CUIILOnE

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ROY l TAYLOR
6ERIILD A. MULLI6XN

<rf- .tg^^lVsi

PART 2 of the FLORA OF THE
QUEEN CHARLOTTE ISLANDS

reports on the cytological aspects of the
vascular plants found on the Islands. It
was planned by James Calder, Roy
Taylor and Gerald Mulligan early in
1963 so that the necessary collections
could be made while work was progressing
on Part 1 , Systematics of the Vascular
Plants. The two companion volumes are
therefore unique, in that they present the
results of a concurrent systematic and
cytological study of a floristic region. In
1963, Taylor made a pilot study of
collecting methods in Western Canada,
and during the summer of 1964 he
collected some 1600 cytological speci-
mens. Taylor and Mulligan later examined
the collections and successfully counted
the chromosomes of 71 percent of the
vascular flora of the Queen Charlottes.
The chromosome numbers given in this
volume are correlated with the systematic
discussion in Part 1 and with previously
published results of cytotaxonomic and
chromosome studies in other regions.
The authors indicate that biologists
have merely scratched the surface of a
vast field of cytological information that
could be made available if cytological
surveys were made a regular part of field
forays. They express the hope that the
publication of these two volumes will
encourage other botantists to pursue
similar studies elsewhere in North
America.

Jacket, cover, title page and text
illustrations by llgvars Steins.

Digitized by tine Internet Arciiive

in 2011 witii funding from

Agriculture and Agri-Food Canada - Agriculture et Agroalimentaire Canada

http://www.archive.org/details/floraofqueenchar02cald

FLORA

OF THE

OUEEN CHARLOTTE ISLANOS

PART 2 C'^TOLOGICAL ASPECTS OF THE VASCULAR PLANTS

ROY L. TIYLOR

and 6ERIIL0 1. MULIIOAH

PLANT RESEARCH INSTITUTE
CENTRAL EXPERIMENTAL FARM
OTTAWA, ONTARIO

1968

RESEARCH BRANCH

CANADA DEPARTMENT OF AGRICULTURE

MONOGRAPH No. 4 PART 2

© Crown Copyrights reserved

Available by mail from the Queen's Printer, Ottawa,

and at the following Canadian Government bookshops:

HALIFAX

1735 Barrington Street

MONTREAL

-■Eterna-Vie Building, 1182 St. Catherine Street West

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Daly Building, corner Mackenzie and Rideau Streets

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or through your bookseller

Price $7.50 Catalogue No. A54-3/4-2

Price subject to change without notice

ROGER DUHAMEL, f.r.s.c.

Queen's Printer and Controller of Stationery

Ottawa, Canada

1968

IV

PREFACE

The systematic portion, or Part 1, of the Flora of the Queen Charlotte
Islands (1968) was initiated after a three-month botanical survey on the Islands
conducted during the summer of 1957. After this survey it became apparent
that a second summer of collecting on the Islands was needed before a system-
atic study of the flora could be completed. When plans were being formulated
for a two-and-a-half-month collecting trip to the Islands during the summer
of 1964, it was decided to collect cytological material of Queen Charlotte Is-
lands plants at the same time. Over 1,600 fixations of cytological material were
collected by Taylor during 1964 and living material of many species was for-
warded to Ottawa for growth in greenhouses and later cytological study. A
representative living plant collection was made by L. C. Sherk, a member of
the staff of the Plant Research Institute, who accompanied the 1964 survey
party for the first two weeks. We subsequently examined the cytological ma-
terial in 1964, 1965, 1966 and early 1967. Dr. R. J. Moore has been respon-
sible for the examination and subsequent discussion of taxa in the genus Carex.

The two companion volumes of Flora of the Queen Charlotte Islands:
Systematics of the Vascular Plants and Cytological Aspects of the Vascular
Plants are unique as they represent the first time that workers have come to-
gether as a team to study the systematic and cytological aspects of a flora con-
currently. Chromosome numbers have appeared in previous floras, for
example, the Flore Laurentienne by Marie-Victorin (1935), Vascular Plants
of the Pacific Northwest by Hitchcock et al. (1955, 1959, 1961, 1964), A
California Flora by Munz and Keck (1959), and Flora Europaea by Tutin et
al. (1964), but the chromosome counts were obtained from the literature.
Other authors have published original chromosome numbers for a large pro-
portion of plants in a geographic area but they have not prepared a systematic
treatment at the same time, for example, Love and Love (1956a) for Iceland,
and J0rgensen ^/ «/. (1958) for Greenland.

This study of chromosome numbers of plants from the Queen Charlotte
Islands has undoubtedly revealed more taxonomic problems than it has solved.
In many instances chromosome numbers obtained from Queen Charlotte plants
differ from those reported for the same taxon from elsewhere. Sometimes the
same taxon may have two or more chromosome races restricted to the Queen
Charlotte Islands. When we have been able to correlate differences in chromo-
some number with morphological differences we have invariably found that
the populations had previously been recognized taxonomically, based solely on
morphological evidence. In most cases, the discovery of two or more chromo-
some races has merely served to point out fruitful areas for more detailed
cytotaxonomic studies. The chromosome numbers found in Queen Charlotte
Islands plants have been useful in determining the base number or numbers
found in various genera and in establishing the presence of polyploidy or

aneuploidy within genera. The percentage of polyploids in the flowering plants
of the Queen Charlotte Islands flora has been calculated and compared with
that of other areas.

A conservative approach has been used to evaluate the taxonomic impli-
cations of differences in chromosome number within recognized species. We
do not consider it useful to come to sweeping taxonomic conclusions based on
a few chromosome numbers any more than most taxonomists consider it
feasible to make taxonomic decisions on the basis of a few herbarium specimens.
Chromosome races must be separable morphologically before they can be
recognized taxonomically, for taxonomy would surely be in chaos without
morphology for classification and the rules of nomenclature for order.

The increase in the number of chromosome counts that have been made
for different species has provided a greater insight into the variation of chromo-
some numbers within and between species. It is becoming more apparent, as
the amount of chromosome number information increases, that a far larger
number of plant taxa possess more than one ploidy level or show small devia-
tions in the number of chromosomes than had previously been recognized. The
overriding conclusion, evident after our work on the Queen Charlotte Islands,
is that we have but scratched the surface of a vast field of cytological information
that could be made available if cytological surveys were incorporated into
regular plant-collecting expeditions. It is apparent that the collection of cyto-
logical material should become standard practice in botanical surveys. The
examination of the material can always be made at a later period, provided
proper care is exercised in the fixation and storage of material. We are still far
from the goal of having sufficient information to make competent cytological
comparisons of floras from widely divergent regions. We hope that our en-
deavors on the botanical problems of the Queen Charlotte Islands will encour-
age other workers to pursue similar studies in other parts of North America.

Roy L. Taylor
Gerald A. Mulligan
Ottawa, 1967

VI

ACKNOWLEDGMENTS

Many people have assisted either directly or indirectly with the develop-
ment and preparation of this volume. Most of them helped with Part 1 too,
but we wish to acknowledge here their special contributions to Part 2.

We would like to express our appreciation to J. A. Calder, who has
contributed in many ways to the development of this volume through his collect-
ing activities and continued interest and constructive criticisms. Particular
thanks are extended to the greenhouse and herbarium technical staffs for their
assistance. We would particularly like to thank C. W. Crompton and A. G.
Moulds for their assistance in processing and preparing cytological material.
Miss J. Horton was responsible for the preparation of the list of literature
cited, the index, and the initial list of citations to previous counts.

vn

CONTENTS

Page

V Preface

vii Acknowledgments

1 The Islands and Their Environment

3 Composition of the Vascular Flora

4 Materials and Methods

8 Polyploidy

9 Format of Cytological Flora

12 Chromosome Numbers of Vascular Plants

121 Literature Cited

139 Index

IX

THE ISLANDS AND THEIR ENVIRONMENT

The Queen Charlotte Islands consist of approximately 150 islands grouped
in a triangle-shaped archipelago that lies off the northwest coast of British
Columbia. Most of the islands are between 52° and 54° N latitude and 131° and
133° W longitude. The Charlottes are approximately 156 miles long with a
maximum width of 52 miles and a combined land mass of nearly 3,600 square
miles. The Islands can be divided into three physiographic areas (Figure 1):
the Queen Charlotte Ranges, the Skidegate Plateau, and the Queen Charlotte
Lowlands. A few peaks of the mountains of the Queen Charlotte Ranges,
between 52° 40' N and 53° 20' N, reach 4,000 feet, with most mountains be-
ing between 2,500 and 3,500 feet high.

The extent of glaciation on the Queen Charlotte Islands has been the
subject of much speculation. In a summary of their findings about glaciation
on the Islands, Brown and Nasmith (1962, p. 218) state that "the evidence
obtained from geological studies clearly indicates that during the Wisconsin
period the Queen Charlotte Islands were buried by glacial ice. This ice was
generated in the mountainous regions of the islands and flowed outward to
join ice from glaciers in the coastal mountains. At the maximum stage of
glaciation probably not more than 3.5 square miles of the land surface stood
above the glacier ice and this small area was subject to severe arctic climate and
swept by snow and rock slides to produce a most inhospitable environment."

As stated by Calder and Taylor in Part 1 , "the indigenous flora of the
Queen Charlotte Islands was to a large degree maintained in refugia on the
headlands and along the fiords of the west coast during maximum Pleistocene
glaciation. The existence of refugia is supported by the presence of clear-cut
endemics, by several examples of disjunct distributions, and finally by the pat-
terns of distribution of the indigenous taxa. An analysis of native plants shows
that the Queen Charlotte Islands represent a center of postglacial plant
radiation similar to the examples given by Hulten (1937) in his discussion of
the theory of equiformal progressive areas. In the postglacial period subsequent
migrations from the south and north took place along the Cordilleran coast. In
addition, a second wave of migrants, the Hypsithermal elements, also invaded
the Queen Charlotte Islands. Subsequent lowering of the mean annual tempera-
tures has left isolated pockets of these elements stranded on the mesic sites
along the east coast and a few drier inland sites. In recent historical times,
introduced taxa have increased the flora about one quarter."

The Queen Charlotte Islands have cool summers and mild winters com-
pared to the rest of Canada. They often have cloudy skies and strong winds
with excessive late fall and early winter rainfall. Annual precipitation exceeds
40 or 50 inches over most parts of the Islands, with 200 to 300 inches on some
of the western slopes. Much of the winter precipitation in the mountains falls
as snow and this snow may last well into the summer at altitudes of 2,000 feet

LANGARA ISLAND

132° 131°

DIXON ENTRANCE

ROSE POINT

GRAHAM
ISLAND

/>^ A SKIDEGATE INLET

HECATE STRAIT

PACIFIC OCEAN

LOUISE ISLAND

TASU SOUND Iv Ti\ / Vr-:::::^

MORESBY

ISLAND

'^"^V^l/ } LYELL ISLA^

JUAN PEREZ
SOUND

LEGEND
PHYSIOGRAPHIC BOUNDARY

132°
I

Figure 1 . Map of the Queen Charlotte Islands showing the three physiographic units: Queen Charlotte
Lowlands, Skidegate Plateau, and Queen Charlotte Ranges.

or more. Near sea level the average temperatures are below 60 degrees
Fahrenheit in the warmest month, but are above freezing in the coldest month.
Average frost-free periods range from 160 days at Masset to over 260 days at
Cape St. James. Days are long during the growing season but the durations of
bright sunshine are among the shortest in Canada.

COMPOSITION OF THE VASCULAR FLORA

The vascular flora of the Queen Charlotte Islands comprises 70 families,
277 genera, and 594 species and subspecific taxa. In Table 1 the flora is tabu-
lated by subdivisions, classes and subclasses.

Table 1. Composition of the flora on the Queen Charlotte Islands

Subdivision

Class

Subclass

Families

Species

and sub-

jenera

specil

fic taxa

Percent of

Number

total flora

1

7

1.2

3

12

2.0

273

574

96.8

14

24

4.1

7

8

1.3

252

542

91.4

59

183

30.9

193

359

60.5

277

593

100.0

Psilopsida —

Sphenopsida 1

Lycopsida 3

Pteropsida 65

Filicinae 3

Gymnospermae 3

Angiospermae 59

Monocotyledoneae 11

Dicotyledoueac .... 48

Total vascular flora 69

A numerical breakdown and the percentage of indigenous and introduced
taxa found on the Queen Charlotte Islands and the number and percentage of
annuals, biennials and perennials are given in Table 2.

Table 2. Introduced and indigenous taxa and their life duration in the flora of the Queen
Charlotte Islands

Annual Biennial Perennial Totals

Indigenous 18 5 453 476

Percentage of indigenous 3.8 1.0 95.2 100

Percentage of flora 3.1 0.8 76.4 80.3

Introduced 51 8 59 118

Percentage of introduced 43.1 6.9 50.0 100

Percentage of flora 8.6 1.3 9.8 19.7

Total taxa in flora 69 13 512 594

Percentage of flora 11.6 2.2 86.2 100

The high percentage of indigenous perennials is not surprising for a flora

found between approximately 52 and 54° N. However, the large number of

annuals, of which most represent introduced taxa, indicates that even on the
Charlottes there are some quite dry environments. Almost without exception,
these annuals, whether they are indigenous or introduced, occur only in the
drier eastern coastal areas of Graham and Moresby islands. Many of these
annuals reach their northern limit on the Charlottes; the next station to the south
is in the dry southeastern section of Vancouver Island. Clearly, the indigenous
annual taxa represent the remnant of a group that was once more widely dis-
tributed during or preceding the Hypsithermal.

A most interesting facet of the flora is the high degree of endemism.
Eleven endemic taxa are found on the Islands and they represent over 2 percent
of the indigenous flora. Of these eleven taxa, nine are restricted to the subalpine-
alpine zones. As the montane flora contains about 120 taxa, the endemics
represent just over 9 percent of this group. This is an extremely high degree of
endemism for such a small area at so northern a latitude.

MATERIALS AND METHODS

The methods used in the cytological survey of the Queen Charlotte Islands
were tested during the summer of 1963 by R. L. Taylor and L. C. Sherk on a
collecting trip in the western Canadian provinces. At that time particular
emphasis was placed on the bulk collection of cytological material under a
wide variety of conditions. Experience was obtained in the use of a cytological
collecting vest and a carton for handling vials in the field and for shipping them
back to Ottawa. Both the shipping carton (Figure 2) and the collecting vest
(Figure 3) were modified for use on the survey of the Islands during the winter
of 1963—1964. It was found that fixatives such as Carnoy's, containing both
alcohol and acetic acid, could be mixed up to 3 days before the time material
was fixed without any obvious deterioration of the meiotic material. It may be
significant that the cytological material obtained in 1963 and 1964 was collected
under moderate climatic conditions (65°— 85° F). The two fixatives used were
Carnoy's solution (6 parts methyl alcohol, 3 parts chloroform, and 1 part glacial
acetic acid) for all flower buds and sporogenous material, and Randolph's
modified Navashin solution (solution A: 1 g chromic acid, 7 cc glacial acetic
acid, and 92 cc distilled water; solution B: 30 cc formalin, and 70 cc distilled
water) for root tips.

Before departing for the Islands in the summer of 1964, we partially filled
vials (57 mm X 28 mm with molded plastic screw caps) with either 4/10
volume chloroform— acetic acid solution of the Carnoy's fixative or 5/10
volume solution A of the Navashin fixative. Bulk solutions of the other parts of
the fixatives were placed in large plastic bottles. In the field, fixative solutions
were prepared up to 3 days before they were used by adding the complimentary
solution to each vial. To avoid mixing the wrong solutions the various solutions
had been prestained with different dyes. The 24 vials carried in the collecting
vest were usually enough for a 2- to 3-hour collecting period; however, on
extensive mountain or hiking trips, extra vials were carried in the collecting
bags. The labels (45 mm X 15 mm) used in the vials were made from 20/1000

Figure 2. Carton for shipping cytological specimens. Corrugated carton 4 inches deep x 1 8 inches long
X 1 2 inches wide. Spacers divide box in 1 04 cubicles for viols. A sheet of Kempock embossed padding is
placed above and below the vials to provide protection. Additional reinforcement and support is pro-
vided by a sheet of masonite board between the embossed padding and the top and bottom of the carton.

Figure 3. Cytological collecting vest. Vest is made of neoprene to safeguard wearer from fixatives
spilled through breakage or leakage of vials. The 24 vials are inserted into elasticized pouches.

gauge "Cobcx" opaque white vinyl. An ordinary lead pencil was used to record
the necessary data on the labels. These vinyl labels have the advantage of not
deteriorating during long storage and the data remain legible throughout the
staining process. When a large amount of cytological material had been fixed
in a vial, the fixative was usually replaced after we returned to the survey base.
All vials were topped with fixative before shipment in cartons by air freight to
Ottawa.

GENUS: S^a^^fi^fl
SPECIES: o«rHOC£ffrt.C
COLLECTION NO,: cXs
SOURCE: T'otiJ HILL.

—

n

2n

Voucher / J? ■/- /
Annotated •*

^«

Photograph
N e g at iv e No .
D r aw ing

Wild/

C ult

Counted by:

~ (^l^fthtflr^ ist/9fvj>

Cross Index

to Flora S.'^lo

AUTHOR & REFERENCE:

rA ^\\\ a,ar>

Vv^^oH^l - Bt-^A->

i W Co\ umnb la

Ro\\>^S \\^\o\^\ - Ca\v-for

r\\ a

/h..C-<-*\2..c^ /I / ^.t-c-/^,^*^

Figure 4. Information card used for accumulation of cytological and collection data.

When the cartons of vials were received in Ottawa, the cytological material
was prepared for storage, or for embedding in paraffin. Each vial was checked
to determine whether it contained meiotic or mitotic material. An information
card (Figure 4) was initiated at this stage and this card was subsequently used
for the accumulation of data, including the recording of special features during
the examination of the material.

Material collected in the field and shipped to Ottawa was processed, and
stored by two general techniques :

(1) Material collected for the examination of mitosis in root tips.

All root tip material was killed and fixed in Randolph's modified
Navashin's solution for a period of approximately two weeks. After
this initial period of fixation, the root tips were removed from the vials
and thoroughly rinsed in water and all extraneous matter was re-
moved from them. Material was put through a series of 30, 50 and 70
percent alcohol changes at one-hour intervals and then stored in a
freezer at -4° C.

(2) Material collected for the examination of meiosis in flower bud and
sporogenous material.

All flower buds and sporogenous material were killed and fixed in
Carnoy's solution. When the material reached the laboratory, it was
rinsed three times at one-hour intervals in 70 percent alcohol, then
stained with an alcoholic hydrochloric acid-carmine technique
described by Snow (1963). After 5 to 7 days, the material was rinsed
three times at one-hour intervals with 70 percent alcohol to remove
excess stain before it was stored in a freezer at —4° C for periods
up to two years.

For examination, material was squashed in 45 percent acetic acid and
preparations were made permanent with carbon dioxide and an
apparatus outlined by Johnson and Janick (1962).

Microslides for examination of mitosis in root tips were prepared by
methods that have been developed by the Plant Research Institute over the
past several years. Some difficulty was experienced in obtaining well-stained
preparations of material that had been in storage for a long time. To overcome
this difficulty, a 1 percent aqueous solution of chromic acid was introduced as
a mordant into the schedule before staining. The addition of this extra chromic
acid replaced the gradual loss of chromic acid that occurs during long periods of
storage. The schedule used for the preparation and staining of the slides follows:

Root tips are dehydrated through a tertiary butyl alcohol series and
embedded in a mixture of 1 part Tissue Mat and 3 parts commercial paraffin.
Material is sectioned at varying thicknesses depending upon the genus, for
example, Carex and J uncus were sectioned at 10 microns, whereas Agrostis
and Alopecurus were sectioned at 30 microns. Serial sections were mounted on
microslides with Mayers adhesive. Paraffin was removed by passing the slides
through a series of xylene, dioxane, and alcohol, and then hydrating through a
descending series of alcohol to water. Slides were mordanted in 1 percent
chromic acid solution for approximately 4 to 5 hours, washed in 30 percent
alcohol for 2 minutes and water for 5 minutes, and stained in crystal violet ( 1
percent aqueous solution) for 45 minutes. After they were stained, the slides
were rinsed quickly in water, and then mordanted in an aqueous iodine solution.
Slides were then transferred to clove oil for differentiation and finally cleared in
xylene. Cover slips were placed on the material with a piccolyte mounting
medium.

A few mitotic examinations were conducted on the squashed floral organs
prepared by the Snow technique.

Although most of the counts were obtained on material collected in the
field, additional counts were obtained on material grown in Plant Research
Institute greenhouses. The cultivated plants were grown from seed collected
in the field in 1964 and from herbarium collections made in 1957, or they
represented transplants of wild material collected in the summer of 1964.
Meiotic examination was made on flower buds or sporogenous material by the
Snow technique. Root tips were either prepared as outlined in the procedure for

8

field material or they were squashed using the oxyquinoline and aceto-orcein
method outlined by Tjio and Levan (1950).

Examination of both meiosis and mitosis was made using either a Reichert
Zetopan research microscope equipped with phase contrast and anoptral con-
trast equipment or a Cari Zeiss research microscope equipped with phase
contrast equipment.

All microslides used in the cytological study are kept at the Plant Research
Institute, Ottawa. The results of the examinations have been transferred to the
herbarium voucher specimens using a standard annotation slip (Figure 5). A
complete set of cytological voucher specimens is housed in the Department
of Agriculture herbarium in the Plant Research Institute in Ottawa. Additional
sets or partial sets of vouchers will be distributed to other herbaria.

Voucher specimen for chromosome number cited in FLORA OF THE QUEEN

LOTTE ISLANDS. II. CYTOLOGICAL ASPECTS OF THE VASCULAR

by Roy L. Taylor and Gerald A. Mulligan.

CHAR-
FLORA

n =

2n =

Figure 5. Annotation slip used for recording cytological information on herbarium voucher specimens.

POLYPLOIDY

The incidence of polyploidy in the angiosperm flora of the Queen Charlotte
Islands is given in Table 3. Species of the genus Carex were excluded from the
calculations because of the unusual chromosome situation found in the genus.

The frequency of polyploids in the angiosperm flora of the Queen Charlotte
Islands was 53.2 percent at 52°— 54° N latitude, which is about the same as that
reported by Johnson and Packer (1965) for Ogotoruk Creek, Alaska: 55.8
percent at 68° N latitude. Neither of these areas is typical of locations at similar
latitudes in western North America. The Queen Charlotte Islands and the Ogo-
toruk Creek areas contain relic floras, whereas most of western North America
at similar latitudes was completely glaciated. Before the significance of poly-
ploidy in western North America can be validly estimated, more information is
needed about the incidence of polyploidy in the adjacent glaciated areas of
western North America and in the large unglaciated areas to the south in the
United States and Mexico.

Table 3. Polyploidy in the angiosperm flora of the Queen Charlotte Islands

Native

Introduced

Total

Diploid

Polyploid

Diploid

Polyploid

Diploid

Polyploid

Monocotyledons

36(37.5%)

60(62.5%)

5(33.3%)

10(66.7%)

41 (36.9%)

70(63.1%)

Dicotyledons

98(47.6%)

108 (52.4%)

40(61.5%)

25(38.5%)

138(50.9%)

133(49.1%)

Angiosperms

134(44.0%)

168(56.0%)

45 (56.2%)

35 (43.8%)

179(46.8%)

203 (53.2%)

Hagerup (1932) first indicated that polyploids are more tolerant of
extreme environments. Tischler (1935) reported that, based on a study of four
European regions, the percentage of polyploidy could be correlated with
latitude. Several other cytological studies followed and the data from 14 such
studies were discussed and summarized in graphic form by Love and Love
(1949). No similar comparison is possible for North America because of the
paucity of cytological information in this floristic region. Furthermore, as
Stebbins (1950) has pointed out, a comparison of polyploidy of different areas
should entail the cytological investigation of the same families or genera in
order to draw valid conclusions on the percentage of polyploidy found in the
floras under consideration.

The percentages of polyploidy in the native and introduced flora of the
Queen Charlotte Islands are very interesting. The native plants of the Islands
are 56.0 percent polyploid, whereas the introduced plants are only 43.8 per-
cent polyploid. This supports the conclusion of Heiser (1950), Heiser and
Whitaker (1948), and Mulligan (1960), that there appears to be no evidence
of any general advantage for introduced weeds to be polyploid.

Of the eleven endemic taxa on the Islands, eight have been counted. Six
are polyploid: four tetraploid, one octoploid and one 16-ploid. Isopyrum
savilei is diploid and Saxifraga taylori is both diploid and tetraploid. All en-
demics are perennial and occur in the montane region.

FORMAT OF CYTOLOGICAL FLORA

The 277 genera included in this flora belong to the division Tracheophyta.
Three subdivisions are represented on the Queen Charlotte Islands, namely,
Sphenopsida, Lycopsida and Pteropsida. The last subdivision, which contains
most of the genera in the flora, is further subdivided into three classes: Filicinae,
Gymnospermae and Angiospermae. The last class contains the two subclasses
Monocotyledonae and Dicotyledonae. Lawrence (1951) has been the guide
for family names and their sequence, which is in turn based on the Englerian
system. In the Angiospermae, the monocotyledonous families precede the
dicotyledonous, the latter ending with the Compositae. Genera and species are
arranged alphabetically under their respective families.

The species number is identical with that used for the same taxon in Part 1.
The citation Part 1 refers to Calder and Taylor (1968), Flora of the Queen
Charlotte Islands, Part 1 , Systematics of the Vascular Plants. An asterisk pre-
ceding the generic name means that a taxon is introduced to the Islands. When
no asterisk is used, the taxon is indigenous to the Islands.

All taxa that appeared in Part I appear in this volume. An additional
species, Vicia cracca L., has been included as species No. 390a. Not all taxa
have chromosome number determinations, but the inclusion of all taxa from
the Islands will facilitate the annotations of other counts when they become
available.

The citations of voucher specimens are grouped into two general geo-
graphic regions, Graham Island and Moresby Island. All specimens collected

10

LANGARA PT

FREDERICK I. {^

ATH
EMPIRE AN

ROSE SPIT

AN PT.
WHITE CREEK MUSKEG
SANGAN R.
TOSH ^•
DOWS V- '

SHIELDS B.

"'^nVBADDECK

^ixMrSLATECHU

r~MT. STAPLETON ( I

DAWSON I
DAWSON HR.
MARBLE l.»
ELLS
MERCER PT.
NEWTON PT.

CHAATL I.

SKIDEGATE CHAN .

KITGORO IN.-

KAISUN
MOSQUITO MTN

HIBBEN ISLAND
PEEL IN.

Lj o Ki Ki a D t jk; v^ • s a N DS P I

NG "-°Tm-A-^V^_-:^?E?4jORREN

LAWN PT.
LAWNHILL

DEAD TREE PT.
MILLAR CR.

QUEEN CHARLOTTE CITY
SKIDEGATE VIL.
SANDSPIT

S I. JEWELL I.

.SKIDEGAT

ALLIFORD
B.

KOOTENAY IN.

MT. RUSSC*''

HORN ROCK
TASU SOUND

ROUSTCHEFF
MUD MARSH
ER B.
HELDENS B.

AY B.

SKIDEGATE

CUMSHEWA

CUMSHEWA IN.

SKEDANS
• SKEDANS I.

LIMESTONE I .

• LOW I .
• south LOW I,
DASS PT.

DE LA TOUCHE
FAIRFAX IN

SUND
KWOO

CRESCENT IN.

LOCKEPORT

RICHARDSON I.

^TUFT IS.
^TARl.

LYELL ISLAND
HOTSPRING I.

0

UAN PEREZ SOUND

GOWGAIA B

MCTORIA L.., ii2l<fe^"»VEASTCOPPE
TZAMTN^-r^AGH*^ SKIN CUTTLE IN.

GOWDAS (.•<;--sJ) /K\ \
UPPER V

BURNABY ISLAND
BOLKUS I.

