mackenzie in Rydb. Fl. Rocky Mts. 137 (1917); Mackenzie in N. Am.
Fl. 18: 369 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 353 (1942).

Shale slide rock on south slope of Red Mt., near summit, alt. 5900’, 9745.

Common; fruit mature early in August. Some of our plants have unusually
long lateral spikes (2 cm.), on peduncles that are somewhat longer. Mackenzie
characterizes the species (N. Am. FI. 18: 369) as having lateral spikes 8-10 mm.
long, on erect peduncles shorter than the spikes. The lowest spike on some of
our specimens is attenuate, with flowers widely separate. This tendency to-
ward elongated spikes is also seen in plants from the Rocky Mountains of the
upper Peace River region (Raup & Abbe 4129). Other specimens in no. 9745
match the typical forms.

Carex atratiformis Britton in Bull. Torr. Bot. Cl. 22: 222 (1895); Mackenzie in N. Am.
Fl. 18: 372 (1935); Raup in Jour. Arn. Arb. 17: 218 (1936); Hultén, Fl. Alaska
& Yuk. Pt. 2: 355 (1942).
This species is occasional in the Wood Buffalo Park, and was collected along
the Frances R. by Dawson in 1887 (Can.). It is to be expected along the upper
Mackenzie and lower Liard Rivers.

Carex Buxbaumii Wahl. in Vet. Akad. Handl. (Stockh.) 24: 163 (1803); Mackenzie in
N. Am. Fl. 18: 374 (1935); Raup in Jour. Arn. Arb. 17: 218 (1936); Hultén, FI.
Alaska & Yuk. Pt. 2: 350 (1942); Porsild in Sargentia 4: 21 (1943).

Known in the Wood Buffalo Park, at Great Bear L., and in Alaska, but not
yet collected along the upper Mackenzie.

Carex lenticularis Michx. Fl. Bor.-Am. 2: 172 (1803); Boott in Hooker, Fl. Bor.-Am.
2: 219 (1839); Macoun, Cat. Can. Pl. Pt. 4: 145 (1888); Mackenzie in N. Am. FI.
18: 385 (1935); Raup in Jour. Arn. Arb. 17: 219 (1936).

This species was noted in Fl. Bor.-Am. as occurring from “Canada to Mack-
enzie River.’ It is occasional in the Athabaska—Great Slave L. region, and there
are several Richardson specimens in Herbs. G and N from Great Bear L. It is
to be expected, therefore, in southwestern Mackenzie.

Carex aquatilis Wahl. in Vet. Akad. Handl. (Stockh.) 24: 165 (1803); Boott in Hooker,
Fl. Bor.-Am. 2: 219 (1839); Macoun, Cat. Can. Pl. Pt. 4: 143 (1888); Mackenzie
in N. Am. FI. 18: 396 (1935); Raup in Jour. Arn, Arb. 17: 219 (1936); Hultén, FI.
Alaska & Yuk. Pt. 2: 338 (1942); Porsild in Sargentia 4: 23 (1943). See Fernald
in Rhod. 44: 295-298 (1942).

Northwest shore of Great Slave L., Bedford (Can.); Rae, Bedford (Can.) ; Mackenzie

R., Richardson (G); wet gravelly and marshy shores of Brintnell L., 9311, 9497; recently

formed mud flat at west end of Brintnell L., 9700.

Common to abundant; fruit mature in late July and early August. A primary
species on marshy ground at the lake shore, where it is associated with Calama-
grostis neglecta and C. canadensis var. robusta... Noted by Richardson (Arct.
Search. Exp. 2: 344-345) at Simpson, and in Fl. Bor.-Am. from “Canada to

138 SARGENTIA [6

Mackenzie River.” Porsild states that it is common throughout the Mackenzie
valley.

Carex aquatilis Wahl. var. stans (Drejer) Ostenf. Fl. Arct. 70, f. 43 (1902); Hultén,
Fl. Alaska & Yuk. Pt. 2: 338 (1942).
Carex stans Drejer, Revis. Crit. Car. 458 (1841); Porsild in Nat. Mus. Can. Bull. 101:
17 (1945).
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11213 (Can.).

Carex microglochin Wahl. in Vet. Akad. Handl. (Stockh.) 24: 140 (1803); Mackenzie
in N. Am. FI. 18: 424 (1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 310 (1942); Porsild

in Sargentia 4: 24 (1943).
Mud flats near west end of Brintnell L., 9870.
Occasional, forming small round colonies in an association of Equisetum varie-
gatum and mosses on the recently formed deposits of glacial mud; found with
mature perigynia in mid-August.

Carex atherodes Spreng. Syst. Veg. 3: 828 (1826); Mackenzie in N. Am. FI. 18: 439
(1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 384 (1942).
Carex trichocarpa Muhl. sensu Boott in Hooker, Fl. Bor.-Am, 2: 222 (1839), in part.
Carex trichocarpa Muhl. var. aristata (R. Br.) Bailey; Raup in Jour. Arn. Arb. 17: 219
(1936).
Lower Slave R., Raup 316; Mackenzie R., Richardson (N); grassy swale near Ft. Liard,
Nowosad (Can.).
This is one of the most abundant sedges in the river lowlands of the Wood
Buffalo Park, and no doubt it extends far into the upper Mackenzie country.
It has also been collected at Whitehorse Rapids in the upper Yukon country

(Hultén, 1. c.).

Carex physocarpa Pres!. Relig. Haenk. 1: 205 (1830); Mackenzie in N. Am. FI. 18: 448
(1935); Hultén, Fl. Alaska & Yuk. Pt. 2: 380 (1942); Porsild in Sargentia 4: 24
(1943), in Nat. Mus. Can. Bull. 101: 17 (1945). See O’Neill & Duman in Rhod.

43: 418-425 (1941).

Marshes and damp stony beaches, shore of Brintnell L., 9195, 9365; wet meadow at west
end of Brintnell L., 9713, 9716; Sekwi R., Canol Rd., mi. 174 E., Porsild 11851 (Can.) ;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11206 (Can.).

Abundant in shallow water along the shores of Brintnell L., but only occasional
in the other habitats mentioned ; mature early in August.

This species usually develops stout rootstocks, with culms rising singly or
few together. Number 9713 is distinctive in forming dense tussocks from very

short crowded rootstocks.

Carex saxatilis L. Sp. Pl. 976 (1753); Raup in Jour. Arn. Arb. 17: 220 (1936); Porsild
in Sargentia 4: 24 (1943).
This species has not yet been collected in the upper Mackenzie country, but it
has been found at Great Slave L. (Raup, 1. c.) and at Great Bear L. (Porsild, i.
c.). There is also a specimen in Herb. G collected by Dutilly on the lower Mack-

enzie.

Carex membranacea Hooker, App. Parry’s 2° Voy. 406 (1825); Boott in Hooker, FI.
Bor.-Am. 2: 220 (1839); Mackenzie in N. Am. FI. 18: 454 (1935); Hultén, FI.
Alaska & Yuk. Pt. 2: 382 (1942); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Carex membranopacta Bailey; Raup in Jour. Arn. Arb. 17: 220 (1936).
Mackenzie, 1886, R. Bell (Can. no. 34724); Mackenzie R., between Wrigley and Black-
water R., Crickmay 128 (Can.) ; between Blackwater R. and Norman, Crickmay 158 (Can.) ;

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 139

wet mossy and gravelly banks on shore of Brintnell L., 9198, 9199, 9338; muskeg thickets
at west end of Brintnell L., 9246, 9247; mud flat at west end of Brintnell L., 9265-4; Bol-
stead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8261, 8262 (Can.); Sekwi R., Canol Rd.,
mi. 174 E., alt. 3625’, Porsild 11850 (Can.) ; Macmillan Pass, Canol Rd., mi. 225 E., Porsild
11202 (Can.).

Occasional to common at Brintnell L., especially in wet muskeg thickets; ma-
ture in July. Noted by Porsild along the Canol Rd., and also at Little Keele R.
and the Plains ef Abraham. The record in Fl. Bor.-Am. reads “Cumberland
House to Arctic Coast.”

Carex rostrata Stokes in Withering, Brit. Pl. ed. 2, 2: 1059 (1787); Macoun, Cat. Can.
Pl. Pt. 4: 170 (1888); Mackenzie in N. Am. FI. 18: 456 (1935); Raup in Jour. Arn.
Arb. 17: 229 (1936); Hultén, Fl. Alaska & Yuk. Pt. 2: 378 (1942).

Carex ampulacea Good. sensu Boott in Hooker, Fl. Bor.-Am. 2: 221 (1839).
Northwest shore of Great Slave L., Bedford (Can.) ; Mackenzie R., Richardson (G).

This species is one of the most abundant in the Athabaska—Great Slave L.
region. Its northern limit is yet unknown, though it undoubtedly extends far
into the Mackenzie valley. The record in Fl. Bor.-Am. reads “Cumberland
Piquee, Bear Lake...

Carex retrorsa Schw. in Ann. Lyc. N. Y. 1: 71 (1824); Raup in Jour. Arn. Arb. 17:
220 (1936).

Grassy swale near Ft. Liard, Nowosad (Can.).

Carex oligosperma Michx. Fl. Bor.-Am. 2: 174 (1803); Boott in Hooker, Fl. Bor.-Am.
2: 220 (1839); Macoun, Cat. Can. Pl. Pt. 4: 168 (1888); Mackenzie in N. Am. FI.
18: 461 (1935); Raup in Jour. Arn. Arb. 17: 220 (1936); Porsild in Sargentia 4:
25 (1943).

A note on this species in Fl. Bor.-Am. says it extends from “Cumberland
House to Bear Lake,” and there is a specimen in Herb. G which bears the ambig‘u-
ous label, “English River, Bear Lake, Norway House.” Judging by later collec-
tions, the plant has a wide but spotty range in the north. It has been collected at
Methye Portage (Macoun), and at several places on the south side of Lake Atha-
baska (Raup, 1. c.). Porsild (1. c.) has recently reported a specimen from the
Yellowknife R., so that eventually the old Fl. Bor.-Am. range may be substan-
tiated.

ARACEAE
Calla palustris L. Sp. Pl. 968 (1753); Raup in Jour. Arn. Arb. 17: 221 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 2: 385 (1942); Porsild in Sargentia 4: 25 (1943).
Southwest and north shores of Great Slave L., Howe 91988 (Can.); northwest shore of
Great Slave L., Bedford (Can.); Liard R. between Nahanni Butte and Simpson, Crickmay
9 (Can.) ; border of lowland slough on west bank of the Mackenzie R. near Simpson, 9895,
Common in the Simpson locality, and also in the Athabaska—Great Slave Lake
region. Collected by Porsild (1. c.) along the lower Mackenzie.

LEMNACEAE
Lemna trisulca L. Sp. Pl. 970 (1753); Raup in Jour. Arn. Arb. 17: 221 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 2: 386 (1942); Porsild in Sargentia 4: 25 (1943).
This species is occasional in the Wood Buffalo Park and has been collected in
the Mackenzie River delta by Porsild (1. c.). It has not been seen along the
upper Mackenzie, though it is to be expected there.

140 SARGENTIA [6

JUNCACEAE

Juncus bufonius L. Sp. Pl. 328 (1753); Hooker, Fl. Bor.-Am, 2: 191 (1839); Macoun
in Cat. Can. Pl. Pt. 4: 60 (1888); Raup in Jour. Arn. Arb. 17: 221 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 3: 422 (1943).

Damp slough along wood road just west of Simpson, 9885.

Very common in this habitat, where it is a primary species on wet soil. It is
also common on damp sand bars along the Mackenzie R. near Simpson. Found
with maturing capsules late in August. Noted in Fl. Bor.-Am. as occurring
“throughout Canada to Bear Lake.”

Juncus biglumis L. Sp. Pl. 328 (1753); Hultén, Fl. Alaska & Yuk. Pt. 3: 421 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11797 (Can.).

Juncus albescens (Lange) Fernald in Rhod. 26: 202 (1924); Porsild in Nat. Mus. Can.
Bull. 101: 17 (1945).

Juncus triglumis L. var. albescens Lange, Conspect. Fl. Groenl. 123 (1880).

Juncus triglumis L., sensu Hultén, Fl. Alaska & Yuk. Pt. 3: 431 (1943), at least in part.
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8263 (Can.).

Noted by Porsild (1. ¢.) also on the Plains of Abraham along the Canol Rd.

Juncus triglumis L. Sp. Pl. 328 (1753); Hultén, Fl. Alaska & Yuk. Pt. 3: 431 (1943),
in part; Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11796 (Can.).
It should be noted that Hulten (1. c.) regards J. albescens as indistinguishable
from J. triglumis in Alaska.

Juncus Vaseyi Engelm. in Trans. St. Louis Acad. 2: 448 (1866).

Upland prairie west of Mackenzie R. near Simpson, 9925,

Common, especially at or near the margins of the prairies, where it is a pri-
mary species in places. In fruit late in August. Not previously noted north of
the Wood Buffalo Park (Raup in Jour. Arn. Arb. 17: 222), although the FI.
Bor.-Am. record (p. 191) for J. tenuis, “Lake Huron, to Bear Lake. . .”, may
refer to it in part.

Juncus balticus Willd. in Mag. Naturf. Fr. Berlin 3: 298 (1809); Hooker, Fl. Bor.-Am.
2: 189 (1839).

Slave R., Richardson (G); east shore of Trout L., C. O. Hage (G); Mackenzie R.,
Onion (N); rocky sandy bank of Mackenzie R. opposite Simpson, 9944; damp sand bar
along Mackenzie R. near Simpson, 9886; wet mossy sloughs and newly formed mud flats
around the west end of Brintnell L., 9705, 9802.

Common ; in fruit at Brintnell L. early in August and at Simpson late in that
month. Iam referring this material to J. balticus in its broad sense, for I am at
present unable to place it satisfactorily in any of the recognized segregates of that
species. The Mackenzie valley plants probably could be placed in var. littoralis
Engelm., but all of the material suggests a transition to some form of J. arcticus
Willd. It all has short anthers (about 1 mm. long), which are about as long as

the filaments.
Juncus Drummondii E. Mey. in Ledeb. Fl. Ross. 4: 235 (1853); Hultén, Fl. Alaska &
Yuk. Pt. 3: 425 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Stony beach on shore of Brintnell L. at mouth of Frost Cr., 9671; Macmillan Pass, Canol
Rd., mi. 225 E., Porsild 11218 (Can.).

Apparently rare at Brintnell L.; in flower early in August.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 141

Our number 9671 contains a single plant which exhibits characters that sug-
gest a varietal segregate of J. Drimmondii, but with such scant material I do not
think it wise to propose it. The plant has one flower in anthesis and another with
a mature capsule. The flowers closely resemble those of J. Drummondiu, though
they are smaller. The valves of the capsule are dark brown and shining, retuse
and mucronate like those of J. Drummondii; likewise the sepals, somewhat
shorter than the capsule, are dark brown and striped with green. The in-
florescences are single-flowered. The plant differs from J. Drummendii chiefly
in having its leaf-sheaths with a distinct, though short blade instead of a bristle,
and in having the auricles at the base of the bract more prominently expanded
above. The lower sheaths have a distinct purplish tinge not seen in the species.

Juncus filiformis L. Sp. Pl. 326 (1753); Hooker, Fl. Bor.-Am, 2: 190 (1839); Macoun,
Cat. Can. Pl. Pt. 4: 55 (1888); Raup in Jour. Arn. Arb. 17: 222 (1936); Hultén,
Fl, Alaska & Yuk. Pt. 3: 427 (1943).

Noted in Fl. Bor.-Am. as occurring from ‘Saskatchewan to Bear Lake .. . ,
but I have not seen specimens from the upper Mackenzie. It is occasional at
Lake Athabaska and in the Wood Buffalo Park, and it has been collected in the
central Yukon district (Hulten, 1. c.).

Juncus nodosus L. Sp. Pl. 466 (1753); Macoun, Cat. Can. Pl. Pt. 4: 63 (1888); Raup
in Jour. Arn. Arb. 17: 223 (1936); Hultén, Fl. Alaska & Vuk. Pt. 3: 429 (1943).

Juncus polycephalus Michx. var. tenuifolius Hooker, Fl. Bor.-Am. 2: 190 (1839).

Fort Smith, Russell 28 (Can.); Mackenzie R., Onion (N); damp sand bars along the
Mackenzie R. near Simpson, 9871, 9904; rocky-sandy bank of Mackenzie R. opposite Simp-
son, 9945,

Common; in fruit late in August. Noted in Fl. Bor.-Am. from “Canada, to
Bear Lake.”

Juncus alpinus Vill. var. rariflorus (Hartm.) Hartm. Scand. Fl. ed. 5: 244 (1849);
Raup in Jour. Arn. Arb. 17: 223 (1936). See Bot. Notiser 1932: 313-372 (1932),
and Rhod. 35: 233-234 (1933).

Juncus rariflorus Hartm. Scand. Fl. ed. 1: 141 (1820).

Juncus alpinus Vill. ssp. nodulosus (Wahl.) Hultén, Fl. Alaska & Yuk. Pt. 3: 417 (1943).

Juncus alpinus Vill. var. insignis Fries; Macoun, Cat. Can. Pl. Pt. 4: 62 (1888).

Juncus acutiflorus sensu Hooker, Fl. Bor.-Am. 2: 190 (1839), in part, non Ehrh.

Juncus Richardsonianus Schultes; Hooker, Fl. Bor.-Am, 2: 191 (1839), in part.

Great Slave L., Richardson (G); Mackenzie R., Onion (N); same, Meyer (G); damp
sand bar along the Mackenzie R. near Simpson, 9884.

Very common at Simpson, where it is a primary species on sand bars. In
fruit late in August. Noted in Fl. Bor.-Am. in the “Woody country to the Mack-
enzie River.”

Juncus castaneus Smith, Fl. Brit. ed. 1, 1: 383 (1800); Hooker, Fl. Bor.-Am. 2: 192
(1838); Macoun, Cat. Can. Pl. Pt. 4: 58 (1888); Raup in Jour. Arn. Arb. 17:

223 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 423 (1943); Porsild in Nat. Mus.
Can. Bull. 101: 17 (1945).

Muskeg along wood road just west of the Mackenzie R. near Simpson, 9933; Mackenzie
R. between Wrigley and Blackwater R., Crickmay 130 (Can.) ; between Blackwater R. and
Norman, Crickmay 160 (Can.) ; mud flats at mouth of glacial stream at west end of Brint-
nell L., 9264, 9703, 9718; muskeg thickets on north shore of Brintnell L., 9361.

Common to abundant; one of the primary species, with Eriophorum Scheuch-
zeri, colonizing the newly formed mud flats at the west end of Brintnell L. Col-
lected in anthesis late in June, and with maturing capsules in July and August,

Noted by Porsild (1. c.) at Little Keele R. along the Canol Rd.

9

142 SARGENTIA [6

Luzula parviflora (Ehrh.) Desv. in Jour. de Bot. 1: 144 (1808); Hultén, Fl. Alaska &
Yuk. Pt. 3: 440 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Juncus parviflorus Ehrh. Beitr. 6: 139 (1791).
Mackenzie R. between Blackwater R. and Norman, Crickmay 16 (Can.).
Noted by Porsild (1. c.) as common in Macmillan Pass along the Canol Rd.

Luzula Wahlenbergii Rupr. Beitr. z. Pflanzenk. Russ. Reich. 2: 58 (1834); Hultén, FI.
Alaska & Yuk. Pt. 3: 443 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Stony shore of Brintnell L. at Camp Pt., 9163; damp grassy place in willow-birch scrub,
upper valley of Frost Cr., alt. 4000’, 9508.

Occasional on the shore of Brintnell L. but common in the willow-birch scrub.
In flower late in June, and beginning to mature its capsules at the higher eleva-
tions about mid-July. Noted by Porsild (1. ¢.) as common in Macmillan Pass
along the Canol Rd.

Luzula spicata (L.) DC. FI. Franc. 3: 161 (1805); Hultén, Fl. Alaska & Yuk. Pt. 3:
442 (1943).

Juncus spicatus L. Sp. Pl. 330 (1753).

Small stream on northeast slope of Red Mt., alt. 4500’, 9285; shale slide rock, west slope
of Terrace Mt., alt. 5000’, 9537; shale slide rock, south slope of Red Mt., alt. 5000’-5900',
9744; shale cliffs at summit of Red Mt., alt. 5900’, north exposure, 9767,

Common, especially at about 5000’. In anthesis late in June, and maturing
capsules by mid-July.

Luzula arcuata (Wahl.) Wahl. FI. Suec. 1: 218 (1825); Hultén, Fl. Alaska & Yuk. Pt.
3: 433 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Juncus arcuatus Wahl. Fl. Lapp. 87, f. 4 (excl. 8) (1812).

Stony beach at Camp Pt., Brintnell L., 9162; rocky bed of small stream on east slope of
Red Mt., alt. 4500’, 9268; mossy bank of rocky stream bed, northeast slope of Red Mt., alt.
4300’, 9733; shale slide rock on south slope of Red Mt., alt. 5800’, 9751; east end of Mac-
millan Pass, Canol Rd., in alpine tundra, alt. 4600’, Porsild 11884 (Can.).

Common in the wetter habitats noted at Brintnell L., but only occasional on
the high slide rock. Plants collected June 20 at the lake shore (9162) were al-
ready maturing their capsules, while those at 4500’ on June 26 were still in
anthesis.

Luzula nivalis (Laestad.) Beurl. in Bot. Notiser 1853: 55 (1853); Hultén, Fl. Alaska
& Yuk. Pt. 3: 438 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Luzula campestris (L.) DC. var. nivalis Laestad. in Vet. Akad. Handl. (Stockh.), 333
(1822).
Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11798 ( Can.)
Luzula confusa Lindb. in Bot. Notiser 1855: 9 (1855); Hultén, Fl. Alaska & Yuk. Pt.
3: 435 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Shale rock-slide, west slope of Terrace Mt., alt. 6000’, 9530; dry shale slope, south side
of Red Mt., alt. 5000’, 9748; ledges of shale cliffs, summit of Red Mt., alt. 5900’, 9764, 9769;
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8264, 8265 (Can.) ; Macmillan Pass,
Canol Rd., mi. 225 E., Porsild 11219 (Can.).

Occasional at Brintnell L.; found with maturing capsules in mid-July and early
August.
Luzula campestris (L.) DC. var. alpina Gand. Agrostol. Helv. 2: 247 (1811); Raup in
Jour. Arn. Arb. 17: 223 (1936).
Evidence for the presence of L. campestris in southwestern Mackenzie is very
slender at present. I have seen but two specimens from the Mackenzie basin,
both of which I have referred to var. alpina. One is from the eastern arm of

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 143

Great Slave L. (Raup 424) and the other is from Great Bear L. (J. M. Bell,
Can.). The record in Fl. Bor.-Am. under L. campestris, “Woody and barren
country,” probably refers at least in part to this variety.

LILIACEAE

Tofieldia palustris Huds. Fl. Angl. ed. 2: 157 (1778); Hooker, Fl. Bor.-Am. 2: 179
(1839); Raup in Jour. Arn. Arb. 17: 224 (1936); Porsild in Nat. Mus. Can. Bull.
101: 17, 30 (1945).

Tofieldia borealis Wahl.; Macoun, Cat. Can. Pl. Pt. 4: 43 (1888).

Southwest and north shores of Great Slave L., Howe 91987 (Can.); northwest shore of
Great Slave L., Bedford (Can.) ; Bear Lake R., Richardson (G); mossy banks and muskeg
thickets at west end of Brintnell L., 9243; wooded lower slope of Colonel Mt., extending
above timber line, 9625; Lone Mt., Wynne-Edwards 8441 (Can.); Bolstead Cr., Canol Rd.,
mi. 111 E., Wynne-Edwards 8269 (Can.); Sekwi R., Canol Rd., mi. 174 E., Porsild 11855
(Can.).

Common in the Brintnell L. shore localities mentioned, but infrequent on the
mountains north of the lake. In flower late in June, and with maturing fruit in
the latter part of July. Noted by Porsild along the Canol Rd., also at Little
Keele R., and the Plains of Abraham.

Tofieldia coccinea Richardson, Bot. App. Frankl. Journ. 736 (1823); Hultén, FI.
Alaska & Yuk. Pt. 3: 444 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17, 30
(1945).

Tofieldia coccinea Richardson var. major Hooker, Fl. Bor.-Am. 2: 179 (1839) ; Macoun
in Cat. Can. Pl. Pt. 4: 44 (1888).

Mackenzie R., Richardson (G); Liard R. between Nahanni Butte and Simpson, Crickmay
14 (Can.); Lone Mt., Wynne-Edwards 8440 (Can.); Bolstead Cr., Canol Rd., mi. 111 E.,
Porsild 11831 (Can.) ; same place, Wynne-Edwards 8268 (Can.).

Also noted by Porsild in Macmillan Pass.

Tofieldia glutinosa (Michx.) Pers. Syn. 1: 399 (1805); Hooker, Fl. Bor.-Am. 2: 179
(1839); Macoun, Cat. Pl. Pt. 4: 44 (1888); Raup in Jour. Arn. Arb. 17: 224
(1936); Porsild in Nat. Mus. Can. Bull. 101: 30 (1945).

Narthecium glutinosum Michx. Fl. Bor.-Am. 1: 210 (1803).

“Cumberland House to Bear Lake,” Richardson (G); Liard R., between Nahanni Butte
and Simpson, Crickmay 13 (Can.) ; north peak, Nahanni Mt., Wynne-Edwards 8625 (Can.).

All of these plants appear to represent the eastern American T. glutinosa
rather than the western 7. occidentalis. See Raup in Contr. Arn. Arb. 6: 137
(1934), and Hultén in Fl. Alaska & Yuk. Pt. 3: 446 (1943), for discussions of
these species.

Zygadenus elegans Pursh, Fl. Am. Sept. 1: 241 (1814); Macoun, Cat. Can. Pl. Pt. 4:
52 (1888); Raup in Jour. Arn. Arb. 17: 224 (1936); Hultén, Fl. Alaska & Yuk.
3: 449 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17, 30, 34 (1945). See Rhod.
37: 256-258 (1935).

Zygadenus chloranthus Richardson; Hooker, Fl. Bor.-Am. 2: 177 (1839).

Clewi R., Wood Buffalo Park, Russell 7 (G); Resolution, E. A. & A. E. Preble 162
(US) ; southwest and north shores of Great Slave L., Howe 91985, 91986 (Can.); Windy
Pt., Great Slave L., Hume 102657 (Can.); northwest shore of Great Slave L., Bedford
(Can.) ; Old Fort Rae, Russell (Can.) ; “Cumberland House to Bear Lake,’ Richardson
(G); Liard R., between Nahanni Butte and Simpson, Crickmay 15, 16 (Can.); Willow
Lake R., 44 mi. above mouth, Crickmay 12 (Can.); Saline R., above Norman, Wynne-
Edwards 8571 (Can.) ; north slope of Colonel Mt., in alpine meadows and subalpine scrub,
9424, 9660; margin of slough in gravel plain near west end of Brintnell L., 9445; Lone Mt.,
W ynne-Edwards 8442 (Can.); Nahanni Mts., Preble & Cary 53, 65 (US); Bolstead Cr.,
Canol Rd., mi. 111 E., Wynne-Edwards 8266 (Can.).

144 SARGENTIA [6

Common at Brintnell L.; in flower early in July. Noted by Porsild along the
Canol Rd. also at Little Keele R.

Veratrum Eschscholtzii Gray in Ann. Lyc. Nat. Hist. N. Y. 4: 119 (1837); Hultén,
Fl. Alaska & Yuk. Pt. 3: 451 (1943); Porsild in Nat. Mus. Can. Bull. 101: 17
(1945).

Noted by Porsild at Macmillan Pass along the Canol Rd.

Allium Schoenoprasum L. var. sibiricum (L.) Hartm. Scand. Fl. ed. 4: 102 (1843);
Raup in Jour. Arn. Arb. 17: 224 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 453
(1943).

Allium sibiricum L. Mant. 562 (1771).
Allium Schoenoprasum L. sensu Hooker, Fl. Bor.-Am. 2: 185 (1839) ; Macoun, Cat.
Can. Pl. Pt. 4: 35 (1888).

Northwest shore of Great Slave L., Bedford (Can.); “Carlton House to Slave Lake,”
Richardson (G); Mackenzie R. at junction with Bell R., McConnell 27421 (Can.); Liard
R., between Nahanni Butte and Simpson, Crickmay 12 (Can.) ; rocky-sandy shore of Mack-
enzie R., opposite Simpson, 9963; in silty soil near Mackenzie R. at Simpson, Nowosad
(Can.) ; Mackenzie R., between Wrigley and Blackwater R., Crickway 131 (Can.).

Common in the Simpson locality; in fruit in the last week of August. Noted
in Fl. Bor.-Am. as occurring “Throughout the woody country to Bear Lake.”

Lloydia serotina (L.) Rchb. Fl. Germ. Excurs. 102 (1830); Porsild in Sargentia 4:
25 (1943); Hultén, Fl. Alaska & Yuk. Pt. 3: 456 (1943); Porsild in Nat. Mus.
Can. Bull. 101: 17 (1945), -

Bulbocodium serotinum L. Sp. Pl. 294 (1753).

Liard R., between Nahanni Butte and Simpson, Crickmay 11 (Can.); birch-willow
thickets on north slope of Colonel Mt., alt. about 4000’, 9385; Plains of Abraham, Canol
Rd., mi. 82 E., Porsild 11799 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards
8267 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11856 (Can.).

Occasional to common in the Colonel Mt. site which had only recently become
free of snow; not seen elsewhere at Brintnell L. In flower the first week in

July.

Smilacina stellata (L.) Desf. in Ann. Mus. Par. 9: 52 (1807); Hooker, Fl. Bor.-Am. 2:
176 (1839); Macoun, Cat. Can. Pl. Pt. 4: 30 (1888); Hultén, Fl. Alaska & Yuk.
Pt. 3: 459 (1943).

Convallaria stellata L. Sp. Pl. 316 (1753).

Scrub timber on south slope of Red Mt., north of Brintnell L., 9224.

Common in the above site, but not seen elsewhere in the Brintnell L. region.
In flower late in June. Common in the Wood Buffalo Park (Raup in Jour. Arn.
Arb. 17: 224) and collected at Great Bear L. by Porsild (Sargentia 4: 26). Our
plants have unusually large flowers and a close, fine pubescence on the under
sides of the leaves.

Smilacina trifolia (L.) Desf. in Ann. Mus. Par. 9: 52 (1807); Hooker, Fl. Bor.-Am. 2:
175 (1839); Macoun, Cat. Can. Pl. Pt. 4: 32 (1888);*Raup in Jour. Arn. Arb. 17:
224 (1936).

Convallaria trifolia L. Sp. Pl. 316 (1753).

Resolution, Kennicott (N); Liard R., between Nahanni Butte and Simpson, Crickmay
17 (Can.); near large lake between southern extremities of Franklin Mts., Crickmay 32
(Can.).

Noted in Fl. Bor.-Am. as occurring in “Swamps throughout Canada, to Bear
aee

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 145

Maianthemum canadense Desf. var. interius Iernald in Rhod. 16: 211 (1914); Raup
in Jour. Arn. Arb. 17: 225 (1936).

Maianthemum bifolia sensu Hooker, Fl. Bor.-Am. 2: 176 (1839), in part, non Roem.
Maianthemum canadense Desf. sensu Macoun, Cat. Can. Pl. Pt. 4: 32 (1888), in part.

Fort Smith, Raup 436; same, Miss E. Taylor 27537 (Can.); same, Seton & Preble
78569 (Can.); “Arctic America,” Richardson (N); Liard R., between Nahanni Butte and
Simpson, Crickmay 18 (Can.); Simpson, Miss E. Taylor (N).

Noted in Fl. Bor.-Am. as extending northward to Great Bear L.

Trillium cernuum L. var. macranthum Eames & Wieg. in Rhod. 25: 191 (1923).

Trillium cernuum L. sensu Hooker, Fl. Bor.-Am. 2: 180 (1839), in part.

Trillium erectum var. declinatum sensu Macoun, Cat. Can. Pl. Pt. 4: 49 (1888), as to

Mackenzie record, non Gray.

There is a specimen in the Gray Herbarium collected by Richardson and
labeled “Mackenzie River.” There is no indication as to where on the Mackenzie
it came from, but presumably it is the basis of the northern range for T. cernuum
given in Fl. Bor.-Am., “. . . from the Saskatchewan to Mackenzie River.” It
is the only evidence for the occurrence of any species of Trillium in the entire
Mackenzie basin.

IRIDACEAE

Sisyrinchium angustifolium Miller, Gard. Dict. ed. 8 (1768); Raup in Jour. Arn. Arb.
17: 225 (1936); Porsild in Nat. Mus. Can. Bull. 101: 30, 33 (1945).

South shore of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards &546 (Can.) ;
Providence, Preble & Cary 40 (US); Liard R., between Nahanni Butte and Simpson,
Crickmay 10 (Can.); dry southerly slope in old field at Simpson, 9124; rocky-sandy shore
of Mackenzie R. opposite Simpson, 9950; Wrigley, Miss E. Taylor 68 (Can.); Norman,
Dutilly 48 (G); “British America,” Richardson (N); north peak, Nahanni Mt., W ynne-
Edwards 8443 (Can.).

Common in the Simpson area; in flower in mid-June; late flowers occasional
in the last week in August, but most of the plants with mature capsules.

ORCHIDACEAE

Cypripedium passerinum Richardson, Bot. App. Frankl. Journ. 34 (1823); Raup in
Jour. Arn. Arb. 17: 225 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 469 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 17, 30 (1945). :

Lower Slave R., Raup 469; northwest shore of Great Slave L., Bedford (Can.); Liard
R., between Nahanni Butte and Simpson, Crickmay 20 (Can.); Wrigley, Miss E. Taylor
11 (Can., N), 17 (G); Willow Lake R., 44 mi. above mouth, Crickmay 17 (Can.) ; Bear
Rock, Norman, R. M. Anderson (N); “British America,” Richardson (N) ; willow-birch
scrub at about 4000’ on north slope of Colonel Mt., 9386; woods and thickets near west end
of Brintnell L., 9447; mossy woods near summit of Lone Mt., Wynne-Edwards 8448 (Can.) ;
north peak of Nahanni Mt., Wynne-Edwards 8449 (Can.) ; Sekwi R., Canol Rd., mi. 174 E.,
alt. 3625’, Porsild 11857 (Can.).

Common to abundant in the Brintnell L. localities; found flowering in the first
half of July.

Cypripedium parviflorum Salisb. in Trans. Linn. Soc. 1: 77, pl. 2, f. 1 (1791); Porsild in
Sargentia 4: 26 (1943), in Nat. Mus. Can. Bull. 101: 30 (1945).

Liard R., between Nahanni Butte and Simpson, Crickmay 21 (Can.) ; Bear Rock, Nor-
man, A. E. & R. T. Porsild 3381 (Can.); north peak of Nahanni Mt., Wynne-Edwards
8447 (Can.).

Apparently rare in the Mackenzie basin. In addition to the above stations
Porsild has collected it along Bear R. and at Great Bear L. It has not yet been
found in the Athabaska—Great Slave L. region.

146 SARGENTIA [6

Cypripedium guttatum Swartz in Vet. Akad. Handl. (Stockh.) 21: 251 (1800); Hooker,
Fl. Bor.-Am. 2: 205 (1839); Macoun, Cat. Can. Pl. Pt. 4: 23 (1888); Raup in
Jour. Arn. Arb. 17: 226 (1936); Porsild in Sargentia 4: 26 (1943); Hultén, FI.
Alaska & Yuk. Pt. 3: 468 (1943); Porsild in Nat. Mus. Can. Bull. 101: 30 (1945).

Great Slave L., R. Bell 23156 (Can.) ; Old Fort Rae, Russell (Can.) ; Liard R., between

Nahanni Butte and Simpson, Crickmay 19 (Can.) ; “River-between-two-mountains,” just

west of Franklin Mts., Crickmay 10 (Can.); Norman, Hume 103449 (Can.) ; mossy spruce

woods near summit of Lone Mt., Wynne-Edwards 8445 (Can.) ; north peak of Nahanni Mt.,

Wynne-Edwards 8446 (Can.); Nahanni Mts., Preble & Cary 77 (US).

Noted in Fl. Bor.-Am. as having been collected at “Fort Franklin, on the

Mackenzie River.”

Cypripedium acaule Ait. Hort. Kew. 3: 303 (1789); Macoun, Cat. Can. Pl. Pt. 4: 22
(1888); Raup in Jour. Arn. Arb. 17: 226 (1936).

Cypripedium humile Salisb.; Hooker, Fl. Bor.-Am. 2: 204 (1839).

This species has not yet been collected in southwestern Mackenzie, but it has
been found at Lake Athabaska and at Great Bear L. A Richardson specimen
in Herb. G is labeled “Fort Franklin,” and evidently is the basis for the range
given in FI. Bor.-Am., “. .. from Canada to Fort Franklin on the Mackenzie
River.”

Orchis rotundifolia Pursh, Fl. Am. Sept. 2: 588 (1814); Raup in Jour. Arn. Arb. 17:
226 (1936); Hultén, Fl, Alaska & Yuk. Pt. 3: 471 (1943); Porsild in Nat. Mus.
Can. Bull. 101: 30 (1945).

Windy Pt., Great Slave L., Hume (Can.); northwest shore of Great Slave L., Bedford
(Can.); Old Fort Rae, Russell (Can.); Liard R., between Nahanni Butte and Simpson,
Crickmay 26 (Can.); Willow Lake R., 44 mi. above mouth, Crickmay 14 (Can.); north
peak of Nahanni Mt., Wynne-Edwards 8450 (Can.) ; damp hollow in summit of Lone Mt.,
Wynne-Edwards 8451 (Can.).

Common in the Athabaska-Great Slave L. region, and collected by Porsild
(Sargentia 4: 26) at Great Bear L. and on the lower Mackenzie,

Habenaria viridis (L.) R. Br. var. bracteata (Muhl.) Gray, Man. ed. 5: 500 (1867).
Satyrium viride L. Sp. Pl. 944 (1753).
Orchis bracteata Muhl. ex Willd. Sp. Pl. 4: 34 (1805).
Habenaria viridis (L.) R. Br. var. interjecta Fernald; Raup in Jour. Arn. Arb. 17: 226
(1936) ; Porsild in Nat. Mus. Can. Bull. 101: 31 (1945).
Coeloglossum viride (L.) Hartm. ssp. bracteatum (Muhl.) Hultén, Fl. Alaska & Yuk.
Pt. 3: 472 (1943).
Liard R., between Nahanni Butte and Simpson, Crickmay 23 (Can.); north peak of
Nahanni Mt., Wynne-Edwards 8455 (Can.).

Habenaria hyperborea (L.) R. Br. in Ait. Hort. Kew. ed. 2, 5: 193 (1813); Macoun,
Cat. Can. Pl. Pt. 4: 14 (1888); Raup in Jour. Arn. Arb. 17: 226 (1936); Porsild
in Nat. Mus. Can. Bull. 101: 30 (1945).
Orchis hyperborea L. Mant. 121 (1767).
Platanthera hyperborea (L.) Lindl., Hooker, Fl. Bor.-Am, 2: 197 (1839) ; Hultén, FI.
Alaska & Yuk. Pt. 3: 479 (1943).

Lower Slave R., Raup 446; Great Slave L., Onion, Kennicott & Hardisty (N); north-
west shore of Great Slave L., Bedford (Can.); damp place in prairie on upland west of
Mackenzie R. near Simpson, 9891; “River-between-two-mountains,” just west of Franklin
Mts., Crickmay 4 (Can.) ; mossy bank on south shore of Brintnell L., 9332; muskeg on north
shore of Brintnell L., 9354; damp woods and thickets at west end of Brintnell L., 9446; north
peak of Nahanni Mt., Wynne-Edwards 9452 (Can.).

Common to abundant in the rich lowland woods about Brintnell L., where it
flowers in late June and early July. Noted in Fl. Bor.-Am., “From the Saskat-

chewan to Fort Franklin.”

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 147

Habenaria obtusata (Pursh) Richardson, Bot. App. Frankl. Journ. 750 (1823); Macoun,
Cat. Can. Pl. Pt. 4: 16 (1888); Raup in Jour. Arn. Arb. 17: 226 (1936); Porsild
in Sargentia 4: 26 (1943), in Nat. Mus. Can. Bull. 101: 30 (1943).
Orchis obtusata Pursh, Fl. Am. Sept. 2: 588 (1814).
Platanthera obtusata (Pursh) Lindl., Hooker, Fl. Bor.-Am. 2: 196 (1839) ; Hultén, FI.
Alaska & Yuk. Pt. 3: 480 (1943).

Lower Slave R., Raup 449; Resolution, Kennicott (N); northwest shore of Great Slave
L., Bedford (Can.) ; Windy Pt., Great Slave L., Hume 102658 (Can.) ; southwest and north
shores of Great Slave L., Howe 91984 (Can.) ; Liard R., between Nahanni Butte and Simp-
son, Crickmay 22 (Can.) ; damp mossy woods at Simpson, 9086, 9105; Willow Lake R., 44
mi. above mouth, Crickmay 20b (Can.) ; muskeg thickets at west end of Brintnell L., 9239;
damp hollow, summit of Lone Mt., Wynne-Edwards 8453 (Can.).

At Brintnell L. this species is occasional in the muskeg thickets at the west
end of the lake and in the mossy open forest of Picea mariana on the north slope
of Colonel Mt., where it reaches an altitude of 3200 feet or more. Found flower-
ing in the latter half of June. Porsild (1943) says it is common throughout the
Mackenzie country north to the tree line and “occasional or rare for a short
distance beyond.”

Habenaria orbiculata (Pursh) Torr. Fl. N. and Mid. U. S. 318 (1826); Porsild in Nat.
Mus. Can. Bull. 101: 30 (1945).

Orchis orbiculata Pursh, Fl. Am. Sept. 2: 588 (1814).
Platanthera orbiculata (Pursh) Lindl., Hultén, Fl. Alaska & Yuk. Pt. 3: 482 (1943).
North peak of Nahanni Mt., in spruce woods, Wynne-Edwards 8454 (Can.).

Spiranthes Romanzoffiana Cham. in Linnaea 3: 32 (1828); Raup in Jour. Arn. Arb. 17:
226 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 485 (1943).

Little Buffalo R., Russell 90 (G) ; Great Slave L., R. Bell 23155 (Can.); same, Onion,
Kennicott & Hardisty (N); Rae, Russell (Can.); Mackenzie R., between Wrigley and
Blackwater R., Crickmay 132 (Can.); near large lake between southern extremities of
Franklin Mts., Crickmay 31 (Can.).

The northern extent of S. Romanzoffiana in the Mackenzie country is not yet
known. Northern specimens upon which the range, “. . . from Lake Huron

_ . to Fort Franklin, on the Mackenzie River,” in Fl. Bor.-Am, (2: 202) un-
der S. gracilis is based may belong to this species.

Goodyera repens (L.) R. Br. var. ophioides Fernald in Rhod. 1: 6 (1899); Raup in
Jour. Arn. Arb. 17: 227 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 490 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 31 (1945).

Goodyera repens (L.) R. Br. sensu Hooker, Fl. Bor.-Am. 2: 203 (1839), in part; Porsild

in Sargentia 4: 26 (1943).

Bank of Mackenzie R. opposite Simpson, Crickmay 38 (Can.) ; in damp lichen-moss mat
in Picea mariana forest on north slope of Colonel Mt., alt. 2640’-4000’, 9657; Lone Mt.,
W ynne-Edwards 8456 (Can.).

Common in the Brintnell L. site noted above but not seen elsewhere in that
district; just beginning to flower in late July. Leaves distinctly mottled.
Hooker states in Fl. Bor.-Am. that G. repens extends “. . . from the Saskat-
chewan to Fort Franklin,” and that its leaves are rarely reticulated. Material
from the Athabaska—Great Slave L. region, however, has mottled leaves.

Listera borealis Morong in Bull. Torr. Bot. Cl. 20: 31 (1893); Raup in Jour. Arn.
Arb. 17: 227 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 486 (1943).

Fort Smith, Miss E. Taylor (N, Can., Type COLL.) ; same, Raup 460; Liard R., between
Nahanni Butte and Simpson, Crickmay 24 (Can.); Willow Lake R., 44 mi. above mouth,
Crickmay 20a (Can.) ; muskeg thicket at west end of Brintnell L., 9240.

148 SARGENTIA [6

Common at Brintnell L.; in flower late in June. Porsild (Sargentia 4: 26)
extends the range to Great Bear L. and the Mackenzie delta.

Calypso bulbosa (L.) Oakes jin Thompson Hist. Vermont Pt. 1: 200 (1842); Raup in
Jour. Arn, Arb. 17: 227 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 494 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 31 (1945).

Cypripedium bulbosum L. Sp. Pl. 951 (1753).
Calypso borealis Salisb.; Hooker, Fl. Bor.-Am. 2: 195 (1839) ; Macoun, Cat. Can. Pl. Pt.
4: 3 (1888).

Resolution, Kennicott (N) ; “Between Forts Simpson and Chipewyan,” Richardson (Can.) ;
Mackenzie R., Kennicott (N); Liard R., between Nahanni Butte and Simpson, Crickmay 25
(Can.) ; Simpson, Kennicott ? (N); same, in rich woods, 9730; north peak of Nahanni Mt.,
Wynne-Edwards 8444 (Can.).

In flower about mid-June. According to Fl. Bor.-Am. it extends northward
“.. to Bear Lake River.”

Corallorrhiza trifida Chat. Spec. Inaug. Corallorrhiza, 8 (1760); Raup in Jour. Arn.
Arb. 17: 227 (1936); Porsild in Sargentia 4: 26 (1943); Hultén, Fl. Alaska &
Yuk. Pt. 3: 492 (1943).

Fort Smith, Miss E. Taylor (G, Can.) ; Resolution, Kennicott (N); mossy woods on
south shore of Brintnell L., 9355; rich woods on lower north slope of Colonel Mt., 9425;
muskeg thickets near west end of Brintnell L., 9458.

Occasional ; in flower early in July. According to Porsild (1. ¢.) this spe-
cies extends throughout the Mackenzie district northward to the tree line or a
short distance beyond.

SALICACEAE

Populus tremuloides Michx. Fl. Bor.-Am. 2: 243 (1803); Hooker, Fl. Bor.-Am. 2:
154 (1839); Macoun, Cat. Can. Pl. Pt. 3: 456 (1886); Raup in Jour. Arn. Arb.
17: 227 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 496 (1943).

Fort Smith, Raup 489; Little Buffalo R., Russell 23 (G); Mackenzie R., Kennicott (N);
spruce-birch forest on lower south slope of Red Mt., tree 40’ high, 5” diam., 9313.

Common to abundant throughout the Mackenzie basin forests. It owes much
of its development to the fires which have burned the timber repeatedly. At
Brintnell Lake it is most abundant on the slopes which gave evidence of having
been burned over in comparatively recent time. It was observed up to about
4000’ on the mountain slopes. Aments had all fallen by mid-June. All the
trembling aspens of this region appear to represent the doubtfully distinct var.
aurea, the leaves of which turn yellow in autumn. This color is a striking and
pleasing feature of the Mackenzie and Yukon landscapes in September.

Populus Tacamahacca Mill. Gard. Dict. ed. 8 (1788); Raup in Jour. Arn. Arb. 17: 228
(1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 495 (1943); Porsild in Nat. Mus. Can.
‘ Bull. 101: 17 (1945).
Populus balsamifera sensu Hooker, Fl. Bor.-Am, 2: 153 (1839); Macoun, Cat. Can. Pl.
Pt. 3: 456 (1886) ; non L,

Fort Smith, Raup 497; Resolution, J. W. Tyrrell 23132 (Can.); northwest shore of
Great Slave L., Bedford (Can.); north of the height of land between Great Slave and
Great Bear Lakes, Bedford (Can.) ; Mackenzie R., Onion (N); Simpson, Preble 304 (US);
flood plain forest along Mackenzie R. near Simpson, 9068; woods along Frost Cr., just
back of camp at Brintnell L., 9315.

Common to abundant on gravelly flats about the west end of Brintnell L., where
it often occurs in nearly pure stands of open forest; also common along stony
mountain streams at low elevations. In the old burn on the south slope of Red

1947] RAUP, THE BOTANY. OF SOUTHWESTERN MACKENZIE 149

Mt. it reaches an elevation of 3000’-4000’. At Simpson it is a primary part of
the flood plain forest, but also occurs on the uplands. Fruits mature at Brintnell
L. in late June, and at Simpson in the second week of June. Preble states that
this tree extends along the entire length of the Mackenzie R., and a note in Fi.
Bor.-Am. says that the Liard R. was so named (“Riviere aux Liards”’) because
of the abundance of poplars on its banks. Porsild states that it was noted as
common bordering creeks and rivers along the Canol Road.

Salix lasiandra Benth. Pl. Hartweg. 335 (1857); Raup in Jour, Arn. Arb. 17: 228
(1936). See Jour. Arn. Arb. 1: 14-20 (1919).

Salix lucida sensu Hooker, Fl. Bor.-Am. 2: 148 (1839), in part; Macoun, Cat. Can. .Pi.
Pt. 3: 450 (1886), in part; non Willd.

Providence, Dutilly (G).

The Providence specimen appears to match material collected in the Athabaska
Lake-Wood Buffalo Park region that I have referred to the typical form. Its
northern limits are yet to be determined. Representatives of the lasiandra com-
plex collected at Simpson I am proposing as the following variety.

Salix lasiandra Benth. var. recomponens var. nov.

A forma typica ramulis pubescentibus et foliis subtus viridibus differt.

Sandbar along Mackenzie R. near Simpson, shrubs 6’-8' high, 9073, m., 9076, f. (Type).

Occasional in the Simpson area; collected with flowers and immature capsules
June 11.

In the absence of a bloom on the lower surfaces of the leaves these plants sug-
gest S. lasiandra var. caudata (Nutt.) Sudw., while the pubescence on the twigs
relates them to var. lancifolia (And.) Bebb (See Schneider in Jour. Arn, Arb.
1: 14-20, 1919, and Ball in Can. Field Nat. 40: 145-150, 1926, for discussions of
this group). In the shape of the leaves, which are lanceolate-acuminate rather than
attenuate at the apex, they resemble the typical form or var. lancifolia, Variety
recomponens, therefore, appears to be formed by a recombination of the charac-
ters by which the above three forms of the species are commonly distinguished.
Ball (1. c.), following Heller (Muhlenb. 2: 186, 1906), maintains S. caudata as
a distinct species, but our plants suggest that it is only a phase in the S. lasiandra
complex, equivalent in taxonomic value to other phases which are recognized only
as varieties.

Hooker, in Fl. Bor.-Am., gives the range of S. lucida . as far north as
Fort Franklin on the Mackenzie River.” This record is probably based upon all
the S. lucida-like willows in the Mackenzie basin (S. serissima, S. lasiandra vars. ).

«é

Salix interior Rowlee var. pedicellata (Anders.) Ball in Can. Field Nat. 40: 175 (1926);
Raup in Jour. Arn. Arb. 17: 229 (1936).

Salix interior Rowlee, sensu Ball in Can. Field Nat. 40: 174 (1926) ; Hultén, Fl. Alaska
& Yuk. Pt. 3: 504 (1943) ; at least in part.

Salix longifolia sensu Hooker, Fl. Bor.-Am. 2: 149 (1839) ; Macoun, Cat. Can. Pl. Pt.
3: 450 (1886) ; non Muhl.

Salix fluviatilis sensu Preble in N. Am. Fauna 27: 521 (1908), non Nutt.

Lower Slave R. near Grande Detour, Harper 99019, 99025 (Can.); Slave R., Kennicott
(N); between Forts Simpson and Chipewyan, Richardson (Can.) ; sand bar along Mack-
enzie R., near Simpson, shrubs 4’ high, 9069, 9070; mouth of north Nahanni R., Preble 362
(US):

Preble states that this species extends throughout the course of the Mackenzie
R. It is common at Simpson, flowering in late May and early June.

150 SARGENTIA [6

Occasional specimens with the broader leaves of the typical form are seen in
the lower Athabaska and Slave R. districts, and the Preble specimen from the
North Nahanni R. has been so determined by Ball (1. ¢.). The broad-leaved
form, however, is not well defined in the north, and I prefer to consider the whole
population as representing var. pedicellata.

Salix reticulata L. Sp. Pl. 1018 (1753); Hooker, Fl. Bor.-Am. 2: 151 (1839); Macoun,
Cat. Can. Pl. Pt. 3: 454 (1886); Preble in N. Am. Fauna 27: 523 (1908); Raup
in Jour. Arn. Arb. 17: 229 (1935); Hultén, Fl. Alaska & Yuk. Pt. 3: 505 (1943);
Porsild in Nat. Mus. Can. Bull, 101: 18, 31 (1945).

Liard R., between Nahanni Butte and Simpson, Crickmay 30 (Can.) ; Smith Cr., 4 mi.
above mouth, Crickmay 26 (Can.) ; mossy muskeg thickets at the western end of Brintnell
L., 9253; east slope of Red Mt., alt. 4500’, 9301; north peak of Nahanni Mt., Wynne-
Edwards 8462 (Can.) ; Lone Mt., W ynne-Edwards 8463 (Can.) ; alpine bog at Bolstead Cr
alt. 5500’, Canol Rd., mi. 112 E., Wynne-Edwards 8278, 8279 (Can.).

Occasional to common in mossy thickets and on rock-slides near the level of
Brintnell L., and one of the commonest of the trailing willows at higher eleva-
tions. It becomes a primary species on the alpine north slopes of Colonel Mt.,
associated in the turf with Dryas integrifolia, Cassiope tetragona, Carex scir-
poidea, Festuca altaica, Vaccinium uliginosum, and Betula glandulosa. Found
flowering late in June. Noted by Porsild (1. ¢.) as common in Macmillan Pass

along the Canol Rd.

Salix polaris Wahl. ssp. pseudopolaris (Flod.) Hultén, Fl. Alaska & Yuk. Pt. 3:50
(1943).
Salix pseudopolaris Flod. in Vet. Akad. Ark. f. Bot. 20A, No. 6: 8 (1926); Porsild in
Nat. Mus. Can. Bull. 101: 18 (1945),

Alpine slope at Bolstead Cr., Canol Rd., alt. 5500’, Wynne-Edwards 8275, 8276, 8277
(Can),

Salix phlebophylla Anders. Ofvers. Vet. Akad. Forhandl. (Stockh.) 15: 131 (1858);
Hultén, Fl. Alaska & Yuk. Pt. 3: 511 (1943); Porsild in Nat. Mus. Can. Bull.
101: 18 (1945).

In limestone rubble, Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11800 (Can.) ;
Bolstead Cr., Canol Rd., Wynne-Edwards 8274 (Can.) ; Sekwi R., Canol Rd., mi. 174 by,
alt. 3625’, Porsild 11861 (Can.).

Porsild states that this species is common everywhere on limestone along the
Canol Rd.

Salix arctica Pall. Fl. Ross. 1, 2: 86 (1788); Raup in Jour. Arn. Arb. 17: 230 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 3: 513 (1943); Porsild in Nat. Mus. Can. Bull.
101: 18 (1945).

Alpine meadows and rock-slides, Colonel Mt., 9410; trailing over rocks along small brook,
west slope of Terrace Mt., alt. 5000’, 9504; along small stream, east slope of Red Mt., alt.
4500’, 9279, 9280; rocky east slopes of Red Mt., alt. 5000’-5800', 9281, 9282, 9760; Plains of
Abraham, Canol Rd., mi. 82 E., Porsild 11801 (Can.) ; Bolstead Cr., Canol Rd., mi. 112 E,,
alt. 4500’-5500’, Wynne-Edwards 8272 (Can.).

Common to abundant on high alpine slopes throughout the Brintnell L. and
Canol Rd. areas ; the commonest of the trailing willows. In flower the last week
of June, and with maturing capsules early in August.

Salix crassijulis Trautv. in Mem. Soc. Nat. Mosc. 2: 308 (1832); Hultén, Fl. Alaska
& Yuk. Pt. 3: 515 (1943).

Alpine meadows and rock-slides, north slope of Colonel Mt., 9410-A,

Growing with S. arctica, but easily distinguished from the latter by its retuse
leaves. Found with mature capsules Aug. 5. It is somewhat doubtful whether

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 151

S. crassijulis is specifically distinct from S. arctica. Hultén (1. c.) has an ex-
cellent discussion of this problem, and concludes that the whole S. arctica com-
plex, including S. crassijulis and others, is one species with several geographic
races. Apparently he maintains S. crassijulis as a species only tentatively until
further studies of the group as a whole, with its intricate nomenclature, have been
made. With this view I am inclined to agree.

Salix arctophila Cockerell ex Heller, Cat. N. Am. Pl. 89 (1910); Raup in Jour. Arn.
Arb. 17: 230 (1936); Porsild in Sargentia 4: 27 (1943). See Schneider in Bot.
Gaz. 66: 140 (1918) and 67: 57 (1919).

Salix Uva-ursi sensu Hooker, Fl. Bor.-Am. 2: 152 (1839), in part?; Macoun, Cat. Can.
Pl. Pt. 3: 455 (1886), in part?; non Pursh.

“Fort Franklin, Mackenzie River,” Richardson (N) (no. 93, H. B. & T.)

Porsild (1. c.) states that this species extends across the tundra country from
Hudson Bay to the Mackenzie delta. I have collected it on the eastern arm of
Great Slave L., however, well within the forested country, and the Richardson
specimen also indicates that it is not limited to the tundra. It probably occurs
in exposed situations in southwestern Mackenzie. Schneider (1. c.) suggests
that the Hooker record for S. Uva-ursi in the northwest, “Woody country be-
tween lat. 54° and 64°,” possibly belongs to S. arctophila.

Salix brachycarpa Nutt., N. Am. Sylva 1: 69 (1843); Porsild in Nat. Mus. Can. Bull.
101: 31 (1945),

Mackenzie R., between Blackwater R. and Norman, Crickmay 162 (Can.); Lone Mt.,
Wynne-Edwards 8459 (Can.).

Salix brachycarpa is occasional in the Wood Buffalo Park and common in the
northern Rocky Mts., usually represented by var. antimima (Schn.) Raup. The
Crickmay specimen cited above is a fragment, uncertainly determined, but the
plant from Lone Mt. is clearly this species.

Salix niphoclada Rydb. in Bull. N. Y. Bot. Gard. 1: 272 (1899); Preble in N. Am.
Fauna 27: 523 (1908); Porsild in Sargentia 4: 28 (1943); Hultén, Fl. Alaska &
Yuk. Pt 3: 530 (1949);

The status of S. niphoclada in the flora of southwestern Mackenzie is doubtful.
In fact, like so many other phases of the S. g/auca-complex, its status as a species
is in doubt. It was described from material collected along the Mackenzie about
30 miles north of the Arctic Circle, and the type (Miss E. Taylor 60) is in the
National Herbarium of Canada. Porsild, who has made frequent comparisons of
northern material with the type, states that the species “appears to be fairly com-
mon in the Palaeozoic country of the northwestern Mackenzie District.’”” He has
collected it at Great Bear L. and on Mt. Charles along the Bear R. Hultén has
tentatively admitted the species to the flora of Alaska and Yukon, suggesting that
it may be a hybrid complex. He gives it a range “from Alaska to Hudson Bay.”
Some specimens collected by Preble at Resolution and Fort Good Hope were
originally determined as S. niphoclada by Rydberg, but most of them have been
otherwise disposed of by later authors (Schneider in Bot. Gaz. 67: 60-62, 1919;
Ball in Univ. Calif. Publ. Bot. 17: 416-417, 1934; Raup in Jour. Arn. Arb. 17:
233, 1936). I have seen one of these specimens in Herb. US, Resolution, Preble
193, which appears to have stood the tests and remains to suggest that S. nipho-
clada extends up the Mackenzie to Great Slave L.

In terms of apparent major trends in the S. glauca group of willows, S. nipho-
clada seems to be intermediate between the typical S. glauca-like plants which are

152 SARGENTIA [6

so common in Mackenzie, the northern Rockies, and Alaska, and the short-cat-
kined S. brachycarpa of the central Cordillera. Its catkins are not shortened as
in S. brachycarpa but it has a habit of growth somewhat like the latter and has
very short petioles similar to those of S. brachycarpa. It is probably but one of
several partial segregates within the S. glauca group, similar in value to S. glauca
vars. acutifolia, Aliceae, Muriei, perstipula, etc. It is probably best for the pres-
ent to maintain the current concept of S. niphoclada, however, until more material
from northern Mackenzie and Yukon is available. In this I agree with Hultén’s
treatment.

Salix glauca L. Sp. Pl. 1019 (1753); Raup in Jour. Arn. Arb. 17: 232 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 3: 525 (1943).

Salix glauca L. var. glabrescens (Anders.) Schn. in Bot. Gaz. 66: 329 (1918), in part.

Salix glaucops Anders. var. glabrescens Anders.; Preble in N. Am. Fauna 27: 522 (1908),
in part.

Northwest shore of Great Slave L. ?, Bedford (Can.) ; Mackenzie R., Richardson (Hb.
Barr.) ; stony bank of the Mackenzie R. a few miles below Providence, shrubs 4’ high,
9976, 9978; “Fort Franklin, Mackenzie R.,” Richardson (N).

Occasional to common along the upper Mackenzie; found with maturing cap-
sules Sept. 9. The leaves turn yellow on the upper surfaces after frost, while the
lower surfaces remain quite glaucous. This material is similar to a large series
of specimens from the Athabaska—Great Slave L. region that I have regarded as
typical S. glauca. There is a great deal of variation in leaf form and pubescence,
and in the form and size of the shrubs, but the whole series seems well within the
range of variation of the Old World species as it is represented in our herbaria.
It is of interest to find that Hultén now bears out the opinion, expressed long ago
by Coville, that many Alaskan representatives of S. glauca are inseparable from
those of northern Europe.

Salix glauca L. var. acutifolia (Anders.) Schneider in Bot. Gaz. 66: 327 (1918); Raup
in Jour. Arn. Arb. 17: 232 (1936). See also Schneider in Bot. Gaz. 67: 60 (1919),
and Raup in Rhod. 33: 241-244 (1931).

Salix villosa Hooker var. acutifolia Hooker, Fl. Bor.-Am. 2: 144 (1838).
Salix glauca L. var. villosa Anders.; Macoun, Cat. Can. Pl. Pt. 3: 448 (1886).
Salix Seemannii Rydb.; Preble in N. Am. Fauna 27: 522 (1908), in part.

Salix desertorum Richardson, Bot. App. Frankl. Jour. 753 (1823) ; Hooker, Fl. Bor.-Am.
2: 151 (1839).

Mackenzie R., Richardson (N); “Fort Franklin, Mackenzie R.,” Richardson (N, Hb.
Barr.) ; Rae, E. A. & A. E. Preble 138 (US); Providence, Preble & Cary 42 (US); rocky
slope near summit of Red Mt., alt. 5800’, 9759; high alpine crevices and rock-slides on the
north slope of Colonel Mt., alt. 5000’-6000', 9409, 9421, 9627.

Common in the Brintnell L. district, usually growing as a depressed shrub,
sometimes with an elaborate root system penetrating the unstable slide rock ;
capsules maturing in August. The validity of var. acutifolia, like that of other
S. glauca segregates, is open to question. I have preferred to keep it up, however,
in order to give expression to a phase of the species which is very common in the
Mackenzie basin. These plants have leaves somewhat smaller and more nar-
rowly elliptic or lanceolate than the typical form. They are usually rather thinly
hairy, with slightly and irregularly denticulate margins. In the Athabaska—
Great Slave L. district var. acutifolia has its greatest development in the pre-
Cambrian country, and in the Brintnell L. region of the Mackenzie Mts. it largely
replaces the typical form. In Alaska Hultén has not recognized it as a valid

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 153

entity. He considers that most of the Alaska and Yukon specimens named var.
acutifolia by Schneider are hybrids of S. glauca and S. pseudomonticola. The
latter species has not been found in that part of the Mackenzie basin where var.
acutifolia is most abundant, and in the Wood Buffalo Park, where S. pseudo-
monticola is occasionally seen, the more typical S. glauca is the commonest form
in its group.

A few comments should be inserted here concerning Salix desertorum. Of all
the S. glauca segregates in the Northwest this was the first to be described
(Richardson, Bot. App. Frankl. Journ. 753, 1823), so that the name might have
claims to priority. In spite of the fact that Richardson gave a fairly good diag-
nostic description of his plant, and that adequate co-type material is in American
herbaria, the species has remained an enigma. Hultén has recently proposed
that Andersson’s designation of S. desertorum as a subspecies of S. glauca (see
Anders. in Ofv. Vet. Akad. Férh. 15: 129, 1858) be revived (Fl. Alaska & Yuk.
527). He looks upon this as “the arctic American S. glauca representative
distributed from Alaska to Newfoundland.” For plants having the lower leaf-
surfaces densely villous he proposes a new variety sericea of ssp. desertorum
Cc);

Hultén does not make clear just what forms he proposes to cover with ssp.
desertorum in the vast subarctic region east of Yukon. After studying the wil-
lows of the Hudson Bay Region, Labrador Peninsula, and northern Newfound-
land I concluded that S. desertorum as represented by our type material was not
found there (Sargentia 4: 102-110). All the eastern specimens that I have seen
so labeled are better placed in S. brachycarpa, S. cordifolia vars., or in some of the
other segregates of S. glauca. In the Mackenzie basin I have seen but one col-
lection that I could match with Richardson’s type. This is my no. 534, from
Taltheilei Narrows in the eastern arm of Great Slave Lake. In fact, an out-
standing feature of the situation in that region is that, in spite of a great deal of
collecting since Richardson's time, his plant has appeared so rarely.

In 1931 I published a brief discussion of the S. glauca forms in the Athabaska-—
Great Slave L. region (Rhod. 33: 241-244, pl. 218) and suggested that Richard-
son’s type of S. desertorum was only a dwaried form of S. glauca, growing in an
exposed rocky situation where it was subject to difficult soil and climatic condi-
tions. My specimens from Great Slave L., which appear to match it so well,
came from just such a spot. Hultén, having only my plate before him, suggested
that the Slave L. plants represented his var. sericea, with the undersides of the
leaves densely villous. This is not the case, however, for the leaves are only
thinly villous to glabrescent beneath. In view of the fact that, in spite of much
further collecting in the North, more authentic material of Richardson’s-plant has
not turned up, I am inclined to hold to my earlier conclusion expressed above, and
to consider S. desertorum of little or no taxonomic significance. There is some
question as to whether it is identical with typical S. glauca or with var. acutifolia.
I am placing it tentatively with the latter.

Salix glauca L. var. Aliceae Ball in Univ. Calif. Publ. Bot. 17: 416, pl. 74 (1934).

Salix glauca L. var. acutifolia (Hook.) Schn, f. poliophylla Schn. in Bot. Gaz. 67: 61
(1919),

Salix glauca L. var. poliophylla (Schn.) Raup in Jour. Arn. Arb. 17: 233 (1936).

Resolution, E. 4. & A. E. Preble 141 (US); same, E. A. Preble 194 (US); Rae, E. A.
& A. E. Preble 139 (US, type of f. poliophylia).

154 SARGENTIA [6

I have not seen the type of Ball’s var. Aliceae, but judging from his description
and cited specimens it is the same as var. poliophylla. Ball mentions in his
discussion the type number of the latter, stating that it would be “cited below,”
but in the citation of specimens the number does not appear. However, I as-
sume from the context that Ball considered his var. Aliceae synonymous with S.
glauca var. acutifolia f. poliophylla of Schneider.

Salix glauca L. var. perstipula var. nov. Fic. 13.

A varietatibus aliis S. glaucae stipulis conspicuis lineari-lanceolatis acuminatis
differt ; stipulis plerumque (interdum ad 5 annos) persistentibus, margine et ventro
basim versus glandulosis, cito revolutis, saepe arcuato-patentibus, interdum quam
petiolis longioribus.

Muskeg on south shore of Brintnell L., shrub 5’ high, 9327 (f.) (Type); along Frost Cr.,
just above Camp Pt., Brintnell L., shrub 6’ high, 9316; open black spruce woods; north slope
of Colonel Mt., alt. about 3000’, shrubs 5’-6’ high, 9632, 9633; upper part of scrub timber,
north slope of Colonel Mt., depressed shrub, 9412; rock-slide, north slope of Colonel Mt., alt.
5000’, shrub 3’ high, 9630; willow-birch scrub in upper valley of Frost Cr., alt. about 4000’,
shrubs up to 6’ high, 9509; south slope of Red Mt., alt. about 4000’, shrubs 6’ high, 9227, 9228;
northeast slope of Red Mt., alt. 5500’, shrub 1’ high, 9300; Lone Mt., Wynne-Edwards 8460
(Can.) ; east end of Macmillan Pass, Porsild 11886 (Can.).

1cm

— iM
VE CROOK

Fic. 13. Salix glauca var. perstipula Raup, var. nov.

Common to abundant about Brintnell L. It is probably the commonest wil-
low in the area—certainly the most conspicuous—and it is a primary species in
the open woods of the lower mountain slopes and in the willow-birch scrub in
the upper valley of Frost Cr. Although found at higher levels, it appears to be
replaced there largely by S. glauca var. acutifolia. Found with flowers and im-
mature capsules late in June, and with bursting capsules about mid-August.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 155

This willow has the habit of growth, glaucous leaves, foliolate peduncles,
woolly capsules, and flavescent bracts of the bewildering complex which
is commonly included in S. glauca; but it is at once distinguished from the
other forms in this group by its conspicuous, lance-attenuate, persistent stip-
ules. Within the S. glauca complex it appears nearest to var. acutifolia (Anders. )
Schn. Schneider, in his rather carefully drawn description of this variety (Bot.
Gaz. 66: 327-329, 1918), states: “Stipulae ut videtur semper evolutae, in ramulis
vegetis maximae, pl. m. lineari- ad semi-cordato-lanceolatae, rarius semi-ovato-
rotundae, satis distincte glanduloso-denticulatae, ut folia pilosa, petiolo duplo
breviores ad 14 vel fere duplo longiores, maximae surculorum ad 3:1 cm. mag-
nae.” Nothing is said here about the persistence of the stipules, but Schneider
separated the entire S. glauca group in his key (Jour. Arn. Arb. 3: 105, 1921)
under a heading “Stipulae numquam persistentes, saepe nullae.” After a care-
ful examination of the numerous specimens of S. glauca and its relatives in the
herbaria at Harvard, I have been unable to find any with persistent stipules of
this nature. While Schneider included a linear-lanceolate form in his descrip-
tion, it is but rarely seen and does not show the acuminate character when dry.
Polunin has recently designated a var. stenolepis (Nat. Mus. Can. Bull. 92: 63,
1940) which is distinguished by these narrowly lanceolate stipules, but he says
nothing about their being persistent.

It is of interest that at least three other willows in the Brintnell L. area are
distinctive for their persistent stipules: S. pulchra, S. Richardsonti, and a new
variety of S. Barrattiana to be described below. The character is not a common
one among American willows and seems to be localized in the extreme north-
western part of the continent, perhaps illustrating a degree of convergence in the
speciation of the region.

Salix mackenzieana Barr. apud Anders. in Ofr. Svensk. Vet. Akad. Forh. 15: 125
(1858); Schneider in Jour. Arn. Arb. 2: 190 (1921); Raup in Jour. Arn. Arb. 17:
233 (1936).

Salix cordata Muhl. var. mackensieana Hooker, Fl. Bor.-Am. 2: 149 (1839); Macoun,
Cat. Can, Pl. Pt. 3: 447 (1886).

Island in Slave R. below Grande Detour, Harper 99026 (Can.); Great Slave L. ?,
Richardson (G); Mackenzie R., Richardson (A, G, Cotype) ; sand bar along Mackenzie R.
near Simpson, shrubs about 5’ high, 9071, 9072, 9074, 9075.

Occasional to common along the Mackenzie; found with maturing capsules
June 11. One of the Simpson plants is notable for having its capsules strongly
tinged with red (no. 9071). More material and field observations are necessary
before it can be determined whether this character deserves taxonomic recogni-
tion. In most descriptions the color of the capsules is not mentioned, though Ball
states (in Coult. & Nels. New Man. FI. Rky. Mts. 133) that they are greenish.
On other specimens collected at Simpson, as well as elsewhere in the Mackenzie
basin, the capsules are greenish ; and those on the type material, of which there is
a fragment in the Arnold Arboretum herbarium, do not show the red color. The
type was collected along the Mackenzie River by Richardson in June, 1826. It
is impossible to say whether the red would persist in very old specimens.

It is probable that at least a part of the material cited in Fl. Bor.-Am. under
S. rigida (2: 149), “Between Fort Franklin and Cumberland House,’ belongs
here, though it may be referable to S. lutea.

156 SARGENTIA [6

Salix lutea Nutt. N. Am. Sylva 1: 63, #. 19 (1843); Raup in Jour. Arn. Arb. 17: 234
(1936).

Lower Slave R., Raup 557.

Salix lutea is common in the Athabaska—Peace delta and along the Slave R.
Its northern extent on the upper Mackenzie is yet unknown.

Salix myrtillifolia Anders. in Ofv. Svensk. Vet. Akad. Foérh. 15: 132 (1858); Preble in
N. Am. Faun. 27: 521 (1908); Schneider in Jour. Arn. Arb. 2: 193 (1921); Raup
in Jour. Arn. Arb. 17: 235 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 537 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 18, 31 (1945).

Salix myrsinites sensu Hooker, Fl. Bor.-Am. 2: 151 (1839), non L.

Salix novae-angliae Anders. var. pseudocordata sensu Macoun, Cat. Can. Pl. Pt. 3: 452
(1886), in part, non Anders.

Salix pseudo-myrsinites sensu Preble in N. Am. Fauna 27: 521 (1908), non Anders.

Resolution, Preble 199, 200 (US); Providence, Preble & Cary. 43, 45 (US): “Fort
Franklin, Mackenzie River,” Richardson (G, N); Smith Cr., 4 mi, above mouth, Crickmay
27 (Can.) ; muskeg thickets near west end of Brintnell L., trailing in wet moss, with branches
sometimes 2’ high, 9254, 9255; Nahanni Mts., Preble & Cary 63 (US); same, Preble 311
(US) ; summit of Lone Mt., Wynne-Edwards 8461 (Can.) ; Sekwi R., Canol Rd., mi. 174 E.,
Porsild 11860 (Can.).

Common in muskegs throughout the region; capsules maturing late in June.
Our no. 9254 is of female specimens in which many of the aments are unusually
elongated and loosely flowered. The fact that some individuals are quite typi-
cal suggests that the elongated aments are abnormalities.

Preble (see N, Am. Fauna 27: 520) collected some specimens on the slopes of
the Nahanni Mts. on June 6, 1904, which were determined by Rydberg as his
Salix padophylla (Bull. Torr. Bot. Cl. 28: 272, 499, 1901). I have not seen
these specimens and consequently can do no more than suggest what they are.
Salix padophylla was originally thought by Rydberg to be in the Sect. Cordatae,
and he compared it with S. mackensieana and S. cordata. Schneider related it
rather to S. adenophylla and S. commutata. Ball has recently made it a variety of
S. pseudomonticola in the Commutatae (Jour. Wash. Acad. Sci. 28: 450, 1938).
It is difficult to estimate Rydberg’s determination of the Nahanni specimens, there-
fore, since it might have been in either of the above sections. Salix monticola is
not known in the north, but S. pseudomonticola has been found in the Wood Buf-
falo Park and in Alaska. Preble’s plants may have been the latter species.

Salix Barclayi Anders. in Ofv. Vet. Akad. Férh. 15: 125 (1858); Hultén, Fl. Alaska &
Yuk. Pt. 3: 533 (1943); Porsild in Nat. Mus. Can. Bull. 101: 18 (1945).

Muskeg thickets at the western end of Brintnell L., shrubs 2’-4’ high, 9258, 9259, 9260;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11224 (Can.).

Noted as occasional at Brintnell L., but common in Macmillan Pass; found
with maturing capsules June 23.

Salix commutata Bebb in Bot. Gaz. 13: 110 (1888); Hultén, Fl. Alaska & Yuk. Pt. 3:
536 (1943).

Marshy muskeg thickets about the western end of Brintnell L., shrubs 2’-3’ high, 9256,
9257, 9499, 9500; gravelly shore of Brintnell L. at Camp Pt., shrub 3’ high, 9307; mud flats
along stream near west end of Brintnell L., shrub 4’ high, 9692; in willow scrub in upper
valley of Frost Cr., alt. 4500’, shrub 3’ high, 9503; west slope of Terrace Mt., alt. 4800’,
depressed shrubs, 9501, 9505.

Abundant on the marshy flats at the west end of the lake, where it is a pri-
mary species in the first thickets which form on the young deposits of mud
brought in by the glacial stream. It is seen sporadically on gravelly lake shores,

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 157

and only occasionally at higher levels on the mountains. About the lake it was
in flower late in June and began to mature its capsules about mid-July. Some
of those collected at 4800’ were in flower about mid-July.

The capsules on nearly all of our plants are strongly tinged with red. When
the fruit is well-formed this is a conspicuous feature, visible from a distance, al-
though the color of the capsules is not mentioned in currently used manuals and
monographs. In the original description of the species, however, Bebb states that
they are “greenish or rufescent,” and he mentions the rufescent character as one
of the features of the species which relates it to S. cordata.

Bebb also described (1. ¢.) a var. denudata, having “young leaves smooth or
nearly so,” and this was kept up provisionally by Schneider (Jour. Arn. Arb. 1:
160-162, 1920). It came from the same locality as the type (Eagle Creek Mts.,
Oregon, Cusick). Four of our numbers may be assigned to this smooth-leaved
form : 9257, 9499, 9500, 9503 ; and it will be noted that they occur with the typical
plants or near them both at lake-level and higher in the mountains. These facts
suggest that var. denudata is without habitat or geographic segregation, and may
not be worthy of the rank of variety in the geographic sense.

Salix Barrattiana Hooker, Fl. Bor.-Am. 2: 146, f. 181 (1839); Hultén, Fl. Alaska &
Yuk. Pt. 3: 542 (1943); Porsild in Nat. Mus. Can. Bull. 101: 18 (1945)

Upper part of willow-birch scrub above timber line on north slope of Colonel Mt., alt.
4000’-5000’, depressed shrubs, 9370, 9371, 9422; Macmillan Pass, Canol Rd., mi. 225 E.,
Porsild 11223 (Can.).

Common on the north slope of Colonel Mt. at Brintnell L., but not seen else-
where in that area. Porsild notes it as common in alpine passes and valley bot-
toms along the Canol Rd. It is a characteristic feature of alpine snow-fed water-
courses. Collected with flowers and maturing capsules July 5.

Salix Barrattiana Hooker var. marcescens var. nov.

A forma typica ramulis dense hirsutis et stipulis lineari- vel semicordato-
lanceolatis persistentibus 1-2.5 cm. longis margine conspicue glanduloso-denti-
culatis differt.

Along small snow-fed stream on the north slope of Colonel Mt., alt. about 5000’, depressed
shrub, 9374 (Type); Bolstead Cr., Canol Rd., mi. 111 E., alt. 5500’, Wynne-Edwards 8273
(Can.) ; Sekwi R., Canol Rd., mi. 174 E., alt. 3625’, Porsild 11841, 11858 (Can.).

Common at the Colonel Mt. site but not seen elsewhere in the Brintnell L.
area; found with maturing capsules July 5; male plants not found.

The exact relationship of this form is not clear, although it appears to belong
in the Section Chrysantheae as interpreted by Schneider (Jour. Arn. Arb, 1: 211,
1920). Within this group it could be regarded as intermediate between S, Bar-
rattiana and S. Richardsonii, two northwestern species that have long been re-
garded as distinct. It has persistent stipules that closely resemble in form, habit,
and degree of persistence those of S. Richardsonti, but its leaves are narrower
(elliptic to narrowly ovate) and persistently hairy or but tardily glabrate. In S.
Richardsonii the leaves are soon glabrous and are commonly broadly ovate and
sometimes almost rounded at the base. Furthermore the leaves of S. Richard-
sonii are usually glandular-dentate toward the base, while in our plants they are
nearly or quite entire. Typical S. Richardsonii has glabrous capsules, while in
our plants the capsules are permanently pubescent.

On the other hand, our material can be separated from S. Barrattiana only by
its persistent stipules and the more densely hairy twigs. It has the character-

158 SARGENTIA [6

istic long, silky, pseudo-terminal aments of this species; and its leaves and habit
of growth also place it here. It should be noted that the character of stipule per-
sistence is not uncommon in this northwestern arctic and subarctic country.
Salix pulchra, S. Richardsonii, and S. glauca var. perstipula (described above)
are distinguished by it.

Salix Richardsonii Hooker, Fl. Bor.-Am. 2: 147, t. 182 (1839); Macoun, Cat. Can. Pl.
Pt. 3: 454 (1886); Hultén, Fl. Alaska & Yuk. Pt. 3: 538 (1943); Porsild in Nat.
Mus. Can. Bull. 101: 18 (1945).

“Fort Franklin, Mackenzie River,” Richardson (G, N); upper part of timber on the
north slope of Colonel Mt., alt. 3200’—4000’, shrubs 4’~6’ high, 9391, 9624; thickets along
small snow-fed stream above timber line, north slope of Colonel Mt., alt. about 5000’, shrubs
4'-5' high, 9372, 9373; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11227 (Can.).

A primary species among the shrubs in the open black spruce forest on the
lower north slopes of Colonel Mt. It extends also some distance above timber
line and is one of the principal species along snow-fed streams. Capsules ma-
ture in the timbered areas early in July, but they are found with flowers at higher
levels at the same time. Salix Richardsonii was observed only once except on
the north slope of Colonel Mt. This was at an elevation of about 3800’ on the
south slope of Terrace Mt., where it was growing in a springy muskeg behind a
small ridge. Porsild notes it also as common in a muskeg along the Canol Rd.
at mi. 51 E,

Salix alaxensis (Anders.) Coy. in Proc. Wash, Acad. Sci. 2: 280 (1900); Preble in N.
Am. Fauna 27: 521 (1908); Schneider in Jour. Arn. Arb. 1: 223 (1920); Raup in
Jour. Arn. Arb. 17: 235 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 539 (1943);
Porsild in Nat. Mus. Can. Bull. 101: 17 (1945).

Salix speciosa Hooker & Arn. sensu Hooker, Fl. Bor.-Am. 2: 145 (1839); Macoun, Cat.
Can. Pl. Pt. 3: 454 (1886) ; non Host.

Salix speciosa var. alaxensis Anders. in DC. Prodr. 162: 275 (1868).

Northwest shore of Great Slave L., Bedford (Can.); Norman, Dutilly 2 (G); “Fort
Franklin, Mackenzie River,” Richardson (G) ; along small snow-fed stream on north slope
of Colonel Mt., alt. about 5000’, shrubs up to 4 high, 9375; Lone Mt., Wynne-Edwards
8457 (Can.) ; alpine bog, Bolstead Cr., Canol Rd., mi. 112 E., Wynne-Edwards 8270 (Can.) ;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11228 (Can.).

Common to abundant along watercourses on the mountain slopes and on stony
or gravelly fans at the shore of Brintnell L. It is a primary species in this phase
of the willow scrub above timber-line on Colonel Mt., and in the upper valley of
Frost Cr. Capsules mature early in July. Such Mackenzie basin plants as I
have seen represent the typical form or var. longistylis, rather than var. obovali-
folia Ball in Jour. Wash. Acad. Sci. 28: 443 (1938). The range of the latter
variety appears to be eastward toward Hudson Bay (See Raup in Sargentia 4:
115), although Ball cites one specimen (Preble 290) from Great Bear Lake.

Salix alaxensis (Anders.) Cov. var. longistylis (Rydb.) Schn. in Jour. Arn. Arb. 1:
225 (1920); Hultén, Fl. Alaska & Yuk. Pt. 3: 541 (1943).

Salix longistylis Rydb. in Bull. N. Y. Bot. Gard. 2: 163 (1901).

Stony shore of Brintnell L. near mouth of Frost Cr., shrub 15’ high, 9149,

This variety apparently is only occasional in our region, and it was not ob-
served above timber line. Capsules mature in the third week of June.

Salix candida Fliigge apud Willd. Sp. Pl. 4: 708 (1805); Hooker, Fl. Bor.-Am. 2: 144
(1838); Macoun, Cat. Can. Pl. Pt. 3: 446 (1886); Schneider in Jour. Arn. Arb.
1: 226 (1920); Raup in Jour. Arn. Arb. 17: 237 (1936); Porsild in Sargentia 4:
27 (1943).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 159

Lobstick Cr., Little Buffalo R., Russell 11 (Can.); Slave R., Richardson (Can.) ; muskeg
thicket on the upland west of the Mackenzie R. near Simpson, 9909.

This species appears to be widespread in the “woody country,” as Hooker
stated in Fl. Bor.-Am., but it is only occasional in occurrence. Porsild reports
having seen it only once, at Great Bear L., during his extensive travels in north-
western Mackenzie. The Simpson specimen was collected late in August and
is sterile.

Salix Bebbiana Sarg. in Garden & Forest 8: 463 (1895); Preble in N. Am. Fauna 7 fe
522 (1908); Schneider in Jour. Arn. Arb. 2: 66 (1920); Raup in Jour. Arn. Arb.
17: 237 (1936); Hultén, Fl. Alaska & Yuk. Pt. 3: 544 (1943).
Salix rostrata Richardson sensu Hooker, Fl. Bor.-Am. 2: 147 (1839) ; Macoun, Cat. Can.
Pl. Pt. 3: 453 (1886); non Thuill.

Fort Smith, Raup 538, 2118; same, Seton & Preble 78311, 78312, 78309, 78310 (Can.) ;
Slave R., Kennicott (N); Resolution, Preble 195, 197 (US); same, Dutilly 108 (G); Rae,
A. E. & E. A. Preble 137 (US); northwest shore of Great Slave L., Bedford (Can.).;
Providence, Dutilly 81 (G); Simpson, Preble 302-A, 306-A, 307 (US); damp lowland
woods near Simpson, shrub 15’ high, 9080; in scrub timber on south slope of Red Mt.,
Brintnell L., alt. about 3500’, shrub 10’ high, 9223; Wrigley, Preble & Cary 85 (TIS) *
“between Simpson and Chipewyan,” Richardson (Can.) ; “between Cumberland House and
Fort Franklin,” Richardson (G, N); “Fort Franklin, Mackenzie River,” Richardson (N).

Salix Bebbiana is one of the commonest willows in the Mackenzie basin. In
the Simpson area it is common to abundant, forming a large part of the shrub
growth in the upland woods west of the river. Preble states that it extends along
the rivers to the lower Mackenzie. In the Brintnell Lake region, however,
though common on the south slope of Red Mt., it was not seen elsewhere except
as the following proposed variety, and then only occasionally. In flower at
Brintnell L. in June.

Here, as in my studies of eastern arctic and subarctic willows (Sargentia 4:
81-127), I am not attempting to separate the Mackenzie basin material into the
commonly recognized varieties. Glabrous- or glabrescent-leaved forms are
common throughout the range and have been called S. perrostrata by Kydb. (>:
Bebbiana var. perrostrata Schn.). There are, however, so many intermediate
forms that a segregation into recognizable entities is nearly if not entirely im-
possible.

According to Schneider (1. c.), Hooker’s record for S. fusca L. in Fl. Bor.-
Am. (2: 151) is referable to S. Bebbiana. It was based on specimens with male
aments collected at “Fort Franklin on the Mackenzie River” by Richardson.

Salix Bebbiana Sarg. var. depilis var. nov.

A forma typica capsulis ramulis foliisque glabris differt.

Gravelly beach on shore of Brintnell L. at Camp Pt., shrub 5’ high, 9306, f. (Type).

Occasional in this type of habitat; collected with immature capsules June 28.
Some of the material collected by Miss Alice Eastwood at Dawson and else-
where in Yukon, in 1914, appears to represent this glabrous form of the species.
Her numbers 238, 519, and 566, in the herbarium of the Arnold Arboretum, may
be cited. Schneider noted these glabrous specimens in his discussion of S. Beb-
biana (Jour. Arn. Arb. 2: 66-72, 1920) but made no attempt to separate them
taxonomically ; nor did he mention the fact that their capsules were glabrous or
nearly so. Variety perrostrata (Rydb.) Schn., described from material collected
in the Black Hills (Bull. N. Y. Bot. Gard. 2: 163, 1901), is distinguished by some
students, and is separated from the species by its glabrescent leaves. Its cap-

160 SARGENTIA [6

sules, however, are hairy like those of the type. Our Brintnell L, plants are
almost completely glabrous, even to the very young leaves and twigs. There are
a few long ciliate hairs on the margins of the floral bracts. Two of the East-
wood numbers cited above (238, 566) are slightly hairy on very young twigs and
petioles, but no. 519 closely resembles our plants.

Salix pedicellaris Pursh var. hypoglauca Fernald in Rhod. 11: 161 (1909); Raup in
Jour. Arn. Arb. 17: 238 (1936).

Salix pedicellaris Pursh; sensu Hooker, Fl. Bor.-Am. 2: 150 (1839), in part; Schneider
in Jour. Arn, Arb, 2: 81 (1920), in part.

Salix myrtilloides sensu Macoun, Cat. Can. Pl. Pt. 3: 451 (1886); Preble in N. Am.
Fauna 27: 521 (1908); non L.

“Fort Franklin, Mackenzie River,” Richardson (N).
Noted in Fl. Bor.-Am. from “Saskatchewan to Fort Franklin. . . ,’ and by

Preble at Norman. In the Athabaska-Great Slave L. region it is common on
the south side of L. Athabaska but was not seen elsewhere.

Salix petiolaris Smith in Trans. Linn. Soc. London 6: 122 (1802); Raup in Jour. Arn.
Arb, 17: 238 (1936).

Fort Smith, Raup 568.

Salix petiolaris is common in the Wood Buffalo Park, but has been collected in
southwestern Mackenzie only at Fort Smith. It will almost certainly be found
in the northern area of the Park, if not along the upper Mackenzie.

|

Salix Scouleriana Barratt var. coetanea Ball in Jour. Wash. Acad. Sci. 24: 73 (1934).

Salix Scouleriana Barratt, sensu Hultén, Fl. Alaska & Yuk. Pt. 3: 550 (1943); Porsild
in Nat. Mus. Can. Bull. 101: 18 (1945).

Stony shore of Brintnell L. and open spruce woods on Camp Pt., shrubs or small trees
15'-20' high, 9146, 9234.

Occasional or common on stony fans on the north side of Brintnell L., and ex-
tending up the warm south-facing slopes of Red and Terrace Mts. to about 4000’.
Not observed south of the lake. Capsules mature in the latter part of June.
On the lower mountain slopes it grows to tree size, with trunks as much as 6” in
diameter. Porsild saw it on a small tributary to Godlin R., along the Canol Rd.,
mi. 168 E., where it was growing to a height of 20’... The extent of the species
east of the mountains is uncertain. Preble collected specimens of a willow at
Simpson which Rydberg determined as S. Nuttallii Sarg., now regarded as a
synonym of S. Scouleriana. I have not seen the Preble specimen, and did not
find the species growing at Simpson when I was there in 1939.

The pubescence on the young leaves and twigs in our plants is rather distinctive
in being silky and somewhat shining instead of more or less opaque as in typical
S. Scouleriana. More material is required before this character can be evaluated,
especially in a species complex that is notably variable in leaf form and pubes-
cence.

Salix pulchra Cham. in Liinnaea 6: 543 (1831); Schneider in Jour. Arn. Arb. 1: 70
(1919); Hultén, Fl. Alaska & Yuk. Pt. 3: 547 (1943); Porsild in Nat. Mus. Can.
Bull. 101: 18 (1945).

“Fort Franklin, Mackenzie River,” Richardson (N, no. 64, H. B. & T.); near summit of
Red Mt., alt. 5800’, depressed shrub 1’ high, 9230; northeast slope of Red Mt., alt. 5500’,
shrub 1’ high, 9299; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11229 (Can.).

Occasional at high elevations on Red Mt., but not seen elsewhere in the Brint-
nell L. area; collected with flowers and immature capsules in the last week of

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 161

June. Schneider (1. c¢.) cites a doubtful specimen collected by Capt. Pullen from
“Fort Simpson to Great Bear Lake.”

Salix planifolia Pursh, Fl. Am. Sept. 2: 611 (1814); Hooker, Fl. Bor.-Am. 2: 150
(1839); Schneider in Jour. Arn. Arb. 1: 75 (1919); Raup in Jour. Arn. Arb. 17:
239 (1936).
Salix chlorophylla sensu Macoun, Cat. Can. Pl. Pt. 3: 446 (1886) ; Preble in N. Am.
Fauna 27: 522 (1908) ; non Anders.

Resolution, E. A. & A. E. Preble 143 (US); about 15 mi. east of Moraine Pt., north-
west shore of Great Slave L., Bedford (Can., doubtful fragment); Rae, E. A. & A. E.
Preble 136 (US); Simpson, Preble 303-B, 305 (US); “Fort Franklin, Mackenzie River,”
Richardson (N); stony shore of Brintnell L. at Camp Pt., shrubs 6’ high, 9148, 9150.

This species is common on the stony fans at Brintnell L., and also in the willow-
birch scrub in the upper valley of Frost Cr. (alt. about 4000’). Preble states
that it is also common at Simpson. At Brintnell L. the capsules were beginning
to mature during the third week in June. Noted in Fl. Bor.-Am. on “Mackenzie
River and Bear Lake.” Schneider (1. c.) cites a Preble specimen (322-A)
from Norman, but I have not seen it.

In my studies of the Athabaska-Great Slave L. flora I determined a few speci-
mens from the Wood Buffalo Park as S. Nelsonii Ball. One of these (Raup
514) came from Fort Smith, which is just in the southeastern corner of the area
treated here. I am doubtful if this form can be maintained as distinct from S.
planifolia in our region. The latter is extremely common in the forested parts
of the Athabaska country and the Wood Buffalo Park, where it inhabits slough
margins and local river flood plains.

Our Mackenzie Mt. plants strongly suggest S. pulchra in general habit, and
although most of the leaves have the characteristic elliptic-lanceolate, lanceolate,
or obovate-oblong shapes of S. planifolia, a few are somewhat rhombic as in S.
pulchra. The plants have no persistent stipules, however, and since this is a
principal character by which S. pulchra is distinguished, it seems best to keep the
material in S. planifolia. Salix pulchra occurs in the region but seems to be con-
fined to high mountain slopes.

Salix arbusculoides Anders. in Vet. Akad. Handl. (Stockh.) 6: 147, t. 8, f. 81 (1867);
Macoun, Cat. Can. Pl. Pt. 5: 358 (1890); Preble in N. Am. Fauna 27: 522 (1908);
Schneider in Jour. Arn. Arb. 2: 84 (1920); Raup in Jour. Arn. Arp: 173-239
(1936); Porsild in Sargentia 4: 27 (1943); Hultén, Fl. Alaska & Yuk. Pt. 3: 549
(1943); Porsild in Nat. Mus. Can. Bull. 101: 18, 31 (1945).

Salix acutifolia sensu Hooker, Fl. Bor.-Am. 2: 150 (1839), non Willd.

Salix argyrocarpa sensu Macoun, Cat. Can. Pl. Pt. 3: 445 (1886), non Anders.

Fort Smith, Raup 2096; lower Slave R., Harper 99027 (Can.) ; Resolution, Preble 201
(US) ; northwest shore of Great Slave L., Bedford (Can.) ; Lake Grant, Preble 248 (WS);
Providence, Dutilly 80 (G); damp woods near Simpson, shrub 15’ high, 9079; river bluff
at Simpson, shrub 6’ high, 9104; Norman, Dutilly (G) ; spruce woods and thickets just back
of Brintnell L. Camp, shrubs or small trees up to 20’ high, 9263, 9317; Nahanni Mts.,
Preble 317 (US); Lone Mt., Wynne-Edwards 8458 (Can.) ; Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8271 (Can.); Sekwi R., Canol Rd., mi. 174 E., Porsild 11859
¢Gan.).

Common; our Simpson specimens were maturing their capsules in mid-June,
and those at Brintnell L. in late June and early July. This species forms tall
slender trunks, up to 2” in diameter, when it grows in spruce woods. At Brint-
nell L. it was not observed more than a hundred feet or so above lake-level.

162 SARGENTIA [6

MYRICACEAE
Myrica Gale L. Sp. Pl. 1024 (1753); Hooker, Fl. Bor.-Am. 2: 160 (1839); Macoun,
Cat. Can. Pl. Pt. 3: 434 (1886); Raup in Jour. Arn. Arb. 17: 240 (1936).

Clewi R., Russell 51 (G); northwest shore of Great Slave L., Bedford (Can.) ; Mack-
enzie R., between Blackwater R. and Norman, Crickmay 163 (Can.).

Preble (N. Am. Fauna 27: 523) reported this species northward to Great Bear
L., and it was noted in Fl. Bor-Am. “Throughout Canada, and to Fort Norman
on the Mackenzie River.” Recently Porsild (Sargentia 4: 28) reports that it is
common at Great Bear L. but rare in the Mackenzie delta.

BETULACEAE

Betula papyrifera Marsh. var. commutata (Regel) Fernald in Rhod. 47: 312 (1945).

Betula alba ssp. occidentalis (Hook.) Regel, 8 commutata Regel in Bull. Soc. Nat. Mosc.
38: 401 (1865).

Betula papyrifera Marsh. var. occidentalis Sarg.; Raup in Jour. Arn. Arb. 17: 241 (1936) ;
Porsild in Sargentia 4: 29 (1943); non B. occidentalis Hooker,

Betula papyrifera Marsh. ssp. occidentalis Hultén, Fl. Alaska & Yuk, Pt. 4: 582 (1944),
non B, occidentalis Hooker.

Liard R. between Nahanni Butte and Simpson, Crickmay 32 (Can.) ; gravelly open woods
along Frost Cr. near Brintnell L. Camp, tree 30’ high, 6” diam., 9566; woods on lower slope
of Red Mt., just back of Camp Pt., trees 30’-40’ high, 4-5” diam., 9314, 9318; mossy shale
ledge in gorge below east end of Brintnell L., tree 30’ high, 2” diam., 9589; same, tree 10’
high, 1.5’" diam., 9578; open spruce wood on rocky knoll near east end of Brintnell L., tree
30’ high, 4” diam., 9587; Bear Rock, near Norman, Porsild (Can.).

Common to abundant in the woods around Brintnell L. and on the lower
mountain slopes. In some places it is a primary part of the forest. It is the
commonest tree birch of the area, usually distinguished by its reddish brown bark
and by the proportionately elongated middle lobes of the floral bracts. Its twigs
are usually glabrous and but little resinous. Between it and var. humilis, how-
ever, there is a maze of intermediate forms. Some of them have the glandular
twigs of the latter but have bark strongly tinged with reddish brown, even in
large trees. Others have brown bark and moderately resinous twigs. Still
others have dark bark and non-resinous twigs but have bracts that look like those
of var. humilis. This is illustrated by no. 9578. A somewhat greater divergence
is illustrated by no. 9587, which has its bark somewhat tinged with red and it
twigs slightly resinous like some of the intermediates. Most of its leaves, how-
ever, are truncate or shallowly cordate at the base, while those of the other forms
are usually broadly cuneate. Its bracts have the elongated terminal lobes as in
var. commutata, but they also have elongated divergent lateral lobes. The seed
and its wings, though immature at the time of collection, show a tendency to be
cuneate at the base. This form may prove worthy of distinction, but the material
at hand (consisting of 3 branchlets with but a single ament among them) is too
scanty for further interpretation.

Betula papyrifera Marsh. var. humilis (Regel) Fernald & Raup in Rhod. 47: 321 (1945).

Betula alba ssp. papyrifera var. humilis Regel in DC. Prodr. 16: 166 (1868).

Betula papyrifera var. neoalaskana (Sarg.) Raup in Contr. Arn. Arb. 6: 152 (1934), in
Jour. Arn. Arb. 17: 241 (1936); Porsild in Sargentia 4: 29 (1943), in Nat. Mus.
Can. Bull. 101: 18 (1945).

Betula papyracea Ait.; Hooker, Fl. Bor.-Am. 2: 155 (1839), in part.

Betula papyrifera sensu Macoun, Cat. Can. Pl. Pt. 3: 436 (1886), in part; Preble in N.
Am. Fauna 27: 523 (1908), in part.

Betula resinifera Britton; Hultén, Fl. Alaska & Yuk. Pt. 4: 583 (1944).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 163

Lower Slave R., Raup 604; northwest shore of Great Slave L., Bedford (Can.) ; Mack-
enzie R., Onion, Kennicott & Hardisty (N); Liard R., between Nahanni Butte and Simp-
son, Crickmay 29 (Can.); ? Simpson, Onion (N); lowland woods near Simpson, tree 20'
high, 9854; shale ledge in gorge below east end of Brintnell L., small tree 6’ high, diam. 1”,
9591; Twitya R. valley, Canol Rd., mi. 124 E., Porsild 11838 (Can.).

This is the common white birch of the Mackenzie valley, but it is only rare
or occasional in the Brintnell L. district (see note under B. papyrifera var. com-
mutata). Porsild states that it is common along the Canol Rd. west of Norman
Wells only to mi. 124, where it was last seen. Our no, 9591 has the characteristic
resinous twigs of var. humilis, and the bark is white, even on young trees. Cat-
kins mature in late summer. In FI. Bor.-Am. the white birch is reported north-
ward only to lat. 65°, but both Preble and Porsild give it a range in the Mack-
enzie basin extending throughout the wooded region.

It proved difficult, during the summer of 1939 at Brintnell L., to secure good
specimens of the larger birches. Very few aments were produced, although the
abundant bare axes of the preceding year’s crop indicated that the scarcity was
not of regular occurrence. It was necessary in some cases to cut down whole
trees or large branches and search them carefully in order to find enough aments
for specimens. Most of the trees were badly infested with a leaf-curling insect.

Betula occidentalis Hooker, Fl. Bor.-Am. 2: 155 (1839); ? Macoun, Cat. Can. Pl. Pt.
3: 437 (1886). See Fernald in Rhod. 47: 312-316 (1945).

Betula microphylla sensu Raup in Jour. Arn. Arb. 17: 242 (1936); Porsild in Sargentia
4: 29 (1943), in Nat. Mus. Can. Bull. 101: 31 (1945) ; non Bunge.

Open spruce woods on rocky knoll near east end of Brintnell L., shrub 5’-6' high, 9590;
north peak of Nahanni Mt., alt. 2700’, Wynne-Edwards 8464 (Can.).

Common at the Brintnell L. station cited above, but observed in only one other
place in the area, on a terrace at about 3700’ on the south slope of Terrace Mt.
Collected with maturing fruit July 24. Occasional in the Athabaska—Great Slave
L. region, and collected by Porsild at Great Bear L. and in the Mackenzie delta.

Hultén, in Fl. Alaska & Yuk. Pt. 4: 575, considers that all Alaskan material
that has been called B. microphylla or B. fontinalis Sarg. is probably composed of
hybrids between B. glandulosa and B. resinifera (= B. papyrifera var. humilis).
He suggests further that all Mackenzie and eastern plants of this category have
had the same origin. Betula occidentalis in our region is a rather well-defined
and easily recognizable form, and until some proof of its hybrid origin is forth-
coming I believe it more useful to maintain it as a species.

Betula pumila L. var. glandulifera Regel in DC. Prod. 162: 171 (1864); Raup in Jour.
Arn. Arb. 17: 242 (1936).

Fort Smith, Dutilly 95 (G).

There are only two specimens of this species known to me from the Mackenzie
basin. One is the Fort Smith plant cited above, and the other came from the
Conibear (Moose) L. district of the Wood Buffalo Park (Raup 2176). It is
probably occasional in the upper Mackenzie valley.

Betula glandulosa Michx. Fl. Bor.-Am. 2: 180 (1803); Hooker, Fl. Bor.-Am, 2: 156
(1839); Macoun, Cat. Can. Pl. Pt. 3: 437 (1886); Preble in N. Am. Fauna 27:
524 (1908); Raup in Jour. Arn. Arb. 17: 242 (1936); Hultén, Fl. Alaska & Yuk.
Pt. 4: 573 (1944); Porsild in Nat. Mus. Can. Bull. 101: 18, 31 (1945).

Great Slave L., R. Bell 23157 (Can.) ; between Forts Simpson and Chipewyan, Richard-
son (Can.) ; Lined Ri between Nahanni Butte and Simpson, Crickmay 31 (Can.); “Fort
Franklin, Mackenzie R.,” Richardson (G); muskeg along south shore of Brintnell L., shrub
3’ high, 9341; open spruce woods on rocky knoll near east end of Brintnell L., shrub 4’ high,

164 SARGENTIA [6

9585; willow-birch scrub in upper valley of Frost Cr., alt. 4000’-4500’, shrubs up to 5’ high,
9510; shale slides on west slope of Terrace Mt., alt. 6000’, depressed shrub, 9502; Lone Mt.,
Wynne-Edwards 8465 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., wet alpine bog, alt.
5500’, Wynne-Edwards 8280 (Can.).

Common to abundant in the Brintnell L. area, forming thickets on lowland
muskegs and watercourses, and extending to high alpine situations. It has its
greatest development in a “zone” just above timber line, ranging in vertical width
from a narrow band to 1000 feet. This zone is broadest on the southward- and
westward-facing slopes of Red and Terrace Mts., and narrowest on the north
slope of Colonel Mt. Betula glandulosa is a primary species in most of the zone,
associated with willows and other shrubs. Found with maturing aments late in
July at lake-level; alpine specimens of that period have very young aments.
Preble’s records for B. nana and B. glandulosa, “. . . practically throughout the
Hudsonian zone,” and “throughout the region . . . northward into the Barren
Grounds,” probably both belong here, and it is noted in Fl. Bor.-Am. from “. . .
Lake Winipeg to Fort Franklin.”

Alnus crispa (Ait.) Pursh, Fl. Am. Sept. 2: 623 (1814); Raup in Jour. Arn. Arb. 17:
243 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 586 (1944); Porsild in Nat. Mus.
Can. Bull. 101: 18 (1945).

Betula crispa Ait. Hort. Kew 3: 339 (1789).

Alnus viridis sensu Hooker, Fl. Bor.-Am. 2: 157 (1839), at least in part; Macoun, Cat.
Can. Pl. Pt. 3: 438 (1886); non DC.

Alnus alnobetula Ehrh.; Preble in N. Am. Fauna 27: 525 (1908).

“Between Forts Simpson and Chipewyan,” Richardson (Can.) ; Willow Lake R. near its
mouth, Crickmay 36 (Can.) ; gravelly fans and shores near Brintnell L. camp, shrubs up to 8’
high, 9147, 9815.

Common to abundant in the Brintnell L. area. It is a primary species in the
open black spruce forest on the lower north slopes of Colonel Mt., and extends
up to timber line or a little above on the mountains north of the lake. Here
it is associated with Betula glandulosa and Salix glauca var. perstipula in dense
timber line scrub. It is one of the commonest shrubs of gravel fans along the
lake. Collected in flower late in June and with maturing cones Aug. 15. Noted
by Preble “. . . throughout the region from the Saskatchewan northward to the
tree limit” ; and in Fl. Bor.-Am. “. . . throughout the woody and barren country
...’ Porsild states that along the Canol Rd. it was common as far west as the
upper Godlin R., mi. 142 E.

Alnus tenuifolia Nutt. Sylva 1: 32, t. 10 (1842).

Alnus incana sensu Hooker, Fl. Bor.-Am. 2: 157 (1839), in part; Macoun, Cat. Can. PI.
Pt. 3: 438 (1886), in part; Preble in N. Am, Fauna 27: 525 (1908); Raup in Jour.
Arn. Arb. 17: 243 (1936); Hultén, Fl. Alaska & Yuk, Pt. 4: 590 (1944); non (L.)
Moench.

Northwest shore of Great Slave L., Bedford (Can.); Willow Lake R. near its mouth,
Crickmay 35 (Can.) ; thickets on low ground about the western end of Brintnell L., shrub
10’ high, 9471.

Alnus tenuifolia is an important component of the lowland flood plain woods
along the Mackenzie at Simpson. At Brintnell L. it is abundant in the situation
noted above, but was not seen elsewhere. Noted in Fl. Bor.-Am. as far north as
Fort Franklin, and by Preble north to the Peel R.

Hultén considers the Alaskan material in the “incana group” to be identical
with the European. Fernald (1. c.), however, has recently shown that the eastern
plants are clearly not the European A. incana, and he has taken up the name A.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 165

rugosa for the more northerly of them. Our plants, though not idential in all
respects with A. rugosa, appear to be nearer that species than to typical A. incana.
Pending further study, therefore, I have preferred to apply Nuttall’s name, A.
tenuifolia, to them.

URTICACEAE

Urtica gracilis Ait. Hort. Kew 3: 341 (1789); Hooker, Fl. Bor.-Am., 2: 141 (1838);
Macoun, Cat. Can. Pl. Pt. 3: 430 (1886); Raup in Jour. Arn. Arb. 17: 244 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 4: 592 (1944). See Fernald in Rhod. 28: 193
(1926).

Clewi R., Russell 13 (G); northwest shore of Great Slave L., Bedford (Can.) ; Wrigley
Harbor, Great Slave L., on gravelly shore, 9980.

Common in the Wood Buffalo Park and probably northward into the Macken-
zie valley. Collected with maturing fruit early in September. In Fi. Bor.-Am.
it is said to extend “. . . throughout the woody country, generally, on the east of
the Rocky Mountains, as far as Fort Franklin, on the Mackenzie River.” Fer-
nald has shown that it has an extensive range in the northern Cordillera as well.

SANTALACEAE
Geocaulon lividum (Richardson) Fernald in Rhod. 30: 23 (1928); Raup in Jour. Arn.
Arb, 17: 244 (1936); Hultén, Fl. Alaska & Yuk, Pt. 4: 593 (1944).

Comandra livida Richardson, Bot. App. Frankl. Journ. 734 (1823) ; Hooker, Fl. Bor.-Am.
2: 139 (1838) ; Macoun, Cat. Can. Pl. Pt. 3: 423 (1886).

Fort Smith, Raup 648; southwest and north shores of Great Slave L., Howe 91983
(Can.) ; northwest shore of Great Slave L., Bedford (Can.) ; east shore Gteurout Li, C.a0s
Hage (G); Liard R. between Nahanni Butte and Simpson, Crickmay 33 (Can.); damp
woods near Simpson, 9100; near Wrigley, Miss E. Taylor 134 (Can.) ; Willow Lake R.
near its mouth, Crickmay 37 (Can.); Willow Lake R., 44 mi. above mouth, Crickmay 16
(Can.) ; woods along shore of Brintnell L., 9302; Nahanni Mts., Preble & Cary 72 (US).

Abundant in low woods; collected in flower during middle and late June.
Noted in Fl. Bor.-Am. northward “to lat. 69° on the Mackenzie River.”

POLYGONACEAE
Rumex mexicanus Meissn. in DC. Prodr. 14: 45 (1856); Raup in Jour. Arn. Arb, 17:
245 (1936).

Rumex salicifolius sensu Hooker, Fl. Bor.-Am. 2: 129 (1838), in part; Macoun, Cat.
Can. Pl. Pt. 3: 416 (1886), in part; non Weinm.

Rumex sibiricus sensu Hultén, Fl. Alaska & Yuk. Pt. 4: 605 (1944), in part, non Hultén.

Northwest shore of Great Slave L., Bedford (Can.); stony shore of Great Slave L. at
Wrigley Harbor, 9983; open field near Simpson, 9838.

Occasional to common in the Simpson district, where it was collected in fruit
late in August. Noted in Fl. Bor.-Am. from “Lake Winnipeg, to Great Bear
Lake and Mackenzie River.”

Rumex mexicanus has been subdivided by Rechinger (Field Mus. Publ. Bot.
17: 58), and the northern representatives referred to R. triangulivalvis (Danser)
Rich. These northern plants are described as having smaller valves and nut-
lets (valves 3 mm., nutlets 2 mm.) than typical R. mexicanus (valves 4 mm., 2.5
mm.). Our plants, however, have the larger valves and nutlets of the latter.
Hultén (1. c.) has recently suggested that at least a part of the Mackenzie basin
material belongs to his R. sibiricus (Fl. Kamtch. 2: 48, f. 7, 1928). Judging by
his figure and description, however, this is not the case, for our plants have pro-
portionately larger and more pointed valves.

166 SARGENTIA [6

The Wrigley Harbor plants are peculiar in having purplish valves which were
maturing without the development of any nutlets. Occasional specimens in the
Gray Herbarium collection show poorly developed or undeveloped nutlets, but
whether the character is worthy of recognition is uncertain. Our specimens
were collected on Sept. 9, after frosts had commenced, so that the growth of the
nutlets may have been halted prematurely.

Rumex maritimus L. var. fueginus (Phil.) Dusén, Svenska Exp. till Magellansl. 3:
194 (1900); Raup in Jour. Arn. Arb. 17: 245 (1936). See Fernald in Rhod. 17:
80 (1915).

Rumex fueginus Phil. in Ann. Univ. Chile 91: 493 (1895) ; Hultén, Fl. Alaska & Yuk.
Pt. 4: 603 (1944).

Little Buffalo R., Russell 29 (G); northwest shore of Great Slave L., Bedford (Can.) ;
damp sand bar along Mackenzie R. near Simpson, 9870; rocky-sandy bank of Mackenzie R.
opposite Simpson, 9937,

Occasional to common in these habitats; collected in fruit the last of August.
Not previously reported below Great Slave L.

Rumex arcticus Trautv. in Middend. Sib. Reise 1: 29 (1856); Porsild in Nat. Mus.
Can. Bull. 101: 18 (1945).

Noted by Porsild as common in Macmillan Pass, Canol Rd., mi. 225 E.

Oxyria digyna (L.) Hill, Hort. Kew. 158 (1768); Hultén, Fl. Alaska & Yuk. Pt. 4:
607 (1944); Porsild in Nat. Mus. Can. Bull. 101: 18 (1945).

Rumex digynus L. Sp. Pl. 337 (1753).

Damp springy ledges and gullies on north slope of Colonel Mt., alt. 3500’-5000', 9381,
9656; Bolstead Cr., Canol Rd., mi. 111 E., alpine peak, alt. 6000'-7000', Wynne-Edwards
8281 (Can.).

Apparently more common on the north-facing slope of Colonel Mt. than else-
where in the Brintnell L. area, occurring both above and below timber line ; col-
lected in flower July 5, and with maturing fruit July 28.

Polygonum aviculare L. Sp. Pl. 362 (1753); Hooker, Fl. Bor.-Am. 2: 132 (1838) ;
Macoun, Cat. Can. Pl. Pt. 3: 407 (1886); Raup in Jour. Arn. Arb. 17: 245 (1936).

Polygonum buxiforme Small, sensu Hultén, Fl. Alaska & Yuk. Pt. 4: 615 (1944), in part.

Fort Smith, Raup 658; northwest shore of Great Slave L., Bedford (Can.); damp sand

bar along Mackenzie R. near Simpson, 9876; along wood road just west of town of Simpson,
9878.

Common in the Simpson area; collected with maturing fruit in the last week
of August. These specimens are referred to P. aviculare provisionally, and using
the name in its broad sense. It is not impossible that two entities are involved,
one native and the other an introduced weed growing around the settlement.
Polygonum aviculare, also considered in the broad sense, was noted by Hooker as
occurring “throughout Canada, to lat. 65°...”

Polygonum viviparum L. Sp. Pl. 360 (1753); Hooker, Fl. Bor.-Am. 2: 130 (1838);
Macoun, Cat. Can. Pl. Pt. 3: 412 (1886); Raup in Jour. Arn. Arb. 17: 245 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 4: 620 (1944); Porsild in Nat. Mus. Can. Bull.
101: 18, 31 (1945).

Windy Pt., Great Slave L., Hume 102660 (Can.) ; Mackenzie R., Onton (N); Liard R.
between Nahanni Butte and Simpson, Crickmay 34 (Can.); Bear R., Miss E. Taylor 124
(Can.) ; stony shore of Brintnell L. at Camp Pt., 9193; alpine meadows and scrub on north
slope of Colonel Mt., 9407; along streams in upper valley of Frost Cr., alt. 4000’-5500’,
9525; turfy east slope of Red Mt., alt. 5000’-5900’, 9736; Lone Mt., W ynne-Edwards 8466
(Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5000’, Wynne-Edwards 8283
(Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 167

Common ; collected in flower at Brintnell L. on June 20, and at the summit of
Red Mt. on Aug. 7. Porsild notes that it is only rare or occasional in Macmillan
Pass.

Polygonum Bistorta L. ssp. plumosum (Small) Hultén, Fl. Alaska & Yuk. Pt. 4: 613
(1944); Porsild in Nat. Mus. Can. Bull. 101: 18 (1945).

Polygonum plumosum Small in Bull. N. Y, Bot. Gard. 2: 166 (1901).
Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5500’, Waynne-Edwards 8282 (Can.).
Polygonum alpinum All. var. lapathifolium Cham. & Schl. in Linnaea 3: 38 (1828);
Porsild in Nat. Mus. Can. Bull. 101: 18 (1945).
Polygonum alaskanum (Small) Wight; Hultén, Fl. Alaska & Yuk. Pt. 4: 610 (1944).
Noted by Porsild as common in Macmillan Pass, along the Canol Rd. It
grows on river bars.

Polygonum lapathifolium L. var. salicifolium Sibth. Fl. Oxon 129 (1794); Raup in
Jour. Arn. Arb. 17: 246 (1936). See Fernald in Rhod. 23: 259 (1921).

Polygonum Persicaria L. var. minus Hooker, Fl. Bor.-Am, 2: 132 (1838).

Damp sand bar along Mackenzie R. near Simpson, 9877.

Common in this habitat, where it usually shows a decumbent habit of growth,
the stems arching over from the base and trailing on the wet sand; collected with
maturing achenes Aug. 22. Occasional in the Wood Buffalo Park district, and
noted by Hooker from “Saskatchewan to Great Bear Lake.’ So far as I am
aware this is the only persicarioid Polygonum in the Mackenzie basin.

Polygonum natans A. Eat. forma Hartwrightii (Gray) Stanford in Rhod. 27: 160
(1925); Raup in Jour. Arn. Arb. 17: 246 (1936).

Polygonum Hartwrightii Gray in Proc. Am. Acad. 8: 294 (1870).

Polygonum amphibium sensu Hooker, Fl. Bor.-Am. 2: 131 (1838), in part, non L.

Fort Smith, Raup 663; northwest shore of Great Slave L., Bedford (Can.); upland
prairie west of Mackenzie R. near Simpson, 9920.

Only sterile specimens were collected near Simpson, late in August; and only
the terrestrial form was seen. Polygonum natans is common in the Wood Buf-
falo Park and probably also in the upper Mackenzie valley. Hooker noted only
the aquatic form as extending northward to Great Slave Lake, but the terrestrial
one is about equally common there.

CHENOPODIACEAE

Chenopodium capitatum (L.) Asch. Fl. Brandenb. 1: 572 (1864); Macoun, Cat. Can.
Pl. Pt. 3: 400 (1886); Raup in Jour. Arn. Arb. 17: 246 (1936); Hultén, Fl. Alaska
& Yuk. Pt. 4: 625 (1944).

Blitum capitatum L. Sp. Pl. 4 (1753); Hooker, Fl. Bor.-Am. 2: 126 (1838).

Lower Slave R., Raup 675; Little Buffalo R., Russell 35 (G); northwest shore of Great
Slave L., Bedford (Can.); Mackenzie R., Kennicott (N); cultivated field on west bank of
Mackenzie R. near Simpson, 9930; Mackenzie R. between Wrigley and Blackwater R.,
Crickmay 132 (Can.).

Occasional to common, chiefly in waste places and in disturbed soil; fruit ma-

ture in late August. In FI. Bor.-Am. it was noted northward only as far as
Great Slave L.

Chenopodium album L. Sp. Pl. 219 (1753); Hooker, Fl. Bor.-Am. 2: 127 (1838);
Macoun, Cat. Can. Pl. Pt. 3: 398 (1886); Raup in Jour. Arn. Arb. 17: 247 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 4: 624 (1944),

Stony bank of Mackenzie R. a few miles below Providence, 9979; rocky-sandy bank of
Mackenzie R. opposite Simpson, 9946.

/

168 SARGENTIA [6

A common weed of settlements and disturbed soils; in fruit late in August
and early in September. Noted in Fl. Bor.-Am. northward as far as Great
Bear L.

Chenopodium leptophyllum Nutt. ex Moq. in DC. Prodr. 132: 71 (1849); Raup in
Jour. Arn, Arb, 17: 247 (1936).

This species has not been seen in southwestern Mackenzie, but it appears to
have a wide though scattered range in the Mackenzie basin, and is to be expected
in our region. I have collected it in a small prairie just west of L. Athabaska,
and there is a Richardson specimen in Herb. G marked “Fort Franklin, Mack-
enzie R.” The latter was probably included in C. album in Fl. Bor.-Am.

Corispermum hyssopifolium L. Sp. Pl. 4 (1753); Hooker, Fl. Bor.-Am. 2: 125 (1838);
Macoun, Cat. Can. Pl. Pt. 3: 403 (1886); Raup in Jour. Arn. Arb. 17: 247 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 4: 632 (1944).

Great Slave L., Richardson (G); windblown sand bars along the Mackenzie R. near

Simpson 9868.

Common at Simpson, forming mat-like growths, usually of individual plants;

a primary species in this unstable habitat.

Salicornia europaea L. Sp. Pl. 3 (1753); Raup in Jour. Arn. Arb. 17: 247 (1936).
Salicornia herbacea L.; Hultén, Fl. Alaska & Yuk. Pt. 4: 633 (1944).
Junction of Nyarling and Little Buffalo Rivers, Russell 72 (G).
Common on the Salt Plains west of the Slave R.

AMARANTHACEAE

Amaranthus retroflexus L. Sp. Pl. 991 (1753).
Weed in a cultivated field on the west bank of the Mackenzie R. near Simpson, 9928.
Common in this situation; fruit mature in late August. So far as I am aware
this species has not previously been reported in the Mackenzie basin except in
the Peace R. agricultural districts.

PORTULACACEAE
Claytonia megarrhiza (Gray) Parry ex Wats. Bibl. Ind. 118 (1878); Porsild in Nat.
‘ Mus. Can. Bull. 101: 18 (1945).
Claytonia arctica Adams var. megarrhiza Gray in Am. Jour. Sci. II. 33: 406 (1862).
Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes of west wall, alt. 4500’-5500', Wynne-
Edwards 8626 (Can.).

CARYOPHYLLACEAE

Stellaria calycantha (Ledeb.) Bong. in Mém. Acad. Imp. Sci. St. Pétersb. Ser. 6,
Math. 2: 127 (1833); Porsild in Sargentia 4: 37 (1943); Hultén, Fl. Alaska &
Yuk. Pt. 4: 648 (1944); Porsild in Nat. Mus. Can. Bull. 101: 19 (1945). See
Rhod. 42: 254 (1940).

Stellaria borealis Bigel.; Hooker, Fl. Bor.-Am. 1: 94 (1830) ; Macoun, Cat. Can. Pl. Pt.

1: 74 (1883) ; Raup in Jour. Arn. Arb. 17: 248 (1936).

Northwest shore of Great Slave L., 12 mi. east of Moraine Pt., Bedford (Can.) ; Mack-
enzie R., Onion (N); Simpson, Onion, Kennicott & Hardisty (N); edge of cultivated field
near Simpson, 9/11; thickets on stony shore of Brintnell L., 9151; wet meadows on mud
flats along glacial stream near west end of Brintnell L., 9693.

Common; in flower at Simpson about mid-June, and from late June to early
August at Brintnell L. Collected by Porsild in the lower Mackenzie country and
at Great Bear L. Also noted by Porsild as common in Macmillan Pass, on

the Canol Rd.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 169

Stellaria crassifolia Ehrh. in Hannov. Mag. 8: 116 (1784); Raup in Jour. Arn. Arb. 17:
248 (1936); Porsild in Sargentia 4: 37 (1943); Hultén, Fl. Alaska & Yuk. Pt. 4:
650 (1944).
This species has not been collected in our region, but it has been found in
the Wood Buffalo Park and in western arctic Canada generally (see Porsild,
ey,

Stellaria longipes Goldie in Edinb. Phil. Jour. 6: 327 (1822); Hooker, Fl. Bor.-Am., 1:
95 (1830); Macoun, Cat. Can. Pl. Pt. 1: 75 (1883); Raup in Jour. Arn. Arb. 17:
248 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 656 (1944); Porsild in Nat. Mus.
Can. Bull. 101: 19 (1945).

Fort Smith, Seton & Preble 78318 (Can.) ; same place, Dutilly 94 (G) ; same place, Raup
717; same place, Miss E. Taylor (G, Can.) ; Lower Slave R., Raup 710; Resolution, Kennt-
cott (N) ; Hay River, Hume 102661 (Can.) ; southwest and north shores of Great Slave L.,
Howe 91981 (Can.) ; Mackenzie R., Onion (N); edge of cultivated field at Simpson, 9110;
“Cumberland House to Bear Lake,” Richardson (G).

The Simpson specimens represent the glaucous form of the plant, which some-
times goes by the name of var. lacta. It is entirely replaced in the Brintnell L.
area by the following variety. Stellaria longipes is noted in Fl. Bor.-Am.“. . . to
the Mackenzie River and Bear Lake.” It is occasional at Simpson, and in
flower about mid-June.

Stellaria longipes Goldie var. Edwardsii (R. Br.) Wats. Bibl. Ind. 113 (1878).

Stellaria Edwardsii R. Br. App. Parry’s 1°* Voy. 271 (1824).

? Stellaria laeta Richardson, sensu Hultén, Fl. Alaska & Yuk. Pt. 4: 654 (1944).

Liard R. between Nahanni Butte and Simpson, Crickmay 36, 37 (Can.); Mackenzie R.
between Blackwater R. and Norman, Crickmay 166 (Can.); “Fort Franklin, Mackenzie
R.,” Coll. ? (N); stony shore of Brintnell L., 9158; mossy bank of small stream at Camp
Pt., Brintnell L., 9191; rock-slide along south shore of Brintnell L., 9349; high alpine slide
rock, north slope of Colonel Mt., 9413; shale cliffs in gorge, north slope of Colonel Mt., alt.
3500’, 9834; alpine meadows on south slope of Red Mt., alt. about 5000’, 9206; Bolstead Cr.,
Canol Rd., mi. 111 E., alpine slopes, Wynne-Edwards 8296, 8297 (Can.).

Occasional to common at Brintnell L., and noted as occasional or rare by
Porsild in Macmillan Pass; most frequent on rock-slides of north-facing slopes.
In flower in late June and early July.

Hultén, in Fl. Alaska & Yuk. Pt. 4, has preferred to recognize two separate
species in the S. longipes complex: S. laeta Richardson and S. longipes Goldie.
From his discussion I presume that his S. laeta is the small, glaucous, 1- to few-
flowered form that I have called S. longipes var. Edwardsii. Our plants have
comparatively broad, very glaucous leaves, and few flowers. They appear to
match the descriptions and plate of S. Edwardsii of Hooker (FI. Bor.-Am. 1: 96,
t. 31) and Richardson (Bot. App. Frankl. Journ. 738) better than those of S.
lacta except for the pubescence of the sepals. Hultén has stressed this character,
using it as diagnostic in this case. In our plants, however, the pubescence on
the sepals is quite variable as to presence and quantity. Plants in the same colony
will vary from sepals pubescent to quite glabrous. My personal experience with
plants of the S. longipes group strongly suggests that entities found in it are
hardly worthy of specific rank. Their geographic segregation, however, sug-
gests that they should be considered as something more than forms (cf. Polunin
in Nat. Mus. Can. Bull. 92: 192, 1940).

Stellaria longifolia Muhl. in Willd. Enum. Pl. Hort. Bot. Berol. 479 (1806); Hooker,
Fl. Bor.-Am. 1: 94 (1830); Macoun, Cat. Can. Pl. Pt. 1: 74 (1883); Raup in Jour.
Arn. Arb. 17: 249 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 655 (1944).

170 SARGENTIA [6

Fort Smith, Raup 718, 723; edge of cultivated field at Simpson, 9109; Norman, Coll.?
(N).

Occasional in the upper Mackenzie valley; in flower about mid-June. Noted
in Fl. Bor.-Am. in the “Woody country from lat. 54° to 64°.”

Stellaria media (L.) Cyrill, Char. Comm. 36 (1784); Hultén, Fl. Alaska & Yuk. Pt.
4: 657 (1944).
Alsine media L. Sp. Pl. 272 (1753).
Damp thickets in a slough near Simpson, 9842.
Common in this situation; capsules maturing late in August. Not previously
noted in the Mackenzie basin north of L. Athabaska (Raup in Jour. Arn. Arb.

17: 249).

Cerastium Beeringianum Cham. & Schl. in Linnaea 1: 62 (1826); Raup in Jour. Arn.
Arb, 17: 249 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 664 (1944); Porsild in
Nat. Mus. Can. Bull. 101: 19 (1945). See Fernald & Wiegand in Rhod. 22: 172
(1920).

Near Fort Smith, Raup 694; gravelly alder thickets on shore of Brintnell L. at Camp
Pt., 9262; dry gravelly watercourse at base of granite mountain west of Brintnell L., 9480;
mossy banks and shale slide rock in gorge on north slope of Colonel Mt., alt. 3200’-3500’,
9825; Bolstead Cr., Canol Rd., high peaks, alt. 5500’, Wynne-Edwards 8295 (Can.).

Common in the Brintnell L. habitats noted, but not observed elsewhere in that
area; in flower late in June, and collected with both flowers and mature fruit
about mid-August. Our no. 9480 appears to be intermediate between the typical
species and var. grandiflorum (Fenzl) Hultén (see Hultén, 1. c. and Fl. Aleut.
Isles. 165-166). It has the large petals of the variety, but the glandular-hairy
stems of the species.

Cerastium arvense L. Sp. PI. 438 (1753); Raup in Jour. Arn, Arb, 17: 250 (1936);
Porsild in Sargentia 4: 31 (1943); Hultén, Fl. Alaska & Yuk. Pt. 4: 663 (1944).
See Fernald & Wiegand in Rhod. 22: 178 (1920).
This species has not yet been collected in southwestern Mackenzie. It is to
be expected, however, for it is common in the Wood Buffalo Park and has been
collected by Porsild at Great Bear L. and in the Mackenzie delta.

Sagina nodosa (L.) Fenzl, Verbreit, Alsin. 18 (1833); Macoun, Cat. Can. Pl. Pt. 1:
79 (1883); Raup in Jour. Arn. Arb. 17: 250 (1936); Porsild in Sargentia 4: 31
(1943), in Nat. Mus. Can. Bull. 101: 33 (1945).

Spergula nodosa L. Sp. Pl. 440 (1753) ; Hooker, Fl. Bor.-Am. 1: 93 (1830).

South bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8547 (Can.); bank
of Mackenzie R. opposite Simpson, Crickmay 39 (Can.).

This species has also been collected in the Athabaska—Great Slave L. region and
at Great Bear L. It is noted in Fl. Bor.-Am. as occuring from ‘““Upper Canada,
to the shores of the Arctic Sea, eastward of the Mackenzie River.”

Sagina Linnaei Presl, Rel. Haenk. 2: 14 (1831); Hultén, Fl. Alaska & Yuk. Pt. 4:
672 (1944).
Stony shore of Brintnell L. at Camp Pt., 9159.
Apparently rare in this district, and observed only in the shale spot cited
above; found with flowers and immature capsules June 21.

Arenaria lateriflora L. Sp. Pl. 423 (1753); Hooker, Fl. Bor.-Am, 1: 102 (1830); Macoun,
Cat. Can. Pl. Pt. 1: 73 (1883); Raup in Jour. Arn. Arb. 17: 250 (1936).

Moehringia lateriflora (L.) Fenzl; Hultén, Fl. Alaska & Yuk. Pt. 4: 689 (1944).

1947 | RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 171

Fort Smith, Raup 683; Resolution, Miss E. Taylor 51 (Can.) ; northwest shore of Great
Slave L., Bedford (Can.); near Rae, Russell (Can.) ; Mackenzie R., Onion (N) ; Provi-
dence, Preble & Cary 28 (US); upper part of timber on south slope of Red Mt., 9205;
muskeg thickets near west end of Brintnell L., 9436.

Common; in flower during late June and the first half of July. Hooker did

not record this plant north of lat. 60°.

Arenaria nardifolia Ledeb. Fl. Alt. 2: 166 (1830); Porsild in Sargentia 4: 30 (1943).

Mackenzie R. between Blackwater R. and Norman, Crickmay 165 (Can.).

Porsild has collected this species also at Great Bear L., on the arctic coast,
and in the Mackenzie delta.

Hultén apparently considers that the American representative of this group
is A. capillaris Poir. (Encycl. Méth. 6: 380, 1804), which he says is distinguished
from A. nardifolia by the subulate apices of the leaves of its sterile shoots.

Arenaria dawsonensis Britton in Bull. N. Y. Bot. Gard. 2: 169 (1901); Raup in Jour.
Arn. Arb. 17: 250 (1936); Porsild in Sargentia 4: 30 (1943), in Nat. Mus. Can.
Bull. 101: 31 (1945).

Minuartia stricta sensu Hultén, Fl. Alaska & Yuk. Pt. 4: 686 (1944), in part, non Hiern.?
Arenaria stricta sensu Hooker, Fl. Bor.-Am. 1: 99 (1830), in part, non Michx.
Arenaria Michauxii sensu Macoun, Cat. Can. Pl. Pt. 1: 72 (1883), in part, non Hooker.
Stony beach on Brintnell L. at Camp Pt., 9/57; dry sand plain near west end of Brintnell

L., 9452; near summit of Lone Mt., Wynne-Edwards 8467 (Gane

Occasional ; in flower in late June and early July. This species is occasional
to common in the Athabaska—Great Slave L. region, and has been collected at

Great Bear L. by Porsild. Hultén (1. c.) has reduced the Alaskan and Yukon

A. dawsonensis to Minuartia stricta of the Old World, and suggests that our

Mackenzie plants as well as those in northeastern America also represent that

species. He does not, however, present a detailed comparison of American and

Eurasian material, and until this is possible it seems best to maintain A. dawso-

nensis as an American species.

Arenaria verna L. var. pubescens (Cham. & Schl.) Fernald in Rhod. 21: 21 (1919);
Raup in Jour. Arn. Arb, 17: 250 (1936); Porsild in Nat. Mus. Can. Bull. 101:
31 (1945). :

Arenaria hirta var. pubescens Cham. & Schl. in Linnaea 1: 56 (1826).

Arenaria propinqua Richardson; Hooker, FI. Bor.-Am. 1: 99 (1830).

Arenaria verna L. var. hirta (Wormskj.) Wats.; Macoun, Cat. Can. Pl. Pt. 1: 71 (1883).

Minuartia rubella (Wahl.) Graebn.; Hultén, Fl. Alaska & Yuk. Pt. 4: 685 (1944), at

least in part.

Liard R. between Nahanni Butte and Simpson, Crickmay 38 (Can.); dry gravelly fan
at base of granite mountain west of Brintnell L., 9487; shale slopes and rock-slides on south
slope of Red Mt., upper part of scrub timber and at 5000’, 9207, 9756; rock-slides and crev-
ices, north slope of Colonel Mt., alt. 4000’-6000’, 9414, 0654; north peak of Nahanni Mt.,
W ynne-Edwards 8468 (Can.).

Common at Brintnell L.; collected in flower late in June, and with maturing
capsules about mid-July. Noted in FI. Bor.-Am. as occurring on “Barren

grounds, north of lat. 54°, to the Arctic Sea.”

Arenaria Rossii R. Br. ex Richardson, Bot. App. Frankl. Journ. 738 (1823); Hooker,
Fl. Bor.-Am. 1: 100 (1830); Porsild in Sargentia 4: 30 (1943), in Nat. Mus. Can.
Bull. 101: 19 (1945).

Shale crevices and slide rock on north slope of Colonel Mt., alt. 4000’-6000', 9416, 9655;

Plains of Abraham, Canol Rd., mi. 82 E., alt. 6000’, Porsild 11804 (Can.); Bolstead Cr.,

Canol Rd., mi. 111 E., alpine peak, alt. 5500’, Wynne-Edwards 8294 (Gana

172 SARGENTIA [6

Occasional to common at Brintnell L., even on the most unstable rock-slides ;
collected in flower early in July, and with maturing capsules late in July; found
only on Colonel Mt. Porsild states that it is a very common species on limestone
rubble of the higher mountain slopes along the Canol Rd.

Arenaria humifusa Wahl. Fl. Lapp. 129 (1812); Raup in Jour. Arn. Arb. 17: 250
(1936); Porsild in Sargentia 4: 30 (1943); Hultén, Fl. Alaska & Yuk. Pt. 4: 689
(1944); Porsild in Nat. Mus. Can. Bull. 101: 19 (1945). See Nordhagen in
Bergens Mus. Arsb. Nat. Rekke 1: 24 (1935); Polunin in Nature 152: 451, map
(1943).

On mossy banks around an iron spring near west end of Brintnell L., 9238; Plains of
Abraham, Canol Rd., alt. 6000’, in wet limestone rubble, Porsild 11803 (Can.) ; Bolstead Ch
Canol Rd., mi. 111 E., alpine slopes, Wynne-Edwards 8291 (Can.).

Common in the single Brintnell L. habitat noted, but not seen elsewhere in that
area. Porsild states that it is scarce on the Plains of Abraham. In flower late
in June.

Arenaria arctica Stev. ex DC. Prodr. 1: 404 (1824); Porsild in Sargentia 4: 30 (1943),
in Nat. Mus. Can. Bull. 101: 19 (1945).

Minuartia arctica (Stev.) Asch. & Graebn.; Hultén, Fl. Alaska & Yuk. Pt. 4: 678 (1944).

Mackenzie R. between Blackwater R. and Norman, Crickmay 164 (Can.); Plains of
Abraham, Canol Rd., mi, 82 E., Porsild 11805 (Can.) ; Bolstead Cr., Canol Ka, mi. 1115...
alpine slopes, Wynne-Edwards 8287, 8288, 8289, 8290 (Can.).