IKEDA

HARRIET HR.

Figure 6. Place names used on the Queen Charlotte Islands.

LOUSCOONE IN

ANTHONY I. '^
C. FANNY

KUNGHiT I.

WOODRU FF B.
'W C. ST. JAMES

11

from land areas north of the center of Skidegate Inlet and Skidegate Channel
are considered to be from Graham Island, and all those collected from land
areas to the south are considered to be from Moresby Island. The place names
for the Queen Charlotte Islands are shown in Figure 6.

Chromosome numbers were determined from examination of somatic and
gametic tissues. The symbol n indicates a gametic chromosome count obtained
by examination of the last two stages of meiosis I, or meiosis 11. The symbol
2n may refer to either a count obtained by examination of meiosis or mitosis.
If the symbol 2n is followed by a number with a subscript, for example 2n =
lOii (the bivalent sign indicates 10 pairs of chromosomes were observed at
either prophase or metaphase of meiosis I) , the count has been determined from
examination of meiosis. If the chromosome number was determined on somatic
tissue, no subscript appears after the chromosome number, for example
2n = 20.

The chromosome number is given in boldface when it is a new or different
count for a taxon.

All specimens cited are deposited in the Department of Agriculture
herbarium at Ottawa. The names of collectors who are members of the Plant
Research Institute are abbreviated as follows: CST (J. A. Calder, D. B. O.
Savile and R. L. Taylor); CTS (J. A. Calder, R. L. Taylor and L. C. Sherk);
S (D. B. O. Savile); and CT (J. A. Calder and R. L. Taylor).

Following the citation of chromosome numbers and voucher specimens,
a discussion of the significance of the chromosome numbers of Queen Charlotte
Islands plants is given. All previous chromosome counts for the taxon concerned
are noted. When many identical chromosome numbers are reported for a taxon
we refer only to the more comprehensive papers and to general compendiums
of chromosome numbers (Delay 1950, Tischler 1950, Darlington and Wylie
1955, Chiarugi 1960, Love and Love 196 la, and Fabbri 1963). Whenever
possible we have used the comprehensive chromosome list of Love and Love
(1961c[).

CHROMOSOME NUMBERS OF VASCULAR PLANTS

Equisetaceae

Equisetum

1. Equisetum arvense L.

2. Equisetum fluviatile L.

3. Equisetum hyemale L. ssp. aflfine (Engelm.) Calder & Taylor

4. Equisetum hyemale X E« variegatum (= X E. trachyodon A. Br.)

5. Equisetum palustre L.

MORESBY ISLAND: In = 108„, Skidegate Lake, CT35150.

This count is the first reported for native North American material and
agrees with previous counts {see Love and Love 1961/?).

6. Equisetum telmateia Ehrh.

7. Equisetum variegatum Schleich. ssp. alaskanum (A. A. Eaton) Hult.

Lycopodiaceae
Lycopodium

8. Lycopodium annotinum L.

9. Lycopodium clavatum L.

10. Lycopodium complanatum L.

11. Lycopodium inundatum L.

12. Lycopodium obscurum L.

13. Lycopodium subinaefolium Willd. ssp. sitchense (Rupr.) Calder & Taylor
14a. Lycopodium selago L. ssp. selago

LYCOPODIUM 13

14b. Lycopodium selago L. ssp. miyoshianum (Makino) Calder & Taylor

14c. Lycopodium selago L. ssp. patens (Beauv.) Calder & Taylor

Selaginellaceae
Selaginella

15. Selaginella selaginoides (L.) Link

16. Selaginella wallacei Heiron

Isoetaceae

ISOETES

17. Isoetes echinospora Dur. ssp. muricata (Dur.) Love & Love

Ophioglossaceae

BOTRYCHIUM

18a. Botrychium lunaria (L.) Sw. ssp. lunaria

18b. Botrychium lunaria (L.) Sw. ssp. minganense (Vict.) Calder & Taylor

19. Botrychium multifidum (Gmel.) Rupr. ssp. silaifolium (Presl) Clausen

MORESBY ISLAND: 2n = 45„, Upper Victoria Lake, CT 35777.

The same chromosome number was previously reported for material of
B. multifidum (Gmel.) Rupr. from Michigan and California (Wagner 1955,
and Wagner and Chen 1964).

Hymenophyllaceae

Mecodium

20. Mecodium wrightii (v.d. Bosch) Copeland

Polypodiaceae

Adiantum

21. Adiantum pedatum L. ssp. aleuticum (Rupr.) Calder & Taylor
GRAHAM ISLAND: 2n = 29u, Long Inlet, CT35997.

14 POLYPODIACEAE

Although several counts of ^7 = 29 or 2^? = 58 have been reported for
the species (see Fabbri 1963), only one previous count has been made on the
western North American ssp, aleuticum. Wagner {in Fabbri 1963) reported
Az = 29 from Washington material.

ASPLENIUM

22. Asplenium trichomanes L.

MORESBY ISLAND: n = ca. 72, Mt. Moresby, CT36381.

Many counts have been made on this widely distributed species, with
diploids, tetraploids, and hexaploids based on ;c = 36 being reported (see
Fabbri 1963).

23. Asplenium viride Huds.

MORESBY ISLAND: n ^ 36, Lang 42 (UBC).

The chromosome count of n = 36 by Taylor and Lang (1963) for ma-
terial from Moresby Island was also reported by many authors on material
from elsewhere (see Fabbri 1963, and Britton 1964).

Athyrium

24. Athyrium filix-femina (L.) Roth. ssp. cyclosorum (Rupr.) C. Chr. in
Hult.

MORESBY ISLAND: 2n = 40n, head of Cumshewa Inlet, CT35226.

The same chromosome number, n = 40 or 2n = 80, was reported by
many workers for material from North America, Europe and Asia (see Love
and Love 1961/?, 1966, and Fabbri 1963). Roy and Pandey (in Fabbri 1963)
reported n = 41 and 2n = 82 for plants of var. pectinata CI. from India.

Blechnum

25. Blechnum spicant (L.) With.

MORESBY ISLAND: 2n = 34n, Upper Victoria Lake CT35778.

All previous counts of this species give the same number as we have re-
ported here. Wagner (in Fabbri 1963) reported « = 34 from plants of Wash-
ington.

Cryptogram MA

26. Cryptogramma crispa (L.) R. Br. ssp. acrostichoides (R. Br.) Christ
MORESBY ISLAND: 2n = 30„, Mt. Moresby, CT 36399.

CRYPTOGRAMMA 15

Wagner (in Fabbri 1963) reported n = 30 for this taxon from Washing-
ton. Manton (1950) reported At = 60 for C. crispa (L.) Hook, from England.
Recently Taylor and Lang (1963) reported n = AO for C. crispa var. acro-
stichoides from Pitt River and Mount Seymour in southwestern British Colum-
bia. However, the number reported appears to be a typographical error
because their discussion implies that their material represented a diploid race
based on x = 30.

Cystopteris

27. Cystopteris fragilis (L.) Bernh.

MORESBY ISLAND: 2n = 84n, Mt. Moresby, CT 36379.

Recent workers have proposed that the base number for this genus be
X = A2 {see Fabbri 1963). The species contains both tetraploids and hexa-
ploids but no diploids have been reported.

Dryopteris

28. Dryopteris austriaca (J acq.) Woynar in Schinz & Thellung

GRAHAM ISLAND: n = ca.Al, Collinson Lake, CT3552L

Numerous counts have been reported for this species and all are based on
X = A\ {see Chiarugi 1960, and Fabbri 1963). There are both diploid and
tetraploid races reported for the species complex.

Gymnocarpium

29. Gymnocarpium dryopteris (L. ) Newman

GRAHAM ISLAND: n = ca. 40, Blackwater Creek, CTS35064.

Wagner {in Fabbri 1963) reported n = 40 for material from Washington.
It is interesting that both populations examined from the Pacific coast region
are diploid with the base number x = 40. Tetraploids with the same base
number have been reported in Europe and eastern North America by a number
of authors {see Chiarugi 1960, Fabbri 1963, and Love and Love 1966).

POLYPODIUM

30. Polypodium scouleri Hook. & Grev.

31a. Polypodium vulgare L. ssp. occidentale (Hook.) Hult.

GRAHAM ISLAND: 2n - 37„, Rose Spit, CT35908; In - 37„,
Yakoun Lake, CT36776.

16 POLYPODIACEAE

MORESBY ISLAND: 2n = 37„, head of Cumshewa Inlet, CT36250;
In = 37n, Kaisun, CT36527.

There are both diploids and tetraploids reported for P. vulgare L. (see
Chiarugi 1960, and Fabbri 1963). All material examined from the Pacific
Northwest is diploid (Taylor and Lang 1963).

31b. Polypodium vulgare L. ssp. columbianum (Gilbert) Hult.

POLYSTICHUM

32a. Polystichum braunii (Spenner) Fee ssp. alaskense (Maxon) Calder &
Taylor

32b. Polystichum braunii (Spenner) Fee ssp. andersonii (Hopkins) Calder &
Taylor

32c. Polystichum braunii (Spenner) Fee ssp. purshii (Femald) Calder &
Taylor

GRAHAM ISLAND: 2n = S2u, near junction of Yakoun River and
Ghost Creek, CT35500.

One previous count of « = 82 has been made on this infraspecific taxon
by Manton and Reichstein (1961). Other counts on F. braunii (Spenner) Fee
by the same authors and by Taylor and Lang ( 1 963 ) are also tetraploid based
on ;t = 41.

33. Polystichum lonchitis (L.) Roth ex Roem.

MORESBY ISLAND: 2n = ca. 82, Takakia Lake, CT36333.

Previous counts on this species by Manton (1950, 1953), Love and Love
(1961/?), Britton {in Fabbri 1963), Britton (1964) and Wagner and Chen
(1964) report a diploid number based on x = 41.

34. Polystichum munitum (Kaulf.) Presl

MORESBY ISLAND: 2n = 41n, Kootenay Inlet, CT36214.

The same number was reported on Pacific Northwest material by Wagner
(1963) and by Taylor and Lang (1963).

Pteridium

35. Pteridium aquilinum (L.) Kuhn ssp. aquilinum var. pubescens Underw.

THELYPTERIS 17

Thelypteris

36. Thelypteris oreopteris (Ehrh.) Slosson in Rydb.

37. Thelypteris phegopteris (L.) Slosson in Rydb.

MORESBY ISLAND: n = ca. 90, head of Cumshewa Inlet, CT35227;
In = 90n, Mt. Moresby, CT36372.

Previous counts by Britton (1953) on plants from Ontario, Canada, and
by Sorsa (1958) on plants from Finland are reported as /i = 90. This is the
only number known for this species and it is probably a hexaploid based on
X = 30.

Taxaceae
Taxus

38. Taxus brevifolia Nutt.

Pinaceae

PiCEA

39. Piceasitchensis (Bong.) Carr.

GRAHAM ISLAND: In = 24, Tlell, CT 34646.

The same chromosome number was reported for North American material
of this species by Thomas {in Munz and Keck 1959).

PiNUS

40. Pinus contorta Dougl. ex Loud.

GRAHAM ISLAND: In = 24, Port Clements, CT 34607.

The only previous count of P. contorta was In = 24 by Langlet (1934).
This number conforms to general cytological findings of all other species of
pines.

TSUGA

41. Tsuga heterophylla (Raf.) Sarg.

GRAHAM ISLAND: In = 24, 5 mi W of Tow Hill, CT34711.

The same chromosome number was recently reported by Taylor and
Brockman (1966) from material collected near Revelstoke, British Columbia.

18 PINACEAE

42. Tsuga mertensiana (Bong.) Sarg.

Cupressaceae
Chamaecyparis

43. Chamaecyparis nootkafensis (Lamb.) Spach

JUNIPERUS

44. Juniperus communis L.

GRAHAM ISLAND: 2n = 22, about 5 mi S of Masset, CT35576.

The same chromosome number has been obtained on North American
and European material (see Love and Love 1961a).

Thuja

45. Thuja plicata D. Don

GRAHAM ISLAND: 2n = 22, 9 mi N of Port Clements, CTS34710.

The same chromosome number has been reported by Sax and Sax (1933),
and Mehra and Khoshoo (1956).

Sparganiaceae

Sparganium

46. Sparganium hyperboreum Laest. ex Beurl.

GRAHAM ISLAND: 2n = 30, Pure Lake CT36098.

MORESBY ISLAND: 2n = 30, Kootenay Inlet, CT36138.

The same mitotic chromosome number was reported from Greenland by
J0rgensen et al. (1958) and from Scandinavian material by Love and Love
(1948).

47. Sparganium minimum E. M. Fries

GRAHAM ISLAND: 2n = 30, Yakoun Lake, CT36766.

MORESBY ISLAND: 2n = 15n, about 2 mi S of Sandspit, CT36085;
2n = 30, Red Mud Marsh, CT36724.

Sparganium minimum was previously reported from Europe to have
2n = 30 (see Love and Love 1961a).

SPARGANIUM 19

48. Sparganium simplex Huds.

GRAHAM ISLAND: In = 30, about 4 mi N of mouth of Oeanda River,
CT35874.

This species has been previously counted from Europe as 2n = 30 (see
Love and Love 1961a). The same number has been recently reported from
Russia by Sokolovskaja (1963 ) .

Zosteraceae
Phyllospadix

49. Phyllospadix scouleri Hook.

POTAMOGETON

50. Potamogeton alpinus Balbis ssp. tenuifolius (Raf. ) Hult.

51a. Potamogeton berchtoldii Fieber ssp. berchtoldii

MORESBY ISLAND: 2n = 26, White Swan Bog, CT35297.

51b. Potamogeton berchtoldii Fieber ssp. tenuissimus (Mert. & Koch) Calder
& Taylor

GRAHAM ISLAND: « = 13, 2 mi NW of Tlell, CT35688.

Previous chromosome numbers reported for this species have appeared
under the name P. pusillus L. and were either n = 13 or 2« = 26 (see Love
and Love 1961fl).

52. Potamogeton epihydrus Raf. ssp. nuttallii (Cham. & Schlecht.) Calder
& Taylor

GRAHAM ISLAND: 2n = 26, 2 mi NW of Tlell, CT35694,
MORESBY ISLAND: 2n = 13„, 2 mi S of Sandspit, CT 36084.

Material of this species from the Queen Charlotte Islands is diploid with
the base number x = XT).

53. Potamogeton gramineus L.

MORESBY ISLAND: n = 26, Skidegate Lake, CT 36056.

All previous counts of this species reported are tetraploid, 2m = 52 {see
Love and Love 196 la, and Stern 1961 ).

20 ZOSTERACEAE

54. Potamogeton natans L.

55. Potamogeton nodosus Poir. in Lam.

MORESBY ISLAND: n = 26, Upper Victoria Lake, CT35769.

Potamogeton nodosus is a tetraploid with the base number ;c = 13. The
chromosome count given for this species in Love and Love (1961a) was
actually obtained on material of P. fluitans Roth by Kuleszanka (1934).

56. Potamogeton pectinatus L.

57. Potamogeton richardsonii (Bennett) Rydb.

58. Potamogeton robbinsii Oakes

RUPPIA

59. Ruppia maritima L.

GRAHAM ISLAND: 2n - 20, Kumdis Creek Delta, CT36125.

The karyotype of this species is a most interesting one and has been lucid-
ly discussed and illustrated by Reese (1962). The chromosome number of
2n = 20 for the Queen Charlotte material supports the decision of Calder and
Taylor in Part 1 in which they indicate that, on the basis of morphology, all
material from the Pacific Northwest should be recognized as R. maritima.
Chromosome numbers given for R. spiralis L. are all tetraploid, 2n = 40 (see
Reese 1962).

ZOSTERA

60. Zostera marina L.

GRAHAM ISLAND: 2n = 12, Kumdis Creek Delta, CT36126.

MORESBY ISLAND: 2n = 12, Richardson Island, CTS34919;
2n = 12, Kootenay Inlet, CT36215.

The same chromosome number was reported for European and Asiatic
material of this species (see Love and Love 1961a).

Juncaginaceae
Triglochin

61. Triglochin maritimum L.

TRIGLOCHIN 21

GRAHAM ISLAND: 2n = 48n, Yakoun River, CT35472.

Many chromosome races have been reported in the Old and New World
populations of T. maritimum agg., In = 12, 24, 30, 36, 48, 96, 120 and 144
{see Love and Love 1961a, and Love and Love 1958). At least three species
are recognized in this complex in North America by Love and Love (1958) :
an eastern North American species, T. elatum Nutt. {In = 144); a dry conti-
nental species, T. debile (Jones) Love and Love {In = 96) \ and a Pacific coast
species, T. concinnum Davy {2n =48). Recent chromesome counts by Packer
(1964) and by the present authors indicate that the pattern described by Love
and Love (1958) is not as clear as they have suggested in their paper. Packer
(1964) has shown that plants recognized as T. debile by Love and Love (1958)
also have the chromosome number 2n = 48. Our count from the Queen Char-
lotte Islands clearly indicates that the northern Pacific coastal population of
T. maritimum has a chromosome number 2n = 96. In contrast, the southern
Pacific coastal entity, T. concinnum, is tetraploid, 2n = 48.

62. Triglochin palustre L.

GRAHAM ISLAND: n = 18, Delkatla Inlet, CT35588.

MORESBY ISLAND: 2n = 12u, Upper Victoria Lake, CT35739.

This species has been counted by many authors and all have reported
n = 12 or 2n = 24 {see Love and Love 1961^). Material from Delkatla
Inlet had irregular meiosis and formed trivalents and (or) univalents at meta-
phase I (e.g. lOn + li + 5in). This population is obviously behaving as a
triploid, although it is a hexaploid with the base number x = 6.

Hydrocharitaceae

Elodea

63. Elodea canadensis Michx.

Gramineae

Agropyron

64. * Agropyron repens (L.) Beauv.

Agrostis

65. Agrostis aequivalvis (Trin.) Trin.

GRAHAM ISLAND: 2n = 14, 3 mi E of Juskatla, CTS35046;
2n = 7ii, about 4 mi N of mouth of Oeanda River, CT35876; 2n = 14,
Mayer Lake, CT36106.

22 GRAMINEAE

MORESBY ISLAND: 2n = 14, Upper Victoria Lake, CT35757;
2n = 14, Sunday Inlet, CT36587.

This Pacific northwestern species, with the closely related A. thurberiana
Hitchc, has been the subject of several taxonomic investigations of generic
placement {see Calder and Taylor, Part 1). Both A. aequivalvis and A. thur-
beriana have been found to have the same diploid chromosome number of
2n = 14.

66. Agrostis borealis Hartm.

GRAHAM ISLAND: 2n = 42, Skidegate Lake, CT23636.

MORESBY ISLAND: 2n = 42, Hotspring Island, CST22294;
2n = 42, Mt. de la Touche, CT23604, CT23607; 2n = 42, Mosquito Lake,
CT23759; 2n = 42, White Swan Bog, CT37409.

Many workers have made chromosome counts on this widely distributed
circumboreal species on material from eastern North America and Europe,
including the Kamchatka Peninsula. All the chromosome numbers reported
were n = 2% ov 2n = 56 {see Bowden 1960^, Sokolovskaja and Strelkova
1962, Sokolovskaja 1962, 1963, and Love and Love 1966). The different
chromosome number of 2n = 42 determined from Queen Charlotte Islands
plants is the first from the Cordillera of North America and indicates that there
are two chromosome races in this wide-ranging species. Further cytotaxonomic
investigations are needed on plants of this species from the southern portion
of the Cordilleran region.

67. Agrostis exarata Trin.

GRAHAM ISLAND: /i = 21, near mouth of Oeanda River, CT35888.

MORESBY ISLAND: 2n = 42, Hotspring Island, CST22310; 2n - 42,
Gray Bay, CST23435; 2n = ca. 42, Mt. de la Touche, CT23608; 2n = 21n,
Kootenay Inlet, CT36198.

This North American species has been counted from California by Steb-
bins and Love (1941) and from Mexico by Beaman et al. (1962). These
reports gave the chromosome number as n = 21. The species is probably
hexaploid based on x = 1 .

68. *Agrostis gigantea Roth

69. Agrostis pallens Trin.

70. *Agrostis palustris Huds.

71. Agrostis scabra Willd.

AGROSTIS 23

GRAHAM ISLAND: « = 21, llVi mi S of Masset, CT36909.
MORESBY ISLAND: n = 21, Skidegate Lake, CT36735.

Numerous counts of 2n = 42 have been made on Canadian and Alaskan
material by Bowden (1960a) and Bjorkman (1951). Sokolovskaja (1937,
1938) also reported In = 42 for this species.

72. Agrostis thurberiana Hitchc.

MORESBY ISLAND: 2n = 7n, between Cumshewa and Peel inlets,
CT35180.

The same chromosome number was obtained on the closely related A.
aequivalvis Trin. {see discussion under the latter species).

AiRA

73. *Aira caryophyllea L.

GRAHAM ISLAND: 2n = 28, between Skidegate and Skidegate Vil-
lage, CT35835.

Bocher and Larsen (1958a) have discussed the distribution and occur-
rence of the diploid and tetraploid chromosome races of this species found in
Europe. The tetraploid race is more widely distributed in southern Europe and
it has been introduced into northern Europe. The Queen Charlotte Islands
population belongs to the tetraploid race.

74. *Aira praecox L.

MORESBY ISLAND: In = 14, Tuft Islets, €7834872; 2n = In, Gray
Bay, CT35245.

The same chromosome number, n ^ 1 or 2n = 14, was obtained on
European material of this species by Hagerup (1939), Maude (1940), Bocher
and Larsen (1958a) and Hedberg and Hedberg (1961).

Alopecurus

75. *AIopecurus gcniculatus L.

GRAHAM ISLAND: 2n = 28, Mamin River Delta, CT35549.

All previous counts on this species arc 2n = 28 {see Love and Love
1961a).

24 GRAMINEAE

Ammophila

76. "^Ammophila arenaria (L.) Link

GRAHAM ISLAND: 2n = 14„, Tlell, CT35426.

This European species has been repeatedly counted as 2n = 28 {see
Love and Love 1961 a, Hedberg and Hedberg 1964, and Kubien 1964).

Anthoxanthum

77. "^Anthoxanthum odoratum L.

GRAHAM ISLAND: In = 20, between Queen Charlotte City and
Skidegate, CT36240.

MORESBY ISLAND: 2n = 20, Sandspit, CT35339.

The nature and origin of this tetraploid, 2n = 20, species from a cyto-
logical viewpoint has been recently reviewed by Jones (1964). He concludes
that A. odoratum is not an autotetraploid but a species of hybrid origin.

AVENA

78. *Avena fatua L.

Bromus

79. *Bronius mollis L.

GRAHAM ISLAND: 2n = 28, Queen Charlotte City, CST23000;
2n = 14„, 2 mi E of Queen Charlotte City, CTS34782.

All previous counts of this species are tetraploid 2n = 2S (see Schulz-
Schaeffer 1956, Bocher and Larsen l95Sa, and Bowden and Senn 1962).

80. Bromus pacificus Shear

GRAHAM ISLAND: 2n = 28, Naden Harbour, CT36848.

MORESBY ISLAND: 2n = 28, mouth of Deena River, CT23789;
2n = 28, Kootenay Inlet, CT36201; 2n = 28, Kaisun, CT36523.

A chromosome number of 2« = 28 for this species was recently reported
by Wilton (1965) from Western Alaska.

BROMUS 25

81. Bromus sitchensis Trin.

GRAHAM ISLAND: 2n = ca. 56, Torrens Island, CT35825.
MORESBY ISLAND: 2n = 28n, Sandspit, CT35327.

Previous counts of 2n = 42 (Stahlin 1929, and Schulz-Schaeffer 1960)
and In = 56 (Schulz-Schaeffer and Markarian 1957, and Schulz-Schaeffer
1960) were reported for this species. These counts were made on botanical
garden material. Our counts from the Islands indicate that the population is
octoploid.

Calamagrostis

82. Calamagrostis canadensis (Michx.) Beauv.

MORESBY ISLAND: 2n = 42, Skidegate Lake, CT35273; 2n = ca. 56,
Skidegate Lake, CT36054; 2n = 56, Mosquito Lake, CT36708.

Bowden (1960^) reviewed the counts that had been reported for this
species. After examination of 27 Canadian collections, he found 15 were hexa-
ploid, 2n = 42, and 12 were octoploid, 2n = 56. There appeared to be no
distinct correlation between either morphological characteristics or geographic
distribution and the differing ploidy levels. Similar conclusions were found in
the Queen Charlotte Islands population of this species.

83. Calamagrostis crassiglumis Thurb.

84. Calamagrostis nutkaensis (Presl) Steud.

GRAHAM ISLAND: 2n = 28, about 15 mi S of Masset, CT35578;
2n = 14„, Delkatla Inlet, CT35593; 2n = 28, about 4 mi N of mouth of
Oeanda River, CT35869; 2n = 28, Mayer Lake, CT36105; 2n = 28, Naden
Harbour, CT36837.

MORESBY ISLAND: 2n = 28, Kootenay Inlet, CT 36204; 2n = 28,
Kaisun, CT36522.

Queen Charlotte Islands plants of this species are tetraploid based on
X = 1. Calder and Taylor in Part I state that "there is no evidence that this
species hybridizes with C. canadensis as inferred by Hulten." The chromosome
number of C. nutkaensis, 2n = 28, and the numbers in C. canadensis, 2n = 42
and 56, certainly suggest that there is little likelihood of introgression between
these two species.

85. Calamagrostis purpurasccns R. Br. ssp. tasuensis Caldcr & Taylor
GRAHAM ISLAND: 2/7 = 28, near Jalun Lake, CT35644.

26 GRAMINEAE

MORESBY ISLAND: 2n - 28, Mt. Russ, CT36J92.

Previous counts made on the wide-ranging Calamagrostis purpurascens R.
Br. have revealed tetraploid, hexaploid, octoploid, and 12-ploid races {see
Bowden 1960/? and Sokolovskaja 1963). The Queen Charlotte Islands popu-
lation of this endemic subspecies is tetraploid based on x = 7.

CiNNA

86. Cinna latifolia (Trev.) Griseb. in Ledeb.

GRAHAM ISLAND: 2n = 28, Blackwater Creek, S3 529; 2n = 28,
Honna River, CT36976.

MORESBY ISLAND: 2n = 28, Mt. de la Touche, CT23609.

This wide-ranging uniform species has been repeatedly reported to have
a chromosome number of 2n = 2S (see Love and Love 1961a).

Cynosurus

87. *Cynosurus cristatus L.

GRAHAM ISLAND: « = 7, 2 mi E of Queen Charlotte City,
CTS34784.

Meiotic examination of this introduced grass revealed regular meiosis with
normal segregation at telophase I. All previous counts are diploid, 2n = \4
(see Love and Love 1961«).

Dactylis

88. *Dactylis glomerata L.

GRAHAM ISLAND: 2n = 28, Torrens Island, CT35826.

Dactylis glomerata has been the subject of detailed cytological investiga-
tion (see Stebbins and Zohary 1959, Jones in Beddows 1959, and Bocher
1961). Two well-established chromosome levels are present in the species,
diploid, 2n = 14, and tetraploid, 2n = 28. At least one of the limited number
of introduced populations on the Queen Charlotte Islands is tetraploid.

Danthonia

89. Danthonia californica Boland.

MORESBY ISLAND: 2n = 36, Mosquito Lake, CT36705.

DANTHONIA 27

Two previous counts on this western North American species have been
made on California material (Stebbins and Love 1941, and Wet 1954). Both
reports gave the chromosome number as tetraploid, 2« = 36.

90. Danthonia intermedia Vasey

GRAHAM ISLAND: 2n = 36, Yakoun Lake, CT36753.
MORESBY ISLAND: 2n =- 36, Mosquito Mtn., CT36439.

The same chromosome number was reported for this species by Packer
(1964) from Jasper National Park, Alberta, and by Wet (1954) from an
undisclosed location in North America. Sokolovskaja (1963) found diploids
with 18 somatic chromosomes in plants from the Kamchatka Peninsula.