This species is also known at Great Bear L., in the Richardson Mts., and along
the arctic coast of Yukon (Porsild, 1. c.).

Arenaria macrocarpa Pursh, Fl. Am. Sept. 1: 318 (1814); Porsild in Sargentia 4: 30
(1943), in Nat. Mus. Can. Bull. 101: 19 (1945),

Minuartia macrocarpa (Pursh) Ostenf.; Hultén, Fl. Alaska & Yuk. Pt. 4: 682 (1944).

Alpine slopes south of Little Keele R., Canol Rd., mi. 55 E., alt. 4000’, Porsild 11785
(Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5500’, Wynne-Edwards 8292,
8293 (Can.).

Arenaria uliginosa Schleich. ex Lam. & DC. Fl. Franc. 4: 786 (1805); Porsild in
Sargentia 4: 31 (1943), in Nat. Mus. Can. Bull. 101: 19 (1945).
Sekwi R., Canol Rd., mi. 174 E., alt. 3625’, Porsild 11862 (Can.).
An eastern arctic species previously collected by Porsild at Great Bear L.
and in Keewatin.

Arenaria sajanensis Willd. ex Schlecht. in Berl. Gesell. Nat. Fr. Mag. 7: 200 (1816);
Porsild in Sargentia 4: 30 (1943), in Nat. Mus. Can. Bull. 101: 19 (1945).

Shale crevices on north slope of Colonel Mt., alt. 4000’-5000’, 9652; northeast slope of
Red Mt., alt. about 4500’, 9288, 9289; mossy banks of small brook on west slope of Terrace
Mt., alt. 5000’, 9561; Plains of Abraham, Canol Rd., mi. 82 E., alt. 6000’, Porsild 11806
(Can.).

Occasional at Brintnell L., and noted by Porsild as occasional in Macmillan
Pass, along the Canol Rd. Found flowering in late June, and with mature cap-
sules in the last week of July.

Spergularia marina (L.) Griseb. Spicil. Fl. Rumel. et Bith. 1: 213 (1843); Rossbach
in Rhod. 42: 123 (1940).

Arenaria rubra var. marina L. Sp. Pl. 423 (1753).

Arenaria rubra L. sensu Hooker, Fl. Bor.-Am. 1: 98 (1830).

Spergularia media sensu Macoun, Cat. Can. Pl. Pt. 1: 80 (1883), in part, non Presl.
Spergularia salina J, & C. Presl; Raup in Jour. Arn. Arb. 17: 251 (1936).
Junction of Nyarling and Little Buffalo Rivers, Russell 72 (G).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 173

This species is common at the margins of saline flats and brine springs on
the Salt Plains west of the Slave R. It was noted in Fl. Bor.-Am. “Between
Cumberland House and Bear Lake,” but I have seen no specimens frorn north of
the Little Buffalo River.

Silene acaulis L. var. exscapa (All.) DC. Fl. Franc. 4: 749 (1805); Raup in Jour. Arn.
Arb. 17: 251 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 694 (1944); Porsild in
Nat. Mus. Can. Bull. 101: 19, 31 (1945), in part. See Rhod. 23: 120 (1921).

Silene exscapa All. Fl. Pedem. 2: 83, t. 79, f. 2 (1785).

Silene acaulis L. sensu Hooker, Fl. Bor.-Am. 1: 87 (1830); Macoun, Cat. Can. Pl. Pt.

1: 68 (1883).

Liard R. between Nahanni Butte and Simpson, Crickmay 35 (Can.) ; alpine meadows and
crevices, north slope of Colonel Mt., 9400; north peak of Nahanni Mt., Wsynne-Edwards
8470 (Can.); Bolstead Cr., Canol Rd., mi. 112 E., alpine slope, alt. 5500’, Wynne-Edwards
8284 (Can.).

At Brintnell L. this variety is abundant on the north slope of Colonel Mt.,
but only occasional elsewhere in the area. On some of the steep rocky slopes
above timber line the thick mats formed by it are a primary part of the ground
cover. It also occurs in similar situations in the upper part of the timber.
Flowering specimens collected July 5.

Silene acaulis L. var. subacaulescens (F. N. Williams) Fernald & St. John in Rhod.
23: 120 (1921); Hultén, Fl. Alaska & Yuk. Pt. 4: 696 (1944).

Silene acaulis L. forma subacaulescens F. N. Williams in Jour. Linn, Soc. 32: 101 (1896).
Silene acaulis L.; Porsild in Nat. Mus. Can. Bull. 101: 31 (1945), in part.
North peak of Nahanni Mt., Wynne-Edwards 8469 (Can.).

Silene Menziesii Hooker, Fl. Bor.-Am. 1: 90, t. 30 (1830). See Porsild in Rhod. 40:
212 (1938).

South-facing slope in old field at Simpson, 9126.

Apparently rather rare; just beginning to flower about mid-June. Hooker
states that Richardson collected this species at “Portage d’embarras, Slave River,”
which is one of the portages at Smith Rapids. Until we collected it at Simpson,
however, I had seen no material from the Mackenzie basin.

Lychnis apetala L. Sp. Pl. 437 (1753); Hooker, Fl. Bor.-Am, 1: 91 (1830), at least in
part; Macoun, Cat. Can. Pl. Pt. 1: 68 (1883), in part.
Melandrium apetalum (L.) Fenzl; Porsild in Sargentia 4: 33 (1943), in Nat. Mus. Can.
Bull. 101: 19 (1945).
Melandrium apetalum (L.) Fenzl ssp. arcticum (Fries) Hultén, Fl. Alaska & Yuk. Pt.
4: 700 (1944).
Shale slide rock on north slope of Colonel Mt., alt. 4500’-6000’, 9659; Plains of Abraham,
Canol Rd., mi. 82 E., Porsild 11802 (Can.).
Common on Colonel Mt., though scattered in the unstable slide rock ; capsules
mature in late July. Not observed on the mountains north of Brintnell L.

Lychnis macrosperma (Porsild) comb. nov.
Melandrium macrospermum Porsild in Rhod. 41: 225, t. 552, f. 1-3 (1939) ; Hultén, FI.
Alaska & Yuk. Pt. 4: 703 (1944); Porsild in Nat. Mus. Can. Bull. 101: 19 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes, Wynne-Edwards 8285, 8286 (Can.).
Collected in flower July 25; previously known only in Alaska.
Lychnis brachycalyx sp. nov. Fic. 14.

Herba perennis, caudice brevi erecto vel ascendente, radicibus longis gracilibus ;
stipitibus singillatim ascendentibus non ramosis, 14-18 cm. altis, erectis vel basi

174 SARGENTIA [6

flexuosis, inferne dense breviterque pubescentibus, superne glanduloso-pubescent-
ibus, valde purpurascentibus, nodis leviter incrassatis; foliis 3 vel 4 paribus,
basalibus oblanceolatis vel spathulatis, 2-5 cm. longis, 4-9 mm. latis, fere ad
petiolum basi contractis, apice obtusis, conspicue uninerviis, utrinque saltem
vetustioribus glabris, margine breviter ciliatis; foliis stipitis inferioris oblanceo-

Seen MMM LES HP ron

HE CROOK

Fic. 14. Lychnis brachycalyx Raup, sp. nov.

latis vel anguste ellipticis, 3-4.5 cm. longis, apice obtusis vel acutis, basi leviter
connatis, praeter costam et margines ciliatos glabris vel glabrescentibus; foliis
stipitis superioris reductis, lanceolatis, 8-24 mm. longis, amplexicaulibus vel
leviter connatis, apice obtusis vel acutis, subtus parce hirsutis, margine dense
glanduloso-ciliatis; floribus singulis, calyce inflato late urceolato glanduloso-
pubescente nervis diffusis purpurascentibus ornato, in sicco plerumque 11.5 mm.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 175

crasso et 9 mm. alto, petalis vix exsertis; seminibus in sicco circiter 1 mm, latis,
triangulari-reniformibus, rubro-fuscis, anguste alatis.

Shale cliffs in gorge on north slope of Colonel Mt., Brintnell L. area, alt. about 3500’,
9821 (Type).

Occasional in this situation, but not seen elsewhere; collected with mature cap-
sules Aug. 16. These plants resemble single-flowered forms of L. furcata (Raf.)
Fern. (see Rhod. 34: 22-5, 1932) in habit, leaf-shape, pubescence, and seeds.
The inflated calyces, however, are broader than high, a character which distin-
guishes L. brachycalyx from L. furcata as well as from other arctic species of
Lychnis in this general affinity. In L. furcata, for instance, the calyx is 9-12 mm.
broad and 12-15 mm. high.

NYMPHAEACEAE

Nuphar variegatum Engelm. acc. to Durand in Ann. Rept. Univ. N. Y. 19: 73 (1866);
Porsild in Sargentia 4: 37 (1943), in Nat. Mus. Can. Bull. 101: 19 (1945).

Nuphar luteum sensu Hooker, Fl. Bor.-Am. 1: 32 (1829), in part; Macoun, Cat. Can.
Pl. Pt. 1: 32 (1883), in part; non Smith.

Nymphozanthus variegatus (Engelm.) Fernald; Raup in Jour. Arn. Arb. 17: 251 (1936).

Northwest shore of Great Slave L., Bedford (Can.); Liard R. between Nahanni Butte
and Simpson, Crickmay 39 (Can.); Norman, Coll.? (N).

Common to abundant in the Athabaska-Great Slave L. region, and probably
also throughout the Mackenzie valley. Hooker gives its range as the “.
wooded country between lat. 54° and 64°,” but Porsild reports it also in the
Eskimo Lakes basin. He also notes it as fairly common at Great Bear L.

RANUNCULACEAE

Actaea rubra (Ait.) Willd. Enum. Pl. 561 (1809); Hooker, Fl. Bor.-Am. 1: 27 (1829);
Raup in Jour. Arn. Arb. 17: 252 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 716
(1944),

Actaea spicata L. var. rubra Ait. Hort. Kew. 2: 221 (1789) ; Macoun, Cat. Can. Pl. Pt.

1: 27 (1883).

Fort Smith, Raup 754, 757; Resolution, Preble 215 (US); same place, E. dA. & A. E.
Preble 159 (US); Hay River, Preble & Cary 21 (US); Mackenzie R., Onion, Kennicott
& Hardisty (N); Simpson, Coll.? (N); damp thickets at Simpson, 9844; thickets along
rocky gullies on lower slopes of granite mountain west of Brintnell L., 9804.

Occasional to common in the habitats above noted ; fruit mature at Brintnell L.
about mid-August. Noted in Fl. Bor.-Am. northward only to lat. 60°.

Aquilegia brevistyla Hooker, Fl. Bor.-Am. 1: 24 (1829); Macoun, Cat. Can. Pl. Pt. i
24 (1883); Raup in Jour. Arn. Arb, 17: 252 (1936); Hultén, Fl. Alaska & Yuk.
Pt. 4: 718 (1944).

Windy Pt., Great Slave L., Hume 102662 (Can.); Old Fort Rae, Russell (Can.) ; dry
water course near base of granite mountain west of Brintnell L., 9473; in scrub timber on
south slope of Red Mt., 9212.

Occasional to common in these situations, but not observed elsewhere in the
Brintnell L. area; collected in flower June 21, and with flowers and maturing
capsules in mid-July. Noted in Fl. Bor.-Am. “As far north as Bear Lake.” I
have seen a Richardson specimen from Great Bear L. (Herb. Can.), and one
from Good Hope collected by Miss E. Taylor (Can.).

Delphinium glaucum Wats. Bot. Calif. 2: 427 (1880); Hultén, Fl. Alaska & Yuk. Pt.
4: 721 (1944); Porsild in Nat. Mus. Can. Bull. 101: 19, 31 (1945).

176 SARGENTIA [6

Delphinium scopulorum Gray var. glaucum (Wats.) Gray; Raup in Jour. Arn. Arb. 17:
252 (1936).

Delphinium Brownii Rydb.; Ewan in Univ. Colorado studies, Ser. D. 2: 133 (1945).

Lower Slave R., Russell 75 (G); Resolution, Kennicott (N); east shore of Trout 1
C. O. Hage (G); Mackenzie R., Onion (N); thickets along dry gravelly watercourse,
north slope of Colonel Mt., alt. 5000’, 9661; crater-like hollow on summit of Lone Mt.,
W ynne-Edwards 8471 (Can.) ; Nahanni Mts., Preble & Cary 78 (US); Bolstead Cr., Canol
Rd., mi. 111 E., Wynne-Edwards 8303 (Can.) ; Macmillan Pass, Canol Rd., mi. 225 E.,
Porsild 11235 (Can.).

Common along watercourses above timber line in the mountains; in flower in
the latter part of July.

Ewan, in his recent “Synopsis of the North American species of Delphinium”
(1. c.), maintains D. Brownii Rydb. as a species separate from D. glaucum. He
gives the former a continental range in Alaska, Yukon, British Columbia, Al-
berta, and Wyoming, while the latter extends northward from California to
western Washington. On page 67 of his monograph he states that they are
distinct, though related; and on page 134 he cites photographic illustrations to
show “the greater sturdiness of this interior species [D. Brownii] over D. glau-
cum . . . the stout rachises and crisp succulence of the leaves is notable.” On
page 223 there is a plate in which representatives of the two species are shown
side by side, with the following note in the legend under D. glaucum: “contrast
raceme and leaves with D. Brownii with which this species has been generally
confused.”

Ewan’s keys, however, are arranged in sections which are determined geograph-
ically, and he does not give a general key to all the species of Delphinium. Fur-
thermore, D. glaucum and D. Brownii do not both appear in any one section, so
that nowhere are they distinguished from one another in the keys. A careful
comparison of his somewhat detailed descriptions of the two (pp. 133, 134) also
fails to reveal any significant differences, and the two plants shown in his plate
appear to show no differences greater than those which commonly occur within
either “species.”

In the absence, therefore, of any clear distinctions I am continuing to consider
D. Brownii as identical with Watson’s D. glaucum, originally described from
California. Ewan is probably correct in maintaining that D. glaucum should not
be related to D. scopulorum. There seems to be nothing inconsistent about the
combined range patterns of Ewan’s two entities as shown by such groups of
plants as are mapped in my Figures 9-a and 10.

Aconitum delphinifolium DC. Syst. Nat. 1: 380 (1818); Hultén, Fl. Alaska & Yuk. Pt.
4: 722 (1944); Porsild in Nat. Mus. Can. Bull. 101: 19 (1945).

Nahanni Butte, Liard R., alt. 3800’, Whittaker 100158 (Can.); Liard R. between Nahanni
Butte and Simpson, Crickmay 40 (Can.) ; gravelly thicket in swampy ground near west end
of Brintnell L., 9432; high alpine rock-slides, north slope of Colonel Mt., 9418; scrub timber
on south slope of Red Mt., 9210; northeast slope of Red Mt., alt. 4500’-5000’, 9275; mossy
bank of brook on west slope of Terrace Mt., alt. 4800’, 9542; Bolstead Cr., Canol Rd., mi.
111 E., alpine bog, alt. 4700’, Wynne-Edwards 8303 (Can.).

Common at Brintnell L.; found flowering in late June and the first half of
July. Wynne-Edwards collected flowering specimens on July 26. Noted by
Porsild at Sekwi R. and Macmillan Pass, along the Canol Rd. The flowers of
our no. 9542 are yellowish white, apparently representing an albino form of the
species. White-flowered individuals have also been observed in northern British

Columbia by Mrs. Mary G. Henry (Nat. Hort. Mag. 13: 271, 1934).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 177

Anemone parviflora Michx. Fl. Bor.-Am. 1: 319 (1803); Hooker, Fl. Bor.-Am. 1: 5
(1829); Macoun, Cat. Can. Pl. Pt. 1: 12 (1883); Raup in Jour. Arn. Arb. 17: 252
(1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 736 (1944); Porsild in Nat. Mus. Can.
Bull. 101: 19, 31 (1945).

Fort Smith, Miss E. Taylor 955 (Can.); Pine Pt., Great Slave L., Brooke (Can.) ;
Mackenzie R., Onion (N); Liard R. between Nahanni Butte and Simpson, Crickmay 44
(Can.) ; mossy banks and muskeg thickets at west end of Brintnell L., 9242; gravelly thickets
and mossy shores, south side of Brintnell L., 9337, 9695; wet ground along snow-fed stream,
north slope of Colonel Mt., alt. 5000’, 9388; damp hollow on Lone Mt., Wynne-Edwards
8473, 8474 (Can.) ; north peak of Nahanni Mt., Wynne-Edwards 8478 (Can.) ; Bolstead Cr.,
Canol Rd., mi. 111 E., high alpine slopes, alt. 4400’-5500', Wynne-Edwards 8301, 8302 (Can.).

Occasional to common; or abundant in wet alpine meadows; mostly in fruit
at low levels late in June, but occasional flowers found later in the summer ;
in flower at high elevations early in July. Wynne-Edwards collected flowering
specimens at Bolstead Cr. as late as July 24 and 25. Porsild notes it as only
occasional in Macmillan Pass. The plants collected around the shores of Brint-
nell L. are notable for their tall habit of growth (up to 35 cm.) and their large
fruiting heads (1.5 cm. high).

Anemone multifida Poir. var. hudsoniana DC. Syst. 1: 209 (1817); Raup in Jour. Arn.
Arb. 17: 253 (1936); Porsild in Nat Mus. Can. Bull. 101: 31 (1945). See Fernald
in Rhod. 19: 140 (1917).

Anemone multifida Poir. sensu Hooker, Fl. Bor-Am. 1: 7 (1829); Hultén, Fl. Alaska
& Yuk. Pt. 4: 730 (1944).

Fort Smith, Miss E. Taylor 48 (Can.) ; same place, Seton & Preble 78313 (Can.) ; same
place, Raup 774; Resolution, Kennicott (N); Windy Pt., Great Slave L., Hume 102663
(Can.) ; Old Fort Rae, Russell (Can.) ; Providence, Preble & Cary 36 (US); same place,
Dutilly 87 (G); Liard R. between Nahanni Butte and Simpson, Crickmay 43 ian.)
Simpson, Onion, Kennicott & Hardisty (N); south-facing slope in old field at Simpson,
9127; dry dolomitic summit of Lone Mt., Wynne-Edwards 8472 (Can.).

Common at Simpson; in flower June 13. Noted in Fl. Bor.-Am. . to
near the shores of the Arctic sea: common.” I have seen specimens from Great
Bear L. (J. M. Bell, Can.) and the lower Mackenzie (Miss E. Taylor, Can.).

Anemone Richardsonii Hooker, Fl. Bor.-Am. 1: 6, t. 4 (1829); Macoun, Cat. Can. Pl.
Pt. 1: 13 (1883); Raup in Jour. Arn. Arb. 17: 253 (1936); Hultén, Fl. Alaska &
Yuk. Pt. 4: 740 (1944); Porsild in Nat. Mus. Can. Bull. 101: 19 (1945).

Wrigley, Miss E. Taylor (G); rich woods on lower south slope of Terrace Mt., alt.
3000’, 9684; wet ground along snow-fed stream, north slope of Colonel Mt., alt. about 5000’,
9389; along stream in upper valley of Frost Cr., alt. 4000’, 9276; along small brook on west
slope of Terrace Mt., alt. 5000’, 9559.

Common to abundant in wet places along streams, particularly at high eleva-
tions; in flower late in June. The FI. Bor.-Am. record for this species is “. . .
barren ground, Rocky Mountains, from lat. 55° to 68°, in wet mossy ground.”
Porsild noted it as common in Macmillan Pass, and I have seen a specimen from
Peel R.

Anemone canadensis L. Syst. ed. 12, 3: App. 231 (1768); Raup in Jour. Arn. Arb. 17:
253 (1936). .

Anemone pennsylvanica L.; Hooker, Fl. Bor.-Am. 1: 8, t. 3, B (1829).

Anemone dichotoma L.; Macoun, Cat. Can. Pl. Pt. 1: 13 (1883).

South bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8548 (Can.) ; Liard
R. between Nahanni Butte and Simpson, Crickmay 42 (Can.); low place in old field at
Simpson, 9132.

Common in the Simpson habitat ; not yet in flower when collected on June 13.
Noted in Fl. Bor. Am. “. . . from the United States to near the mouth of Mack-

ce
.

178 SARGENTIA [6

enzie’s River ; not found in the barren grounds.” Common in the Wood Buffalo
Park, but I have seen no material from north of Simpson. There is a Richard-
son specimen in Herb. G marked “Arctic Nt". America,” but without specific
locality data.

Anemone narcissiflora L. Sp. Pl. 542 (1753); Porsild in Sargentia 4: 38 (1943), in Nat.
Mus. Can. Bull. 101: 19 (1945).

Anemone narcissiflora L. ssp. villosissima (DC.) Hultén, Fl. Alaska & Yuk. Pt. 4: 732
(1944),

Alpine meadows, east slope of Red Mt., alt. 4500’-5500’, 9217; meadow in dwarf birch
scrub on southwest slope of Terrace Mt., alt. 4500’, 9685.

Common to abundant in high alpine meadows at Brintnell L., and noted by
Porsild in Macmillan Pass; in flower in the latter part of June, and in fruit
early in August. Hultén has divided the American representatives of A. narcissi-
flora into four subspecies based upon the shape and cutting of leaves, the number
of flowers per stem, and relative hairiness. All of these are notoriously fickle
characters, and Hultén admits that the lines of demarkation are not clear. On
the basis of geographic position our plants should represent his ssp. interior, but
when keyed out they fall rather into his ssp. villosissima.

Pulsatilla ludoviciana (Nutt.) Heller, Cat. N. Am. Pl. ed. 2, 4 (1900); Raup in Jour.
Arn, Arb, 17: 253 (1936); Porsild in Nat. Mus. Can. Bull. 101: 20 (1945).

Anemone ludoviciana Nutt. Gen. Am. Pl. 2: 20 (1818).

Anemone patens sensu Hooker, Fl. Bor.-Am. 1: 4 (1829), non L.

Anemone patens L. var. Nuttalliana Gray; Macoun, Cat. Can. Pl. Pt. 1: 12 (1883).

Anemone patens L. ssp. multifida (Pritzel) Zamels; Hultén, Fl. Alaska & Yuk. Pt. 4:

_ 738 (1944).

Fort Smith, Seton & Preble 78314 (Can.); same place, Raup 766, 2403; Mackenzie R.,
Onion, Kennicott & Hardisty (N); Liard R. between Nahanni Butte and Simpson, Crick-
may 41 (Can.).

Common in the central Mackenzie basin, and probably also far northward.
It was noted in Fl. Bor.-Am. as occurring “. . . in the central limestone tracts
from lat. 45° to 67° on the Mackenzie.” I have seen specimens from Great Bear
L. (J. M. Bell, Can.) and the lower Mackenzie (McConnell, Jones, Kennicott,
Can.). Porsild noted it in Macmillan Pass.

Ranunculus trichophyllus Chaix var. typicus Drew in Rhod. 38: 18 (1936); Raup in
Jour. Arn. Arb. 17: 254 (1936); Porsild in Sargentia 4: 39 (1943); Hultén, FI.
Alaska & Yuk. Pt. 4: 770 (1944).

Ranunculus aquatilis L.; Hooker, Fl. Bor.-Am. 1: 10 (1829), in part.

Ranunculus aquatilis L. var. capillaceus DC. sensu Raup in Nat. Mus. Can. Bull. 74: 134

(1935).

Little Buffalo R., Russell 91 (G); northwest shore of Great Slave L., Bedford (Can.).

Common in the central Mackenzie basin and probably also far northward
through the Mackenzie valley. Like most aquatics, it has been very little col-
lected. Porsild has found it in the Mackenzie delta and in the Eskimo Lakes
basin. In FI. Bor.-Am. it is noted as extending northward ‘. . . to near the

Arctic Sea, lat. 68°,” but part of the material upon which this range was based

probably represents other varieties of FR. trichophyllus (See Porsild, 1. c.).

Ranunculus Gmelini DC. var. Purshii (Richardson) Hara in Rhodora 41: 386 (1939).

Ranunculus Purshi Richardson, Bot. App. Frankl. Journ. ed. 1, 741 (1823) ; Hooker, FI.
Bor.-Am. 1: 15 (1829); Raup in Jour. Arn. Arb. 17: 254 (1936); Hultén, Fl. Alaska
& Yuk. Pt. 4: 765 (1944).

Ranunculus multifidus sensu Macoun, Cat. Can. Pl. Pt. 1: 16 (1883), non DC.

1947] RAUP, THE BOTANY OF SOUTHWESTERN ‘MACKENZIE 179

Ranunculus Gmelinii DC. var. terrestris (Ledeb.) Benson in Bull. Torr. Bot. Cl. 69:
313 (1942).

Smith Portage, Miss E. Taylor 1109 (Can.); Resolution, Miss E. Taylor 22 (G);
northwest shore of Great Slave L., Bedford (Can.); Mackenzie R., Coll.? (N); Liard R.
between Nahanni Butte and Simpson, Crickmay 46 (Can.) ; swamp in wet woods, Simpson,
9144; old stream channel in mud flats at west end of Brintnell L., 9709, 9805.

Occasional to common; in flower at Brintnell L. about mid-August, and at
Simpson about mid-June. Noted in Fl. Bor.-Am. “. . . from Canada to near the
Arctic Sea.”

Ranunculus hyperboreus Rottb. in Acta Univ. Hafn. 10: 458 (1770); Raup in Jour.
Arn. Arb, 17: 254 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 757 (1944).

Pools in wet thickets at west end of Brintnell L., 9719.

Occasional in the above habitat, and apparently rare in the Mackenzie basin.
I have seen only one other specimen, from the Wood Buffalo Park (Raup 2368).
Collected with maturing achenes early in August at Brintnell L.

Ranunculus reptans L. Sp. Pl. 549 (1753); Raup in Jour. Arn. Arb. 17: 254 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 4: 767 (1944); Porsild in Nat. Mus. Can. Bull.
101: 33 (1945).
Ranunculus Flammula L. var. major and var. intermedia Hooker, Fl. Bor.-Am. 1: 11
(1829).
Ranunculus Flammula L. var. reptans (L.) Meyer; Macoun, Cat. Can. Pl. Pt. 1: 17
(1883).

Northwest. shore of Great Slave L., Bedford (Can.); Lake Grant, between Great Slave
and Great Bear Lakes, Preble 248 (US); south bank of Mackenzie R., 10 mi. below Mills
L., Wynne-Edwards 8550 (Can.).

Occasional to common in the central Mackenzie basin, and probably also in the
upper Mackenzie valley. Hooker notes it as occurring “On the gravelly banks of
rivers from Canada to lat. 69°.” I have seen no specimens, however, from the
country north of the areas cited above.

Ranunculus pygmaeus Wahl. Fl. Lapp. 157 (1812); Hultén, Fl. Alaska & Yuk. Pt. 4:
765 (1944); Porsild in Nat. Mus. Can. Bull. 101: 20 (1945).
Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes, Wynne-Edwards 8300 (Can.) ; Mac-
millan Pass, Canol Rd., mi. 225 E., Porsild 11234 (Can.).
Not otherwise known in the Mackenzie basin except in the mountains of north-
ern British Columbia.

Ranunculus nivalis L. Sp. Pl. 553 (1753); Hultén, Fl. Alaska & Yuk. Pt. 4: 760 (1944);
Porsild in Nat. Mus. Can. Bull. 101: 20 (1945).

Along small brook on west slope of Terrace Mt., alt. 4800’, 9562; Bolstead Cr., Canol
Rd., mi. 111 E., alpine slopes, alt. 6000’, Wynne-Edwards 8299 (Can.); Macmillan Pass,
Canol Rd., mi. 225 E., Porsild 11233 (Can.).

Apparently rare in the Brintnell L. area, but more common along the Canol
Rd.; in flower in mid-July.

Ranunculus Eschscholtzii Schl. Animadv. Bot. Ranunc. Candollii 2: 16, ¢. 1 (1820) ;
Hultén, Fl. Alaska & Yuk. Pt. 4: 751 (1944); Porsild in Nat. Mus. Can. Bull.
101: 20 (1945).

Wet moss along small brook on west slope of Terrace Mt., alt. 4800'—5000', 9539.

Occasional in this situation, but not seen elsewhere in the Brintnell L. region ;
in flower about mid-July. Noted by Porsild as common in Macmillan Pass.

Ranunculus lapponicus L. Sp. Pl. 553 (1753); Hooker, Fl. Bor.-Am. 1: 16 (1829);

Macoun, Cat. Can. Pl. Pt. 1: 19 (1883); Raup in Jour. Arn. Arb. 17: 254 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 4: 758 (1944).

180 SARGENTIA [6

Mossy muskeg thickets at west end of Brintnell Ts 09237579712:
Common in the habitat noted above; in flower late in June, and with mature
fruit early in August. Noted in Fl. Bor.-Am. “. . . from lat. 50° to the Arctic

2F

sea.” I have seen a specimen from Arctic Red River (Dutilly, G).

Ranunculus sceleratus L. var. multifidus Nutt. in Torr. & Gray, Fl. N. Am. 1:,19
(1838). See Benson in Bull. Torr. Bot. Cl. 69: 312 (1942).

Ranunculus sceleratus L. sensu Hooker, Fl. Bor.-Am. 1: 15 (1829) ; Macoun, Cat. Can.
Pl. Pt. 1: 19 (1883) ; Raup in Jour. Arn. Arb. 17: 255 (1936).
Ranunculus sceleratus L. ssp. multifidus (Nutt.) Hultén, Fl. Alaska & Yuk. Pt. 4: 768

(1944).
Northwest shore of Great Slave L., Bedford (Can.).
Noted in FI. Bor.-Am. as extending “. . . from Canada to lat. 67°,” but I have

seen no material from below Great Slave L.

Ranunculus Macounii Britton in Trans. N. Y. Acad. Sci. 12: 3 (1892); Raup in Jour.
Arn, Arb. 17: 255 (1936); Hultén, Fl. Alaska & Yuk. Pt. 4: 759 (1944); Porsild

in Nat. Mus. Can. Bull. 101: 33 (1945).
Ranunculus hispidus Michx. sensu Hooker, Fl. Bor.-Am. 1: 19 (1829), at least in part.

Ranunculus repens var. hispidus (Michx.) T. & G., sensu Macoun, Cat. Can. Pl. Pt. 1:
21 (1883), at least in part.

Forth Smith, Raup 741; northwest shore of Great Slave L., Bedford (Can.) +; south bank
of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8549 (Can.) ; Liard R. between
Nahanni Butte and Simpson, Crickmay 45 (Can.) ; Simpson, Coll.? (N); damp thicket
along wood road just west of Simpson, 9881; damp place in upland prairie west of the
Mackenzie R. near Simpson, 9914.

Occasional northward in the Mackenzie valley at least as far as Simpson.
Noted in Fl. Bor.-Am. as occurring on “Banks of rivers from Canada to near the
mouth of the Mackenzie River, lat. 67°.” Hooker states that R. pennsylvanicus
L. is “found with the last mentioned species [R. hispidus = R. Macounii|, and it
is equally diffused. . . .” I have seen no evidence of R., pennsylvanica, however,
from north of the Athabaska—Peace delta region.

Ranunculus acris L. Sp. Pl. 554 (1753); Raup in Jour. Arn. Arb. 17: 255 (1936).
Fort Smith, Russell 30 (G).
Common as a weed around settlements in the central Mackenzie basin north-
ward at least to Fort Smith, and probably also along the upper Mackenzie,

Ranunculus Cymbalaria Pursh, Fl. Am. Sept. 2: 392 (1814); Hooker, Fl. Bor.-Am. 1:
11 (1829); Macoun, Cat. Can. Pl. Pt. 1: 17 (1883); Raup in Jour. Arn. Arb. 17:
256 (1936).

Fort Smith, Raup 733; Little Buffalo R., Russell 65 (G).

I have seen no authentic material from north of the Little Buffalo R., though
there is a specimen in Herb. N collected by Onion, Kennicott & Hardisty which
may have come from the Mackenzie. The species is noted in Fl. Bor.-Am. as
occurring “. . . from Canada to near the Arctic Sea, lat. 68°. . .”

Thalictrum alpinum L. Sp. Pl. 545 (1753); Hultén, Fl. Alaska & Yuk. Pt. 4: 773 (1944) ;
Porsild in Nat. Mus. Can. Bull. 101: 20, 31 (1945).
North Peak of Nahanni Mt., Wynne-Edwards 8476 (Can.) ; Bolstead Cr., Canol Rd., mi.
111 E., high alpine slopes, alt. 5500’, Wynne-Edwards 8298 (Can.).
Porsild noted it as common in alpine tundra near the eastern end of Macmillan

Pass, Canol Rd., mi. 220 E.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 181

Thalictrum occidentale Gray in Proc. Am. Acad. 8: 372 (1873).

Liard R. between Nahanni Butte and Simpson, Crickmay 47 (Can.); fence row by
plowed field at Simpson, 9133, 9134.

Common in the Simpson habitat ; in flower about mid-June. These specimens,
representing both staminate and pistillate plants, are too immature for definite
determination. They closely resemble 7. occidentale, however, in habit, leaves,

and flowers. I am unable also to place them in one of the varieties of T. oc-
cidentale recognized by Boivin (Rhod. 46: 454-457, 1944).

Thalictrum venulosum Trel. in Proc. Bost. Soc. Nat. Hist. 23: 302 (1886); Raup in
Jour. Arn. Arb. 17: 256 (1936); Boivin in Rhod. 46: 439 (1944); Porsild in Nat.
Mus. Can. Bull. 101: 33 (1945).

Thalictrum dioicum sensu Hooker, Fl. Bor.-Am. 1: 3 (1829), in part; Macoun, Cat. Can.
Pl. Pt. 1: 14 (1883), in part; non L.

North shove of Great Slave L., Hume 102664 (Can.); south bank of Mackenzie R., 10
mi. below Mills L., Wynne-Edwards 8551 (Can.).

In Fl. Bor.-Am. T. dioicum is said to range “. . . from Canada to the banks of
the Mackenzie R., in lat. 67°.” That this may involve some other species than
T. venulosum is suggested by the tentative determination of our Simpson speci-
mens as T. occidentale,

Caltha palustris L. Sp. Pl. 558 (1753); Raup in Jour. Arn. Arb. 17: 256 (1936).

Caltha palustris L. var. arctica (R. Br.) Hutch.; Hultén, Fl. Alaska & Yuk. Pt. 4: 711

(1944).

Near Pine Pt., Great Slave L., Brooke (Can.) ; near Big Island, Great Slave L., Onion,
Kennicott & Hardisty (N); Wrigley Harbor, Great Slave L., Kindle (Can.).

Hultén maintains, perhaps rightly, that our northern American plants do not
exactly match the typical European. He takes up for them the name var. arctica,
based upon a much dwarfed plant from Melville Island described by Robert
Brown. He considers this to be only an extremely reduced form of the Ameri-
can plant.

PAPAVERACEAE
Papaver radicatum Rottb. in Skr. Kjgbenh. Selsk. Laerd. Vidensk. 10: 455, t. 8 (1770);
Porsild in Sargentia 4: 39 (1943); Hultén, Fl. Alaska & Yuk. Pt. 5: 805 (1945).
Mackenzie R. between Blackwater R. and Norman, Crickmay 167 (Can.).
Noted by Porsild at Great Bear L.

Papaver microcarpum DC. Syst. Veg. 2: 71 (1821); Porsild in Sargentia 4: 39 (1943).

This species was collected at Norman on the Mackenzie by Hume (103443,
Can.), just at the northern boundary of our area. Porsild reports it in the Rich-
ardson Mts., and on the arctic coast west of the Mackenzie.

Papaver Keelei A. E. Porsild in Nat. Mus. Can. Bull. 101: 20 (1945).

Canol Rd., wet limestone rubble, Plains of Abraham, alt. 6000’, mi. 82 E., Porstld 11807
(Can.) ; small tributary to Little Keele R., mi. 51 E., alt. 2200’, Porsild 11782 (Can., TYPE
COLL.) ; wet alpine meadow, Bolstead Cr., mi. 111 E., alt. 5500’, Wynne-Edwards 8305, 8307
(Can.) ; same place, alpine peak, alt. 7000’, Wynne-Edwards 8306.

Porsild states that this species is common in places, apparently restricted to
calcareous soils and preferring moist, springy places. Number 8306 was found in
flower July 26, and the others were collected later and were in fruit.

182 SARGENTIA [6

Corydalis pauciflora (Steph.) Pers. Syn. Pl. 2: 269 (1807); Hultén, Fl. Alaska & Yuk.
Pt. 5: 810 (1945).

Fumaria pauciflora Steph. in Willd. Sp. Pl. 3: 861 (1800).

Mossy banks of a small brook on west slope of Terrace Mt., alt. 4800’-5500', 9526.

Common in this place but not seen elsewhere in the Brintnell L. area. Col-
lected with flowers and maturing capsules July 18.

Corydalis sempervirens (L.) Pers. Syn. Pl. 2: 269 (1807); Raup in Jour. Arn. Arb. 17:
256 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 812 (1945).
Fumaria sempervirens L. Sp. Pl. 700 (1753).
Corydalis glauca Pursh; Hooker, Fl. Bor.-Am. 1: 37 (1829) ; Macoun, Cat. Can. Pl. Pt.
1: 36 (1883).
Southwest and north shores of Great Slave L., Howe 91980 (Can.) ; northwest shore of
Great Slave L., Bedford (Can.) ; east shore of Trout L., C. O, Hage (G); Mackenzie R.
between Wrigley and Blackwater R., Crickmay 133 (Can.).

Noted in Fl. Bor.-Am. as occurring “Throughout Canada, as far as lat, 64°.”

Corydalis aurea Willd. Enum. Pl. Hort. Berol. 2: 740 (1809); Hooker, Fl. Bor.-Am. 1:
37 (1829); Macoun, Cat. Can. Pl. Pt. 1: 36 (1883); Raup in Jour. Arn. Arb. 17:
256 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 809 (1945).

Lower Slave R., Raup 781; Resolution, Kennicott (N) ; Providence, Dutilly 63 (G);
Simpson, Onion, Kennicott & Hardisty (N); sandy bank of Mackenzie R. opposite Simpson,
9947; Mackenzie R. between Wrigley and Blackwater R., Crickmay 134 (Can.).

Occasional ; collected with flowers and mature capsules at Simpson late in

August. Noted in Fl. Bor-Am. “Throughout Canada to lat. 64°. . . in the
woody country.”

CRUCIFERAE
Draba alpina L. Sp. Pl. 642 (1753); Hooker, Fl. Bor.-Am. 1: 50 (1830); Macoun, Cat.
Can. Pl. Pt. 1: 49 (1883); Hultén, Fl. Alaska & Yuk. Pt. 5: 846 (1945).
Liard R. between Nahanni Butte and Simpson, Crickmay 48 (Can.).

Draba alpina L. var. nana Hooker in Trans. Linn. Soc, 14: 363 (1825). See Fernald
in Rhod. 36: 285 (1934).

Draba Bellii Holm; Porsild in Nat. Mus. Can. Bull. 101: 20 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11811 (Can.) ; high ridge east of pump
station at Bolstead Cr., Carol Rd., mi. 111 E., alt. 6000’, Wynne-Edwards 8314 (Can.) ;
same locality, west slope of valley, Waynne-Edwards 8312, 8313 (Can.).

Draba fladnizensis Wulf. var. heterotricha (Lindblom) Ball in Bull, Soc. Bot. Fr. 7:
230 (1860). See Fernald in Rhod. 36: 286 (1934).
Draba lactea Adams.; Hultén, Fl. Alaska & Yuk. Pt. 5: 857 (1945).

South slope of Red Mt., north of Brintnell L., near summit, alt. about 5900’, 9203; shale
cliffs above timber line on north slope of Colonel Mt., 9620.

Occasional ; collected in flower June 21 and in fruit July 28.

Draba longipes Raup in Contr. Arn. Arb. 6: 165, pl. 6 (1934); Hitche. in Univ. Wash.
Publ. Bot. 11: 25 (1941); Hultén, Fl. Alaska & Yuk. Pt. 5: 858 (1945).

Gravelly beach on south shore of Brintnell L., 9336; dry gravelly watercourse on north
slope of Colonel Mt., alt. 5000’, 9616, 9617; alpine meadow along brook, west slope of Ter-
race Mt., alt. 5000’, 9548.

Common in the alpine habitats but rare at lake level; siliques mature in the
latter half of July.

Draba nivalis Liljebl. var. elongata Wats. Proc. Am. Acad, 23: 258, f. 5le (1888). See
Hitche, in Univ. Wash. Publ. Bot. 11: 84 (1941).

Shale crevices and slide rock, south slope of Red Mt., alt. 5000’-5500’, 9785.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 183

Apparently rare or only occasional in our region; found with shattered
siliques early in August. This material appears rather intermediate between D.
nivalis and its var. elongata (= D. lonchocarpa Rydb.). Its siliques are variable
in length from those similar to D. nivalis to those of the variety.

Draba incerta Payson in Am. Jour. Bot. 4: 261 (1917); Porsild in Sargentia 4: 42
(1943).
Liard R. between Nahanni Butte and Simpson, Crickmay 49 (Can.).

Noted at Great Bear L. by Porsild (1. c.).

Draba aurea M. Vahl in Hornem. Fors. Oecan. Plantel. ed. 2: 599 (1806); Raup in
Jour. Arn. Arb. 17: 257 (1936); Porsild in Sargentia 4: 41 (1943); Hultén, FI.
Alaska & Yuk. Pt. 5: 847 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 50 (Can.).

The Crickmay specimen is the only one yet available from southwestern Mack-
enzie. The species has been collected at L. Athabaska, and doubtful specimens
have been collected by Porsild in the Mackenzie delta.

Draba glabella Pursh, Fl. Am. Sept. 2: 434 (1814); Fernald in Rhod. 36: 333 (1934);
Raup in Jour. Arn. Arb..17: 257 (1936); Porsild in Nat. Mus. Can. Bull. 101:
20 (1945).
Draba hirta L. sensu Hultén, Fl. Alaska & Yuk. Pt. 5: 853 (1945), in part.
Shale slopes on south side of Red Mt., alt. 5000’, 9786, 9786-A; Bolstead Cr., Canol Rd.,
mi. 111 E., in alpine bog, alt. 5000’, Wynne-Edwards 8310, 8311 (Can.).

Occasional ; siliques mature early in August.

Draba praealta Greene, Pittonia 3: 306 (1898); Porsild in Sargentia 4: 43 (1943).
Dry sandy plain near west end of Brintnell L., 9459; shale cliffs in gorge on north slope
of Colonel Mt., alt. 3500’, 9829; alpine rock-slide, north slope of Colonel Mt., 9405-A.
Occasional to common; mature siliques collected from mid-July to mid-August.
Found at Great Bear L. by Porsild.

Draba McCallae Rydb. in Bull. Torr. Bot. Cl. 29: 241 (1902). See Fernald in Rhod.
36: 341-342 (1934); Hitche. in Univ. Wash. Publ. Bot. 11: 26 (1941).

Draba aurea M. Vahl, sensu Hitchce., |. c., in part.

Thicket on gravelly shore of Brintnell L. at Camp Pt., 9145; along small brook on west
slope of Terrace Mt., alt. 4500’-5000’, 9514.

Occasional; collected with flowers and maturing siliques at lake-level on June
20, and with mature fruit at the higher altitudes about mid-July.

Draba lanceolata Royle, Illustr. Bot. Himal. Mts. 1: 72 (1839); Fernald in Rhod. 36:
357 (1934); Raup in Jour. Arn. Arb. 17: 257 (1936); Hultén, Fl. Alaska & Yuk.
Pt. 5: 857 (1945).

Alpine rock-slide, north slope of Colonel Mt., 9405; shale cliffs in gorge on north slope
of Colonel Mt., alt. 3500’, 9818; shale crevices and rock-slides, north slope of Colonel Mt.,
alt. 5000’-6000’, 9618.

Occasional to common, sometimes inhabiting very loose slide rock; collected
with mature siliques in late July and the first half of August.

Nos. 9818 and 9618 have the habit of growth, pubescence, and commonly the
twisted pods of typical D. lanceolata, but they differ from the latter in having the
pods quite glabrous.

Records in Fl. Bor.-Am. for Draba confusa and D. incana in the Mackenzie
basin may be referable to D. lanceolata. The first is noted as occurring “. . .
north in the plains, nearly to the mouth of the Mackenzie River,” and the second
at “Bear Lake and the Mackenzie River.”

184 SARGENTIA [6

Draba cinerea Adams. in Mem. Soc. Nat. Mosc. 5: 103 (1817); Raup in Jour. Arn.
Arb. 17: 257 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 849 (1945),

Shale slide rock, north slope of Colonel Mt., alt. 5000’-6000’, 9609.
Occasional in this type of habitat; collected with mature siliques July 28.
Also known in the Athabaska—Great Slave L. district.

Draba nemorosa L. Sp. Pl. 643 (1753); Macoun, Cat. Can. Pl. Pt. 1: 52 (1883); Raup
in Jour. Arn. Arb. 17: 257 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 861 (1945).
See Fernald in Rhod. 36: 365 (1934); Hitchce. in Univ. Wash. Publ. Bot. 11:
109 (1941).

Fort Smith, Raup 808; same place, Seton & Preble 78316 (Can.) ; Resolution, Kennicott
(N); cleared land on river bluff at Simpson, 9115,

Common weed around settlements. Collected at Simpson with flowers and
maturing siliques June 12. Both the typical form and the glabrous-fruited var.
lejocarpa Lindbl. (Linnaea 13: 33, 1839) are found in the Mackenzie basin,
though the Simpson plants all represent the variety. The latter, under D. lutea
Gilib., is noted in FI. Bor.-Am. as extending “. . . from lat. 56° to 66°... .”

Lesquerella arctica (Wormsk.) Wats. in Proc. Am. Acad. 23: 254 (1888); Porsild in
Nat. Mus. Can. Bull. 101: 33 (1945); Hultén, Fl. Alaska & Yuk. Pt. 5: 841 (1945).

Alyssum arcticum Wormsk. in Fl. Dan. 9, Fasc. 26, #. 1520, p. 3 (1814).

Vesicaria arctica (Wormsk.) Richardson; Hooker, Fl. Bor.-Am. 1: 48 (1829) ; Macoun,
Cat. Can. Pl. Pt. 1: 54 (1883).

Outwash gravel plain near mouth of North Nahanni R., Wynne-Edwards 8536 (Can.).

Noted in Fl. Bor.-Am. “. . . on the summit of a hill 700 feet high, at Bear
Lake River.”

Lepidium apetalum Willd. Sp. Pl. 3: 439 (1801); Raup in Jour. Arn, Arb. 17: 257
(1936).

Lepidium ruderale sensu Hooker, Fl. Bor.-Am. 1: 68 (1830), in part, non L.

Lepidium densiflorum Schrad. of auth., in part.

Mackenzie R., Kennicott (N).

Noted in Fl. Bor.-Am. as occurring “. . . from lat. 50° ‘to 68°.” There is a
Hooker specimen, presumably collected during Franklin's Journeys, in Herb.
N, but it has no locality data.

Capsella Bursa-pastoris (L.) Medic. PA. Gatt. 1: 85 (1792); Hooker, Fl. Bor.-Am. 1:
69 (1830); Raup in Jour. Arn. Arb, 17: 257 (1936); Hultén, Fl. Alaska & Yuk.
Pt. 5: 842 (1945).

Fort Smith, Raup 797; Resolution, Preble 213 (US); cleared land on river bluff at
Simpson, 97/4.

A common weed around settlements ; collected with flowers and siliques about
mid-June.
Neslia paniculata (L.) Desv. in Jour. Bot. 3: 162 (1814); Raup, Jour. Arn. Arb. 17:
258 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 843 (1945).
Cultivated field west of the Mackenzie R. near Simpson, 9924,
Occasional as a weed around settlements; siliques mature in late August.
Brassica Kaber (DC.) Wheeler var. pinnatifida (Stokes) Wheeler in Rhod. 40: 308
(1938).
Sinapis arvensis L. 8 pinnatifida Stokes, Bot.- Mater. Med. 3: 478 (1812).
Cultivated field on west bank of Mackenzie R. near Simpson, 9929,
Occasional as a weed in fields; collected with flowers and maturing fruit the
last week in August.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 185

Thellungiella salsuginea (Pall.) O. E. Schulz, Pflanzenr. Crucif. Sisymbr. 252 (1924);
Porsild in Sargentia 4: 40 (1943).

Sisymbrium salsugineum Pall. Reise 2: App. 114, ¢. 5 (1773); Raup in Jour. Arn. Arb.
17: 258 (1936) (misspelled “salsuginosum”) ; Payson in Univ. Wyo. Publ. Sci. 1:
17 (1922).
Turritis ? diffusa Hooker, Fl. Bor.-Am. 1: 41 (1829) ; Macoun, Cat. Can. Pl. Pt. 1: 47
(1883).
This rare species has not been collected in the Mackenzie valley, but I have
found it on the salt plains west of the upper Slave R., and Porsild has it from
Liverpool Bay on the arctic coast east of the Mackenzie.

Descurainia sophioides (Fisch.) O. E. Schulz, Pflanzenr. Crucif. Sisymbr. 316 (1924);
Raup in Jour. Arn. Arb. 17: 258 (1936); Hultén, Fl. Alaska & Yuk. Pt. a: Gee
(1945).

Sisymbrium sophioides Fisch. apud Hooker, Fl. Bor.-Am. 1: 61 (1830) ; Macoun, Cat.
Can. Pl. Pt. 1: 46 (1883).
Twelve mi. east of Moraine Pt., northwest shore of Great Slave L., Bedford (Can.).

This species was observed growing on the river bluff at Simpson. Its range
extends, according to Fl. Bor.-Am., ‘“. . . to the shores of the Arctic sea, west-
ward of the Mackenzie River.” I have seen specimens from Fort Good Hope and
Peel River.

Descurainia Richardsonii (Sweet) O. E. Schulz, Pflanzenr. Crucif. Sisymbr. 318 (1924);
Raup in Jour. Arn. Arb. 17: 258 (1936); Hultén, Fl. Alaska & Yok Pt.5: 82)
(1945).

Sisymbrium Richardsonii Sweet, Hort. Brit. ed. 2: 30 (1830).
Sisymbrium canescens sensu Hooker, Fl. Bor.-Am. 1: 62 (1830), in part; Macoun, Cat.
Can. Pl. Pt. 1: 46 (1883), in part; non Nutt.
Sisymbrium brachycarpum Richardson, sensu Hooker, Fl. Bor.-Am. 1: 62 (1830), in part.
Resolution, E. A. & A. E. Preble 148 (US); Providence, Preble & Cary 31 (US): dry
places, Simpson, Miss E. Taylor (Can., US); Norman, Dutilly 49 (G); Mackenzie R.,
Onion & Ross (N).

Braya humilis (C. A. Mey.) Robins. Syn. Fl. N. Am. 1: 141 (1895); Raup in Jour.
Arn. Arb. 17: 258 (1936); Porsild in Sargentia 4: 45 (1943), in Nat. Mus. Can.
Bull. 101: 31 (1945).

Sisymbrium humile C. A. Mey. in Ledeb. Icon. Pl. Ross. 2: 16, ¢. 147 (1830) ; Hooker,

Fl. Bor.-Am. 1: 62 (1830) ; Macoun, Cat. Can. Pl. Pt. 1: 47 (1883).
Torularia humilis (C. A. Mey.) O. E. Schulz; Hultén, Fl. Alaska & Yuk. Pt. 5: 889
(1945).

Northwest shore of Great Slave L., Bedford (Can.); Liard R. between Nahanni Butte
and Simpson, Crickmay 51, 54 (Can.) ; Simpson, Onion, Kennicott & Hardisty (N); steep
slope of river bluff at Simpson 9178; same place, Wynne-Edwards 8601 (Can.) ; sandy under-
cut bank of Mackenzie R. west of Simpson, 9918; Bear Rock, Norman, A. £. & R. T.
Porsild 3383 (Can.) ; Lone Mt., Wynne-Edwards 8477 (Can.) ; north peak of Nahanni Mt.,
W ynne-Edwards 8478 (Can.).

Occasional to common in the Simpson district ; collected in flower about mid-

June, and with flowers and mature siliques late in August.

Braya Henryae Raup in Contr. Arn. Arb. 6: 167, t. 7, fig. 1 (1934).
Braya purpurascens sensu Porsild in Nat. Mus. Can. Bull. 101: 21 (1945), non Bunge.
Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11812 (Can.) ; Bolstead Cr., Canol Rd.,
mi. 111 E., alpine slope, Wynne-Edwards 8309 (Can.).
Braya Richardsonii (Rydb.) Fernald in Rhod. 20: 203 (1918); Porsild in Nat. Mus.
Can. Bull. 101: 31 (1945).
Pilosclla Richardsonii Rydb. in Torreya 7: 159 (1907).

186 SARGENTIA [6

Porsild reports a doubtfully determined specimen of this species collected by
W’ynne-Edwards on the north peak of Nahanni Mt., 8479.

Erysimum cheiranthoides L. Sp. Pl. 661 (1753); Hooker, Fl. Bor.-Am. 1: 64 (1830);
Macoun, Cat. Can. Pl. Pt. 1: 45 (1883); Raup in Jour. Arn. Arb. 17: 258 (1936);
Hultén, Fl. Alaska & Yuk. Pt. 5: 884 (1945).

Fort Smith, Raup 799; Lower Slave R., Raup 798; Great Slave L., Onion, Kennicott &
Hardisty (N); Hay River, Hume 102666 (Can.) ; northwest shore of Great Slave L.,
Bedford (Can.).

Noted in Fl. Bor.-Am. “Throughout Canada . . . to lat. 67° on the Mack-
enzie.” I have seen specimens from Great Bear L., Good Hope, and Peel R.

Erysimum inconspicuum (Wats.) MacMillan, Metasperm. Minn. Valley, 268 (1892);
Hultén, Fl. Alaska & Yuk. Pt. 5: 885 (1945),

Erysimum asperum DC. var. inconspicuum Wats. in King, Geol. Expl. 40°" Par. 5: 24
(1871). .

Erysimum lanceolatum sensu Hooker, Fl. Bor.-Am. 1: 64 (1830).

Erysimum parviflorum Nutt. (non Pers.) ; Macoun, Cat. Can. Pl. Pt. 1: 45 (1883);
Raup in Jour. Arn. Arb. 17: 258 (1936).

Mackenzie R., Richardson (Can.).

This species is only occasional in the Wood Buffalo Park, and judging by the
rarity of more northern collections of it, it presumably is also occasional in the
Mackenzie country. I have seen a specimen in Herb. Can. from the lower Mack-
enzie collected by McConnell.

Erysimum Pallasii (Pursh) Fernald in Rhod. 27: 171 (1925); Hultén, Fl. Alaska &
Yuk. Pt. 5: 885 (1945).

Cheiranthus Pallasii Pursh. Fl. Am. Sept. 2: 436 (1814) ; Macoun, Cat. Can. Pl. Pt. 1:
48 (1883).

Hesperis pygmaeus Hooker, Fl. Bor.-Am. 1: 60 (1830).
On a mountain above timber line, June 3, 1908, Keele (Can.).

This rare plant is known on the arctic coast and at Great Bear L. The above
record is the only one to date in our region.

Rorippa obtusa (Nutt.) Britton in Mem. Torr. Bot. Cl. 5: 169 (1894); Raup in Jour.
Arn. Arb. 17: 259 (1936); Porsild in Sargentia 4: 40 (1943).

Nasturtium obtusum Nutt. in T. & G. Fl. N. Am. 1: 74 (1838).

Not yet collected on the upper Mackenzie, but found in the Wood Buffalo Park
and in the Mackenzie delta (Porsild, 1. ¢.).

Rorippa islandica (Oeder) Borbas var. microcarpa (Regel) Fernald in Rhod. 42: 271
(1940). See Rhod. 42: 25-32 (1940).

Nasturtium palustre DC. var. microcarpa Regel in Mém. Acad. Imp. Sci. St. Pétersb.
VII. 4, no. 4 (Tent. Fl. Ussur.): 20 (1861).

Nasturtium palustre DC. sensu Hooker, Fl. Bor.-Am. 1: 39 (1829), in part.
Rorippa palustris (L.) Bess. var. glabrata (Lunell) Victorin; Raup in Jour. Arn. Arb.
17: 259 (1936).

Resolution, Onion, Kennicott & Hardisty (N); 12 mi. east of Moraine Pt., northwest
shore of Great Slave L., Bedford (Can.); stony shore of Brintnell L. at Camp Pt., 9377;
wet mossy shore of Brintnell L. near Camp Pt., 9489,

Occasional to common; collected with flowers and maturing capsules about
mid-July. Noted in Fl. Bor.-Am. “. . . to the shores of the Arctic Sea. . . .”

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 187

Barbarea orthoceras Ledeb. in Index Sem. Hort. Dorpat (1824); Raup in Jour. Arn.
Arb. 17: 259 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 825 (1945).

Barbarea praecox sensu Hooker, FI. Bor.-Am. 1: 39 (1829), in part, non Sm.

Barbarea vulgaris R. Br. var. stricta sensu Macoun, Cat. Can. Pl. Pt. 1: 45 (1883), in
part, non Regel.

Northwest shore of Great Slave L., Bedford (Can.); Rae, Russell (Can.); Wrigley
Harbor, Great Slave L., Kindle (Can.) ; “Canada to Lat. 68°,” “Mack. River,” Richardson
(Can.).

Cardamine bellidifolia L. Sp. Pl. 654 (1753); Hultén, Fl. Alaska & Yuk. Pt. 5: 832
(1945).

Stony beaches and thickets near shore of Brintnell L. at Camp Pt., 9152; shale cliffs with
north exposure, summit of Red Mt., alt. 5900’, 9765; crevices and stony stream channels,
northeast slope of Red Mt., alt. 4500’-5500’, 9272; mossy banks of small brook on west slope
of Terrace Mt., alt. 5000’, 9541.

Occasional to common; siliques beginning to mature at lake-level late in June,
but plants found in flower at high elevations as late as mid-July. Noted in Fl.
Bor.-Am. as occurring “Throughout Arctic America, sparingly.”

Cardamine pratensis L. Sp. Pl. 656 (1753); Hooker, Fl. Bor.-Am. 1: 45 (1829), in part;
Macoun, Cat. Can. Pl. Pt. 1: 41 (1883), in part; Hultén, Fl. Alaska & Yuk. ith
5: 834 (1945); Porsild in Nat. Mus. Can. Bull. 101: 20 (1945). See Rhod. 22:
11-14: (1920).

Muskeg thickets at west end of Brintnell L., 9236.

Occasional in this habitat; collected with flowers and immature siliques June
23. Noted in Fl. Bor.-Am. as occurring “Throughout Arctic America.” Por-
sild reports that he observed it in Macmillan Pass, on the Canol Rd., mi. 225 E.

The Brintnell L. plants are near var. angustifolia Hooker (1. c.), but the lateral
leaflets of some of the basal leaves are distinctly broad ovate as in the typical form.
It is of interest that the lower leaves and petioles have the strigose pubescence
mentioned by Porsild as characteristic of his Alaskan material (Rhod. 41: 234,
1939). This appears to relate the plants to some northern Eurasian and Euro-
pean alpine forms rather than to the northeastern American forms which lack the
pubescence.

Cardamine pratensis L. var. palustris Wimm. & Grab. Fl. Siles. 2: Pt. 1, 266 (1829);
Raup in Jour. Arn, Arb. 17: 259 (1936).
Northwest shore of Great Slave L., Bedford (Can.); wet swamp in low woods near
Simpson, 9101.
Common at the Simpson locality, but not seen elsewhere; in flower June 12.
I have seen a specimen also from Peel R., in the lower Mackenzie region.

Cardamine pennsylvanica Muhl. apud Willd. Sp. Pl. 3: 486 (1800); Raup in Jour. Arn.
Arb. 17: 259 (1936).

Cardamine hirsuta sensu Hooker, Fl. Bor.-Am., 1: 45 (1829) ; Macoun, Cat. Can. Pl. Pt.
1: 41 (1883); in part, non L.

This species has not been collected along the upper Mackenzie, but it is com-
mon in upland muskegs in the Wood Buffalo Park. In FI. Bor.-Am. it is noted
as extending “. . . to the shores of the Arctic Sea.” There are Richardson
specimens in Herbs. N and Can. labeled “Franklins Journey” and “Arctic N.
A.,” both originally marked C. hirsuta, but without specific locality data.

-Eutrema Edwardsii R. Br., Suppl. to App. Parry’s 1°* Voy. 267, t. A (1824); Hultén,

Fl. Alaska & Yuk. Pt. 5: 819 (1945); Porsild in Nat. Mus. Can. Bull. 101: 20
(1945).

188 SARGENTIA [6

Plains of Abraham, Canol Rd., mi. 82 E., alt. 6000’, Porsild 11808 (Can.) ; alpine meadow,
Bolstead Cr., Canol Rd., mi. 111 E., alt. 4500'-5500', Wynne-Edwards 8308 (Can.).

Noted by Porsild as rare in the Cassiope tetragona communities on the Plains
of Abraham.

Parrya nudicaulis (L.) Regel in Bull. Mosc. 34: Pt. 2, 176 (1861); Hultén, Fl. Alaska
& Yuk. Pt. 5: 890 (1945), in part; Porsild in Nat. Mus. Can. Bull. 101: 21 (1945).

Cardamine nudicaulis L. Sp. Pl. 654 (1753).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11810 (Can.) ; Bolstead Cr., Canol Rd.,
mi. 111 E., Wynne-Edwards 8316, 8317 (Can.); Sekwi R., Canol Rd., mi. 174 E., Porsild
11963 (Can.).

Hultén (1. c.) has proposed a ssp. interior for plants with leaves lanceolate
or broadly lanceolate, obtuse or subacute, entire or nearly so; with filaments
dilated at the middle or toward the base; and with slender pods and long styles.
The above specimens may represent this subspecies.

Melanidion boreale Greene in Ottawa Nat. 25: 146 (1912); Porsild in Nat. Mus. Can.
Bull. 101: 21 (1945),

Ermania borealis (Greene) Hultén, Fl. Alaska & Yuk. Pt. 5: 878 (1945),

Barren wind-swept hill in limestone rubble, Plains of Abraham, Canol Rd., mi. 82 E.,
alt. 6000’, Porsild 11809 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., in limestone scree,
Wynne-Edwards 8318 (Can.); Sekwi R., Canol Rd., mi. 174 E., alt. 3625’, in limestone
scree, Porsild 11865 (Can.).

The recent discussions by Hultén and Porsild (1. c.) have greatly enlarged
our understanding of this rare species. Hultén has reduced Melanidion to the
genus Ermania, which was described by Chamisso in 1831 (Linnaea 6: 533)
from material collected in Kamtchatka. The type species of Ermania was E.
parryoides, which was placed in the genus Parrya by Hultén in his Flora of
Kamtchatka (p. 172, 1928). For this plant Hultén took up the name Parrya
Ermani Ledeb. (Fl. Ross. 1: 132, 1842). It is very difficult for me to identify
the unique Melanidion, which has striking racemes of purple flowers and short
pods, with anything in the genus Parrya, including the Parrya Ermani of
Kamtchatka. I am therefore following Porsild in retaining Greene’s genus.

Arabis lyrata L. var. kamchatica Fisch. ex DC. Syst. Veg. 2: 231 (1821). See Rollins
in Rhod. 43: 360 (1941).

Sisymbrium arabidoides Hooker, Fl. Bor.-Am. 1: 63 (1830).

Arabis lyrata L. var. glabra (DC.) Hopkins in Rhod. 39: 93 (1937); Porsild in Nat.
Mus. Can. Bull. 101: 21 (1945).

Arabis lyrata L. ssp. kamchatica (Fisch.) Hultén, Fl. Aleut. I, 202 (1937) ; Fl. Alaska
& Yuk. Pt. 5: 875 (1945).

Stony beach of Brintnell L. at Camp Pt., 9153, 9694; shale crevices and alpine rock-slides,
north slope of Colonel Mt., alt. 5000’-6000’, 9408, 9619; along stream in upper valley of
Frost Cr., alt. 4000’, 9564; shaly south slope of Red Mt., alt. 5000’, 9723.

Occasional to common, especially in alpine slide rock; found with flowers
and mature siliques at lake-level June 21, and in flower above timber line as late
as mid-July. Noted by Porsild (1. c.) in Macmillan Pass along the Canol Rd.,
and in Fl. Bor.-Am, “. . . from Canada to lat. 68°.” There are specimens in
Herbs. G, N, and Can. labeled “FI. Bor.-Am.,” “Franklin’s Journey,” and “Arctic
Canada,” but none have specific locality data.

The occurrence of typical Arabis lyrata in southwestern Mackenzie is somewhat
in doubt. Hopkins (1. c., 91) regarded the typical form as mainly confined to
eastern America, but considered one of my numbers from the Wood Buffalo Park

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 189

(2485) as representing it. Porsild (Sargentia 4: 43) collected several numbers
at Great Bear L., all of which he listed as A. lyrata. Rollins (1. c.), on the
other hand, does not recognize typical A. lyrata west of the 100" meridian, al-
though he regards var. kamchaticg as a poorly defined segregate. The appear-
ance of the eastern form in the Mackenzie country is, of course, not inconsistent
with other range patterns.

Arabis hirsuta (L.) Scop. var. pycnocarpa (Hopkins) Rollins in Rhod. 43: 318 (1941).
Turritis hirsuta L. Sp. Pl. 2: 666 (1753).
Arabis pycnocarpa Hopkins in Rhod. 39: 112 (1937).
Arabis hirsuta (L.) Scop., sensu Hooker, Fl. Bor.-Am. 1: 42 (1829), in part; Macoun,
Cat. Can. Pl. Pt. 1: 42 (1883), in part; Raup in Jour. Arn. Arb. 17: 260 (1936).
Arabis hirsuta (L.) Scop. ssp. pycnocarpa (Hopkins) Hultén, Fl. Alaska & Yuk. Pt. 5:
873 (1945).
Resolution, Dutilly 100 (G); Mackenzie R., Kennicott (N); clearing at Simpson, 9108;
upland prairie west of the Mackenzie R. near Simpson, 9897.
Occasional; in flower about mid-June, and found with mature and shattered
siliques late in August. Noted in Fl. Bor.-Am, “. . . from Canada to lat. 68°,”
and I have seen a specimen from Good Hope (Onion, N).

Arabis Holboellii Hornem. in FI. Dan. 11: 5, t. 1879 (1827); Porsild in Sargentia 4:
43 (1943); Hultén, Fl. Alaska & Yuk. Pt. 5: 874 (1945). See Rollins in Rhod.
43: 439 (1941).

Mossy thickets on stony shore of Brintnell L., 9755; gravelly slope in gorge on north
slope of Colonel Mt., alt. 3300’, 9837; scrub timber on south slope of Red Mt., alt. about
4000’, 9219, 9778.

Occasional; collected with flowers and immature siliques late in June, and with
mature fruit early in August.

Arabis Holboellii Hornem. var. retrofracta (Graham) Rydb. in Contr. U. S. Nat. Herb.
3: 484 (1896); Rollins in Rhod. 43: 441 (1941); Hultén, Fl. Alaska & Yuk. Pt. 5:
874 (1945).
Arabis retrofracta Graham in Edinb. New Phil. Jour. 344 (1829); Macoun, Cat. Can.
Pl. Pt. 1: 43 (1883); Hopkins in Rhod. 39: 179 (1937); Raup in Jour. Arn, Arb.
17: 260 (1936).

Turritis retrofracta (Graham) Hooker, Fl. Bor.-Am. 1: 41 (1829).

Fort Smith, Miss E. Taylor 1683 (Can.); Windy Pt., Great Slave L., Hume 102665
(Can.); Mackenzie R., Onion, Kennicott & Hardisty (N); Liard R. between Nahanni
Butte and Simpson, Crickmay 52 (Can.).

Noted in Fl. Bor.-Am. “. . . from Canada to lat. 68° ; at the Mackenzie River.”
I have seen a specimen from Good Hope (Onion, N).

Arabis divaricarpa A. Nels. in Bot. Gaz. 30: 193 (1900); Raup in Jour. Arn. Arb. 17:
260 (1936); Hopkins in Rhod. 39: 129 (1937); Rollins in Rhod. 43: 374 (1941);
Hultén, Fl. Alaska & Yuk. Pt. 5: 871 (1945).

Providence, Preble & Cary 32 (US); dry slopes, Simpson, Miss E. Taylor 120 (Can.) ;
south-facing slope in old field, Simpson, 9125; Liard R. between Nahanni Butte and Simpson,
Crickmay 53 (Can.).

Occasional; in flower about mid-June. All of the above specimens represent
the typical form.

Arabis Drummondii Gray in Proc. Am. Acad. 6: 187 (1866); Raup in Jour. Arn. Arb.
17: 260 (1936); Hopkins in Rhod. 39: 136 (1937); Rollins in Rhod. 43: 369 (1941);
Porsild in Sargentia 4: 43 (1943); Hultén, Fl. Alaska & Yuk. Pt. 5: 871 (1945).

Dry openings in upper part of timber, south slope of Red Mt., 9677; on undercut banks

in upper valley of Frost Cr., alt. 4000’, 9521.

190 SARGENTIA [6

Occasional ; found with maturing capsules and a few flowers about mid-July.
Occasional in the Wood Buffalo Park, and collected by Porsild at Great Bear L.
and in the Richardson Mts. ,

DROSERACEAE

Drosera rotundifolia L. Sp. Pl. 281 (1753); Hooker, Fl. Bor.-Am, 1: 81 (1830); Macoun,
Cat. Can. Pl. Pt. 1: 165 (1883); Raup in Jour. Arn. Arb. 17: 261 (1936); Hultén,
Fl. Alaska & Yuk. Pt. 5: 893 (1945).
Great Slave L., R. Bell 23170 (Can.); southwest and north shores of Great Slave L.,
Howe 91976 (Can.).

Noted in Fl. Bor.-Am. “. . . as far north as the Arctic Circle.”

Drosera anglica Huds. Fl. Angl. ed. 2: 135 (1778); Raup in Jour. Arn. Arb. 17: 261
(1936); Porsild in Sargentia 4: 47 (1943); Hultén, Fl. Alaska & Yuk. Pt. 5:
893 (1945).
This species has not yet been observed on the upper Mackenzie, but I have
collected it at L. Athabaska, and Porsild has it from Great Bear L.

CRASSULACEAE
Sedum roseum (L.) Scop. var. integrifolium (Raf.) Berger in Engl. & Prantl, Nat.
Pflanzenf. 2 Aufl. 18a: 440 (1930).

Rhodiola integrifolia Raf. Atl. Jour. 1: 146 (1832); Porsild in Nat. Mus. Can. Bull.
101: 22 (1945).

Sedum roseum (L.) Scop. ssp. integrifolium (Raf.) Hultén, Fl. Alaska & Yuk. Pt. 5:
895 (1945).

Dry gravelly watercourse on north slope of Colonel Mt., alt. 5000’, 9622; south slope of
Red Mt., alt. 4000’, 9226,

Common on alpine summits and rocky slopes at Brintnell L.; in flower late in
June, and with maturing capsules late in July. Porsild also notes it as common
in Macmillan Pass, along the Canol Rd.

SAXIFRAGACEAE ‘aio
Saxifraga rivularis L. Sp. Pl. 404 (1753); Hultén, Fl. Alaska & Yuk. Pt. 5: 935 (1945);
Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).
Wet mossy banks of small brook on west slope of Terrace Mt., alt. 4800’-5500’, 9524;
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8333 (Can.).
Common at the Terrace Mt. station but not seen elsewhere in the Brintnell L.
area. Porsild notes it as common in Macmillan Pass.

Saxifraga cernua L. Sp. Pl. 403 (1753); Hultén, Fl. Alaska & Yuk. Pt. 5: 910 (1945);
Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8321 (Can.).

Saxifraga sibirica L. Syst. ed. 10: 1027 (1759).

Damp and shaded mossy bank in gorge on north slope of Colonel Mt., alt. 3300’, 9836.

Apparently rare, and found only in this single locality ; collected with immature
capsules and the dried remains of flowers on Aug. 16.

This species does not seem to have been reported previously on the North
American continent, and it is not recorded in the Bering Sea region or neighbor-
ing parts of eastern Siberia. It has a wide range in central Asia, extending, ac-
cording to Hultén (Dist. of Arct. and Bor. Biota, 18), westward to the Urals.
It is also known in the Caucasus. Our plants are a good match for many speci-
mens of S. sibirica in the Gray Herbarium. They have rather long, obovate-

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 191

cuneate petals (1 cm. long) and ovate-oblong sepals (3.5 mm. long), and their
rounded-reniform leaves have lobes that are rounded to almost broadly truncate
at the apex. The stems, peduncles, and hypanthia, as well as the calyx-lobes in
part, are finely glandular-pubescent.

The nearest American relative of S. sibirica is S. debilis Engelm., described
from a collection by Parry in the Rocky Mountains in 1862 (ex Gray, Proc. Acad.
Phil., 1863, p. 62). The type of this species (in the Gray Herbarium) is almost
completely glabrous, with ovate sepals 1.5-2 mm. long, and cuneate to oblong
petals 3-7 mm. long. The lobes of its leaves are not so rounded as in S. sibirica,
in some even being acute; also the lobes are not so rounded below or overlapping
in the sinuses as in the latter species. Occasional specimens of S. debilis in the
Gray Herbarium have finely pubescent stems, but are not glandular except on
the hypanthium (See A. Nelson 2447, Dome L., Wyo.). The range of S.
debilis does not appear to extend north of Montana and Mt. Rainier (See N. Am.
Fi; 22: Pt! 231287;

Engler and Irmscher (Pflanzenreich 67 [IV. 117.1]: 262) distinguish S.
sibirica and some other Old World species from S. debilis as follows:

Caules floriferi pluriflori, interdum 1-2-flori, floribus breviter pedicellatis...... S. sibirica.

Caules floriferi 1—-2-flori, floribus longius pedicellatis.........seeeeeeeeee ences S. debilis.
To these differences might well be added the size of the flowers, for the American
species has consistently shorter petals and sepals than S. sibirica. Many Asiatic
specimens of S. sibirica in the Gray Herbarium collection have long-pediceled
flowers, 1 or 2 to the plant. Following Engler & Irmscher’s subdivision of S.
sibirica (1. c., pp. 264-267), these forms fall into var. pekinensis (Maxim.) Engl.
& Irmsch., and are most nearly related to our Mackenzie Mountain plants, each
of which has only 1 or 2 flowers, on peduncles 2.5-3 cm. long.

Saxifraga nivalis L. Sp. Pl. 401 (1753); Hultén, Fl. Alaska & Yuk. Pt. 5: 924 (1945).
Shale cliffs on upper slopes and summit of Red Mt., alt. 5500’-5900’, 9766.

Apparently rare; capsules mature early in August.

Saxifraga hieracifolia Waldst. & Kit. Pl. Rar. Hung. 1: 17, t. 18 (1802); Hultén, FI.

Alaska & Yuk. Pt. 5: 918 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., alpine ridge, alt. 5000’-6000’, Wynne-Edwards 8323,
8324 (Can.).

Saxifraga Lyallii Engler in Verh. Zool.-Bot. Ges. Wien. 542 (1869); Hultén, FI.
Alaska & Yuk. Pt. 5: 922 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Among soil polygons near iron spring in upper valley of Frost Cr., north of Brintnell L.,
alt. 4500’, 9560; Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes up to alt. 7000’, Wynne-
Edwards 8325, 8326, 8327 (Can.); Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11240
(Can.).

Apparently rare or occasional at Brintnell L., but more common along the
Canol Rd. Among the soil polygons mentioned above it was the only vascular
plant growing in the frost-heaved finer soils. Collected with flowers and im-

mature capsules July 18.

Saxifraga punctata L. Sp. Pl. 401 (1753); Hultén, Fl. Alaska & Yuk. Pt. 5: 928 (1945);
Porsild in Nat. Mus. Can. Bull. 101: 22 (1945). See Hultén, Fl. Aleut. I. 213-214
(1937), and Porsild in Rhod. 41: 242 (1939) for recent discussions of this species.

Along stream in upper valley of Frost Cr., alt. 4500’, 9515; along small stream on east
slope of Red Mt., and in high shale crevices, alt. 4000’-5500', 9273; mossy banks of brook,

west slope of Terrace Mt., alt. 4800’-5500’, 9551.

192 SARGENTIA [6

In general rare or occasional, but abundant locally; collected with flowers
and maturing fruit about mid-July. Our plants answer the description of those
collected by Porsild in Alaska, and seem to approach the ssp. insularis proposed
by Hultén. Noted by Porsild as common in Macmillan Pass, along the Canol
Rd.

Saxifraga radiata Small in N. Am. FI. 22: 128 (1905); Hultén, Fl. Alaska & Yuk. Pt.
5: 933 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).
Gravelly thickets, south shore of Brintnell L., 9335; Macmillan Pass, Canol Rd., mi. 225
E., Porsild 11239 (Can.).
Apparently rare or occasional at Brintnell L., and noted by Porsild as rare in
an alpine meadow in Macmillan Pass. Collected in flower June 30.

Saxifraga aizoides L. Sp. Pl. 403 (1753); Raup in Jour. Arn. Arb. 17: 261 (1936);
Porsild in Nat. Mus. Can. Bull. 101: 22, 31 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 56-A (Can.); Smith Cr., 4 mi.
above mouth, Crickmay 24 (Can.); dry gravelly fan at base of granite mountain west of
Brintnell L., 9477; alpine crevices and rock-slides, north slope of Colonel Mt., alt. 4000’—
6000’, 9406, 9651; Lone Mt., Wynne-Edwards 8480 (Can.) ; Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8322 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11867 (Can.).

Common to abundant on Colonel Mt., at Brintnell L., but apparently rare or oc-
casional elsewhere in that area; in flower about mid-July.

Saxifraga tricuspidata Rottb. in Skr. Kjébenh. Selsk. Laerd. Vidensk. 10: 446, t. 6, f. 21
(1770); Hooker, Fl. Bor.-Am. 1: 254 (1834); Macoun, Cat. Can. Pl. Pt. 1: 154
(1883); Raup in Jour. Arn. Arb. 17: 261 (1936); Hultén, Fl. Alaska & Yuk. Pt.
5: 939 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22, 31 (1945).

Resolution, Kennicott (N); same place, Preble 235 (US); southwest and north shores
of Great Slave L., Howe 91970 (Can.) ; Windy Pt., Great Slave L., Hume 102667 (Can.) ;
northwest shore of Great Slave L., Bedford (Can.); Liard R. between Nahanni Butte and
Simpson, Crickmay 58 (Can.) ; Mackenzie R. between Blackwater R. and Norman, Crick-
may 168 (Can.); gravelly thickets on shore of Brintnell L., near camp, 9312; south slope
of Red Mt., 9225; Lone Mt., Wynne-Edwards 8482 (Can.) ; north peak of Nahanni Mt.,
Wynne-Edwards 8483 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8320
(Can.).

One of the most abundant species on high alpine slopes and in unstable situa-
tions at lower altitudes. On many high ledges and loose rocky slopes it is one
of the primary plants of the scanty cover. Collected in flower in middle and late
June. Noted in Fl. Bor.-Am. “From Lake Winipeg . . . to the Arctic Sea . . .”

Saxifraga serpyllifolia Pursh, Fl. Am. Sept. 1: 310 (1814); Hultén, Fl. Alaska & Yuk.
Pt. 5: 937 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., on alpine ridge, alt. 7000’, Wynne-Edwards 8334
(Can:

Saxifraga Hirculus L. Sp. Pl. 402 (1753); Hooker, Fl. Bor.-Am. 1: 252 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 154 (1883); Hultén, Fl. Alaska & Yuk. Pt. 5: 920 (1945);
Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes up to alt. 7000’, Wynne-Edwards 8328,
8329 (Can.).

Noted in Fl. Bor.-Am. as occurring “From the Saskatchewan to the Arctic Sea-
shore... .” I have seen a specimen from Great Bear L., but the species has not
been reported in the Athabaska—Great Slave L. region.

Saxifraga flagellaris Willd. ex Sternb. Rev. Saxifr. 25, t. 6 (1810); Hultén, Fl. Alaska
& Yuk. Pt. 5: 915 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 193

Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes up to alt. 7000’, Wynne-Edwards 8330,
8331 (Can.).

Saxifraga oppositifolia L. Sp. Pl. 402 (1753); Hultén, Fl. Alaska & Yuk. Pt. 5: 926
(1945); Porsild in Nat. Mus. Can. Bull. 101: 22, 31 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 57 (Can.); alpine meadows,
shale cliffs and rock-slides, north slope of Colonel Mt., 9397, 9623; Lone Mt., Wynne-
Edwards 8481 (Can.); Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11813 (Can.) ;
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8332 (Can.).

In the Brintnell L. area this species is common on Colonel Mt. but was not
observed elsewhere ; found with flowers and immature capsules early in July, and
with ripe fruit late in July.

Heuchera Richardsonii R. Br. in Richardson, Bot. App. Frankl. Journ. 766, t. 29 (1823);
Hooker, Fl. Bor.-Am. 1: 237 (1834); Raup in Jour. Arn. Arb. 17: 262 (1936).
See Rosendahl, Butters, & Lakela in Rhod. 35: 111-118 (1933).

Heuchera hispida sensu Macoun, Cat. Can. Pl. Pt. 1: 158 (1883), non Pursh.

Providence, Dutilly 74 (G).

Common in the Wood Buffalo Park and probably so along the upper Mack-
enzie. Noted in Fl. Bor.-Am. on “Dry banks of Rivers, from lat. 54° to 64°
North.”

Mitella nuda L. Sp. Pl. 406 (1753); Hooker, Fl. Bor.-Am. 1: 240 (1834); Macoun, Cat.
Can. Pl. Pt. 1: 157 (1883); Raup in Jour. Arn. Arb, 17: 262 (1936); Hultén, FI.
Alaska & Yuk. Pt. 5: 947 (1945).

Fort Smith, Miss E. Taylor 21 (G, Can.) ; Simpson, Hardisty & McFarlane (N); same,

Onion, Kennicott & Hardisty (N); damp mossy lowland woods near Simpson, 9099.

Common in the Simpson area, and probably also in rich woods throughout
the river lowlands. In flower about mid-June. Noted in Fl. Bor.-Am. “From

Lake Huron to the shores of the Arctic Sea... .”

Chrysosplenium tetrandrum (Lund) Th. Fries, in Bot. Notiser 1858: 193 (1858); Raup
in Jour. Arn. Arb. 17: 262 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 949 (1945);
Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Chrysosplenium alterniflorum sensu Hooker, FI. Bor.-Am. 1: 241 (1834); Macoun, Cat.
Can. Pl. Pt. 1: 159 (1883) ; non L.

Thickets in wet muskeg at west end of Brintnell L., 9241; stony beach on shore of Brint-
nell L. at Camp Pt., 9160.

Common in the muskeg habitat; capsules mature in the latter part of June.
Noted by Porsild as occasional or rare in Macmillan Pass. In Fl. Bor.-Am. it
is said to occur “In the woody, but principally the barren country of British
North America.”

Parnassia fimbriata Koenig in Ann. Bot. 1: 391 (1804); Hultén, Fl. Alaska & Yuk.
Pt. 5: 952 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Alpine meadow along small brook on west slope of Terrace Mt., alt. 5000’, 9553; alpine
meadows in Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11238 (Can.):.

Noted by Porsild as common in Macmillan Pass; but at Brintnell L. it was
found in only one place. In flower about mid-July.

Parnassia palustris L. var. neogaea Fernald in Rhod. 39: 311 (1937); Hultén, Fl.
Alaska & Yuk. 5: 956 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22, 31, 35
(1945).

Parnassia palustris L. sensu Hooker, Fl. Bor.-Am., 1: 82 (1830), in part; Macoun, Cat.
Can. Pl. Pt. 1: 159 (1883).

\

194 SARGENTIA [6

Parnassia multiseta, sensu Fernald in Rhod. 28: 211 (1926), non P. palustris var. multi-
seta Ledeb.; Raup in Jour. Arn. Arb, 17: 262 (1936).

Slave R., Miss E. Taylor 25 (G); Little Buffalo R., Russell 77 (G); Great Slave L.,
R. Bell 23164 (Can.) ; southwest and north shores of Great Slave L., Howe 91969 (Can.) ;
northwest shore of Great Slave L., Bedford (Can.); east shore of Trout L.,, C. O. Hage
(G) ; Mackenzie R., Kennicott (N); Liard R. between Nahanni Butte and Simpson, Crick-
may 55 (Can.) ; Mackenzie R., 5 mi. above old Ft. Norman, Wynne-Edwards 8562 (Can.) ;
thickets in upland woods west of Mackenzie R. near Simpson, 9916; muskeg thickets along
north shore of Brintnell L., 9344, 9571; gravelly thickets near west end of Brintnell 1 BAS
9440; Lone Mt., Wynne-Edwards 8484 (Can.).

Common at Brintnell L. and Simpson, and noted by Porsild as common along
a small tributary to Little Keele R., Canol Rd., mi. 51 E. In Fl. Bor.-Am. its

range is given as extending “. . . as far north as the Arctic Circle... .”

Parnassia montanensis Fernald & Rydb. in N. Am. FI. 22: 79 (1905); Porsild in Nat.
Mus. Can. Bull. 101: 33 (1945).

South bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8552 (Can.).
This species is occasional in the Athabaska—Great Slave L. region. It is not

included in Hultén’s Fl. Alaska & Yuk., but was collected in the mountains of
southwestern Yukon by Bakewell (Rhod. 45: 311, 1943).

Parnassia Kotzebuei Cham. & Schl. in Linnaea 1: 549 (1826); Hultén, Fl. Alaska &
Yuk. Pt. 5: 953 (1945); Porsild in Nat. Mus. Can. Bull. 101: 22 (1945).

Mossy banks and muskegs along north shore of Brintnell L., 9154, 9343; Bolstead Cr.,
Canol Rd., mi. 111 E., Wynne-Edwards 8319 (Can.).

Occasional to common at Brintnell L., found flowering in late June and early
July. Porsild notes it as common in Macmillan Pass.

Ribes oxyacanthoides L. Sp. PI. 201 (1753); Preble in N. Am. Fauna 27: 525 (1908) ;
sina Jour. Arn. Arb. 17: 263 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 962
(1945).

Fort Smith, Seton & Preble 78578 (Can.) ; lower Slave R., Raup 854; Resolution, Kenni-
cott (N); same place, Preble 205 (US); Hay River, Preble & Cary 26 (US) ; east shore
of Trout L., C. O. Hage (G); Simpson, Preble 300, 309 (US); damp woods near Simpson,
shrubs up to 3’ high, 9081; open woods and thickets on gravel plain west of Brintnell L.,
shrub 2’ high, 9806.

Common in the lowlands, and only occasional at Brintnell L.; in flower at
Simpson about mid-June, and in fruit at Brintnell L. in mid-August. Preble
states that it “is the commonest representative of the genus throughout the re-
gion... .” He noted it at Great Bear L.

Ribes hudsonianum Richardson, Bot. App. Frankl. Journ. ed. 2: 6 (1824); Preble in
N. Am. Fauna 27: 526 (1908); Raup in Jour. Arn. Arb. 17: 263 (1936); Porsild in
Sargentia 4: 49 (1943); Hultén, Fl. Alaska & Yuk. Pt. 5: 959 (1945).

Fort Smith, Miss E. Taylor (G, Can.) ; same place, Seton & Preble 78326 (Can.) ;
Resolution, Onion, Kennicott & Hardisty (N); southwest shore of Great Slave L., Brooke
(Can.) ; northwest shore of Great Slave L., Bedford (Can.) ; east shore of Trout 1 Ga.
Hage (G); Mackenzie R., Kennicott (N); Liard R. between Nahanni Butte and Simpson,
Crickmay 56 (Can.); damp woods at Simpson, shrubs 3’ high, 9078; mossy woods along
shore of Brintnell L. near camp, shrubs up to 3’ high, 9189, 9598.

Common to abundant, particularly at the edge of woods or in open spaces;
in flower about mid-June and in fruit in the latter part of July. Richardson
noted it only as far north as lat. 57°, but Preble reports it at “. . . various
places from the Athabaska nearly to Great Bear Lake.” Porsild collected it in
the Mackenzie delta, and states that it is common at Great Bear Lia

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 195

Ribes lacustre (Pers.) Poir. in Lam. Encycl. Méth. Suppl. 2: 856 (1812); Hooker, FI.
Bor.-Am. 1: 232 (1834); Macoun, Cat. Can. Pl. Pt. 1: 161 (1883); Porsild in
Sargentia 4: 48 (1943); Hultén, Fl. Alaska & Yuk. Pt. 5: 960 (1945).

Ribes oxyacanthoides var. lacustris Pers. Syn. Pl. 1: 252 (1805).

Damp lowland woods near Simpson, decumbent shrub trailing in moss, 9077 ; open woods
and thickets on gravel plain at west end of Brintnell L., low shrub, 9807.

Occasional; in flower at Simpson about mid-June, and with mature fruit in
the mountains about mid-August. Hooker gives the northern range of this spe-
cies as “Throughout Canada to Fort Franklin and Bear Lake, near the Arctic
circle.” Porsild, however, maintains that the Fl. Bor.-Am. record for Great Bear
L. is R. oxyacanthoides, but he reports R. lacustre at Gordon L. on the Yellow-
knife R. Preble apparently did not see R. lacustre anywhére in the Mackenzie
basin.

Ribes glandulosum Grauer, Pl. Min. Cog. 2 (1784); Raup in Jour. Arn. Arb. 17: 263
(1936).
Ribes prostratum L’Her.; Preble in N. Am, Fauna 27: 526 (1908) ; Hultén, Fl. Alaska
& Yuk. Pt. 5: 962 (1945).
Slave R., Miss E. Taylor 8666 (Can.); Resolution, Kennicott (N); mossy woods on
shore of Brintnell L. near camp, decumbent shrub, 9182, 9601.
Common in the Brintnell L. habitat; collected in flower June 20, and with
mature fruit July 26. Noted by Preble as occurring “. . . throughout the coun-
try from the Athabaska north at least to Great Bear Lake.”

Ribes triste Pall. in Nova Acta Acad. Petrop. 10: 378 (1797); Raup in Jour. Arn. Arb.
17: 263 (1936); Hultén, Fl. Alaska & Yuk. Pt. 5: 963 (1945).

Ribes rubrum sensu Hooker, Fl. Bor.-Am. 1: 232 (1834); Macoun, Cat. Can. Pl. Pt. 1:
162 (1883); Preble in N. Am. FI. 27: 526 (1908) ; non L.

Fort Smith, Seton & Preble 78328 (Can.); Mackenzie R., Onion (N); near Wrigley,
Miss E. Taylor 125 (Can.); rich woods on shore of Brintnell L., near camp, low shrubs
up to 2’ high, 9188, 9488.

Common in the Brintnell L. habitat; collected with late flowers and immature
fruit June 20, and with ripe berries in mid-July. Noted in Fil. Bor.-Am. “
to the mouth of the Mackenzie River;” and by Preble, “. . . throughout the
region north at least to Great Bear Lake.” I have seen specimens from the
lower Mackenzie collected by Kennicott (N) and McConnell (Can.).

ROSACEAE

Spiraea Beauverdiana Schn. in Bull. Herb. Boiss. II. 5: 348 (1905); Porsild in Nat.
Mus. Can. Bull. 101: 23 (1945). See also Porsild in Rhod. 41: 244 (1939), and
in Sargentia 4: 48 (1943). ;

Mackenzie R. between Blackwater R. and Norman, Crickmay 169 (Can.).
This species was noted by Porsild as very common in Macmillan Pass, along

the Canol Rd., mi. 225 E.

Amelanchier florida Lindl. in Bot. Reg. ¢. 1589 (1833); Raup in Jour. Arn. Arb. 17:
264 (1936).
Amelanchier ovalis sensu Hooker, Fl. Bor.-Am. 1: 202 (1834), in part, non Willd.
Amelanchier alnifolia sensu Preble in N. Am. Fauna 27: 528 (1908), ? non Nutt.
“Amelanchier canadensis var. (?) oblongifolia” sensu Macoun, Cat. Can. Pl. Pt. 1: 149
(1883), in part, non T. & G.

Fort Smith, Raup 930; same place, Seton & Preble 78587 (Can.) ; Resolution, Kennicott
(N); along road in upland woods west of Mackenzie R. near Simpson, 9906; south slope
of Red Mt., near timber line, shrub 2’-3' high, 9222.

196 SARGENTIA [6

Occasional to common near Simpson, but aparently rare in the Mountains;
in flower at Brintnell L. June 21. Part of the records for A. ovalis in Fl. Bor.
Am., “. .. to the Saskatchewan and Mackenzie Rivers” probably refers to
this species, as does also Preble’s note that it occurs “. . . northward to the
limit of the woods.” Porsild (Sargentia 4: 48) regards the Amelanchier of Great
Bear L. and the lower Mackenzie as A. sanguinea (Pursh) DC.

Our Simpson and Brintnell L. plants approach the A. humilis described by
Wiegand in Rhod. 14: 141 (1912), but there appears to be considerable doubt
whether this species is separable from the widespread A. florida of the northwest.

Rubus strigosus Michx. Fl. Bor.-Am. 1: 297 (1803); Preble in N. Am. Fauna 27: 527
(1908). See L. MH. Bailey in Gentes Herb, 5: 861-878 (1945).

Rubus idaeus L. var. canadensis Richardson ex Fernald in Rhod. 21: 89-98 (1919) ; Raup
in Jour. Arn. Arb. 17: 264 (1936), in part.

Rubus idaeus L. var. strigosus (Michx.) Maxim.; Fernald, |. c.; Raup in Jour. Arn. Arb.
17: 265 (1936), in part.

Fort Smith, Raup 885; lower Slave R., Raup 882; Rae, Russell (Can.); east shore of
Trout L., C. O. Hage (G); small sandy clearing near eastern end of Brintnell L., shrubs
2’ high, 9350.

Occasional to common in gravelly or sandy thickets around the shore of Brint-
nell L., and observed at about 4000’ on the warm southerly slope of Terrace Mt.;
in flower early in July. This species is common to abundant in the Athabaska—
Great Slave Lake region, and it is strange that in Hooker’s Fl. Bor.-Am. it is not
noted north of the Saskatchewan. Preble gives its range as “. . . nearly through-
out the forested region.” I have seen specimens of it from the lower Mackenzie
(Good Hope, Onion, N; Peel R., Miss E. Taylor, Can.). Bailey, in his recent
revision of the American representatives of R. idacus, does not recognize the
two entities that were defined by Fernald as R. idaeus var. canadensis and R. idacus
var. strigosus. In the former the primocanes are invested with a fine pubescence
beneath the prickles, while in the latter the stems are smooth beneath the armature.
In the Mackenzie basin “var. canadensis” is by far the most abundant, though
“var. strigosus” is actually widespread within the same area. In the east, how-
ever, the former is somewhat more northern in its range, and should perhaps be
kept separate.

Rubus Chamaemorus L. Sp. Pl. 494 (1753); Hooker, Fl. Bor.-Am. 1: 183 (1834);
Macoun, Cat. Can. Pl. Pt. 1: 128 (1883); Preble in N. Am. Fauna 27: 526 (1908);
Raup in Jour. Arn. Arb. 17: 265 (1936).

Resolution, Kennicott (N); same place, Miss E. Taylor (G, N, Can.) ; southwest and
north shores of Great Slave L., Howe 91975 (Can.); mouth of Little Buffalo R., Brooke
(Can.) ; 12 mi. E. of Moraine Pt., Great Slave L., Bedford (Can.); near Rae, Russell
(Can.); Liard R. between Nahanni Butte and Simpson, Crickmay 59 (Can.); Willow
Lake R., 44 mi. above mouth, Crickway 19 (Can.) ; muskeg on south shore of Brintnell
L., 9322.

Common in muskegs; collected in flower at Brintnell L. the last of June.
Noted by Hooker “From Lake Winipeg . . . to the shores of the Arctic Sea,”
and Preble gives it a similar range.

Rubus pubescens Raf. Med. Repos. 2: 333 (1811); Raup in Jour. Arn. Arb. 17: 265
(1936).
Slave R., 100 mi. below Ft. Smith, Preble & Cary 14 (US); east shore of Trout L.,
C. O. Hage (G); Mackenzie R., Kennicott (N); Simpson, Onion (N); damp lowland
woods at Simpson, 9094; Willow Lake R., 44 mi. above mouth, Crickmay 15 (Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 197

Common northward at least as far as Simpson, and collected in the Mackenzie
delta by Porsild (Sargentia 4: 48). In flower in late May and the first half of
June.

Rubus acaulis Michx. Fl. Bor.-Am. 1: 298 (1803); Hooker, FI. Bor.-Am. 1: 182 (1834);
Raup in Jour. Arn. Arb. 17: 265 (1936); Porsild in Nat. Mus. Can. Bull. 101:
23 (1945).
Rubus arcticus L. var. grandiflorus sensu Macoun, Cat. Can. PI. Pt. 1: 129 (1883), non
Ledeb.

Rubus arcticus sensu Preble in N. Am. Fauna 27: 526 (1908), non L.

Fort Smith, Miss E. Taylor (G, Can.) ; Little Buffalo R., Russell 79 (G); lower Slave
R., Brooke (Can.); Resolution, Kennicott (N); Hay River, Preble & Cary 20 (US);
southwest and north shores of Great Slave L., Howe 91974 (Can.); northwest shore of
Great Slave L., Bedford (Can.); Liard R. between Nahanni Butte and Simpson, Crickmay
60 (Can.) ; Simpson, Kennicott (N) ; same, Onion (N); damp place in lowland woods near
Simpson, 9106; Willow Lake R., 44 mi. above mouth, Crickmay 18 (Can.); damp moss
in woods on shore of Brintnell L., 9179; Nahanni Mts., Preble & Cary 60° (US).

Common; in flower about mid-June. Porsild notes it as common in Macmillan
Pass, along the Canol Rd. Its range is given in FI. Bor.-Am., “From the Sas-
katchewan. . . to Fort Franklin. .. .” Preble states that it has about the same
range as R. Chamaemorus in the Mackenzie basin.

I have seen a specimen from Hay River, on the southwest shore of Great Slave
L. (Preble & Cary 25, US), which appears to be intermediate between FR. pu-
bescens and R. acaulis. This form has been-observed before, and is regarded by
some students as a hybrid. L. H. Bailey, however, has described such an inter-
mediate from central Saskatchewan and Alberta as R. paracaulis (Gent. Herb. 5:
33, 1941). The status of the Hay River plant must remain uncertain until fur-
ther material is available.

Fragaria glauca (Wats.) Rydb. Mem. Dept. Bot. Columbia Univ. 2: 183 (1898); Raup
in Jour. Arn. Arb. 17: 266 (1936).
Fragaria virginiana Duchesne ? glauca Wats. Bot. King’s Expl. 85 (1871).
Fragaria virginiana sensu Hooker, FI. Bor.-Am. 1: 184 (1834), in part; Macoun, Cat.
Can. Pl. Pt. 1: 135 (1883), in part; non Duchesne.

Fragaria canadensis sensu Hooker, FI. Bor.-Am. 1: 185 (1834), in part, non Michx.

Fragaria pauciflora Rydb. in Mem. Dept. Bot. Columbia Univ. 2: 183 (1898), at least

in part.

Forth Smith, Raup 969; same place, Miss E. Taylor 7169 (Can.) ; same place, Seton &
Preble 78574 (Can.) ; Little Buffalo R., Russell 37 (G) ; east shore of Trout L., C. O. Hage
(G); Simpson, Onion (N); old field at Simpson, 9128; damp thickets near west end of
Brintnell L., 9437.

Occasional to common; found flowering at Simpson about mid-June, and at
Brintnell L. early in July. Noted in Fl. Bor.-Am. in the “Woody country be-
tween lat. 52° and 64°.” Our no. 9437 is referred to F. glauca with some ques-
tion, for it has unusually narrow, obovate-cuneate leaflets (some of them 5:2
cm.). This tendency can be seen in occasional specimens throughout the range
of the species, but in our plants nearly all of the. leaves show’ it.

Fragaria vesca L. var. americana Porter in Bull. Torr. Bot. Cl. 17: 15 (1890).

Resolution, Preble 208, 209 (US); along road in damp lowland woods near Simpson,
9087.

Occasional ; found in flower June 11.

198 SARGENTIA [6

Potentilla arguta Pursh, Fl. Am. Sept. 736 (1814); Hooker, Fl. Bor.-Am. 1: 186 (1834);
Macoun, Cat. Can. Pl. Pt. 1: 136 (1883); Raup in Jour. Arn. Arb. 17: 266 (1936).

Drymocallis agrimonioides (Pursh) Rydb. in N. Am. Fl. 22: 368 (1908).

This species has not been collected in recent years north of the Wood Buffalo
Park, where’it is common in dry prairies and rock crevices. Hooker gives its
range in Fl, Bor.-Am. as “Woody country as far as lat. 65°, north... .” Ryd-
berg states that it extends to Simpson.

Potentilla norvegica L. Sp. Pl. 499 (1753); Hooker, FI. Bor.-Am. 1: 193 (1834);
Macoun, Cat. Can. Pl. Pt. 1: 136 (1883); Raup in Jour. Arn. Arb. 17: 266 (1936).

Fort Smith, Raup 966; Resolution, Preble 214 (US); southwest and north shores of
Great Slave L., Howe 91973 (Can.) ; east shore of Trout L., C. O. Hage (G); Providence,
Dutilly 68 (G) ; Liard R. between Nahanni Butte and Simpson, Crickmay 62 (Can.) ; damp
place in open field at Simpson, 9851 ; along dry watercourse on south-facing slope of granite
mountain west of Brintnell L., 9478.

Apparently rather common in the low country, but rare or occasional at Brint-
nell L.; in flower about mid-July. Noted in Fl. Bor.-Am, as occurring
“Throughout Canada; as far north as the Bear Lake.”

Potentilla diversifolia Lehm. var. glaucophylla (Lehm.) Lehm. Stirp. Pug. 9: 44 (1851);
Porsild in Nat. Mus. Can. Bull. 101: 23 (1945),

Potentilla glaucophylla Lehm. Delect. Sem. Hort. Bot. Hamb. 1836: 7 (1836).

Liard R. between Nahanni Butte and Simpson, Crickmay 66 (Can.) ; alpine meadows
and turfy shale slopes, south side of Red Mt., alt. about 5000’, 9208, 9724; along small brook
on west slope of Terrace Mt., alt. 4800-5000’, 9545 > Macmillan Pass, Canol Rd., mi. 225 E.,
Porsild 11245, 11246 (Can.).

Common on south- and west-facing slopes above Brintnell L., but not seen
elsewhere in that area; collected in flower in late June and mid-July. Porsild
reports it as common in Macmillan Pass,

Potentilla emarginata Pursh, Fl. Am. Sept. 353 (1814); Porsild in Nat. Mus. Can. Bull.
101: 23 (1945).

High alpine rock crevices, east slopes of Red Mt., alt. 5500’-5900’, 9293, 9726; shale
slopes on west side of Terrace Mt., alt. 5000’-6000’, 9520; Bolstead Cr., Canol Rd., mi. 111
E., alpine ridge, alt. 6000’, Wynne-Edwards 8337 (Can.).

Apparently rare or occasional in the Brintnell L. area; in flower late in June,
and with maturing achenes by mid-July.

Potentilla nivea L. Sp. Pl. 499 (1753); Hooker, Fl. Bor.-Am. 1: 195 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 139 (1883); Raup in Jour. Arn. Arb, 17: 267 (1936).

Liard R. between Nahanni Butte and Simpson, Crickmay 64 (Can.) ; alpine meadows
and loose shale, south slope of Red Mt., alt. 5500’, 9209, 9757; shale cliffs on south slope of
Terrace Mt., alt. 5000’, 9682; Nahanni Mts., Preble 314 (US).

Common locally, but not widely distributed at Brintnell L.; observed only
on south-facing slopes. In flower in the latter part of June. Noted in FI.
Bor.-Am, “. . . to the shores of the Arctic Sea.” Most of the specimens cited
above appear to represent var. subquinata Lange (Consp. Fl. Groenl. 9, 1880)
(= P. nipharga Rydb. in N. Am. FI. 22: 332, 1908).

Potentilla biflora Willd. in Schlecht. Ges. Nat. Freunde Berlin Mag. 7: 297 (1815);
Porsild in Nat. Mus. Can. Bull. 101: 23 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., on rocky slope with Cassiope tetragona, alt.
6000’, Porsild 11814 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., on alpine ridge, alt. 5500’,
W ynne-Edwards 8339, 8340 (Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 199

Potentilla elegans Cham. & Schl. in Linnaea 2: 22 (1827); Porsild in Nat. Mus. Can.
Bull. 101: 23 (1945).

Alpine rocky slopes south of Little Keele R., Canol Rd., mi. 55 E., alt. 4000’, Porsild
11786 (Can.).

Potentilla uniflora Ledeb. in Mém. Acad. St. Pétersb. 5: 543 (1812); Porsild in Nat.
Mus. Can. Bull. 101: 23 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 67 (Can.); shale cliffs and
slide rock, south slope of Red Mt., alt. 5500’, 9753; Lone Mt., Wynne-Edwards 8486 (Can.) ;
Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11815 (Can.); Bolstead Cr., Canol Rd.,
mi. 111 E., Wynne-Edwards 8338 (Can.).

Common at the site of the single Brintnell L. collection, but not seen else-
where in that area; found with maturing achenes and a few late flowers Aug. 7.

Potentilla multifida L. Sp. Pl. 496 (1753); Raup in Jour. Arn. Arb. 17: 267 (1936).
Fort Smith, Seton & Preble (N, Can.) ; Liard R. between Nahanni Butte and Simpson,
Crickmay 68 (Can.) ; roadways and dooryards at Simpson, 9142; Norman, Dutilly 55 (G).
A common weed at Simpson; found with a few early flowers in mid-June. I
have seen a specimen also from Good Hope ( Miss E. Taylor, Can.).

Potentilla pennsylvanica L. Mant. 76 (1767); Hooker, Fl. Bor.-Am. 1: 187 (1834);
Macoun, Cat. Can, Pl. Pt. 1: 136 (1883); Raup in Jour. Arn. Arb. 17: 267 (1936).

Resolution, Onion, Kennicott & Hardisty (N); dry bank in open field at Simpson, 9850.

Common in this habitat at Simpson; achenes mature in late August. Noted in
Fl. Bor.-Am. as occurring “About the Saskatchewan, and as far north as the
Bear Lake... .”’ Porsild (Sargentia 4: 48) has collected it in the Richardson
Mts. and in the Eskimo Lakes basin.

Potentilla pectinata Raf. Aut. Bot. 164 (1840); Raup in Jour. Arn. Arb. 17: 268 (1936);
Porsild in Sargentia 4: 48 (1943).

Liard R. between Nahanni Butte and Simpson, Crickmay 69 (Can.).

This species is apparently widespread in the Mackenzie basin, though scattered.
It has been found at a few places in the Athabaska—Great Slave L. region, and
Porsild has reported a plant from the arctic coast which closely resembles it.

Potentilla Anserina L. Sp. Pl. 495 (1753); Hooker, Fl. Bor.-Am. 1: 189 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 141 (1883); Raup in Jour. Arn. Arb. 17: 268 (1936). See
Fernald in Rhod. 35: 273 (1933).

Fort Smith, Raup 943; along the Nyarling R., Russell 52 (G); Resolution, Miss E,
Taylor 100931 (Can.); Hay River, Hume 102668 (Can.); same place, Preble & Cary 23
(US); Liard R. between Nahanni Butte and Simpson, Crickmay 63 (Can.); Simpson,
Onion (N); same, Onion, Kennicott & Hardisty (N); cleared ground at Simpson, 9116;
Mackenzie R. near Wrigley, Miss E. Taylor 98 (Can.) and 34 (G); Willow Lake R., 44
mi. above mouth, Crickmay 5 (Can.).

An abundant weed in villages; in flower about mid-June. Noted in FI. Bor.-
Am. from “Lake Huron to the extreme Arctic regions. .. .”’ Forma sericea
(Hayne) Fernald (1. c.) is of common occurrence with the typical species.

Potentilla palustris (L.) Scop. Fl. Carn. ed. 2, 1: 359 (1772); Hooker, Fl. Bor.-Am. 1:
187 (1834); Macoun, Cat. Can. Pl. Pt. 1: 140 (1883); Raup in Jour. Arn. Arb.
17: 268 (1936); Porsild in Nat. Mus. Can. Bull. 101: 23 (1945).

Comarum palustre L. Sp. Pl. 502 (1753).

Rae, Russell (Can.) ; noted by Porsild in ponds in Macmillan Pass along the Canol Rd.,
mi. 225 E.

Hooker gives the range of this species as extending “. .. as far north as
Bear Lake.” There are two specimens in Herb. N. collected by Onion, Kennicott

200 SARGENTIA [6

& Hardisty without locality data, but they probably came from the upper Mack-
enzie. I have seen a specimen from Great Bear L. (J. M. Bell, Can.).

Potentilla fruticosa L. Sp. Pl. 495 (1753); Hooker, Fl. Bor.-Am. 1: 186 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 141 (1883); Raup in Jour. Arn. Arb. 17: 268 (1936); Porsild
in Nat. Mus. Can. Bull. 101: 23, 31 (1945).

Dasiphora fruticosa (L.) Rydb.; Preble in N. Am. Fauna 27: 527 (1908).

Fort Smith, Raup 961; Little Buffalo R., Russell 88 (G); southwest and north shores of
Great Slave L., Howe 91972 (Can.) ; northwest shore of Great Slave L., Bedford (Can.);
Rae, Russell (Can.); Great Slave L., R. Bell 23158 (Can.); same, Kennicott (N); east
shore of Trout L., C. O. Hage (G); Providence, Preble & Cary 35, 49 (US); Mackenzie
R., Kennicott (N); Liard R. between Nahanni Butte and Simpson, Crickmay 65 (Can.) ;
Wrigley, Miss E. Taylor 32 (N), 41 (G); muskeg thickets at west end of Brintnell L.,
9251; woods on shore of Brintnell L. near Camp Pt., 9183; rocky south and east slopes of
Red Mt., in scrub timber and near summit, 9213, 9727; north peak of Nahanni Mt., Wynne-
Edwards 8485 (Can.).

Common throughout the region; found flowering from late June to mid-
August. Preble gives its range as “. . . throughout the region north to the
limit of the forest.” I have seen specimens from Great Bear L. (J. M. Bell,
Can.) and the lower Mackenzie (Onion, N). Porsild reports the species com-
mon all along the Canol Rd.

Potentilla tridentata Ait. Hort. Kew. 2: 216 (1789); Hooker, Fl. Bor.-Am. 1: 195
(1834); Macoun, Cat. Can. Pl. Pt. 1: 141 (1883); Raup in Jour. Arn. Arb. 17:
268 (1936).

This species has not been collected in recent years northward of Lake Atha-
baska and the Wood Buffalo Park, but in Fl. Bor.-Am. its range is noted as
follows: “. . . Canada, and throughout the woody country, from Lake Huron

. as far as lat. 64°.” There is a Richardson specimen in Herb. N., but it is
without locality data.

Sibbaldia procumbens L. Sp. PI. 284 (1753); Porsild in Sargentia 4: 49 (1943), in Nat.
Mus. Can. Bull. 101: 23 (1945).

Thickets on stony beach at Camp Pt., Brintnell L., 9169; along small brook on east slope
of Red Mt., alt. 4500’, 9271.

Occasional to common in the Brintnell L. area, and noted by Porsild as very
common in Macmillan Pass along the Canol Rd. Found with maturing achenes
at low elevations on June 20, and with flowers higher in the mountains late in
June.

Geum macrophyllum Willd. var. perincisum (Rydb.) Raup in Rhod. 33: 176 (1931),
in Jour. Arn. Arb. 17: 269 (1936); Porsild in Nat. Mus. Can. Bull. 101: 23 (1945).

Geum perincisum Rydb. in N. Am. FI. 22: 405 (1913).

Geum oregonense sensu Rydb. in op. cit. 404, non Scheutz.

Fort Smith, Raup 922, 925, 926; lower Slave R., Raup 924; Mackenzie R., Onion, Kenni-
cott & Hardisty (N); Simpson, Onion, Kennicott & Hardisty (N); upland prairie west of
the Mackenzie R. near Simpson, 9890; gravelly and marshy thickets on south shore of
Brintnell L., 9331, 9493; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11241 (Can.).

Common at the Simpson and Brintnell L. stations; in flower at the latter

place the last of June.

Geum aleppicum Jacq. var. strictum (Ait.) Fernald in Rhod. 37: 294 (1935).
Geum strictum Ait. Hort. Kew. 1: 217 (1789); Raup in Jour. Arn. Arb. 17: 269 (1936).
Simpson, Miss E. Taylor 122 (Can.); damp thickets at Simpson, 9845.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 201

Common at Simpson; in fruit in the latter part of August. This appears to
be a northwestern extension of the range of this species. It is occasional in the
Wood Buffalo Park and southward.

Dryas Drummondii Richardson in Hook. Bot. Mag. pl. 2972 (1830); Hooker, Fl. Bor.-
Am. 1: 174 (1834); Macoun, Cat. Can. Pl. Pt. 1: 132 (1883); Preble in N. Am.
Fauna 27: 528 (1908); Raup in Jour. Arn. Arb, 17: 269 (1936); Porsild in Nat.
Mus. Can. Bull. 101: 23, 31, 33 (1945).

Resolution, Kennicott (N); Jones Pt., Great Slave L., Hume 102669 (Can.) ; northwest
shore of Great Slave L., Bedford (Can.) ; Simpson, Onion (N); outwash plain near mouth
of North Nahanni R., Wynne-Edwards 8537 (Can.) ; gravel plains in low ground near west
end of Brintnell L., 9443; rock-slides on north slope of Colonel Mt., alt. 5000’, 9631; Lone
Mt., Wynne-Edwards 8487 (Can.); Nahanni Mts., Preble & Cary 79 (US); Sekwi R.,
Canol Rd., mi. 174 E., Porsild 11868 (Can.).

Common to abundant on the low gravel plains at Brintnell L., where it forms’
broad mats to the exclusion of nearly all other species; only occasional on high
mountain slopes; in flower early in July. Noted in Fl. Bor.-Am., “In the woody
country between lat. 54° and 64°, and about Slave Lake and Bear Lake, as far
as the mouth of the River Bear Lake on the shores of the Arctic Sea in lat. 68° ;”
and by Preble on the “. . . summit of the Nahanni Mountains. .. .”

Dryas integrifolia Vahl in Skr. Nat. Selsk. Kjgb. 42: 171 (1798); Hooker, FI. Bor.-Am.
1: 174 (1834); Macoun, Cat. Can. Pl. Pt. 1: 132 (1883); Preble in N. Am. Fauna
27: 527 (1908); Raup in Jour. Arn. Arb. 17: 269 (1936); Porsild in Nat. Mus. Can.
Bull. 101: 23, 31 (1945).

Great Slave L., R. Bell 23156 (Can.); southwest and north shores of Great Slave L.,
Howe 91971 (Can.); northwest shore of Great Slave L., Bedford (Can.); Liard R. be-
tween Nahanni Butte and Simpson, Crickmay 61 (Can.); Lone Mt., W ynne-Edwards 8488,
8489 (Can.); on Cap Mt., 4000’ above Mackenzie R., Rk. A. Brooke (Can.); steep rocky
slopes and gravel beaches, south shore of Brintnell L., 9324, 9342; newly-formed mud-flats
at west end of Brintnell L., 9811; rock-slides on north slope of Colonel Mt., alt. 5000’, 9628 ;
Bolstead Cr., Canol Rd., mi. 111 E., on alpine ridge, alt. 5000’-5500’, Wynne-Edwards 8336
(Can.) ; Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11816 (Can.); Sekwi R., Canol
Rd., mi. 174 E., Porsild 11869, 11870 (Can.).

One of the most abundant and characteristic species of high alpine slide rock,
upon which it exerts a stabilizing influence by the formation of dense mats.
Though generally distributed over all the mountains of the Brintnell L. area, it
appeared most abundant on the north slope of Colonel Mt. and the steep cliffs
on the northeast side of Red Mt. It is also abundant on cliffs and ledges on
the lower timbered slopes of Colonel Mt. On parts of the newly-formed mud-flats
at the mouth of the glacial stream it forms dense broad mats. In flower in late
June or early July. Porsild reports it common in many places along the Canol
Rd., and it is noted in Fl. Bor.-Am. as occurring “In dry rocky places from
lat. 54° to the shores and islands of the Arctic Sea... .”’ Preble states that
at the level of the Mackenzie River he first saw it, on his way upstream, at Good

Hope.

Dryas octopetala L. Sp. Pl. 501 (1753); Porsild in Nat. Mus. Can. Bull. 101: 23 (1945).

Meadows and shaly slopes, south side of Red Mt., alt. 5000’-5500’, 9211, 9750; high shale
rock-slides on west slope of Terrace Mt., alt. 6000’, 9556; Bolstead Cr., Canol Rd., mi. 111
E., alpine slope, Wynne-Edwards 8334 (Can.).

An abundant primary species on some high-alpine, semi-stabilized slide rock
at Brintnell L., particularly in the mountains north of the lake, where it appears
to supersede D. integrifolia. It was not observed below timber line, and was
collected in full flower June 21.

202 SARGENTIA [6

Forms intermediate between D. octopetala and D, integrifolia are common.
They have been variously designated by different authors. Polunin regards
those he saw in the eastern Arctic as D. integrifolia £. intermedia ( Nath.)
Polunin, while Porsild states that one of his numbers cited under D. integrifolia
(11870) is the var. intermedia Nath. of that species. Still another view is that
of Hartz, who makes integrifolia a subspecies of D. octopetala, and intermedia
a form of the subspecies. Rydberg, in N. Am. Fl, 22: 400, considered all the
intermediates simply as hybrids of D. octopetala and D. integrifolia, and did not
give them a separate name. Among our Brintnell Lake numbers 9556 and 9211
show some intermediate characters.

Rosa acicularis Lindl. Ros. Monogr. 44 (1820); Preble in N. Am. Fauna 27: 528 (1908);
Raup in Jour. Arn. Arb. 17: 270 (1936).

Rosa majalis sensu Borrer in Hooker, Fl. Bor.-Am. 1: 200 (1834), at least in part, non
Herrm.

Fort Smith, Seton & Preble 78325 (Can.) ; same place, Raup 901; Resolution, Preble 223
(US) ; 12 mi. E. of Moraine Pt., Great Slave L., Bedford (Can.) ; near Rae, Russell (Can.) ;
east shore of Trout L., C. O. Hage (G); Mackenzie R., Richardson (G); Providence,
Preble & Cary 29 (US); Simpson, Onion, Kennicott & Hardisty (N); Wrigley, Miss E.
Taylor 69 (Can.) ; Willow Lake R. at main Forks, 50 mi. above mouth, Crickmay 23 (Can):
Nahanni Mts., Preble & Cary 58 (US); mossy spruce woods on Camp Pt., Brintnell L.,
shrubs 1’-4’ high, 9304, 9812, 9813, 9814, 9816; thickets on gravel plain near west end of
Brintnell L., shrub 3’ high, 9808; open spruce woods on rocky-sandy knoll near east end of
Brintnell L., shrub 2’ high, 9583; clearing in spruce woods along north shore of lake just
east of Camp Pt., shrubs 1-3’ high, 9817.

Common to abundant throughout the forested parts of the region. At Brint-
nell L. it was observed as high as 4800’ on the warm southerly slopes of Red Mt.
In flower late in June, and with maturing fruit about mid-August.

The wild roses of the Brintnell L. region proved to be extremely variable, espe-
cially in the shape of the ripened fruits. As soon as the latter began to mature,
collections were made to illustrate as many forms as possible. All of them are
quite clearly in the acicularis group, the variations of which have been studied
with great care by Mrs. Erlanson (See Papers Mich. Acad. Sci. Arts & Lett. 5:
77-94, 1925). The following excerpts from my field notes will illustrate the
confusion in our material:

“In the mossy spruce woods west of the creek (Frost Creek, which enters the
lake at Camp Pt.) the fruits are mostly elongate-elliptic or cylindric (no. 9812),
but occasionally one finds a spherical fruit.” “East of the creek, in the mossy
spruce woods at the margin of an old stream channel area, we found a form
with distinctly pear-shaped hips (no. 9816). Here also spherical fruits some-
times appear (no. 9814). Along the lake east of camp is a shrubby clearing in
the spruce woods. It is grown up to Alnus crispa, Ribes triste, R. hudsonianum,
and Rosa. Practically all of the last have robust spherical fruits with scarcely
any elongation (no, 9817).”

It will be noted that in this small area (probably not over 1/, square mile),
there appeared to be a certain amount of local segregation into groups with
spherical, or with pyriform to cylindric or elliptic fruits. On one bush, how-
ever, no. 9513, all of these forms were found! Using Mrs. Erlanson’s suggested
organization of the group, which involves also the glandularity and pubescence
of the leaves, nos. 9814 and 9817 would represent var. Bourgeauiana Crépin, while
nos. 9583, 9808, 9812, 9816 would fall into var. lacorum Erlanson (1..6)y00r

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 203

possibly into typical R. acicularis, which has obovate fruits. Similar mixtures,
even within small areas, have been noted by Mrs. Erlanson.

It should be mentioned that the forms with elongated fruits are much the
most abundant in our region. They are also common in the Athabaska L. district
(Raup, 1936, p. 270), where definitely cylindric fruits have been found. It is
doubtful, however, if these variations can be maintained as varieties or subspecies
in a geographic sense until more is known of their distribution.

Still another form in the R. acicularis group, described below, grows on another
type of site in the Brintnell L. area, and appears to make no genetic contribution
to the mixture noted above. At any rate nothing approaching the shape of its
fruits is to be seen in the general mixture.

KE ROK

Fic. 15. Rosa acicularis var. cucurbiformis Raup, var. nov.

Rosa acicularis Lindl. var. cucurbiformis var. nov. Fic. 15.

A forma typica fructibus longis ampullaceis (2.5 cm. longis et 1 cm. crassis)
cum collo attenuato longitudine fructus trientem saepe aequante et plerumque
deflexo differt.

Mossy open spruce woods at margin of gravel fan on the lower north slope of Colonel
Mt., near the south shore of Brintnell L., shrubs 1’-3’ high, 9826 (Type) ; thicket on rocky
north-facing hillside, Tanana valley, Alaska, along Alaska Highway near mi. 1380, H. M-:
& L. C. Raup 12662.

204 SARGENTIA [6

Common at the Brintnell L. site, but not seen elsewhere except at the Alaska
station above noted. Fruit maturing in mid-August.

In other characters these plants are clearly related to R. acicularis s. 1. Their
leaves have glandular serrate teeth and are glandular-granuliferous and somewhat
pubescent beneath especially on the veins and mid-rib. Among the segregates
recognized by Mrs. Erlanson (1. c.) they are most nearly related to var. lacorum
or to R. Engelmanii S. Wats. However, the form of the fruits seems entirely
unique. There is a tendency, particularly on the upper leaves, to sharply acumi-
nate leaflets. This tendency can be seen also in Alaskan and Eastern Asiatic
material of FR. acicularis, and may eventually require further recognition ; but ex-
cept for the cited specimens I have seen no forms that have the peculiar fruits de-
scribed here.

Rosa Woodsii Lindl. Ros. Monogr. 21 (1820); Borrer in Hooker, Fl. Bor.-Am. 1: 199
(1834), in part; Raup in Jour. Arn. Arb. 17: 270 (1936).

Rosa blanda sensu Borrer in Hooker, Fl. Bor.-Am. 1: 199 (1834), in part; Porsild in
Nat. Mus. Can. Bull. 101: 34 (1945); non Ait.

Willow thickets on sand bars along Mackenzie R. near Simpson, 9864; thickets on rocky-
sandy bank of Mackenzie R. opposite Simpson, shrubs 1’-2’ high, 9938; Wrigley, Wynne-
(on) 8602 (Can.) ; Mackenzie R. between Wrigley and Blackwater R., Crickmay 135
(Can.).

Common in these habitats; fruit maturing in late August. Both R. Woodsii
and &. blanda are noted in FI. Bor.-Am. as ranging northward as far as Great
Bear L., but the Mackenzie plants I have seen are fairly uniform and similar
to those of the Wood Buffalo Park. The specimens in our no. 9938 and in
Wynne-Edwards’ no. 8602 are notable for having only 1 or 2 flowers together in
an inflorescence. This is not unusual in the more southern parts of the range
of Rk. Woodsii, but along the Mackenzie it is very common.

Prunus virginiana L. var. demissa (T. & G.) Torr. in Wilkes, U. S. Explor. Bxp. ys
284 (1874).

Cerasus demissa Nutt. ex T. & G. Fl. N. Am. 1: 411 (1840).
Prunus demissa (Nutt.) D. Dietr.; Raup in Jour. Arn. Arb. 17: 270 (1936).

Cerasus virginiana L. sensu Hooker, Fl. Bor.-Am. 1: 169 (1834), in part; Macoun, Cat.
Can. Pl. Pt. 1: 125 (1883), in part.

Cerasus serotina sensu Hooker, Fl. Bor.-Am. 1: 169 (1834), in part, non Ehrh.

Both Cerasus virginiana and C. serotina are noted in Fl. Bor.-Am. as far
north as lat. 62°, the first to “Great Slave Lake, lat. 62°, where it attains a
height of five feet only,” and the second simply, “as far as north lat. 62°.” Both
are probably referable to P. virginiana var. demissa, but I have seen no other
evidence that this plant extends beyond the lower Athabaska R. Preble and
Seton also placed its northern limit on the lower Athabaska.

Prunus pennsylvanica L. f. Suppl. 252 (1781); Preble in N. Am. Fauna 27: 528 (1908) ;
Raup in Jour. Arn. Arb. 17: 271 (1936).
Liard R. between Nahanni Butte and Simpson, Crickmay 70 (Can.).

LEGUMINOSAE

Lupinus arcticus Wats. in Proc. Am. Acad. 8: 526 (1873); Macoun, Cat. Can. Pl. Pt.
1: 102 (1883); Porsild in Nat. Mus. Can. Bull. 101: 23 (1945).
Liard R. between Nahanni Butte and Simpson, Crickmay 80 (Can.); mossy woods on

north shore of Brintnell L., 9180; open spruce woods on north slope of Colonel Mt., alt.
3000’, 9645; south slope of Red Mt., above timber line, 9221.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 205

Common to abundant in high mountain meadows, particularly on the slopes
north of Brintnell L., and common also throughout the more open timbered areas ;
in flower in late June, and maturing its legumes late in July. Porsild noted it
as common in a muskeg along the Canol Rd. near Little Keele R., mi. HG Sea
have seen specimens from Great Bear L. and the lower Mackenzie.

Astragalus frigidus (L.) Gray var. americanus (Hooker) Wats. Bibl. Ind. 193 (1878);
Macoun, Cat. Can. Pl. Pt. 1: 113 (1883); Raup in Jour. Arn. Arb. 17; 271 (1936).
Phaca frigida L. var. americana Hooker, FI. Bor.-Am. 1: 140 (1830).
Providence, Dutilly (G); Liard R. between Nahanni Butte and Simpson, Crickmay 71
(Can.) ; thicket at margin of upland prairie west of Mackenzie R. near Simpson, 9919.

Common locally ; collected in fruit Aug. 28.

Astragalus frigidus (L.) Gray var. littoralis (Hooker) Wats. Bibl. Ind. 193 (1878);
Porsild in Nat. Mus. Can. Bull. 101: 23 (1945). See Rhod. 41: 250 (1939); Contr.
Lab. Bot. Univ. Mont. 24: 48 (1933).

Phaca frigida L. var. littoralis Hooker, F'. Bor.-Am. 1: 140 (1830).

Open spruce woods near timber line and in alpine meadows, north slope of Colonel Mt.,
alt. 3500’-4300’, 9387, 9658; along small brook on west slope of Terrace Mt., alt. 4800’,
9557; Bolstead Cr., Canol Rd., mi. 111 E., alpine turfy slopes, alt. 4500'-5500’, Wynne-
Edwards 8347, 8348 (Can.).

Abundant on the middle north slopes of Colonel Mt. at Brintnell L., but ap-
parently rare elsewhere in that area. Collected with flowers and immature fruit
early in July, and with ripe legumes late in July.

Astragalus striatus Nutt. in T. & G. FI. N. Am, 1: 330 (1838); Porsild in Nat. Mus.
Can. Bull. 101: 33 (1945).

Astragalus adsurgens sensu Raup in Jour. Arn. Arb. 17: 271 (1936), non Pall.

Outwash gravel plain near mouth of North Nahanni R., Wynne-Edwards 8538 (Can.).

This species was not previously known to extend north of the Wood Buffalo
Park.

Astragalus goniatus Nutt. in T. & G. Fl. N. Am. 1: 330 (1838).

Astragalus hypoglottis sensu Raup in Jour. Arn. Arb. 17: 271 (1936).

Junction of the Nyarling and Little Buffalo Rivers, Russell 5 (G).

Common in the Wood Buffalo Park, and probably having a wider range in the
upper Mackenzie country than the scanty records show. In flower about mid-
June.

Astragalus alpinus L. Sp. Pl. 760 (1753); Macoun, Cat. Can. Pl. Pt. 1: 112 (1883);
Raup in Jour. Arn. Arb. 17: 271 (1936); Porsild in Nat. Mus. Can. Bull. 101: 33,
34 (1945).

Phaca astragalina DC.; sensu Hooker, Fi. Bor.-Am. 1: 145 (1834).

Fort Smith, Raup 994 (Coll. Mrs. Conibear), 995; Lobstick Cr., Little Buffalo R., Rus-
sell 4 (G); Mackenzie R., Onion (N) ; south side of Mackenzie R., 10 mi. below Mills L.,
Wynne-Edwards 8555 (Can.) ; Liard R. between Nahanni Butte and Simpson, Crickmay
77,78 (Can.) ; rocky-sandy bank of Mackenzie R. opposite Simpson, 9974; Wrigley, Wynne-
Edwards 8604, 8605 (Can.) ; Saline R., above Norman, Wynne-Edwards 8572 (Can.) ; high
alpine rock crevice, north slope of Colonel Mt., Brintnell L., alt. about 6000’, 9428.

Apparently rare in the Brintnell L. area, but common in the lower country ; in
flower in the mountains early in July. Our Simpson specimens approach var.
Brunetianus Fernald (Rhod. 10: 51) in habit, with long vine-like zig-zag stems
up to 3.5 dm. in length. The ripe legumes, however, are straight and have the
rather long, black, partly spreading pubescence of the typical form.

206 SARGENTIA [6

Astragalus aboriginum Richardson, Bot. App. Frankl. Journ. 756 (1823); Macoun, Cat.
Can. Pl. Pt. 1: 111 (1883).

Phaca aboriginum (Richardson) Hooker, Fl. Bor.-Am, 1: 143 (1830).
Astragalus vaginatus sensu Hooker, Fl. Bor.-Am. 1: 149 (1834), at least in part, non
Pall.

“Gravelly ridges, north to Bear Lake,” Richardson (Can.); river bluff at Simpson,
? 9113.

There appear to have been no recent collections of this species in the Mack-
enzie basin except along the upper Peace River (Raup in Contr. Arn. Arb. 6:
179). In Richardson’s original description the only indication of range is ““W”
meaning “wooded country from latitude 54° to 64° north.” Hooker, in Fl. Bor.-
Am., gave the range as extending “From Lake Winipeg to the Rocky Mountains,
and as far north as Bear Lake, in lat. 66°.” Astragalus vaginatus is given a
similar northern range, “Woody country, between lat. 54° and 64° north.” Our
specimens from Simpson are doubtfully determined. They were collected about
the middle of June when the inflorescences were just forming. The stems are
nearly glabrous below and finely appressed-pubescent above, and the leaves are
glabrous above.

Astragalus eucosmus Robinson in Rhod. 10: 33 (1908); Raup in Jour. Arn. Arb, 17:
271 (1936); Porsild in Sargentia 4: 50 (1943).

Phaca elegans Hooker, Bor.-Am. 1: 144 (1830).

This species has not yet been collected along the upper Mackenzie, though it
is known in the Wood Buffalo Park and at Great Bear L. (J. M. Bell 22899,
Can.).

Astragalus tenellus Pursh, Fl. Am. Sept. 2: 473 (1814); Raup in Jour. Arn. Arb. 17:
272 (1936).

Phaca nigrescens Hooker, Fl. Bor.-Am. 1: 143 (1830), at least in part.

Astragalus multiflorus (Pursh) Gray, sensu Macoun, Cat. Can. Pl. Pt. 1: 114 (1883),
at least in part.

Rocky-sandy bank of Mackenzie R. opposite Simpson, 9973; “Prairies, Ft. Franklin,
Mackenzie R.,” Richardson (Can.).

Common at the Simpson locality ; found with flowers and mature pods late in
August. I have seen specimens from as far north as Good Hope (Miss E.
Taylor, Can.).

Astragalus linearis (Rydb.) Porsild in Rhod. 41: 250 (1939); Porsild in Sargentia 4:
50 (1943).

Atelophragma lineare Rydb. in Bull. Torr. Bot. Cl. 40: 50 (1913), in N. Am. Fl. 24:
368 (1929).

I have seen no specimens of A. linearis from the central part of the Mack-
enzie basin. However, Porsild collected it in the Richardson Mts. and on the
Coppermine R., and I have seen a specimen of it collected in the Lesser Slave L.
district (Brinkman 4073). It is therefore to be expected along the Mackenzie.

Astragalus yukonis M. E. Jones, Rev. Astr. 83, ¢. 7, f. 28 (1923); Raup in Jour. Arn.
Arb. 17: 272 (1936); Porsild in Sargentia 4: 51 (1943).

Rocky-sandy bank of Mackenzie R. opposite Simpson, 9972; willow thickets on Mackenzie
R. sand bars near Simpson, 9085, 9859.

Common to abundant in these habitats; collected in flower about mid-June,
and with mature legumes late in August. Astragalus yukonis was based upon
plants collected by Gorman (no. 1082) on the Yukon near Ft. Selkirk and by

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 207

Miss Alice Eastwood at Whitehorse. It is probably identical with Rydberg’s
Homalobus retusus, the type of which was also collected at Ft. Selkirk by Tarl-
ton (nos. 128-A, 128-B) (Bull. Torr. Bot. Cl. 50: 186, 1923). The Gorman
number was cited provisionally by Rydberg as identical with his type of Homalo-
bus retusus. Although A. ywkonis antedates the latter by some months, it is not
mentioned by Rydberg in his later monograph (N. Am. FI. 24: 266, 1929).

Both of the above descriptions call for a plant with black-hairy calyces and
legumes, but ours have considerable white hair among the black. This is also
true of specimens collected along the lower Athabaska R. (Raup 990). The
species has also been collected in the Mackenzie delta by Porsild.

Oxytropis retrorsa Fernald in Rhod. 30: 140 (1928); Raup in Jour. Arn. Arb. 17: 272
(1936); Porsild in Sargentia 4: 55 (1943).

This species has not been collected in southwestern Mackenzie, but is to be
expected there. Porsild has collected it at Great Bear L. and in the Eskimo Lakes
basin, while it is known on the lower Athabaska (Raup 1. c.) and in the Peace R.
region (Raup in Jour. Arn. Arb. 23: 20, 1942).

Oxytropis foliolosa Hooker, Fl. Bor.-Am. 1: 146 (1834). See Fernald in Rhod. 30:
139 (1928).

Grayelly thickets at west end of Brintnell L., 9431.
Common on this site, but not observed elsewhere; found with flowers and
maturing pods about mid-July.

Oxytropis hudsonica (Greene) Fernald in Rhod. 30: 142 (1928); Raup in Jour. Arn.
Arb. 17: 272 (1936); Porsild in Sargentia 4: 53 (1943), in Nat. Mus. Can. Bull.
101: 23, 32 (1945).

Aragallus hudsonicus Greene in Proc. Biol. Soc. Wash. 18: 17 (1905).

Gravelly-stony fan on south shore of Brintnell L., 9586; dry rock gully on south-facing
slope of granite mountain west of Brintnell L., 9482; Lone Mt., Wynne-Edwards 8491
(Can.) ; Bolstead Cr., Canol Rd., mi. 111 E.. Wynne-Edwards 8342 (Can.); same place,
Porsild 11832 (Can.).

Occasional to common in the Brintnell L. habitats noted; found with flowers
and mature legumes in late July. Known also from Great Slave L. (Raup, l.c.),
Great Bear L. (J. M. Bell, Can.; Porsild, Can.), the arctic coast (Porsild), and
the Richardson Mts. (Porsild).

Oxytropis Maydelliana Trautv. in Acta Horti Petrop. 6: 16 (1879); Porsild in Nat.
Mus. Can. Bull. 101: 23 (1945). See Fernald in Rhod. 30: 142 (1928).

Alpine crevices, ledges, and rock-slides, north slope of Colonel Mt., 9426; Plains of Abra-
ham, Canol Rd., mi. 82 E., Porsild 11818 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alpine
ridge, alt. 6500’-7000’, Wynne-Edwards 8341 (Can.); Sekwi R., Canol Rd., mi. 174 E.,
Porsild 11872 (Can.).

Very common on rocky slopes and local summits on Colonel Mt., but not seen
elsewhere in the vicinity of Brintnell L.; collected with flowers and maturing fruit
early in July.

Oxytropis podocarpa Gray in Proc. Am. Acad. 6: 234 (1864). See Fernald in Rhod.

30: 154 (1928).
Liard R. between Nahanni Butte and Simpson, Crickmay 76 (Can.).

Oxytropis pygmaea (Pall.) Fernald in Rhod. 30: 153 (1928); Porsild in Nat. Mus. Can.
Bull. 101: 23 (1945).

Astragalus pygmaeus Pall. Sp. Astrag. 66, t. 54 (1800).

208 SARGENTIA (6

Shale crevices, ledges and slide rock on north slope of Colonel Mt., above timber line,
9398, 9636; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 6000’, Wynne-Edwards
8345 (Can.).