Deschampsia

91. Deschampsia caespitosa (L.) Beauv. ssp. beringensis (Hult.) Lawr.

GRAHAM ISLAND: 2n = 26, Long Inlet, CT 36004.

MORESBY ISLAND: In = 13„, Gray Bay, CT35248; In = 26, Upper
Victoria Lake, CT35733; In = 26, Kootenay Inlet, CT36206; 2n = 26, Mt.
Moresby, CT36438; 2n = 26, Sunday Inlet, CT36590; 2n = 26, Yakulanas
Bay, CT36645; 2n = 26, Takakia Lake, CT37408.

A complete summary and discussion of the cytological aspects of Des-
champsia caespitosa (L.) Beauv. has been given by Kawano (1963, 1966).
The Queen Charlotte Islands population has a uniform, presumably derived,
tetraploid number of 2n = 26. Similar counts were reported on numerous
Canadian and American populations by Kawano (1963) and Lawrence
(1945).

92. Deschampsia elongata (Hook.) Munro ex Benth.

GRAHAM ISLAND: 2n = Hu, 2 mi E of Queen Charlotte City,
CTS34785; 2n = 26, 21/2 mi S of Tlell, CT35946.

MORESBY ISLAND: 2/t = 13„, Gray Bay, CT35249B.

Two previous counts have been made on this species. Stebbins {in Myers
1947) reported 2n - 26 for presumably western United States material and
Bowden ( 1960/?) reported 2n = 26 for material from southern British Colum-
bia. It is presumed that I), elongata represents a derived tetraploid based on
X = 1 {see Kawano 1963 for discussion of a similar situation in D.
caespitosa ) .

28 GRAMINEAE

Elymus
93a. Elymus glaucus Buckl. ssp. glaucus

93b. Elymus glaucus Buckl. ssp. virescens (Piper) Gould

GRAHAM ISLAND: 2n = 28, Torrens Island, CST22435; In - 28, 4
Jewell Island, CST22457; In = 28, Una Island, CST22915; 2n = 28, Tlell,
CT35918.

MORESBY ISLAND: 2n = 28, Hotspring Island, CST22273.

The taxonomic and cytological aspects of Elymus glaucus Buckl. and the
varieties glaucus and virescens have been discussed by Bowden (1964). All
counts reported by Bowden {op. cit.) and by Hartung (1946) are tetraploid,
2n = 28 based onx = 1.

94. Elymus hirsutus Presl

GRAHAM ISLAND: 2n = 28, Mamin River Delta, CT35553; 2n = 28,
Kumdis Creek Delta, CT36121.

MORESBY ISLAND: 2n = 28, Mt. de la Touche, CT23601.

The only counts made on this species have been on plants from British
Columbia. Bowden (1964) reported 2n = 28 on three populations, including
the count reported here from Mount de la Touche. The species is a tetraploid
based on jc = 7.

95. Elymus hirsutus Presl X Hordeum brachyantherum Nevski

MORESBY ISLAND: 2n = 28, head of Cumshewa Inlet, CT 36491.

The chromosome number 2n = 28 was reported for this material from
the Queen Charlotte Islands by Bowden (1967). The hybrid was named X
Elymordeum schaakianum by Bowden (1958) and he cited two specimens
from the Aleutian Islands, Alaska, including the holotype of his hybrid species
from Attn Island. The hybrid occurs at one other location on the Queen Char-
lotte Islands, at Kumdis Creek Delta on Graham Island. The anthers of the
hybrids from the Queen Charlotte Islands, like those from the Aleutian
Islands, had completely aborted pollen grains that remained inside the anthers.

96. Elymus mollis Trin. in Sprengel

GRAHAM ISLAND: 2n = 28, about 4 mi N of mouth of Oeanda River,
CT3585L

ELYMUS 29

This species has been counted many times from Iceland, Greenland,
Canada and the United States. The same chromosome number, n = 14 or
2n = 28, was reported in each case {see Bowden 1957).

Festuca

97. *Festuca arundinacea Schreb.

GRAHAM ISLAND: 2n = 42, near mouth of KHki Creek, CT3682L

The same chromosome number, n = 21 or 2n = 42, has been reported
by many authors (see Love and Love 1961(3, and Thomas 1962). Thomas
found 2n = 70 in native plants of this species from North Africa.

98. *Festuca dertonensis (All.) Asch. & Graebn.

MORESBY ISLAND: 2n = 14, Hotspring Island, CST22309B.

This weedy European annual grass has a diploid, 2n = 14, chromosome
number based on x = 7.

99. Festuca elatior L.

MORESBY ISLAND: 2n = 14, Copper Bay, CT36080; 2n = 14, near
Alliford Bay, CT36262.

Previous authors have obtained a diploid chromosome number of 2ai = 14
(see Love and Love 1961a). A hexaploid count was reported from Middle
East material by Sakamoto and Muramatsu (1962).

100. Festuca megalura Nutt.

101. *Festuca myuros L.

MORESBY ISLAND: 2n - 42, Mosquito Lake, CT3671L

Two chromosome races have been reported for this species (see Love
and Love 196 la). The diploid race, 2n = 14, has been recognized as Vidpia
(Festuca) myuros s. str., whereas the hexaploid race, 2n = 42, has been
recognized as Vulpia myuros (L). Gmel. var. major (Rohl.) B. de Lesd. by
Litardiere (1948) or as a separate species, Vulpia major (Rohl.) Love &
Love by Love and Love (1961c).

102. Festuca occidcntalis Hook.

30 GRAMINEAE

103. Festuca prolifera (Piper) Fernald

MORESBY ISLAND: 2n = ca. 70, Mt. Moresby, CT36418.

This species of Festuca is probably 10-ploid based on ;c = 7. See the
discussion of F. prolifera in Calder and Taylor Part 1 .

104. Festuca rubra L.

GRAHAM ISLAND: 2n = 21„, Tlell, CTS34640; In - 21„, Image
Pt., CTS34674; 2n = 21„, Masset Spit, CTS34729; 2n = 21„, Dawson Inlet,
CTS35138.

Many workers have studied this species cytologically and have reported
the numbers 2n = 14, 28, 42, 45, 53, 56 and 70 {see Love and Love 1961a).
The number most commonly found by previous workers is 2n = 42, the same
number was determined on all the populations sampled from the Queen Char-
lotte Islands. The uniformity of chromosome number on the Islands is of in-
terest considering the morphological variability that Calder and Taylor {Part
1 ) encountered in surveying this species on the Islands.

105. Festuca subulata Trin. in Bong.

GRAHAM ISLAND: n = 14, Honna River, CT35402; « = 14, 6 mi S
of Juskatla, CT35488.

MORESBY ISLAND: 2n = 28, Skidegate Lake, CT36075.

This species is tetraploid with the base number x = 1 . Material from
Juskatla showed configurations that varied from 2n = 12n + In to 14ii at
diakinesis.

Glyceria

106. Glyceria occidentalis (Piper) J. C. Nels.

GRAHAM ISLAND: 2n = ca. 40, AV2 mi S of Port Clements,
CT35469; 2n = 20„, 4 mi SW of Port Clements, CT35562; 2n = ca. 20, Ya-
koun Lake, CT36789.

Studies of the genus Glyceria by Church ( 1942, 1949) have revealed two
chromosome numbers in the section Euglyceria Griseb., 2n = 20 and 2/7 = 40.
He reported a chromosome number of 2n = 40 for Glyceria occidentalis
from northern California. Two chromosome races were found in this species
on the Queen Charlotte Islands. The paucity of counts known for this species,
as well as for other species of the section Euglyceria for the Pacific Northwest,
indicate that considerably more chromosome number information is needed to
clarify the occurrence of chromosome races in any one species of the genus.

HIEROCHLOE 31

HiEROCHLOE

107. Hierochloe odorata (L.) Beauv.

GRAHAM ISLAND: 2n = 56, Tlell, CT35929.

Many authors have examined this species cytologically and they have
reported three chromosome numbers, 2n = 28, 42 and 56 (see Love and
Love 1961a). Bowden (1960Z?) found that plants from five widely separated
locations in Canada had the chromosome number, 2n = 56, the same number
that we found for plants from the Queen Charlotte Islands.

HOLCUS

108. *HoIcus lanatus L.

GRAHAM ISLAND: 2n = In, Image Pt., CTS34678; 2n = 14,
CT37397; 2n = 14, Tlell, CT37423.

The chromosome number of this species has been consistently reported
by many workers as « = 7 or 2ai = 14 (see Love and Love 1961 a, Hedberg
and Hedberg 1961, and Carroll and Jones 1962).

HORDEUM

109. Hordeum brachyantherum Nevski

GRAHAM ISLAND: 2n = 28, Kumdis Creek Delta, CT36120;
2n = 28, Naden Harbour, CT36862.

Previous counts by Covas (1952), Rajhathy and Morrison (1959) and
Bowden (1962) were 2n = 28. See Part 1 for the discussion of the taxonomy
of this species.

LOLIUM

110. *Loliuni perenne L.

GRAHAM ISLAND: 2n = 7„, Tlell, CTS34653.

This species has been widely introduced throughout North America as a
constituent of lawn grass and has been investigated cytologically in Europe,
Asia and North America by many workers (see Love and Love 1961(3). All
chromosome numbers reported for L. perenne were n = 1 ox 2n ^ 14.

Melica

111. Melica subulata (Griseb.) Scribn.

32 GRAMINEAE

MORESBY ISLAND: 2n = 11, between Cumshewa and Peel inlets,
CT35205.

One chromosome count of 2a7 = 18 was reported for this species from
material collected in Plumas County, California (Stebbins and Love 1941).
Since the base number of the genus Melica is x =^ 9 (Love and Love 1961(3),
plants from California are diploid and those from the Queen Charlotte Islands
are triploid. The triploid count of 2n = 27 from the Islands suggests that this
population is apomictic.

Phalaris

112. ^Phalaris arundinacea L.

Phleum

113. Phleum alpinum L.

MORESBY ISLAND: 2n = 28, Mt. de la Touche, CT23611; 2n = 28,
Mosquito Mtn., CT23736; 2n = 14„, Takakia Lake, CT36319.

Two chromosome races, 2^7 = 14 and 2n = 28, are found in Phleum
alpinium s. lat. {see Love and Love 1961a). North American material counted
has been 2n = 28 {see Bowden 1960/?). Sokolovskaja (1963) reported
2n = 28 from Kamchatka.

114. *Phleum pratense L.

POA

115. *Poa annua L.

GRAHAM ISLAND: 2n = 28, Masset Spit, CST21267.

Many earlier counts have been made on this species and all have been
tetraploid, 2n = 28 {see Love and Love 1961(3).

116. Poa confinis Vasey

GRAHAM ISLAND: n = 2\, Tlell, CTS34637.

One previous count has been made on this species by Hartung (1946) on
western United States material. She obtained a chromosome number of
2n = 42. Our count was obtained by examination of mitosis in microgameto-
genesis.

117. Poa douglasii Nees ssp. macrantha (Vasey) Keck

GRAHAM ISLAND: 2n = 28, mouth of Sangan River, CST22765.

POA 33

The typical subspecies of this Pacific coast species has been counted as
2n = 28 by Stebbins and Love (1941) and Hartung (1946).

118. *Poa interior Rydb.

119. Poa laxiflora Buckl.

GRAHAM ISLAND: In = ca. 98, Tow Hill, CST22676.

Plants of this species from Tow Hill are probably 14-ploid with the base
number x = 1 . No previous counts have been made on this species.

120. Poa leptocoma Trin.

121. *Poa pratensis L.

GRAHAM ISLAND: In = 84, Langara Island, CST22563.

Many authors have counted plants of this species complex and nearly all
the chromosome numbers possible from In = 28 to 2n = 124 have been ob-
tained {see Love and Love \96\a). Our material from the Queen Charlotte
Islands is 12-ploid with the base number x = 1 .

Ill, Poa stenantha Trin.

MORESBY ISLAND: n = 42, Mt. Moresby, CT36417.

Meiotic examination revealed normal tetrad development, however, there
was some lagging and irregular pairing association in metaphase I. Hartung
(1946) reported 2a? = 81, 84, and 86 on material from western North Ameri-
ca. The report by Bowden (1961) oiln = 42 on plants from Yellowknife in
the Northwest Territories of Canada and attributed to P. stenantha should be
referred to P. interior Rydberg.

123. *Poa trivialis L.

GRAHAM ISLAND: 2n = Vu, Honna River, CT35401.
MORESBY ISLAND: 2n = lu, Gray Bay, CT35249A.

Two chromosome numbers have been reported for this species: a diploid,
2n = 14; and a tetraploid, 2n = 28 (see Love and Love 1961a). The North
American introductions are diploid on the basis of the few counts reported.

PUCCINELLIA

124. Puccincllia borcalis Swallen

GRAHAM ISLAND: 2n 42, Nadcn Harbour, CT 36864.

34 GRAMINEAE

MORESBY ISLAND: 2n = 42, Skedans Islands, CST22397.

The counts of 2n = 42 for Canadian material of P. borealis by Bowden
(1961) should be referred to P. interior S0rcnsen {see discussion in Calder
and Taylor, Part 1).

125. Puccinellia nutkaensis (Presl) Fern. & Weath.

GRAHAM ISLAND: 2n = 56, NW comer Graham Island, CST22629;
2n = 56, Masset Spit, CST22651; 2n = 28, 1 mi W of Queen Charlotte City,
CT35419; 2n = 42, Naden Harbour, CT 36874.

MORESBY ISLAND: 2n = 56, Gray Bay, CT35238.

The taxonomic relationships of this species to P. pumila are not clear
when plants are examined in their natural environment and they become even
more complex when only herbarium material is studied. The morphological
complexity is mirrored in our cytological investigations. Three chromosome
numbers have been obtained for the two species P. nutkaensis and P. pumila;
namely a tetraploid 2n = 28, a hexaploid 2n = 42 and an octoploid 2n = 56,
all based on .;c = 7. This is one group of grasses that will need detailed
biometrical and cytological investigations before the delimitation of species
can be successfully achieved on a biological basis.

126. Puccinellia pumila (Vasey) Hitchc.

GRAHAM ISLAND: 2n = 56, Queen Charlotte City, CST22433; 2n -
42, Delkatla Slough, CT36917.

MORESBY ISLAND: n = 21, Yakulanas Bay, CT 36644.
The discussion of these new counts will be found under Puccinellia nut-
kaensis.

TORREYOCHLOA

127. Torreyochloa pauciflora (Presl) Church

GRAHAM ISLAND: 2n = 14, Yakan Pt., CT36818.

Three diploid counts of 2n = 14 were made by Church (1949) on ma-
terial from California. Our studies on Queen Charlotte Islands material have
supported his findings and his proposal that the genus Torreyochloa with base
number x = 1 ht separated from that of Glyceria with x = 10.

Trisetum

128. Trisetum cernuum Trin.

GRAHAM ISLAND: 2n = 42, about 1 mi W of Queen Charlotte City,
CST22462; In = 42, Langara Island, CST22555; 2n = 42, Tow Hill,

TRISETUM 35

CST22696; n = 21, about 1 mi W of Queen Charlotte City, CTS34800;
2n = 42, about 2 mi W of Queen Charlotte City, CT34806.

The only previous chromosome count 2n = 42, was determined by
Stebbins and reported in Myers (1947). This count was presumably made on
plants from California and can undoubtedly be referred to T. cernuum Trin.
ssp. canescens (Buckl.) Calder & Taylor. For a discussion of the taxonomy of
this species see Part 1 .

129. Trisetum spicatum (L.) Richt.

Vahlodea

130. Vahlodea atropurpurea (Wahlenb.) Fr. ssp. paramushirensis (Kudo)
Hult.

GRAHAM ISLAND: 2n = 14, Takakia Lake, CT36311.
MORESBY ISLAND: 2/7 = 14, Kootenay Inlet, CT36153.

The same chromosome number was previously reported for other sub-
species by Nygren {in Love and Love 1948) and J0rgensen et al. (1958).

Cyperaceae
Carex

131. Carex anthoxanthea Presl

GRAHAM ISLAND: 2/7 = ca. 56, Jalun Lake, CT35617.
MORESBY ISLAND: 2/2 = 54, Upper Victoria Lake, CT35813.

This species and C. circinata C. A. Meyer are the only North American
members of the section Circinatae Meinsh. Examination of Queen Charlotte
Islands material of these two species revealed the chromosome numbers
In = 54 to 56.

132. Carex arcta Boott

MORESBY ISLAND: n = 30, Skidegate Lake, CT 36069.

The same chromosome number of 2/7 -- 60 was reported for the first
time on this species by Moore and Calder (1964) on material from Pontiac
County, Quebec.

133. Carex arcnicola Schmidt ssp. pansa (Bailey) Koyama & Calder

36 CYPERACEAE

134. Carex brevicaulis Mackenzie

GRAHAM ISLAND: 2n - 28, Towustasin Hill, CT35524.

The chromosome numbers of other species in the section Montanae Fries
are among the lowest known in the genus Carex (Wahl 1940). The chromo-
some number of this western North American species from the Islands falls
within the range of the chromosome numbers already reported for the section.

135. Carex brunnescens (Pers.) Poir. in Lam.

GRAHAM ISLAND: at = 28, 4 mi SE of Port Clements, CST22824A;
n = 27, 51/2 mi SE of Port Clements, CTS34590; In = ca. 56, llVi mi S of
Masset, CT36902.

Previous counts of either n = 28or2« == 56 have been reported {see
Love and Love 1961a). The same count was made on eastern North American
plants by Wahl (1940) and Love and Love (1966). It would appear that the
chromosome number for the species is n = 28. However, in examining ma-
terial from near Port Clements, 27 small spherical bodies were clearly seen.
There is a possibility that a quadrivalent was misinterpreted as a bivalent or a
variation in the chromosome number may exist in the species. Examination of
configurations at metaphase II revealed 28 bodies, one of them very small.

136. Carex buxbaumii Wahlenb.

137a. Carex canescens L. ssp. canescens

MORESBY ISLAND: In = 56, White Swan Bog, CT35298.

137b. Carex canescens L. ssp. arctaeformis (Mackenzie) Calder & Taylor

MORESBY ISLAND: 2n = 56, between Moresby and Aero logging
camps, CT35289; n =28, White Swan Bog, CT35299.

Earlier counts have been made on Carex canescens, at = 26 (Okuno
1939, 1940), AT = 27 (Wahl 1940) and 2n = 54 and 56 (Love and Love
1942a, 19566, 1966). These counts and the results of cytological examination
of plants from the Queen Charlotte Islands indicate that the chromosome num-
ber for this species is variable, ranging from at = 26 to /? = 28. There appears
to be no correlation between the cytological findings and morphological seg-
regation of infraspecific taxa.

138. Carex circinata C. A. Meyer

GRAHAM ISLAND: 2at = ca. 60, Long Inlet CT35982.
See discussion under C anthoxanthea Presl.

CAREX 37

139. Carex cusickii Mackenzie

140. Carex deweyana Schw. ssp. leptopoda (Mackenzie) Calder & Taylor

GRAHAM ISLAND: n = 27, 41/2 mi S of Port Clements, CT35044;
In = ca. 54, Mamin River Delta, CT35556.

The only previous count reported for C. deweyana was made by Wahl
(1940) on material from the eastern United States. He obtained a count of
n = 21.

141. Carex disperma Dewey

GRAHAM ISLAND: 2n = 70, Mamin River about 8 mi SSW of Jus-
katla, CT35484; 2n = 70, about 4 mi N of Oeanda River, CT35857.

The same chromosome number was obtained by Wahl (1940) and by
Moore and Calder (1964) from the Kenai Peninsula in Alaska. Love and
Ritchie (1966) reported 2n = 70 for plants of this species from Manitoba.

142. Carex enanderi Hult.

143. Carex exsiccata L. H. Bailey

144. Carex glareosa Wahlenb.

MORESBY ISLAND: 2n - 66, Kootenay Inlet, CT36202.

All previous counts for this species are 2n = 66 (see Love and Love
1961a).

145. Carex gmelinii H. & A.

146. Carex kelloggii W. Boott in S. Wats.

GRAHAM ISLAND: n = 44, near Tow Hill, CST22659.

Previous counts in the section Acutae Fries by Wahl (1940) indicate a
range of numbers from ^7 = 33 to 38.

147. Carex laeviculmis Meinsh.

GRAHAM ISLAND: a? = 28, 3 mi E of Juskatla, CTS35048; 2n - 56,
Mamin River Delta, CT35555.

Other counts in the section Stellulatae Kunth. reported by Wahl (1940)
were n = 22,n = 26 and n = 27.

38 CYPERACEAE

148. Carex leptalea Wahlenb. ssp. pacifica Calder & Taylor

MORESBY ISLAND: 2n = 52, White Swan Bog, CT35303; In^ca. 50,
Upper Victoria Lake, CT35727; 2n = 52, Mosquito Lake, CT367 16.

Love and Ritchie (1966) reported the same chromosome number for
material of the typical subspecies from Manitoba.

149. Carex livida (Wahlenb.) Willd.

GRAHAM ISLAND 2n = 52, Jalun Lake, CT35655; 2n = 50, Upper
Victoria Lake, CT35737.

The taxonomic position of the infraspecific taxa in Carex livida has been
discussed by Calder and Taylor in Part 1. Love and Love (1944a, 1956a)
reported 2n = 32 for this species, as did Moore and Calder (1964). However,
according to Moore (personal communication), they observed the chromo-
some number 2/7 = ca. 54, not 2n ^ 32, in material of C. livida from Ontario.

150. Carex lyngbyei Homem.

151. Carex macloviana d'Urv. ssp. pachystachya (Cham.) Hult.

GRAHAM ISLAND: n = 38, Torrens Island, CST22441; n = 38, near
Yakoun River Delta, CST23499; n = 38, Tlell, CT35943.

MORESBY ISLAND: n = 38, between Skidegate Lake and Copper Bay,
CT35261.

Previous cytological studies on this species were made on plants of North-
ern Europe {see Love and Love \96\a). The numbers reported for European
plants give a somatic number of approximately 86. Our studies on plants from
the Queen Charlotte Islands indicate that the western North American sub-
species pachystacha is cytologically distinct. Future taxonomic studies, pro-
posed by Calder and Taylor in Part 1 , should include not only morphological
investigations but also cytological studies so that an attempt can be made to
correlate the different chromosome races now known to exist in the species.

Examination of plants from Torrens Island revealed that meiosis was
irregular and one metaphase plate showed a configuration of 35n + 2i + In-.

152. Carex macrocephala Willd. in Spreng.

GRAHAM ISLAND: n - 39, Tlell, CTS35074; 2n = 74, Tlell,
CT S3 464 3.

153. Carex macrochaeta C. A. Meyer

GRAHAM ISLAND: 2n - 60, Long Inlet, CT35983.

CAREX 39

One previous count of 2n = 60 was made by Moore and Calder (1964)
on material from southern British Columbia. The problem of whether or not
this species should be placed in section Limosae Tuckerm. or section Atratae
Kunth., as discussed by Calder and Taylor in Part 1, is not clarified by the
available cytological information because chromosome numbers range between
2n = 54 and 62 for both sections.

154. Carex mertensii Prescott in Bong.

GRAHAM ISLAND: « = 31, 41/2 mi S of Port Clements, CT35043;
2n = 62, near Juskatla, CT35554.

The only previous count for this species was made by Moore and Calder
(1964) on material from the Kenai Peninsula of Alaska. They reported a
somatic chromosome number of 2n = 62.

155. Carex nigricans C. A. Meyer

MORESBY ISLAND: 2n = ca. 72, Takakia Lake, CT 36286.

156. Carex obnupta Bailey

157. Carex pauciflora Lightf.

GRAHAM ISLAND: 2n = ca. 74, 3 mi E of Juskatla, CTS35045.

158. Carex phyllomanica W. Boott in S. Wats.

GRAHAM ISLAND: 2n = 54, 4 mi N of mouth of Oeanda River,

CT35858.

MORESBY ISLAND: 2n = ca. 54, Kootenay Inlet, CT 36177.

Although Calder and Taylor in Fart 1 consider Carex phyllomanica to be
one of the most distinct species in section Stellulatae Kunth., the chromosome
number obtained on Queen Charlotte Islands material, 2n = 54, differs from
that reported for plants from Kenai Peninsula, Alaska, 2/7 = 70, by Moore
and Calder (1964). Three species that are placed in this section are reported
to have haploid numbers of 22, 26 and 27 by Wahl (1940).

159. Carex physocarpa Presl

160. Carex pluriflora Hult.

GRAHAM ISLAND: 2n - ca. 50, 3 mi E of Juskatla, CST35047.

161. Carex pyrenaica Wahlenb. ssp. micropoda (C. A. Meyer) Hult.

40 CYPERACEAE

162. Carex scirpoidea Michx.

MORESBY ISLAND: 2n = 62, Takakia Lake, CT36292.

Several chromosome numbers have been reported for this species,
2n = 68 (Heilbom 1939), 2n = 62 (J0rgensen et al 1958, Love and Love
1964, 1966, and Love and Ritchie 1966) and 2n = 64 (Moore and Calder
1964). Somatic chromosomes ranged from small spherical bodies to oval
bodies of about three times the volume.

163. Carex sitchensis Prescott in Bong.

164. Carex stylosa C. A. Meyer

165. Carex tracyi Mackenzie

GRAHAM ISLAND: 2n = 62, near Yakoun River Delta, CT35465.

166. Carex viridula Michx.

MORESBY ISLAND: 2n = 70, Upper Victoria Lake, CT35758;
2n = 70-72, Kootenay Inlet, CT36174.

J0rgensen et al. (1958) reported 2n = ca. 70 for plants of this species
from Greenland, and Moore and Calder (1964) observed 2n = ca. 72 in
somatic material of this species from Hastings County, Ontario.

Eleocharis

167. Eleocharis acicularis (L.) R. & S.

MORESBY ISLAND: n ^ 10, 2ai = 20, Skidegate Lake, CT36065.

Previous counts have all been 2n = 20 or n = 10 {see Love and Love
1961a). Harms (1964) recently reported n = 10 from material collected in
Kansas. The only other North American count was made by Hicks (1929) but
the numbers reported, n = 15-19 and 2n = 56, indicate that a misinterpre-
tation was made of unusual behavior exhibited by this genus at the conclusion
of meiosis (^ee Strandhede 1965).

168. Eleocharis kamtschatica (C. A. Meyer) Komarov

GRAHAM ISLAND: 2n = 12, about 21/2 mi E of Tow Hill, CST22737;
2n = ca. 12, near mouth of Oeanda River, CT35906.

169. Eleocharis macrostachya Britt. in Small

MORESBY ISLAND: 2n = 8„, 2n = 16, White Swan Bog, CT23666.

ELEOCHARIS 41

The diploid chromosome number of 2n = 16 for plants from the Queen
Charlotte Islands is the same number as that found by Strandhede (1958) for
plants of Scandinavia. Subsequent research was conducted by Strandhede
(1960) on the correlation of the two chromosome numbers 2n = 16 and
2/1 = 38, known for the Scirpus palustris (=Eleocharis palustris) complex.
He was able to morphologically compare Linnaean specimens with specimens
of known chromosome number. As a result, Strandhede has chosen a lectotype
for Scirpus palustris (^^Eleocharis palustris) and has shown that S. palustris
belongs to the In = \6 group. On the basis of Strandhede's research, the taxon
on the Queen Charlotte Islands would be best treated as Eleocharis palustris, a
proposal that Calder and Taylor made in Part 1.

170. Eleocharis obtusa (Willd.) Schultes

MORESBY ISLAND: n = 5„, 2« = 10, Skidegate Lake, CT36063.

The same chromosome number was reported for North American material
of this species by Lewis et al. ( 1 962 ) and Harms ( 1 964 ) .