Common at the Brintnell L. locality cited, but not seen elsewhere in that re-
gion; collected with flowers and maturing legumes early in July. Noted by
Porsild in the Richardson Mts. and at the Mackenzie delta (Sargentia 4: 55).

Oxytropis hyperborea A. E. Porsild in Sargentia 4: 53 (1943), in Nat. Mus. Can. Bull.
101: 23, 31 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 75 (Can.) (? doubtful) ;
gravelly thickets near west end of Brintnell L., 9449; alpine meadows and open places in
willow-birch scrub, north slope of Colonel Mt., 9427; north peak of Nahanni Mt., Wynne-
Edwards 8490 (Can.) ; Bolstead Cr., Canol Rd., mi, 111 E., alpine bog, alt. 5000’, Wynne-
Edwards 8343, 8344 (Can.).

Common in the Brintnell L. habitats noted; in flower early in July, those at
lake-level with maturing pods July 10.

Oxytropis terrae-novae Fernald in Rhod. 30: 147, pl. 174 (1928); Porsild in Nat. Mus.
Can. Bull. 101: 24 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11817 (Can.) ; Bolstead Cr., Canol Rd.,
mi. 111 E., Porsild 11833 (Can.) ; same place, alpine slope, alt. 6000’, Wynne-Edwards 8346
(Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11873 (Can.).

Found in flower at Bolstead Cr. July 25, and in fruit Sept. 6. Porsild states
that this species “appears to be one of the most common members of this genus
on the upper slopes of mountains east of the Yukon—Mackenzie divide where it
grows in turfy places on limestone barrens.”

Oxytropis campestris (L.) DC. (s. str.) Astrag. 74 (1802); Porsild in Sargentia 4:
52 (1943); ? Porsild in Nat. Mus. Can. Bull. 101: 23 (1945),

Astragalus campestris L. Sp. Pl. 761 (1753).

Oxytropis gracilis sensu Raup in Jour. Arn. Arb, 17: 272 (1936), non Nels.

Northwest shore of Great Slave L., Bedford (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E.,
Porsild 11834 (Can.) (doubtfully determined).

Porsild (Sargentia 4: 52) believes that some northern specimens in this group
are a close match for the northern European O. campestris. Another from the
Mackenzie basin was collected at Good Hope by Miss E. Taylor (5353, Can.).
Two other specimens from Great Slave L. may be referred here tentatively :
southwest and north shores, Howe 91977, 91978 (Can.) (as O. Lamberti in
Raup, 1. c.).

Oxytropis spicata (Hooker) Standley in Contr. U. S. Nat. Herb. 22: 373 (1921).
Oxytropis campestris (L.) DC. var. spicata Hooker, Fl. Bor.-Am, 1: 147 (1834).
Fragmentary specimens collected by Crickmay (73, 74, Can.) along the Liard

R. between Nahanni Butte and Simpson are referred somewhat doubtfully to

this species. They consist only of inflorescenses. I have collected O. spicata

along the upper Peace R., and have seen specimens from near the height of land
between the Peace and the Nelson (Raup in Contr. Arn. Arb. 6: 180). Con-
sequently it might well be expected to appear on the Liard.

Oxytropis viscidula (Rydb.) Tidestr. in Contr. U. S. Nat. Herb. 25: 332 (1925); Raup
in Jour. Arn. Arb. 17: 272 (1936).

Aragallus viscidulus Rydb. in Mem. N. Y. Bot. Gard. 1: 253 (1900).
Northwest shore of Great Slave L., Bedford (Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 209

Oxytropis ixodes Butters & Abbe in Rhod. 45: 2, t. 745 (1943).

Mossy banks in woods on south shore of Brintnell L., 9325; alpine meadows and crevices,
north slope of Colonel Mt., alt. 4000’-6000’, 9399, 9635,

Common to abundant in the above sites; in flower late in June, and found
with mature legumes July 28.

When we collected this species at Brintnell Lake in 1939 it was soon recog-
nized as unique, and a description of it was drawn up. I have seen a cotype of ©
O. ixodes, and Butters & Abbe’s description and plate so nearly cover our
Mackenzie Mountain plants that I do not hesitate to identify the latter with
the Minnesota material. In support of this, and in order further to emphasize
the characters of the species, I venture to include here the original description
that I wrote earlier.

Subacaulescent herb with thick, branched caudex; stipules yellowish-straw-
colored, ovate, acuminate, long-ciliate on the margins, glabrous; leaves 3-15 cm.
long ; leaflets 23-27, rather thick, lanceolate or oblong, 3-11 mm. long, revolute,
ciliate and viscid-glandular at the margins; rachis thinly hirsute with white,
spreading hairs, viscid; scapes equaling or somewhat exceeding the leaves,
loosely hairy with mixed black and white hairs; inflorescence 3- to 10-flowered,
loosely subcapitate; bracts lance-attentuate, black- and white-ciliate, viscid, up
to 1 cm. long; calyx densely black-hairy with a few white hairs mixed, 6-8 mm.
long, tinged with blue at least when dry, the lobes lance-attenuate, 4-5.5 mm.
long, viscid-glandular, thinly black-hairy or glabrous; corolla blue to pale purple,
up to 18 mm. long; legume narrowly ovoid, the body about 1.5 cm. long, black-
and white-hairy and glandular-verrucose, tapering to a long slender beak.

I agree with Butters and Abbe that the affinities of Oxytropis trodes are
clearly with the primarily eastern arctic O. hudsonica (Greene) Fernald (see
Rhod. 30: 142), and with the cordilleran O. viscida Natt. or O. viscidula
(Rydb.) Tidestrom. All of these are characterized by their viscid glandularity.

I have seen a few other northwestern specimens that appear to be identical
with the Mackenzie Mt. plants. They were collected by A. E. Porsild at
Castner Glacier, Alaska Range, no. 495 (see Rhod. 41: 251), and by Miss
Edith Scamman in Mt. McKinley Park, no. 651 (see Rhod. 42: 333). Another
specimen in the Gray Herbarium, collected by Aven and Ruth Nelson in Mt.
McKinley Park, no. 3582, was distributed as O. viscida but belongs here. A
specimen collected by G. M. Dawson on the upper Liard R. may also be refer-
able to O. ixodes.

Oxytropis splendens Dougl.’in Hooker, Fl. Bor.-Am. 1: 147 (1834); Raup in Jour. Arn.
Arb. 17: 272 (1936); Porsild in Nat. Mus. Can. Bull. 101: 34 (1945).

Oxytropis splendens Dougl. var. Richardsonii Hooker, Fl. Bor.-Am. 1: 148 (1834) ;
Macoun, Cat. Can. Pl. Pt. 1: 116 (1883).

First point north of Gypsum Pt., Great Slave L., Hume 102671 (Can.); Liard R. be-
tween Nahanni Butte and Simpson, Crickmay 72 (Can.) ; Saline R., above Norman, Wynne-
Edwards 8573 (Can.).

Noted in Fl. Bor.-Am. as occurring “From Cumberland House . . . to Fort
Franklin and the Bear Lake.”

Hedysarum alpinum L. var. americanum Michx. ex Pursh, Fl. Am. Sept. 2: 484 (1814);
Raup in Jour. Arn. Arb. 17: 273 (1936); Rollins in Rhod. 42: 222 (1940); Porsild
in Nat. Mus. Can. Bull. 101: 32 (1945).
Hedysarum alpinum americanum Michx. Fl. Bor.-Am. 2: 74 (1803).
Hedysarum boreale sensu Hooker, Fl. Bor,-Am, 1: 155 (1834) ; Macoun, Cat. Can. PI.
Pt. 1: 117 (1883).

210 ' SARGENTIA [6

Southwest and north shores of Great Slave L., Howe 91991 (Can.); northwest shore of
Great Slave L., Bedford (Can.) ; Mackenzie R., Onion & Ross (N); Liard R. between
Nahanni Butte and Simpson, Crickmay 79 (Can.); among willows on Mackenzie R. sand
bar near Simpson, 9084; gravelly thickets near west end of Brintnell L., 9442; mossy bank
on south shore of Brintnell L., 9328; mossy ledges in upper part of timber and in alpine
meadows and scrub, north slope of Colonel Mt., alt. 3000’-4000’, 9423, 9637; north peak of
Nahanni Mt., Wynne-Edwards 8492 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alt. 5000’,
alpine slopes, Wynne-Edwards 8350 (Can.).

Common in the Brintnell L. area, but like H. Machenzii it was not seen on the
mountains north of the lake; in flower in late June and early July; fruit maturing
in middle and late July. Noted in Fl. Bor.-Am. “, . . from the Saskatchewan to
the Arctic Circle.” I have seen specimens from Great Bear L. (J. M. Bell, Can.),
and from the lower Mackenzie (McConnell, Miss E. Taylor, Can.).

Hedysarum Mackenzii Richardson, Bot. App. Frankl. Journ. 745 (1823); Raup in Jour.
Arn. Arb. 17: 273 (1936); Porsild in Nat. Mus. Can. Bull. 101: 24 (1945). See
Rollins in Rhod. 42: 237 (1940).

Northwest shore of Great Slave L., Bedford (Can.) ; Mackenzie R., Richardson (Can.) ;
rocky-sandy bank of Mackenzie R. opposite Simpson, 9942; gravel beaches and fans, south
shore of Brintnell L., 9346, 9592; alpine meadows, rock-slides and crevices, north slope of
Colonel Mt., alt. 4500’-5500’, 9390, 9638; Bolstead Cr., Canol R. mi. 111 E., Wynne-
Edwards 8349 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11871 (Can.).

Common on the south shore of Brintnell L. and on Colonel Mt., but not ob-
served north of the lake; in flower early in July, and with mature fruit the last
week in July. Fruiting specimens of this species from the Brintnell L. district
(9592, 9638) have unusually large loments. Rollins (1. c.) gives the size of the
individual articles of the loments as 4-6 mm. wide and 5-8 mm. long. On our
plants they are commonly 7 mm. wide and 9 (sometimes 9.5-10 mm.) long.

Vicia americana Muhl. in Willd. Sp. Pl. 3: 1096 (1803); Hooker, Fl. Bor.-Am. 1: 157
(1834); Macoun, Cat. Can. Pl. Pt. 1: 121 (1883); Raup in Jour. Arn. Arb. 17: 273
(1936); Porsild in Nat. Mus. Can. Bull. 101: 33, 34 (1945).

Fort Smith, Raup 1006; Hay River, Great Slave L., Hume 102670 (Can.) ; Providence,
Dutilly 60 (G); south bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8553
(Can.); Wrigley, Wynne-Edwards 8603 (Can.); Saline R., above Norman, W ynne-
Edwards 8624 (Can.).

Noted in Fl. Bor.-Am. as extending to Great Bear L. The narrow-leaved form
of the species (var. angustifolia Nees) is occasional in the Wood Buffalo Park
(Raup, 1. c.), and has been found along the lower Slave R. (Raup 1009).

Lathyrus ochroleucus Hooker, Fl. Bor.-Am. 1: 159 (1834); Macoun, Cat. Can. PI.
Pt. 1: 123 (1883); Raup in Jour. Arn. Arb. 17: 273 (1936).

Fort Smith, Raup 986; same place, Miss E. Taylor 5507 (Can.) ; same place, Seton &
Preble 78319, 78320 (Can.) ; Resolution, Onion, Kennicott & Hardisty (N); Providence,
Preble & Cary 41 (US); along roadside in lowland woods, Simpson, 9097; Mackenzie R.,
Onion (N).

Noted in Fl. Bor.-Am. as occurring “. . . throughout the whole woody coun-
try to Bear Lake, in lat. 66°.”

GERANIACEAE
Geranium Bicknellii Britton in Bull. Torr. Bot. Cl. 24: 92 (1897); Raup in Jour. Arn.
Arb. 17: 274 (1936). See Fernald in Rhod. 37: 296 (1935).
Fort Smith, Raup 1011.
This species is occasional to common in the Wood Buffalo Park, particularly
in recently burned forest. It is probably widespread in the upper Mackenzie
country.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 211

LINACEAE
Linum Lewisii Pursh, Fl. Am. Sept. 1: 210 (1814); Raup in Jour. Arn, Arb. 17: 274
(1936); Porsild in Nat. Mus. Can. Bull. 101: 34 (1945).

Linum perenne sensu Hooker, Fl. Bor.-Am. 1: 106 (1830); Macoun, Cat. Can. Pl. Pt.
1: 89 (1883); non L.

Mackenzie R., Onion (N) ; Liard R. between Nahanni Butte and Simpson, Crickmay 82
(Can.) ; thickets on rocky-sandy bank of Mackenzie R. opposite Simpson, 9970; Wrigley,
W ynne-Edwards 8608 (Can.) ; Norman, Kindle (Can.); same place, Dutilly (G).

Common at Simpson; found with mature fruit and occasional late flowers Aug.
29. Noted in Fl. Bor.-Am. “. . . as far north as the shores of the Arctic Sea,”
but I have seen no material from beyond Great Bear L. (J. M. Bell, Can.).

CALLITRICHACEAE
Callitriche verna L., emend. Loennr. Obs. crit. pl. Suec. illustr. Upsaliae. 17 (1854);
Porsild in Sargentia 4: 55 (1943).

Callitriche palustris L. sensu Raup in Jour. Arn. Arb. 17: 274 (1936).

Northwest shore of Great Slave L., Bedford (Can.).

Porsild has collected this species in the Keewatin District, at Great Bear L.,
and in the Eskimo Lakes basin. It is probably common throughout the lowland
parts of our region.

Callitriche autumnalis L. Fl. Suec. ed. 2, 4 (1755); Hooker, Fl. Bor.-Am. 1: 217 (1834);
Porsild in Sargentia 4: 55 (1943).

Callitriche hermaphroditica L. sensu Raup in Jour. Arn. Arb. 17: 274 (1936).

This species has not yet been collected in the upper Mackenzie country. It
is common in the Wood Buffalo Park, however, and Porsild has collected it in
several places at Great Bear L. and in the lower Mackenzie region. It is noted
in Fl. Bor.-Am. as occurring “. . . as far north as Bear Lake, lat. 66°.” There
is a Richardson specimen in Herb. Can. marked “North of the Saskatchewan,”
but without further locality data.

EMPETRACEAE

Empetrum nigrum L. Sp. Pl. 1022 (1753); Hooker, Fl. Bor.-Am. 2: 140 (1838); Macoun,
Cat. Can. Pl. Pt. 3: 458 (1886); Preble in N. Am. Fauna 27: 529 (1908); Raup in
Jour. Arn. Arb. 17: 274 (1936); Porsild in Nat. Mus. Can. Bull. 101: 24 (1945).

Southwest and north shores of Great Slave L., Howe 91979 (Can.); 12 mi. E. of
Moraine Pt., Great Slave L., Bedford (Can.); east shore of Trout L., C. O. Hage (G);
mossy thickets along shore of Brintnell L., 9186; alpine meadows on east slope of Red Mt.,
alt. 5000’, 9743; Bolstead Cr., Canol R., mi. 111 E., alpine slopes, alt. 5500’, Wynne-Edwards
8352 (Can.).

Common to abundant on turfy alpine slopes at Brintnell L., and occasional in
the woods at lake-level ; a primary species in parts of the scrub and alpine meadow
communities on the east slope of Red Mt., all the way to the summit. Mature
fruit collected Aug. 7. Noted in Fl. Bor.-Am. “Throughout the woody and
barren country, abundant... .” Preble states that it “occurs nearly throughout
the region,” and that “It reaches its greatest luxuriance on the northern border of
the forest... .”

BALSAMINACEAE
Impatiens Noli-tangere L. Sp. Pl. 938 (1753); Raup in Jour. Arn. Arb. 17: 274 (1936).
See Raup in Nat. Mus. Can. Bull. 74: 149 (1935).

Impatiens fulva sensu Hooker, Fl. Bor.-Am. 1: 117 (1830), in part; Macoun, Cat. Can.
Pl. Pt. 1: 92 (1883), in part; non Nutt.

212 SARGENTIA [6

This species appears to be rather rare in the Mackenzie basin. Tt has been
found around the western end of L. Athabaska (Raup, 1. c.) and in the upper
Peace R. country (Raup in Contr. Arn. Arb. 6: 183). The only basis for in-
cluding it in the present flora is the range given by Hooker in Fl. Bor.-Am.,
“Throughout Canada; and as far north as Bear Lake, lat. 66°.” There is a
Richardson specimen in Herb. Can., originally labeled J. fulva and marked “Arc-
tic Nth. Am.,” but without more specific locality data.

CISTACEAE
Hudsonia tomentosa Nutt. var. intermedia Peck in 45'" Rept. N. Y. State Mus. 26
(1893); Raup in Jour. Arn. Arb. 17: 275 (1936).
Hudsonia tomentosa Nutt. sensu Hooker, Fl. Bor.-Am. 1: 73 (1830), in part; Macoun,
Cat. Can. Pl. Pt. 1: 60 (1883), in part.
Noted in Fl. Bor.-Am. as occurring “as far north as Slave Lake.” It is
abundant on sandy beach ridges and dunes around L. Athabaska, and will prob-
ably be found again in similar situations farther north.

VIOLACEAE

Viola (glean Greene, Pittonia 3: 144 (1896); Raup in Jour. Arn. Arb. 17: 275
(1936).

The only basis for including this species in the present flora is the record in
Fl. Bor.-Am, (2: 75) for the closely related V. cucullata Ait., “Throughout
Canada . . . and as far north as the Mackenzie River and Bear Lake.” Viola
cucullata has never been collected in the Mackenzie basin, but V. nephrophylla
occurs in the Wood Buffalo Park, and in the upper Peace R. region (Raup in
Contr. Arn, Arb. 6: 183). I have seen no Richardson specimens from the north,
however, nor any FI. Bor.-Am. material marked V’. cucullata.

Viola epipsila Ledeb. Ind. Sem. Hort. Dorpat, 5 (1820). See Fernald in Mem. Am.
Acad. 15: 268, footnote 5 (1925).

Damp places along streams, upper valley of Frost Cr., north of Brintnell L., alt>4000’—
4500’, 9287, 9550.

Rare or occasional ; found flowering in late June and mid-July.

Viola renifolia Gray var. Brainerdii (Greene) Fernald in Rhod. 14: 88 (1912); Raup in
Jour. Arn. Arb. 17: 276 (1936); Porsild in-Sargentia 4: 55 (1943), in Nat. Mus.
Can. Bull. 101: 32 (1945).

Viola blanda sensu Hooker, Fl. Bor.-Am. 1: 76 (1830), in part; Macoun, Cat. Can. PI.
Pt. 1: 62 (1883), in part; non Willd.

Mackenzie R., Onion (N); Simpson, Onion, Kennicott & Hardisty (N); same place,
damp places, Miss E. Taylor 2365 (Can.), 25 (G) ; same place, damp lowland woods, 9093,
9107; Lone Mt., Wynne-Edwards 8493 (Can.).

Occasional to common at Simpson; in flower about mid-June. Noted in Fl.
Bor.-Am. as extending “as high north as Fort Franklin, in lat, 66°.” Porsild
has collected it at Great Bear L.

Viola adunca J. E. Sm. in Rees Cyclop. 38: no. 63 (1817); Raup in Jour. Arn. Arb.
17: 276 (1936).

Mackenzie R., Hardisty (N); same, Kennicott (N); Simpson, Preble 310 (US); old
field at Simpson, 9129; Willow Lake R., 44 mi. above mouth, Crickmay 1 (Can.).

Abundant at Simpson; in flower about mid-June.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 213

ELAEAGNACEAE

Elaeagnus commutata Bernh. in Allg. Thuer. Gartenz. 2: 137 (1843); Raup in Jour.
Arn. Arb. 17: 276 (1936); Porsild in Nat. Mus. Can. Bull. 101: 34 (1945).

Elaeagnus argentea Pursh, sensu Hooker, Fl. Bor.-Am. 2: 137 (1838) ; Macoun, Cat.
Can. Pl. Pt. 3: 420 (1886); Preble in N. Am. Fauna 27: 530 (1908) ; non Moench.

Hay River, Great Slave L., Preble & Cary 18 (US); shale bluff of Mackenzie R. oppo-
site Simpson, shrubs up to 4’ high, 9977; Wrigley, Miss E. Taylor (N), 18 (Can.), 13 (G);
Norman, Dutilly 47 (G); Saline R., above Norman, Wynne-Edwards 8574 (Can.).

Common at Simpson; in fruit late in August. Noted in Fl. Bor.-Am. as oc-
curring northward “. . . to lat. 69° near the coast.” Preble does not report it
beyond the Norman district.

Shepherdia canadensis (L.) Nutt. Gen. 2: 240 (1818); Hooker, FI. Bor.-Am. 2: 138
(1838); Macoun, Cat. Can. Pl. Pt. 3: 421 (1886); Raup in Jour. Arn. Arb. 17:
276 (1936); Porsild in Sargentia 4: 55 (1943).

Elaeagnus canadensis L. Sp. Pl. 1024 (1753).

Lepargyrea canadensis (L.) Greene; Preble in N. Am. Fauna 27: 530 (1908).

Fort Smith, Seton & Preble 78389 (Can.); same place, Raup 1034; Bear R., Russell
(G); Resolution, E. A. & A. E. Preble 144 (US) ; southwest and north shores of Great
Slave L., Howe 91982 (Can.); northwest shore, Great Slave L., Bedford (Can.); damp
lowland woods at Simpson, shrub 4’ high, 9083; alpine scrub above timber line on north
slope of Colonel Mt., 9369.

Common in rich woods around Brintnell L. and high up on the slope of Colonel
Mt., but only occasional on the mountains north of the lake; in flower in the
mountains early in July. Noted in Fl. Bor.-Am. “Throughout Canada to Fort
Franklin on the Mackenzie River . . .”; and by Preble “. . . throughout the
country north to the edge of the timber.” I have seen a specimen from Good
Hope (Miss E. Taylor, Can.), and Porsild has collected it in the Mackenzie delta.

ONAGRACEAE

Epilobium angustifolium L. Sp. Pl. 347 (1753); Hooker, FI. Bor.-Am. 1: 205 (1834);
Macoun, Cat. Can. Pl. Pt. 1: 168 (1883); Raup in Jour. Arn. Arb. 17: 277 (1936);
Porsild in Nat. Mus. Can. Bull. 101: 24 (1945).

Fort Smith, Raup 1053 (Mrs. Conibear, coll.) ; Clewi R., Russell 76 (G); Hay River,
Great Slave L., Hume 102672 (Can.); 12 mi. E. of Moraine Pt., Great Slave L., Bedford
(Can.) ; east shore of Trout L., C. O. Hage (G) ; Mackenzie R., Kennicott (N); Liard R.
between Nahanni Butte and Simpson, Crickmay 81 (Can.) ; gravelly thickets on Camp Pts
Brintnell L., 9574; turfy slopes on south side of Red Mt., alt. 5000’, 9763; Nahanni Mts.,
Preble & Cary 56 (US).

Common around Brintnell L., and on the mountains north of the lake, but not
observed on Colonel Mt.; in flower in late July and early August. Noted in FI.
Bor.-Am. “. . . to north lat. 69°.” Porsild states that it is common in Macmillan
Pass, on the Canol Rd.

Our Brintnell L. plants, though tall, have rather short, narrowly lanceolate

leaves, approaching var. intermedium (Wormsk.) Fernald (in Rhod. 20: 5,
1918).

Epilobium latifolium L. Sp. Pl. 347 (1753); Hooker, FI. Bor.-Am. 1: 205 (1834);
Macoun, Cat. Can. Pl. Pt. 1: 169 (1883); Raup in Jour. Arn. Atb. 17:2/77-(1936);
Porsild in Nat. Mus. Can. Bull. 101: 24, 34 (1945).

Northwest shore of Great Slave L., Bedford (Can.); Saline R., above Norman, W ynne-
Edwards 8575 (Can.); muskegs and gravel beaches along shores of Brintnell L., 9348,
9329; open spruce woods on north slope of Colonel Mt., alt. 3000’, 9646; Bolstead Cr., Canol
Rd., mi. 111 E., alpine bog, alt. 4700’, Wynne-Edwards Ssoe (Can).

214 SARGENTIA [6

At Brintnell L. this species is common to abundant on gravel beaches and on
the north slope of Colonel Mt. up to an altitude of about 6000’; rare or occasional
on the mountains north of the lake. In flower late in June, and with maturing
fruit late in July. Porsild reports it as common throughout the route of the
Canol Road through the Mackenzie Mts.

Epilobium palustre L. Sp. Pl. 348 (1753); Raup in Jour. Arn. Arb, 17: 277 (1936);
Porsild in Sargentia 4: 56 (1943), in part.

Little Buffalo R., Russell 76 (G).
Common in the Wood Buffalo Park (Raup in Nat. Mus. Can. Bull. 74: 151),
and known also at Great Bear L. (J. M. Bell, Porsild, Can.).

Epilobium palustre L. var. monticola Haussk. in Oest. Bot. Zeit. 29: 119 (1879); Raup
in Jour. Arn. Arb. 17: 277 (1936).

Epilobium palustre L. var. albiflora Hooker, Fl. Bor.-Am. 1: 207 (1834).

Epilobium palustre L. var. lineare Gray, sensu Macoun, Cat. Can. Pl. Pt. 1: 170 (1883),

at least in part.

This is the small, strict, narrow-leaved form of the species. It is occasional
in the Athabaska L. and Wood Buffalo Park districts, though I have seen no ma-
terial from southwestern Mackenzie. Hooker, in Fl. Bor.-Am., gives its range
as “Throughout Canada, and as far north as lat. 64°... .”

Epilobium glandulosum Lehm. var. adenocaulon (Haussk.) Fernald in Rhod. 20: 35
(1918); Raup in Jour. Arn, Arb. 17: 277 (1936).

Epilobium adenocaulon Haussk. in Oest. Bot. Zeit. 29: 119 (1879).

Epilobium tetragonum sensu Hooker, Fl. Bor.-Am. 1: 206 (1834), in part; Macoun, Cat.
Can. Pl. Pt. 1: 169 (1883), in part; non L.

Northwest shore of Great Slave L., Bedford (Can.) ; bank of Mackenzie R. a few miles
below Providence, 9977 ; muskeg along road on west bank of Mackenzie R. near Simpson,
9911.

Common along the upper Mackenzie; collected with flowers and maturing cap-
sules Aug. 24. Hooker’s record for E. tetragonum, “. . . throughout the plains
to lat. 64°. . . ,” probably should be referred to this species, at least in part.

Epilobium davuricum Fisch. ex Hornem. Suppl. Hort. Bot. Hafn. 44 (1819); Porsild
in Sargentia 4: 56 (1943).

Mossy muskeg thickets at west end of Brintnell L., 9439, 9717.
Common in this habitat; in flower early in July, and in fruit early in August.
Collected in the Mackenzie delta and at Great Bear L. by Porsild.

Epilobium lactiflorum Haussk. in Oest. Bot. Zeit. 29: 89 (1879); Porsild in Nat. Mus.
Can. Bull. 101: 24 (1945),
Gravelly thickets on south shore of Brintnell L., 9334; Macmillan Pass, Canol Rd., mi.
225 E., Porsild 11249 (Can.).

Common locally at Brintnell L., in the habitat mentioned above; in flower June

30.

Epilobium anagallidifolium Lam. Encycl. Méth. 2: 376 (1786); Porsild in Nat. Mus.
Can. Bull. 101: 24 (1945).

Along small brook on west slope of Terrace Mt., north of Brintnell L., alt. 5000’, 9549;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11250 (Can.).

Noted as rare or occasional both at Brintnell L. and Macmillan Pass; collected
in flower July 18.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 215

HALORHAGIDACEAE

Myriophyllum exalbescens Fernald in Rhod. 21: 120 (1919); Raup in Jour. Arn. Arb.
17: 278 (1936); Porsild in Sargentia 4: 56 (1943).

Myriophyllum spicatum sensu Hooker, Fl. Bor.-Am. 1: 216 (1834); Macoun, Cat. Can.
Pl. Pt. 1: 166 (1883) ; non L.

This species has not yet been collected in southwestern Mackenzie, though it
is undoubtedly widespread in the upper Mackenzie valley. It is abundant in the
Wood Buffalo Park, and Porsild has collected it in the Mackenzie delta and in
the Eskimo Lakes basin. In Fl. Bor.-Am. it was noted “. . . as far north as
Bear Lake.”

HIPPURIDACEAE

Hippuris vulgaris L. Sp. Pl. 4 (1753); Hooker, Fl. Bor.-Am. 1: 217 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 167 (1883); Raup in Jour. Arn. Arb. 17: 278 (1936).

Sass Cr., Russell 39 (G); 12 mi. E. of Moraine Pt., Great Slave L., Bedford (Can.) ;
Liard R. between Nahanni Butte and Simpson, Crickmay 83 (Can.) ; Norman, (Coll.?) (N).

Hooker noted this species as occurring northward only to lat. 60°. Its extent
below Norman is unknown.

Hippuris montana Ledeb. in Reichenb. Icon. Bot. 1: 71, t. 86, f. 181 (1823).

In pools in muskeg thickets at west end of Brintnell L., 9720.

Occasional in this habitat, partly floating and partly emergent in pools formed
by beaver dams; only sterile plants found.

ARALIACEAE

Aralia nudicaulis L. Sp. Pl. 274 (1753); Hooker, Fl. Bor.-Am. 1: 274 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 189 (1883); Raup in Jour. Arn. Arb. 17: 278 (1936).
This species is abundant in open woods throughout the Wood Buffalo Park,
and was reported in Fl. Bor.-Am. as extending northward to lat. 64°. I have
seen no specimens, however, from the upper Mackenzie valley.

UMBELLIFERAE

Cicuta Douglasii (DC.) Coult. & Rose in Contr. U. S. Nat. Herb. 7: 95 (1900). See
Mathias & Constance in N. Am. FI. 28B: 155 (1944).

Cicuta occidentalis Greene; Raup in Jour. Arn. Arb. 17: 279 (1936).
Cicuta maculata sensu Hooker, Fl. Bor.-Am. 1: 259 (1834), in part; Macoun, Cat. Can.
Pl. Pt. 1: 181 (1883), in part; non L.

This species is occasional in wet meadows throughout the lower Athabaska
R. region and the Wood Buffalo Park. As C. maculata it was noted in FI. Bor.-
Am. “between lat. 54° and 64° North.” I have seen no specimens from beyond
the 60th parallel, though the species probably extends into the upper Mackenzie
valley.

Cicuta mackenzieana Raup in Jour. Arn. Arb. 17: 279, pl. 197 (1936); Porsild in
Sargentia 4: 56 (1943).

Cicuta virosa sensu Hooker, Fl. Bor.-Am. 1: 259 (1834), in part; Macoun, Cat. Can. PI.
Pt. 1: 182 (1883), in part; non L.

Liard R. between Nahanni Butte and Simpson, Crickmay 84 (Can.).

The material cited under C. virosa in Fl. Bor.-Am., “. . . between lat. 54° and
64°” is probably for the most part referable to this species. It has been collected
as far north as the Eskimo Lakes basin (Porsild, 1. c.).

216 SARGENTIA [6

Sium suave Walt. Fl. Car. 115 (1788); Raup in Jour. Arn. Arb. 17: 280 (1936). See
Rhod. 17: 131 (1915).

Cicuta virosa sensu Hooker, Fl. Bor.-Am. 1: 259 (1834), in part, non L.

Resolution, Kennicott (N); northwest shore of Great Slave L., Bedford (Can.); Rae,
Bedford (Can.) ; sand bar along Mackénzie R. at mouth of small creek opposite Simpson,
9943,

Occasional at the Simpson locality ; fruit mature in late August. A Richardson
specimen of S. suave at Herb. Can. was originally labeled Cicuta virosa, indicat-
ing Hooker’s confusion of these two species.

Heracleum lanatum Michx. Fl. Bor.-Am. 1: 166 (1803); Hooker, Fl. Bor.-Am. 1: 269
(1834); Macoun, Cat. Can. Pl. Pt. 1: 187 (1883); Raup in Jour. Arn. Arb. 17:
280 (1936); Porsild in Nat. Mus. Can. Bull. 101: 24 (1945).

Stony-gravelly shore of Great Slave L. at Wrigley Harbor, 9982; east shore of Trout LG
C. O. Hage (G); thickets along glacial stream near west end of Brintnell L., 9702; Mac-
millan Pass, Canol Rd., mi. 225 E., Porsild 11252 (Can.).

Common along the south shore of Brintnell L. near the western end, but not
seen elsewhere. Fruit mature early in August. Noted in Fl. Bor.-Am. “. .
as far north as the Mackenzie River. . . .”

CORNACEAE

Cornus canadensis L. Sp. Pl. 118 (1753); Hooker, Fl. Bor.-Am. 1: 277 (1834); Macoun,
Cat. Can. Pl. Pt. 1: 190 (1883); Preble in N. Am. Fauna 27: 530 (1908); Raup in
Jour. Arn. Arb. 17: 281 (1936); Porsild in Sargentia 4: 56 (1943), in Nat. Mus.
Can. Bull. 101: 24 (1945).

Fort Smith, Miss E. Taylor (G, N, Can.) ; same place, Seton & Preble 78332 (Can.) ;
Little Buffalo R., Russell 70 (G); Hay River, Great Slave L., Hume 102673 (Can.) ; same
place, Preble & Cary 22 (US); northwest shore of Great Slave L., Bedford (Can.); near
Rae, Russell (Can.) ; east shore of Trout L., C. O. Hage (G); Liard R. between Nahanni
Butte and Simpson, Crickmay 85 (Can.) ; Simpson, Hardisty (N); same place, Onion, Ken-
nicott & Hardisty (N); rich lowland woods at Simpson, 9098; on Cap Mt., Wrigley,
Brooke (Can.); mossy spruce woods on Camp Pt., Brintnell L., 9310; Nahanni Mts.,
Preble & Cary 66 (US).

Abundant in woods throughout the region, and extending up to timber line on
Red Mt. at Brintnell L.; in flower the last of Juné. Porsild noted it as common
in Macmillan Pass, along the Canol Rd., and states that he did not see it at Great
Bear L., although he collected it on Mt. Charles along the Bear River.

Cornus canadensis L. forma purpurascens (Miyabe & Tatewaki) Hara in Rhod. 44:
20 (1942). See also Fernald in Rhod. 43: 156 (1941).

Chamaepericlymenum canadense forma purpurascens Miyabe & Tatewaki in Trans. Sap-
poro Nat. Hist. Soc. 15: 43 (1937).

In willow-birch scrub along stream in upper valley of Frost Cr., north of Brintnell L.,
alt. 4000’, 9506.

Common in this situation but not observed elsewhere; in flower July 18.
This material is peculiar in having the pink color of the bracts arranged in
longitudinal streaks, sometimes from near the base. Occasional plants have the
leaves scattered down the stem, suggesting var. intermedia Farr, but nearly all
of them have only one whorl of leaves.

Cornus stolonifera Michx. Fl. Bor.-Am. 1: 92 (1803); Macoun, Cat. Can. Pl. Pt. 1:
191 (1883); Preble in N. Am. Fauna 27: 530 (1908); Raup in Jour. Arn. Arb. 17:
281 (1936); Porsild in Sargentia 4: 57 (1943).

Cornus alba sensu Hooker, Fl. Bor.-Am. 1: 276 (1834), in part, non L.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 237

Providence, Preble & Cary 48 (US); Simpson, Hardisty (N); same place, Miss E.
Taylor 6 (Can.).; thickets near Simpson, shrubs up to 6’ high, 9855; Bear R., Miss E.
Taylor 74 (Can.); along dry gully on south-facing slope of granite mountain west of
Brintnell L., shrubs up to 5’ high, 9475.

Common in the lowlands and occasional in the mountains ; in flower at Brintnell
L. about mid-July, and with ripe fruit at Simpson the last of August. Preble
states that it is “a conspicuous shrub along streams throughout the wooded coun-
try,” and in Fl. Bor.-Am. it is said to range northward to lat. 69°. Porsild,
however, appears to be doubtful whether it extends to the lower Mackenzie, and
suggests that Hooker’s “69°” is a misprint for 59°.

The plants from Simpson and Brintnell L. have somewhat woolly inflorescences.
thus suggesting var. Bailey: (Coult. & Evans) Drescher; but the hairs on the
lower surfaces of the leaves are nearly all appressed as in the typical form.

PYROLACEAE
Moneses uniflora (L.) Gray, Man. ed. 1: 273 (1848); Macoun, Cat. Can. Pl. Pt. 2: 306

(1884); Raup in Jour. Arn. Arb. 17: 281 (1936); Porsild in Nat. Mus. Can. Bull.
101: 24 (1945).

Pyrola uniflora L. Sp. Pl. 397 (1753) ; Hooker, Fl. Bor.-Am. 2: 45 (1834).

Little Buffalo R., Russell 68 (G); Great Slave L., R. Bell 23203 (Can.) ; southwest and
north shores of Great Slave L., Howe 91962 (Can.); Windy Pt., Great Slave L., Hume
102676" (Can.); Rae, Russell (Can.); Mackenzie R., Kennicott (N), and Onion (N);
Liard R. between Nahanni Butte and Simpson, Crickmay 95 (Can.); Wrigley, Miss E.
Taylor 17 (Can.), 31 (G, N); lowland woods and thickets near west end of Brintnell L.,
9463.

Common; collected in flower July 10 at Brintnell L. Observed by Porsild
along the Canol Rd. at a tributary of Godlin R., mi. 168 E. Noted in FI. Bor.-
Am. “. . . throughout Canada to lat. 64°.”

Pyrola minor L. Sp. Pl. 396 (1753); Porsild in Sargentia 4: 57 (1943), in Nat. Mus.
Can. Bull. 101: 24 (1945).

Along small brook on west slope of Terrace Mt., north of Brintnell L., 9540.

Apparently rare in this area; collected in flower July 18. Porsild notes it as
common in Macmillan Pass, along the Canol Rd. It has been found at L. Atha-
baska, Great Bear L., the Mackenzie delta, and in the Richardson Mts.

Pyrola secunda L. Sp. Pl. 396 (1753); Hooker, Fl. Bor.-Am. 2: 45 (1834); Macoun,
Cat. Can. Pl. Pt. 2: 304 (1884); Raup in Jour. Arn. Arb. 17: 282 (1936); Porsild
in Nat. Mus. Can. Bull. 101: 32 (1945).

Little Buffalo R., Russell 71 (G); Great Slave L., Kennicott (N); northwest shore of
Great Slave L., Bedford (Can.); Liard R. between Nahanni Butte and Simpson, Crickmay
93 (Can.); Wrigley and vicinity, Miss E. Taylor 34, 84, 85 (Can.); Norman, Dutilly 40
(G) ; woods and thickets in lowland at west end of Brintnell L., 9462; turfy shale slope on
north side of Colonel Mt., alt. 4000’, 9653; north peak of Nahanni Mt., Wynne-Edwards
8496 (Can.).

Common in lowland woods, and occasional in alpine meadows up to 5000’—
5500’ at Brintnell L.; in flower the second week in July. Some of the Brintnell
L. plants appear to be intermediate between the typical form and the following

variety.

Pyrola secunda L. var. obtusata Turcz. Fl. Baical.-Dahur. 2': 213 (1856); Porsild in
Sargentia 4: 57 (1943), in Nat. Mus. Can. Bull. 101: 24 (1945). See also Rhod.
41: 274 (1939).
Sekwi R., Canol Rd., mi. 174 E., Porsild 11875 (Can.) ; Macmillan Pass, Canol Rd., mi.
225 E., Porsild 11253 (Can.).

218 SARGENTIA [6

Porsild states that this is the prevailing form of the species in Keewatin and
northern Mackenzie. It also appears occasionally in exposed situations in the
Athabaska—Great Slave L. region. Intermediates between species and variety are
frequent, often with the characters of both appearing on the same plant.

Pyrola virens Schweigg. in Schweigg. & Koerte, Fl. Erlang. 1: 154 (1804). See Fer-
nald in Rhod. 43: 167 (1941).

Pyrola chlorantha Sw.; Hooker, Fl. Bor.-Am. 2: 46 (1834) ; Macoun, Cat. Can. Pl. Pt.
2: 304 (1884); Raup in Jour. Arn. Arb, 17: 282 (1936) ; Porsild in Sargentia 4:
57 (1943).

Great Slave L., Richardson (G); northwest shore of Great Slave L., Bedford (Can.) ;
Wrigley, Miss E. Taylor 31 (Can.), 33 (G); Bear Rock, Norman, Porsild 3389 (Can.) ;
lowland woods near west erid of Brintnell L., 9435,

Common in low woods; in flower at Brintnell L. the second week in July.
Noted in Fl. Bor.-Am. as occurring northward to Great Bear L., where Porsild
has collected it in recent years. The latter has found it along the Mackenzie
northward at least to Sans Sault Rapid.

Pyrola grandiflora Radius, Diss. Pyrola 27 (1821); Raup in Jour. Arn. Arb. 17: 282
(1936); Porsild in Nat. Mus. Can. Bull. 101: 24, 32 (1945),
Pyrola rotundifolia sensu Hooker, Fl. Bor.-Am. 2: 46 (1834), in part; Macoun, Cat. Can.
Pl, Pt. 2: 305 (1884), in part; non L,

Slave R., Richardson (G); southwest and north shores of Great Slave L., Howe 91963
(Can.) ; Summit of Lone Mt., Wynne-Edwards 8495 (Can.) ; Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8353 (Can.); Sekwi R., Canol Rd., mi. 174 E., Porsild 11874
(Can.).

Pyrola grandiflora Radius var. canadensis (Andres) A. E. Porsild in Rhod. 41: 273
(1939).

Pyrola canadensis Andres in Oest. Bot. Zeit, 63: 250 (1913).

Pyrola borealis Rydb. in N. Am. Fl. 29: 24 (1914).

Pyrola rotundifolia sensu Hooker, Fl. Bor.-Am. 2: 46 (1834), in part; Macoun, Cat. Can.

Pl. Pt. 2: 305 (1884), in part; non L.

Fort Smith, Miss E. Taylor (Can.); Rae, Russell (Can.) ; Mackenzie R., Onion (N)
(type of P. borealis) ; damp lowland woods near Simpson, 9096; Bear Rock, Norman, Hume
103428 (Can.) ; mossy woods on shore of Brintnell L., 9/81.

Common ; collected in flower at Simpson on June 11, and at Brintnell L. on

June 20.

Pyrola asarifolia Michx. Fl. Bor.-Am. 1: 251 (1803); Raup in Jour. Arn. Arb. 17: 283
(1936).
Fort Smith, Raup 1111.
This typical form of the species is common in the southern part of the Mack-
enzie basin and is known in our region just at the southeastern corner.

Pyrola asarifolia Michx. var. incarnata (DC.) Fernald in Rhod. 6: 178 (1904); Raup
in Jour. Arn. Arb. 17: 283 (1936); Porsild in Nat. Mus. Can. Bull. 101: 24, 32
(1945).

Pyrola rotundifolia L. var. incarnata DC. Prodr. 7: 773 (1839) ; Hooker, Fl. Bor.-Am.

2: 46 (1834) ; Macoun, Cat. Can. Pl. Pt. 2: 305 (1884).

Pyrola incarnata Fisch. ex DC., as syn.

Liard R. between Nahanni Butte and Simpson, Crickmay 94 (Can.); lowland woods
and thickets near west end of Brintnell L., 9461; along Frost Cr., in upper valley north of
Brintnell L., alt. 4000’, 9274; turfy east slope of Red Mt., alt. 5500’, 9737; north peak of
Nahanni Mt., Wynne-Edwards 8494 (Can.); Nahanni Mts., Preble & Cary 64 (US);
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8354 (Can.); Macmillan Pass, Canol
Rd., mi. 225 E., Porsild 11254 (Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 219

Common in lowland woods at Brintnell L., but only occasional at higher levels ;
found flowering in late June and early July. Our no. 9737 has deep pink to red
petals, purplish or brownish anthers, and a relatively short, slightly depressed
style; furthermore it was found in flower much later in the season (Aug. 7).
In some ways it seems intermediate between this species and P. minor, and it is
left here tentatively. Its calyx and corolla closely resemble those of P. asarifolia
var. incarnata.

Monotropa uniflora L. Sp. Pl. 387 (1753); Raup in Jour. Arn. Arb. 17: 283 (1936).

The presence of Monotropa uniflora in southwestern Mackenzie is uncertain.
It was not found anywhere in the Mackenzie basin by the earlier collectors until
Miss Elizabeth Taylor collected it in 1892. Her labels (specimens in Herbs. G.
and Can.) read simply “Slave River,” with no indication as to where on the Slave
River she found her specimens. It was not collected again until 1935, when we
got it near Chipewyan and on the south shore of L. Athabaska.

ERICACEAE

Ledum groenlandicum Oeder, FI. Dan. pl. 567 (1771); Preble in N. Am. Fauna 27:
530 (1908); Raup in Jour. Arn. Arb. 17: 283 (1936); Porsild in Nat. Mus. Can.
Bull. 101: 24 (1945).

Ledum palustre L. var. latifolium (Ait.) Michx. sensu Hooker, Fl. Bor.-Am. 2: 44

(1834).

Ledum latifolium Ait.; Macoun, Cat. Can. Pl. Pt. 2: 301 (1884).

Smith Portage, Miss E. Taylor 59851 (Can.); 12 mi. east of Moraine Pt., Great Slave
L., Bedford (Can.); southwest and north shores of Great Slave L., Howe 91964 (Can.) ;
east shore of Trout L., C. O. Hage (G); Mackenzie R., Onion (N); mossy banks along
shore of Brintnell L., 9787; turfy slope near summit of Red Mt., alt. 5500’-5900', 9740;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11257 (Can.).

Common to abundant in timbered muskegs throughout the region. At Brint-
nell L. it is abundant in the open timber on the north slope of Colonel Mt. nearly
to timber line, and it is occasional in the high tundra. In flower in late June.

Ledum palustre L. var. decumbens Ait. Hort. Kew. 2: 65 (1789); Raup in Jour. Arn.
Arb. 17: 284 (1936).
Ledum palustre L., sensu Preble in N. Am. Fauna 27: 531 (1908).
Ledum decumbens (Ait.) Lodd. ex Steud.; Porsild in Nat. Mus. Can. Bull. 101: 24
(1945).

South shore of Great Slave L., Brooke (Can.); Mackenzie R., Onion (N); Liard R.
between Nahanni Butte and Simpson, Crickmay 90 (Can.); Mackenzie R. between Black-
water R. and Norman, Crickmay 170 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., alpine
slope, alt. 5000’, Wynne-Edwards 8357 (Can.); Trout Cr., Canol Rd. mi. 118 E., Porsild
11836 (Can.).

Preble notes this species as occurring “On the Barren Grounds and in the
northern part of the wooded country.” I have collected it as far south in the
Mackenzie basin as the northwest shore of L. Athabaska.

Rhododendron lapponicum (L.) Wahl. Fl. Suec. 249 (1824); Preble in N. Am. Fauna
27: 531 (1908); Raup in Jour. Arn. Arb. 17: 284 (1936); Porsild in Nat. Mus. Can.
Bull. 101: 25, 32 (1945).

Azalea lapponica L. Sp. Pl. 151 (1753).
Liard R. between Nahanni Butte and Simpson, Crickmay 87 (Can.); Lone Mt., Wynne-

Edwards 8497 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5000’, Wynne-

Edwards 8358 (Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11877 (Can.).

An arctic species which comes southward only a short distance into the forest.

220 SARGENTIA [6

Loiseleuria procumbens (L.) Desv. in Jour. Botanique 3: 35 (1813); Raup in Jour.
Arn. Arb. 17: 284 (1936); Porsild in Sargentia 4: 57 (1943), in Nat. Mus. Can.
Bull. 101: 25 (1945).

Azalea procumbens L. Sp. Pl. 151 (1753).

Liard R. between Nahanni Butte and Simpson, Crickmay 92 (Can.); Macmillan Pass,
Canol Rd., mi. 225 E., Porsild 11260 (Can.).

An arctic species which has been collected by Porsild also in Keewatin, at
Great Bear L., and in the Richardson Mts.

Phyllodoce empetriformis (Sm.) D. Don in Edinb. New Philos. Jour. 17: 160 (1834);
Porsild in Nat. Mus. Can. Bull. 101: 25 (1945).

Mensziesia empetriformis Sm. in Trans. Linn. Soc. 10: 380 (1811).

Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11259 (Can.).

A Pacific slope species which apparently extends eastward only to the Yukon—
Mackenzie divide.

Phyllodoce glanduliflora (Hook.) Cov. in Mazama 1: 196 (1897),

Menziesia glanduliflora Hooker, Fl. Bor.-Am. 2: 40 (1834).

Turfy east and northeast slopes of Red Mt., north of Brintnell L., alt. 4500’—5000’, 9277,
9749,

Abundant on the above site, and extending up to the summit of Red Mt.; but
only rare or occasional elsewhere on the mountains north of the lake, and not seen
at all south of the lake. Collected in flower late in June, and with maturing fruit
early in August.

Kalmia polifolia Wang. in Beob. Ges. Nat. Freunde Berlin 2: 130 (1788); Raup in
Jour. Arn, Arb. 17: 284 (1936); Porsild in Sargentia 4: 57 (1943), in Nat. Mus.
Can. Bull. 101: 25 (1945).

Kalmia glauca Ait.; Hooker, Fl. Bor.-Am. 2: 41 (1834); Macoun, Cat. Can. Pl. Pt. 2:
300 (1884) ; Preble in N. Am. Fauna 27: 531 (1908).

Resolution, Kennicott (N) ; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11258 (Can.).

The range of Kalmia polifolia in the Mackenzie basin is not clearly defined,
but it appears to be most abundant near and on both sides of the arctic timber line.
It is occasional on the shores of L. Athabaska, but has not been collected around
the eastern arm of Great Slave L. It has been found at Great Bear L. and in the
Keewatin District by Porsild. In Fl. Bor.-Am. it is noted as “. . . extending
beyond the woody regions, but scarcely, I believe, to the Arctic Circle”; and
Preble states that it was “. . . noted south of Smith Landing [Fitzgerald] . . .”
It apparently extends north to the Barren Grounds. Porsild says it is “A plant
of northern muskegs that in Keewatin extends some distance beyond the limit

of trees; rare in the lower Mackenzie basin, north to Bear Lake and in Central
Yukon.”

Cassiope tetragona (L.) D. Don in Edinb. New Philos. Jour. 17: 158 (1834); Macoun,
Cat. Can. Pl. Pt. 2: 297 (1884); Raup in Jour. Arn. Arb. 17: 285 (1936); Porsild
in Nat. Mus. Can. Bull. 101: 25, 32 (1945).

Andromeda tetragona L. Sp. Pl. 393 (1753) ; Hooker, Fl. Bor.-Am. 2: 38 (1834).

Mossy banks on south shore of Brintnell L., 9340; northeast slope of Red Mt., alt. 5000’—
5500’, 9295; north peak of Nahanni Mt., Wynne-Edwards 8498 (Can.); Lone Mt., Wynne-
Edwards 8499 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8355, 8356
(Can.).

Abundant on alpine slopes, where it is a primary part of the plant cover in
many places. On the north slope of Colonel Mt. it extends down through the
open timber to the shore of the lake; but it is rare on the southward-facing slopes

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 221

of Red Mt. In addition to the Canol Rd. localities mentioned above, Porsild
reports it as common on the Plains of Abraham and in Macmillan Pass.

Cassiope saximontana Small in N. Am. Fl. 20: 59 (1914).

Liard R. between Nahanni Butte and Simpson, Crickmay 86 (Can.).

This appears to be a northeastern range extension for an otherwise Rocky
Mountain and upper Yukon species.

Andromeda Polifolia L. Sp. Pl. 393 (1753); Hooker, FI. Bor.-Am. 2: 38 (1834);
Macoun, Cat. Can. Pl. Pt. 2: 297 (1884); Preble in N. Am. Fauna 27: 532 (1908);
Raup in Jour. Arn. Arb. 17: 285 (1936); Porsild in Nat. Mus. Can. Bull. 101:
25, 32 (1945).

Resolution, Kennicott (N); southwest and north shores of Great Slave L., Howe 91965
(Can.) ; northwest shore of Great Slave L., Bedford (Can.) ; Mackenzie R., Onion (N);
north of the height of land between Great Slave and Great Bear Lakes, Bedford (Can.) ;
Liard R. between Nahanni Butte and Simpson, Crickmay 89 (Can.) ; muskeg along Willow
Lake R., 60 mi. above mouth, Crickmay 21 (Can.) ; mossy banks on shores of Brintnell L.,
9185, 9252, 9327; north peak of Nahanni Mt., Wynne-Edwards 8500 (Can.); Bolstead Cr.,
Canol Rd., mi. 111 E., alpine slope, alt. 5000’, Wynne-Edwards 8359 (Can.); Sekwi R.,
Canol Rd., mi. 174 E., Porsild 11876 (Can.).

Occasional to common; collected in flower the last week of June. Noted in
Fl. Bor.-Am. northward “. . . to the Arctic sea-shore”; and by Preble “. . .
from Great Slave Lake northward. . . .” It is common in muskegs throughout
the pre-Cambrian parts of the Athabaska—Great Slave L. region.

Chamaedaphne calyculata (L.) Moench, Meth. 457 (1794); Preble in N. Am. Fauna 27:
532 (1908); Raup in Jour. Arn. Arb. 17: 285 (1936); Porsild in Sargentia 4: 58
(1943).

Andromeda calyculata L. Sp. Pl. 394 (1753) ; Hooker, Fl. Bor.-Am. 2: 39 (1834).

Cassandra calyculata (L.) D. Don.; Macoun, Cat. Can. Pl. Pt. 2: 296 (1884).

Resolution, Kennicott (N); northwest shore of Great Slave L., Bedford (Can.); north
of the height of land between Great Slave and Great Bear Lakes, Bedford (Can.); Liard

R. between Nahanni Butte and Simpson, Crickmay®88 (Can.); upland muskeg west of the

Mackenzie R. near Simpson, 9926; muskeg along Willow Lake R., 60 mi. above mouth,

Crickmay 22 (Can.).

Common at the Simpson station; found with mature fruit late in August.

Preble reports it at Norman, and Porsild has collected it at Great Bear L., in the

Eskimo Lakes basin, and in the Richardson Mts.

Arctostaphylos Uva-ursi (L.) Spreng. Syst. 2: 287 (1825); Macoun, Cat. Can. Bl. sets
2: 295 (1884); Preble in N. Am. Fauna 27: 532 (1908); Raup in Jour. Arn. Arb.
17: 285 (1936); Porsild in Nat. Mus. Can. Bull. 101: 25 (1945).

Arbutus Uva-ursi L. Sp. Pl. 395 (1753) ; Hooker, Fl. Bor.-Am. 2: 37 (1834).

Fort Smith, Seton & Preble 78352 (Can.); Resolution, Kennicott (N); southwest and
north shores of Great Slave L., Howe 91968 (Can.); northwest shore of Great Slave L.,
Bedford (Can.); east shore of Trout L., C. O. Hage (G); Mackenzie R., Onion (N);
stony fan on south shore of Brintnell L., 9579; Nahanni Mts., Preble & Cary 76 (US).

Common on dry stony fans at Brintnell L., and on warm south-facing slopes
up to timber line; fruit mature in late summer. Porsild noted the species as
common along a small tributary of Keele R., on the Canol Rd., mi. 51 FE. In Fi.
Bor.-Am. it is said to extend northward “. . . as far as Fort Franklin on the
Mackenzie River’; but Preble says he saw it “. . . throughout the territory
covered by our observations, and . . . also into the Barren Grounds.”

Most of the plants in the Brintnell L. area represent the typical form, but no.
9579 may be referred provisionally to var. adenotricha Fernald & McBr. (in

222 SARGENTIA [6

Rhod. 16: 211-213). It is somewhat more long-hirsute than this variety, how-
ever, and rather less glandular,

Arctostaphylos alpina (L.) Spreng. Syst. 2: 287 (1825); Macoun, Cat. Can. Pl. Pt. 2:
294 (1884), in part; Porsild in Sargentia 4: 58 (1943), in Nat. Mus. Can. Bull.
101: 25 (1945).

Arbutus alpina L. Sp. Pl. 395 (1753) ; Hooker, Fl. Bor.-Am. 2: 37 (1834), in part.

Mairainia alpina (L.) Desv.; Preble in N. Am. Fauna 27: 532 (1908), in part.

This typical, black-berried form of the species was noted as common in Mac-
millan Pass by Porsild. It was observed by Preble at Great Bear L.; and more
recently Porsild has reported it in Keewatin, at Great Bear L., in the Eskimo
Lakes Basin, and the Mackenzie R. delta. Farther south in the Mackenzie basin
it seems to be replaced entirely by the following red-fruited species.

Arctostaphylos rubra (Rehder & Wils.) Fernald in Rhod. 16: 32 (1914); Raup in
Jour. Arn. Arb. 17: 286 (1936); Porsild in Sargentia 4: 58 (1943), in Nat. Mus.
Can. Bull. 101: 25 (1945).

Arctous alpina (L.) Niedzu var. ruber Rehder & Wils. in Sargent, Pl. Wils. 3: 556
(1913).

Arbutus alpina L.; Hooker, Fl. Bor.-Am. 2: 37 (1834), in part.

Mairainia alpina (L.) Desv.; Preble in N. Am. Fauna 27: 532 (1908), in part.

Arctostaphylos alpina (L.) Spreng.; Macoun, Cat. Can. Pl. Pt. 2: 204 (1884), in part.

Southwest and north shores of Great Slave L., Howe 91967 (Can.); northwest shore of
Great Slave L., Bedford (Can.) ; woods near Simpson, Crickmay 41 (Can.); “River-
between-two-mountains,” just west of Franklin Mts., Crickmay 11 (Can.) ; gravelly thickets
and lowland woods near west end of Brintnell L., 9450; sphagnum hummocks in muskeg near
east end of Brintnell L., 9594.

Common to abundant in the Brintnell L. habitats mentioned, and extending up
through the open timber on the north-facing slope of Colonel Mt. into the alpine
meadows. The fruit turns red about mid-July in that area, Porsild reports the
species as only occasional in Macmillan Pass, along the Canol Rd. This is the
common form in the Athabaska—Great Slave L. region, and Porsild states that
it is also common on the Arctic coast and at Great Bear Ls

Vaccinium canadense Kalm ex Richardson, Bot. App. Frankl. Journ. 736 (1823);
Hooker, Fl. Bor.-Am. 2: 32 (1834); Macoun, Cat. Can. Pl. Pt. 2: 290 (1884);
Preble in N. Am. Fauna 27: 533 (1908); Raup in Jour. Arn. Arb. 17: 286 (1936).

No specimens of this species are yet available from southwestern Mackenzie,
though it undoubtedly extends at least into the country southwest of Great Slave

L. It is abundant in many parts of the Wood Buffalo Park. Hooker gives its

range as extending northward to Great Bear L., but there is no recent evidence

that it goes that far. Preble did not report it north of Fitzgerald, on the Slave R.

Vaccinium uliginosum L. Sp. Pl. 350 (1753); Hooker, Fl. Bor.-Am. 2: 32 (1834), in
part; Macoun, Cat. Can. Pl. Pt, 2: 291 (1884), in part; Preble in N. Am. Fauna
27: 533 (1908), in part; Raup in Jour. Arn. Arb. 17: 286 (1936); Porsild in Nat.
Mus. Can. Bull. 101: 32 (1945).

Northwest shore of Great Slave L., Bedford (Can.) ; east shore of Trout L., C. O. Hage
(G); Mackenzie R., Onion (N); mossy wooded bank on shore of Brintnell L., at Camp
Pt., shrubs 2’ high, 9190; Lone Mt., Wynne-Edwards 8502 (Can.); Nahanni Mts., Preble
& Cary 69 (US).

Common at low altitudes throughout much of the region. Noted by Preble as
. . . the commonest blueberry throughout the region north to the Arctic Sea.”
This is true for southwestern Mackenzie, but hardly so for the whole Athabaska—

“ce

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE £23

Mackenzie basin. Vaccinium canadense is probably the commonest form in the
Athabaska country.

No doubt a series of transition forms between this and var. alpinum could be
found, but they are sufficiently different in the Brintnell L. region, in habit,
site-selection, and fruit, to warrant their separation. The typical form is an
upright shrub growing at or near lake-level, and bearing an abundance of large
juicy: berries, while var. alpinum is a depressed shrub characteristic of high
alpine situations, and bearing small berries. The typical form flowers in the
third week of June and begins to mature its fruit about the first week in
August. The berries vary greatly in shape from depressed spherical to spherical
to pyriform or even cylindric.

Vaccinium uliginosum L. var. alpinum Bigel. Fl. Bost. ed. 2: 153 (1824); Porsild in
Nat. Mus. Can. Bull. 101: 25, 32 (1945).

Shale slide rock on west slope of Terrace Mt., alt. 6000’, depressed shrubs, 9511; north
peak of Nahanni Mt., Wynne-Edwards 8501 (Can.); Bolstead Cr., Canol Rd., mi. 111 E.,
alpine bog, Wynne-Edwards 8360 (Can.). |

A primary part of the vegetation on high mountain slopes and summits, where
it forms a large proportion of the plant cover, associated with Dryas and dwarf
Betula. Collected in flower at 6000’ on July 18. Porsild reports it as common
along the Canol Road.

Vaccinium Vitis-Idaea L. var. minus Lodd. Bot. Cab. p/. 1023 (1825); Raup in Jour.
Arn. Arb. 17: 287 (1936); Porsild in Nat. Mus. Can. Bull. 101: 25 (1945).

Vaccinium Vitis-Idaca L. sensu Hooker, Fl. Bor.-Am. 2: 34 (1834); Macoun, Cat. Can.
Pl. Pt. 23293 (1884).

Vitis-Idaca Vitis-Idaea (L.) Britt. sensu Preble in N. Am. Fauna 27: 532 (1908).

Fort Smith, Seton & Preble 78384 (Can.); same place, Miss E. Taylor 15459 (Can.) ;
same place, Raup 1194; northwest shore of Great Slave L., Bedford (Can.) ; Old Ft. Rae,
Russell (Can.) ; Witaley Harbor, Great Slave L., Kindle. (Can): east shore of Trout L.,
C. O. Hage (G); rich woods along shore of Brintnell L., 9303:

Common to abundant in woods and muskegs throughout the region. At Brint-
nell L. it reaches far above timber line, sometimes as much as 2000 feet. Porsild
noted it as a dominant species in the heath at Macmillan Pass. In flower late in
June. ;

Vaccinium microcarpum (Turcz. ex Rupr.) Hooker f. in Trans. Linn. Soc. 23: 334
(1861).

Oxycoccus microcarpus Turcz. ex Rupr. in Beitr. z. Pflanzenk. des Russ. Reich 4: *
(1845); Porsild in Can. Field Nat. 52: 117 (1938), in Sargentia 4: 58 (1943),
Nat. Mus. Can. Bull. 101: 25 (1945).

Vaccinium Oxycoccus sensu Hooker, Fl. Bor.-Am. 2: 34 (1834), in part; Raup in Jour.
Arn. Arb. 17: 287 (1936); non L.

Oxycoccus Oxycoccus sensu Preble in N. Am. Fauna 27: 533 (1908), non MacMill.

Southwest and north shores of Great Slave L., Howe 91966 (Can.); near Rae, Russell

(Can.); Liard R. between Nahanni Butte and Simpson, Crickmay 91 (Can.); sphagnum
hummocks in muskeg near east end of Brintnell L., 9593; mossy banks in lower part of
timber on north slope of Colonel Mt., 9415; Wacniiitan Pass, Canol Rd., mi. 225 E., Porsild
71256: (Can.).

Common in muskegs; in flower early in July and with ripening fruit in late

July. Porsild states that it is mostly limited to the forested country in the Mack-
enzie basin, extending only a short distance beyond the tree line.

224 SARGENTIA [6

PRIMULACEAE

Primula incana M. E. Jones in Proc. Cal, Acad. II. 5: 706 (1895); Fernald in Rhod.
30: 72 (1928); Raup in Jour. Arn, Arb, 17: 287 (1936); Porsild in Sargentia 4:
59 (1943).

Primula farinosa sensu Hooker, Fl. Bor.-Am. 2: 120 (1838), in part, non L.

Great Slave L., Richardson (G); Bear Rock, Norman, Porsild 3390 (Cans).

Occasional to common in the Wood Buffalo Park, and probably also through
the upper Mackenzie country. In Fl. Bor.-Am. P. farinosa, something of a
mixture as treated there, was noted as occurring northward “. . . to lat. 66° on
the Mackenzie River.”

Primula mistassinica Michx. Fl. Bor.-Am. 1: 124 (1803); Macoun, Cat. Can. Pl. Pt.
2: 309 (1884); Fernald in Rhod. 30: 88 (1928); Raup in Jour. Arn. Arb. 17: 288
(1936); Porsild in Nat. Mus. Can. Bull. 101: 33 (1945).

Primula Hornemanniana sensu Hooker, Fl. Bor.-Am, 2: 120 (1838), in part, non Lehm,

Southwest and north shores of Great Slave L., Howe 91961 (Can.); Pine Pt., Great
Slave L., Brooke (Can.); south bank of Mackenzie R., 10 mi. below: Mills L., Wynne-
Edwards 8556 (Can.); Mackenzie R., Richardson (G); north of the height of land be-
tween Great Slave and Great Bear Lakes, Bedford (Can.); Liard R. between Nahanni
Butte and Simpson, Crickmay 96 (Can.) ; Willow Lake R., 80 mi. above mouth, Crickmay
3 (Gan.).

Common in the Athabaska—Great Slave L. country, and I have seen a specimen
from Great Bear L. (J. M. Bell, Can.). All our specimens represent var. fypica
Fernald (1. ¢.).

Primula egaliksensis Wormsk. in Hornem. FI. Dan, 9: ¢. 1511 (1816); Fernald in Rhod.
30: 97 (1928); Raup in Jour. Arn. Arb. 17: 288 (1936),

Primula sibirica sensu Hooker, Fl. Bor.-Am. 2: 121 (1838), in large part; Macoun, Cat.
Can. Pl. Pt. 2: 310 (1884), in part; non Jacq.
This species has not yet been collected in the upper Mackenzie region, though
it is to be expected there. It is occasional on the eastern arm of Great Slave L.,
and has been collected at Great Bear L. by J. M. Bell (Can.). The range of
P. sibirica is given in Fl. Bor.-Am. as “Barren country between lat. 60° and 69°,
in clayey soil, by river banks and in marshy grounds.”

Primula stricta Hornem. in FI. Dan. 8: #. 1385 (1810); Fernald in Rhod. 30: 66 (1928);
Raup in Jour. Arn. Arb,.17: 288 (1936); Porsild in Nat. Mus. Can. Bull. 101:
35 (1945).
Primula sibirica sensu Hooker, Fl. Bor.-Am. 2: 120 (1838), in small part, non Jacq.
South shore of Great Slave L., R. Bell 23158 (Can.) ; Mackenzie R., Richardson (G) ;
Providence, Dutilly 65 (G); Smith Cr., about 13 mi. above Wrigley, Crickmay 2 (Can.) ;
Mackenzie R., 5 mi. above Old Fort Norman, Wynne-Edwards 8563 (Can.); Bear R., Miss
E, Taylor 87 (Can.).
Apparently common or occasional along the Mackenzie from Great Slave L.
northward. Also known on the arctic coast.

Androsace septentrionalis L. Sp. Pl. 142 (1753); Hooker, Fl. Bor.-Am. 2: 119 (1838) ;
Macoun, Cat. Can. Pl. Pt. 2: 311 (1884); Raup in Jour. Arn. Arb. 17: 288 (1936).

Fort Smith, Raup 1232, 3006; Resolution, Kennicott (N); Mackenzie R., Kennicott GR).
Simpson, Onion, Kennicott & Hardisty (N) ; south-facing slope in old field at Simpson,
9123.

Common in the Simpson habitat mentioned above; in flower about mid-June.
Its northward extent is given in Fl. Bor.-Am. as, “. . . to lat. 68° on the Mack-
enzie River,” and I have seen a specimen from Good Hope (Miss E. Taylor.

Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 225

Androsace Chamaejasme Host var. arctica Knuth in Pax & Knuth, in Pflanzenr, 22
(IV. 237): 190 (1905); Porsild in Nat. Mus. Can. Bull. 101: 25, 32 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 97 (Can.); Norman, Coll.?
(G): ? Cap Mt. near Wrigley, R. A. Brooke (Can.) ; north peak of Nahanni Mt., Wynne-
Edwards 8503 (Can.); Lone Mt., Wynne-Edwards 8504 (Can.); Bolstead Cr., Canol Rd.,
mi, 111 E., alpine slope, Wynne-Edwards 8361 (Can.); Sekwi R., Canol Rd., mi. 174 E.,
Porsild 11878 (Can.).

A western arctic species, noted in Fl. Bor.-Am. as far south as Norman, but
now known in the Mackenzie and Franklin Mts.

Dodecatheon pauciflorum (Durand) Greene in Pittonia 2: 72 (1890); Raup in Jour.
Arn. Arb. 17: 288 (1936).
Dodecatheon Meadia L. var. pauciflorum Durand, Pl. Pratten. 95 (1855).
Junction of Nyarling and Little Buffalo Rivers, Russell 1 (G); Resolution, Kennicott
(N).
Abundant in the prairies of the Wood Buffalo Park; in flower about mid-June.