Eriophorum

171. Eriophorum angustifolium Honck.

172a. Eriophorum chamissonis C. A. Meyer ex Ledeb.
172b. Eriophorum chamissonis X Eriophorum russeolum

Rhynchospora

173. Rhynchospora alba (L.) Vahl

MORESBY ISLAND: n=U, Sunday Inlet, CT36593.

Two chromosome numbers are reported for European material of this
species, « - 13 or 2« = 26 by Scheerer (1939, 1940) and Gadella and
Kliphuis (1963), and Al - 21 or 2/1 = 42 by Love and Love (1942^).

Scirpus

174. Scirpus atrocinctus Femald

175. Scirpus cernuus Vahl

GRAHAM ISLAND: 2ai - 30„, Kumdis Creek Delta, CT23802.

176. Scirpus cespitosus L.

42 CYPERACEAE

177. Scirpus lacustris L. ssp. glaucus (Smith) Hartm.

178. Scirpus sylvaticus L. ssp. digynus (Boeckl.) Koyama

GRAHAM ISLAND: 2n = 32„, Yakoun River about AV2 mi S of Port
Clements, CT S3 5032.

Two chromosome numbers have been reported for Scirpus sylvaticus L.
Hakansson (1928) reported 2n = 62 for Swedish material and Ehrenberg
(1945) found 2n = 64 on plants from Skane, Sweden. Our count from the
Queen Charlotte Islands represents the first meiotic count made on this species
and the first reported for the western North American subspecies of this wide-
ranging species.

Araceae

Lysichiton

179. Lysichiton americanum Hult. & St. John

GRAHAM ISLAND: 2n = 28, Tlell, CTS34648.

The same chromosome number was obtained on cultivated western North
American material of this species by Love and Kawano (1961 ). These authors
discussed the relationship of L. americanum to L. camtschatcense (L.) Schott.

Juncaceae

JUNCUS

180. Juncus alpinus Vill. ssp. nodulosus (Wahlenb.) Lindm.

MORESBY ISLAND: 2n = 20„, Mosquito Lake, CT35311.

J0rgensen et al. (1958) reported 2n = 40 for material from Greenland
and gave a brief discussion of the complexity in the /. alpinus Vill. aggregate.
Vaarma and Love and Love reported 2n = 80 {in Love and Love 1948) for
this subspecies.

181a. Juncus arcticus Willd. ssp. sitchensis Engelm.

GRAHAM ISLAND: 2n = 40„, Lepas Bay, CST22619; n = 40, Daw-
son Inlet, CST35139.

Earlier counts for Juncus arcticus Willd. are 2n = 80 {see Love and Love
1961fl, and Love and Ritchie 1966) and 2n = 74 (Sokolovskaja and Strel-
kova 1948a).

JUNCUS 43

181b. Juncus arcticus Willd. ssp. ater (Rydb.) Hult.

182. Juncus articulatus L.

GRAHAM ISLAND: 2n = 40„, 4 mi NW of Tlell, CT35458; n = 40,
2 mi NW of Tlell, CT35692.

All earlier counts were 2n = 80 for this species {see Love and Love
1961fl). The report by Wulff (1937fl, 1938) of In = ca. 60 and n = ca. 30,
respectively, should be referred to another species.

183. *Juncus bufonius L.

GRAHAM ISLAND: n = 17, Delkatla Inlet, CT35584.
MORESBY ISLAND: n = 17, Gray Bay, CT35244.

Many authors have published chromosome numbers for this species and
they have reported 2n = 30, 60, 80, 104, 106 and 120 {see Love and Love
1961a). The chromosome number for plants from the Queen Charlotte Islands
differs greatly from those previously reported for this species.

184. Juncus drummondii E. Meyer in Ledeb.

185a. Juncus effusus L. var. brunneus Engelm.

185b. Juncus effusus L. var. gracilis Hook.

MORESBY ISLAND: n = 40, Alliford Bay, CST23234.

Previous counts made on Juncus effusus L. were 2n = 40 (Love and
Love 1944b, and Sasaki 1937).

185c. Juncus effusus L. var. pacificus Fern. & Wieg.

186. Juncus ensifolius Wikstr.

GRAHAM ISLAND: 2n - 40, Blackwater Creek, CT35058; 2n = 40,
2 mi S of Cape Chroustcheff, CT37351 .

MORESBY ISLAND: 2n - 40, between Sandspit and Copper Bay,
CST23195; n = 20, 2n - 40, Mosquito Lake, CT3680L

Snogerup (1963) reported 2n = 40 for material from the Aleutian
Islands. The plants from the Queen Charlotte Islands showed the two small
chromosomes in each complement that were noted by Snogerup in his study
of the section Ensifolii (Rydb.) Snogerup.

44 JUNCACEAE

187. Juncus falcatus E. Meyer

MORESBY ISLAND: 2n = 38, Mosquito Lake, CT 36706.

Snogerup (1963) reported counts for two other species in the section
Graminijolii (Buch.) Vierh. as 2n = 38. One of these, /. covillei Piper, is
considered to be closely related to /. falcatus.

188. Juncus filiformis L.

MORESBY ISLAND: n - 40, E end of Skidegate Lake, CT35264.

This chromosome number agrees with that of Wulff (1937a, 1938) for
German material and of Love and Love (1944/?, 1956a) and J0rgensen et al.
(1958) for Scandinavian plants. Vaarama (in Love and Love 1948) reported
finding 2n = 40 for material from Finland. Love and Love (1966) reported
2n = 80 for plants of this species from Mount Washington in the eastern
United States.

189. Juncus leseurii Boland.

190. Juncus mertensianus Bong.

MORESBY ISLAND: n = 20, between Cumshewa and Peel inlets,
CT35169; n = 20, Takakia Lake, CT36322.

In a recent review of chromosome numbers in the genus Juncus by Sno-
gerup (1963), the base number oi x = 20 is proposed for the section Septati
(Buch.) Vierh. On the basis of this proposal, the Queen Charlotte Islands
material of /. mertensianus is diploid.

191. Juncus oreganus S. Wats.

MORESBY ISLAND: n = ca. 50-60, between Aero and Moresby log-
ging camps, CT35285; n = ca. 30, Upper Victoria Lake CT35762; n = ca.
50-60, Skidegate Lake Bridge, CT36060; 2n = ca. 112, 2 mi S of Cape
Chroustcheff, CT37350.

Meiosis in the material examined was very irregular, consequently, we
were unable to make an accurate count in pollen mother cells. The mitotic
chromosome number given is also only approximate because of the large num-
ber of small chromosomes in the plates studied.

192. *Juncus tenuis Willd.

GRAHAM ISLAND: 2n = 40, 7 mi N of Port Clements, CT37473.

JUNCUS 45

Two previous counts have been reported for this species. Sasaki (1937)
reported In = 32 whereas Love and Love (1948) reported 2n = 30. Snoger-
up (1963) gives a base number of x = 20 and 21 for the section Genuini
(Buch.) Vierh. to which /. tenuis belongs.

193. Juncus triglumis L.

MORESBY ISLAND: In = 22„, Upper Victoria Lake, CT35802.

Previous counts on this species do not agree with the chromosome num-
ber obtained on plants from the Queen Charlotte Islands. Love and Love
(1944b) reported 2n = 50 for plants from Iceland, and Holmen (1952) and
J0rgensen et al. (1958) found 2n = ca. 134 and 2n = ca. 130 for plants from
Greenland. J0rgensen et al. (op. cit.) did not consider it likely that two chro-
mosome numbers, 2n = 50 and 2n = ca. 130, exist within /. triglumis and
suggested that the specimen counted by Love and Love ( 1 944^ ) should be
placed in another species. The chromosome count oi n = 22 for Queen Char-
lotte Islands material supports the conclusion of Calder and Taylor in Part 1
that, based on morphology, plants from the Islands should not be referred to
/. albescens (Lange) Femald, a taxon based on a specimen collected in Green-
land. However, there are several chromosome races in the /. triglumis complex,
which suggests that this group needs a thorough biosystematic study before its
taxonomy can be clarified.

LUZULA

194a. Luzula multiflora (Retz.) Lejeune

GRAHAM ISLAND: 2n = 36, Langara Island, CST22570; 2n = 36,
Tlell, CST23164.

MORESBY ISLAND: 2n = 12, Mt. de la Touche, CT23602.

194b. Luzula multiflora (Retz.) Lejeune ssp. comosa (E. Meyer) Hult.

GRAHAM ISLAND: 2n = 24, between Skidegate and Skidegate Village,
CST21398; 2n = 24, Queen Charlotte City, CST22480.

MORESBY ISLAND: 2n = 24, near Alliford Bay, CST21850;
In = 24, East Copper Island, CST22257; 2n = 24, Hotspring Island,
CST22285; 2n = 24, Limestone Island, CST22415.

The taxonomy of the Luzula multiflora complex is not well understood
and the disposition of the taxa on the Queen Charlotte Islands has been the
subject of considerable discussion by* Calder and Taylor in Part I. The low-
land subspecies comosa has been recognized as a distinct entity and the same
chromosome number of 2n ^ 24 has been obtained on all populations sam-
pled. The only previous counts made on this taxon were 2n = \2 and 24 by

46 JUNCACEAE

Nordenskiold (1951). The disposition of the montane taxon (taxa) is not
clear. Two chromosome numbers have been found, 2« = 12 and 2n = 36.
The cytology corroborates the findings of Calder and Taylor that the popula-
tions of Luzula multifiora that cannot be referred to ssp. comosa represent a
complex that cannot be clearly distinguished as a distinct infraspecific taxon.
Previous counts of L. multifiora reported are based on .;c = 12, but variation
in ploidy levels range from diploid to octoploid (see Love and Love 1961a).
The disposition of the western North American segregants of this complex
must await a detailed systematic study including cytological analyses.

195. Luzula parviflora (Ehrh.) Desv.

MORESBY ISLAND: 2n = 24, between Aero and Moresby logging
camps, CT36513.

Earlier counts for this species complex have been reported as 2n = 24
(see Love and Love 1961<3, 1966, and Packer 1964). It is unfortunate that
populations of the alpine and the lowland phases described in Part 1 were not
obtained for cytological analyses because such analyses might have decided
whether variation in chromosome number correlates with variation in mor-
phology between the two entities. The plants counted belong to the lowland
phase of this species.

Liliaceae
Allium

196. *Alliuin schoenoprasum L.

Fritillaria

197. Fritillaria camschatcensis (L.) Ker-Gawl.

GRAHAM ISLAND: 2n = 12n, Tlell, CTS35082.
MORESBY ISLAND: 2n = 24, Tuft Islets, CTS34862.

North American material from Washington has been reported to have
n = 12 (Omduff and Kruckeberg 1957). The Queen Charlotte Islands popu-
lation is also diploid, based on x = 12.

Matsuura (1935) reported diploid counts for a depauperate alpine form
from Japan and triploid counts for the robust lowland coastal entity from
northern Japan and the Kuriles. In 1963, Matsuura and Toyokuni recognized
the alpine diploid as a new subspecies, F. camschatcensis (L.) Ker-Gawl. ssp.
alpina Matsuura & Toyokuni. The lowland triploid was recognized as the typi-
cal subspecies based on the original species description given by Linnaeus for
Kamchatka plants. The plants from the Queen Charlotte Islands are robust
and would be classified as the typical subspecies; however, our plants are

FRITILLARIA 47

diploid and viable seed was obtained from several populations sampled on the
Islands. On the basis of our observations, we propose that the typical sub-
species has two chromosome races, but segregation of the two lowland
chromosome races on morphological bases cannot be made.

Lloydia

198. Lloydia serotina (L.) Reichenb. ssp. flava Calder & Taylor
GRAHAM ISLAND: 2n = 24, Jalun Lake, CT35646.

MORESBY ISLAND: 2n = 24, Yatza Mtn., CT35715; In = 24, Taka-
kia Lake, CT36266.

The chromosome number obtained on Queen Charlotte Islands material
of ssp. flava, In = 24, was also reported for the typical subspecies in Europe
and Asia by Sokolovskaja and Strelkova (1960) and by other workers (see
Love and Love 1961fl).

Maianthemum

199. Maianthemum dilatatum (Wood) Nelson & Macbr.

GRAHAM ISLAND: 2n = 36, 4 mi N of Oeanda River, CT35862.
MORESBY ISLAND: In = ca. 36, Upper Victoria Lake, CT35810.

The chromosome number «=18or2« = 36 was reported by Therman
(1956) on material from botanical gardens, by Palmgren (1943) on plants
from Alaska, and by Sokolovskaja (1963) on material from Kamchatka.
Sato (1942) reported In = 54 for the species based on plants grown from
seed from a botanical garden. Although there are two chromosome races in
this species, it is impossible to determine where the two races occur because
of the paucity of counts and the fact that some of the previous chromosome
number determinations were on botanical garden plants of an undisclosed
origin.

Streptopus

200. Streptopus amplexifolius (L.) DC.

GRAHAM ISLAND: 2n = 32, 1 mi W of Queen Charlotte City,
CTS34797; 2n = ca. 32, Dawson Inlet, CTS35085; 2n = 32, Long Inlet,
CT35951; 2n = 32, near mouth of Kliki Creek, CT36825.

MORESBY ISLAND: 2n = 32, Crescent Inlet, CT3498I; 2n - 32,
Kootenay Inlet, CT36227.

Many authors have reported the same chromosome number, 2/7 32,
for North American, Asiatic and European material of this species (see Love
and Love I96\a, Love and Harries 1963, and Sokolovskaja 1963).

48 LILIACEAE

201. Streptopus roseus Michx. ssp. curvipes (Vail) Hult.

GRAHAM ISLAND: 2n - ca. 48, Collinson Lake, CT35516; 2n = 48,
Jalun Lake, CT3561 1 .

MORESBY ISLAND: 2n = 48, Yatza Mtn., CT35749.

The only chromosome counts on this species were reported by Therman
(1956) and by Love and Harries (1963) as In = 16 for material from botan-
ical gardens in Sweden and the northeastern United States, respectively.
Queen Charlotte Islands material is hexaploid, with the base number x = ^.
On the basis of the two counts made on North American material, there is an
apparent cytological distinction between the western subspecies curvipes and
the diploid eastern North American taxon.

202. Streptopus streptopoides (Ledeb.) Frye & Rigg ssp. brevipes (Baker)
Calder & Taylor

GRAHAM ISLAND: In = 16, about 8 mi SSW of Juskatla, CT35479.

This species is diploid with a base number of j: = 8.

TOFIELDIA

203a. Tofieldia glutinosa (Michx.) Pers. ssp. glutinosa

MORESBY ISLAND: In = 30, White Swan Bog, CT35295; 2n = 30,
Upper Victoria Lake, CT35764.

203b. Tofieldia glutinosa (Michx.) Pers. ssp. brevistyla C. L. Hitchc.

GRAHAM ISLAND: 2n = 30, Yakoun Lake, CT36769.
MORESBY ISLAND: 2n = ca. 30, Red Mud Marsh, CT36725.

Eariier counts reported for the genus were 2n = 28 (Miller 1930),
2n = 30 (Sato 1942, Love and Love 1961a, and Taylor and Brockman 1966)
and 2n = 60 (Sato 1942). Cave (1966) obtained n = 15 for material of
ssp. brevistyla from Oregon. The two subspecies on the Islands are diploid,
based on a: = 15.

Veratrum
204. Veratrum eschscholtzii A. Gray

GRAHAM ISLAND: « = 16, Honna River, CT35405.

MORESBY ISLAND: 2n = 32, Upper Victoria Lake, CT35806;
2n = 32, Takakia Lake, CT36363.

VERATRUM 49

The same chromosome number n ^ \6 or In = 2>2, was previously
reported for North American material of this species by Love and Love

(1965) and by Taylor and Brockman (1966). See discussion of the taxonomy
of V. viride Ait. and V. eschschoitzii in Calder and Taylor, Part 1, and in Tay-
lor and Brockman (1966).

Iridaceae

SiSYRINCHIUM

205. Sisyrinchium littorale Greene

GRAHAM ISLAND: 2n = ca. 96, Juskatla, S3517.

MORESBY ISLAND: n = 48, Skidegate Lake, CT35279; In = 48„,
Mosquito Lake, CT35304.

This coastal species is closely related to the Sisyrinchium montanum
Greene aggregate. A recent cytotaxonomical study on Sisyrinchium by Bocher

(1966) gives a chromosome number of In = 96 for this group. The coastal
S. littorale is probably 12-ploid based on jc = 8.

Orchidaceae

Calypso

206. Calypso bulbosa (L.) Oakes ssp. occidentalis (Holz.) Calder & Taylor

CORALLORHIZA

207. Corallorhiza maculata Raf. ssp. mertensiana (Bong.) Calder & Taylor

GRAHAM ISLAND: In = 20ii, near junction of Yakoun River and
Ghost Creek, Cri557 (9.

The previous base number of a' = 7 for the genus Corallorhiza was based
on counts of 2n = 42 on C. trifida Chat, (see Love and Love 1961a) and C.
innata R. Br. (Miduno 1940). No previous counts have been made on North
American members of the genus. The meiotic configurations at metaphase I
clearly showed 20 pairs and we are proposing that x = 10 be an additional
base number for this genus.

GOODYERA

208. Goodyera oblongifolia Raf.

GRAHAM ISLAND: n 15, Haida Pt., CT36679; n 15, about
IVi mi S of Jungle Beach, CT3674L

50 ORCHIDACEAE

The base numbers for this genus are x = 11, 14 and 15 (see DarHngton
and Wyhe 1955). The material from the Queen Charlotte Islands is diploid
and belongs to the group of species that have the base number x = 15.

Habenaria

209. Habenaria chorisiana Cham.

GRAHAM ISLAND: 2n = 42, Jalun Lake, CT35668.
MORESBY ISLAND: 2n = 21„, Takakia Lake, CT36277.

The Queen Charlotte Islands population is diploid based on jc = 21.

210. Habenaria dilatata (Pursh) Hook.

MORESBY ISLAND: n = 2\, 2n = A2, Upper Victoria Lake,
CT35729.

The only other counts made on this species, 2n = 42 (Humphrey 1934,
and Love and Love 1966), were also from North America. The species is pre-
sumably diploid based on x = 21. Regular meiosis was observed in our ma-
terial.

211. Habenaria saccata Greene

MORESBY ISLAND: n = 21, Bigsby Inlet, CT 34880; 2n = 42, Upper
Victoria Lake, CT35788; n = ca. 21, Kootenay Inlet, CT36161; 2n = 21ii,
Takakia Lake, CT36300.

The Queen Charlotte Islands population is diploid based on x = 21.
Meiosis was regular with normal development of microspores.

212a. Habenaria unalascensis (Spreng.) S. Wats. ssp. unalascensis

212b. Habenaria unalascensis (Spreng.) S. Wats. ssp. maritima (Greene)
Calder & Taylor

GRAHAM ISLAND: 2n = 21n, Torrens Island, CT35820; n = 21,
HaidaPt, Cr5665a

The Queen Charlotte Islands population is diploid based on x = 21.
Meiotic examination revealed regular segregation and production of tetrads.

LiSTERA

213. Listera caurina Piper

LISTERA 51

GRAHAM ISLAND: 2n = 34, Long Inlet, CT35970; 2n = 34, Koote-
nay Inlet, CT36224.

MORESBY ISLAND: 2n = 17„, Anna Inlet, CT34940.

Several base numbers, including x = 17, have been proposed for the
genus. The Queen Charlotte Islands populations are diploid, based on :t = 17.

214. Listera cordata (L.) R. Br. in Ait.

GRAHAM ISLAND: 2n = 38, 9 mi N of Port Clements, CTS34708;
2n = ca. 38, about 4 mi N of mouth of Oeanda River, CT35875.

MORESBY ISLAND: 2n = 38, Upper Victoria Lake, CT35807;
2n = ca. 38, Kootenay Inlet, CT36222.

Queen Charlotte Islands plants all had a diploid number oi 2n = 38,
based on x = 19. Chromosome numbers of 2n = 38, 40 and 42 have been
reported for this species {see Love and Love 1961a, Kliphuis 1963, and Shoji
1963). Love and Love (1966) reported 2n = 36 for plants from Mount
Washington in eastern United States.

Malaxis

215. Malaxis paludosa (L.) Sw.

GRAHAM ISLAND: n = 14, about 3 mi NW of Tlell, CT36111.

Meiotic examination showed normal meiosis and regular segregation at
anaphase I with subsequent formation of normal tetrads. The same chromo-
some number has been reported for this species by Hagerup (1944) and
Kliphuis (1963).

Spiranthes

216. Spiranthes romanzoffiana Cham. & Schlecht.

GRAHAM ISLAND: 2n = 15„, Masset Spit, CT 36920.

Cytological examination of material from the Queen Charlotte Islands
reveals that the species is diploid based on x = 15. Heslop-Harrison {in Love
and Love 196 la) reported a tetraploid count of 2n = 60 for material from
Northern Ireland. The question of the taxonomy of these two widely separated
populations should perhaps be reconsidered on the basis of this new evidence.

Salicaceae

Salix

217. Salix hookcriana Barratt in Hook.

52 SALICACEAE

218. Salix lasiandra Benth.

219. Salix reticulata L. ssp. glabellicarpa Argus

220. Salix scouleriana Barratt in Hook.

GRAHAM ISLAND: 2n = ca. 1 14, IVi mi E of Masset, CT35605.

The determination of exact mitotic counts on polyploid species of Salix
are extremely difficult because of the small size and variable morphology of
the chromosomes. This coastal species of the Charlottes is presumably a hexa-
ploid based onx = 19.

221. Salix sitchensis Sanson in Bong.

GRAHAM ISLAND: 2n = 38, Mamin River Delta, CT35538.

This species is widely distributed along the Pacific slope of the Cordilleran
region and is diploid with the base number x = \9.

Myricaceae

Myrica

222. Myrica gale L.

MORESBY ISLAND: 2n = ca. 96, Skidegate Lake, CT35147.

European material of this species has the chromosome number n = 24 or
2n = 48 according to Hagerup (1941) and Hakansson (1955). Since species
of the genus Myrica have the base number x = S (Darlington and Wylie
1955), European plants of M. gale are hexaploid. Queen Charlotte Islands
plants of the same species are 12-ploid based on jc = 8, the highest level of
polyploidy known for a species of the genus Myrica.

Betulaceae

Alnus

223. Alnus crispa (Ait.) Pursh ssp. sinuata (Regel) Hult.

MORESBY ISLAND: 2n = 28, Mt. Moresby, CT37547.

There are too few counts on this species from North America to make
any generalizations about chromosome numbers for the species. Love and
Love (1964) reported 2n = 28 for Alnus crispa (Ait.) Pursh from Manitoba.
Our count from the Queen Charlotte Islands represents the first report of the
Pacific coast race of the wide-ranging ^4 . crispa.

ALNUS 53

224. Alnus rubra Bong.

GRAHAM ISLAND: 2n = 28, 21/2 mi SE of Port Clements, CT34621;
2n = 28, Long Inlet, CT35965.

MORESBY ISLAND: 2n = 28, Crescent Inlet, CTS34978; In = 28,
Bigsby Inlet, CTS34896; In = 28, Anna Inlet, CT37426.

Only two earlier counts have been made on this species and both were
reported as 2n = 28 (Gram et al. 1941, and Wetzel in Darlington and Wylie
1955).

Moraceae

HUMULUS

225. "'"Humulus lupulus L.

Urticaceae

Urtica

226. *Urtica dioica L.

GRAHAM ISLAND: 2n = 52, Queen Charlotte City, CST23006.

Two chromosome numbers, 2n = 48 and 2n = 52, have previously been
reported for European material of this species (see Love and Love 196 la,
Sorsa 1962, and Gadella and Kliphuis 1963).

Loranthaceae

Arceuthobium

227. Arceuthobium campylopodum Engelm. in Gray.

Polygonaceae

OXYRIA

228. Oxyria digyna (L.) Hill

GRAHAM ISLAND: n ^1, Long Inlet, CT35957.
MORESBY ISLAND: 2n = In, Mount Moresby, CT 36388.

The same chromosome number has been obtained on Canadian, Europ-
ean and Asiatic material of this species by many workers. Recent counts have
been reported by J0rgensen et al. (1958), Wcislo in Skaiinska et al. (1959),
Mooney and Billings (1961), Sokolovskaja and Strclkova (1962), Sorsa
(1963), Packer (1964), and Love and Love (1966). Mciotic examination of
plants from Graham Island revealed regularity at telophase I.

54 POLYGONACEAE

Polygonum

229. ^Polygonum aviculare L.

GRAHAM ISLAND: 2n = 30„, Tlell, CT35915; In = 30„, Delkatla
Inlet, CT36916.

MORESBY ISLAND: 2n = 20n, Sandspit, CT 36019.

Two chromosome counts, tetraploid and hexaploid, are reported by Love
and Love (196 let) in their compendium for the P. aviculare complex. Doida
(1962) and Styles (1962) report 2/7 = 60 for plants of this complex. Styles
(1962) recognizes four species in this complex, two of which, P. aviculare s.
sir. and P. arenastrum Bor., are widely distributed and well established in
North America. In Britain, he found that P. aviculare s. str. always has 60
somatic chromosomes and P. arenastrum the number 2n = 40. Mertens and
Raven (1965) consider P. arenastrum to be much commoner in North
America than P. aviculare s. str. They found 2n = 40 for plants of P. arenas-
trum from California and Indiana and 2n = 60 for two collections of P.
aviculare s. str. from California. However, they state that "they would have
been most difficult to determine as P. aviculare had their chromosome number
not been determined." Both tetraploid and hexaploid populations of the P.
aviculare complex occur on the Queen Charlotte Islands. Based strictly on
chromosome number, these plants would be P. arenastrum and P. aviculare s.
str., respectively. However, all specimens examined cytologically would prob-
ably be classified as P. arenastrum, using the characters given by Styles
(1962), but the fruits vary considerably in the aspect of the sides, and all
fruits have rows of punctate dots. There seems little doubt that the morpho-
logical characters given by Styles for British material cannot always be cor-
related with chromosome number of American plants. We are therefore sup-
porting the decision of C alder and Taylor in Part 1 in referring all plants from
the Islands to the collective name P. aviculare.

230. *Polygonum convolvulus L.

MORESBY ISLAND: 2n = 20„, Sandspit, CT36011, CT37013.

This chromosome number was previously reported for Canadian and
European material of this species {see Mulligan 1961).

231. Polygonum fowleri B. L. Robinson

GRAHAM ISLAND: n = 20, Masset Spit, CT35705.

Love and Love (1956/?) reported the same chromosome number,
2n = 40, on material from the east and west coasts of Canada. Meiosis in
plants from Graham Island was regular at telophase II.

POLYGONUM 55

232. ^Polygonum persicaria L.

GRAHAM ISLAND: 2n = 22n, Delkatla Inlet, CT36918.

The same chromosome number has been reported for European material
by Jaretzky (1927, 1928fl), Andersson in Love and Love (19426) and
Pauwels (1958). Doida (1960, 1962) has reported 2n = 40 from mitotic
examination of Japanese material.

233. ^Polygonum polystachyum Wall.

GRAHAM ISLAND: 2n = 11„, betv^een Queen Charlotte City and
Skidegate Village, CT36947.

This garden escape is sometimes confused with P. cuspidatum Sieb. &
Zucc. According to Jaretzky (1928^), P. cuspidatum has the chromosome
number n = ca. 44.

234. ^Polygonum scabrum Moench

MORESBY ISLAND: 2n = lln, Sandspit, CT 37004.

The same chromosome number was reported on North American and
European material of this species {see Mulligan 1961 ).

235. Polygonum viviparum L.

MORESBY ISLAND: 2n = > 100, Takakia Lake, CT 36326.