Lysimachia thyrsiflora L. Sp. Pl. 147 (1753); Hooker, Fl. Bor.-Am. 2: 122 (1838);
Macoun, Cat. Can. Pl. Pt. 2: 314 (1884); Raup in Jour. Arn. Arb, 17: 288 (1936).

Sass Cr., Russell 26 (G); Old Fort Rae, Russell (G).

Noted in Fl. Bor.-Am. as extending from “. . . Lake Huron to the Mackenzie
River.”
Trientalis europaea L. var. arctica (Fisch. ex Hooker) Ledeb. Fl. Ross. 3: 25 (1851);

Raup in Jour. Arn, Arb. 17: 289 (1936).

Trientalis arctica Fisch. ex Hooker, Fl. Bor.-Am. 2: 121 (1838).

Junction of Nyarling and Little Buffalo Rivers Russell 57 (Can.).

This species is apparently common or abundant very locally in the Wood

Buffalo Park, and has been collected also at Lesser Slave L. (Brinkman 4023,
4279, N) and near Waterways, Alta. (Raup &Soper 9037).

Glaux maritima L. Sp. Pl. 207 (1753); Raup in Jour. Arn. Arb. 17: 289 (1936).
Junction of Nyarling and Little Buffalo Rivers, Russell 72 (G); Sulfur Bay, Great Slave
L., Hume 102677 (Can.); Mackenzie R., Kennicott (N).
Common at the margins of saline flats in the Wood Buffalo Park; in flower
about mid-June.

PLUMBAGINACEAE

Armeria maritima Willd. var. purpurea (Koch) Lawrence in Gent. Herb. 4: 405 (1940).

Armeria vulgaris Willd. ssp. arctica Hultén, sensu Porsild in Nat. Mus. Can. Bull. 101:

25 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 98 (Can.) ; Plains of Abraham,
Canol Rd., mi. 82 E., Porsild 11819 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., wet alpine
slope, alt. 5500’, Wynne-Edwards 8364 (Cant)

Porsild reports the species in flower late in July, and with mature fruit early in
September.

GENTIANACEAE
Gentiana Amarella L. Sp. Pl. 230 (1753); Raup in Jour. Arn. Arb. 17: 290 (1936);
Porsild in Sargentia 4: 59 (1943).
Gentiana acuta Michx. sensu Griseb. in Hooker, Fl. Bor.-Am. 2: 63 (1838).
Gentiana Amarella L. var. acuta (Michx.) Herder, sensu Macoun, Cat. Can. Pl. Pt. 2:
322 (1884).
Little Buffalo R., Russell 45 (G); southwest and north shores of Great Slave L., Howe
91959 (Can.) ; northwest shore of Great Slave L., Bedford (Can.); Rae, Russell (Can.) ;

226 SARGENTIA [6

Mackenzie R., Onion (N); along wood road near Simpson, 9883; Mackenzie R. between
Wrigley and Blackwater R., Crickmay 136 (Can.) ; between Blackwater R. and Norman,
Crickmay 171 (Can.) ; Norman, Coll.? (N).

Apparently common in the upper Mackenzie valley. I have seen specimens
from as far north as Good Hope (Onion, N), and Porsild has collected it at
Great Bear L. Collected with late flowers and maturing capsules at Simpson
Aug. 22.

Gentiana propinqua Richardson, Bot. App. Frankl. Journ. 734 (1823); Griseb. in
Hooker, Fl. Bor.-Am. 2: 62 (1838); Macoun, Cat. Can. Pl. Pt. 2: 322 (1884);
Porsild in Nat. Mus. Can. Bull, 101: 25, 32 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 99 (Can.) ; Mackenzie R.,
Onion (N) ; gravelly plains near west end of Brintnell L., 9460; mossy banks on north shore
of Brintnell L., 9347; alpine meadows and turfy shale slopes on north side of Colonel Mt.,
alt. 4000’-6000', 9642; turfy alpine slopes of Red Mt., alt. 5000’, 9761; Lone Mt., Wynne-
Edwards 8505 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5500’, Wynne-
Edwards 8363.

Common in the Brintnell L. area; found flowering in late June and in July.
Its range in Fl. Bor.-Am. is given as “Cumberland House to Bear Lake, and
Bear Lake River.”

Gentiana arctophila Griseb. in Hooker, FI. Bor.-Am. 2: 61 (1838); Porsild in Sargentia
4: 59 (1943), in Nat. Mus. Can. Bull. 101: 25 (1945).

Mossy banks on north shore of Brintnell L., 9161; alpine meadows and turfy shale slopes
on north side of Colonel Mt., alt. 5000’, 9417, 9608; Macmillan Pass, Canol Rd., mi, 225 E.,
Porsild 11262 (Can.).

Occasional in the Brintnell L. area, and apparently also on the Canol Rd. ;
in flower during late June and July. Our no. 9608 is an albino form with flowers
greenish white or barely tinged with pink.

Gentiana glauca Pall. Fl. Ross. 2: 104, t. 93, f. 2 (1788); Porsild in Nat. Mus. Can.
Bull. 101: 25 (1945),

Northeast slope of Red Mt., alt. 5000’-5900', 9291, 9735; along small brook on west slope
of Terrace Mt., alt. 5000’, 9543; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, Wynne-
Edwards 8362 (Can.).

Common in the Brintnell L. localities cited above, but only occasional else-
where in that area; found flowering in late June and mid-July, and fruiting early
in August. Porsild reports it as common in Macmillan Pass.

Gentiana Raupii A. E. Porsild in Sargentia 4: 60 (1943), in Nat. Mus. Can. Bull. 101:
35 (1945).

Gentiana detonsa sensu Griseb, in Hooker, FI. Bor.-Am. 2: 64 (1838), non Fries.

Gentiana serrata sensu Macoun, Cat. Can. Pl. Pt. 2: 321 (1884), non Gunner.

Gentiana elegans sensu Raup in Jour. Arn. Arb. 17: 290 (1936), non A. Nels.

Junction of Nyarling and Little Buffalo Rivers, Russell 66 (G); southwest and north
shores of Great Slave L., Howe 91958 (Can.) ; Sulfur Bay, Great Slave L., Hume 102678
(Can.) ; Mackenzie R., Jones 62299 (Can.) ; same, Onion (N); Simpson, near sandy beach
west of island, Nowosad (Can.); Mackenzie R. between Wrigley and Blackwater R.,
Crickmay 137 (Can.); Blackwater R., Porsild 6571 (Tyre) ; 5 mi. above Old Ft, Norman,
W ynne-Edwards 8564 (Can.); Norman, Coll.? (N); same place, Kindle (Can.) ; “Cum-
berland House to Bear Lake,” Richardson (G).

Lomatogonium rotatum (L.) Fries ex Nyman, Consp. Fl. Eur. 3: 500 (1881); Raup in
Jour. Arn. Arb. 17: 290 (1936); Porsild in Sargentia 4: 60 (1943). See Fernald
in Rhod, 21: 193 (1919),
Swertia rotata L. Sp. Pl. 226 (1753).
Pleurogyne rotata (L.) Griseb.; Macoun, Cat. Can. Pl. Pt. 2: 325 (1884),

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 227

Little Buffalo R., Russell 93 (G); Clewi R., Russell 96 (G); southwest and north shores
of Great Slave L., Howe 91960 (Can.) ; Mackenzie R., Onion (N).

Apparently widespread in the Mackenzie basin, though not abundant in any one
place. Porsild has collected it at Great Bear L. and on the arctic coast.

Menyanthes trifoliata L. Sp. Pl. 145 (1753); Macoun, Cat. Can. Pl. Pt. 2: 327 (1884);
Raup in Jour. Arn, Arb, 17: 290 (1936); Porsild in Sargentia 4: 61 (1943). See
Fernald in Rhod. 31: 195 (1929).

Northwest shore of Great Slave L., Bedford (Can.) ; Mackenzie R., Onion (N); same,
Kennicott (N).

Common in wet muskegs in the Athabaska—Great Slave L. region, and collected
by Porsild at Great Bear L. and in the lower Mackenzie country.

APOCYNACEAE
Apocynum androsaemifolium L. Sp. Pl. 213 (1753); Hooker, Fl. Bor.-Am. 2: 51 (1838);
Raup in Jour. Arn. Arb. 17: 290 (1936).
Fort Smith, Raup 1239 (Mrs. Conibear, coll.) ; northwest shore of Great Slave L., Bed-
ford (Can.).
Noted in Fl. Bor.-Am. as occurring “. . . throughout the woody country,”
but I have seen no specimens from north of Great Slave L.

Apocynum sibiricum Jacq. Hort. Vind. 3: 37, pl. 66 (1776); Raup in Jour. Arn. Arb.
17: 291 (1936). See Fernald in Rhod. 37: 327 (1935).

Rocky-sandy bluff along Mackenzie R. opposite Simpson, 9941.

Common in this habitat; collected Aug. 29, with fruit either fallen or not
developed, and the leaves turned yellow. Not previously reported in the Mack-
enzie basin north of the Wood Buffalo Park, where it is only occasional.

POLEMONIACEAE
Polemonium acutiflorum Willd. ex R. & S. Syst. 4: 792 (1819); Porsild in Sargentia
4: 61 (1943), in Nat. Mus. Can. Bull. 101: 25 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 100 (Can.); thickets on old
rock-slides in gorge, north slope of Colonel Mt., alt. 3300’, 9831; willow-birch scrub, north
slope of Colonel Mt., 9382; birch scrub on northeast slope of Red Mt., alt. 4000’-5000’, 9292 ;
Bolstead Cr., Canol Rd., mi. 111 E., wet alpine slope, alt. 4700’, Wynne-Edwards 8365 (Can.).

Common locally at Brintnell L., mostly confined to the willow-birch scrub
zones on the mountains ; found flowering in late June and early July; fruit mature
in mid-August. Porsild reports it common in Macmillan Pass.

Polemonium boreale Adams. Mem. Soc. Nat. Mosc. 5: 92 (1817); Porsild in Nat. Mus.
Can. Bull. 101: 25 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., on high alpine ridge, alt. 6000’-7000', Wynne-Edwards
8366 (Can.).

Flowering specimens were gathered here on July 25.

Collomia linearis Nutt. Gen. 1: 126 (1818); Hooker, Fl. Bor.-Am. 2: 76 (1838); Macoun,
Cat. Can. Pl. Pt. 2: 329 (1884); Raup in Jour. Arn. Arb. 17: 291 (1936).

Mackenzie R., ? Richardson (G); open field at Simpson, 9856.
Common at Simpson; collected with mature fruit and occasional flowers Aug.
21. Noted in Fl. Bor.-Am., “From the Saskatchewan to Fort Frankin. . . .”

HYDROPHYLLACEAE

Phacelia Franklinii (R. Br.) Gray, Man. ed. 2: 329 (1858); Macoun, Cat. Can. Pl. Pt.
2: 333 (1884); Raup in Jour. Arn. Arb. 17: 291 (1936).

228 SARGENTIA [6

Eutoca Franklinii R. Br, in Richardson, Bot. App. Frankl. Journ. (Repr. 51) t. 27 (1823) ;
Hooker, Fl. Bor.-Am. 2: 79 (1838).

Resolution, Onion, Kennicott & Hardisty (N); Windy Pt., Great Slave L., Hume 102679
(Can.) ; Old Fort Rae, Russell (Can.); “Cumberland House to Bear Lake, June—Aug.,”
Richardson (G).

Phacelia Franklinii is common in clearings and natural openings in the Atha-
baska-Great Slave L. district. It is especially abundant as a fireweed in re-
cently burned forest. In Fl. Bor.-Am. it is said to extend “From the Saskat-
chewan to the Bear Lake and English Rivers,” but I have seen no specimens
from beyond Great Slave L. except the Richardson collection with the somewhat
equivocal label.

BORAGINACEAE

Lappula Redowskii (Hornem.) Greene var. occidentalis (Wats.) Rydb. in Contr. U. S.
Nat. Herb. 3: 170 (1895); Raup in Jour. Arn. Arb. 17: 291 (1936): Porsild in
Sargentia 4: 62 (1943),

Echinospermum Redowskii var. occidentale Wats. Bot. King Exped. 246, ¢. 23, f. 9-10

(1871) ; Macoun, Cat. Can. Pl. Pt. 2: 336 (1884).
Echinospermwm patulum sensu Hooker, Fl. Bor.-Am, 2: 84 (1838), at least in part, non
Lehm.

Fort Smith, Seton & Preble 78385 (Can.) ; on south-facing slope in old field at Simpson,
9122.

Common around an old house site at Simpson, and at the edge of a cultivated
field; collected in flower about mid-June. Noted in Fl. Bor.-Am. from “Cum-
berland House to Bear Lake,” and collected in the Eskimo Lakes basin by
Porsild.

Myosotis alpestris Schmidt ssp. asiatica Vesterg. in Hultén, Fl. Kamtch. 4: 80 (1930);
Porsild in Nat. Mus. Can. Bull. 101: 25 (1945),
Myosotis sylvatica Hoffm. var. alpestris Koch, sensu Macoun, Cat. Can. Pl. Pt, 2: 340
(1884). ¢
Alpine meadows on south- and east-facing slopes of Red Mt., alt. about 5000’, 9202, 9294;
meadows on west slope of Terrace Mt., alt. 4800’-5500', 9544; Macmillan Pass, Canol Rd.,
mi. 225 E., Porsild 11263 (Can.).
Common to abundant above timber line; in flower in late June and at least
through mid-July. In Macoun’s Catalogue there is a record for this species at
the “Mackenzie River near Fort Simpson, Lat. 62° 30’ (McGill Coll. Herb.).”

Mertensia paniculata (Ait.) G. Don, Gen. Hist. 4: 318 (1838); Macoun, Cat. Can. PI.
Pt. 2: 339 (1884); Raup in Jour. Arn. Arb. 17: 292 (1936); Williams in Ann. Mo.
Bot. Gard. 24: 42 (1937); Porsild in Sargentia 4: 62 (1943), in Nat. Mus. Can.
Bull. 101: 25 (1945).

Pulmonaria paniculata Ait. Hort. Kew. 1: 181 (1789).

Lithospermum paniculatum Lehm.; Hooker, Fl. Bor.-Am. 2: 87 (1838).

Hay River, Great Slave L., Preble & Cary 24 (US); same place, Hume 102680 (Can.) ;
east shore of Trout L., C. O. Hage (G); Mackenzie R., Onion (N); Simpson, Onion (N) ;
gravelly thickets at Camp Pt., Brintnell L., 9261; dry gravelly watercourse, north slope of
Colonel Mt., alt. 5000’, 9647; alpine meadows on south slope of Red Mt., 9218.

Common to abundant, from low levels to high alpine meadows; in late June
its blue flowers make banks of color on the high slopes. Porsild noted it as
common in Macmillan Pass, and has collected it along the arctic coast of Yukon
Territory. In Fl. Bor.-Am. its range is given as “Saskatchewan to Hudson’s

Bay and Bear Lake.”

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 229

LABIATAE
Scutellaria epilobiifolia Hamilton, Monogr. Gen. Scut. 32 (1832); Raup in Jour. Arn.
Arb. 17: 292 (1936).

Scutellaria galericulata sensu Hooker, Fl. Bor.-Am. 2: 114 (1838) ; Macoun, Cat. ‘Can:
Pl. Pt. 2: 388 (1884) ; non L.

Sass Cr., Russell 16 (G); southwest and north shores of Great Slave L., "Howe 91955
(Can.); Mackenzie R., Onion (N), Ross (N), Kennicott (N); damp thicket along wood
road near Simpson, 9880.

Occasional ; found with maturing fruit and a few flowers Aug. 22. Noted in
Fl. Bor.-Am. “. . . as far north as Fort Franklin on the Mackenzie River.” .

Dracocephalum parviflorum Nutt. Gen. 2: 35 (1818); Hooker, Fl. Bor.-Am. 2: 115
(1838); Macoun, Cat. Can. Pl. Pt. 2: 387 (1884); Raup in Jour. Arn. Arb. 17:
292 (1936).

Resolution, Kennicott (N); “Between Bear Lake and Cumberland House,” Richardson
(G).
Common in dry openings and clearings in the Wood Buffalo Park, and abun-
dant in recently burned woods.
Agastache Foeniculum (Pursh) Ktze. Rev. Gen. Pl. 511 (1891); Raup in Jour. Arn.
Arb. 17: 292 (1936).
Stachys Foeniculum Pursh, Fl. Am. Sept. 2: 407 (1814).

Lophanthus anisatus (Nutt.) Benth.; Hooker, Fl. Bor.-Am. 2: 115 (1838) ; Macoun, Cat.
Can. Pl. Pt. 2: 386 (1884).

Resolution, Kennicott (N).

The northern range of this species is given by Hooker as “Plains of Saskatche-
wan ... to Fort Franklin on the Mackenzie River.” In addition to the above
specimen, however, I have seen it only at McMurray on the lower Athabaska R.

Stachys palustris L. var. pilosa (Nutt.) Fernald in Rhod. 45: 474 (1943).
Stachys pilosa Nutt. in Jour. Acad. Phil. 7: 48 (1834).
Stachys palustris L. sensu Hooker, Fl. Bor.-Am. 2: 116 (1838); Macoun, Cat. Can. PI.
Pt. 2: 390 (1884); Porsild in Sargentia 4: 62 (1943), in Nat. Mus. Can. Bull. 101:
33 (1945).

Stachys scopulorum Greene; Raup in Jour. Arn. Arb. 17: 293 (1936).

Junction of Nyarling and Little Buffalo Rivers, Russell 67 (G); Resolution, Onion,
Kennicott & Hardisty (N); Mackenzie R., Onion, Kennicott & Hardisty (N); south bank
of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8554 (Can.); grassy swale near
Ft. Liard, Nowosad (Can.); open field at Simpson, 9847; rocky-sandy bank of Mackenzie
R. opposite Simpson, 9954; upland prairie west of Mackenzie R. near Simpson, 9893;
Mackenzie R. between Wrigley and Blackwater R., Crickmay 140 (Can.).

Common to occasional in the upper Mackenzie valley; collected in fruit late
in August. Noted in Fl. Bor.-Am., “Throughout Canada to Fort Franklin on
the Mackenzie River.”

Mentha canadensis L. var. glabrata Benth. in DC. Prodr. 12: 173 (1848); Macoun, Cat,
Can. Pl. Pt. 2: 381 (1884); Raup in Jour. Arn, Arb. 17: 293 (1936).

Mentha canadensis L. var. glabrior Hooker, Fl. Bor.-Am, 2: 111 (1838).

Northwest shore of Great Slave L., Bedford (Can.); Mackenzie R., Onion (N); Simp-
son, Miss E. Taylor 119 (Can.); damp lowland thickets and sloughs near Simpson, 9843,
9879: rocky-sandy bank of Mackenzie R. opposite Simpson 9955; damp places in upland
prairie west of Mackenzie R. near Simpson, 9921; grassy swale near Ft. Liard, Nowosad
(Can oh:

Common in the Simpson area; collected in flower and with maturing fruit late
in August. This is the commonest mint of the Athabaska—Great Slave L. region,
but its northern limit in the Mackenzie valley is not yet known.

230 SARGENTIA [6

SCROPHULARIACEAE
Limosella aquatica L. Sp. Pl. 631 (1753); Raup in Jour. Arn, Arb. 17: 293 (1936);
Porsild in Sargentia 4: 62 (1943).
This species has not yet been found in southwestern Mackenzie, but it has
been collected at Lake Athabaska and at Great Bear L. (Porsild, 1. ¢.). Small
and inconspicuous, it is easily overlooked.

Veronica scutellata L. Sp. Pl. 12 (1753); Raup in Jour. Arn. Arb. 17: 294 (1936).
Near Fort Smith, Raup 1252; northwest shore of Great Slave L., Bedford (Can.).
Veronica alpina L. var. alterniflora Fernald in Rhod. 41: 455, t. 567 (1939): Porsild in
Nat. Mus. Can. Bull. 101: 25 (1945).
Noted by Porsild as common in Macmillan Pass, Canol Rd., mi. 225 E. Mr.
Porsild apparently did not collect specimens in the Pass itself, but specimens
from the Yukon side were determined as var. alterniflora by Fernald.

Veronica alpina L. var. unalaschcensis Cham. & Schl. in Linnaea 2: 556 (1827); Porsild
in Sargentia 4: 62 (1943). See Fernald in Rhod. 41: 450 (1939),

Willow-birch scrub and alpine meadows, upper valley of Frost Cr., and on west slope
of Terrace Mt., north of Brintnell L., alt. 4000'-5500’, 9554.

Common ; observed also on the east slopes of Red Mt., but not on Colonel Mt. ;
in flower about mid-July.

Some of these plants approach var. geminiflora Fernald (In Rhod. 41: 454).
They have the short styles and rather elongate fruiting racemes of this variety,
with the lowermost capsules sometimes opposite. Others have the close inflo-
rescence of var. unalaschcensis.

_Veronica peregrina L. var. xalapensis (H.B.K.) Pennell in Torreya 19: 167 (1919);
Raup in Jour. Arn. Arb. 17: 294 (1936); Porsild in Nat. Mus. Can. Bull. 101:
33 (1945).

Veronica xalapensis H.B.K. Nov. Gen. & Sp. 2: 389 (1818).

Veronica peregrina L. sensu Hooker, Fl. Bor.-Am. 2: 101 (1838) ; Macoun, Cat. Can.
Pl. Pt. 2: 362 (1884).

South bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8558 (Can.) : sandy
bank of Mackenzie R. opposite Simpson, 9939,

Occasional at Simpson; found with mature capsules late in August. Its
northern range is given in Fl. Bor.-Am., as “Throughout Canada to Mackenzie
River.”

Euphrasia subarctica Raup in Rhod. 36: 87 (1934), in Jour, Arn. Arb. 17: 294 (1936).
This species has been found at Lake Athabaska, on the eastern arm of Great
Slave L., and in Alaska. Its occurrence in the upper Mackenzie valley is prob-
lematical, but it should be looked for in that region. In Fl. Bor.-Am., E. offi-
cinalis L, is said to range northward to lat. 64°, and it may be that part of the
material upon which this range was based is E. subarctica. On the other hand
Porsild (Sargentia 4: 64) has collected specimens at Great Bear L. which he has
named £. disjuncta, so that Hooker may have had both species in his concept of
E. officinalis.
Castilleja pallida L. ssp. caudata Pennell in Proc. Acad. Nat. Sci. Phil. 86: 524 (1934);
Porsild in Sargentia 4: 63 (1943), in Nat. Mus. Can. Bull. 101: 32 (1945),
North Peak of Nahanni Mt., Wynne-Edwards 8506 (Catinn
Previously collected in the Mackenzie basin only on Peel River and in the
Mackenzie delta.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 231

Castilleja pallida L. ssp. elegans (Malte) Pennell in Proc. Acad. Nat. Sci. Phil. 86:
526 (1934); Porsild in Sargentia 4: 63 (1943), in Nat. Mus. Can. Bull. 101: 25
(1945).

Castilleja elegans Malte in Rhod. 36: 187 (1934).
Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes, Wynne-Edwards 8375 (Can.).

An arctic subspecies, whose principal range is north of the tree line.

Castilleja Raupii Pennell in Proc. Acad. Nat. Sci. Phil. 86: 528 (1934); Raup in Jour.
Arn. Arb, 17: 294 (1936); Porsild in Sargentia 4: 63 (1943), in Nat. Mus. Can.
Bull. 101: 35 (1945).

Fort Smith, Seton & Preble 78386 (Can.); same place, Raup 1258; lower Slave R.,
Raup 1262; southwest and north shores of Great Slave L., Howe 91957 (Can.) ; Windy Pt.,
Great Slave L., Hume 102682 (Can.); Pine Pt., Great Slave L., Brooke (Can.); near Rae,
Russell (Can.); Hay River, Preble & Cary 16 (US); Wrigley Harbor, Great Slave L.,
Kindle (Can.); east shore of Trout L., C. O. Hage (G); Mackenzie R., Onion (N);
Liard R. between Nahanni Butte and Simpson, Crickmay 101 (Can.); along wood roads
around Simpson, 9088, 9143; upland woods west of Mackenzie R. near Simpson, 9915;
Nahanni Mts., Preble & Cary 71 (US).

Common in the Athabaska-Great Slave L. region and the upper Mackenzie
valley; in flower about mid-June.

Castilleja Raupii Pennell ssp. ursina Pennell in Proc. Acad. Nat. Sci. Phil. 86: 530
(1934); Porsild in Sargentia 4: 64 (1943), in Nat. Mus. Can. Bull. 101: 33, 34,-35
(1945).

South bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8557 (Can.) ;
Wrigley, Miss E. Taylor 35 (Can.); Saline R., above Norman, Wynne-Edwards 8576
(Can.) ; Bear R., Miss E, Taylor 86 (Can.) (Type).

Porsild has collected this subspecies in the Mackenzie delta, and Wynne-Ed-
wards got it a few miles below Good Hope.

Rhinanthus groenlandicus Chab. in Bull. Herb. Boiss. 7: 515 (1899); Porsild in
Sargentia 4: 64 (1943), in Nat. Mus. Can. Bull. 101: 34, 35 (1945).
Rhinanthus Crista-galli sensu Hooker, Fl. Bor.-Am. 2: 106 (1838) ; Macoun, Cat. Can.
Pl. Pt. 2: 371 (1884); non L.

Rhinanthus Kyrollae Chab. sensu Raup in Jour. Arn. Arb. 17: 294 (1936).

Rocky-sandy bank of Mackenzie R. opposite Simpson, 9936; Wrigley, W ynne-Edwards
8606 (Can.); Mackenzie R. between Wrigley and Blackwater R., Crickmay 138 (Can.) ;
along Mackenzie R. near Root R., Porsild 6562 (Can.); 5 mi. above Old Fort Norman,
W ynne-Edwards 8565 (Can.).

Common locally at Simpson; collected in fruit late in August. The determi-
nation of our material is provisional, not only because of the incompleteness of
the specimens (dry stems and capsules, but no leaves), but also because of
uncertainty of the species limits within the genus. Rhinanthus Crista-galli is
noted by Hooker as extending “. . . to the shores of the Slave Lake, and to Fort
Franklin.” P

Pedicularis labradorica Wirsing, Ecolog. Bot. [2] ¢. 10 (1778); Raup in Jour. Arn. Arb.
17: 295 (1936), name wrongly attributed to Houttuyn; Porsild in Nat. Mus. Can.
Bull. 101: 26, 33 (1945). See Fernald in Rhod. 33: 193 (1931); Merrill in Rhod.
40: 292 (1938).

Pedicularis euphrasioides Stephan; Hooker, Fl. Bor.-Am. 2: 109 (1838); Macoun, Cat.

Can. Pl. Pt. 2: 368 (1884).

Windy Pt., Great Slave L., Hume 102681 (Can.); northwest shore of Great Slave Le
Bedford (Can.); Rae, Russell (Can.); Mackenzie R., Onion (Can.); Liard R. between
Nahanni Butte and Simpson, Crickmay 103 (Can.); outwash gravel plain near mouth of
North Nahanni R., Wynne-Edwards 8539 (Can.); Norman, Dutilly 44 (G); upper slopes
of Red Mt., north of Brintnell L., alt. 4000’-5000’, 9204, 9290; Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8373 (Can.).

232 Tut SARGENTIA: [6

Occasional to common in the Brintnell L. district; found below timber line on
the north slope of Colonel Mt.; in flower late in June. Noted in Fl. Bor.-Am.,
“North of the Saskatchewan to the Arctic Sea-shore.” I have seen specimens
from Great Bear L. (J. M. Bell, Can.) and the lower Mackenzie (Miss E. Taylor,
Can.).

Pedicularis sudetica Willd. Sp. Pl. 3: 209 (1800); Porsild in Nat. Mus. Can. Bull. 101:
26 (1945).

Swampy muskeg thickets around the shores of Brintnell L., 9192, 9235, 9326; Plains of
Abraham, Canol Rd., mi. 82 E., Porsild 11820 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E.,
wet alpine slopes, alt. 5000’, Wynne-Edwards 8370, 8371, 8372 (Canny

Occasional in the Brintnell L. habitat noted, though widespread around the
lake ; found flowering in late June.

Pedicularis lanata Cham. & Schl. in Linnaea 2: 583 (1827); Porsild in Nat. Mus. Can.
Bull, 101: 26, 32 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 102 (Can.); alpine meadows
and turfy ledges, north slope of Colonel Mt., 9401; summit and higher alpine slopes of Red
Mt., 9201, 9762; Nahanni Mts., Preble 315 (US); Lone Mt., Wynne-Edwards 8507 (Can.) ;
north peak of Nahanni Mt., Wynne-Edwards 8508 (Can.) ; Plains of Abraham, Canol Rd.,
mi. 82 E., Porsild 11821 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, alt. 5000’,
Wynne-Edwards 8369 (Can.).

Common in the Brintnell L. area; collected in flower June 21, and with matur-

ing capsules early in August.

Pedicularis arctica R. Br. in Suppl. App. Parry’s 1** Voy. 280 (1824), Chlor. Melv. 32
(1824); Porsild in Nat. Mus. Can. Bull. 101: 26 (1945). See Porsild in Sar-
gentia 4: 64 (1943).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11822 (Can.) ; Bolstead Cr., Canol Rd.,

mi. 111 E., alpine slope, alt. 5000’-6000’, Wynne-Edwards 8367, 8368 (Can.).

Pedicularis parviflora Smith in Rees, Cycl. 26 (1813); Raup in Jour. Arn. Arb, 17:
295 (1936).

This species is included here provisionally. It is a western arctic and north
Pacific coast species which I collected at Lake Athabaska in 1935 (1. ¢.), thus
greatly extending its known range. In Macoun’s Catalogue of Canadian Plants
(Pt. 2: 369, 1884), under P. palustris L. var. Wlassoviana Bunge, is the note
“On the Mackenzie River. (McGill Coll. Herb.).” Other than my own col-
lection this is the only indication we have of P. parviflora in the Mackenzie basin.
It suggests, however, that the plant may occur somewhere in the Mackenzie valley.

Pedicularis capitata Adams in Mém. Soc. Nat. Mosc. 5: 100 (1817); Porsild in Nat.
Mus. Can. Bull. 101: 26 (1945).

Mackenzie R., Jones 62305 (Can.); mossy bank on south shore of Brintnell L.,. 9357 ati
willow-birch scrub above timber line on north slope of Colonel Mt., 9378; Bolstead Cr.,
Canol Rd., mi. 111 E., alpine slope, Wynne-Edwards 8374 ean).

At Brintnell L. this species was seen only on the mountain sides south of the
lake, where it was common ; found flowering early in July.

OROBANCHACEAE

Boschniakia rossica (Cham. & Schl.) Fedtch. in Fedtch. & Flerov. Fl. Eur. Ross. 896
(1910); Raup in Jour. Arn. Arb. 17: 295 (1936); Porsild in Sargentia 4: 65 (1943),
in Nat. Mus. Can. Bull. 101: 33 (1945).

Orobanche rossica Cham. & Schl. in Linnaea 4: 132 (1828).
Orobanche glabra Hooker, Fl. Bor.-Am. 2: 91 (1838). :
Boschniakia glabra (Hooker) C. A. Mey.; Macoun, Cat. Can. Pl. Pt. 2: 374 (1884).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 233

Great Slave L., R. Bell 23149 (Can.) ; Simpson, Wynne-Edwards 8610 (Can.) ; willow-
poplar-alder thickets near Simpson (under Alnus crispa), 9103; Norman, Dutilly 1 (G);
mossy banks on south shore of Brintnell L. (under Alnus crispa), 9358; in alder thickets
on lower north slope of Colonel Mt., 9393.

Common to abundant; in flower early in July. Noted in Fl. Bor.-Am. from
“Great Slave Lake to Fort Franklin on the Mackenzie River.” It appears to be
common in woods and alder thickets, however, throughout the lower Mackenzie
country. Porsild has collected it in the Mackenzie delta, in the Eskimo Lakes

basin, and at Great Bear L.

LENTIBULARIACEAE

Pinguicula vulgaris L. Sp. Pl. 17 (1753); Hooker, Fl. Bor.-Am. 2: 118 (1838); Macoun,
Cat. Can. Pl. Pt. 2: 376 (1884); Raup in Jour. Arn. Arb. 17: 295 (1936); Porsild
in Nat. Mus. Can. Bull. 101: 26, 32 (1945).

Southwest and north shores of Great Slave L., Howe 91956 (Can.) ; northwest shore of
Great Slave L., Bedford (Can.) ; Rae, Russell (Can.) ; Mackenzie R. between Wrigley and
Blackwater R., Crickmay 139 (Can.); Lone Mt., Wynne-Edwards 8509 (Can.) ; Bolstead
Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8376 (Can.).

Noted in Fl. Bor.-Am. as occurring in “woody and barren territories to
Mackenzie’s River.” The species is common around Great Slave L., and I have
seen specimens from Great Bear L. (J. M. Bell, Can.) and Good Hope (M iss

E. Taylor, G. Can.).

Pinguicula villosa L. Sp. Pl. 17 (1753); Raup in Jour. Arn. Arb. 17: 296 (1936); Porsild
in Sargentia 4: 65 (1943).

On hummocks of Sphagnum in muskeg near eastern end of Brintnell L., 9595.

Though widespread in northern Mackenzie, this species appears to be only
occasional in occurence. At Brintnell L. it was common in the single spot noted,
but was not seen elsewhere in that area. Porsild states that it is “Rare or oc-
casional in sphagnum bogs of the wooded country north to the limit of trees or
slightly beyond.”

Utricularia vulgaris L. Sp. Pl. 18 (1753); Raup in Jour. Arn. Arb. 17: 296 (1936).
Utricularia vulgaris L. var. americana Gray, sensu Porsild in Sargentia 4: 66 (1943).
Grande Detour Portage, lower Slave R., Russell 74 (G); northwest shore of Great Slave

L., Bedford (Can.) ; Rae, Russell (Can.) ; near large lake between southern extremities of

Franklin Mts., Crickmay 34 (Can.).

Common to abundant in the Athabaska-Great Slave L. district, and probably
far northward. Porsild has collected it in the Mackenzie delta, the Eskimo Lakes
basin, and at Great Bear L.

Porsild, perhaps correctly, has assigned all the Mackenzie basin specimens to
var. americana Gray (Man. ed. 5: 318, 1867). Northwestern American plants,
however, approach the typical European, and since I have seen but little flower-
ing material from the Mackenzie country I think it best at present to regard
our plants as representing the species in its broad sense.

Utricularia minor L. Sp. Pl. 18 (1753); Raup in Jour. Arn. Arb. 17: 296 (1936); Porsild
in Sargentia 4: 65 (1943).
This species has not been collected in the upper Mackenzie valley, though
it is probably common there. I have collected it at Athabaska and Great Slave
Lakes, and Porsild has it from several places around Great Bear L.

Utricularia intermedia Hayne in Schrad. Jour. Bot. 1800: 18 (1801); Hooker, FI. Bor.-
Am. 2: 118 (1838); Macoun, Cat. Can. Pl. Pt. 2: 375 (1884); Raup in Jour. Arn.
Arb. 17: 296 (1936); Porsild in Sargentia 4: 65 (1943).

234 SARGENTIA [6

Not yet collected in the upper Mackenzie country, though it unquestionably
is common there. Like so many aquatics it has been picked up by only a few
collectors. It was noted in Fl. Bor.-Am. from “Lake Huron to Bear Lake,”
where Porsild has found it common. Porsild also has it, however, from the
Mackenzie delta and from the Eskimo Lakes basin. It is occasional to common
around Athabaska and Great Slave Lakes.

PLANTAGINACEAE

Plantago major L. Sp. Pl. 112 (1753); Hooker, Fl. Bor.-Am. 2: 123 (1838).

Plantago major L. var. asiatica (L.) Decaisne, sensu Macoun, Cat. Can. Pl. Pt. 2: 392
(1884) ; Raup in Jour. Arn. Arb. 17: 296 (1936).

Fort Smith, Raup 1294; lowland woods near Simpson, 9089; “To the Arctic Sea,”
Richardson (G).

Common at Simpson; in fruit late in August. Noted in Fl. Bor.-Am., “From
Lake Huron . . . to lat. 68°.” The relationship of the Mackenzie basin plants
to the typical form is uncertain. In detailed structural characteristics they
resemble the common weed of waste places very closely. In our region, however,
their habit of growth and selection of habitats strongly suggest a native species.
Their leaves usually are rather stiff and upright, and they appear in woods and
thickets well away from much frequented paths.

Plantago eriopoda Torr. in Ann. Lyc. N. Y. 2: 237 (1827); Raup in Jour. Arn, Arb.
17: 296 (1936); Porsild in Sargentia 4: 66 (1943).

Plantago eriopoda is common on saline flats in the Wood Buffalo Park, and it
has been collected by Porsild on the arctic coast east of the Mackenzie delta.
It has not been collected in southwestern Mackenzie, but undoubtedly occurs
there, in areas watered by salt springs from the Silurian limestones.

Plantago septata Morris in Bull. N. Y. Bot. Gard. 2: 182 (1901); Porsifd in Sargentia
4: 66 (1943), in Nat. Mus. Can. Bull. 101: 33 (1945).

Plantago lanceolata L. var. 8 Hooker, Fl. Bor.-Am. 2: 123 (1838).

Mackenzie R., Onion (G) ; Liard R. between Nahanni Butte and Simpson, Crickmay 104
(Can.) ; Simpson, Wynne-Edwards 8609 (Can.) ; Norman, Dutilly 46 (G).

Noted in Fl. Bor.-Am. as occurring from “Fort Norman to Arctic Sea-shore,”
but we now have it considerably farther south, at Simpson and on the lower
Liard R.

RUBIACEAE

Galium boreale L. Sp. Pl. 108 (1753); Hooker, Fl. Bor.-Am. 1: 289 (1834); Macoun,
Cat. Can. Pl. Pt. 2: 203 (1884); Raup in Jour. Arn. Arb. 17: 297 (1936); Porsild
in Sargentia 4: 66 (1943), in Nat. Mus. Can. Bull. 101: 32, 34 (1945).

Fort Smith, Raup 1304; junction of Nyarling and Little Buffalo Rivers, Russell 55 (isis
Sulfur Bay, Great Slave L., Hume 102685 (Can.) ; northwest shore of Great Slave L.,
Bedford (Can.); Old Fort Rae, Russell (Can.); Mackenzie R., Onion, Kennicott &
Hardisty (N); Providence, Preble & Cary 30 (US); Liard R. between Nahanni Butte
and Simpson, Crickmay 105 (Can.) ; open field at Simpson, 9848; Saline R., above Norman,
Wynne-Edwards 8577 (Can.) ; Nahanni Mts., Preble & Cary 54 (Can.); north peak of
Nahanni Mt., Wynne-Edwards 8510 (Can.).

Common ; collected with flowers and maturing fruit Aug. 21. Hooker gives
its range “. . . to lat. 68° North ... ,” and Porsild has collected it both in the
Mackenzie delta and at Great Bear L. It is abundant in the central Mackenzie
basin.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 235

Galium trifidum L. Sp. Pl. 105 (1753); Macoun, Cat. Can. Pl. Pt. 2: 201 (1884); Raup
in Jour. Arn. Arb. 17: 297 (1936); Porsild in Sargentia 4: 67 (1943), in Nat. Mus.
Can. Bull. 101: 26 (1945).
Galium Claytoni sensu Hooker, Fl. Bor.-Am. 1: 288 (1834), in part, non Michx.

Fort Smith, Raup 1304; Little Buffalo R., Russell 86 (G) ; damp place in upland prairie
west of Mackenzie R. near Simpson, 9913; Macmillan Pass, Canol Rd., mi. 225 E., Porsild
11264 (Can.).

Common locally, north to Great Bear L. and the lower Mackenzie (Hooker,
1. c.; Porsild, 1. c.). Collected with mature fruit at Simpson late in August.

CAPRIFOLIACEAE
Viburnum edule (Michx.) Raf. Med. Repos. Lex. II. 5: 354 (1808); Hooker, Fl. Bor-.
Am. 1: 281 (1834), in part; Porsild in Sargentia 4: 67 (1943). See Fernald in
Rhod. 43: 481 (1941).
Viburnum Opulus L. var. edule Michx. Fl. Bor.-Am, 1: 18 (1803).

Viburnum pauciflorum Raf.; Macoun, Cat. Can. Pl. Pt. 2: 195 (1884) ; Preble in N. Am.
Fauna 27: 534 (1908); Raup in Jour. Arn. Arb. 17: 297 (1936).

Viburnum Oxycoccus sensu Hooker, Fl. Bor.-Am. 1: 281 (1834), in part, non Pursh.

Fort Smith, Seton & Preble 78333 (Can.) ; same place, Raup 1324; Resolution, Kennicott
(N); same place, Preble 204 (US); same place, Preble & Cary 152, 153 (US); Hay
River, Great Slave L., Preble & Gary 19 (US) ; northwest shore of Great Slave L., Bedford
(Can.) ; east shore of Trout L., C. O. Hage (G); Mackenzie R., Onion (N); Simpson,
Onion, Kennicott & Hardisty (N); damp lowland woods at Simpson, shrubs up to 6’ high,
9082; Wrigley, Preble & Cary 87 (US); rich spruce woods along shore of Brintnell L.,
shrubs up to 6’ high, 9498; Nahanni Mts., alt. 700’ and 1500’, Preble & Cary 59, 81 (US).

Common to abundant in woods throughout the Athabaska-Great Slave L.
region and the upper Mackenzie country. Porsild states that it is only occasional
along the lower Mackenzie and at Great Bear L. At Brintnell L. we found it
in stunted form as high as 4100’ on the south slope of Terrace Mt. In flower at
Simpson on June 11, and collected with unripe berries at Brintnell L. in mid-July.

Symphoricarpos, occidentalis Hooker, Fl. Bor.-Am. 1: 285 (1834); Macoun, Cat. Can.
Pl. Pt. 2: 195 (1884); Raup in Jour. Arn. Arb, 17: 298 (1936).

Resolution, Kennicott (N).

This species is noted in Fl. Bor.-Am. as occurring in the “Woody country
between lat. 54° and 64°,” but I have seen no material from north of Resolution.
It is rather abundant in the drier prairies of the Wood Buffalo Park, and prob-
ably extends to similar situations along the upper Mackenzie.

Symphoricarpos albus (L.) Blake var. pauciflorus (Robbins) Blake in Rhod. 16: 119
(1914); Raup in Jour. Arn. Arb. 17: 298 (1936).

Vaccinium album L. Sp. Pl. 1: 350 (1753), in part.

Symphoricarpos racemosus Michx. var. pauciflorus Robbins in Gray, Man. ed. 5203
(1867), in part; emend. Fernald in Rhod. 7: 167 (1905).

Fort Smith, Raup 1320; Little Buffalo R., Russell 102 (G).

Common to abundant in the open woods and prairies of the Wood Buffalo
Park, and probably also in similar situations along the upper Mackenzie. It
has not been collected, however, from beyond the lower Little Buffalo Rk.

Linnaea borealis L. var. americana (Forbes) Rehder in Rhod. 6: 56 (1904); Raup in
Jour. Arn. Arb. 17: 298 (1936).
Linnaea americana Forbes, Hort. Woburn. 135 (1825).
Linnaea borealis L. sensu Hooker, Fl. Bor.-Am. 1: 285 (1834) ; Macoun, Cat. Can. PI.
Pt. 2: 195 (1884).

236 SARGENTIA. : [6

Fort Smith, Miss E. Taylor 9572 (Can.); same place, Raup 1311; Little Buffalo R.,
Russell 69 (G); Windy Pt., Great Slave L., Hume 102684 (Can.) ; southwest and north
shores of Great Slave L., Howe 91954 (Can.) ; northwest shore of Great Slave L., Bedford
(Can.) ; Old Fort Rae, Russell (Can.) ; east shore of Trout L., C. O. Hage (G); gravelly
woods near shore of Brintnell L., 9366; shrubby thicket above timber line on north slope of
Colonel Mt., alt. 4300’, 9634; Nahanni Mts., Preble & Cary 67 (US).

Abundant throughout the forests of the central and southern parts of the
Mackenzie basin, and probably far northward. It is common to abundant in the
region of Simpson and at Brintnell L., where it is occasional even above timber
line. We found it at an altitude of between 5000’ and 5500’ on the eastern slope
of Red Mt. Noted in Fl. Bor.-Am. as occurring northward to the arctic circle ;
but I have seen specimens from the lower Peel R. (Miss E. Taylor, G), while
Porsild has collected it in the Mackenzie delta and on the neighboring parts of
the arctic coast.

Lonicera glaucescens Rydb. in Bull. Torr. Bot. Cl. 24: 90 (1897); Preble in N. Am.
Fauna 27: 534 (1908); Raup in Jour. Arn. Arb, 17: 298 (1936).

Fort Smith, Miss E. Taylor 9658 (Can.); same place, Seton & Preble 78334 (Can.);
same place, Raup 1318; Mackenzie R., Onion (N); open lowland woods along road near
Simpson, shrub 3’ high, 9117.

Rare at Simpson; collected with flower buds gn June 12. This species is
common in the open woods of the Wood Buffalo Park, but it is not definitely
known north of Simpson.

Hooker, in Fl. Bor.-Am., notes two species of Lonicera in the Mackenzie basin,
neither of which, it seems, could have been confused with L. glaucescens, the
only one now known to be common there. Lonicera caerulea he ranges
“Throughout the woody country, as far North at lat. 66°.” Neither this species
nor any close relative of it is now known in the Mackenzie basin. The other
species noted by Hooker is L. involucrata, for which he gives the somewhat
equivocal range, “Woody country between lat. 54° and 64°, (but probably con-
fined to the vicinity of the Saskatchewan).” This species actually occurs in the
Mackenzie basin, but so far as known at present only in the southern and south-
western districts (Lac la Biche, Lesser Slave L., upper Peace River). It was
found by Preble (1. c.) along the lower Athabaska as far north as Pelican Rapid.
The significance of these two records for honeysuckles in the upper Mackenzie
valley, therefore, remains uncertain.

Hooker’s record for Adoxa Moschatellina L. presents a similar enigma. It
also is said to range in the woody country between lat. 54° and 64°. It has been
collected along the lower Athabaska R. at Cascade Rapids above McMurray
(Miss E. Taylor, G, N, Can.), and in the Lesser Slave Lake district (Brinkman),
but is unknown farther northward except for the above record.

VALERIANACEAE

Valeriana septentrionalis Rydb. in Mem. N. Y. Bot. Gard. 1: 375 (1900); Raup in Jour,
Arn. Arb. 17: 299 (1936); Porsild in Sargentia 4: 67 (1943).

Mossy thickets near mouth of small stream at west end of Brintnell L., 9333.
Occasional in the above habitat; in flower late in June. Occasional also in
the Wood Buffalo Park, and collected by Porsild at Great Bear L.

Valeriana sitchensis Bong. Veg. Sitch. 145, repr. 27 (1832); Porsild in Nat. Mus. Can.
Bull. 101: 26 (1945).

Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11265 (Gans).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 237

CAMPANULACEAE

Campanula rotundifolia L. Sp. Pl. 163 (1753); Hooker, FI. Bor.-Am. 2: 27 (1834), in
part; Macoun, Cat. Can. Pl. Pt. 2: 288 (1884); Raup in Jour. Arn. Arb. 17: 299
(1936); Porsild in Nat. Mus. Can. Bull. 101: 32 (1945).

Campanula linifolia sensu Hooker, Fl. Bor.-Am. 2: 27 (1834), in part, non a Dis.

Little Buffalo R., Russell 89 (G); Resolution, E. A. & A. E. Preble 163 (US) ; south-
west and north shores of Great Slave L., Howe 91953 (Can.) ; Windy Pt., Great Slave D5
Hume 102683 (Can.); northwest shore of Great Slave L., Bedford (Can.); Rae, Russell
(Can.) ; Liard R. between Nahanni Butte and Simpson, Crickmay 108, 109 (Can.) ; Nahanni
Mts., Preble & Cary 55 (US); north peak of Nahanni Mt., W ynne-Edwards 8513 (Can.).

Common in the Wood Buffalo Park, but apparently becoming rare north of
Great Slave L. Noted in Fl. Bor.-Am., “Woody country between lat. 54° and

64°.”

Campanula lasiocarpa Cham. in Linnaea 4: 39 (1829).

Mackenzie R. between Blackwater R. and Norman, Crickmay 172 (Can.) ; gravelly
thickets at Camp Pt. and about the western end of Brintnell L., 9456, 9491; turfy shale
slopes on north slope of Colonel Mt., alt. 4000’-5000’, 9643; rocky stream banks and shale
rock-slides in upper valley of Frost Cr., and on west slope of Terrace Mt., alt. 4300’-5500’,
9547.

Occasional to common on alpine slopes at Brintnell L., but rare at lake-level ;

collected in flower during middle and late July.

Campanula aurita Greene in Pittonia 1: 221 (1888); Porsild in Nat. Mus. Can. Bull.
101: 26, 32 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 106, 107 (Can.) ; west ridge of
Franklin Mts. at “River-between-two-mountains,” alt. 1000’, Crickmay 6 (Can.) ; southern
extreme of Franklin Mts., alt. 800’, Crickmay 30 (Can.); Lone Mt., Wynne-Edwards 8511,
8512 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., alpine slope, Wynne-Edwards 8377
(Can.) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11879 (Can.).

LOBELIACEAE

Lobelia Kalmii L. Sp. Pl. 930 (1753); Hooker, Fl. Bor.-Am. 2: 30 (1834); Macoun,
Cat. Can. Pl. Pt. 2: 286 (1884); Raup in Jour. Arn. Arb. 17: 299 (1936).

Sandy muskeg, north arm of Great Slave L., Hume 102686 (Can.) ; near large lake be-
tween southern extremities of Franklin Mts., Crickmay 33 (Can.).

Noted in Fl. Bor.-Am. “. . . in saline marshes in lat. 60°.” Apparently rare
in the Mackenzie basin, and known east of the mountains only from the above
records. We have recently collected it in the valley of the Liard R. near the
lower crossing of the Alaska Highway.

COMPOSITAE

Grindelia perennis A. Nels. in Bull. Torr. Bot. Cl. 26: 355 (1899); Raup in Jour. Arn,
Arb. 17: 299 (1936). See Steyermark in Ann. Mo. Bot. Gard. 21: 485 (1934).

Grindelia squarrosa sensu Macoun, Cat. Can. Pl. Pt. 2: 208 (1884), in part, non (Pursh)
Dunal.

Donia squarrosa sensu Hooker, Fl. Bor.-Am. 2: 25 (1834), in part, non Pursh.

Little Buffalo R., Russell (Can.).

Hooker gives the northern range of this species as extending between lat. 54°
and 64°. It is rather common on the Salt Plains of the Wood Buffalo Park, and
Russell’s specimen came from such a situation just north of the 60th parallel.
It is probable that it extends over at least a part of the region southwest of Great
Slave L., if not into the upper Mackenzie valley.

238 SARGENTIA [6

Solidago multiradiata Ait. Hort. Kew. 3: 218 (1789); Hooker, Fl. Bor.-Am. 2: 5
(1834), in part; Macoun, Cat. Can. Pl. Pt. 2: 212 (1884); Raup in Jour. Arn. Arb.
17: 300 (1936); Porsild in Nat. Mus. Can. Bull. 101: 26 (1945).

Northwest shore of Great Slave L., Bedford (Can.) ; lowland woods and gravelly thickets
at west end of Brintnell L., 9430; gravelly beach on south shore of Brintnell L., 9356; alpine
meadows and rock-slides, north slope of Colonel Mt., 9380; rocky slopes of Red Mt., in
scrub timber and at 5500’, 9220, 9728; Bolstead Cr., Canol Rd., alpine slopes, alt. 5000'—
6000’, Wynne-Edwards 8378 (Can.); Macmillan Pass, Canol Rd., mi. 225 E., Porsild
11266 (Can.).

Common on alpine slopes, and occasional on the shore of Brintnell L. This
is the only goldenrod observed in the Brintnell L. area: in flower early in July.
Noted in Fl. Bor.-Am. at “Mackenzie R. and Bear Lake.”

Solidago multiradiata Ait. var. scopulorum Gray in Proc. Am. Acad. 17: 187 (1882) ;
Raup in Jour. Arn. Arb. 17: 300 (1936); Porsild in Nat. Mus. Can. Bull. 101:
32 (1945).
? Solidago Virgaurea sensu Hooker, Fl. Bor.-Am. 2: 5 (1834), in part, non L.
Resolution, Kennicott (N); Lone Mt., Wynne-Edwards 8514 (Can.) ; Sekwi R., Canol
Rd., mi. 174 E., Porsild 11883 (Can.).
This is the common form of the species in the central part of the Mackenzie
basin. It is probable that Hooker’s record for S. Virgaurea, “Woody country
between lat. 54° and 64°,” is applicable to it.

Solidago decumbens Greene var. oreophila (Rydb.) Fernald in Rhod. 38: 202 (1936);
Raup in Jour. Arn. Arb. 17: 300 (1936).
Solidago oreophila Rydb. in Mem. N. Y. Bot. Gard. 1: 387 (1900).
Solidago decumbens sensu Porsild in Nat. Mus. Can. Bull. 101: 34 (1945), in part, non
Greene. °

Solidago stricta sensu Hooker, Fl. Bor.-Am. 2: 4 (1834), in part, non Ait.

Fort Smith, Raup 1367 (Coll. Mrs. Conibear) ; Little Buffalo R., Russell 46 (G);
Resolution, Onion, Kennicott & Hardisty (N); Liard R. between Nahanni Butte and
Simpson, Crickmay 123 (Can.) ; upland prairie west of Mackenzie R. near Simpson, 9912;
Saline R., above Norman, Wynne-Edwards 8578 (Can.).

Common in the Wood Buffalo Park and in the Simpson locality above men-
tioned; achenes mature in late August. On the lower Mackenzie this variety
probably merges insensibly with the typical form.

Solidago lepida DC. var. elongata (Nutt.) Fernald in Rhod, 17: 9 (1915); Raup in
Jour. Arn. Arb. 17: 300 (1936); Porsild in Nat. Mus. Can. Bull. 101: 34 (1945).

Solidago elongata Nutt. in Trans. Am. Phil. Soc. 7: 327 (1840) ; Macoun, Cat. Can. Pl.
Pt. 2: 216 (1884), in part.

Solidago canadensis sensu Hooker, Fl. Bor.-Am. 2: 1 (1834), in part; Macoun, Cat. Can.
Pl. Pt. 2: 216 (1884), in part; Raup in Nat. Mus. Can. Bull. 74: 165 (1935), in part;
non L,

Solidago procera sensu Hooker, 1. c. 2, in part, non Ait.

Great Slave L., Richardson (G) ; willow thickets on sand bars along Mackenzie R. near
Simpson, 9861; rocky-sandy bank of Mackenzie R. opposite Simpson, 9965; Mackenzie R.
between Wrigley and Blackwater R., Crickmay 145 (Can.) ; near Wrigley, Miss E. Taylor
136 (Can.) ; Saline R. above Norman, Wynne-Edwards 8579 (Can.).

Common throughout the central Mackenzie basin and the upper Mackenzie
valley. The range of S. canadensis is given, “. . . from Lake Huron to Fort
Franklin, on the Mackenzie River” ; and of S. procera, “Woody country between
lat. 54° and 64°.”

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 239

Solidago lepida DC. var. fallax Fernald in Rhod. 17: 9 (1915); Raup in Jour. Arn.
Arb. 17: 301 (1936).
Lower Slave R., Raup 1357; Resolution, Kennicott (N).

Solidago graminifolia (L.) Salisb. var. major (Michx.) Fernald in Rhod. 46: 330 (1944).
Chrysocoma graminifolia L. Sp. Pl. 841 (1753), in part.
Solidago lanceolata L. var. major Michx. Fl. Bor.-Am, 2: 116 (1803) ; Hooker, Fl. Bor.-
Am. 2: 6 (1834).
Solidago graminifolia (L.) Salisb. var. camporum (Greene) Fernald, sensu Raup in
Jour. Arn. Arb. 17: 301 (1936).
Rocky-sandy bank of Mackenzie R. opposite Simpson, 9952.
Occasional in the Wood Buffalo Park region, but apparently rare along the
Mackenzie. Hooker gave its northern range, however, “Throughout Canada to

lat. 64°.”

Aster alpinus L. ssp. Vierhapperi Onno in Bibl. Bot. 26: 25, t. 6, f. 4 (1932); Porsild in
Sargentia 4: 67 (1943), in Nat. Mus. Can. Bull. 101: 32 (1945).

Liard R. between Nahanni Butte and Simpson, Crickmay 120 (Can.) ; north peak of
Nahanni Mt., Wynne-Edwards 8516 (Can.); dry summit of Lone Mt., Wynne-Edwards
8517 (Can.).

Porsild has collected this species at Great Bear L. and on Mt. Charles and Bear

Rock along Bear R.

Aster sibiricus L. Sp. Pl. 872 (1753); Porsild in Nat. Mus. Can. Bull. 101: 26, 32, 33
(1945).
Aster Richardsonii Spreng.; Hooker, Fl. Bor.-Am. 2: 7 (1834); Raup in Jour. Arn.
Arb. 17: 301 (1936).

Aster montanus Richardson; Macoun, Cat. Can. Pl. Pt. 2: 219 (1884).

Great Slave L., R. Bell 23159 (Can.) ; southwest and north shores of Great Slave L.,
Howe 91990 (Can.); northwest shore of Great Slave L., Bedford (Can.); Rae, Russell
(Can.); Liard R. between Nahanni Butte and Simpson, Crickmay 115 (Can.); willow
thickets on sandbar in Mackenzie R. near Simpson, 9865; Simpson, near sandy beach west
of island, Nowosad (Can.); outwash gravel plain near mouth of N. Nahanni R., Wynne-
Edwards 8540 (Can.) ; Mackenzie R. between Wrigley and Blackwater R., Crickmay 142
(Can.) ; between Blackwater R. and Norman, Crickmay 174 (Can.); Norman, Coll.? (N) ;
Bear R. at Norman, Miss E. Taylor 14 (G); mossy banks of south shore of Brintnell L.,
9597; damp gravelly thickets at west end of Brintnell L., 9434; dry rocky gully on south
slope of granite mountain west of Brintnell L., 9485; shale rock-slides on north slope of
Colonel Mt., alt. 5000’-6000', 9640; shale cliffs on south slope of Red Mt., alt. 5500’, 9741;
north peak of Nahanni Mt., Wynne-Edwards 8515 (Can.) ; Bolstead Cr., Canol Rd., mi.
111 E., Wynne-Edwards 8379 (Can.) ; Macmillan Pass, Canol Rd., mi. 225 E., Porsild
11278 (Can.).

Occasional to common throughout the Mackenzie valley and high in the
Mackenzie Mts.; collected in flower during July and early August.

Mackenzie basin and northern Rocky Mt. material of this complex, commonly
reported as A. Richardsonii Spreng., is apparently not distinct from A. sibiricus
of the Bering Sea region (See Porsild in Rhod. 41: 291; Scamman in Rhod. 42:
339; Hultén, Fl. Kamtch. 4: 157-158, and Fl. Aleut. I. 317-318). Our speci-
mens are variable in the serration of the leaves and in the size of the heads and
shape of the involucral bracts, but the variations do not seem worthy of taxonomic
distinction, as noted by Hultén (Fl. Kamtch.). Most of our Brintnell L. material
is low in stature, and has one to few rather small flowers to the stem, though no.
9485 is taller (3-4 dm.) and has single large heads.

In 1934 I proposed that the more southern Rocky Mt. relatives of this species,
commonly called A. meritus A. Nels., be considered a variety of the northern

240 SARGENTIA [6

A. Richardsonii (See Contr. Arn, Arb. 6: 202-204). The southern plants are
taller, less pubescent, and have much-branched stems with many short-pediceled
heads. In view of the above realignment of A. Richardsonii, the following new
combination is necessary: Aster sibiricus L. var. meritus (A. Nels.) comb. nov.
(Aster meritus A. Nels. in Bot. Gaz. 37: 268-269, 1904).

Aster foliaceus Lindl. in DC. Prodr. 5: 228 (1836).
Trout Lake, C. O. Hage (G).

This material matches plants collected in the Peace River Pass and in the
Lesser Slave L. district (see Raup, 1934).

Aster Lindleyanus T. & G. Fl. N. Am. 2: 122 (1841); Macoun, Cat. Can. Pl. Pt. 2: 222
(1884), at least in part; Raup in Jour. Arn. Arb, 17: 301 (1936); Porsild in Sar-
gentia 4: 68 (1943).

Aster paniculatus sensu Hooker, Fl. Bor.-Am. 2: 8 (1834), in part, non Nees.

Aster praecox sensu Hooker, Fl. Bor.-Am. 2: 9 (1834), in part, non Nees.

Fort Smith, Raup 1353 (Mrs. Conibear coll.) ; Resolution, Kennicott (N); Great Slave
L., Howe 91065 (Can.) ; thickets along wood road near Simpson, 9882; upland woods and
prairies west of Mackenzie R. near Simpson, 9922, 9923.

Occasional to common in the upper Mackenzie country, and collected by
Porsild at Mt. Charles along the Bear R. Among the ranges given by Hooker
for his segregates in this complex, the farthest north is “Fort Franklin, on the
Mackenzie River.” Collected with flowers and maturing achenes at Simpson
late in August.

Aster Lindleyanus T. & G. var. ciliolatus (Lindl.) Gray, Syn. Fl. 1: Pt. 2, 182 (1884);
Raup in Jour. Arn. Arb. 17: 302 (1936).

Aster ciliolatus Lindl. in Hooker, Fl. Bor.-Am. 2: 9 (1834).
Great Slave L., Richardson (G).
This specimen is probably part of the type collection of A. ciliolatus.

Aster Lindleyanus T. & G. var. comatus Fernald in Eggleston (FI. Vt.) in Rhod. 6:
142 (1904); Raup in Jour. Arn. Arb. 17: 302 (1936).

Mackenzie R., Kennicott (N).

This variety is occasional in the Wood Buffalo Park. Its presence on the
Mackenzie, at an unknown locality, is based upon the single specimen of Kenni-
cott.

Aster ericoides L. Sp. Pl. 875 (1753); Raup in Jour. Arn. Arb. 17: 302 (1936). See
Mackenzie in Rhod. 28: 65 (1926), and Blake in Rhod. 32: 136-140 (1930).

Little Buffalo R. near Lobstick Cr., Russell 2 (G).
Common in dry prairies of the Wood Buffalo Park, and probably extending
into similar habitats in the upper Mackenzie valley.

Aster falcatus Lindl. in Hooker, Fl. Bor.-Am. 2: 12 (1834); DC. Prodr. 5: 241 (1836);
Macoun, Cat. Can, PI. Pt. 2: 223 (1884); Raup in Jour. Arn. Arb. 17: 303 (1936).

Aster ramulosus Lindl. in Hooker, Fl. Bor.-Am. 2: 13 (1834).

Mackenzie R., Onion & Ross (N); willow thickets on sand bars in Mackenzie R. near
Simpson, 9866; rocky-sandy bank of Mackenzie R. opposite Simpson, 9964; Simpson, near
sandy beach west of island, Nowosad (Can.); Wrigley, Kindle (Can.); Saline R., above
Norman, Wynne-Edwards 8580 (Can.).

Common in the Simpson habitats ; achenes maturing in late August. With its
long, linear, glabrescent stem-leaves and but little imbricated involucral bracts,
this material resembles a part of the type of A. falcatus at the Gray Herbarium,

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 241

The latter is a rather strict plant, with relatively few heads, while ours are much
branched and have many heads. The Simpson specimens are a good match for
those which I have collected at McMurray (7069) and Chipewyan (4684). Aster
falcatus is noted in Fl. Bor.-Am. simply as occurring in “Arctic America”; but
A. ramulosus, which appears to be identical with it, is said to extend from “Sas-
katchewan to Fort Franklin on the Mackenzie River. . . .”

Aster junceus Ait. Hort. Kew. 3: 204 (1789); Macoun, Cat. Can. Pl. Pt. 2: 225 (1884);
Raup in Jour. Arn. Arb. 17: 302 (1936).
Aster laxifolius sensu Lindl. in Hooker, Fl. Bor.-Am. 2: 10 (1834), in large part, non
Nees.

Aster Franklinianus Rydb. in Bull. Torr. Bot. Cl. 37: 141 (1910).

Little Buffalo R., Russell 42, 78 (G); Great Slave L., Richardson (G, N); Resolution,
Onion, Kennicott & Hardisty (N); Rae, Russell (Can.) ; Mackenzie R., Kennicott (N);
fence-row thicket at Simpson, 9846; willow thickets on sand bars in Mackenzie R. near
Simpson, 9862, 9863; rocky-sandy bank of Mackenzie R. opposite Simpson, 9953; Simpson,
near sandy beach west of island, Nowosad (Can.) ; Mackenzie R. between Wrigley and
Blackwater R., Crickmay 146 (Can.).

This is one of the most abundant asters in the central Mackenzie basin. It is
common at Simpson, and apparently throughout the upper Mackenzie valley.
Collected with flowers and maturing achenes late in August. The concept of
Aster junceus here used is a broad one. Much of our Mackenzie basin material
seems to represent the species in its strict sense, but there is wide variation which
may, with more specimens and more study, justify subdivision.

Aster spathulatus Lindl. in Hooker, Fl. Bor.-Am. 2: 8 (1834).

Aster ? ptarmicoides sensu Porsild in Nat. Mus. Can. Bull. 101: 34 (1945), non T. & G.

Mackenzie R. between Wrigley and Blackwater R., Crickmay 143 (Can.); Saline R.,
above Norman, Wynne-Edwards 8581 (Can.).

The above specimens are a good match for Richardson specimens from Great
Bear L. in Herbs. G and N. These plants are evidently part of the type ma-
terial, which was stated in Fl. Bor.-Am. to have come from “Bear Lake and Fort
Franklin, on the Mackenzie River. Dr. Richardson.”

Aster angustus (Lindl.) T. & G. Fl. N. Am. 2: 162 (1841).

Tripolium angustum Lindl. in Hooker, Fl. Bor-Am. 2: 15 (1834).

Stony bank of Mackenzie R. a few miles below Providence, 9975.

Occasional or rare; found with maturing achenes Sept. 9. The type of Aster
angustus “8. ramis polycephalis” apparently came from Great Slave Lake, col-
lected by Richardson. Our plants from the upper Mackenzie answer the de-
scription very closely. So far as I am aware, the species had not been collected
in the Mackenzie basin since Richardson’s time except in the upper Peace
River region (Raup in Jour. Arn, Arb. 23: 25, 1942).

Erigeron angulosus Gaudin var. kamtschaticus (DC.) Hara in Rhod. 41: 389 (1939).
See also Fernald in Rhod. 40: 346 (1938).
Erigeron kamtschaticus DC. Prodr. 5: 290 (1836).
Erigeron glabratus sensu Hooker, Fl. Bor.-Am, 2: 18 (1834), non Hoppe.
Erigeron acris L, var. droebachensis Blytt sensu Macoun, Cat. Can. Pit 24 235 (1884).
Erigeron acris L. var. asteroides (Andrz.) DC.; Raup in Jour. Arn. Arb. 17: 305 (1936) ;
Porsild in Sargentia 4: 68 (1943), in Nat. Mus. Can. Bull. 101: 34 (1945).

Northwest shore of Great Slave L., Bedford (Can.); Saline R., above Norman, Wynne-
Edwards 8582 (Can.) ; gravelly banks and thickets along shore of Brintnell L. at Camp Pt.,
9573; in old cabin clearing near east end of Brintnell L., 9596.

242 SARGENTIA, [6

Occasional to common in the habitats mentioned. It is common in the cen-
tral Mackenzie basin, and Porsild has collected it at Great Bear L. Noted in
Fl. Bor.-Am. “. . . from the Saskatchewan to Fort Franklin on the Mackenzie
River.”

Erigeron elatus Greene in Pittonia 3: 164 (1897). See Fernald in Rhod. 40: 347 (1938).

Erigeron acris L. var. arcuans Fernald; Raup in Jour. Arn. Arb. 17: 303 (1936).

Fort Smith, Miss E. Taylor (N); Little Buffalo R., Russell 77 (G); Mackenzie kee
Onion (N); muskeg thickets on the north and west shores of Brintnell L., 9352, 9707:
cleared muskeg at east end of Brintnell L., 9351.

Common in the Brintnell L. habitats noted ; in flower early in July, and achenes
mature early in August. This species is common in the Athabaska—Great Slave
L. region, and I have seen a specimen from as far north as Good Hope (Miss E.
Taylor, Can.).

Erigeron jucundus Greene in Pittonia 3: 165 (1897).
Erigeron acris L. var. debilis Gray; Raup in Jour. Arn. Arb. 17: 303 (1936). See Rhod.
12: 225 (1910).

Shale cliffs on south slope of Red Mt., alt. 5500’, 9739, 9755; shale crevices and slide rock
on south slope of Terrace Mt., alt. 5000’, 9687.

Occasional to common above timber line at Brintnell L.; in flower early in
August. Apparently rare in the Mackenzie basin. I have collected it only in
the mountains along the upper Peace R., and at the eastern end of Great Slave L.
Part of our Brintnell L. material is peculiar in having rather thick stems, but
otherwise it seems fairly typical.