The chromosome number obtained on the Queen Charlotte Islands is
similar to chromosome numbers previously obtained on European and other
Canadian material. It is difficult to count the many small chromosomes
of this species either in mitosis or meiosis. Previous counts reported are:
n = ca. 55 (Jaretzky 1928a), 2n = ca. 88, n = 44, 2n = 83-88 (Sokolov-
skaja and Strelkova 1938), 2ai = ca. 100, n = ca. 50 (Flovik 1940),
2/2 = ca. 110, /I = ca. 55 (Love and Love 1948), 2/7 = ca. 132 (Skalinska
1950), 2/z =>100 (Sokolovskaja and Strelkova 1960), 2n = 100 ± 10
(Mosquin and Hayley 1966), and 2/7 = 120 (Love and Love 1966, and
Love and Ritchie 1966).

Rum EX

236. *Rumex acetosella L.

GRAHAM ISLAND: 2/7 42, Queen Charlotte City, CST22485,
CST23J59; 2/7 = 42, Tlcll, CT 37461.

56 POLYGONACEAE

This is the same chromosome number reported for this material from
two locations in Ontario, one in Saskatchewan, and three on the mainland of
British Columbia (Mulligan 1959). Love (1941, 1944) placed the chromo-
some races of the R. acetosella complex in the following species: R. angiocar-
pus Murbeck {2n = 14), R. tenuijolius (Wallr.) Love {In = 28), R.
acetosella L. s. str. (2n = 42), and R. graminifolius Lambert {2n = 56).
However, apparently the only morphological character separating R. angio-
carpus from R. acetosella s. str. is the fusion of the valve to the seed in the
former and the nonfusion of valves to seed in the latter species. Queen Char-
lotte Islands material and some of the plants from elsewhere in Canada have
the valves fused to the seed and 42 somatic chromosomes. It is because of
this confusion that Calder and Taylor used the aggregate name R. acetosella
for the plants from the Islands in Part 1 .

237. *Rumex crispus L.

GRAHAM ISLAND: In = 30„, Jungle Beach, CT35449; n = 30, west
of Queen Charlotte City, CT36938.

MORESBY ISLAND: 2n = 30n, Sandspit, CT35333.

Many counts on this species have been made on North American, Asian,
and European material (see Mulligan 1957). In all reports the chromosome
number of.2n = 60 was given.

238. '''Rumex obtusifolius L.

GRAHAM ISLAND: 2n = 20n, between Skidegate and Skidegate Vil-
lage, CT35840.

MORESBY ISLAND: 2n = 40, Sandspit, CT35355.

Earlier mitotic and meiotic examinations on Canadian, European and
Asiatic material of this species have revealed the same chromosome numbers
(^^6 Mulligan 1959).

239. Rumex occidentalis S. Wats.

MORESBY ISLAND: 2n = ca. 140, Anna Inlet, CTS34926.

Two other counts of this native Rumex species are known. Mulligan
(1957) reported 2n = ca. 140 from Saskatchewan plant and Love and
Ritchie (1966) reported 2n = 200 for plants from Manitoba.

240. Rumex transitorius Rech. f .

GRAHAM ISLAND: 2n = 20, Jungle Beach, CST23398.

MORESBY ISLAND: 2n = 20, near Alliford Bay, CST23230;
2n = 20, head of Cumshewa Inlet, CT23648; 2n = 20, Gray Bay, CT35287.

RUMEX 57

Sarkar (1958) and Mulligan (1959) reported the same chromosome
number for plants from the coastal region of British Columbia.

Chenopodiaceae

Atriplex

241. Atriplex patula L. ssp. obtusa (Cham.) Hall & Clements.
MORESBY ISLAND: 2n = 11 u. Gray Bay, CT35254.

This is the first chromosome count on ssp. obtusa and the first indication
of a hexaploid in A. patula L. Earlier chromosome numbers reported for A.
patula are 2n = 18 (Kjellmark 1934 and Witte 1947) and 2n = 36 (Winge
1917, Love and Love 1956fl, and Hulme 1958). Flora Europaea by Tutin et
al. (1964) lists only 2/i = 36 for /i. patula and 2« = 18 for the closely re-
lated A . hastata L.

The hexaploid chromosome number reported for the western North
American ssp. obtusa supports the taxonomic segregation of this entity from
typical A. patula. Extensive morphological and cytological investigation of the
three Cordilleran subspecies recognized by Hall and Clements and discussed
by Calder and Taylor in Part 1 may lead to the recognition of separate species
for this complex.

Chenopodium

242. "^Chenopodium album L.

MORESBY ISLAND: In = 54, Sandspit, CST23216.

There are several chromosome numbers given in the literature for Cheno-
podium album, however, there is good reason to believe that C. album has
only one chromosome number, 2n = 54. This is the same chromosome num-
ber obtained on material from the Queen Charlotte Islands.

Cole (1962) reports 2n = 54 for material of C. album from Great
Britain, France, Switzerland, Germany, Denmark, Italy, New Zealand, Aus-
tralia, U.S.A., and Canada. Cole states that the other chromosome numbers
reported for this species are errors arising from the taxonomic misidentification
of the original material used, mistakes easily made in this taxonomically diffi-
cult genus.

Salicornia

243. Salicornia pacifica Standley

GRAHAM ISLAND: 2n = 18,i, 1 mi W of Queen Charlotte City,
CST22432; n = 18, Masset Spit, CT35699.

MORESBY ISLAND: 2n^ 18,,, Moresby logging camp, CST21991.

58 CHENOPODIACEAE

The only chromosome numbers that have been reported for other species
of this genus are either diploid, In ^ 18, or tetraploid. In = 36 {see Love
and Love 1961a).

SUAEDA

244. Suaeda depressa (Pursh) S. Wats.

GRAHAM ISLAND: In = 11 n, 1 mi W of Queen Charlotte City,
CT35421; 2n - 27n, Tlell, CT35938; 2n = 27„, Naden Harbour, CT36870;
2n = 27ii, Masset, CT36919.

Hulten (1944) considered Pacific coast plants to be S. maritima (L.)
Dumort, a cosmopolitan species described from Europe. Calder and Taylor in
Part 1 found that western prairie and Pacific plants tend to have seeds ranging
from 1.0 to 1.3 mm long, whereas the eastern Canadian and European popu-
lations have seeds approximately 1,5 to 2.2 mm long. They state that the
flowers of the western plants tend to be smaller and have more prominently
corniculate perianth lobes than those of eastern Canada and Europe. They
decided to tentatively refer both inland and coastal British Columbia plants
to the western species S. depressa. The chromosome number obtained on
Queen Charlotte Islands material {n = 27) and reports of the chromosome
number 2n = 36 for European and Asiatic plants of S. maritima (Jinno 1956
and other workers ) tend to support this position.

Nyctaginaceae

Abronia

245. Abronia latifolia Eschsch.

GRAHAM ISLAND: 2n = 44-46„, Tlell, CT35428.

The small meiotic chromosomes and the difficulties in squashing caused
by the presence of raphides in the anther walls made examination of the
material difficult.

Portulacaceae
Montia

246. Montia fontana L.

247. Montia parvifolia (Moc.) Greene

248. Montia sibirica (L.) Howell

GRAHAM ISLAND: ^7 = 12, Tlell, CTS34656; 2n = 12„, Tow Hill,
CTS34753; 2n = 6„, 1 mi W of Queen Charlotte City, CTS34795; 2n = 12„,
N end of Dawson Inlet, CT37346; 2n = 12„, 12 mi N of Port Clements,
CT37453; 2n = 12„, 2 mi W of Queen Charlotte City, CT 37481.

MONTIA 59

MORESBY ISLAND: 2n = 6„, Anna Inlet, CTS34922; 2n = 12n,
Bigsby Inlet, CT37464.

According to Lewis (1962, 1963) M. sibirica has tetraploid and hexa-
ploid races in California. It has been introduced into Great Britain where it
is tetraploid according to Anderson (1963). On the Queen Charlotte Islands
both diploid and tetraploid races occur; the tetraploid race is more common.
It would be interesting to establish the distribution of the three chromosome
races in the native range of this species and to determine whether or not only
one of the chromosome races has been introduced into Great Britain.

Caryophyllaceae

Arenaria

249. Arenaria lateriflora (L. ) Fenzl

250. Arenaria peploides L. ssp. major (Hook.) Calder & Taylor

GRAHAM ISLAND: 2n = 34„, Torrens Island, CT35830.

Meiotic examination of this species revealed regular formation of 34 bi-
valents at metaphase I, with regular segregation leading to normal telophase I
configurations. Mailing (1957) reported n = 34 and 2n = 68 on European
material of A. peploides s. lat. under Honckenya peploides (L. ) Ehrh. Other
chromosome numbers obtained from European material are 2n = 66 (Flovik
1940), n = 2A and 32 (Rohweder 1939) and 2/7 = 70 (Sokolovskaja and
Strelkova 1962).

251. Arenaria stricta Michx. ssp. macra (Nels. & Macbr. ) Maguire

MORESBY ISLAND: 2/7 - 12ii, between Cumshewa and Peel inlets,
CT35172.

Cerastium

252. Cerastium arvense L.

GRAHAM ISLAND: 2/7 = ISn, Towustasin Hill, CT35518.
MORESBY ISLAND: /? - 18, Limestone Island, CTS34830.

Plants on the Islands arc tetraploid, based on .v = 9, and form 18 biva-
lents at metaphase I. This is a cosmopolitan species widely distributed in
temperate regions. In Europe two chromosome races, tetraploid and octo-
ploid, are known to occur (Brett 1952, Blackburn and Morton 1957, and
others). Plants from Limestone Island were regular at telophase I.

60 CARYOPHYLLACEAE

253. Cerastium fischerianum Ser. in DC.

254. Cerastium tomentosum L.

255. *Cerastiuin viscosum L.

GRAHAM ISLAND: 2n = 36„, Tlell, CTS34645.

This European adventive has the same chromosome number on the
Queen Charlotte Islands as has been obtained on European material by Larsen
(1960) and many other workers.

256. *Cerastium vulgatum L.

GRAHAM ISLAND: 2n = 70„, 3 mi N of Lawnhill, CST21726;
2n = 70n, Langara Island, CST22571; 2n = ca. 72n, Honna River,
CT35409.

Blackburn and Morton (1957) reported 2n = 72, 126, 144 and 180 for
European material of this species. The number 2n = 144 was also obtained
on European plants by Mattick {in Tischler 1950), Sollner (1952, 1954),
Brett (1953) and Love and Love (1956fl). Brett (1955) examined selfed
and F2 plants from seed originally collected in Sussex, England, and obtained
chromosome numbers ranging from 136 to 152 with 136 the most frequent
number. Brett (1955) also studied Fi plants collected from Mickleham, Eng-
land, and obtained numbers ranging from 137 to 147, with 2n = 144 the
most common number. She states that plants of this species are difficult to
study because multivalents are present. She considers it probable that different
numbers of chromosomes pass to each pole. Plants from the Queen Char-
lotte Islands were either n = 10 or probably 72. In all cases only bivalents
were seen at metaphase I.

Sagina

257. Sagina maxima A. Gray

GRAHAM ISLAND: 2n = 33n, Dawson Inlet, CTS35144.
MORESBY ISLAND: 2n = 33„, Kootenay Inlet, CT 36221.

The Queen Charlotte Islands material is hexaploid based on x = 11.
Mizushima (1960fl) reported 2n = 42 or 44 for the same species from an
unstated locality in Japan. In all likelihood his material is a tetraploid with
2n = 44 and a base number of 1 1 .

258. *Sagina procumbens L.

MORESBY ISLAND: 2n = lln, between Cumshewa and Peel inlets,
CT35173.

SAGINA 61

This widespread North Temperate species has a uniform chromosome
number of 2n = 22 based on examination of European, North American and
Asiatic material by Blackburn and Morton (1957), Mizushima (1960^), and
others.

SiLENE

259. Silene acaulis L. ssp. subacaulescens (F. N. Williams) Hitchc. & Maguire

260. '''Silene noctiflora L.

Spergula

261. *Spergula arvensis L.

GRAHAM ISLAND: In = 9n, Queen Charlotte City, CST23806.

This adventive from Europe has the same chromosome number on the
Queen Charlotte Islands as it has in Europe, 2n = 18 (Blackburn and Mor-
ton 1957, and others).

Spergularia

262. Spergularia canadensis (Pers. ) G. Don

GRAHAM ISLAND: 2n = 36, Shields Bay, CT23383; n = 18, Jus-
katla, S3515.

Plants from Iceland (Love and Love 1956fl) and Europe (Ratter 1964)
have the same chromosome number.

Stellaria

263. Stellaria calycantha Bong.

MORESBY ISLAND: 2n = 13„, Takakia Lake, CT36357.

European material of this species has the chromosome numbers 2n = 52
(Love and Love 1956a) and 2n = 44-48 (Peterson 1936). A recent count
by Love and Love (1966) on plants from Mount Washington was reported as
2n = 56, which indicates that a tetraploid race of this species occurs in eastern
North America. Plants from the Queen Charlotte Islands arc diploid, based on
jc = 13. The count by Love and Love ( 1956a, 1966) on this species indicates
there are diploids and tctraploids in the S. calycantha complex. It seems likely
that the plants examined by Peterson were probably tetraploid, 2n = 52, and
the base number in this species complex is a' = 13.

62 CARYOPHYLLACEAE

264. Stellaria crispa Cham. & Schlecht.

GRAHAM ISLAND: 2n = 13„, IVi mi SE of Port Clements,
CTS34600; 2n = 13„, Dawson Inlet, CTS35128.

On the basis of our new counts on S. crispa, three of our four native
species of Stellaria on the Islands can now be considered to have the same base
number of x = 13.

265. *SteIlaria graminea L.

GRAHAM ISLAND: n = 26, Queen Charlotte City, CT36516.

This adventive from Europe has the same chromosome number on the
Queen Charlotte Islands as it has in Europe (Blackburn and Morton 1957,
and other workers). Plants from the Islands had regular meiosis at telophase I.

266. Stellaria humifusa Rottb.

GRAHAM ISLAND: 2n = 13n, Kumdis Creek Delta, CT36127.

Plants on the Queen Charlotte Islands have the same chromosome num-
ber as those from northern Europe and Greenland (Flovik 1940, S0rensen
and Westergaard in Love and Love 1948, Love and Love 1956fl, and J0rgen-
scnetal. 1958).

267. Stellaria longipes G oldie

268. *Stellaria media (L.) Vill.

GRAHAM ISLAND: 2n = 20n, Tlell, CrS35080.
MORESBY ISLAND: 2n = 20n, Kaisun, CT36537.

This introduced weed has the same chromosome number on the Islands
that was reported for this species from Ontario, Canada (Mulligan 1961 ). The
same and different counts of 2/7 = 28, 42 and 44 have been reported by other
workers from Europe and Asia {see Peterson 1936, Gyorffy 1940, and Pal
1953). It is interesting that only one chromosome race has been found in
North America.

Nymphaeaceae

NUPHAR

269. Nuphar luteum (L.) Sibth. & Sm. ssp. polysepalum (Engelm.) Beal

GRAHAM ISLAND: /i = 17, 10 mi N of Port Clements, CTS34713;
2n = 34, 4 mi NW of Tlell, CT3546L

NUPHAR 63

MORESBY ISLAND: 2n = 34, Upper Victoria Lake, CT35790.

These counts represent the first chromosome determinations made on
North American material. The same number was obtained on European
material of other subspecies of N. luteum (L.) Sibth. & Sm. by Rosenberg
(1909), Langlet and Soderberg (1927) and Heslop-Harrison (1953).

Ranunculaceae

ACONITUM

270. Aconitum delphinifolium DC.

MORESBY ISLAND: 2n = 16, Takakia Lake, CT36317.

Material of this species from the alpine Takakia Lake was diploid with
16 somatic chromosomes. This count establishes the presence of two chromo-
some races in this species. The only previous chromosome count was 2n = 32
by Sokolovskaja (1963) on plants of A. delphinifolium from Kamchatka.
Morphological discontinuities in the species and the recognition of them as
subspecific entities is discussed by Calder and Taylor in Part 1 .

ACTAEA

271. Actaea rubra (Ait.) Willd. ssp. arguta (Nutt.) Hult.

Anemone

272. Anemone multifida Poir. in Lam.

MORESBY ISLAND: 2n = 32, Limestone Island, CTS34814.

Heimburger (1959) and others have obtained the same chromosome
number on North and South American populations of this species.

273. Anemone narcissiflora L. ssp. alaskana Hult.

GRAHAM ISLAND: 2n = 14, Jalun Lake, CT35636.

The same chromosome number was obtained on European and Asiatic
material of this species by Kurita (1955) and others. The count on the Queen
Charlotte Islands material represents the first report for this subspecies. The
chromosomx count of Sokolovskaja and Strelkova (1948/?) of 2ai = 16 may
be in error.

274. Anemone parviflora Michx.

64 RANUNCULACEAE

Aquilegia

275. Aquilegia formosa Fisch.

GRAHAM ISLAND: n = 1, between Skidegate and Skidegate Village,
CTS34664.

MORESBY ISLAND: « = 7, South Low Island, CTS34845; 2n = 14,
South Low Island, CT37434.

The chromosome number determined on Queen Charlotte Islands
material is apparently the first documented report for this species. The Vascu-
lar Plants of the Pacific Northwest report 2n = 14 for the species but no docu-
mentation is given. In Darlington and Wylie (1955), 2n = 14 is reported for
A. formosa under the name A. truncata Fisch. & Mey. but there is no indica-
tion by Skalinska ( 1931 ) of the source of the plant examined.

Caltha

276. Caltha biflora DC.

GRAHAM ISLAND: 2n = 48, Blackwater Creek, CTS35073.

MORESBY ISLAND: 2n = 48, Bigsby Inlet, CT S3 4988; 2n - 48, Sun-
day Inlet, CT36607.

The counts obtained on Queen Charlotte Islands material indicate C.
biflora is a hexaploid with the common base number for the genus .x = 8. The
closely related C. leptosepala DC. is also a hexaploid.

277. Caltha palustris L. ssp. asarifolia (DC.) Hult.

COPTIS

278. Coptis asplenifolia Salisb.

MORESBY ISLAND: 2n = 18, between Cumshewa and Peel inlets,
CT35171; 2n = 18, Kootenay Inlet, CT36155.

This is the third species of Coptis to be counted. All members of the
genus studied cytologically have a base number x = 9 and are diploid.

279. Coptis trifolia (L.) Salisb.

GRAHAM ISLAND: 2n = 9„, about 6I/2 mi SE Port Clements,
CT34615.

All previous counts reported for this species are 2n = IS (Langlet 1932,
Kurita 1958, Sokolovskaja 1963, and Love and Love 1966).

ISOPYRUM 65

ISOPYRUM

280. Isopyrum savilei Calder & Taylor

GRAHAM ISLAND: n = l,2n = 14, Long Inlet, CT35963.
MORESBY ISLAND: 2n = 7„, Takakia Lake, CT36330.

This endemic species is restricted to the central montane region on the
Islands. It is diploid based on x = 7 as are other members of the genus count-
ed. Meiosis was regular with 7 bivalents formed at metaphase I and normal
segregation at telophase I.

Ranunculus

281. '''Ranunculus acris L.

GRAHAM ISLAND: 2n - 7„, Image Pt., CTS34691.

Previous counts on this introduced North American weed were also
2n = 14. Many similar counts have been made on European material, but
there have been some counts that differ, for example 2n = 12, 16, 28 and 56
(see Mulligan 1959). It appears from the large number of counts made on
this species that x = 1 is the most common base number and most populations
are diploid. Only diploids have been found in North America.

282. Ranunculus aquatilis L.

GRAHAM ISLAND: 2n = 32, Collinson Lake, CT 35537.

MORESBY ISLAND: 2n = ca. 32, Skidegate Lake, CJ 35274;
2n = 16„, Skidegate Lake, CT35309.

The R. aquatilis — R. circinatus group of the subgenus Batrachium is
taxonomically complex in the Old and New World. Diploids, tetraploids and
hexaploids, based on jc = 8, are known in this group. Only tetraploids occur on
the Queen Charlotte Islands.

283. Ranunculus cooleyae Vasey & Rose

GRAHAM ISLAND: 2n = 16, Jalun Lake, CT35635.

MORESBY ISLAND: 2n = 16, Mt. Russ, CT36I42; 2n ^ 16, Takakia
Lake, CT36308.

This distinctive North American Cordilleran montane species is diploid,
based on jc = 8.

66 RANUNCULACEAE

284. Ranunculus eschscholtzii Schiccht.

MORESBY ISLAND: 2n = 32, Mt. Moresby, CT36404; In = 32,
Mosquito Mtn., CT3646L

This variable species is tetraploid on the Queen Charlotte Islands. Other
members of the polymorphic section Epirotes (Prantl) L. Benson from North
America that have been counted are diploid.

285a. Ranunculus flammula L. var. flammula

MORESBY ISLAND: 2n = 16„, between Copper Bay and Skidegate
Lake, CT35258; 2n = 32, W of Moresby logging camp, CT37331.

285b. Ranunculus flammula L. var. filiformis (Michx.) Hook.

GRAHAM ISLAND: In = 32, Yakoun Lake, CT36762.
MORESBY ISLAND: 2n = 32, Skidegate Lake, CT35265.

285c. Ranunculus flammula L. var. ovalis (Bigel.) L. Benson

GRAHAM ISLAND: 2« = 32, Yakoun River, 16 mi S of Juskatla,
CT35498; 2n = ca. 32, near mouth of Oeanda River, CT35900.

Calder and Taylor in Part 1 tentatively recognized three varieties in R.
flammula on the Queen Charlotte Islands. All three varieties have the same
chromosome number, n = 16 or 2« = 32. Plants that Calder and Taylor
would place in R. flammula have been examined cytologically by many workers
(Sokolovskaja and Strelkova 1960, and others) and all had the chromosome
number n = 16 or 2n = 32.

286. Ranunculus hyperboreus Rottb.

GRAHAM ISLAND: 2n = 16„, 2n = 32, about 4 mi N of mouth of
Oeanda River, CT35871; 2n = 32, Wi mi W of Yakan Pt, CT36819.

This species of aquatic or damp terrestrial habitats is tetraploid, based on
jc = 8, on the Islands. The same chromosome number was obtained on Euro-
pean and Asian material of the species {see J0rgensen et al. 1958, Sokolov-
skaja and Strelkova 1960, 1962).

287. Ranunculus occidentalis Nutt. ssp. occidentalis

GRAHAM ISLAND: 2n = 28, Dawson Inlet, CTS35135; 2n = 28,
Image Pt., near Skidegate, CST20872.

MORESBY ISLAND: 2n = 28, Copper Bay, CST21906; 2n = 28, islet
off Bolkus Islands, CST22223; 2n = 28, South Low Island, CTS34842;
2n = 28, between Sandspit and Cape Chroustcheff, CT35160.

RANUNCULUS 67

288. Ranunculus orthorhynchus Hook.

GRAHAM ISLAND: n = 16, Tlell, CTS34651; In = 32, Yakoun
River Delta, CT35464.

289. Ranunculus pygmaeus Wahlenb.

290. *Ranunculus repens L.

GRAHAM ISLAND: n = \6, 2n = 32, Image Pt., CTS34692;
In = 32, Image Pt., CT35384.

This species, introduced into North America, is tetraploid on the Islands,
based on x == 8. It is extremely variable morphologically in the Old World,
where it is known to have at least two chromosome races, diploid and tetra-
ploid {see Love and Love 1961a, and Sokolovskaja 1963).

291. Ranunculus uncinatus D. Don in G. Don

GRAHAM ISLAND: 2n = ca. 28, Queen Charlotte City, CTS34798.

MORESBY ISLAND: 2n = 14„, Crescent Inlet, CTS34980; In = 28,
Sandspit, CT35356.

This species is a tetraploid on the Islands with a base number of x = 7.
It is interesting that this species and R. occidentalis Nutt., both previously un-
counted, have the same ploidy level and the same base number. Both species
belong to the section Chrysanthe (Spach) L. Benson.

Thalictrum

292. Thalictrum alpinum L.

MORESBY ISLAND: n = 1, Mosquito Mtn., CT 36448.

This circumboreal species reaches its most southern limit in the Pacific
Northwest on the Queen Charlotte Islands. Plants of this species from Europe
and Asia are also diploid, based on x = 1 (Sokolovskaja 1963, and many
other workers).

Cruciferae

Arabis

293. Arabis glabra (L.) Bcrnh.

MORESBY ISLAND: In - 6,,, Kaisun, CT36526.

68 CRUCIFERAE

Arabis glabra, native to North America and Eurasia, has several chromo-
some numbers. The n = 6 number obtained on material from the Queen
Charlotte Islands was also found in plants of this species elsewhere in Canada
(Mulligan 1964) and from Denmark, England and Canada by Bocher and
Larsen (1955). Manton (1932) and Mattick {in Tischler 1950) reported
2« = 16 and Jaretzky (1928/?) reported 2n = 32 for European plants of this
species.

294. Arabis hirsuta (L.) Scop. ssp. eschscholtziana (Andrz.) Hult.

GRAHAM ISLAND: 2n = 32„, Tow Hill, CTS34754.

Arabis hirsuta is native both to North America and Eurasia. This large-
flowered subspecies of the Pacific coast of North America is octoploid, based
on X = 8. Rollins (1941) also found octoploid plants of A. hirsuta in Utah.
Mulligan (1964) reported the presence of tetraploid plants from northern
Canada, and Rollins (1941) reported tetraploids in Colorado. Diploids (Mat-
tick in Tischler 1950, Baksay 1957, Novotna 1962, and Titz 1964) and
tetraploids (Jaretzky 1928/?, Novotna 1962, and Titz 1964) have been found
in Europe.

295. Arabis lyrata L. ssp. kamchatica (Fisch.) Hult.

MORESBY ISLAND: 2n = 32, Mt. Moresby, CT36389.

The same chromosome number. In = 32, was obtained on material of
this western subspecies of A . lyrata from Yukon Territory and British Colum-
bia by Mulligan (1964). Rollins (1966) reported 2/i = 16 for plants of ssp.
kamchatica from Alaska. Plants of the more eastern race of A. lyrata from
Saskatchewan (Mulligan 1964), Wisconsin (Smith 1938) and Connecticut
(Rollins 1941) were all diploid, In = 16.

Barbarea

296. Barbarea orthoceras Ledeb.

GRAHAM ISLAND: 2n = 8ii, Tow Hill, CTS34756.

MORESBY ISLAND: 2n = 16, South Low Island, CTS34847;
2n = 16, Kaisun, CT 37490.

Queen Charlotte Islands material of this native species had the same
chromosome number, 2n == 16, as previously reported by Mulligan (1964)
and Rollins (1966) on material from northern British Columbia and Cali-
fornia, respectively.

BRASSICA 69

Brassica

297. *Brassica campestris L.

MORESBY ISLAND: 2n = 10„, Sandspit, CT 36051.

This introduced weed has the same chromosome number, n = \0,m the
Queen Charlotte Islands as it has elsewhere in North America, Europe and
Asia (Mulligan 1959, and others).

Cakile

298. Cakile edentula (Bigel.) Hook.

MORESBY ISLAND: 2n = 9„, Sandspit, CT35336,

Material of this species from Vancouver Island, Ontario and Quebec
(Mulligan 1964) and California (Kruckeberg 1948) had the same chromo-
some number (« = 9) as Queen Charlotte Islands plants. Pobedimova (1963)
places the Pacific population of this species under C. calijornica Heller, Atlan-
tic populations under C. edentula and the Great Lake population under C
lacustris (Fern.) Pobed. The morphological characters she uses to distinguish
the three populations are obscure and unreliable. It is interesting to note that
plants from these three areas are diploid (Mulligan 1964).

299. *Cakile maritima Scop.

GRAHAM ISLAND: « = 9, Tlell, CT35430.

This introduced species has the same chromosome number {n = 9) in
the Queen Charlotte Islands as it has in Europe and Asia (Skalinska et al.
1961, and others).

Camelina

300. * Camelina sativa (L.) Crantz

Capsella

301. "^Capsella bursa-pastoris (L. ) Medic.

MORESBY ISLAND: In = 32, Sandspit, CT35I68.

Capsella bursa-pastoris is a widely distributed adventivc in North America
with the same chromosome number as found in Europe, In ^ 32 {see Mulli-
gan 1957).