Erigeron unalaschkensis (DC.) Vierh. in Beih. Bot. Centralbl. 19: 492 (1906); Porsild
in Nat. Mus, Can. Bull. 101: 26 (1945).
Erigeron pulchellum var. unalaschkense DC. Prodr. 5: 287 (1836).

Turfy thickets and mossy banks on north slope of Colonel Mt., alt. 4000’-5000', 9403,
9639; rocky places along brook on west slope of Terrace Mt., alt. 5000’, 9523; Plains of
Abraham, Canol Rd., mi. 82 E., Porsild 11826 (Can.); Bolstead Cr., Canol Rd., mi. 111
E., Wynne-Edwards 8380 (Can.).

Rare to occasional in the Brintnell L. area; found flowering during most of
July.
Erigeron eriocephalus J. Vahl in FI. Dan. 13: ¢, 2200 (1840). See Polunin in Nat.

Mus. Can. Bull. 92: 343-345 (1940); Malte in Rhod. 36: 192 (1934).

Rocky places along small brook on west slope of Terrace Mt., alt. 5000’, 9522; mountain
range west of Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8382 (Can.).

Common on the first-mentioned site; in flower about mid-July. Erigeron
eriocephalus is here growing with E. unalaschkensis, from which it is easily dis-
tinguished in the field.

Erigeron compositus Pursh var. trifidus (Hooker) Gray in Proc. Am. Acad. 16: 90
(1880); Raup in Jour. Arn. Arb. 17: 303 (1936); Porsild in Sargentia 4: 68 (1943).
See Contr. Gray Herb. 49: 72-79 (1917); Univ. Wyo. Publ. Bot. 1: 176 (1926).

Erigeron trifidus Hooker, Fl. Bor.-Am. 2: 17 (1834).

Liard R. between Nahanni Butte and Simpson, Crickmay 117 (Can.).

An arctic and alpine species, found in the central Mackenzie basin only on
Great Slave L. Porsild has identified variety trifidus at Great Bear L. and on
Mt. Charles, but assigns most of his western arctic material to the typical form.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 243

Erigeron glabellus Nutt. Gen. Am. Pl. 2: 148 (1818); Hooker, Fl. Bor.-Am. 2: 19
(1834), in part; Macoun, Cat. Can. Pl. Pt. 2: 232 (1884); Raup in Jour. Arn. Arb.
17: 304 (1936).

Erigeron pulchellus sensu Hooker, Fl. Bor.-Am. 2: 19 (1834), in part, non Michx.

Resolution, Kennicott (N); Liard R. between Nahanni Butte and Simpson, Crickmay
118, 119, 179 (Can.) ; near Wrigley, Miss E. Taylor 121 (Can.) ; same place, Kindle (Can.).

This species is rather common on dry prairies in the Wood Buffalo Park, and
probably extends into similar habitats in the upper Mackenzie valley. Most of
the material cited in Fl. Bor.-Am. which may be referred here is said to range
northward in the “woody country” to lat. 64°.

Erigeron philadelphicus L. Sp. Pl. 863 (1753); Hooker, Fl. Bor.-Am. 2: 19 (1834),
in part; Macoun, Cat. Can. Pl. Pt. 2: 233 (1884); Raup in Jour. Arn. Arb. 17:
304 (1936); Porsild in Nat. Mus. Can. Bull. 101: 33 (1945).

Erigeron purpureus Ait. sensu Hooker, Fl. Bor.-Am. 2: 19 (1834), in part.

Lower Slave R., Raup 1344; Mackenzie R., Kennicott (N); south bank of Mackenzie
R., 10 mi. below Mills L., Wynne-Edwards 8559 (Can.) ; sand bar along. Mackenzie R. near
Simpson, 9858; old clearing at Simpson, 9131; Wrigley, Miss E. Taylor 43 (G).

Occasional; found in flower both early and late in the summer. In FI. Bor.-
Am. this species is said to range northward to lat. 64°, while E. purpureus is
noted “From Lake Huron . . . to the Arctic Circle.”

Erigeron hyssopifolius Michx. Fl. Bor.-Am. 2: 123 (1803); Raup in Jour. Arn. Arb. 17:
304 (1936); Porsild in Nat. Mus. Can. Bull. 101: 26, 32 (1945).

Smith Portage, Miss E. Taylor 11157 (Can.); Little Buffalo R., Russell 81 (G); Pine
Pt., Great Slave L., Brooke (Can.); Windy Pt., Great Slave L., Hume 102687 (Can.) ;
northwest shore of Great Slave L., Bedford (Can.) ; Mackenzie R., Ross (N); and Onion
& Ross (N); Liard R. between Nahanni Butte and Simpson, Crickmay 116 (Can.); north
peak of Nahanni Mt., Wynne-Edwards 8518 (Can...

This species was noted by Porsild as common along a small tributary of
Little Keele R., Canol Rd., mi. 51 E. It is occasional in the Wood Buffalo Park,
and I have seen a specimen from Great Bear L. (J. M. Bell, Can.).

Antennaria monocephala DC. Prodr. 6: 269 (1837); Porsild in Nat. Mus. Can. Bull.
101: 27 (1945). See Rhod. 36: 101-117 (1934).

Mossy bank on stony shore of Brintnell L. at Camp Pt., 9156; turfy shale on north slope
of Colonel Mt., alt. 4300’, 9662; south and east slopes of Red Mt., alt. 5000’-5500’, 9286,
9722, 9758; high shale slopes on west side of Terrace Mt., alt. 6000’, 9552; Bolstead Cr.,
Canol Rd., mi. 111 E., alpine slopes, alt. 5000’-6000', W ynne-Edwards 8386, 8387 (Can.).

Common on alpine slopes at Brintnell L., but rare at lake-level; found flower-
ing in late June, and with mature heads early in August. Porsild notes it as
common also in Macmillan Pass.

Antennaria densifolia A. E. Porsild in Nat. Mus. Can. Bull. 101: 26 (1945).

Nahanni Mt., Wynne-Edwards 8521 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., alt.
about 6000’, Wynne-Edwards 8384 (Can., TYPE coLL.) ; Sekwi R., Canol Rd., mi. 174 E.,
on dry turfy limestone scree, alt. 3625’, Porsild 11881 (Can.).

Porsild reports this new species also from Mt. Charles, along the Bear R.

Antennaria Ekmaniana A. E. Porsild in Sargentia 4: 69 (1943), in Nat. Mus. Can.
Bull. 101: 27 (1945).
Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11823 (Can.) ; Bolstead Cr., Canol Rd.,
mi. 111 E., rocky slope, Wynne-Edwards 8385 (Can.).

Antennaria isolepis Greene in Ottawa Nat. 25: 41 (1911); Porsild in Sargentia 4: 71
(1943), in Nat. Mus. Can. Bull. 101: 27, 32 (1945). See Malte in Rhod. 36:
108 (1934). :

244 SARGENTIA IG

Damp margin of lagoon in gravel plain at west end of Brintnell L., 9454; on gravelly cut
bank of Frost Cr., in upper valley, alt. 4500’, 9512; dry dolomitic summit of Lone Mt.,
Wynne-Edwards 8519 (Can.); Bolstead Cr., Canol Rd. mi. 111 E., W ynne-Edwards
8384-A (Can.).

Common locally at Brintnell L.; in flower in late June and mid-July.

Antennaria media Greene in Pittonia 3: 286 (1898), 4: 84 (1899),
Turfy shale slopes on hogback, north slope of Colonel Mt., alt. 4300’, 9648.
Occasional ; in flower late in July.

Antennaria rosea (D. C. Eat.) Greene in Pittonia 3: 281 (1898); Raup in Jour. Arn.
Arb. 17: 304 (1936); Porsild in Nat. Mus. Can. Bull. 101: 33 (1945).

Fort Smith, Raup 1379; lower Slave R., Raup 1380; Great Slave L., R. Bell 23160
(Can.) ; southwest and north shores of Great Slave L., Howe 91995 (Can.); northwest
shore of Great Slave L., Bedford (Can.) ; south bank of Mackenzie R., 10 mi. below Mills
L., Wynne-Edwards 8561 (Can.) ; Providence, Preble & Cary 39 (US); Liard R. between
’ Nahanni Butte and Simpson, Crickmay 111 (Can.).

Common in the Athabaska—Great Slave L. region, and at least occasional in
the upper Mackenzie country.

Antennaria sp.
In scrub timber on south slope of Red Mt., north of Brintnell L., alt. 3000’-4000', 9214.
Occasional ; collected with young flowers on June 21.

Antennaria subviscosa Fernald in Rhod. 16: 131 (1914). See also Fernald in Rhod.
35: 334 (1933), and 47: 232, t. 916, 917 (1945).

Scrub timber on south slope of Red Mt., north of Brintnell L., 9215.
This is an eastern species, previously known only from the Gaspé Peninsula.

Antennaria nitida Greene in Pittonia 3: 283 (1898); Raup in Jour. Arn. Arb. 17: 305
(1936); Porsild in Sargentia 4: 71 (1943), in Nat. Mus. Can. Bull. 101: 33 (1945).
South bank of Mackenzie R., 10 mi. below Mills L., Wynne-Edwards 8560 (Can.) ;
Providence, Dutilly 67, 69 (G); upland prairie west of Mackenzie R. near Simpson, 9910,
Common in the Athabaska—Great Slave L. district, and at least occasional in
the upper Mackenzie valley. Porsild has collected it at Bear Rock, along the
Mackenzie, and at Great Bear L. At Simpson the heads are mature in late
August.

Antennaria campestris Rydb. in Bull. Torr. Bot. Cl. 24: 304 (1897); Raup in Jour.
Arn. Arb. 17: 305 (1936).

Southwest and north shores of Great Slave L., Howe 91996 (Can.).
Occasional in dry prairies of the Wood Buffalo Park, and probably extending
into the upper Mackenzie country. In flower about mid-June.

Antennaria oxyphylla Greene in Pittonia 4: 284 (1901); Raup in Jour. Arn. Arb. 17:
305 (1936).

Fort Smith, Raup 1375.

So far as is known this species just reaches the southeastern corner of our
region. It is probable, however, that it will be found farther to the northwest-
ward in the upper Mackenzie valley. It is occasional in dry upland woods in
the Athabaska Lake region.

Antennaria pulcherrima (Hooker) Greene in Pittonia 3: 176 (1897); Raup in Jour. Arn.
Arb. 17: 304 (1936); Porsild in Sargentia 4: 72 (1943), in Nat. Mus. Can. Bull.
101: 32 (1945).
Antennaria carpathica var. pulcherrima Hooker, Fl. Bor.-Am, 1: 329 (1834).
Lone Mt., Wynne-Edwards 8520 (Can.).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 245

Rare or occasional in the central Mackenzie basin, and apparently also in the
upper Mackenzie valley. Porsild collected it at several places on Great Bear L.,
however. In flower about mid-July.

Helianthus annuus L. Sp. Pl. 904 (1753).

Weed in a grain field on the west bank of the Mackenzie R. near Simpson, 9932.

This species is doubtfully included in the spontaneous flora. It was in flower
late in August, and whether or not it was able to mature its seed is unknown.
It probably came in with seed grain, and is nothing more than a waif.

Helenium autumnale L. Sp. Pl. 886 (1753); Hooker, Fl. Bor.-Am. 1: 317 (1834);
Macoun, Cat. Can. Pl. Pt. 2: 249 (1884).

Mackenzie R., Onion, Kennicott & Hardisty (N); rocky-sandy bank of Mackenzie R.
opposite Simpson, 9956; in silty soil near Mackenzie R. at Simpson, Nowosad (Can.).

Common at the Simpson locality; collected with maturing heads and a few
flowers late in August. The northern form of H. autumnale has been designated
by some students as var. grandiflorum (Nutt.) Gray (H. macranthum Rydb.)
(See Gray, Syn. Fl. 1: Pt. 2, 349). The size of the heads, however, is so vari-
able, both in the same colony and on the same plant, that the separation seems
hardly justifiable. This species was noted in Fl. Bor.-Am., “Throughout British
North America, from Lake Huron nearly to the Arctic Circle... ,” and there
is a sheet in Herb. G labeled “High northern specimen Coll. in Franklin Exped.”
There is also one in Herb. Can. marked “British N. Am. Richardson.” With
the exception of the Onion, Kennicott & Hardisty specimen, our material from
Simpson is the first authentication of the earlier records of the species in the en-
tire Mackenzie basin. The Herb. N specimen may well have come from the
vicinity of Simpson, as I. S. Onion, one of the principal contributors, was lo-
cated there for some years.

Achillea sibirica Ledeb. Ind. Sem. Hort. Dorpat (1811), Fl. Ross. 2: 528 (1844); Raup
in Jour. Arn. Arb. 17: 305 (1936); Porsild in Sargentia 4: 73 (1943), in Nat.
Mus. Can. Bull. 101: 35 (1945).

Achillea multiflora Hooker, Fl. Bor.-Am. 1: 318 (1834) ; Macoun, Cat. Can. Pl. Pt. Y
251 (1884).
Mackenzie R., Onion (N); rocky-sandy bank of Mackenzie R. opposite Simpson, 9948;
five mi. above Old Fort Norman, Wynne-Edwards 8566 (Can.):
Occasional to common along the main rivers; found with maturing heads late
in August. Noted in Fl. Bor.-Am., “Throughout the woody country, and as far
north as Fort Franklin.” Porsild has collected it in the Mackenzie River delta.

Achillea Millefolium L. Sp. Pl. 899 (1753); Hooker, Fl. Bor.-Am. 1: 318 (1834), in
part; Macoun, Cat. Can. Pl. Pt. 2: 251 (1884), in part; Raup in Jour. Arn. Arb.
17: 306 (1936).

Fort Smith, Raup 1398, 1399 (Coll. Mrs. Conibear) ; Little Buffalo R. near 60th parallel,
Russell 85 (G) ; junction of Little Buffalo and Nyarling Rivers, Russell 8 (G) ; Mackenzie
R., Onion (N); rocky-sandy bank of Mackenzie R. opposite Simpson, 9949; Bear R., Miss
E. Taylor 88 (Can.). ;

Common throughout the central Mackenzie basin and at least the upper Mack-
enzie valley; collected with flowers and maturing heads late in August.

Achillea borealis Bong. in Mém. Acad. Imp. Sci. St. Pétersb. I1V. Math. 2: 149 (1833);
Raup in Jour. Arn. Arb. 17: 306 (1936); Porsild in Nat. Mus. Can. Bull. 101:
27, 34, 35 (1945).
Achillea Millefolium sensu Hooker, Fl. Bor.-Am. 1: 318 (1834), in part; Macoun, Gat.
Can. Pl. Pt. 2: 251 (1884), in part; non L,

246 SARGENTIA: [6

Liard R. between Nahanni Butte and Simpson, Crickmay 110 (Can.); Saline R., above
Norman, Wynne-Edwards 8583 (Can.) ; Macmillan Pass, Canol Rd., mi. 225 E., Porsild
11279 (Can.).

Matricaria ambigua (Ledeb.) Kryl. Fl. Alt. 3: 625 (1904); Porsild in Sargentia 4:
73 (1943). See M. P. Porsild in Medd. om Grgnl. 92, 1: 72 (1932).

Pyrethrum inodorum sensu Hooker, Fl. Bor.-Am. 1: 320 (1834), non Sm.

Matricaria inodora sensu Macoun, Cat. Can. Pl. Pt. 2: 253 (1884), non L.

Pyrethrum ambiguum Ledeb. Fl. Alt. 4: 118 (1833).

I include this species with much uncertainty. Porsild (1. c.) states that it is
“A common species along the Arctic Coast and thus far not collected in the in-
terior of our district.” Nevertheless it is noted in Fl. Bor.-Am. as occurring
“. . . as far north as Bear Lake,” indicating that it was collected somewhere along
the route of the Franklin expeditions as they traveled northward. There are
Richardson specimens in Herbs. G and N, but all of them are without specific
locality data.

Chrysanthemum integrifolium Richardson, Bot. App. Frankl. Jour. 749 (1823); Porsild
in Nat. Mus. Can. Bull. 101: 27 (1945).
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8398, 8399 (Can.) ; Sekwi R., Canol
Rd., mi. 174 E., Porsild 11866 (Can.).

Tanacetum huronense Nutt. var. bifarium Fernald in Rhod. 37: 334, t. 377, f. 3, 4 (1935);
Raup in Jour. Arn. Arb. 17: 307 (1936).

Tanacetum pauciflorum Richardson, sensu Hooker, Fl. Bor.-Am. 1: 327 (1834), in part.

This plant is somewhat doubtfully included here. It is common on the sandy
beaches around L. Athabaska, but I have seen no material from north of that
region. In Fl. Bor.-Am., however, it is noted as occurring in the “Woody coun-
try, between lat. 54° and 64°, North.” Whether this record is based entirely
upon specimens that may have been collected at L. Athabaska is unknown, but the
species is to be expected farther north.

Artemisia canadensis Michx. Fl. Bor.-Am. 2: 129 (1803); Macoun, Cat. Can. Pl. Pt.
2: 256 (1884); Raup in Jour. Arn. Arb. 17: 308 (1936); Porsild in Nat. Mus.
Can. Bull. 101: 34 (1945).

Artemisia desertorum Bess. in Hooker, Fl. Bor.-Am. 1: 325 (1834), in part.
Fort Smith, Raup 1388 (Coll. Mrs. Conibear) ; Resolution, Onion, Kennicott & Hardisty

(N); Saline R. above Norman, Wynne-Edwards 8584 ( Canzy.

Occasional to common in the central Mackenzie basin, though apparently less

common northward. I have seen a specimen from the Mackenzie R. about 30

mi. north of the Arctic Circle (Miss E. Taylor, Can.).

Artemisia biennis Willd. Phytogr. 11 (1794); Hooker, Fl. Bor.-Am. 1: 325 (1834);
Macoun, Cat. Can. Pl. Pt. 2: 259 (1884); Raup in Jour. Arn. Arb, 17: 308 (1936);
Porsild in Sargentia 4: 73 (1943).

Mackenzie R., Onion & Ross (N); open field at Simpson, 9857; sand bars along Mack-

enzie R. near Simpson, 9875,

Common in the Simpson area; in flower late in August. Porsild has collected
it as far north along the Mackenzie as the Ramparts above Good Hope.

Artemisia frigida Willd. Sp. Pl. 3: 1838 (1804); Macoun, Cat. Can. Pl. Pt. 2: 259
(1884); Raup in Jour. Arn. Arb. 17: 308 (1936); Porsild in Sargentia 4: 73 (1943).

East bank of Mackenzie R. about 7 mi. above Wrigley, Crickmay 28 (Can.).
This species has been collected only once in the upper Mackenzie valley,
though it is common at L. Athabaska and in the Wood Buffalo Park, and I have

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 247

seen specimens from the lower Mackenzie (M cConnell, Can.; Miss E. Taylor,
Can.). Porsild has collected it in the Mackenzie delta and on the arctic coast.
The only earlier record we have is in Macoun’s Catalogue: “Mackenzie River.
(McGill Coll. Herb.).”

Artemisia arctica Less. in Linnaea 6: 213 (1831); Porsild in Sargentia 4: 73 (1943),
in Nat. Mus. Can. Bull. 101: 27 (1945). See Hultén in Fl. Kamtch. 4: 176-180
(1930).

Turfy shale slopes on north side of Colonel Mt., alt. 4000’-6000', 9641; damp grassy
place in willow-birch scrub, upper valley of Frost Cr., alt. 4000’, 9555; summit and upper
slopes of Red Mt., 9216, 9738; Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes, alt. 5000’,
W ynne-Edwards 8388, 8389 (Can.).

Common to abundant in alpine areas at Brintnell L., and noted by Porsild
as a “dominant species” in Macmillan Pass. In flower during most of July and
early August. Porsild reports it common also in the Richardson Mts., and along
the arctic coast west of the Mackenzie.

Artemisia Tilesii Ledeb. in Mém. Acad. Imp. Sci. St. Pétersb. 5: 568 (1812); Raup in
Jour. Arn. Arb, 17: 308 (1936); Porsild in Nat. Mus. Can. Bull. 101: 27 (1945).

Artemisia vulgaris sensu Hooker, Fl. Bor.-Am. 1: 322 (1834), in part; Macoun, Cat.
Can. Pl. Pt. 2: 257 (1884), in part; non L.

Stony-gravelly shore of Great Slave L. at Wrigley Harbor, 9981; east shore of Trout
L., C. O. Hage (G); Mackenzie R., Kennicott (N) ; willow thickets on sand bars along
Mackenzie R. near Simpson, 9860; Bear R., Miss E. Taylor (Can.); thickets on gravel
beaches at Brintnell L., 9305, 9691; dry gully on lower slope of granite mountain west of
Brintnell L., 9472; gravelly thicket in gorge on north slope of Colonel Mt., alt. 3000’, 9832;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11277 (Can.).

Common in the habitats cited above, and occasional on river banks and lake
shores in the Peace and Athabaska R. regions. I have seen specimens also from
Great Bear L. (Richardson, Can.; J. M. Bell, Can.), and from Good Hope
(Onion, N). In flower about mid-July (see below).

The specimens collected by us in 1939 fall rather clearly into three groups
which are left provisionally in A. Tilesii sens. lat. Numbers 9981 and 9860 are
tall slender plants with moderate-sized, upright, more or less ovoid heads (in-
volucres about 4 mm. high, somewhat arachnoid) arranged in elongated panicu-
late inflorescences which overtop the leaves. The leaves are long-lanceolate,
glabrescent and bright green above, entire or with a few divergent acute lobes.
These plants are from Ft. Simpson and the western end of Great Slave Lake,
and they strongly resemble specimens from the Peace and Athabaska Rivers
(Raup 6001, 7074; Raup & Abbe 4359, 4393), as well as specimens from Hud-
son Bay (Robt. Bell 100; J. M. Macoun 79263). From the size of the heads
they might suggest A. Douglasiana Bess. (A. vulgaris subsp. heterophylla; see
Clausen, Keck & Heisey in Carn. Inst. Wash, Publ. no. 520: 327, and Hall &
Clements, ibid., Publ. no. 326: 76), but the leaves and habit seem to relate them
to the A. Tilesii group, perhaps to forms like the type of A. Gormani Rydberg,
which came from the L. Iliamna region of Alaska (Gorman 281).

A second group includes nos. 9472, 9832 and 9691, all from the vicinity of
Brintnell L. These are also tall plants whose leaves are glabrescent and green
above. The leaves are more deeply lobed than in group 1, however, though in
no. 9832 most of the upper ones are entire. The inflorescences are more loosely
paniculate, the fewer and much larger heads with involucral bracts up to 6 mm.
long, slightly arachnoid or nearly glabrous. The dry, pressed heads are up to
1 cm. across, and are nodding, on pedicels as much as 2 cm. long. These speci-

248 SARGENTIA [6

mens are a good match for material collected in Alaska (Scamman 848, 893,
1035; Porsild 1484, 1633), but no such large-headed forms have been seen in the
Mackenzie country. They appear to be nearest to A. Tilesii s. 1. as understood
by Hall & Clements (Carn. Inst. Wash. Publ. no. 326: 71-72), and to the var.
elatior T. & G. (A. Tilesii subsp. unalaschcensis (Bess.) Hultén in Fl. Aleut. T.
327 ; see also Clausen, Keck & Heisey in Carn. Inst. Wash. Publ. no. 520: 325—
344).

The third group contains only one number, 9305, but it shows the greatest di-
vergence from the concept of A. Tilesii as outlined above. The plants are much
lower in stature (up to 4 dm.), with simple, racemose inflorescences of a few
nodding heads. The involucral bracts are 4-5 mm. long, arachnoid, and the
heads are 6-7 mm. in diameter. The leaves, except the uppermost, are deeply
divided into linear-lanceolate lobes, and are arachnoid pubescent on both sides,
though darker green above. The closest relationship seems to be with the sec-
ond group mentioned above (A. Tilesii var. elatior), but the simpler inflores-
cences and more deeply lobed leaves, pubescent above, suggest a distinctive form.

Petasites frigidus (L.) Fries Summa Veg. Scand. 182 (1845); Macoun, Cat. Can. PI.
Pt. 2: 260 (1884); Porsild in Sargentia 4: 74 (1943), in Nat. Mus. Can. Bull. 101:
27 (1945).

Tussilago frigida L. Sp. Pl. 865 (4753).

Nardosmia frigida (L.) Hooker, Fl. Bor.-Am. 1: 307 (1834).

Nardosmia sagittata sensu Hooker, Fl. Bor.-Am. 1: 307 (1834), in part, non Pursh.

Nardosmia corymbosa (R. Br.) Hooker, Fl. Bor.-Am. 1: 307 (1834), in part.

Wet grassy place in willow scrub, upper valley of Frost Cr., north of Brintnell La ante
4000’, 9519; Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes, alt. 4500’-5000', Wynne-
Edwards 8397 (Can.) ; Macmillan Pass, Canol. Rd., mi. 225 E., Porsild 11270 (Can.).

At Brintnell L. this plant was found only in the single locality cited, though it
was common there. I have seen no material from the upper Mackenzie valley,
though it appears to be common along the lower Mackenzie, in the Richardson
Mts., and on the Arctic coast (Porsild, 1943). Petasites frigidus in our region is
readily distinguished from P. vitifolius, P. sagittatus, and P. palmatus, in the
absence of leaves, by its deeply alveolate receptacle. The spaces between the
alveolae consist of sharpened and often toothed ridges, whereas in the other three
species the pits are sunken in a more or less flat surface, Eurasian specimens
of P. frigida show the same character, but the ridges tend to be lower, blunter,
and less toothed.

Petasites palmatus (Ait.) Gray, Bot. Calif. 1: 407 (1876); Macoun, Cat. Can. Pl. Pt.
2: 260 (1884); Raup in Jour. Arn. Arb, 17: 309 (1936); Porsild in Nat. Mus. Can.
Bull. 101: 32 (1945).

Tussilago palmata Ait. Hort. Kew. 3: 188 (1789).

Nardosmia palmata (Ait.) Hooker, Fl. Bor.-Am. 1: 308 (1834).

Fort Smith, Raup 1435; summit of Lone Mt., Wynne-Edwards 8522 (Can.).

Noted in Fl. Bor.-Am., “From Lake Huron . . . to the Bear Lake, lat. 67°,
north,” and I have seen a specimen from the latter place (J. M. Bell, Can.). The
species is common in the central Mackenzie basin, but apparently rare or occa-
sional along the upper Mackenzie R.

Petasites vitifolius Greene in Leaflets 1: 180 (1906); Raup in Jour. Arn. Arb. 17: 309
(1936).

Resolution, Kennicott ? (N); Mackenzie R., Onion (N); Simpson, May 25, 1860, Coll.?
(N).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 249

This species is occasional to common in the Athabaska—Great Slave L. region,
but apparently less common northward. I have seen no authentic material from
north of Simpson.

Petasites sagittatus (Banks) Gray, Bot. Calif. 1: 407 (1876); Macoun, Cat. Can. Pl. Pt.
2: 260 (1884); Raup in Jour. Arn. Arb. 17: 309 (1936); Porsild in Sargentia 4:
75 (1943), in Nat. Mus. Can. Bull. 101: 27 (1945).
Tussilago sagittata Banks ex Pursh, Fl. Am. Sept. 2: 531 (1814).
Nardosmia sagittata (Banks) Hooker, Fl. Bor.-Am. 1: 307 (1834).
Clewi R., Russell 15 (G); old clearing in lowland woods near Simpson, 9102.

This species is abundant in the Wood Buffalo Park and at least occasional
northward to Great Bear L. and the Mackenzie delta. The Simpson specimens
were found with maturing achenes on June 12. Porsild noted it as rare in Mac-
millan Pass. .

All the species of Petasites were much confused (as Nardosmia) in FI. Bor.-
Am. This is obvious from the original labels on Hooker duplicates in Ameri-
can herbaria, and from photographs of specimens in the British Museum. Por-
sild has greatly clarified the tangle in Sargentia 4: 74, 75.

Arnica alpina (L.) Olin ssp. genuina Maguire in Brittonia 4: 408 (1943).

Gravel plain near shore of Brintnell L., 9200; rich mossy spruce woods on fan at base of
Colonel Mt., 9492; alpine meadows and scrub, north slope of Colonel Mt., 9384; birch scrub
on south slope of Terrace Mt., alt. 4500’, 9686.

Common on the high north slopes of Colonel Mt., but rare or occasional else-
where in the district; in flower in late June and early July. Maguire does not
admit genuine A. alpina to the American flora, but our Brintnell L. plants so
nearly match many northern European specimens of A. alpina in the Gray Her-
barium that I am forced to identify them with it. Most of these plants have nar-
row, lance-attenuate leaves which suggest ssp. attenuata, but they are not so gland-
ular as in the latter. Furthermore, the usual pair of small leaves high up on the
stem in ssp. attenuata is almost entirely wanting. The ligules of the flowers are
much more deeply cut than in any American material that I have seen (in some
cases to a depth of 5 mm.), thus again suggesting the European plant. Maguire
makes a key character of the glandularity on the involucre and upper part of the
stem. In ssp. genuina the stipitate glands are lacking or very short and ob-
scured, while in American varieties they are usually well developed. In some
of our plants they are almost entirely wanting, while in others they are prominent
only on the upper part of the stem. The latter apparently approach ssp. an-
gustifolia,

Arnica alpina (L.) Olin ssp. angustifolia (Vahl.) Maguire in Madrofio 6: 153 (1942);
Porsild in Sargentia 4: 75 (1943), in Nat. Mus. Can. Bull. 101: 27 (1945).

Arnica angustifolia Vahl, Fl. Dan. 926: 5 (1816).

; Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11824 (Can.) ; Bolstead Cr., Canol Rd.,
mi. 111 E., Wynne-Edwards 8392 (Can.).

Arnica alpina (L.) Olin ssp. attenuata (Greene) Maguire in Madrofio 6: 153 (1942);
Porsild in Sargentia 4: 76 (1943), in Nat. Mus. Can. Bull. 101: 32 (1945). See
also Maguire in Brittonia 4: 412 (1943).
Arnica attenuata Greene in Pittonia 4: 170 (1900); Raup in Jour. Arn. Arb. 17: 309
(1936).

North peak of Nahanni Mt., Wynne-Edwards 8523 (Can.).

250 SARGENTIA [6

This subspecies has been collected in the upper Mackenzie region only in the
locality cited above, but it is to be expected elsewhere. It is occasional around
Athabaska and Great Slave Lakes, and Porsild has collected it in the Mackenzie
delta and the Richardson Mts. Porsild also has specimens from Great Bear L.
which appear to be intermediate between this and ssp. angustifolia.

Arnica alpina (L.) Olin ssp. tomentosa (J. M. Macoun) Maguire in Madrofio 6: 153
(1942); Porsild in Sargentia 4: 76 (1943). See also Maguire in Brittonia 4:
413 (1943),
Arnica tomentosa J. M. Macoun in Ottawa Nat. 13: 168 (1899).
Liard R. between Nahanni Butte and Simpson, Crickmay 113 (Can.); loose shale slide
rock on south slope of Red Mt., north of Brintnell L., alt. 5500’, 9730.
Common in the one Brintnell L. habitat noted, but not seen elsewhere; in flower
early in August.

Arnica Louiseana Farr ssp. frigida (Meyer ex Ijin) Maguire in Madrofio 6: 153 (1942);
Porsild in Sargentia 4: 76 (1943), in Nat. Mus. Can. Bull. 101: 27, 32 (1945). See
also Maguire in Brittonia 4: 417 (1943).
Arnica frigida Meyer ex Iljin in Trav. Mus. Bot. Acad. U. R. S. S. 19: 112°(1926),
Dry limestone summit of Lone Mt., Wynne-Edwards 8526 (Can.); Bolstead Cr., Canol
Rd., mi. 111 E., Wynne-Edwards 8391 (Can.).
A western arctic subspecies, known also in the Richardson Mts., the Mackenzie
delta, and at Great Bear L.

Arnica Snyderi sp. nov. Fic. 16.

Herba perennis, rhizomate gracili, ramuloso horizontali; culmis scaposis vel
basim versus foliorum pare praeditis, pilos cellulares patentes flavo-fusco geren-
tibus, apicem versus lanatis, vix glandulosis, sub anthesi 6-15 cm. altis et 1—1.5
mm. crassis ; foliis satis crassis, anguste ovatis vel ellipticis vel valde rhomboideo-
ovatis, ad 6 cm. longis, costa prominente, pare unico nervorum lateralium satis
inconspicuo, apice obtusis vel acutis, ad basim contractis, infimis petiolis alatis,
supra tantum rugulosis vel breviter glanduloso-papillosis, subtus glabris pallidior-
ibus, margine integerrimis vel raro dentibus glandulosis paucis ornatis et breviter
hispido-ciliatis ; foliis inferioribus culmorum basalibus similibus, superioribus
valde reductis vel defectis; capitulis solitariis sub anthesi nutantibus, in fructu
erectis ; involucro 9-12 mm. alto, bracteis circiter 10 late lanceolatis, apice obtusis
vel acutis, marginem versus purpureo-fuscis, interdum medio subviridibus,
basim versus lanatis, superne extus glabris vel subglabris intus ciliato-pilosis ;
acheniis 44.5 mm. longis parce hirsutis ; pappo albo; floribus disci flavis circiter
7 mm. longis basim versus villosis, lobis parce villosis, antheris flavo-fuscis ;
floribus radii flavis 1.5-2.2 cm. longis ad 5 mm. latis, apice breviter lobatis.

Alpine crevices and rock-slides, north slope of Colonel Mt., July 5, 9383 (Type); shale
cliffs in gorge on north slope of Colonel Mt., alt. 3500’, 9823; loose slide rock on south
slope of Red Mt., alt. 5500’-5900', 9731; high shale slopes on west side of Terrace Mt., alt.
5500’-6000’, 9563.

Common in the Brintnell L. area; found flowering in July and early August.
This beautiful species has the single nodding heads and semi-scapose habit of the
far-northwestern group which contains A, obtusifolia and A. Lessingii. It is
separated from these, however, by its yellowish brown anthers and white pappus.
In these characters it resembles A. Louiseana Farr, which has proved to be of
wide range in boreal America (see Fernald in Rhod. 35: 368-369 ; Porsild in
Rhod. 41: 298; Maguire, 1. c.). When compared with an isotype of A. Louise-
ana at the Gray Herbarium, our plants prove to be much more scapose, with the
leaves nearly or quite entire, whereas they are saliently dentate in A. Louiseana.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 251

In the latter the leaves are glandular puberulent on both sides, and ovate- or obo-
vate-lanceolate, while in A. Snyderi they are glabrous beneath, glabrous or spar-
ingly glandular-papillose above, and are commonly broadly ovate to rhombic-
ovate. In leaf shape they resemble some small forms of the eastern representative
of A. Louiseana originally described by Fernald as A. Griscomi (Rhod. 26: 105,
1924), but the latter, now reduced to a subspecies of A. Louiseana, has the char-

KE CRO00k

Fic. 16. Arnica Snyderi Raup, sp. nov.

acteristically denticulate leaves. Furthermore, our plants have the upper parts
of the stem and the base of the involucre yellowish brown-lanate, while in A.
Louiseana ssp. Griscomi they are merely villous or villous-hirsute with white
hairs. According to Fernald (Rhod. 35: 368) and Maguire (1. c.), A, Louiseana
belongs in the group which centers around A. alpina, and its heads are not truly
nodding but only bent over in drying or wilting. All of our field notes, however,
state that the peduncles of A. Snyderi were bent while the flowers were in anthesis,

252 SARGENTIA [6

straightening later. It is true, however, that the bend is not a sharp one as in
A, Lessingii, and it may be that our plants’ affinities are with the A. alpina group.

Arnica lonchophylla Greene ssp. genuina Maguire in Brittonia 4: 430 (1943); Porsild
in Nat. Mus. Can. Bull. 101: 32 (1945).

Arnica chionopappa sensu Raup in Jour. Arn. Arb. 17: 309 (1936), non Fernald.

Fort Smith, Seton & Preble 78293 (Can.); same place, Raup 1408; lower Slave R.,
Raup 1409; Windy Pt., Great Slave L., Hume 102689 (Can.) ; northwest shore of Great
Slave L., Bedford (Can.); Old Fort Rae, Russell (Can.) ; north peak of Nahanni Mt.,
Wynne-Edwards 8524 (Can.); Lone Mt., Wynne-Edwards 8525 (Can,),.

Common in dry upland woods in the Athabaska-Great Slave L. region, but
apparently rarer northward. I have seen specimens from as far north as Good
Hope and Arctic Red River.

Arnica cordifolia Hooker ssp. genuina Maguire in Madrojio 6: 154 (1942).
Liard R. between Nahanni Butte and Simpson, Crickmay 114 (Can.).

Arnica Chamissonis Less. ssp. foliosa Maguire in Rhod. 41:. 508 (1939). See also
Maguire in Brittonia 4: 463, f. 14 (1943).

Arnica rhizomata A. Nels. sensu Raup in Jour. Arn. Arb, 17: 309 (1936).

Mackenzie R., Kennicott (N).

The Kennicott specimen is referred to this subspecies somewhat doubtfully.
Maguire (1. c.) has recorded ssp. foliosa for two places in southern Mackenzie,
one on the south shore of Great Slave L., and the other along the Mackenzie R.
near Simpson, but I have seen neither of these specimens. Maguire also has a
map citation of his 4. Chamissonis ssp. genuina in southwestern Mackenzie. I
presume, from the position of the dot, that this is based upon material from Simp-
son or the Nahanni Mts. He states, however, that this is a transition region
wherein it is difficult to define the subspecies.

Arnica mollis Hooker, Fl. Bor.-Am. 1: 331 (1834).
Loose shale slide rock on south slope of Red Mt., alt. 5800’, 9732.
Occasional in this habitat, but not seen elsewhere; in flower early in August.

Arnica Lessingii Greene in Pittonia 4: 167 (1900); Porsild in Sargentia 4: 76 (1943),
in Nat. Mus. Can. Bull. 101: 27 (1945). See also Maguire in Brittonia 4: 486
(1943).

Alpine crevices and slide rock, north slope of Colonel Mt., 9383-A; along brook on west
slope of Terrace Mt., alt. 4800’, 9573; Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards

8390 (Can.) ; Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11268 (Can.).

Common locally ; in flower during early and mid-July.

Senecio eremophilus Richardson, Bot. App. Frankl. Journ, ed. 2, repr. 31 (1823);
Hooker, Fl. Bor.-Am.°1: 334 (1834); Macoun, Cat. Can. Pl. Pt. 2: 266 (1884);
Raup in Jour. Arn. Arb. 17: 309 (1936).

This species has not yet been collected in southwestern Mackenzie, but is to
be expected there. It is occasional in the Athabaska and Slave R. lowlands, and
is said by Hooker to extend northward “to Fort Franklin, on the Mackenzie
River.”

Senecio triangularis Hooker, Fl. Bor.-Am. 1: 332, t. 115 (1834); Porsild in Nat. Mus.
Can. Bull. 101: 28 (1945).

Damp grassy place in willow-birch scrub, upper valley of Frost Cr., North of Brintnell L.,
alt. 4000’, 9516.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 253

Abundant in the locality noted, but not seen elsewhere in the Brintnell L. area ;
in flower about mid-July. Porsild states that it is very common in Macmillan
Pass, along the Canol Rd.

Senecio lugens Richardson, Bot. App. Frankl. Journ. ed. 2, repr. 31 (1823); Porsild
in Nat. Mus. Can. Bull. 101: 28, 32 (1945).

Mackenzie R., Onion (N); Mackenzie R. between Blackwater R. and Norman, Crickmay
177 (Can.) ; wet swamps and mossy banks on shore of Brintnell L., 9323, 9345; lowland
woods and thickets at west end of Brintnell L., 9429; alpine meadows and scrub, north slope
of Colonel Mt., 9379; shale cliffs and slides in gorge on north slope of Colonel Mt., alt.
3500’, 9820; damp hollow in summit of Lone Mt., Wynne-Edwards 8530 (Can.); north
peak of Nahanni Mt., Wynne-Edwards 8531 (Can.) ; Bolstead Cr., Canol Rd., mi. 111 E.,
Wynne-Edwards 8393 (Can.).

Common in the Mackenzie Mts. and at least occasional along the Mackenzie
R.; in flower during late June and early July. Porsild noted it as common in
a muskeg along a small tributary to the Little Keele R., and also in Macmillan
Pass.

Senecio cymbalarioides Nutt. var. borealis (T. & G.) Greenm. in Ann. Mo. Bot. Gard.
3: 177 (1916); Raup in Jour. Arn. Arb. 17: 310 (1936).

Senecio aureus sensu Hooker, Fl. Bor.-Am. 1: 333 (1834), in part, non L.

Senecio aureus L. var. borealis T. & G. Fl. N. Am. 2: 443 (1841) ; Macoun, Cat. Can. Pl.
Pt. 2: 265 (1884).

Senecio cymbalarioides sensu Porsild in Nat. Mus. Can. Bull. 101: 32 (1945).

Fort Smith, Miss E. Taylor 14790 (Can.); same place, Seton & Preble 78580 (Can.) ;
same place, Raup 1424, 1429; Windy Pt. Great Slave L., Hume 102688 (Can.) ; southwest
and north shores of Great Slave L., Howe 91998 (Can.) ; northwest shore of Great Slave L.,
Bedford (Can.); Mackenzie R., Onion & Ross (N); Providence, Dutilly 77 (G); Liard
R. between Nahanni Butte and Simpson, Crickmay 122 (Can.) ; Mackenzie R. between
Wrigley and Blackwater R., Crickmay 141 (Can.) ; north peak of Nahanni Mt., Wynne-
Edwards 8527 (Can.).

Common in the Athabaska-Great Slave L. region and the upper Mackenzie
valley.

Senecio indecorus Greene, Fl. Francisc. 470 (1897); Raup in Jour. Arn. Arb. 17: 310
(1936). See Fernald in Rhod. 26: 120 (1924).

Mackenzie R., Onion (N); “Mackenzie R., Ft. Franklin,” Richardson (Can.) ; muskeg
along road just west of Mackenzie R. near Simpson, 9900; muskeg thickets at west end of
Brintnell L., 9433, 9704.

Occasional to common; found flowering about mid-July and early in August.
Most of our plants have radiate heads. Common in the Wood Buffalo Park, but
its northern limits are yet unknown.

Senecio pauperculus Michx. Fl. Bor.-Am. 2: 120 (1803); Raup in Jour. Arn. Arb. 17:
310 (1936); Porsild in Sargentia 4: 77°(1943), in Nat. Mus. Can. Bull. 101: 32,
34 (1945). ;
Senecio aureus sensu Hooker, Fl. Bor.-Am. 1: 333 (1834), in part, non L.
Senecio Balsamitae sensu Hooker, Fl. Bor.-Am. 1: 332 (1834), in part, non Muhl.

Senecio aureus L. var. Balsamitae sensu Macoun, Cat. Can. Pl. Pt. 2: 265 (1884), in
part, non T. & G.

Lower Slave R., Raup 1417; Saline R., above Norman, Wynne-Edwards 8585 (Can.) ;
muskeg at west end of Brintnell L., 9438; north peak of Nahanni Mt., Wynne-Edwards
8532 (Can.).

Common in the Athabaska-—Great Slave L. region, but apparently only occa-
sional in the upper Mackenzie country. In flower about mid-July. Its known

254 SARGENTIA [6

northern limit on the Mackenzie appears to be in the neighborhood of Norman,
and it has been found at Great Bear L. by Porsild.

Senecio pauciflorus Pursh, Fl. Am. Sept. 2: 529 (1814); Raup in Jour. Arn. Arb, 17:
310 (1936). See Fernald in Rhod. 26: 119 (1924).

Senecio aureus L. var. discoideus Hooker, Fl. Bor.-Am. 1: 333 (1834) ; Macoun, Cat.
Can. Pl. Pt. 2: 265 (1884).

Resolution, Harper 90128 (Can.); Rae, Russell (Ca9.),

This species is aparently only occasional at Great Slave L. and on the upper
Mackenzie. I have seen only the specimens cited above, but the type of S.
aureus var. discoideus came from the “Mackenzie River,” according to Hooker.
In the Gray Herbarium is a tracing of this type, with the label, “Mackenzie River,
Fort Franklin, Dr. Richardson.”

Senecio atropurpureus (Ledeb.) Fedtsch. in Fedtsch. & Fler. Fl. Eur. Russ. 992
(1910); Porsild in Nat. Mus. Can. Bull. 101: 27 (1945).

Cineraria atropurpurea Ledeb. in Mém. Acad. St. Pétersb. 5: 574 (1814).

Liard R. between Nahanni Butte and Simpson, Crickmay 121 (Can.) ; Bolstead Cr., Canol
Rd., mi. 111 E., Wynne-Edwards 8395 (Can.). Porsild states that it is common in Mac-
millan Pass.

Senecio hyperborealis Greenm. in Ann. Mo. Bot. Gard. 3: 98 (1916); Porsild in Sar-
gentia 4: 76 (1943), in Nat. Mus. Can. Bull. 101: 32 (1945).

Senecio resedifolius sensu Hooker, Fl. Bor.-Am. 1: 333 (1834), in part, non Less., acc.
to Greenm.

Dry summit of Lone Mt., Wynne-Edwards 8528, 8529 (Can.).

Porsild has collected this species also on Mt. Charles along Bear R., in the
Mackenzie delta, and along the arctic coast. He states that it is apparently
endemic in Yukon and northwestern Mackenzie.

Senecio Kjellmanii A. E. Porsild in Rhod. 41: 299 (1939), Porsild in Sargentia 4: 76
(1943), in Nat. Mus. Can. Bull. 101: 27 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., alpine slopes, alt. 5000'-6000', Wynne-Edwards 8394
(Can.).

This species was based upon Cineraria frigida Richardson f. tomentosa Kjellm.,
Vega Exp. Vetensk, Iakttag 2: 13, t. 7. (1883), the type of which came from St.
Lawrence Island. Porsild records several Alaskan specimens, and a number
from the Richardson Mts. west of the Mackenzie delta.

Senecio Lindstroemii (Ostenf.) A. E. Porsild in Nat. Mus. Can, Bull. 101: 27 (1945).
Noted by Porsild on the Little Keele R., along the Canol Rd. near mi. 60 E.

Senecio resedifolius Less. in Linnaea 6: 243 (1831); Porsild in Sargentia 4: 77 (1943),
in Nat. Mus. Can. Bull. 101: 28 (1945).

Plains of Abraham, Canol Rd., mi. 82 E., Porsild 11825 (Can.) ; Sekwi R., Canol Rd., mi.
174 E., Porsild 11882 (Can.); Bolstead Cr., Canol Rd., mi. 111 E., high rocky ridge, alt.
5000'-6000', Wynne-Edwards 8396 (Can.).

Senecio congestus (R. Br.) DC. var. palustris (L.) Fernald in Rhod. 47: 256 (1945).
Cineraria congesta R. Br. in Parry’s 1** Voy. App. 279 (1824), in part.
Cineraria palustris L. Sp. Pl. ed. 2: 1243 (1763).
Senecio palustris (L.) Hooker, Fl. Bor.-Am. 1: 334 (1834) ; Macoun, Cat. Can. Pl. Pt.
2: 263 (1884) ; Raup in Jour. Arn. Arb. 17: 310 (1936).
Little Buffalo R., Russell 40 (G); northwest shore of Great Slave L., Bedford (Can.) ;
Mackenzie R., Kennicott (N).

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 255

Common in lowland sloughs, wet meadows, and mud bars along the main
streams.

Cirsium Drummondii T. & G. Fl. N. Am. 2: 459 (1842); Raup in Jour. Arn. Arb. 17:
311 (1936).

Cnicus Drummondii Gray; Macoun, Cat. Can. Pl. Pt. 2: 270 (1884).

Clewi R., Russell 49 (G).

This species is rather common in prairies of the Wood Buffalo Park, and
there is a Richardson specimen in Herb. G marked “Ft. Franklin.”

Saussurea angustifolia DC. in Ann. Mus. Paris 16: 200, ¢. 10-13 (1810), and Prodr.
6: 535 (1837); Porsild in Sargentia 4: 77 (1943), in Nat. Mus. Can. Bull. 101:
28 (1945).

Saussurea alpina DC. var. remotifolia Hooker, Fl. Bor.-Am. 1: 303 (1834), in part.

Saussurea alpina var, Ledebouri sensu Macoun, Cat. Can. Pl. Pt. 2: 272 (1884), in part,

non Gray.

Saussurea alpina sensu Raup in Jour. Arn. Arb. 17: 311 (1936), non DC.

Mackenzie R., Hardisty (G) ; Sekwi R., Canol Rd., mi. 174 E., Porsild 11880 (Can.).

In Fl. Bor.-Am. it is stated that S. alpina var. remotifolia ranges “North
of the Saskatchewan to the Bear Lake, in lat. 66° North.” So far as [ am aware,
no Saussurea has been collected in the central or southern parts of the Mackenzie
basin except for a single number (J. W. Tyrell 23186, Can.) from Great Slave
L., which should be assigned to S. angustifolia. The Hardisty specimen could
have come from anywhere on the Mackenzie R. Porsild has found S. angusti-
folia to be the common form in Keewatin, at Great Bear L., in the Mackenzie
delta, and in the Richardson Mts.

Saussurea angustifolia DC. var. yukonensis A. E. Porsild in Nat. Mus. Can. Bull. 101:
28 (1945).

Bolstead Cr., Canol Rd., mi. 111 E., high alpine ridge, alt. 6000’, Wynne-Edwards 8403
(Can.).

This was the type collection of var. ywkonensis. It was found with fruiting
heads July 25. Porsild cites additional specimens from the Dawson Range and
the Mayo district, in Yukon. The variety is distinguished by its very short stem
(3-8 cm.), and by its densely crowded heads.

Crepis elegans Hooker, FI. Bor.-Am. 1: 297 (1834); Porsild in Sargentia 4: 78 (1943),
in Nat. Mus. Can. Bull. 101: 33 (1945). .

Steep shale slope of high bluff on east bank of Mackenzie R. opposite Simpson, 9940;
outwash gravel plain near mouth of N. Nahanni R., W ynne-Edwards 8542 (Can.) ; Norman,
Coll.? (N).

Common at the Simpson locality; achenes maturing in late August. Porsild
has collected it also along the Bear R.

Crepis nana Richardson, Bot. App. Frankl. Journ. 746 (1823); Porsild in Sargentia
4: 78 (1943), in Nat. Mus. Can. Bull. 101: 28 (1945).
Mackenzie R., McFarlane (N); gravel fan at base of a granite mountain west of Brint-
nell L., 9476; shale rock-slides, north slope of Colonel Mt., alt. 5000’-6000’, 9402, 9650;
Bolstead Cr., Canol Rd., mi. 111 E., Wynne-Edwards 8400 (Can.).

Common in alpine slide rock; in flower during middle and late July.

Hieracium canadense Michx. Fl. Bor.-Am. 2: 86 (1803); Raup in Jour. Arn. Arb. 17:
311 (1936); Porsild in Sargentia 4: 78 (1943), in Nat. Mus. Can. Bull. 101: 34
(1945).

Hieracium umbellatum L. sensu Hooker, Fl. Bor.-Am. 1: 300 (1834) ; Macoun, Cat.
Can. Pl. Pt. 2: 275 (1884).

256 SARGENTIA [6

Southwest and north shores of Great Slave L., Howe 91997 (Can.); northwest shore of
Great Slave L., Bedford (Can.) ; east shore of Trout L., C. O. Hage (G); Simpson, Onion
(N); sandy roadway in upland woods west of Mackenzie R. near Simpson, 9892; in silty
soil near Mackenzie R. at Simpson, Nowosad (Can.) ; Mack€nzie R. between Wrigley and
Blackwater R., Crickmay 144 (Can.); Saline R., above Norman, Wynne-Edwards 8586
(Can.).

Occasional to common; collected with flowers and mature achenes Aug. 24.
Noted in Fl. Bor.-Am. as occurring “From Lake Huron ... to Fort Franklin,
on the Mackenzie River, in lat. 66°, north, abundant.” Porsild has collected it
in recent years on Mt. Charles and at Great Bear L.

Hieracium gracile Hooker, Fl. Bor.-Am. 1: 298 (1834); Porsild in Sargentia 4: 78
(1943), in Nat. Mus. Can. Bull. 101: 28 (1945).

Turfy shale slopes on north side of Colonel Mt., alt. 4300’, 9649; rocky stream bed in
upper valley of Frost Cr., alt. 4300’, 9546; east slope of Red Mt., alt. 4300-5000’, 9734;
Macmillan Pass, Canol Rd., mi. 225 E., Porsild 11267 (Can.).

Occasional in the Brintnell L. region ; found with flowers and maturing achenes
in middle and late July. Part of our material, notably no. 9734, seems to be in-
termediate between H. gracile and the very hairy H. triste Willd. It appears
to be nearest, however, to the former. See Hultén, Fl. Aleut. Isl. 340.

Taraxacum alaskanum Rydb. in Bull, Torr. Bot. Cl. 28: 512 (1901); Porsild in Sar-
gentia 4: 77 (1943), in Nat. Mus. Can. Bull. 101: 28 (1945).

Mackenzie R. between Blackwater R. and Norman, Crickmay 173 (Can.) ; gravelly
thicket at Camp Pt., Brintnell L., 9320; sandy plain near west end of Brintnell L.,, 94555
among rocks along small brook on west slope of Terrace Mt., alt. 4800’-5500’, 9558; Bol-
stead Cr., Canol Rd., mi. 111 E., alpine ridge, alt. 6000’, Wynne-Edwards 8401 (Can);

Rare at Brintnell L., and noted by Porsild as rare also in Macmillan Pass,
where it inhabits river bars. Found flowering at Brintnell L. in late June and
mid-July, and with ripe achenes July 18. Porsild has collected it also in the
Richardson Mts. and at the Mackenzie delta.

Taraxacum ceratophorum (Ledeb.) DC. Prodr. 7: 146 (1838); Raup in Jour. Arn.
Arb. 17: 311 (1936). See Fernald in Rhod. 35: 373 (1933).

Leontodon ceratophorus Ledeb. Icon. Pl. Fl. Ross. 1: 9, t. 34 (1829).

Mackenzie R., Onion (N) ?; Mackenzie R. between Blackwater R, and Norman, Crick-
may 175 (Can.); gravelly thickets near shore of Brintnell L. at Camp Pt., 9367; muskeg
thickets near west end of Brintnell L., 9444-A.

Occasional at Brintnell L.; collected with flowers and maturing achenes in the
first two weeks of July.

Taraxacum lacerum Greene in Pittonia 4: 230 (1901); Raup in Jour. Arn. Arb. 17:
312 (1936); Porsild in Sargentia 4: 77 (1943), in Nat. Mus. Can. Bull. 101: 33
(1945). See Fernald in Rhod. 35: 378 (1933).

Outwash gravel plain near mouth of North Nahanni R., Wynne-Edwards 8541 (Can.) ;
gravelly thickets on shore of Brintnell L. at Camp Pt., 9319; mossy muskeg at west end of
Brintnell L., 9441,

Occasional or common in the above habitats; achenes maturing in late June
and early July. Our no, 9441 has many of the involucral bracts ecorniculate,
resembling 7. lapponicum in this respect. It is therefore placed here provision-
ally.

Taraxacum lapponicum Kihlm. in Meddel. Soc. Faun. Fl. Fenn. 11: 108 (1884); Raup
in Jour, Arn. Arb. 17: 312 (1936). See Fernald in Rhod. 35: 383 (1933).

Gravel plain near west end of Brintnell L., 9457; muskeg thickets near west end of
Brintnell L., 9444,

Common in the muskeg habitat ; achenes maturing in the second week of July.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 257

Lactuca pulchella (Pursh) DC. Prodr. 7: 134 (1838); Macoun, Cat. Can. Pl. Pt. 2:
281 (1884); Raup in Jour. Arn. Arb. 17: 312 (1936).

Sonchus pulchellus Pursh, Fl. Am. Sept. 2: 502 (1814).

Sonchus sibiricus sensu Hooker, Fl. Bor.-Am. 1: 293 (1834), non tie

Lower Slave R., Raup 1444; open fields at Simpson, 9852; Mackenzie R. between Black-
water R. and Norman, Crickmay 178 (Can.); Bear R.,, Miss E. Taylor 89 (Can.).

Common on stream banks and in clearings throughout the Mackenzie basin
northward at least to Norman. At Simpson it was found with flowers and ma-
turing achenes Aug. 21.

Sonchus oleraceus L. Sp. Pl. 794 (1753).
Weed in a cultivated field on the west bank of the Mackenzie R. near Simpson, 9931.

Occasional ; found with flowers and mature achenes Aug. 24.

LITERATURE CITED

Appr, Ernst C. 1936. Botanical results of the Grenfell-Forbes Northern Labrador Ex-
pedition, 1931. Rhod. 38: 102-164.

Arsricut, W. D. 1933a. Gardens of the Mackenzie. Geogr. Rev. 23: 1-22.

——, 1933b. Crop growth in high latitudes. Geogr. Rev. 23: 608-620.

——. 1937. Agriculture and horticulture. In Bethune, W. D., Canada’s western northland.

Anperson, J. P. 1943-45. Flora of Alaska and adjacent parts of Canada. Pt. 1, Pteri-
dophyta and’ Gymnospermae, Iowa State Coll. Jour. Sci. 18: 137-175 (1943); Pt. 2,
Typhaceae to Poaceae, ibid. 18: 381-445 (1944); Pt. 3, Cyperaceae to Orchidaceae,
ibid. 19: 133-205 (1945).

Anperson, R. M. 1937. Faunas of Canada. In Canada Year Book for 1937, pp. 29-52.
Can. Dept. Trade and Comm.

Antevs, E. 1931. Late-Glacial correlations and ice recession in Manitoba. Geol. Surv.
Can. Mem. 168. ‘

——. 1932. Alpine zone of Mt. Washington Range. Publ. by the author.

——._ 1935. Correlations of Late Quaternary chronologies. Rept. 16th Intern. Geol. Cong.
1933.

Bett, J. M. 1929. Great Slave Lake. Roy. Soc. Can. Proc. 23: Sec. 4.

Bett, Rosert. 1900. Preliminary report of explorations about Great Slave Lake in 1899.
Geol. Surv. Can. Sum. Rept. for 1899, Pt. A, pp. 103-110.

Beruune, W. C. 1937. Canada’s western northland, its history, resources, population and
administration. Dept. Mines and Resources, Can., Lands, Parks and Forests Branch.
Bisuop, Harrow. 1930. The Austin collection from the Labrador coast. Rhod. 32: 59-62.
Biancuet, G. H. 1926. Great Slave Lake area, Northwest Territories. Dept. of Int.,

Can., N. W. T. and Yukon Branch.

Bocuer, T. W. 1933a. Phytogeographical studies of the Greenland flora. Meddel. Grgnl.
104 (3).

—._ 1933b. Studies of the vegetation of the east coast of Greenland between Scoresby
Sound and Angmagssalik. Meddel. Grgnl. 104 (4).

Britton, N. L. & P. A. Rypperc. 1901. An enumeration of the flowering plants collected
by R. S. Williams and by J. B. Tarleton (Contributions to the botany of the Yukon
territory, 4). Bull. N. Y. Bot. Gard. 2: 149-187.

Carn, Stantey A. 1944. Principles of Plant Geography. New York.

Cameron, A. E. 1917. Reconnaissance on Great Slave Lake. Geol. Surv. Can. Sum. Rept.
for 1916.

——. 1918. Explorations in the vicinity of Great Slave Lake. Geol. Surv. Can. Sum. Rept.
for 1917.

__ 1922a. Post-Glacial lakes in the Mackenzie River basin, Northwest Territories,
Canada. Jour. Geol. 30: 337-353.

——. 1922b. Hay and Buffalo Rivers, Great Slave Lake, and adjacent country Geol. Surv.
Can. Sum. Rept. for 1921, Pt. B, pp. 1-44.

——. 1936. The South Nahanni River. Can. Geogr. Jour. 13: 37-45.

— and P. S. WarrEN. 1938. Geology of South Nahanni River, N. W. T. Can. Field-
Nat. 52; 15-18.

258 SARGENTIA [6

CAMSELL, Cuar.es, 1906. Report on the Peel River and tributaries, Yukon and Mackenzie.
Geol. Surv. Can. Ann. Rept. for 1904, Pt. CC, pp. 1-49.

—. 1936. Flying through northwestern Canada. Can. Geogr. Jour. 12: 112-122.

—— and Wyarr Matcotm. 1921. The Mackenzie River basin. Geol. Surv. Can. Mem. 108.

Connor, A. J. 1937. Climate and weather of the Arctic. In Bethune, W. D., Canada’s
western northland, pp. 44-47. :

——. 1938. The climates of North America, Part 2, Canada. Handb. Klimat. 2: (J).

Dawson, G. M. 1881. Report on an exploration from Port Simpson on the Pacific coast
to Edmonton on the Saskatchewan, etc. Geol. Surv. Can. Rept. Prog. for 1879-80,

——. 1888a. Report on an exploration in the Yukon District, N. W. T. and adjacent north-
ern portion of British Columbia. Geol. Surv. Can. Ann. Rept. for 1887, Pt. B.

——. 1888b. Forest trees of the far Northwest. Garden and Forest 1: 58-59.

——. 1896, In Pike, Warburton, 1896.

Dowtinc, D. B. 1922. Geological structure of the Mackenzie River region. Geol. Surv.
Can. Sum. Rept. for 1921, Pt. B., pp. 79-90,

Farr, Epirn M. 1907. Contributions to a catalogue of the flora of the Canadian Rocky
Mountains and the Selkirk Range. Contr. Bot. Lab. Univ. Penn. 3: 1-88.

FERNALD, M.L. 1925. Persistence of plants in unglaciated areas of boreal America. (Mem.
Gray Herb. 2); Mem. Am. Acad. Arte, Sct. 18 (3).

FRANKLIN, JoHN. 1823. Narrative of a journey to the shores of the Polar Sea in the years
1819, 20, 21, and 22, with an Appendix on various subjects relating to science and
natural history. London.

——. 1828. Narrative of a second expedition to the shores of the Polar Sea in the years
1825, 1826, and 1827, including an account of the progress of a detachment to the east-
ward by John Richardson. London.

GeLtInc, P. 1934. Studies of the vascular plants of east Greenland between Franz Joseph
Fjord and Dove Bay. Meddel. Grgnl. 101 (2).

GraHAM, ANnGus. 1935. The golden grindstone; the adventures of George M. Mitchell.
London.

Grices, R. F. 1934. The problem of arctic vegetation. Jour. Wash. Acad. Sci, 24: 153-175.

Gr¢gNTVED, Jons. . 1936. Vascular plants from arctic North America collected by the Fifth
Thule Expedition, 1921-24. Rept. Fifth Thule Exped. 2: 1-93. Copenhagen.

Gtssow, W. C. 1933. Contributions to the knowledge of the flora of northern Manitoba
and the Northwest Territories, Dominion of Canada. Can. Field—Nat. 47: 116-119,

Hatitway, W. E. D. 1937. A forest classification for Canada. Dept. Mines and Re-
sources, Can. For. Serv. Bull. 89,

—— and A. W. A. Brown. 1943. The distribution of some important forest trees in Can-
ada. Ecology 24: 353-373.

Henry, J. K. 1915. Flora of. southern British Columbia and Vancouver Island, ete.
Toronto.

Hooker, W. J. 1829-1840. Flora Boreali-Americana. London.

Huttén, Eric. 1937a. Flora of the Aleutian Islands. Stockholm.

——. 1937b. Outline of the history of arctic and boreal biota during the Quaternary Pe-
riod. Stockholm.

——. 1941-1945. Flora of Alaska and Yukon. Pt. 1, Lunds Univ. Arssk. N. F. Avd. 2
Bd. 37, Nr. 1 (1941) ; Pt. 2, ibid. Bd. 38, Nr. 1 (1942) ; Pt. 3, ibid. Bd. 39, Nr. 1 (1943) ;
Pt. 4, ibid. Bd. 40, Nr. 1 (1944); Pt. 5, ibid. Bd. 41, Nr. 1 (1945).

Hume, G. S. 1922. North Nahanni and Root Rivers area, and Caribou Island, Mackenzie
River District. Geol. Surv. Can. Sum. Rept. for 1921, Pt. B.

——. 1923. Geology of the Norman oil fields and a reconnaissance of a part of Liard
River. Geol. Surv. Can. Sum. Rept. for 1922, Pt. B., pp. 47-64.

——. 1924. Mackenzie River area, District of Mackenzie, Northwest Territories. Geol.
Surv. Can. Sum. Rept. for 1923, Pt. B., pp. 1-15;

Jones, G. N. 1936. A_ botanical survey of the Olympic Peninsula, Washington. Univ.
Wash. Publ. Bot. 5: 1-286.

——. 1938. The flowering plants and ferns of Mount Rainier. Univ. Wash. Publ. Bot. 7:
1-192.

KEELE, JosePpH. 1906. The upper Stewart River region, Yukon. Geol. Surv. Can. Ann.
Rept. for 1904, Pt. C., pp. 1-23.

—. 1910. A reconnaissance across the Mackenzie Mountains on the Pelly, Ross, and
Gravel Rivers, Yukon and Northwest Territories. Geol. Surv. Can. Publ. no. 1097.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 259

Kennicott, Ropert. 1869. Biography of Robert Kennicott, by a Committee of the Chicago
Academy of Sciences. Trans. Chi. Acad. Sci. 1: Art. 6, pp. 133-226.

Kerr, Forrest A. 1934. Nahanni-Frances River district. Can. Mining Jour. 55 (Mar.).

ee E. M. 1918. Notes on sedimentation in the Mackenzie basin. Jour. Geol. 26: 341-

—. 1920. Arrival and departure of winter conditions in the Mackenzie River basin.
Geogr. Rev. 10: 388-399.

—. 1921. Mackenzie River driftwood. Geogr. Rev. 11: 50-53.

Lerroy, J. H. 1857. In Minutes of evidence, report from the Select Committee on the
Hudson’s Bay Company, Senate Sessions, 1857.

——. 1883. Diary of a magnetic survey of a portion of the Dominion of Canada, chiefly in
the North West Territories. London.

—. 1886. Report upon the depth of permanently frozen soil in the polar regions, its
geographical limits, and relations to the present poles of greatest cold. Proc. Roy.
Geogr. Soc. 8: 740-746.

McConnett, R. G. 1891. Report on an exploration in the Yukon and Mackenzie basins,
N. W. T. Geol. Sury. Can. Ann. Rept. for 1888-9, Pt. D.

——. 1906. The Macmillan River, Yukon District. Geol. Surv. Can. Sum. Rept. for
1902, Pt. A., 22-38.

MACKENzIE, ALEXANDER. 1801. Voyages from Montreal on the River St. Laurence through
the continent of North America to the Frozen and Pacific Oceans, etc. London.

Macoun, J. M. 1896. List of plants known to occur on the coast and in the interior of
the Labrador Peninsula. App. 6, Rept. on Expl. Lab. Penins. by A. P. Low. Geol.
Surv. Can. Ann. Rept.

——. 1911. Flora and fauna of the west coast of Hudson Bay. Geol. Surv. Can. Sum.
Rept. for 1909-10: 281-283.

—— and Turopore Horm. 1921. Report of the Canadian Arctic Expedition 5: Botany,
Pt. A: Vascular plants. 1-50.

Macoun, Joun. 1877. List of plants in a Report on Exploration in British Columbia by
Alfred R. C. Selwyn. Geol. Surv. Can. Rept. Prog. for 1875-1876, pp. 110-232.

——, 1883-1890. Catalogue of Canadian Plants. Montreal.

Meek, F. B. 1869. Remarks on the geology of the Mackenzie River, with figures and
descriptions of fossils in the Smithsonian Institution chiefly collected by the late Robert
Kennicott, Esq. Trans. Chi. Acad. Sci. 1: 61-114.

Ocitviz, Witt1AM. 1890. Exploratory survey of part of the Lewes, Taton-duc, Porcupine,
Bell, Trout, Peel, and Mackenzie Rivers. Dept. Int. Can. Ann. Rept. for 1889. Sess.
Papers vol. 23, no. 11.

Petitot, Emre. 1875. Géographie de l’Athabaskaw-Mackenzie et des grandes lacs du
bassin arctique. Bull. Soc. Geogr. Paris 10 (Ser. 6).

——. 1876. Monographie des Déné-dindjié. Paris.

——. 1883. On the Athabaska District of the Canadian Northwest Territory. Proc. Roy.
Geogr. Soc. and Monthly Rec. Geogr. no. 11, pp. 633-655.

_ 1885. On the Athabaska District of the Canadian North-West Territory. Can. Rec.
Sci. 1: 27-53.

——. 1887. En Route pour le Mer Glacial. Paris, 3rd ed.

—. 1888. Letter in The Great Mackenzie basin. See Schultz, John, 1888.

——, 1891. Autour du Grande Lac des Esclaves. Paris, 2nd ed.

——. 1893. Exploration de la region du Grande Lac des Ours. Paris.

Pixe, Warsurton. 1896. Through the subarctic forest. London.