70 CRUCIFERAE

Cardamine

302. Cardamine an^ulata Hook.

GRAHAM ISLAND: 2n - 20,,, Image Pt., CTS34681; In = 40, Ya-
koun River bridge, about 41/2 mi S of Port Clements, CTS35030; In = 40, 16
mi S of Juskatla, CT35497.

This chromosome number was previously reported for Queen Charlotte
Islands plants by Mulligan ( 1965<^). Pollen mother cells examined from plants
collected at one location formed 20 bivalents at metaphase I and subsequent
telophase stages were regular. Mulligan suggested that C. angulata is almost
certainly an allotetraploid species with the base number x = 10. Cardamine
angulata was probably derived from a diploid species with the common base
number of Cardamine, x == 8, and a tetraploid with the base number x ^ 6.

303. Cardamine bellidifolia L.

MORESBY ISLAND: 2n = 8,,, Mosquito Mtn., CT36447.

The same chromosome number was previously reported by Mulligan
{\965a) for plants from the Queen Charlotte Islands, Vancouver Island and
the Northwest Territories, by Packer (1964) on plants from Jasper, Alberta,
and by Love and Love (1966) on plants from Mount Washington. European
material of this species also has the chromosome number n = S (Jaretzky
1928fl, S0rensen and Westergaard in Love and Love 1948, Holmen 1952,
Love and Love 1956a, J0rgensen et al. 1958, and Sokolovskaja and Strelkova
1960).

304. Cardamine occidentalis (S. Wats.) Howell

MORESBY ISLAND: 2n = 32„, In = 64, Skidegate Lake, CT35271.

The only previous chromosome count on this species was reported by
Mulligan (1965fl) on material from the Queen Charlotte Islands. Calder and
Taylor in Part 1 state that "Cardamine occidentalis is probably the most poorly
represented western North American member of the genus Cardamine in her-
baria." They state further that "it is frequently overlooked in field surveys as
it closely resembles C. iimbellata.'' It is interesting to note that C. occidentalis
has the chromosome number 2n = 64, whereas C. umbellata has the number
2n = 48.

305. Cardamine oligosperma Nutt.

GRAHAM ISLAND: 2n = 16, 2^/2 mi S of Tlell, CST20895.

MORESBY ISLAND: 2n = 16, Alliford Bay, CST21073; 2n = 16,
East Copper Island, CST22219A.

CARD AMINE 71

Plants from the Queen Charlotte Islands and Vancouver Island have the
same chromosome number, 2n = 16 (Mulligan 1965a). This contrasts with
the chromosome number, 2n = 48, for the closely related species C. umbellata
Greene {see discussion under C. umbellata) .

306. Cardamine pensylvanica Muhl. ex Willd.

GRAHAM ISLAND: In = 32, Langara Island, CST22580.
MORESBY ISLAND: 2n = 16„, Skidegate Lake, CT37514.

Tetraploid, 2n = 32, plants of this species occur in New Brunswick,
Quebec, Yukon Territory, British Columbia and Colorado (Mulligan 1965a)
and octoploids, 2n = 64, occur in Ontario (Mulligan 1965fl) and Wisconsin
(Smith 1938).

307. Cardamine umbellata Greene

GRAHAM ISLAND: 2n = 24„, Masset Spit, CTS34730; 2n = 24„,
Tow Hill, CTS34755; 2n = 24„, Long Inlet, CT35966.

MORESBY ISLAND: 2n = 48, islet off Bolkus Islands, CST22226;
2n = 24n, Takakia Lake, CT36297.

Cardamine umbellata from five locations in the Queen Charlotte Islands
was hexaploid (« = 24 or 2/t = 48). The same chromosome number was ob-
tained on material of this species from elsewhere in British Columbia by
Mulligan (1965a). In contrast, the closely related species C. oligosperma Nutt.
is diploid {2n = 16). This difference in chromosome number supports mor-
phological evidence of Calder and Taylor in Part 1 that these two taxa should
be recognized at the specific level rather than the view of Hitchcock et al.
(1964) that C. umbellata should only be considered a variety of C. oligo-
sperma.

COCHLEARIA

308. Cochlearia officinalis L. ssp. oblongifolia (DC.) Hult.

GRAHAM ISLAND: n = 1, Masset Spit, CTS34727; 2n = In. Daw-
son Inlet, C75J5/47.

MORESBY ISLAND: 2n = 14, islet off Bolkus Islands, CST2223L

The somatic chromosome number of 14 reported for C. oblongifolia DC.
from Japan by Matsuura and Suto (1935) may have been determined on
material of this subspecies. The mciotic chromosomes segregated regularly at
telophase I.

72 CRUCIFERAE

Descurainia

309. *Descurainia sophia (L.) Webb ex Prantl in Engler & Prantl

MORESBY ISLAND: 2n = 28, Sandspit, CT 36036.

This same chromosome number has been obtained on North American
and European material by MulHgan (1961) and many other workers. The
2n = 56 count often credited to Manton (1932) is incorrect. In her paper, she
states that only a few octoploid cells occurred in normally tetraploid plants.

Draba

310. Draba hyperborea (L.) Desv.

MORESBY ISLAND: 2n = 36, islet off Bolkus Islands, CST22222;
2n = 19n, Gowdas Islands, CT 36570.

The 2/1 = 36 chromosome number for plants of this Pacific coast species
from the Bolkus Islands agrees with the number obtained from Vancouver
Island plants by Mulligan (1966). The material from the Gowdas Islands
sometimes formed 17 bivalents and 1 quadrivalent at metaphase I of meiosis
in pollen mother cells, which indicates that one chromosome pair might be
duplicated in plants with 38 somatic chromosomes.

311. Draba lonchocarpa Rydb. ssp. kamtschatica (Ledeb.) Calder & Taylor

MORESBY ISLAND: 2n = 8„, Mt. Moresby, CT36382.

This is the only known chromosome count for subspecies kamtschatica,
the coastal race of the Cordilleran D. lonchocarpa Rydb. It has the same
chromosome number, « = 8, as the typical subspecies of D. lonchocarpa from
interior British Columbia (Mulligan 1966).

Erysimum

312. ^Erysimum cheiranthoides L.

MORESBY ISLAND: 2n = 8n, Sandspit, CT 36026.

The chromosome number of this introduced weed is the same as that
obtained on North American and European material of this species {see Mulli-
gan 1957).

Hesperis

313. *Hesperis matronalis L.

LEPIDIUM 73

Lepidium

314. "^Lepidium campestre (L.) R. Br. in Ait.

MORESBY ISLAND: 2n = 8„, Sandspit, CT36012.

This is the same chromosome number, « = 8, that was obtained on North
American and European plants by many workers {see MulHgan 1957, and
Easterly 1963).

315. *Lepidiuiii densiflorum Schrad.

Neslia

316. '''Neslia paniculata (L.) Desv.

MORESBY ISLAND: In = 7„ Sandspit, CT36018.
The same chromosome number was also obtained from North American
and European material by Mulligan (1957) and other workers.

Raphanus

317. ^Raphanus raphanistrum L.

RORIPPA

318. *Rorippa islandica (Oeder) Borbas

GRAHAM ISLAND: 2n = 8„, IVi mi S of Tlell, CT35945.

This is the first diploid chromosome number found in material of the
species from North America. Mulligan (1964) reported the presence of tetra-
ploid plants in widely separated locations from Nova Scotia to British Colum-
bia and northward to Yukon and Alaska. Both diploid and tetraploid races of
this species are known to occur in Europe (J0rgensen et al. 1958 and other
workers ) .

SiNAPIS

319. *Sinapis arvensis L.

MORESBY ISLAND: 2n = 9„, Sandspit, CT36014.

This chromosome number, az = 9, was also found in European and
Asiatic material by Love and Love (1956^) and many other workers.

Sisymbrium

320. * Sisymbrium altissimum L.

MORESBY ISLAND: « - 7, Sandspit, CT 36043.

74 CRUCIFERAE

The same chromosome number, n = 1, was obtained by Mulligan
(1961), Easterly (1963) and by other workers on North American and
European plants of this introduced Eurasian species.

321. ^Sisymbrium officinale (L.) Scop.

GRAHAM ISLAND: 2n = 14, Queen Charlotte City, CT35912.

Easterly (1963) and other workers found the same chromosome number,
2n = 14, on North American and European material of this introduced Eura-
sian—African species.

SUBULARIA

322. Subularia aquatica L. ssp. americana Mulligan & Calder

MORESBY ISLAND: In = ISn, Mosquito Lake, CT23653.

Mulligan (1964) found approximately 28 somatic chromosomes in a
plant of subspecies americana from Quebec. Since his preparation was very
poor, this plant may have had 30 somatic chromosomes. Love and Love
(1956a) reported about 36 somatic chromosomes in material of subspecies
aquatica from Iceland. Hedberg (1957) found 2n = 28 in plants of 5. monti-
cola A. Br. from Mount Kenya in Africa. We need more chromosome numbers
of Subularia species before we can determine the basic chromosome number or
numbers of this genus.

Thlaspi

323. *Thlaspi arvense L.

Droseraceae
Drosera

324. Drosera- rotundifolia L.

GRAHAM ISLAND: In = 10„, 8 mi NW of Tlell, CT35695.

MORESBY ISLAND: 2n = lOn, between Aero and Moresby logging
camps, CT35293; n = 10, Kootenay Inlet, CT36135.

This widely distributed species has been counted many times and all
counts are 2n = 20 {see Love and Love 1961a).

Crassulaceae

Sedum

325. Sedum divergens S. Wats.

GRAHAM ISLAND: n = ^, Image Pt., CTS34683.

SEDUM 75

MORESBY ISLAND: 2n = 8ii, Limestone Island, CST22406.

These are the first counts reported for this species and a base number of
a: = 8 is proposed. There is an earher count of 2n = 28 for this species by
Baldwin (1935), but he questioned the identification of his material. His count
should be referred to another species, because the material was collected in
the southeastern United States. We now know that S. divergens is restricted to
northwestern United States and to adjacent Canadian regions.

326. Sedum roseum (L.) Scop.

MORESBY ISLAND: In = 18„, Kaisun, CT36550; In = 36, Gowdas
Islands, €736643.

Sedum roseum is known to have three chromosome races in Europe and
Asia, 2/7 = 16, 22 and 33 (Uhl 1952, J0rgensen et al. 1958, Banach-Pogan
1958, and Sokolovskaja and Strelkova 1960). Uhl (1952) surveyed the
chromosome numbers of this species in North America and found that only
two chromosome races. In = 22 and 36, occurred on this continent. He found
that the race with 22 somatic chromosomes is confined to the northeast, where-
as the race with 36 somatic chromosomes has a wider distribution, occurring
in central New York, in Minnesota and in western North America. Our counts
of 2/1 = 36 from the Queen Charlotte Islands answer in part the question
asked by Uhl about the race(s) present in the Pacific Northwest. The counts
from the Islands support his general statement on the distribution of the two
chromosome races in North America.

Saxifragaceae

Heuchera

327. Heuchera chlorantha Piper

GRAHAM ISLAND: n = 7,2n = 14, 2 mi E of Queen Charlotte City,
CTS34787; 2n = 14, Queen Charlotte City, CT36975.

MORESBY ISLAND: /2 = 7, Limestone Island, CST22402.

328. Heuchera glabra Willd. ex R. & S.

GRAHAM ISLAND: /? = 7, Blackwater Creek, CTS3507I; n = 7,
Dawson Inlet, C75J5/ 7 9.

This species was counted earlier by Packer (1964) from Jasper National
Park, Alberta.

Leptarrhena

329. Leptarrhena pyrolifolia (D. Don) R. Br.

76 SAXIFRAGACEAE

MiTELLA

330. Mitclla pentandra Hook.

GRAHAM ISLAND: 2n = 14, Jalun Lake, CT35654.
MORESBY ISLAND: 2n = 14, Takakia Lake, CT36299.

Previous chromosome counts were made by Schoennagel (1931) and
Packer (1964). The latter author counted material from Jasper National
Park, Alberta.

Parnassia

331. Parnassia fimbriata Konig

GRAHAM ISLAND: 2n = 36, Long Inlet, CT35959.

MORESBY ISLAND: 2n = 36, Takakia Lake, CT36273.

The same chromosome number was obtained by Packer (1964) on
material from Jasper National Park, Alberta, and by Taylor and Brockman
(1966) on material from Kokanee Glacier Provincial Park, in the interior of
British Columbia.

RiBES

332. Ribes bracteosum Dougl. in Hook.

GRAHAM ISLAND: 2n = 16, Dawson Inlet, CTS35124.

The same chromosome number was obtained by Zielinski (1953) on
material from Oregon.

333. Ribes lacustre (Pers.) Poir. in Lam.

GRAHAM ISLAND: 2n = 16, Haida Pt, C ST 2 3450.

Two previous workers have counted R. lacustre. Tischler (1927), and
more recently Zielinski (1953) and Love and Love (1966) have reported
2« = 16 for this species from North America.

334. Ribes laxiflorum Pursh

GRAHAM ISLAND: 2n = 16, Naden Harbour, CT 36847.
MORESBY ISLAND: 2n = 16, Sandspit, CST21846.

Two earlier counts with the same number have been made on this west-
em North American species. Zielinski (1953) counted material from Oregon,
and Anderson (1963) examined meiosis of material from Humboldt County
in California.

SAXIFRAGA 77

Saxifraga

335. Saxifraga caespitosa L.

336. Saxifraga ferruginea Grab.

GRAHAM ISLAND: 2/7 = 10„, Dawson Inlet, CTS35111; In = 10„,
Jalun Lake, CT35652; In = 10„, Long Inlet, CT35996.

MORESBY ISLAND: n = 10, Anna Inlet, CTS34942, CT35314;
2n = lOn, 2n = 20, Takakia Lake, CT36325; In = 10„, Mosquito Mtn.,
CT36469.

The previous number oi n = 19 for this species was reported by Beamish
(1960, 1961). Unfortunately the plates shown in her 1961 paper do not sup-
port the recognition of 19 chromosomes as the haploid complement. Plates 14
and 15 of this paper represent poor preparations and 19 chromosomes are not
visible in each of the segregating complements. Plate 13 is said to be a polar
view of a meiotic configuration. However, we would suggest that the illustra-
tion represents either the early stage of anaphase I or metaphase II because,
from our experience with this species, it is clearly not metaphase I. We have
counted not only the material from the Queen Charlotte Islands but also
material from inland British Columbia and it seems evident that the haploid
number for this species \s n = 10.

337. Saxifraga lyallii Engler ssp. hultenii Calder & Savile

MORESBY ISLAND: 2n = ca. 56, Mt. Moresby, CT 36403;
n - ca. 28, Mosquito Mtn., CT36462.

The chromosome number of this species is diflficult to determine from
either mitosis or meiosis because of the variation of the chromosome morphol-
ogy and pairing configurations. The only other species counted in this section
of Saxifraga is 5. odontoloma Piper, which has a chromosome number of

2n = 48.

338. Saxifraga mertensiana Bong.

GRAHAM ISLAND: 2n - ca. 50, Dawson Inlet, CTS35089.

MORESBY ISLAND: 2n = ca. 48-50, Anna Inlet, CTS34928;
2n = ca. 48, Blackwatcr Creek, CTS35053.

The only previous count for this species was made on Vancouver Island
material by Beamish (1961). She reported that it was difficult to obtain a
meiotic count and was only able to make an approximate mitotic count of 48
from ovular tissue. Similarly, wc have found it difficult to obtain an accurate
count, since the somatic chromosomes arc small and arc of several dilTcrcnt
sizes with indistinct outlines.

78 SAXIFRAGACEAE

339. Saxifraga oppositifolia L.

MORESBY ISLAND: « = 13, Mt. Russ, CT36I96.

Many other workers have obtained the same chromosome number for this
species. Recently Packer (1964) reported 2n = 26 for material from Jasper
National Park, and Mosquin and Hayley (1966) examined plants from Mel-
ville Island in the Canadian Arctic. The only other counts for S. oppositifolia
are 2n = 39 and 52. Both of these counts are discussed by J0rgensen et al.
(1958).

340a. Saxifraga punctata L. ssp. carlottae Calder & Savile

MORESBY ISLAND: n = 36, Takakia Lake, CT36348; 2n = 36„,
Mosquito Mtn., CT36463.

The counts obtained on material from the Queen Charlotte Islands agree
with the count obtained by Moore {in Calder and Savile 1960) on ssp. por-
sildiana Calder & Savile {2n = ca. 72). Packer (1964) has recently counted
ssp. nelsoniana (D. Don) Hult. from the Richardson Mountains in northwest-
em Canada and found the chromosome number to be 2n = 28. A similar
number is reported for S. punctata s. lat. in Russia by Sokolovskaja and Strel-
kova (1938).

340b. Saxifraga punctata L. ssp. cascadensis Calder & Savile

341. Saxifraga taylori Calder & Savile

GRAHAM ISLAND: 2n = 26„, Jalun Lake, CT35641.

MORESBY ISLAND: n = 13, Yatza Mtn., CT35712; 2n - 13„, Taka-
kia Lake, CT36272; 2n = 26„, Mt. Moresby, CT36455.

This species was previously counted by Moore (1959) on plants from
Mount de la Touche on the Queen Charlotte Islands {CT23511) but he only
made an approximate mitotic count of 2n = 54-56 on a leaf primordium
squash. Meiotic examination of this species has shown that the haploid num-
ber is either 13 or 26 and undoubtedly Moore's count should be considered as
2n = 52.

Saxifraga taylori is restricted to the central montane region on the Islands
and is one of the conspicuous endemic elements of the Charlottes. The pres-
ence of two chromosome races, based on x = 13, on the Islands is a most
interesting facet of this species. Saxifraga taylori is the only member of the
section Trachyphyllum Gaudin of Saxifraga on the Islands. It is suggested that
the tetraploid race represents a natural autotetraploid. Morphologically the
two races are identical. Examination of meiosis in the tetraploid reveals the
regular formation of 26 bivalents in diakinesis and metaphase I. This pairing
behavior indicates that some genetic mechanism is operative in this polyploid
to prevent the formation of multivalents.

SAXIFRAGA 79

342. Saxifraga tolmiei T. & G.

Tellima

343. Tellima grandiflora (Pursh) Dougl. in Lindl.

TiARELLA

344. Tiarella trifoliata L.

GRAHAM ISLAND: 2n = 7„, Image Pt., CTS34680; n = 1, Black-
water Creek, CTS35067; 2n = lu, Dawson Inlet, CTS35086.

The same chromosome number has been reported for this species by Packer
(1964) from Whitecourt, Alberta, and by Kern (1966) from an undisclosed
locality in the Pacific Northwest.

345. Tiarella unifoliata Hook.

GRAHAM ISLAND: 2n = In, Dawson Inlet, CT35087.
MORESBY ISLAND: 2n = 7n, Takakia Lake, CT 36320.

Kern (1966) reported n = 1 for this same entity under the name T. tri-
foliata L. ssp. unifoliata (Hook.) Kern. No documentation for the count was
given. Taylor (unpublished) has made a number of counts, all n = 7, for this
species from Canadian and United States localities.

TOLMIEA

346. Tolmiea menziesii (Pursh) T. & G.

GRAHAM ISLAND: 2n = 28, Yakoun Lake, CT 36777.

MORESBY ISLAND: 2/7 = 28, Crescent Inlet, CTS34979; n = 14, be-
tween Cumshewa and Peel inlets, CT35178.

Other workers have reported the same number for this western North
American species (Schoennagel 1931, Skovsted 1934, Hamel 1963, and
Rogers 1965).

Rosaceae

Amelanchier

347. Amelanchier florida Lindl.

GRAHAM ISLAND: 2n = 34„, Image Pt., CTS3467I.

This is the first count reported for the western North American scgregant
of the A. alnifolia Nutt. complex. According to Sax (1931 ), the base number
of Amelanchier is jc = 17. Queen Charlotte Islands plants of A. florida are
tetraploid.

80 ROSACEAE

Aphanes

348. Aphanes occidentalis (Nutt. ) Rydb.

GRAHAM ISLAND: 2n = 16, Queen Charlotte City, CST23020.

Aruncus

349. Aruncus Sylvester Kostel.

GRAHAM ISLAND: 2n = 9„, Dawson Inlet, CTS35120.
MORESBY ISLAND: n = 9, head of Cumshewa Inlet, CT35224.

Chromosome counts on other species of Aruncus in North America are
the same as for the western North American A . Sylvester.

Crataegus

350. Crataegus douglasii Lindl.

GRAHAM ISLAND: n = ca. 17, 1 mi W of Queen Charlotte City,
CTS34796.

The haploid number of 17 agrees with counts obtained for other species
of Crataegus and for the Pomoideae Focke in general by most authors. The
only other count reported for C. douglasii is « = 24 by Longley (1924) and
he considered it a triploid with a base number of ;c = 16.

Meiosis in the material from the Islands was irregular with univalents
present at metaphase I and lagging chromosomes during anaphase and telo-
phase I.

351. * Crataegus monogyna J acq.

Fragaria
352a. Fragaria chiloensis (L.) Duchesne ssp. lucida (E. Vilm.) Staudt

352b. Fragaria chiloensis (L. ) Duchesne ssp. pacifica Staudt

GRAHAM ISLAND: In = 56, Masset Spit, CST21223.

Staudt (1962) has reported that he obtained an octoploid somatic count
of 2/1 = 56 on 45 collections of this species. He made one count on material
collected during the 1957 Plant Research Institute survey of the Islands. The
octoploid number is in agreement with the many counts made on this species
by other workers {see Love and Love 1961 a).

GEUM 81

Geum

353. Geum calthifolium Smith in Rees

GRAHAM ISLAND: 2n = 42, Jalun Lake, CT35671.

MORESBY ISLAND: n = 21, Bigsby Inlet, CTS34899; In = 42, Anna
Inlet, CTS34931; 2n = 42, Yatza Mtn., CT35710; 2n = 42, Mt. Russ,
CT36168; In = 42, Sunday Inlet, CT36610.

This western native Geum is considered to be a hexaploid.

354. Geum macrophyllum Willd.

MORESBY ISLAND: 2n = 42, head of Cumshewa Inlet, CT36244.

This chromosome number has been repeatedly observed in the same
species (5^e Gajewski 1957).

355. Geum schofieldii Calder & Taylor

MORESBY ISLAND: 2n = 111, Mt. Russ, CT36144.

Geum schofieldii is an endemic of the montane regions on the Islands.
This species is 16-ploid and has the highest chromosome number known in
the genus. The evolution of this distinct species is vague and the cytological
information does not provide any real clue to its position or relationship to
other members of the subgenus Acomastylis (Greene) Gajewski {see discus-
sion in Part 1 ) .

LUETKEA

356. Luetkea pectinata (Pursh) Kuntze

MORESBY ISLAND: n = 9, Mt. Moresby, CT37017.

This count agrees with the recent report of 2a? =: 18 by Packer (1964)
on plants from Jasper National Park, Alberta. Telophase I was regular.

POTENTILLA

357. Potentilla pacifica Howell

GRAHAM ISLAND: 2n = 28, Yakoun River Delta, CT35471;
2n = 28, Masset Spit, CT35703.

MORESBY ISLAND: 2n - 28, Gray Bay, CT35239; 2n 28, Mike
ln\Q{, CT3666J.

82 ROSACEAE

In a recent general review of the Potentilla anserina L. complex, Rousi
(1965) reports that all counts for the Pacific coast entity that we recognize
as P. pacifica are tetraploid, 2n = 28. On the basis of his extensive study, he
recognizes the west coast taxon as a new subspecies, P. anserina L. ssp. paci-
fica (Howell) Rousi.

358. Potentilla palustris (L.) Scop.

MORESBY ISLAND: 2n = 35, Skidegate Lake, CT35148.

The only other chromosome count on this species is 2n = 28 reported
by Ehrenberg (1945) on Scandinavian material. The 2n = 35 count on the
Queen Charlotte Islands population strongly suggests that it is apomictic.

359. Potentilla villosa Pallas ex Pursh

GRAHAM ISLAND: 2n = 14, Newton Pt., CST22942.
MORESBY ISLAND: 2n = 14, Hotspring Island, CST22279.

Previous counts of 2n = 14, 42 and 49 have been reported for this
species. The chromosome counts of 42 and 49 have been attributed incorrectly
to P. villosa through a misconception of a statement by MUntzing (1931) ". . .
P. villosa Zimm. is probably synonymous with P. CrantzU Beck. According to
Tischler P. villosa has n = 1; my Crantzii-stTains were hexaploid and hepta-
ploid." Unfortunately MUntzing's misleading proposal that P. villosa, a Berin-
gian species, was probably synonymous with P. crantzii, a Eurasian species,
has led to the present confusion about chromosome numbers in P. villosa. The
exact source of the material that Tischler (1929) counted is not known, but
his report of a haploid number of « = 7 corresponds to our own diploid
counts of 2« = 14 from the Queen Charlotte Islands.

Pyrus

360. Pyrus fusca Raf.

Rosa

361. Rosa nutkana Presl.

RUBUS

362. Rubus chamaemorus L.

363. *Rubus laciniatus Willd.

RUBUS 83

364. Rubus parviflorus Nutt.

GRAHAM ISLAND: 2n = \4, Queen Charlotte City, CST23012;
2n = 14, Image Pt., CTS34685; 2n = 14, Haida Pt., CT36686.

These counts from the Queen Charlotte Islands agree with counts by
Darrow and Longley (1933), Vaarama (1954), and Jinno (1958).

365. Rubus pedatus Smith

GRAHAM ISLAND: In = 14, 5 mi N of Port Clements, CTS34695.

366. *Rubus procerus P. J. Mueller ex Boulay

367. Rubus spectabilis Pursh

GRAHAM ISLAND: 2« = 14, about 8 mi SSW from Juskatla,
CT35559; 2n = 14, 5 mi W of Tow Hill, CT37471.

MORESBY ISLAND: 2n = U, Bigsby Inlet, CT37463.

The same chromosome number was obtained by Darrow and Longley
(1933) from western North American material.

368. Rubus ursinus Cham. & Schlecht.

Sanguisorba

369. Sanguisorba canadensis L. ssp. latifolia (Hook.) Calder & Taylor

GRAHAM ISLAND: 2n = 28, Long Inlet, CT35958.

MORESBY ISLAND: 2n = 28, Mt. Russ, CT36147; 2n = 28, Taka-
kia Lake, CT36318; 2n = 28, Bigsby Inlet, CT37400.

The only previous count reported for Sanguisorba canadensis L. is
2n = 56 (Larsen 1959; Nordborg 1963, and Love and Love in Love and Love
1961a). Larsen's count was obtained on botanical garden material. The octo-
ploid chromosome number of 2n = 56 reported by these authors should prob-
ably be referred to the typical variety found in eastern North America. The
western subspecies, which is found on the Queen Charlotte Islands, is a
tetraploid.

370. Sanguisorba menzicsii Rydb.

84 ROSACEAE

371. San^uisorba officinalis L. ssp. microcephala (Presi) Calder & Taylor

GRAHAM ISLAND: 2/? = 14„, 2« - 28, 15 m S of Masset,
CT35565.

MORESBY ISLAND: 2n = 28, Red Mud Marsh, CST23187.

Nordborg (1963), in a review of S. officinalis L., reports two cytotypes
(tetraploid and octoploid) based on x = 1. Her information was based on
studies conducted on European and Asiatic material. The tetraploid number
from the Queen Charlotte Islands represents the first count for this species in
North America. It would be interesting to know the chromosome number of
the typical subspecies found in the dry interior valleys of western Yukon and
central Alaska.

SiBBALDIA

372. Sibbaldia procumbens L.

MORESBY ISLAND: 2n = 14, Mt. Moresby, CT36375; In = 14,
Mosquito Mtn., CT36446.

Many counts have been made on the species and all have been reported
as either n = 1 or 2n = 14. Three counts have been reported from North
America (Wiens and Halleck 1962, Packer 1964, and Love and Love 1966).

SORBUS

373. *Sorbus aucuparia L.

GRAHAM ISLAND: 2n = 34, Masset, CT 36831.

This same number has been reported by several authors {see Love and
Love \96\a).

374a. Sorbus sitchensis M. Roemer ssp. sitchensis

374b. Sorbus sitchensis M. Roemer ssp. grayi (Wenzig) Calder & Taylor

MORESBY ISLAND: 2n - ca. 34, Takakia Lake, CT3632L

Material of this species from the Queen Charlotte Islands is probably
diploid with the base number x = 17.

Spiraea
375. Spiraea douglasii Hook. ssp. menziesii (Hook.) Calder & Taylor
MORESBY ISLAND: n = 18, Skidegate Lake, CT 36062.

SPIRAEA 85

Sax (1936) reported n = IS for S. douglasii Hook., but was unable to
obtain a count for 5. menziesii Hook. Our examination shows that the more
northern ssp. menziesii has the same chromosome number as the more wide-
spread southern ssp. douglasii.

Leguminosae

Cytisus

376. * Cytisus scoparius (L.) Link

MORESBY ISLAND: 2n = 23„, Skidegate Lake, CT35152.

Our examination of material of this European species has revealed the
same meiotic configuration found by Bocher and Larsen (1958/?). These two
authors have given a lucid discussion of the discrepancies in counts previously
reported for this species; namely, that counts of 2n = 48 reported from
mitotic examinations are undoubtedly in error because of the presence of two
large satellited chromosomes that usually give the appearance of four chromo-
somes. However, our examinations and those by Bocher and Larsen have
substantiated the earHer count of Maude (1940).

Lathyrus

377. Lathyrus japonicus Willd.

MORESBY ISLAND: 2n = U, Sandspit, CT37008.

All previous counts for this species give the same chromosome number
(see J0rgensen et al. 1958, and Brightmore and White 1963).

378. Lathyrus littoralis (Nutt.) Engl.

GRAHAM ISLAND: 2n = 14, Tlell, CT35432.

This interesting coastal strand and beach species occurs only along the
central Pacific coast of North America. It has the same base number of jc = 7
as previously determined for other members of the genus.

379. Lathyrus ochroleucus Hook.

380. Lathyrus palustris L.

LUPINUS

38L Lupinus littoralis Dougl. in Lindl.

GRAHAM ISLAND: 2n = 24„, mouth of Sangan River, CT35597.

A single count of /? ^ 24 on material from the coast of Oregon was re-
ported by Phillips ( 1957) for this western American tetraploid species.

86 LEGUMINOSAE

382. Lupinus nootkatensis Donn ex Sims

GRAHAM ISLAND: 2n - 24„, between Skidegate and Queen Char-
lotte City, CTS34777.

Previous counts by Maude (1939, 1940) and Bragdo (1957) are tetra-
ploid, 2n = 48, and were made on naturalized or cultivated material. The
count from the Queen Charlotte Islands represents the first time native
material of this species has been counted.

Melilotus

383. *MeIilotus alba Desr. in Lam.

MORESBY ISLAND: n = 8, Sandspit, CT 36032.

Many other diploid numbers based on ;c = 8 have been reported for
this species. Two other counts are reported, 2n = 24 (Atwood 1936) and
2n = 32 (Atwood 1936, and Wipf 1939). However, these counts were made
on experimentally induced polyploids or the examination was made a somatic
tissue found in root nodules.

384. *Melilotus officinalis (L.) Lam.

MORESBY ISLAND: 2n = 8„, Sandspit, CT36031.

All earlier counts of this species are either n = ^ or 2n = \6 {see Love
and Love 1961<3).

OXYTROPIS

385. Oxytropis campestris (L.) DC.

Trifolium

386. *Trifolium dubium Sibth.

387. *Trifolium pratense L.

MORESBY ISLAND: 2n = In, Sandspit, CT35346.

The many counts made on this species are either /i = 7 or 2^? = 14,
except for those made on induced tetraploids used in plant-breeding experi-
ments (Wipf and Cooper 1938).

388. *Trifolium repens L.

MORESBY ISLAND: n ^ \6, Sandspit, CT35165.

TRIFOLIUM 87

This widely distributed clover species has been counted many times and
all counts, except those for experimentally produced polyploids, are tetraploid,
2n = 32 {see Love and Love 1961a).

389. Trifolium wormskjoldii Lehm.

GRAHAM ISLAND: 2n = 32, Tlell, CST22098; 2n = 32, Image Pt.,
CT37398; 2n = 32, Haida Village, CT37417; 2n = 32, N end of Dawson
Inlet, CT37483; 2n = 32, between Millar Creek and Skidegate Mission,
CT37546.

MORESBY ISLAND: 2n = 32, Anna Inlet, CTS34921; 2n = 16„,
Kootenay Inlet, CT 36199; 2n = 32, Moresby logging camp, CT36510.

The previous count by Wexelsen (1928) of 2n = ca. 48 for this species
is probably an error, as was pointed out by Mosquin and Gillett (1965) in a
review of chromosome numbers in Trifolium.

390. tilex europaeus L.

Ulex

ViCIA

390a. *Vicia cracca L.

GRAHAM ISLAND: 2/7 = 14„, Port Clements, CT36113.

Rousi (1961) has demonstrated that three chromosome races exist in
this species: 2n = 12, 14 and 28. All populations examined in North America
were tetraploid with a base number of 7. The Queen Charlotte Islands count
represents the first count made on a western North American adventive popu-
lation.

391. Vicia gigantea Hook.

GRAHAM ISLAND: a? = 7, between Skidegate and Skidegate Village,
CTS34665.

Three basic chromosome numbers of x = 5, 6 and 7 have been reported
for the genus. A review of the chromosome numbers of western North Ameri-
can species indicates that most species are diploid and have a base number of

392. *Vicia sativa L.

MORESBY ISLAND: n = 6,2n = 12, Sandspit, CT35348.

This widely introduced European adventive has been repeatedly reported
to be a diploid with a base number of jc = 6 {see Srivastava 1963, and Huzi-
wara and Kondo 1963).

88 LEGUMINOSAE

393. *Vicia villosa Roth

MORESBY ISLAND: 2n = 7„, Sandspit, CT36081.

All previous counts for this series are diploid, 2n = \4 (see Love and
Love 1961a).

Geraniaceae

Geranium

394. *Geranium dissectum L.

MORESBY ISLAND: 2n = 11„, Sandspit, CT35329.

Three earlier counts on this species have shown a uniform chromosome
number of ^z = 11 (Gauger 1937, Warburg 1938, and Love and Love
1944fl).

395. ^Geranium molle L.

MORESBY ISLAND: 2n = 13n, Sandspit, CT35330.

The same chromosome number was obtained on Canadian and European
material of this species (see Mulligan 1959).

396. Geranium richardsonii Fisch. & Trautv.

MORESBY ISLAND: n = 26, Limestone Island, CTS3483L

Our count of « ^ 26 agrees with that of Shaw (1952) on material from
Utah. Warburg (1938) reported "« = 28?," which suggests that he was un-
certain of his count. This species is probably tetraploid with the base number
X = 13.

Linaceae

LiNUM

397. Linum bienne Mill.

Callitrichaceae

Callitriche

398. Callitriche heterophylla Pursh ssp. bolanderi (Hegelm.) Calder & Taylor

GRAHAM ISLAND: 2n = 20, between Juskatla and Port Clements,
CT S3 5 08 3.

MORESBY ISLAND: 2n - 20, between Moresby and Aero logging
camps, CT35291; 2n = 20, Skidegate Lake, CT37515.

CALLITRICHE 89

The base numbers for this genus are x = 3, 5 and 9. The Queen Char-
lotte Islands population is presumably tetraploid, based on ;t = 5.

Enipetraceae
Empetrum

399. Empetrum nigrum L.

Violaceae

Viola

400. Viola adunca Smith in Rees

GRAHAM ISLAND: 2n = 20, IVi mi S of Jungle Beach, CST23460.

The same chromosome number, n = 10 or 2/i = 20, was obtained on
material of this species from California, Nevada, Oregon, Idaho, Colorado and
Vancouver Island, British Columbia, by Gershoy (1934) and Clausen et al.
(1940). It is interesting to note that plants with the chromosome number
n "^ 20 or 2n = 40 occur in Alberta and Manitoba (Packer 1964, and Taylor
andBrockman 1966).

401. Viola biflora L. ssp. carlottae Calder & Taylor

GRAHAM ISLAND: 2n = 48, Long Inlet, CT35977.

MORESBY ISLAND: 2n = 48, Bigsby Inlet, CTS34893; 2n = 48,
Anna Inlet, CTS34924; 2n = 48, Yatza Mtn., CT35711; 2n = 48, Takakia
Lake, CT36306; 2n = 48, Mosquito Mtn., CT36449.

The widely distributed V. biflora L. has been reported to have many
counts oi 2n = 12 {see Love and Love \96\a) and one count oi 2n = 18
(Sokolovskaja and Strelkova 1962). Miyaji (1929) has reported n = 6 iov
V. biflora and 2n = 48 for the Japanese species V. crassa Makino, an entity
that has been treated as a northern Asiatic alpine race of V. biflora found in
central and northern Honshu and Hokkaido in Japan and also in the Kuriles
and Kamchatka. The recent English translation of the revised Japanese flora
by Ohwi (1965) segregates the latter race from V. biflora as a separate species
V. crassa. The members of Viola section Dischidium Ging. present some inter-
esting systematic problems that await research.

402. Viola glabella Nutt in T. & G.

MORESBY ISLAND: 2n - 24, Takakia Lake, CT 36298.

90 VIOLACEAE

Two previous counts have been made on this species. Gershoy (1934)
reported 2n =^ 24 on material from an unstated locality and Taylor and Brock-
man (1966) reported 2« = 24 from a population in southern British Co-
lumbia.

403. Viola Ian gsdorffii (Regel) Fisch.

MORESBY ISLAND: 2n = 60„, Mount Moresby, CT36425; In = ca.
120, 60n, Anna Inlet, CTS34963.

The only previous count attributed to V. langsdorffii was made on culti-
vated material by Miyaji (1929), who found Japanese plants to be octoploid,
2/7 = 96. Sokolovskaja (1963) reported 2/t = 96 for V. kamtschadalorum
W. Becker & Hult. from Kamchatka. This species is closely related to V.
langsdorffii and has been considered by Ohwi (1965) to be synonymous with
it. The disposition of the morphological variants of V. langsdorffii s. lat. is not
clear. The decaploid chromosome number of 2n = ca. 120 obtained for the
Queen Charlotte Islands material may indicate that the eastern Pacific popu-
lation of V. langsdorffii differs from the western Pacific population.

404. Viola palustris L.

GRAHAM ISLAND: 2n = 48, Dawson Inlet, CT37345.
MORESBY ISLAND: 2n = ca. 48, Skidegate Lake, CT37411.

Many counts have been made on this species and all counts reported are
2n = 48 (see Gadella 1963, and Love and Love 1961a).

Onagraceae

CiRCAEA

405a. Circaea alpina L. ssp. alpina

MORESBY ISLAND: n = 11, Mosquito Lake, CT36710.

405b. Circaea alpina L. ssp. pacifica (Asch. & Magnus) Raven in Calder &
Taylor

GRAHAM ISLAND: n = 11, Honna River, CTS34802; n = 11, AVi
mi S of Port Clements, CTS35038; n = 11, 8 mi SSW of Juskatla, CT35476;
n - 11, Long Inlet, CT35968; n = 11, Honna River, CT 36934.

Previous counts on this species have revealed both subspecies to be di-
ploid, 2n = 22 (Lewis et al. 1958). A recent count from western Canada of
2n = 22 was reported by Packer (1964).

EPILOBIUM 91

Epilobium

406. Epilobium anagallidifolium Lam.

MORESBY ISLAND: 2n = 36, 3 mi E of Skidegate Lake, CST21919;
2n = 36, Takakia Lake, CST2305L

This widely reported occurring species has been counted by many authors
and all but one report give the chromosome number as either a? = 18 or
2n = 36. The report by Mattick {in Tischler 1950) oi n = 9 may be incorrect.

407. Epilobium angustifolium L.

GRAHAM ISLAND: n = 36, between Millar Creek and Skidegate Vil-
lage, CrJ (5697.

MORESBY ISLAND: /? = 36, Tuft Islets, CT S3 4865; n = 36, Sands-
pit, CT36072; « = 36, Horn Rock, CT36517.

These counts have been discussed by Mosquin (1966). In this paper
dealing with a worldwide study of the species, the Queen Charlotte Islands
population has been recognized as part of his newly proposed subspecies cir-
cumvagum. A recent report of In = 72 by Love and Love (1966) has been
reported under a new species E. platyphyllum (Daniels) Love and Love.

408. Epilobium brevistylum Barbey in Brewer & Wats.
MORESBY ISLAND: In = 36, near Copper Bay, CST21926.

409. Epilobium delicatum Trel.

GRAHAM ISLAND: In = 36, Tow Hill, CST22667; 2n = 36, be-
tween Ells and Mercer pts., CST22906; 2n = 36, Shields Bay, CT23343.

MORESBY ISLAND: 2n = 36, below Newcombe Peak, CST22049;
2n = 36, Mt. de la Touche, CT23594; 2n = 36, between Cumshewa and
Peel inlets, CrJ57 55^.

410. Epilobium glandulosum Lehm.

GRAHAM ISLAND: 2n = 36, Jungle Beach, CST23393; 2n - 36,
about 2^/2 mi S of Tlell, CST23410.

MORESBY ISLAND: 2n = 36. between Sandspit and Copper Bay,
CST23192; 2n = 36, Skidegate Lake, CT23639.

Sokolovskaja (1963) counted material from Kamchatka and reported
2n = 36. No previous counts have been reported from native North American
material.

92 ONAGRACEAE

411. Epilobium latifolium L.

412. Epilobium minutum Lindl. ex Hook.

GRAHAM ISLAND: 2n = 26, between Skidegate and Skidegate Vil-
lage, CST21399.

MORESBY ISLAND: In = 26, Hotspring Island, CST22302;
2n = 26, Limestone Island, CST22414.

In 1962, Kurabayashi et al. examined meiosis of material from California
and reported « = 13.

413. Epilobium palustre L.

Haloragidaceae

Myriophyllum

414. Myriophyllum spicatum L.

GRAHAM ISLAND: In = 28, near mouth of Honna River, CT 36945.

Previous counts for this species have been made by Love and Love on
material from Iceland. Their first report in 1948 gave the chromosome num-
ber as 2n = 36, but subsequent reexamination by Love in 1954 gave the
number as 2« = 28. However, both these reported counts were refuted by A.
Love in 1961 when another examination of new material from Iceland gave
the number as 2n = 42. A count of 2n = 42 is reported for material of this
species from Manitoba by Love and Ritchie (1966).

Hippuridaceae

HiPPURIS

415. Hippuris montana Ledeb.

MORESBY ISLAND: 2n = 16, Takakia Lake, CT 36315.

This count oi2n = 16 is the first report of a diploid species in the genus
and substantiates the previously proposed base number of x = 8. All earlier
counts on Hippuris were polyploid.

416. Hippuris vulgaris L.

GRAHAM ISLAND: 2n = 32, about 4 mi N of mouth of Oeanda River,
CT35853.

HIPPURIS 93

Many similar counts have been made on this species {see Love and Love
1961a, Sokolovskaja 1963, Gadella and KHphuis 1963, Sorsa 1963, and Love
and Ritchie 1966). The report of 2n = 30 by Harada (1952) on Japanese
material may be in error.

Araliaceae

Oplopanax

417. Oplopanax horridus (Sm.) Miq.

Umbelliferae
Angelica

418. Angelica lucida L.

MORESBY ISLAND: 2n = 14„, 2n = 28, Takakia Lake, CT36336;
In = 14n, Gowdas Islands, CT36572.

These counts establish the presence of another tetraploid species in the
Apioideae Drude with a base number oi x = 7.

CONIOSELINUM

419. Conioselinum pacificum (S. Wats.) Coult. & Rose

GRAHAM ISLAND: n = 22, Haida Pt., CT35396; 2n = 22„, Torrens
Island, CT35829; 2n = 22„, 21/2 mi S of Jungle Beach, CT 36696.

MORESBY ISLAND: 2n = 44, Tuft Islets, CTS34864; 2n = 22„,
Sandspit, CST23210; 2n = 22„, Horn Rock, CT36518; 2n = 22n, Kaisun,
CT36536; 2n = 44, Gowdas Islands, CT3657L

Previous counts for this taxon are reported under C. chinense (L. ) B.S.P.
by Bell and Constance (1966) and all are n = 22. For a discussion of the
taxonomic considerations see Calder and Taylor in Part 1 .

Glehnia

420. Glehnia littoralis Schmidt ssp. leiocarpa (Mathias) Hult.

GRAHAM ISLAND: n = 1 1, Tlell, CT35438.

All earlier counts of this species have given a haploid chromosome num-
ber oi n = 11. Jinno (1956) and Liu ct al. (1961) obtained this chromosome
number on material of the typical subspecies from Japan and from Taiwan;
Bell and Constance (1966) found a? 1 1 in plants of subspecies leiocarpa
from Oregon.

94 UMBELLIFERAE

Heracleum

421. Heracleum lanatum Michx.

GRAHAM ISLAND: 2n = 11„, N end of Dawson Inlet, CST22852.

MORESBY ISLAND: 2n = 22, W of Moresby logging camp, CT37343,
CT37441.

The count reported here agrees with other previous counts of H. lanatum
made by Bell and Constance (1957) and Love and Love (1966) on material
from North America.

LiGUSTICUM

422. Ligusticum calderi Math. & Const.

GRAHAM ISLAND: In = 33n, Jalun Lake, CT35634.

MORESBY ISLAND: 2n = 33„, Bigsby Inlet, CTS34903; In = 33„,
In = ca. 66, Anna Inlet, CT34985; In = 33„, Yatza Mtn., CT35728.

This endemic species is the only hexaploid member of Ligusticum found
on the Pacific coast of North America. The same chromosome number has
been reported by Bell and Constance (1966) for L. filicinum S. Wats, from
Wyoming.

423. Ligusticum scoticum L. ssp. hultenii (Fernald) Calder & Taylor

LiLAEOPSIS

424. Lilaeopsis occidentalis Coult. & Rose

GRAHAM ISLAND: 2« = 44, Kumdis River Delta, CT23803; 2n = 44,
Yakoun River, CT35468; 2n = 22„, Delkatla Inlet, CT35587.

MORESBY ISLAND: 2n = 22„, Skidegate Lake, CT 36064; 2n = 44,
Skidegate Lake, CT36064.

This species is a tetraploid with the base number x = W.

Oenanthe

425. Oenanthe saimentosa Presl in DC.

GRAHAM ISLAND: 2n = 44, Tlell, CT35925.

Bell and Constance (1957) obtained the same chromosome number,
n = 22, from material of this species collected in California.

OSMORHIZA 95

OSMORHIZA

426. Osmorhiza chilensis H. & A.

427. Osmorhiza purpurea (Coult. & Rose) Suksd.

GRAHAM ISLAND: 2n = 22, Dawson Inlet, CT37356.

This diploid number of O. purpurea places this species at the same ploidy
level as other Osmorhiza counted by Bell and Constance (1957, 1960 and
1966).

Sanicula

428. Sanicula crassicaulis Poepp. ex DC.

GRAHAM ISLAND: 2n = 16„, Image Pt, CTS34672.

Bell (1954) has fully discussed the three chromosome races (tetraploid,
hexaploid and octoploid) in this species for western North America. The count
from the Queen Charlotte Islands indicates that the tetraploid race is present
near the northern limit of the typical subspecies along the Pacific coast.

Cornaceae

CORNUS

429. Cornus stolonifera Michx.

GRAHAM ISLAND: 2n = 22, near junction of Yakoun River and
Ghost Creek, CT35508.

This count agrees with other counts made on this species from Western
Canada by Taylor and Brockman (1966).

430. Cornus unalaschkensis Ledeb.

GRAHAM ISLAND: 2n = 22„, near Masset Spit, CTS34726.

A previous count by Taylor and Brockman (1966), cited under C. inter-
media (Farr. ) Calder & Taylor, gave 2n = 1 In for this species. Counts on the
closely related C. canadensis L. have shown the presence of two chromosome
races, a diploid 2n = 22 (Packer 1964) and a tetraploid 2n = 44 (Derman
1932). It appears that two races are also present in C. unalaschkensis.

Pyrolaceae
Hypopitys

431. Hypopitys monotropa Crantz

96 PYROLACEAE

MONESES

432. Moneses uniflora (L.) Gray ssp. reticulata (Nutt.) Calder & Taylor

GRAHAM ISLAND: 2n = 26, Jungle Beach, CTS34669.

The typical subspecies was reported to have 2n = 26 by Hagerup
(1941). The Queen Charlotte Islands record is the first count for North
American material.

Pyrola

433. Pyrola secunda L.

Ericaceae
Andromeda

434. Andromeda polifolia L.

Arctostaphylos

435. Arctostaphylos uva-ursi (L.) Sprengel

Cassiope

436. Cassiope lycopodioides (Pall.) D. Don ssp. cristapilosa Calder & Taylor

437. Cassiope mertensiana (Bong.) G. Don

438. Cassiope stelleriana (Pall.) DC.

Cladothamnus

439. Cladothamnus pyrolaeflorus Bong.

Gaultheria

440. Gaultheria shallon Pursh

Kalmia

441. Kalmia polifolia Wang.

Ledum

442. Ledum palustre L. ssp. groenlandicum (Oeder) Hull

LOISELEURIA

443. Loiseleuria procumbens (L.) Desv.

MENZIESIA 97

Menziesia

444. Menziesia ferruginea Smith

Phyllodoce

445. Phyllodoce glanduliflora (Hook.) Coville

MORESBY ISLAND: 2n = 12u, Takakia Lake, CT36276,

This count represents the second species counted in the genus Phyllodoce.

Vaccinium

446. Vaccinium alaskense Howell

447. Vaccinium caespitosum Michx.

448. Vaccinium ovalifolium Smith in Rees

449. Vaccinium oxycoccus L.

450. Vaccinium parvifolium Smith in Rees

451. Vaccinium scoparium Leiberg

452. Vaccinium uliginosum L.

453. Vaccinium vitis-idaea L. ssp. minus (Lodd.) Hult.

GRAHAM ISLAND: 2n = 12„, 5V^2 mi SE of Port Clements,
CTS3459L

All earlier counts on this species have been reported as 2n = 24 (see
Love and Love 1961a).

Primulaceae

Dodecatheon

454. Dodecatheon jeffreyi Van Houtte

GRAHAM ISLAND: 2n = 43„, White Creek Muskeg, CTS34742;
2n = ca. 86, Jalun Lake, CT35624.

MORESBY ISLAND: 2n = ca. 86, Anna Inlet, CTS34936; 2n = ca. 86,
Upper Victoria Lake, CT35736; 2n = ca. 86, Mt. Russ, CT36J60.

98 PRIMULACEAE

In his biosystematic study of Dodecatheon, Thompson (1953) stated, in
the discussion on the cytology of D. jefjreyi ssp. jefjreyi, that mitotic counts
varied from 2n = 42 to 44. In a later paper, Beamish (1955) reported that
plants from Douglas Island, Alaska, Prince Rupert and Vancouver Island, in
British Columbia, belong to ssp. jefjreyi and had either n = A2> ovln = ca. 86.
One collection from Stevens Pass in Washington was reported as ^z = 22
by Beamish. The northern Cordilleran population of D. jeffreyi appears to
constitute a uniform cytological race presumably equivalent to an octoploid
race that has lost a pair of chromosomes.

455. Dodecatheon pulchellum (Raf.) Merrill

MORESBY ISLAND: 2n - ca. 88, South Low Island, CTS34831.

Beamish (1955) previously reported counts for this species under D.
radicatum Greene. Plants from Saskatchewan and British Columbia (Fair-
mont Springs and Dutch Creek) had n = 22 or 2n = ca. 44. Other plants
examined from Lulu Island, British Columbia, and Anchorage, Alaska, were
n = ca. 44 and one population from Victoria, British Columbia, was deter-
mined as n = ca. 66.

DOUGLASIA

456. Douglasia laevigata Gray ssp. ciliolata (Constance) Calder & Taylor

Glaux

457. Glaux maritima L. ssp. obtusifolia (Femald) Boivin

GRAHAM ISLAND: 2n = 30, Dawson Inlet, CTS35143; 2n = 30,
Naden Harbour, CT36865.

Previous counts on G. maritima of 2n = 30 have been reported from
Europe (^^e Kress 1963).

Lysimachia

458. *Lysimachia punctata L.

GRAHAM ISLAND: n = 15, Skidegate Village, CT 36954.

The haploid number on the material from the Queen Charlotte Islands
was made from examination of telophase I. Meiosis was regular. Reese (1953)
reported 2n = 30 for this species from Europe.

Trientalis

459. Trientalis europaea L.

GRAHAM ISLAND: n = ca. 44, between 6 and 8 mi SE of Port Cle-
ments, CTS34616.

TRIENTALIS 99

We found it impossible to determine an exact chromosome count for this
species because of irregularities in meiosis. The chromosome number
n = ca. 44 was obtained from cells at anaphase I, a stage that exhibited
lagging chromosomes. Earlier attempts by several authors to determine the
chromosome number of this species have resulted in approximate counts of
2n = ca. 160 (see Love and Love 1961a) and 2n = ca. 170 (Sokolovskaja
1963). It appears that the chromosome number of plants from the Queen
Charlotte Islands is significantly lower than those previously reported for this
species from other areas.

Gentianaceae

Gentiana

460. Gentiana douglasiana Bong.

MORESBY ISLAND: In = 13„, Bigsby Inlet, CST22174; n = 13,
Kootenay Inlet, CT36184.

Meiosis was regular in all material examined.

461. Gentiana platypetala Griseb.

Gentianella

462. Gentianella amarella (L.) Borner ssp. acuta (Michx.) J. M. Gillett

SWERTIA

463. Swertia perennis L.

MORESBY ISLAND: 2n = 28, Mt. Russ, CT36156; 2n = 28, Taka-
kia Lake, CT36265.

This is the same chromosome number reported for this species by Sakai
(1935) and Favarger (1952) on Japanese and European material, respec-
tively. The count of n = 12 or 2« = 24 was obtained by Woycicki (1937)
should be ignored unless it can be confirmed by other workers.

Menyanthaceae

Faltria

464. Fauria crista-galli (Menzies) Makino

MORESBY ISLAND: 2n = ca. 102, Upper Victoria Lake, CT35814.

100 MENYANTHACEAE

The chromosome number of 2n ^ 102 was previously obtained for this
species from material collected at Terrace, British Columbia, by Mulligan
(1965^). Matsuura and Suto (in Darlington and Wylie 1955) reported
2n = 68 for plants of this species from Japan. Their chromosome count prob-
ably should be referred to the species F. japonica Franchet.

Menyanthes

465. Menyanthes trifoliata L.

GRAHAM ISLAND: 2n = ca. 54, Jalun Lake, CT35662.

MORESBY ISLAND: 2n = ca. 54, White Swan Bog, CT35300;
2n = 27ii, Red Mud Marsh, CT35365; 2n = ca. 54, Upper Victoria Lake,
CT35768; 2n = ca. 54, Sunday Inlet, CT36583.

The same chromosome number for this species was reported earlier by
many workers {see Love and Love 1961a, Sorsa 1962, and Love and Ritchie
1966).

Convolvulaceae
Convolvulus

466. *Convolvulus sepium L.

467. Convolvulus soldanella L.

GRAHAM ISLAND: 2n = 22, Tlell, CT35427.

One previous count on the species was made by Kano (1929) in which
n = W was reported.

Polemoniaceae

POLEMONIUM

468. Polemonium pulcherrimum Hook.

Hydrophyllaceae

ROMANZOFFIA

469. Romanzoffia sitchensis Bong.

MORESBY ISLAND: 2n = lln, Takakia Lake, CT23094; 2n = 11„,
Mt. Moresby, CT36396.