Piper, C. V. 1906. Flora of the State of Washington. Contr. U. S. Nat. Herb. 11: 1-637.

Potunin, Nicuoras. 1934-1935. The vegetation of Akpatok Island. Jour. Ecology 22:
337-395 ; 23: 161-209.

——, 1940. Botany of the Canadian Eastern Arctic. Pt. 1. Pteridophyta and Spermato-
phyta. Nat. Mus. Can. Bull. 92: (Biol. Ser. 24) 1-408.

Porsitp, A. E. 1937. Flora of the Northwest Territories. In Bethune, W. D., Canada’s
western northland, pp. 130-141. Can. Dept. Mines and Res., Lands, Parks and Forests
Branch,

———. 1939. Contributions to the flora of Alaska. Rhod. 41: 141-301.

_ 1943. Materials for a flora of the continental Northwest Territories of Canada.
Sargentia 4: 1-79.

——. 1945. The alpine flora of the east slope of the Mackenzie Mountains, N. W. T.
Nat. Mus. Can. Bull. 101, Biol. Ser. 30: 1-35.

260 SARGENTIA [6

Porsitp, M. P. 1932. Alien plants and apophytes of Greenland. Meddel. Grgnl. 92: no. 1.

Potter, Davin. 1934. Plants collected in the southern region of James Bay. Rhod. 36:
274-284.

Presiz, E, A. 1908. A biological investigation of the Athabaska—Mackenzie region. U. S.
Dept. Agr., Bur. Biol. Surv., N. Amer. Fauna No. 27.

Putten, W. J. S. 1852. Notes in British Arctic Blue Book, vol. 50, Brit. Parl. Sess.
Papers of Feb. 3 to July 1, 1852.

Raup, Hucu M. 1930. The distribution and affinities of the vegetation of the Athabaska—
Great Slave Lake region. Rhod. 32: 187-208.

—. 1933. Range conditions in the Wood Buffalo Park of western Canada, with notes on
the history of the wood bison. Spec. Publ. Am. Comm. for Intern. Wild Life Protect.
stir 2:

——. 1934. Phytogeographic studies in the Peace and upper Liard River regions, Canada,
with a catalogue of the vascular plants. Contr. Arn. Arb, 6.

——. 1935. Botanical investigations in Wood Buffalo Park. Nat. Mus. Can. Bull. 74,
Biol. Ser. 20. .

——. 1936. Phytogeographic studies in the Athabaska—Great Slave Lake region, I.
Catalogue of the vascular plants. Jour. Arn. Arb. 17: 180-315.

—. 1939. Botanical exploration of the Mackenzie Mountains. Arn. Arb. Bull. Pop.
Inform. IV. 7: 69-72.

——. 1941. Botanical problems in boreal America. Bot. Rev. 7: 147-248.

——. 1942. Additions to a catalogue of the vascular plants of the Peace and upper Liard
River regions. Jour. Arn. Arb. 23: 1-28.

—. 1943. The willows of the Hudson Bay region and the Labrador Peninsula. Sar-
gentia 4: 83-127.

——. 1945a. Forests and gardens along the Alaska Highway. Geogr. Rev. 35: 22-48.

——. 1945b. Expeditions to the Alaska Military Highway 1943-1944. Arnoldia 4: 65-72.

—. 1946. Phytogeographic studies in the Athabaska—Great Slave Lake region, II,
Forests. Jour. Arn. Arb. 27: 1-85.

RicHarpson, Joun. 1825. Climate and vegetable productions of the Hudson’s Bay coun-
tries. Edinb. Phil. Jour. 12: 197-234.

—. 1851. Arctic searching expedition: a journal of a boat-voyage through Rupert’s Land
and the Arctic Sea, in search of the discovery ships under command of Sir John Franklin.
London. 2 vols.

——. 1861. The polar regions. Edinburgh.

Russett, Frank. 1898. Explorations in the far north. Univ. of Iowa.

Rypperc, P. A. 1922. Flora of the Rocky Mountains and adjacent plains, Ed. 2. New
York.

——. 1932. Flora of the prairies and plains of central North America. New York.

St. Joun, Harortp. 1937. Flora of southeastern Washington and of adjacent Idaho.
Pullman.

ScAMMAN, Epitn. 1940. A list of plants from interior Alaska. Rhod. 42: 309-343.

SEEMANN, B. 1852-1857. Flora of western Esquimaux-Land. In Botany of the voyage of
H. M. S. Herald, 1845-51, London.

Stmmons, H. G. 1906. The vascular plants in the flora of Ellesmereland. Rept. 2nd.
Norwegian Arct. Exped. in the “F ram,” 1898-1902, no. 2.

—. 1913. A survey of the phytogeography of the arctic American archipelago. Lunds
Univ. Arssk, N. F. Afd. 2, 9 (19).

Snyper, Harry. 1937. Exploring upper Nahanni River and Snyder Mountains in 1937.
Can. Geogr. Jour. 15: 169-202.

Soper, J. Dewey. 1941. History, range, and home life of the northern bison. Ecol.
Monog. 11: 347-412.

STANDLEY, P. C. 1921. Flora of Glacier National Park, Montana. Contr. U. S. Nat.
Herb. 22: 235-438.

Sressins, G. L. 1942. The genetic approach to problems of rare and endemic species.
Madrojfio 6: 241-258.

StOrMer, Per. 1933. Plants collected by Frits Johansen in 1929, at Hudson Bay Railway
and Port Churchill in arctic Canada. Nyt. Magaz. for Naturvid. 73: 259-272.

Tyrrett, J. B. 1896. Report on the country between Athabasca Lake and Churchill River,
etc. Geol. Surv. Can. Ann. Rept. for 1894. Vol. 8: Pt. D.

1947] RAUP, THE BOTANY OF SOUTHWESTERN MACKENZIE 261

——. 1897. Report on the Doobaunt, Kazan and Ferguson Rivers, and the north-west
coast of Hudson Bay, and on two overland routes from Hudson Bay to Lake Winnipeg.
Geol. Surv. Can. Ann. Rept. for 1895. Vol. 9: Pt. F.

Tyrre.t, J. W. 1902. Report on an exploratory survey between Great Slave Lake and
Hudson Bay, Districts of Mackenzie and Keewatin. Dept. Int. Can. Ann. Rept. for
1900-1901: 98-131.

Victortn, Fr. Marie-. 1935. Flore Laurentienne.

Wetmore, R. H. 1923. Plants of Labrador. Rhod. 25: 4-12.

Wuirtaker, E. J. 1922. Mackenzie River district between Great Slave Lake and Simpson.
Geol. Surv. Sum. Rept. for 1921, Pt. B., pp. 45-55.

——. 1923, Mackenzie River district between Providence and Simpson, N. W. T. Geol.
Surv. Can. Sum. Rept. for 1922, Pt. B., pp. 88-100.

Witams, M. Y. 1922. Exploration east of Mackenzie River between Simpson and Wrig-
ley. Geol. Surv. Can. Sum. Rept. for 1921, Pt. B., pp. 56-66.

——. 1923. Reconnaissance across northeastern British Columbia and the geology of the
northern extension of the Franklin Mountains, N. W. T. Geol. Surv. Can. Sum. Rept.,
Pt. B., pp. 65-87.

Woopwortn, R. H. 1927. Interesting plants of northern Labrador. Rhod. 29: 54-57.

EXPLANATION OF PLATES

Prate I

Air view of Brintnell Lake and vicinity, looking south, with South Nahanni River in
valley at left. (Reproduced through the courtesy of the Department of National Defense,
Ottawa, Canada.)

Pirate II

(Upper) Aerial photo of mountains along upper South Nahanni River. Note ancient
erosion surface in foreground.

(Lower) Air view of Simpson and the Mackenzie River, looking west. Note abandoned
channels on the island flood plain, and semi-open grasslands on the upland west of the river.
(Reproduced through the courtesy of the Department of National Defense, Ottawa, Canada.)

Prate III

(Upper) View of western end of Brintnell Lake; Red Mt. at right and Mt. Harrison
Smith at left. Frost Creek enters at the right. Note contrast between shale and granite
slopes.

(Lower) Panorama from summit of Red Mt., alt. about 5900 feet. The views extend
from northwest through northeast, and to southeast. The valley in the central part is the
upper valley of Frost Creek, and the mountain immediately back of it is Terrace Mt. The
distant valley at the right is that of the South Nahanni River. Note the expanse of sub-
alpine scrub on the till terrace just east of Frost Creek.

Piate IV

(Upper) View of Brintnell Lake from near summit of Red Mt., showing truncated hog-
backs and fans on the north slope of Colonel Mt. Note shoulders of old till just above
timber line at the right of the photo (see Plate V).

(Lower) Glacier and valley west and southwest of Brintnell Lake. View from south
slope of Red Mt.

PLATE V
Exposure of cemented till on north slope of Colonel Mt., alt. about 3800 feet.

Priate VI

(Upper) Bank of unconsolidated till in upper valley of Frost Creek, alt. about 4400 feet.
(Lower) Stone rings in upper valley of Frost Creek, alt. about 4500 feet.

262 SARGENTIA [6

PLATE VII

(Left) Forest of white spruce (Picea glauca var. albertiana) on fan at mouth of Frost
Creek (Camp Point).

(Right) Forest of white spruce and birch (Betula papyrifera vars. commutata and
humilis) in old burn on lower south slope of Red Mt., back of Camp Point.

Pirate VIII

(Left) Open forest of white spruce (Picea glauca var. Porsildii) near timber line on
south-facing slope of Red Mt., alt. about 4000 feet. The predominant grass in the fore-
ground is Festuca altaica.

(Right) Open forest of black spruce (Picea mariana) on north slope of Colonel Mt.,
alt. about 3200 feet.

Piate IX

Partially stabilized slide rock on north slope of Colonel Mt. View from an altitude of
about 4800 feet. Note turf-banked terraces and stone stripes.

Pirate X

(Upper) View northwest across upper valley of Frost Creek, from an altitude of about
5000 feet on the west slope of Terrace Mt. Note rounded alpine surfaces of shale mountains
in contrast to rugged granite peaks.

(Lower) Cliffs just below summit on north slope of Red Mt.

Piate XI

(Upper) Mud flat at western end of Brintnell Lake, showing association of Eriophorum
Scheuchzeri and Juncus castaneus.

(Lower) Natural prairie opening on the upland west of the Mackenzie River near
Simpson (see Plate IT).

Piate XII

(Left) Picea glauca var. Porsildii,

(Right) Picea glauca var. albertiana.

Both photographs were taken on the gravel fan at Camp Point, Brintnell Lake. Note
differences in the forms of the trees and the texture of the bark.

Piates XIII-XXXVII

Distribution maps of species as indicated. These maps represent the entire known vascular
flora of the Brintnell Lake area with the exception of 12 species which are of very limited
local range or are described as new in this paper.

Tue Botany oF SOUTHWESTERN MACKENZIE

LUvS

Tr

IA VIIN

I WvIg

SARGENTIA VI Priate II

Tue Botany oF SOUTHWESTERN MACKENZIE

SARGENTIA VI

Piate IIf

MACKENZIE

STERN

Tue Botany or SouTHW

SARGENTIA VI Pirate IV

Tue Botany oF SOUTHWESTERN MACKENZIE

SARGENTIA VI PLate V

Tue Botany oF SOUTHWESTERN MACKENZIE

SARGENTIA VI Piate VI

Ture BotaANy oF SOUTHWESTERN MACKENZIE

a
8 Be,

Tue Botrany

OF SOUTHWESTERN MACKENZIE

ADUVS

IA VILN

Pirate VIII

MACKENZIE

SOUTH WESTERN

THE BoTANY OF

SARGENTIA VI

Pirate IX

MACKENZIE

RN

OUTHWESTE

ep

BoTANY OF

THE

SARGENTIA VI PLATE X

Tue Botany oF SOUTHWESTERN MACKENZIE

SARGENTIA VI ; Pirate XI

Tue Botany or SOUTHWESTERN MACKENZIE

BoTANY OF

SOUTHWESTERN MACKENZIE

ADUVS

TA VILN

[IX dLVIg

SARGENTIA VI PLATE XIII

Equisetum arvense

77 7 oe

hl
xh
Jar PLY,

atile Equisetum scirpoides

PLATE XIV

SARGENTIA VI

+ +—4
Lycopodium Selago

hyon

Lycopodiu

i

m alpinum

Picea mariana

Abies lasiocarpa

alaskensis

SARGENTIA VI

PLATE XV

ae |

Festuca brachyphylla

7 7 7 Ao
/ f | |
NW } Safa
J 3
\) LA / / “Na oe
S / 3 i =
/ ‘iad a 9 we

Poa pratensis s.l.

< cy ee.
ia Guuh Y 7
J Ct RNa ae

—

Teh
kK?

‘=~

Poa nemoralis

Poa glauca

Poa Buckleyana

Agropyron trachycaulum var. novae-angliae

PLATE XVI

SARGENTIA VI

Agropyron latiglume

*

Calamagrostis lapponica 5.1.

Calamagrostis canadensis var. robusta

Agrostis scabra

atts

Arctagrostis arundinacea

PLATE XVII

SARGENTIA VI y
‘€ a FA Bim

\e Lar of B ib
Be; q 1, Sah gar
ERS (DP oe

~~ Oe,

“ee
QJ
if s\

4
sao , «2 :
Nao ., gs
Geo 4 :
N cA ; ’ ag \

1a

ere

ra

Merochloe alpina

Eriophoraish Scheuch

\ Sapa
79 he 7

Carex diandra

—

Carex gynocrates Carez leptalea

PLATE XVIII

SARGENTIA VI

ie

Carex scirpoidea

Carex aurea
a aie

Carex nesophila Carex podocarpa

te ) 7

Carex atrata ssp.
atrosquama

Carex albo-nigra

SARGENTIA VI PLATE XIX

nase DY as ; = ema ee a et (4

Carex aquatilis, incl. var. stans Carex microglochin

Carex physocarpa

: Z F: tot ua Sei ae FZ ’
J 2S em’ 3

ath Wwe 4

ie | heey) KX
es / y pd iy ¢/} Sy ~ <>
| / Y , 1a . i ~
we! HS by
: aN t

Ae
<< *

i “Lae, F my c 4
Ln NES eA OL

Luzula arcuata

Luzula spicata Luzula confusa

PLAT 3X

SARGENTIA VI

: A ae!
. de y wes ‘ KA ~ Se Eabas
sR ea

Smilacina stellata

Corallorrhiza trifida

ay aie \ \\

f FT

Populus tremuloides

SARGENTIA VI

PLATE XXI

as ay

Populus Tacamahacca

Salix retic

ulata

Salix glau

a

n BUS >
ANS

va
Ke

ca s.l.

Salix Barclayt

Salix Richardsonit

Salix alaxensis s.l.

rae

—_—

5 f ~
" J 4 1s ¢ 4
S pie yan
" —- —-
Bg a J

Salix Bebbiana s.l.

i

ar

Y/Y \

Salix Scouleriana s.l.

Salix pulchra s.l.

PLATE XXII

SARGENTIA VI

—

Betula papyrifera var. humilis

se
Betula occidentalis

‘ A f f It
y, f /

Alnus tenutfolia

rey
“ee

SARGENTIA VI

PLATE XXIII

a a” ya

aS

p
C/ ES

ML AS

Arenaria verna var. pubescens

Arenaria Rossii

= ais 9 UY

Pp

Arenaria humifusa

PLATE XXIV SARGENTIA VI
: WAL
\ } >~
eee
EC yy
Gane we
XN oe RICK
oe * Wi), if
MASE?
/ % i
~\ e , OE i J
y
ies «sae 5s ae ow ye. ee Pee HA
sh [Lo a ve) 43 F ‘ er? :

Silene acaulis var. exscapa

: Be
Lychnis apetala Actaea rubra
yuh

S/d
fe Kn SP r

> la
NR

/

Aconitum delphi

nefolium

Ps — - ee,
Anemone parviflora

aS

Anemone narcissiflora

SARGENTIA VI

AI yf
iy nna A le

nae.
—+9 Y
| \

Ranunculus hyperboreus

Sy

Ve Z a |

\ pFte YA a,
ede

~i—

AN

Ranunculus Eschscholtzit

LY RAE RS

‘Ranunculus lapponicus

hapa

KR
Corydalis pauciflora

. PAS
Draba longipes

WY

Draba lanceolata

a Hy rx <ha
Ly ew ems OY a GAD

Draba cinerea

PLATE XXVI

SARGENTLA VI

f Weep

% | “<< rt Js 4 a Ske tA) te 3

Bagel | TS ON io
OR Se Rt
tfolia

Cardamine bellid

Tht Far

Sedum roseum var,

Saxifraga Ly

alliz

integrifolium
rE’ Y /

wong. RY ac SoA
J / Z \ K ie i.
/Tir)g eo S75,
~ me / AL 7 P
fe te’ bd
Jy fy oe
ee cte ew! NetL

Saxifraga nivalis

am) ary | Ye? =
= — . lH \S

Saxifraga punctata

Saxrtfraga radiata

SARGENTIA VI

PLATE XXVII

+4
Tt 3,

Sazifraga aizoides

Mat el Nes
Us SIS

JO
NEL

Ribes hudsonianum

PLATE XXVIII SARGENTIA VI

Ae)

ef of Sen

fy B | i , i. \ a 7 \ 4 a ; % ™" %
KA 7 van f ee ) / / 4 ACA gt lf E/ b "
Y Le LR UA Cac Sefer - | NEP EN aA Se Fle es
a bea iy "7 = : nad oa Me? r\

Amelanchier florida Rubus strigosus

é! i
Potentilla diversifolia
var. glaucophylla

TT SSSSt oe

Potentilla emarginata Potentilla nivea s.1.

SARGENTIA VI

PLATE XXIX

~~

Potentilla uniflora

1 q f \ r 5 i -
+ { ' ' is a \ ° €
i | t AN -_ ‘
Tags eel ae) Bs * = yy
: vat + Aas i
oN Ts aot

e
Geum macrophyllum var. perincisum

or) er:
Rosa acicularis s.l.

Astragalus frigidus

Lupinus arcticus : ,
P var. littoralis

7 Ty Ie

PUATE- XXX

SARGENTIA VI

Oxytropis foliolosa

Oxytropis hyperborea

Oxytropis ixode

ae

To

Viola epipsila

Shepherdia canadensis

SARGENTIA VI

PLATE XXXI

Epilobium angustifolium

ss
° |
afk ct /
1 Fifa
‘ 4~] /
n i
4, | o~\
a a
zi
et |
i ee Led

x : . Net. )
= = Pr

— as |

aoe

Epilobium lactiflorum

le as |

Hippuris montana

4 MY eS J
man’s a At SOAS ok
— a

Cornus canadensis

Cornus canadensis f. purpurascens

PLATE XXXII

SARGENTIA VI

Moneses uniflora

~

Pyrola virens

Cassiope tetragona s.l.

i~ va Ate }
ig |

t

Andromeda Polifolia

. { \t$-~~ Vy atl Ay “4
MAY Me
RT IRA

. Vacqua Se
bey \

SARGENTIA VI PLATE XXXIII

hs peg ag <8 LY ak \ S iy
Arctostaphylos rubra

r P2 / /
7 /

A: ean ge’
" ve hse ‘he AP
e 1 —19 a, et (i vy
—_— aed n
Vaccinium microcarpum Gentiana propinqua

yy ay, 7 J mar f

Gentiana arctophila Gentiana glauca

PLATE AAXAILV

SARGENTIA VI

WR
Myosotis alpestris

» Y

<= 7

nS “4

Pedicularis lanata

~FES

Pinguicula villosa

SARGENTIA VI

PLATE XXXV

x, es

s +
Valeriana septentrionalis

_—

Campanula lasiocarpa

Ca hy
~a Ny

Erigeron eriocephalus

Antennaria isolepis

PLATE XXXVI

SARGENTIA VI

Thi BK

LS
Antennaria monocephala

/
/
*,

Antennaria subviscosa

eR,
Artemisia arctica

= 7

Petasites frigidus

j ry ae ‘
Arnica alpina

ssp. tomentosa Arnica mollis

e

Senecio triangularis

Senecio pauperculus

Crepis nana

SARGENTILA VI PLATE XXXVII

las pay .<
pom wid \J

Taraxacum lacerum

INDEX

Synonyms are printed in italics, new names in bold-face type.

Abies balsamea, 103
lasiocarpa, 46, 48, 71, 81, 104
mariana, 104
subalpina, 104
Acer negundo, 38
Achillea borealis, 245
Millefolium, 38, 245
Millefolium, 245
multiflora, 245
sibirica, 38, 245
Aconitum delphinifolium, 42, 45, 48, 52, 54,
70, 73, 78, 83, 176
Acrostichum ilvense, 97
Actaea rubra, 41, 64, 80-82, 175
spicata rubra, 175
Adoxa Moschatellina, 236
Agastache Foeniculum, 229
Agropyron dasystachyon, 77, 117
latiglume, 45, 52, 59, 67, 73, 83, 116, 117
pilosiglume, 37, 116
novae-angliae, 116
subsecundum, 116
trachycaulum, 116, 117
Langsdorfi, 58
novae-angliae, 42, 45, 58, 66, 81, 116
typicum, 37, 38, 115
unilaterale, 58, 66, 80, 81, 116
unilaterale, 116
violaceum latiglume, 116
Agrostis latifolia, 122
scabra, 36, 42, 48, 58, 64, 80, 122
Tuckermani, 122
Aira cespitosa, 118
spicata, 118
Allium Schoenoprasum, 144
Schoenoprasum sibiricum, 38, 144
sibiricum, 144
Allosurus sitchensis, 99
Alnus alnobetula, 164
crispa, 41-43, 45-47, 58, 64, 80, 93, 164, 202
incana, 164
incana, 164
rugosa, 93, 164
tenuifolia, 35, 38, 45, 47, 58, 66, 81, 93, 164
viridis, 164
Alopecurus aequalis, 123
alpinus, 123
aristulatus, 123
geniculatus aristulatus, 123
Alsine media, 170
Alyssum arcticum, 184
Amaranthus retroflexus, 39, 168

263

Amelanchier alnifolia, 195
canadensis oblongtfolia, 195
florida, 42, 66, 81, 195
humilis, 196
humilis, 66
ovalis, 195
sanguinea, 196
Andromeda calyculata, 221
Polifolia, 46, 47, 68, 85, 221
tetragona, 220
Androsace Chamaejasme arctica, 225
septentrionalis, 39, 224
Anemone canadensis, 39, 177
dichotoma, 177
ludoviciana, 178
multifida, 177
multifida hudsoniana, 39, 177
narcissiflora, 48, 52, 70, 83, 178
interior, 178
villosissima, 178
villosissima, 178
parviflora, 45, 47, 52, 57, 58, 68, 73, 84, 177
patens, 178
multifida, 178
Nuttalliana, 178
pennsylvanica, 177
Richardsonii, 42, 48, 52, 68, 73, 78, 79, 83,
84, 177
Antennaria campestris, 244
_carpathica pulcherrima, 244
densifolia, 243
Ekmaniana, 243
isolepis, 42, 48, 49, 59, 69, 84, 243
media, 53, 71, 81, 244
monocephala, 47, 52, 53, 70, 73, 78, 83, 243
nitida, 36, 244
oxyphylla, 244
pulcherrima, 244
rosea, 244
sp., 42, 58, 63, 244
subviscosa, 65, 84, 244
Apocynum androsaemifolium, 227
sibiricum, 38, 227
Aquilegia brevistyla, 42, 66, 77, 81, 175
Arabis divaricarpa, 39, 189
Drummondii, 42, 48, 49, 65, 81, 189
hirsuta, 189
hirsuta pycnocarpa, 36, 39, 189
pycnocarpa, 189
Holboellii, 42, 45, 65, 81, 189
retrofracta, 189

264

Arabis lyrata, 188
glabra, 188
kamchatica, 48, 54, 56, 70, 73, 80, 188
kamchatica, 188
pycnocarpa, 189
retrofracta, 189
Aragallus hudsonicus, 207
viscidulus, 208
Aralia nudicaulis, 215
Arbutus alpina, 222
Uva-ursi, 221
Arctagrostis arundinacea, 45-47, 51, 58, 70,
73, 78, 100, 121, 122
latifolia, 56, 57, 122, 123
arundinacea, 122
poaeoides, 123
Arctophila fulva, 117
Arctostaphylos alpina, 222
alpina, 222
rubra, 41, 45-47, 53, 58, 69, 73, 82, 83, 222
Uva-ursi, 36, 42, 65, 85, 221
adenotricha, 65, 81, 221
coactilis, 65
Arctous alpina ruber, 222
Arenaria arctica, 56, 172
capillaris, 171
dawsonensis, 59, 65, 81, 171
hirta pubescens, 171
humifusa, 56, 57, 68, 78, 83, 84, 172
lateriflora, 42, 47, 64, 85, 170
macrocarpa, 172
Michauxiti, 171
nardifolia, 171
propinqua, 171
Rossii, 51, 54, 56, 68, 84, 171
rubra, 172
marina, 172
sajanensis, 52, 54, 56, 57, 69, 83, 84, 172
stricta, 171
uliginosa, 172
verna hirta, 171
pubescens, 42, 52, 54, 56, 67, 73, 83, 171
Armeria maritima purpurea, 225
vulgaris arctica, 225
Arnica alpina, 41, 48, 53, 63, 73, 90, 249, 251
252
angustifolia, 249
attenuata, 249
genuina, 249
tomentosa, 51, 54, 71, 81, 250
angustifolia, 249
attenuata, 249
Chamissonis foliosa, 252
genuina, 252
chionopappa, 252
cordifolia genuina, 252
frigida, 250
Griscomi, 251
Lessingii, 52, 54, 56, 70, 78, 250, 252
lonchophylla genuina, 252

,

SARGENTIA

[6

Arnica Louiseana, 250, 251
frigida, 250
Griscomi, 251
mollis, 51, 54, 71, 81, 252
obtusifolia, 250
rhizomata, 252
Snyderi, 53, 54, 56, 63, 73, 250
tomentosa, 250
Artemisia arctica, 42, 48, 52, 53, 70, 73, 83,
247
biennis, 37, 39, 246
canadensis, 246
desertorum, 246
Douglasiana, 247
frigida, 246
Gormani, 247
Tilesii, 37, 45, 58, 70, 83, 247
elatior, 248
unalaschcensis, 248
vulgaris, 247
vulgaris heterophylla, 247
Arundo canadensis, 121
Langsdorfi, 121
lapponica, 120
neglecta, 122
Aspidium fragrans, 98
Asplenium viride, 98
Aster alpinus Vierhapperi, 239
angustus, 38, 241
ciliolatus, 240
ericoides, 240
falcatus, 37, 38, 240
foliaceus, 240
Franklinianus, 241
junceus, 37-39, 241
laxifolius, 241
Lindleyanus, 36, 240
ciliolatus, 240
comatus, 240
meritus, 239, 240
montanus, 239
paniculatus, 240
praecox, 240
ptarmicoides, 241
ramulosus, 240
Richardsonti, 239
sibiricus, 37, 47, 54, 56, 59, 70, 73, 83, 85,
239
meritus, 240
spathulatus, 241
Astragalus aboriginum, 38, 206
adsurgens, 205
alpinus, 38, 56, 67, 73, 84, 205
Brunetianus, 205
campestris, 208
eucosmus, 206
frigidus americanus, 36, 205
littoralis, 46, 48, 51, 52, 70, 73, 78, 205
goniatus, 205
hypoglottis, 205

1947]

Astragalus linearis, 206
multiflorus, 206
pygmaeus, 207
striatus, 205
tenellus, 38, 206
vaginatus, 206
yukonis, 37, 38, 206
Atelophragma lineare, 206
Avena mollis, 118
salea lapponica, 219
procumbens, 220
Barbarea orthoceras, 187
praecox, 187
vulgaris stricta, 187
Beckmannia erucaeformis, 124
Syzigachne, 37, 38, 124
Betula alba occidentalis commutata, 162
papyrifera humilis, 162
crispa, 164
fontinalis, 163
glandulosa, 42, 43, 46-48, 51-53, 58, 64, a5;
80, 81, 93, 104, 163, 164
microphylla, 163
nana, 164
occidentalis, 42, 66, 82, 84, 163
papyracea, 162
papyrifera, 162
papyrifera commutata, 41, 42, 45, 65, 81,
162
humilis, 35, 41, 45, 46, 70, 77, 81, 162
humilis, 163
neoalaskana, 162
occidentalis, 162
pumila glandulifera, 163
resinifera, 162, 163
sibirica, 80
Blitum capitatum, 167
Boschniakia glabra, 232
rossica, 35, 46, 47, 70, 81, 82, 232
Botrychium Lunaria, 96
minganense, 96
minganense, 96
multifidum, 36, 96
virginianum europaeum, 96
Brassica Kaber pinnatifida, 39, 184
Braya Henryae, 185
humilis, 38, 56, 57, 185
purpurascens, 185
Richardsonii, 185
Bromus ciliatus, 109
Pumpellianus, 36, 109
arcticus, 109
purgans vulgaris, 109
Bulbocodium serotinum, 144
Calamagrostis canadensis, 121
Langsdorfi, 58 68, 83-85, 121
Langsdorfi, 121
robusta, 45, 47, 48, 58, 66, 85, 121, 137
scabra, 121
coarctata, 122

INDEX

265

Calamagrostis hyperborea, 120
inexpansa, 121
inexpansa brevior, 38, 121
lapponica, 52, 68, 83, 90, 120
brevipilis, 120
nearctica, 120
neglecta, 36, 37, 58, 64, 85, 120, 122, 137
purpurascens, 42, 52, 56, 57, 67, 73, 84, 119,
120
stricta brevior, 121
Calla palustris, 139
Callitriche autumnalis, 211
hermaphroditica, 211
palustris, 211
verna, 211
Caltha palustris, 181
arctica, 181
Calypso borealis, 148
bulbosa, 35, 148
Campanula aurita, 237
lasiocarpa, 45, 52-54, 59, 70, 73, 81, 82, 237
linifolia, 237
rotundifolia, 237
Capsella Bursa-pastoris, 39, 184
Cardamine bellidifolia, 45, 52, 56, 67, 73, 84,
187
hirsuta, 187
nudicaulis, 188
pennsylvanica, 187
pratensis, 47, 67, 83, 187
angustifolia, 187
palustris, 187
Carex aenea, 39, 130
albo-nigra, 51, 54, 71, 81, 137
alpina, 136
ampulacea, 139
aquatilis, 47, 58, 63, 73, 85, 100, 137
stans, 138
atherodes, 138
atrata, 42, 45, 136
atrosquama, 48, 51, 71, 73, 81, 136
atratiformis, 137
atrofusca, 56, 57, 135
atrosquama, 136
aurea, 36, 47, 65, 81, 134
Bebbii, 36, 120
bipartita, 52, 67, 84, 128
bonanzensis, 35, 128
Buxbaumii, 137
canescens, 64, 85, 128
subloliacea, 129
subloliacea, 129
capillaris, 47, 67, 73, 84, 134
capitata, 127
chordorrhiza, 127
concinna, 47, 66, 80, 81, 133
Crawfordii, 130
curta, 128, 129
deflexa, 51, 69, 73, 81, 132
diandra, 47, 66, 85, 127
dioica, 129

\

266

Carex disperma, 128

eburnea, 133
festiva, 129
festivella, 130
Garberi, 64
bifaria, 81, 133
glacialis, 133
gynocrates, 47, 65, 81, 129
Hasset, 133
Hepburnii, 127
incurva, 127
Lachenalii, 128
lanuginosa, 135
lasiocarpa, 135
lasiocarpa, 135
americana, 135
lenticularis, 137
leptalea, 36, 47, 64, 80, 130
limosa, 135 .
irrigua, 135
loliacea, 47, 85, 70, 128
Macloviana, 129
pachystachya, 129
magellanica, 135
media, 35, 47, 66, 82, 84, 136
membranacea, 47, 56, 57, 68, 78, 138
membranopacta, 138
microglochin, 58, 69, 83-85, 138
misandra, 56, 135
nardina Hepburnii, 52, 54, 56, 57, 71, 84,
127
nesophila, 51, 54, 70, 76, 136
norvegica inferalpina, 136
obtusata, 132
Oederi pumila, 134
viridula, 134
oligosperma, 139
paupercula, 135
paupercula irrigua, 135
petricosa, 135
phaeocephala, 48, 52, 54, 71, 80, 129
bhaeostachya, 134
physocarpa, 47, 70, 77, 81, 82, 138
platylepis, 129
podocarpa, 48, 51, 52, 54, 70, 78, 136
praticola, 39, 130
pyrenaica, 48, 52, 71, 73, 80, 127
pyriformis, 134
rariflora, 135
retrorsa, 139
Richardsonii, 133
Rossii, 132
rostrata, 36, 139
rupestris, 57, 133
saltuensis, 134
saxatilis, 138
scirpoidea, 46-48, 52, 53, 57, 66, 73, 84, 132
scoparia, 130
siccata, 36, 127, 130
simpliciuscula, 127
Spaniocarpa, 132

SARGENTIA

Carex Soperi, 48, 63, 129
stans, 138
supina, 132
supina spaniocarpa, 132
tenuiflora, 128
tonsa, 132
trichocarpa, 138
aristata, 138
umbellata, 132
tonsa, 132
vaginata, 46-48, 64, 85, 134
Vahlii, 136
inferalpina, 136
viridula, 134
Cassandra calyculata, 221
Cassiope saximontana, 221

[6

tetragona, 46-48, 52-54, 56, 67, 73, 83, 84,

188, 220
saximontana, 84
Castilleja elegans, 231
pallida caudata, 230
elegans, 231
Raupii, 35, 231
ursina, 231
Cerastium arvense, 170
Beeringianum, 45, 68, 83, 170
grandiflorum, 170
Cerasus demissa, 204
serotina, 204
virginiana, 204
Cetraria, 42, 46, 53
Chamaedaphne calyculata, 221
Chamaepericlymenum canadense purpura-
scens, 216
Cheiranthus Pallasti, 186
Chenopodium album, 38, 167, 168
capitatum, 39, 167
leptophyllum, 168
Chrysanthemum integrifolium, 246
Chrysocoma graminifolia, 239
Chrysosplenium alterniflorum, 193
tetrandrum, 47, 67, 83-85, 193
Cicuta Douglasii, 215
mackenzieana, 215
maculata, 215
occidentalis, 215
virosa, 215, 216
Cineraria atropurpurea, 254
congesta, 254
frigida tomentosa, 254
palustris, 254
Cinna latifolia, 37, 122
Cirsium Drummondii, 255
Cladonia, 42, 46, 53
Claytonia arctica megarrhiza, 168
megarrhiza, 168
Cnicus Drummondii, 255
Coeloglossum viride bracteatum, 146
Collomia linearis, 39, 227
Colpodium arundinaceum, 122
latifolium, 122

1947]

Comandra livida, 165
Comarum palustre, 199
Convallaria stellata, 144
trifolia, 144
Corallorrhiza trifida, 41, 46, 47, 64, 85, 148
Corispermum hyssopifolium, 37, 38, 168
Cornus alba, 216
canadensis, 35, 41, 42, 64, 80, 216
purpurascens, 48, 69, 73, 84, 216
stolonifera, 35, 64, 77, 81, 94, 216
Baileyi, 217
Corydalis aurea, 38, 182
glauca, 182
pauciflora, 51, 52, 70, 83, 182
sempervirens, 77, 182
Crepis elegans, 38, 255
nana, 51, 54, 68, 73, 84, 255
Cryptogramma acrostichoides, 99
acrostichoides, 98
crispa, 99
acrostichoides, 98, 99
acrostichoides, 98
sitchensis, 70, 80, 99
Cypripedium acaule, 146
bulbosum, 148
guttatum, 146
humile, 146
parviflorum, 145
passerinum, 41, 47, 48, 58, 65, 77, 81, 145
Cystopteris fragilis, 41, 56, 63, 73, 79, 85, 97
montana, 97
Dasiphora fruticosa, 200
Delphinium Brownti, 176
glaucum, 48, 52, 70, 73, 81, 175
scopulorum, 176
glaucum, 176
Deschampsia alpina, 119
atropurpurea paramushirensis, 119
brevifolia, 119
cespitosa, 36-38, 118
glauca, 118
Descurainia Richardsonii, 185
sophioides, 185
Deyeuxia canadensis, 121
Dodecatheon pauciflorum, 225
Meadia pauciflorum, 225
Donia squarrosa, 237
Draba alpina, 182
nana, 56, 182
aurea, 183
Bellti, 182
cinerea, 54, 56, 67, 78, 79, 83, 84, 18
confusa, 183
fladnizensis, 84
heterotricha, 52, 68, 84, 182
glabella, 52, 68, 84, 183
hirta, 183
incana, 183
incerta, 183
lactea, 182
lanceolata, 54, 56, 64, 73, 85, 183

INDEX 267

Draba lonchocarpa, 80, 183
longipes, 52, 70, 73, 81, 182
lutea, 184
McCallae, 45, 52, 71, 73, 81, 183
nemorosa, 39, 184
lejocarpa, 184
nivalis, 183
elongata, 54, 56, 71, 80, 182
praealta, 54, 56, 59, 71, 73, 81, 183
Dracocephalum parviflorum, 229
Drosera rotundifolia, 190
Dryas Drummondii, 42, 54, 58, 59, 65, 73, 81,
201
integrifolia, 46, 48, 52-54, 56, 58, 67, 73, 83,
84, 201, 202
intermedia, 202
octopetala, 52-57, 67, 84, 201
integrifolia, 202
Drymocallis agrimontoides, 198
Dryopteris austriaca, 97
fragrans, 41, 67, 73, 83, 98
remotiuscula, 98
Linnaeana, 41, 64, 85, 97
Robertiana, 98
spinulosa, 98
Echinospermum patulum, 228
Redowskii occidentale, 228
Elaeagnus argentea, 213
canadensis, 213
commutata, 38, 213
Eleocharis acicularis, 126
caespitosa, 126
palustris, 37, 38, 126
typica, 126
pauciflora, 126
Elymus arenarius mollis, 117
villosus, 117
canadensis, 117
innovatus, 117
mollis, 117
Empetrum nigrum, 46-48, 52, 53, 66, 73, 85,
211
Epilobium adenocaulon, 214
anagallidifolium, 52, 67, 84, 214
angustifolium, 36, 42, 45, 52, 64, 73, 85, 213
intermedium, 213
davuricum, 47, 68, 82, 84, 214
glandulosum adenocaulon, 38, 214
lactiflorum, 45, 69, 85, 214
latifolium, 46, 47, 51, 52, 67, 73, 84, 213
palustre, 214
albiflora, 214
lineare, 214
monticola, 214
tetragonum, 214
Equisetum arvense, 37, 38, 51, 63, 73, 79, 85,
99
fluyiatile, 37, 38, 58, 64, 85, 100
limosum, 100
palustre, 45, 58, 64, 85, 99
pratense, 35, 45, 99

268

Equisetum prealtum, 36, 100
scirpoides, 35, 46, 47, 64, 85, 100
sylvaticum, 65

pauciramosum, 47, 85, 99
variegatum, 47, 51, 57, 58, 66, 73, 85, 100,
138
Erigeron acris arcuans, 242
asteroides, 241
debilis, 242
droebachensis, 241
angulosus kamtschaticus, 45, 65, 85, 241
compositus trifidus, 242
elatus, 47, 65, 81, 242
eriocephalus, 51, 52, 68, 78, 242
glabellus, 243
glabratus, 241
hyssopifolius, 243
jucundus, 51, 54, 56, 71, 81, 242
kamtschaticus, 241
philadelphicus, 37, 39, 243
pulchellum unalaschkense, 242
pulchellus, 243
purpureus, 243
trifidus, 242
unalaschkensis, 48, 52, 57, 67, 84, 242

Eriophorum angustifolium, 125
brachyantherum, 47, 65, 85, 125
capitatum, 125
Chamissonis albidum, 125
gracile, 126
opacum, 125
russeolum albidum, 125

leucothrix, 125
Scheuchzeri, 58, 67, 82, 84, 100, 125, 141
spissum, 125
vaginatum, 125

spissum, 125

Ermania, 188
borealis, 188
parryotdes, 188

Erysimum cheiranthoides, 186
inconspicuum, 186
lanceolatum, 186
Pallasii, 186
parviflorum, 186

Euphrasia disjuncta, 230
officinalis, 230
subarctica, 230

Eutoca Franklinii, 228

Eutrema Edwardsii, 56, 187

Festuca altaica, 42, 43, 46, 48, 49, 51-53, 58,

70, 73, 83, 110
arenaria, 110
brachyphylla, 42, 45, 52, 53, 59, 66, 73, 84,
109
saxtmontana, 110
ovina, 110
rubra, 110 1%
arenaria, 110
Saximontana, 77, 110

SARGENTIA 6

Fragaria canadensis, 197
glauca, 39, 59, 66, 81, 197
pauciflora, 197
vesca americana, 35, 197
virginiana, 197
glauca, 197
Fumaria pauciflora, 182
sempervirens, 182
Galium boreale, 36, 39, 224
Claytoni, 235
trifidum, 36, 235
Gentiana acuta, 225
Amarella, 225
acuta, 225
arctophila, 70, 84, 226
detonsa, 226
elegans, 226 ;
glauca, 51-53, 70, 81, 226
propinqua, 47, 53, 56, 57, 59, 69, 73, 81, 226
Raupii, 226
serrata, 226
Geocaulum lividum, 35, 41, 46, 65, 81, 165
Geranium Bicknellii, 210
Geum aleppicum strictum, 200
macrophyllum perincisum, 36, 45, 47, 58,
65, 77, 81, 200
oregonense, 200
perincisum, 200
strictum, 200
Glaux maritima, 225
Glyceria grandis, 110
maxima grandis, 110
pulchella, 110
Goodyera repens, 66
ophioides, 46, 85, 147
Grindelia perennis, 237
squarrosa, 237
Habenaria hyperborea, 36, 41, 47, 58, 64, 80,
146
obtusata, 35, 46, 47, 64, 81, 147
orbiculata, 147
viridis bracteata, 146
interjecta, 146
Hedysarum alpinum americanum, 37, 45-48,
51, 52, 58, 64, 73, 81, 209
boreale, 209
Mackenzii, 38, 52, 54, 56, 69, 73, 84, 210
Helenium autumnale, 38, 245
Helianthus annuus, 39, 245
Heracleum lanatum, 58, 65, 80, 216
Hesperis pygmaeus, 186
Heuchera hispida, 193
Richardsonii, 193
Hieracium canadense, 36, 255
gracile, 48, 52, 53, 71, 80, 256
triste, 256
umbellatum, 255
Hierochloé alpina, 51, 52, 67, 83, 124
odorata, 39, 124
Hippuris montana, 47, 71, 80, 215

1947]

Holcus alpinus, 124
odoratus, 124
Homalobus retusus, 207
Hordeum jubatum, 39, 117
Hudsonia tomentosa, 212
tomentosa intermedia, 212
Hypnum spp., 41, 42, 45, 46
Impatiens biflora, 77
fulva, 211
Noli-tangere, 211
Isoétes Braunii, 102
Juncus acutiflorus, 141
albescens, 140
alpinus, 38
insignis, 141
nodulosus, 141
rariflorus, 37, 141
arcticus, 140
arcuatus, 142
balticus, 37, 38, 47, 64, 85, 140
littoralis, 140
biglumis, 56, 57, 140
bufonius, 39, 140
castaneus, 47, 58, 67, 84, 100, 125, 141
Drummondii, 71, 80, 140
filiformis, 141
nodosus, 37, 38, 141
parviflorus, 142
polycephalus tenutfolius, 141
rariflorus, 141
Richardsonianus, 141
spicatus, 142
tenuis, 140
triglumis, 56, 57, 140
albescens, 140
Vaseyi, 36, 37, 140
Juniperus communis nana, 106
montana, 42, 48, 52, 56, 64, 73, 85, 106
horizontalis, 106
nana, 106
sabina, 106
Kalmia glauca, 220
polifolia, 93, 220
microphylla, 93
Kobresia simpliciuscula, 56, 57, 127
Lactuca pulchella, 39, 257
Lappula Redowskii occidentalis, 39, 228
Larix alaskensis, 105
laricina, 77, 105
alaskana, 47, 70, 78, 105
Lathyrus ochroleucus, 210
Ledum groenlandicum, 46, 47, 53, 64, 73, 81,
82, 219
latifolium, 219
palustre decumbens, 219
latifolium, 219
Lemna trisulca, 139
Leontodon ceratophorus, 256
Lepargyrea canadensis, 213

INDEX

269

Lepidium apetalum, 184
densiflorum, 184
ruderale, 184
Lesquerella arctica, 184
Limosella aquatica, 230
Linnaea americana, 235
borealis, 65, 85
americana, 41, 42, 48, 53, 73, 235
Linum Lewisii, 38, 211
perenne, 211
Listera borealis, 47, 66, 80, 81, 147
Lithospermum paniculatum, 228
Lloydia serotina, 48, 51, 56, 57, 83, 144
Lobelia Kalmii, 237
Loiseleuria procumbens, 220
Lomatogonium rotatum, 226
Lonicera caerulea, 236
glaucescens, 35, 236
involucrata, 236
Lophanthus anisatus, 229
Lupinus arcticus, 41, 42, 46, 52, 70, 73, 78,
204
Luzula arcuata, 51, 52, 54, 70, 73, 82, 84, 142
campestris, 142
alpina, 142
nivalis, 142
confusa, 51, 52, 54, 56, 68, 84, 142
nivalis, 56, 142
parviflora, 142
saltuensis, 77
serotina, 70
spicata, 51, 52, 54, 56, 67, 84, 142
Wahlenbergii, 48, 67, 78, 79, 142
Lychnis apetala, 51, 54, 68, 84, 173
brachycalyx, 57, 63, 173
furcata, 175
macrosperma, 173
Lycopodium alpinum, 48, 51, 52, 68, 83, 101
annotinum, 64, 85, 101
alpestre, 52, 100
pungens, 41, 101
clavatum monostachyon, 42, 65, 85, 101
complanatum, 41, 42, 48, 52, 65, 73, 85
canadense, 101
obscurum dendroideum, 101
selaginoides, 101
Selago, 51, 52, 66, 83-85, 100
Lysimachia thysiflora, 225
Maianthemum bifolia, 145
canadense, 145
canadense interius, 145
Mairainia alpina, 222
Matricaria ambigua, 246
inodora, 246
Melandrium apetalum, 173
arcticum, 173
macrospermum, 173
Melanidion borealis, 57, 188
Mentha canadensis glabrata, 36, 38, 229
glabrior, 229
Menyanthes trifoliata, 227

270

Mensziesia empetriformis, 220
glanduliflora, 220
Mertensia paniculata, 42, 45, 46, 52, 53, 66,
73, 77, 81, 228
Minuartia arctica, 172
macrocarpa, 172
rubella, 171
stricta, 171
Mitella nuda, 35, 193
Moehringia lateriflora, 170
Moneses uniflora, 41, 45, 64, 85, 217
Monotropa uniflora, 219
Muhlenbergia Richardsonis, 124
Myosotis alpestris, 70, 83
asiatica, 52, 53, 228
sylvatica alpestris, 228
Myrica Gale, 93, 162
Myriophyllum exalbescens, 215
spicatum, 215
Nardosnua corymbosa, 248
frigida, 248
palmata, 248
sagittata, 248, 249
Narthecium glutinosum, 143
Nasturtium obtusum, 186
palustre, 186
microcarpa, 186
Neslia paniculata, 39, 184
Nuphar /utewm, 175
variegatum, 175
Nymphosanthus variegatus, 175
Orchis bracteata, 146
hyperborea, 146
obtusata, 147
orbiculata, 147
rotundifolia, 77, 146
Orobanche glabra, 232
rossica, 232
Osmunda crispa, 98
Lunaria, 96
multifida, 96
virginiana, 96
Oxycoccus microcarpus, 223
Oxycoccus, 223
Oxyria digyna, 46, 48, 51, 52, 67, 73, 84, 166
Oxytropis campestris, 208
spicata, 208
foliolosa, 45, 67, 82, 84, 207
gracilis, 208
hudsonica, 69, 84, 207, 209
hyperborea, 45, 48, 51, 52, 70, 73, 84, 208
ixodes, 46, 47, 51, 52, 56, 69, 73, 84, 209
Maydelliana, 51, 52, 54, 56, 57, 68, 78, 207
podocarpa, 207
pygmaea, 51, 52, 54, 56, 70, 78, 207
retrorsa, 207
spicata, 208
splendens, 209
Richardsonti, 209
terrae-novae, 57, 208
viscida, 209

SARGENTIA [6

Oxytropis viscidula, 208, 209
Panicularia pulchella, 110
Papaver Keelei, 56, 181
microcarpum, 181
radicatum, 181
Parnassia fimbriata, 52, 71, 80, 193
Kotzebuei, 47, 52, 69, 73, 77, 81, 194
montanensis, 194
multiseta, 194
palustris, 193
palustris neogaea, 45, 47, 58, 65, 85, 193
Parrya, 188
Ermani, 188
nudicaulis, 56, 188
interior, 188
Pedicularis arctica, 232
capitata, 46-48, 51, 53, 57, 67, 73, 83, 232
euphrasioides, 231
labradorica, 42, 46, 48, 52, 53, 69, 73, 82,
83, 231
lanata, 53, 68, 83, 232
palustris Wlassoviana, 232
parviflora, 232
sudetica, 45, 47, 57, 67, 83, 232
Pellaea glabella, 98 —
Peltigera aphthosa, 41
spp., 42
Petasites frigidus, 48, 70, 83, 85, 248
palmatus, 248
Sagittatus, 39, 248, 249
vitifolius, 248
Phaca aboriginum, 206
astragalina, 205
elegans, 206
frigida americana, 205
littoralis, 205
nigrescens, 206
Phacelia Franklinii, 227
Phalaris arundinacea, 36, 125
Phegopteris Dryopteris, 97
Phleum alpinum, 124
americanum, 124
Phyllodoce empetriformis, 220
glanduliflora, 48, 51, 53, 71, 80, 220
Picea albertiana, 102
canadensis, 102
glauca, 66, 77, 81, 102, 103
albertiana, 35, 36, 41, 42, 45, 46, 48, 52,
102, 103
Porsildii, 36, 42, 45, 63, 102
mariana, 45-47, 66, 77, 81, 102, 104, 147
Pilosella Richardsonti, 185
Pinguicula villosa, 46, 47, 69, 78, 79, 233
vulgaris, 233
Pinus alba, 102
Banksiana, 36, 105, 106
contorta latifolia, 106
Murrayana, 106
divaricata, 105
lasiocarpa, 104
microcarpa, 105

1947]

Pinus Murrayana, 106
nigra, 104
Plantago eriopoda, 234
lanceolata, 234
major, 37, 234
astatica, 234
septata, 234
Platanthera hyperborea, 146
obtusata, 147
orbiculata, 147
Pleurogyne rotata, 226
Poa airoides, 111
alpigena, 111
alpina, 45, 48, 52, 56, 66, 73, 84, 111
arctica, 45, 48, 51-53, 67, 73, 82, 83, 112-
114
longiculmis, 114
Williamsii, 112
Brintnellii, 45, 57, 63, 112
Buckleyana, 42, 71, 81, 115
cenisia, 114
cenisia, 112
flexuosa, 112
fulva, 117
glauca, 38, 39, 45, 52, 56, 57, 67, 73, 78, 79,
83, 84, 115
laxa, 115
leptocoma, 114
nemoralis, 42, 64, 85, 114
nervosa, 114
Nuttalliana, 111
palustris, 36, 114
paucispicula, 56, 70, 73, 78, 114
pratensis, 36, 37, 39, 45, 47, 58, 64, 85, 111
alpigena, 111
scabrella, 115
Vahliana, 111
Polemonium acutiflorum, 48, 52, 53, 70, 73,
227
boreale, 227
Polygonum alaskanum, 167
alpinum lapathifolium, 167
amphibium, 167
aviculare, 37, 39, 166
Bistorta plumosum, 167
buxiforme, 166
Hartwrightii, 167
lapathifolium salicifolium, 37, 38, 167
natans Hartwrightii, 36, 167
Persicaria minus, 167
plumosum, 167
viviparum, 48, 52, 57, 67, 73, 84, 166
Polypodium Dryopteris, 97
fragile, 97
fragrans, 98
montanum, 97
Robertianum, 98
spinulosa, 98
virginianum, 99
Populus balsamifera, 148
Tacamahacca, 35, 36, 38, 59, 65, 80, 81, 148

INDEX

271

Populus tremuloides, 36, 65, 80, 81, 148
aurea, 148
Potamogeton alpinus tenuifolius, 108
tenuifolius, 108
borealis, 107
filiformis borealis, 107
foliosus macellus, 108
Friesii, 108
gramineus graminifolius, 108
maximus, 108
typicus, 108
pectinatus, 108
perfoliatus Richardsonii, 108, 109
praelongus, 108
Richardsonii, 108
tenuifolius, 108
vaginatus, 108
Potentilla Anserina, 39, 199
sericea, 199
arguta, 198
biflora, 56, 198
diversifolia, 71, 81
glaucophylla, 52, 198
elegans, 199
emarginata, 52, 56, 68, 78, 83, 84, 198
fruticosa, 41, 42, 47, 52, 64, 73, 85, 93, 200
glaucophylla, 198
multifida, 39, 199
nipharga, 198
nivea, 52, 56, 67, 84, 198
subquinata, 198
norvegica, 36, 39, 65, 85, 198
palustris, 199
pectinata, 199
pennsylvanica, 39, 199
tridentata, 200
uniflora, 52, 56, 57, 70, 83, 199
Primula egaliksensis, 224
farinosa, 224
Hornemanniana, 224
incana, 224
mistassinica, 77, 224
sibirica, 224
stricta, 224
Prunus demissa, 204
pennsylvanica, 204
virginiana demissa, 204
Pteris atropurpurea, 98
Puccinellia Nuttalliana, 111
Vahliana, 56, 57, 111
Pulmonaria paniculata, 228
Pulsatilla ludoviciana, 178
Pyrethrum ambiguum, 246
inodorum, 246
Pyrola asarifolia, 218
incarnata, 41, 53, 58, 64, 73, 81, 82, 218
borealis, 218
canadensis, 218
chlorantha, 218
grandiflora, 67, 78, 83, 84, 218
canadensis, 35, 41, 218
incarnata, 218

272

Pyrola minor, 52, 64, 217, 219
rotundifolia, 218
incarnata, 218
secunda, 41, 45, 46, 53, 64, 73, 85, 217
obtusata, 217
uniflora, 217
virens, 41, 65, 85, 218
Ranunculus acris, 180
aquatilis, 178
capillaceus, 178
Cymbalaria, 180
Eschscholtzii, 52, 71, 80, 179
Flammula intermedia, 179
major, 179
reptans, 179
Gmelini Purshii, 47, 64, 81, 178
terrestris, 179
hispidus, 180
hyperboreus, 47, 67, 84, 179
lapponicus, 47, 64, 85, 179
Macounii, 36, 180
multifidus, 178
nivalis, 52, 68, 78, 179
pennsylvanicus, 180
Purshti, 178
pygmaeus, 179
repens hispidus, 180
reptans, 179
sceleratus, 180
multifidus, 180
trichophyllus, 178
typicus, 178
Rhinanthus Crista-galli, 231
groenlandicus, 38, 231
Kyrollae, 231
Rhodiola integrifolia, 190
Rhododendron lapponicum, 219
Ribes glandulosum, 41, 66, 77, 81, 195
hudsonianum, 35, 41, 47, 65, 77, 81, 194,
202
lacustre, 35, 45, 64, 81, 82, 195
oxyacanthoides, 35, 45, 66, 81, 194, 195
lacustris, 195
prostratum, 195
rubrum, 195
triste, 41, 58, 64, 81, 195, 202
Rorippa islandica microcarpa, 47, 66, 85, 186
obtusa, 186
palustris glabrata, 186
Rosa acicularis, 35, 36, 41, 42, 45, 52, 58, 65,
73, 85, 93, 202, 204
Bourgeauiana, 202
cucurbiformis, 46, 63, 203
lacorum, 93, 202, 204
blanda, 204
Engelmanii, 204
mayjalis, 202
Woodsii, 37, 38, 204
Rubus acaulis, 35, 41, 47, 66, 81, 197
arcticus, 197
grandiflorus, 197

SARGENTIA [6

Rubus Chamaemorus, 47, 64, 85, 196, 197
idaeus, 196
canadensis, 196
strigosus, 196
paracaulis, 197
pubescens, 35, 196, 197
strigosus, 36, 42, 45, 66, 85, 196
Rumex arcticus, 166
digynus, 166
fueginus, 166
maritimus fueginus, 37, 38, 166
mexicanus, 39, 165
salicifolius, 165
sibiricus, 165
triangulivalvis, 165
Sagina Linnaei, 69, 83, 84, 170
nodosa, 170
Sagittaria cuneata, 109
Salicornia europaea, 168
herbacéa, 168
Salix acutifolia, 161
adenophylla, 156
alaxensis, 46, 48, 49, 51, 70, 73, 78, 158
longistylis, 45, 158
obovalifolia, 158
arbusculoides, 35, 38, 41-43, 70, 77, 81, 161
arctica, 48, 51, 52, 54, 56, 57, 66, 78, 150, .
151
arctophila, 151
argyrocarpa, 161
Barclayi, 47, 71, 80, 156
Barrattiana, 48, 51, 71, 80, 81, 155, 157
marcescens, 48, 63, 157
Bebbiana, 35, 36, 42, 45, 64, 80, 81, 93, 159
depilis, 63, 159
perrostrata, 159
brachycarpa, 151-153
antimima, 151
candida, 93, 158
caudata, 149
chlorophylla, 161
commutata, 45, 47-49, 58, 71, 80, 156
cordata, 156, 157
denudata, 157
mackensieana, 155
cordifolia, 84, 153
crassijulius, 150
desertorum, 152, 153
exigua, 93
fluviatilis, 149
fusca, 159
glauca, 38, 69, 83, 84, 151-153, 155
acutifolia, 52-54, 56, 152, 154, 155
poliophylla, 153
Aliceae, 152, 153
desertorum, 153
sericea, 153
glabrescens, 152
Muriei, 152
perstipula, 42, 43, 46-49, 51, 52, 54, 63,
73, 152, 154, 158, 164

1947]

Salix glauca poliophylla, 153
stenolepis, 155
villosa, 152
glaucops glabrescens, 152
interior, 149
interior, 38, 93
pedicellata, 37, 38, 149
lasiandra, 38, 93, 149
caudata, 149
lancifolia, 149
recomponens, 37, 149
longifolia, 149
longistylis, 158
lucida, 149
lutea, 155, 156
mackenzieana, 37, 38, 93, 155, 156
monticola, 156
myrsinites, 156
myrtillifolia, 47, 66, 80, 81, 93, 156
myrtilloides, 160
Nelsonii, 161
niphoclada, 151
novae-angliae pseudocordata, 156
Nuttallii, 160
padophylla, 156
pedicellaris, 160
pedicellaris hypoglauca, 160
pellita, 94
perrostrata, 159
petiolaris, 160
phlebophylla, 56, 150
planifolia, 45, 48, 49, 64, 82, 84, 93, 161
polaris pseudopolaris, 150
pseudomonticola, 153, 156
pseudo-myrsinites, 156
pseudopolaris, 150
pulchra, 52, 54, 70, 78, 155, 158, 160, 161
reticulata, 46-48, 51-53, 58, 67, 73, 83, 150
.Richardsonii, 46-48, 51, 67, 73, 78, 155, 157,
158
rigida, 155
rostrata, 159
Scouleriana, 160
Scouleriana, 80, 160
coetanea, 41-43, 45, 71, 160
Seemannit, 152
serissima, 93, 149
speciosa, 158
alaxensis, 158
subcoerulea, 94
Uva-ursi, 151
villosa acutifolia, 152
Satyrium viride, 146
Saussurea alpina, 255
Ledebouri, 255
remotifolia, 255
angustifolia, 255
yukonensis, 255
Saxifraga aizoides, 51, 54, 56, 67, 73, 78, 83,
84, 192
cernua, 190

INDEX

273

Saxifraga debilis, 191
flagellaris, 192
hieracifolia, 191
Hirculus, 57, 192
Lyallii, 48, 55, 71, 80, 191
nivalis, 56, 68, 83, 191
oppositifolia, 51, 52, 56, 67, 84, 193
punctata, 48, 52, 70, 83, 191
insularis, 192
radiata, 45, 70, 78, 192
rivularisy 52, 67, 84, 190
serpyllifolia, 192
sibirica, 57, 63, 90, 190, 191
pekinensis, 191
tricuspidata, 42, 43, 45, 46, 48, 52, 54, 56,
67, 73, 77, 83, 192
Schizachne purpurascens, 115
Scirpus acicularis, 126
caespitosus callosus, 126
microcarpus, 126
. palustris, 126
pauciflorus, 126
validus, 126
Scutellaria epilobiifolia, 229
galericulata, 229
Sedum roseum integrifolium, 48, 52, 70, 83,
190
Selaginella selaginoides, 47, 58, 64, 85, 101
Senecio atropurpureus, 57, 254
aureus, 253
Balsamitae, 253
borealis, 253
discoideus, 254
Balsamitae, 253
congestus palustris, 254
cymbalarioides, 253
cymbalarioides borealis, 253
eremophilus, 252
hyperborealis, 254
indecorus, 47, 65, 81, 253
Kjellmanii, 254
Lindstroemii, 254
lugens, 41, 47, 48, 51, 53, 70, 73, 81, 253
palustris, 254
pauciflorus, 254
pauperculus, 47, 65, 81, 253
resedifolius, 57, 254
resedifolius, 254
triangularis, 48, 71, 80, 252
Shepherdia canadensis, 35, 41, 42, 45, 46, 48,
52, 64, 73, 81, 213
Sibbaldia procumbens, 45, 52, 68, 73, 84, 200
Silene acaulis, 173
acaulis exscapa, 46, 48, 51, 52, 56, 67, 73,
84, 173
subacaulescens, 173
subacaulescens, 173
exscapa, 173
Menziesii, 39, 173
Sinapis arvensis pinnatifida, 184

274

Sisymbrium arabidoides, 188
brachycarpum, 185
canescens, 185
humile, 185
Richardsonii, 185
salsugineum, 185
sophioides, 185

Sisyrinchium angustifolium, 38, 39, 145

Sium suave, 37, 38, 216

Smilacina stellata, 42, 64, 81, 144
trifolia, 144 «

Solidago canadensis, 238
decumbens, 238
decumbens oreophila, 36, 238
elongata, 238
graminifolia camporum, 239
graminifolia major, 38, 239
lanceolata major, 239
lepida elongata, 37, 38, 238

fallax, 239
multiradiata, 41, 42, 45, 53, 54, 69, 73, 77,
81, 238
scopulorum, 238
oreophila, 238
procera, 238
stricta, 238
Virgaurea, 238

Sonchus oleraceus, 39, 257
pulchellus, 257
sibiricus, 257

Sparganium angustifolium, 107
eurycarpum, 107
minimum, 107
multipedunculatum, 107
simplex, 107

multipedunculatum, 107

Spartina gracilis, 124
pectinata, 124
polystachya, 124

Spergula nodosa, 170

Spergularia marina, 172
media, 172
salina, 172

Sphagnum, 47

Spiraea Beauverdiana, 195

Spiranthes gracilis, 147
Romanzoffiana, 147

Stachys Foeniculum, 229
palustris, 229
palustris pilosa, 36, 38, 39, 229
pilosa, 229
scopulorum, 229

Stellaria borealis, 58
borealis, 168
calycantha, 39, 45, 64, 85, 168
crassifolia, 169
Edwardsti, 169
laeta, 169
longifolia, 39, 169

SARGENTIA [6

Stellaria longipes, 39, 169
Edwardsii, 46, 51, 52, 54, 68, 73, 84, 169
laeta, 169
media, 39, 170
Stereocaulon, 53
Swertia rotata, 226
Symphoricarpos albus pauciflorus, 235
occidentalis, 235
racemosus pauciflorus, 235
Tanacetum huronense bifarium, 246
pauciflorum, 246
Taraxacum alaskanum, 45, 52, 59, 70, 73, 78,
256
ceratophorum, 45, 47, 69, 82, 84, 256
lacerum, 45, 47, 67, 78, 256
lapponicum, 47, 59, 69, 83, 84, 256
Thalictrum alpinum, 56, 57, 180
dioicum, 181
occidentale, 39, 181
venulosum, 181
Thellungiella salsuginea, 185
Thelypteris fragrans Hookeriana, 98
Tofieldia borealis, 143
coccinea, 143
major, 143
glutinosa, 143
occidentalis, 143
palustris, 46, 47, 51, 52, 56-58, 69, 73, 84,
143
Torularia humilis, 185
Trientalis arctica, 225
europaea arctica, 225
Triglochin maritimum, 109
palustre, 38, 109
Trillium cernuum, 145
cernuum macranthum, 145
erectum declinatum, 145
Tripolium angustum, 241
Trisetum purpurascens, 115
spicatum, 56, 67, 118
Maidenii, 42, 48, 52, 56, 73, 84, 118
molle, 118
subspicatum, 118
Maidenti, 118
molle, 118 F
Triticum trachycaulum, 115
Turritis diffusa, 185
hirsuta, 189
retrofracta, 189 .
Tussilago frigida, 248
palmata, 248
sagittata, 249
Typha latifolia, 107
Urtica gracilis, 165
Utricularia intermedia, 233
minor, 233
vulgaris, 233
americana, 233
Vaccinium album, 235
canadense, 222, 223
microcarpum, 46, 47, 68, 85, 223

1947] INDEX 21D

Vaccinium O-rycoccus, 223 Viburnum Opulus edule, 235
uliginosum, 42, 43, 46, 47, 68, 85, 222 Oxycoccus, 235
alpinum, 53, 68, 83, 223 pauciflorum, 235
oe 223 eee er Ge Vicia americana, 36, 210
iftis-Ldaea@ minus, —49, 40, 46, J, 4 ifolia, 210
73, 85, 223 angustifoha,

Vilfa arundinacea, 122
Richardsonis, 124
Viola adunca, 39, 212+

Vahlodea atropurpurea paramushirensis, 119
Valeriana septentrionalis, 47, 66, 82, 84, 236
sitchensis, 236

Veratrum Eschscholtzii, 144 blanda, 212
Veronica alpina alterniflora, 230 cucullata, 212
geminiflora, 230 epipsila, 48, 70, 81, 82, 212
unalaschcensis, 48, 51-53, 69, 82, 84, 230 nephrophylla, 212
peregrina, 230 renifolia Brainerdii, 35, 212

peregrina xalapensis, 38, 230

scutellata, 230 7itis-Idaea Vitis-Idaea, 223

xalapensis, 230 Woodsia glabella, 97
Vesicaria arctica, 184 ilvensis, 52, 54, 56, 68, 85, 97

Viburnum edule, 35, 36, 41, 42, 45, 58, 65, 80, | Zygadenus chloranthus, 143
93, 235 elegans, 48, 51, 52, 54, 59, 70, 73, 81, 143

=~

CONTRIBUTIONS FROM THE ARNOLD ARBORETUM
OF HARVARD UNIVERSITY

No. 1. The Hypodermataceae of Conifers. By Grant Dooxs DARKER.
131 pp. 27 pl. June 15, 1932. Price $3.00

No. 2. Taxonomy and Geographical Distribution of the Genus Milesia. By
JoserH Horace FauLt. 138 pp. 2 figs.,9 pl. Oct. 1, 1932. Price $3.00

No. 3. Studies in the Boraginaceae, IX. By Ivan M. JoHNsTON. 102 pp.
Dec. 15, 1932. Price $2.00

No. 4. Ligneous Plants Collected in North Queensland for the Arnold
Arboretum by S. F. Kajewskiin 1929. ByC.T.Wuite. 113pp. 9pl. April 1,
1933. Price $2.75

No. 5. Flora of Barro Colorado Island, Panama. By Paut C. STANDLEY.
178 pp. 21 pl. and map. Oct. 1, 1933. Price $3.50

No. 6. Phytogeographic Studies in the Peace and Upper Liard River Regions,
Canada. With a Catalogue of the Vascular Plants. By Hucu M. Ravp.
230 pp. 9pl.and map. Feb. 15, 1934. Price $2.50

No. 7. The Beech Bark Disease; a Nectria Disease of Fagus following Crypto-
coccus Fagi (Baer.). By JoHN Euriicu. 104 pp. 9 pl. Sept. 29, 1934.
Price $2.00

No. 8. An Enumeration of Plants Collected in Sumatra by W. N. and C. M.
Bangham. By E.D. MERRILL. 178 pp. 14 pl. Aug. 25, 1934. Price $2.50

No. 9. The Species of Tradescantia Indigenous to the United States. By
EpcGar ANDERSON and Rosert E. Woopson. 132 pp. 12 pl. Aug. 30, 1935,
Price $2.25

No. 10. The Cephalosporium Disease of Elms. By Don BAKER CREAGER.
91 pp. 16 pl. July 1937. Price $2.00

No. 11. Taxonomy and Geographical Distribution of the Genus Uredinopsis.
By JosepH Horace Fauty. 120 pp. 6 pl. Sept. 30, 1938. Price $2.00

List of other publications issued by the Arnold Arboretum
will be sent on request.