The recent review of the chromosome numbers in the Hydrophyllaceae
by Constance (1963) states that all taxa of this small genus have /i = 11.
Previous counts of R. sitchensis were made by Cave and Constance (1950).

AMSINCKIA 101

Boraginaceae

Amsinckia

470. Amsinckia spectabilis Fisch. & Mey.

MORESBY ISLAND: n^ 5, Sandspit, CT35349.

Examination revealed regular meiosis. Ray (1954) and Ray and Chisaki
(1957) reported the same chromosome number for material from Pacific
coast regions.

Lappula

471. *LappuIa echinata Gilib.

MORESBY ISLAND: 2n = 23n, Sandspit, CT36025.

Strey (1931) and Mulligan (1957) reported 2n = 48 on material from
Europe and Canada, respectively. The chromosome count of a7 = 23 on ma-
terial from the Islands suggests either that there are two chromosome races in
this species or that the counts of previous authors may be incorrect.

LiTHOSPERMUM

472. *Lithospermuni officinale L.

Mertensia

473. Mertensia maritima (L.) S. F. Gray

GRAHAM ISLAND: 2n = 12„, Masset Spit, CTS34724.

The same chromosome number was obtained on Icelandic material of
this species by Love and Love (1948, 1956«). This is the first count on
North American plants.

Myosotis

474. *Myosotis arvensis (L.) Hill

GRAHAM ISLAND: 2n = 24„, Queen Charlotte City, CT35844.

Strey (1931 ) reported n = ca. 24 and 2n =^ ca. 48 for European plants
of this species. Other chromosome races of M. arvensis are known to occur in
Europe: 2n = 24 (Mattick in Tischler 1950), 2n = ca. 50 (Sorsa 1962),
2n = 52 (Merxmullcr and Grau 1963) and 2n = 54 (Gcitler 1936, and
Love and Love 1956c/).

102 BORAGINACEAE

475. Myosotis laxa Lchm.

GRAHAM ISLAND: n = ca. 42, near Yakoun River Delta, CT35462.

MORESBY ISLAND: 2n = 42n, between Skidegate Lake and Copper
Bay, CT35276.

This species of Myosotis is probably 12-ploid, based on ;c = 7.

476. *Myosotis scorpioides L.

GRAHAM ISLAND: 2n = 32„, Skidegate Village, CT 36961.

The same chromosome number has been obtained on European material
(^see Love and Love 1961a, and Gadella and Kliphuis 1963).

Labiatae

Galeopsis

477. ^Galeopsis tetrahit L.

GRAHAM ISLAND: n = 16, Masset, CT 36832,

The chromosome number obtained for plants from the Queen Charlotte
Islands was « = 16, the same number that was reported for plants from else-
where in Canada and in Europe {see Mulligan 1959, and Gadella and Kli-
phuis 1963).

Lycopus

478. Lycopus uniflorus Michx.

MORESBY ISLAND: In = 22, Skidegate Lake Bridge, CT35151.

This widely distributed species of North America, Siberia and Japan is
diploid with the base number x = \\.

Mentha

479. Mentha arvensis L.

MORESBY ISLAND: 2az = 36, Sandspit, CT 36039; n = 46, Skidegate
Lake, CT36736.

Many chromosome races occur in European populations of the M. arven-
sis complex, 2n = 12, 64, 60-62, 72, ca. 90 and 92 (see Morton 1956).
It is interesting to note that two chromosome races are found in the limited
population of this species that occurs on the Queen Charlotte Islands. The
Sandspit population growing in a gravel pit may represent an introduction of
European origin, whereas the other populations collected, including the one
from Skidegate Lake, presumably represent native stands of this species.

MENTHA 103

480. Mentha citrata Ehrh.

Prunella

481. Prunella vulgaris L.

GRAHAM ISLAND: 2n = 28, Long Inlet, CT35993.
MORESBY ISLAND: 2n ^ 14„, Mosquito Lake, CT35305.

Two chromosome numbers, 2n = 28 and 2n = 32, have been reported
for this species. According to Bocher (1949), the 2n = 32 counts made by
many workers are incorrect because the somatic chromosomes are of different
sizes and some large bent-over chromosomes have been counted as two chrom-
osomes. In their chromosome list. Love and Love (1961fl) agree with Bocher;
they list the 2n = 32 counts for this species in parentheses. After counting 50
populations of P. vulgaris in North America, Nelson (1964) agreed with
Bocher that the chromosome number for P. vulgaris is 2n = 28 and that the
reported number 2n = 32 is incorrect.

Stachys

482. Stachys cooleyae A. Heller

MORESBY ISLAND: 2n = 32n, Sandspit, CT36035; 2n = 32„, near
Moresby logging camp, CT36704.

Counts of n = 16 have been reported for other native North American
species of this genus (Beaman et al. 1962, and Lewis et al. 1962). On the
basis of these counts and our own from the Queen Charlotte Islands, it would
appear that the genus has a base number of ;c = 8 in North America.

Scrophulariaceae

Castilleja

483. Castilleja hyetophila Pennell

484. Castilleja parviflora Bong.

485. Castilleja unalaschensis (Cham. &Schlecht.) Make

MORESBY ISLAND: n = ca. 48, Tuft Islets, CTS34860; n - 48, be-
tween Cumshewa and Peel inlets, CT35183.

Previous counts on other species' of Castilleja indicate that the base num-
ber for the genus is x - 6 (Beaman et al. 1962, and Hcckard 1964). Castil-
leja unalaschensis is an octoploid and has one of the highest ploidy levels in
the genus in North America.

1 04 SCROPHULARI ACEAE

COLLINSIA

486. Collinsia parviflora Dougl. ex Lindl.

GRAHAM ISLAND: 2n = 14„, Haida Pt., CST20873.

One previous count of /t = 7, a diploid, was reported by Garber (1956)
but the source of the material was not given. He reported at this time that 50
percent of the species of this genus had been counted and all were diploid.
The population from the Queen Charlotte Islands departs from the general
diploid nature of the genus and is a tetraploid.

Digitalis

487. *Digitalis purpurea L.

MORESBY ISLAND: n = 28, Skidegate Lake, CT35281.

Many counts have been made on this species and all authors have re-
ported either n = 2S or 2n = 56 (see Love and Love 1961«).

MiMULUS

488a. Mimulus guttatus DC. ssp. guttatus

GRAHAM ISLAND: 2m = 28, Queen Charlotte City, CST22481.

MORESBY ISLAND: 2n = 28, Kaisun, CT36535; 2n = 28, South
Low Island, Cr57452.

488b. Mimulus guttatus DC. ssp. haidensis Calder & Taylor

GRAHAM ISLAND: 2n = ca. 56, Dawson Inlet, CT35061; 2n = 28n,
Blackwater Creek, CT35131 .

MORESBY ISLAND: 2n = ca. 56, between Cumshewa and Peel inlets,
CT35184; 2n = 56, Takakia Lake, CT 36284; 2n = 56, Mt. Moresby,
CT37337.

Recent extensive studies by Vickery and his students in Utah have shown
that M. guttatus has both diploid (2/t = 28) and tetraploid {2n = 56) popu-
lations {see Mia et al. 1964). However, only one tetraploid count was obtained
in the 40 to 50 populations sampled. The tetraploid was found in Verde Val-
ley, Yavapi County, Arizona. Our studies on the two subspecies in the Queen
Charlotte Islands have revealed that the endemic ssp. haidensis found in the
montane regions is consistently tetraploid, whereas the lowland ssp. guttatus,
which occurs extensively throughout most of western North America, is dip-
loid as has been well documented by Vickery et al.

PEDICULARIS 105

Pedicularis

489. Pedicularis lanata Cham. & Schlecht.

490. Pedicularis oederi Vahl in Homem.

GRAHAM ISLAND: 2n = 16, Jalun Lake, CT35639.

Plants of this species from the Queen Charlotte Islands have the same
chromosome number, 2n = 16, that has been reported for European plants
of this species by Knaben (1950) and Mattick (in Tischler 1950).

491. Pedicularis ornithorhyncha Benth. in Hook.
MORESBY ISLAND: 2n = 8„, Takakia Lake, CT36275.

492. Pedicularis pennellii Hult. ssp. insularis Calder & Taylor

493. Pedicularis verticillata L.

MORESBY ISLAND: 2n = 6„, Takakia Lake, CT3630L

One previous count on this species, on European material, by Favarger
(1963) was reported as 2n = 12. The other two species counted from Queen
Charlotte Islands material were diploid based on x = 8. Pedicularis verticil-
lata is also diploid but has the base number x = 6. Only two base numbers
are known in the genus Pedicularis and plants with both base numbers occur
on the Islands.

Rhinanthus

494. Rhinanthus crista-galli L.

GRAHAM ISLAND: 2n = 7„, Tlell area, CT35927.

Earlier counts on the Rhinanthus crista-galli complex have shown
2n = 22. However, in a discussion of this diploid number by Hambler (1958),
he reports that the chromosome complement is made up of 14 large chrom-
osomes and 8 much smaller chromosomes. Our examination of meiosis of
material from the Queen Charlotte Islands revealed normal meiosis with seven
pairs regularly formed at metaphase I and regular segregation at telophase I
and II. We are recognizing the base number jc = 7 for this western North
American Rhinanthus.

Veronica

495. Veronica americana Schwein. ex Benth. in DC.
MORESBY ISLAND: 2n - 36, Skidegate Mission, CT 36958.

1 06 SCROPHULARI ACEAE

Two previous counts of 2n = 36 have been made on this species (Sch-
lenker 1937, and Sokolovskaja 1963). Sokolovskaja counted material from
Kamchatka.

496. *Veronica arvensis L.

MORESBY ISLAND: 2« = 16, Alliford Bay, CST21054.

Two diploid chromosome numbers have been reported for this species,
2n = 14 and 2n = 16 (see Love and Love 1961a). Material from the Queen
Charlotte Islands was diploid with the base number x = 8.

497. * Veronica filiformis Sm.

498. Veronica peregrina L. ssp. xalapensis (H.B.K.) Pennell

499. Veronica scutellata L.

MORESBY ISLAND: 2« = 18, Red Mud Marsh, CT35364.

All earlier counts of this species are either n =^ 9 or 2n = 18 (see Love
and Love 1961a, and Gadella and Kliphuis 1963).

500. *Veronica serpyllifolia L.

GRAHAM ISLAND: 2n ^ 14, Image Pt., CTS37340.

This species has often been reported to have a diploid number of
2n = 14 (see Love and Love 1961a, and Sokolovskaja 1963).

501. Veronica wormskjoldii R. & S.

MORESBY ISLAND: 2az - 18, Takakia Lake, CT36327.

The Veronica alpina complex which includes V. wormskjoldii is not
clearly understood (see Hulten 1958). The western North American plant
has been recognized as V. wormskjoldii by most authors. Two chromosome
numbers are known for the species, 2« = 36 (Bocher and Larsen 1950,
J0rgensen et al. 1958, Sokolovskaja and Strelkova 1960, and Love and Love
1966) and 2« = 18 (Packer 1964). The diploid number reported by Packer
was determined on plants from Jasper National Park in Western Canada,
whereas the tetraploid count of Love and Love (1966) was obtained on plants
from the eastern United States. The presence of the diploid race on the Queen
Charlotte Islands would indicate that there may be only a diploid race of
this species in Western Canada.

PINGUICULA 107

Lentibulariaceae

PiNGUICULA

502. Pinguicula villosa L.

503. Pinguicula vulgaris L. ssp. macroceras (Link) Calder & Taylor

Utricularia

504. Utricularia intermedia Hayne in Schrader

505. Utricularia minor L.

506. Utricularia vulgaris L.

Plantaginaceae
Plantago

507. *Plantago lanceolata L.

GRAHAM ISLAND: 2n = 12, Masset Spit, CST22639; n = 6, Masset
Spit, CT S3 473 3.

MORESBY ISLAND: n = 6, Sandspit, CT35158.

Many counts have been made on this species and the number most fre-
quently reported is diploid, 2n = \2 (Rahn 1957). The counts of 2n = 24
and 96 by McCullagh (1934) probably refer to other species. Aneuploidy
occurs in the species, with the result that 2n = 13 has been found (see dis-
cussion in Rahn). Meiosis was regular in material examined from the Islands.

508. Plantago macrocarpa Cham. & Schlecht.

MORESBY ISLAND: 2n = 24, Upper Victoria Lake, CT35745.

The same chromosome number has been obtained on material from Van-
couver Island, British Columbia (Bassett 1967).

509. *Plantago major L.

GRAHAM ISLAND: ai - 6, 21/2 mi S of Tlell, CTS34662; 2n = 61,,
2 mi NW of Tlell, CT35690; 2/7 - 12, Tlell, CT3594L

MORESBY ISLAND: 2« = 12, Anna Inlet, CT37472.

1 08 PL ANT AG I N ACE AE

This species has been counted many times on North American, Euro-
pean and Asiatic material and the chromosome number usually reported is
n = 6 or 2n = 12. Three tetraploid counts reported by Ishikawa (1916),
Ikeno (1929) and Takahashi (in Kihara et al. 1931) were on Japanese ma-
terial identified as Plantago major var. asiatica (L.) Decne. It was suggested by
Mulligan (1959) that if tetraploids exist in Canada, they may be expected to
occur along the Pacific coast. The counts on Queen Charlotte Islands material
suggest that Pacific coast populations are diploid.

510. Plantago maritima L.

GRAHAM ISLAND: 2n = \2, Tlell, CST23178; n = 6, Tlell,
CTS34632; n = 6, between Skidegate and Skidegate Village, CTS34670;
n = 6, Dawson Inlet, CTS35142.

MORESBY ISLAND: In = \2, Moresby logging camp, CST21981.

Plantago maritima is a widely distributed polymorphic species with many
segregates. Chromosome numbers reported for the species indicate diploid and
tetraploid races (see Love and Love 1961, and Sorsa 1962). The Queen
Charlotte Islands population is diploid.

Rubiaceae
Galium

511. Galium aparine L.

GRAHAM ISLAND: n = ca. 32, Image Pt., CTS34690; n = ca. 32,
21/2 mi S of Jungle Beach, CT36700.

Many chromosome numbers have been reported for this species:
2n = 42, 44, 63, 64, 66, 86 and 88 (see Love and Love 1961(3, Lewis 1962,
andKliphuis 1962).

512. Galium kamtschaticum Steller ex R. & S.

GRAHAM ISLAND: « = 22, 8 mi SSW of Juskatla, CT35475.

The same chromosome number was reported by Love and Love (1966)
for plants from eastern United States.

513. Galium trifidum L.

GRAHAM ISLAND: 2n = 12„, Lawn Pt, CT35447.

All previous counts on this species have been on European material (see
Love and Love 1961^) and are either n = 12 or 2m = 24.

GALIUM 109

514. Galium triflorum Michx.

GRAHAM ISLAND: 2n == 66, Jungle Beach, CT37357.

MORESBY ISLAND: 2n = 66, Mosquito Lake, CT37509; In = 66,
Crescent Inlet, CT37516.

This circumpolar species has recently been examined cytologically from
three widely separated regions and each region has a different chromosome
race based on x = H. Sorsa (1963) reported 2/t = ca. 44 on material from
Finland, Khoshoo and Bhatia (1963) reported n = \\ from Himalayan
plants, and our own counts from the Queen Charlotte Islands constitute a
hexaploid race.

Sherardia

515. *Sherardia arvensis L.

MORESBY ISLAND: n = l\,2n = 22, Sandspit, CT35328.

Many earlier cytological investigations on this species have consistendy
revealed a diploid number of 2n = 22 (see Love and Love 1961a, and Piotro-
wicz in Skalinska et al. 1964).

Caprifoliaceae

LiNNAEA

516. Linnaea borealis L. ssp. longiflora Hult.

GRAHAM ISLAND: 2n = 32, 15 mi S of Masset, CT35569.

Many counts have been reported for Linnaea borealis L. and all have
been n ^ 16 or 2« = 32 (see Love and Love 1961 a, 1966, Sorsa 1962,
Czapik in Skalinska et al. 1964, Packer 1964, Taylor and Brockman 1966, and
Love and Ritchie 1966). The taxonomic relationship of the western North
American material to the typical subspecies is discussed by Calder and Taylor
in Part 1 .

LONICERA

517. *Lonicera etrusca Santi

518. Lonicera involucrata (Rich.) Banks ex Sprengcl

GRAHAM ISLAND: « = 9, Tlell area, CTS35077; 2n = 18, Tlell area,
CT35434,

One previous count, 2n ^- 18, has been made on L. involucrata by
Janaki-Ammal and Saunders (1952). Mciosis was regular in the material
examined from the Islands.

110 CAPRIFOLIACEAE

Sambucus

519. Sambucus racemosa L. ssp. pubens (Michx.) House

MORESBY ISLAND: 2n = 36, Gray Bay, CT35246.

The same somatic number has been reported by several authors (see
Love and Love 1961, 1966, and Sorsa 1962). A recent report of 2n = 19n
by Taylor and Brockman (1966) from material collected in Yoho National
Park in British Columbia indicates that an increase in the normal chromosome
complement by one pair of chromosomes may have taken place during the
evolution of this tetraploid population.

Symphoricarpos

520. Symphoricarpos albus (L.) Blake

GRAHAM ISLAND: 2n = 36u, between Queen Charlotte City and
Skidegate Village, CT36685.

MORESBY ISLAND: n = 36, Sandspit, CT35351.

This wide-ranging, polymorphic species appears to have more than one
cytological race. Sax and Kribs (1930) reported a hexaploid, n = 27, Parrish
(1957) an octoploid under the name S. rivularis Suksd., and Taylor and
Brockman (1966) a tetraploid from McLean, Saskatchewan. The Queen
Charlotte Islands population, which is very limited in its distribution, is octo-
ploid.

Viburnum

521. Viburnum edule (Michx.) Raf.

Valerianaceae

Plectritis

522. Plectritis congesta (Lindl.) DC.

GRAHAM ISLAND: n = 16, In = 32, Image Pt., CTS34677.
MORESBY ISLAND: 2n - 32, South Low Island, CTS34838.

The same chromosome number was obtained on material of this species
from four locations in southwestern British Columbia by Taylor and Brock-
man (1966). They also counted 2n = 32 for P. macrocera T. & G. from Ver-
non, British Columbia, and proposed the base number .^ = 8 for the genus
Plectritis.

VALERIANA 111

Valeriana

523. Valeriana sitchensis Bong.

GRAHAM ISLAND: n = ca. 48, Long Inlet, CT35956.

The information obtained on other closely related species indicates that
the base number for this genus is x = 8. The Queen Charlotte Islands popu-
lation is 12-ploid based on x = 8 and represents the highest ploidy number
known for the genus. Meiotic examination revealed some univalents at meta-
phase I.

Campanulaceae

Campanula

524. Campanula alaskana (A. Gray) Wight ex J. P. Anderson

MORESBY ISLAND: 2n = 102, Tuft Islets, CTS34861; n = 51,
2n = ca. 102, Kootenay Inlet, CT36235.

Sugiura (1940) reported n = 17 for C. alaskana on material from a
European botanical garden. Other workers have reported both diploids and
tetraploids for the closely related C. rotundiflora, based on x = 17 (Sugiura
1938, Bocher 1938, Flovik 1940, and Vaarama in Love and Love 1948). The
count attributed to this species by Sugiura (1940) may apply to C. rotundi-
folia, because C. alaskana has a distribution restricted to the northern Bering-
ian region. The Queen Charlotte Islands population has the highest number
recorded for the C. rotundijolia complex.

525. Campanula lasiocarpa Cham.

Lobelia

526. Lobelia dortmanna L.

MORESBY ISLAND: In = 14, Mosquito Lake, CT36722.

All previous counts of the species were diploid, In = 14, based on x = 7
(^^eBowden 1959).

Compositae

Achillea

527. Achillea millefolium L.

GRAHAM ISLAND: In = 54, N end of Dawson Inlet, CT35132.

112 COMPOSITAE

MORESBY ISLAND: 2n = 54, Middle Tuft Islet, CT37364; In = 54,
Gowdas Islands, CT37392; 2n = 54, South Low Island, CT37437.

This hexaploid count substantiates the discussion by Calder and Taylor
in Part 1, that the Achillea millefolium complex is represented on the Queen
Charlotte Islands by the native coastal race that is referred to A. borealis
Bong, by many authors.

Anaphalis

528. Anaphalis margaritacea (L.) Benth. & Hook.

GRAHAM ISLAND: 2n = 28, White Creek Muskeg, CT37527.

The same chromosome number has been reported on European and
Asian material by Maude (1940), Sokolovskaja (1960), Love and Love
(1964), and Zhukova (1964). Arano (1956) obtained 2n = 27 on Japanese
plants of this species.

Anthemis

529. *Anthemis cotula L.

MORESBY ISLAND: n = 9, Sandspit, CT36071.

Meiotic examinations show normal meiosis with nine chromosomes pres-
ent in each dyad of telophase I. Other counts on this species have been diploid,
2« = 18 {see Love and Love 1961a).

Apargidium

530. Apargidium boreale (Bong.) T. & G.

GRAHAM ISLAND: 2« - 18, 12 mi N of Pt. Clements, CT 34721;
2n= 18, 4 mi NW of Tlell, CT35455, CT37373.

MORESBY ISLAND: 2n = 18, Echo Harbour, CST22368; 2n = 18,
Anna Inlet, CTS34945.

The only previous count on this species was 2n = 18 by Stebbins et al.
(1953) on material from California. The many counts we have made on the
Queen Charlotte Islands clearly establish the diploid nature of this species.

Arctium

531. *Arctium minus (Hill) Bernh.

GRAHAM ISLAND: 2n = 18n, between Millar Creek and Skidegate
Village, Cr3669M.

ARCTIUM 113

The same chromosome number was reported for Canadian and for Euro-
pean material of this species (see Mulligan 1961). Some of the plants ex-
amined from the Queen Charlotte Islands had one small, weakly stained
chromosome in addition to 18 haploid chromosomes.

Arnica

532. Arnica amplexicaulis Nutt.

GRAHAM ISLAND: n = ca. 28, 2n - 56, Blackwater Creek,
CTS35060; n = ca. 28, Mt. Moresby, CT36440.

MORESBY ISLAND: In = 56, Upper Victoria Lake, CT35815,
CT37383.

Mitotic examination clearly showed 56 somatic chromosomes. It was not
possible to make an accurate count from pollen mother cells as meiosis was
extremely irregular (for example, one cell had 38i -f 9ii at metaphase I).

533. Arnica latifolia Bong.

GRAHAM ISLAND: 2n = 19„, Long Inlet, CT35961.
MORESBY ISLAND: n = ca. 19, Takakia Lake, CT 36296.

This is the same chromosome number that was reported for this species
on material from two locations in Washington by Omduff et al. (1963). Love
and Love (1964) obtained 2n = 112 for plants of A. puberula Rydb. {=A.
latifolia var. gracilis (Rydb.) Cronq.) from Alberta. Queen Charlotte Islands
plants of this species clearly belong to the typical variety.

Artemisia

534. Artemisia arctica Less.

MORESBY ISLAND: 2n = 36, Takakia Lake, CT36309.

Three previous counts have been made on this species. Sokolovskaja

(1963) reported 2n = 36 for material from Kamchatka, Zhukova (1964)
reported 2n = ca. 18 from botanical garden material, and Kawatani and Ohno

(1964) reported 2n = 36 for Japanese material. The Queen Charlotte Islands
population is tetraploid, based on jc = 9..

Aster

535. Aster siibspicatus Nees

114 COMPOSITAE

Bellis

536. *Bellis perennis L.

MORESBY ISLAND: 2n = 9ii, between Sandspit and Cape Chroust-
cheff, CT35159.

The same chromosome number has often been reported for North
American, European and Asiatic material {see Love and Love 1961a, Gadel-
la and KHphuis 1963, and Turner and King 1964).

Chrysanthemum

537. * Chrysanthemum leucanthemum L.

GRAHAM ISLAND: 2n = 9n, 21/2 mi S of Masset, CT35580.

The same diploid chromosome number, based on x = 9, has been ob-
tained on North American and European material of this species (see Love
and Love 1961a). Tetraploid counts reported for this species should be refer-
red to C ircudanum Turcz.

CiRSIUM

538. *Cirsium arvense (L.) Scop.

GRAHAM ISLAND: n = \1 , Jungle Beach, CT35450.

This introduced species has the same chromosome number on the Queen
Charlotte Islands as has been reported for European material {see Love and
Love 1961a, and Gadella and Kliphuis 1963).

539. *Cirsium brevistylum Cronq.

540. *Cirsium vulgare (Savi) Airy-Shaw

MORESBY ISLAND: In = 68, 2 mi S of Cape Chroustcheff, CT37425;
In = 68, Skidegate Lake, CT37501.

The same chromosome number was reported for this species on European
and North American material by many authors {see Moore and Frankton
1962).

COTULA

541. *Cotula coronopifolia L.

GRAHAM ISLAND: 2n = 20, Queen Charlotte City, CT 36939.

Earlier counts on 2n = 20 have been made on C. coronopifolia from dif-
ferent areas: Wulff (1937/?), Rodrigues (1953), Castro and Pontes (1946),
and Hair (1962).

CREPIS 115

Crepis

542. *Crepis capillaris (L.) Wallr.

GRAHAM ISLAND: 2n = 3ii, between Queen Charlotte City and
SkidegatQ, CTS34781.

This species has often been counted (see Babcock 1941a, 1941 b, and
Love and Love 1961a) as 2n = 6. Meiosis was regular in the material ex-
amined from the Queen Charlotte Islands.

Erigeron

543. Erigeron humilis Grah.

MORESBY ISLAND: 2n = 36, Takakia Lake, CT36340.

European and Greenland material of this species has been reported to
have a chromosome number of 2n = 36. Packer (1964) reported the same
number from plants of the Richardson Mountains in Yukon.

544. Erigeron peregrinus (Pursh) Greene

MORESBY ISLAND: 2n = IS, Upper Victoria Lake, CT35722;
2n = 18, Kootenay Inlet, CT36191; 2n = 18, Sunday Inlet, CT36611.

Erigeron peregrinus has recently been counted by Packer (1964) from
plants from Jasper National Park, Alberta. He reported a mitotic chromosome
number of 2« = 18.

Franseria

545. Franseria chamissonis Less.

GRAHAM ISLAND: 2n = 36, Masset Spit, CT 34766.

This species has been consistently reported to have a tetraploid chromo-
some number of 2n = 36 by Wiggins and Stockwell (1937) and Payne
(1964).

Gnaphalium

546. "^Gnaphalium uliginosum L.

GRAHAM ISLAND: 2/7 = 7„, about 21/2 mi S of Tlell, €735947.

The same chromosome number obtained on material from the Queen
Charlotte Islands was reported for European and Asiatic material of this
species by WulfT (1938), Love and Love (1956a), and Arano and Nakamura
(1964).

116 COMPOSITAE

Grindelia

547. Grindelia integrifolia DC.

GRAHAM ISLAND: 2n = 24, Delkatla Inlet, CT35594.
MORESBY ISLAND: In =- 24, head of Cumshewa Inlet, CT2368I.

In a recent study on Grindelia, Dunford (1964) reported a tetraploid
number of 2n = 24 for G. striata DC., a species we consider synonymous
with G. integrifolia. His count was made on material from Marin County in
California.

HiERACIUM

548. Hieracium albiflorum Hook.

GRAHAM ISLAND: 2n = 9n, between Queen Charlotte City and
Skidegate Village, CTS34786.

For discussion, see H. triste Willd.

549. * Hieracium aurantiacum L.

MORESBY ISLAND: In = 36, Alliford Bay, €736090.

Most earlier counts of this adventive of European origin have been re-
ported as «= 18 or 2/7 = 36 {see Love and Love \96\a). A recent report
on cultivated garden material by Zhukova (1964) gives a somatic count from
root tips as 2/2 = 30.

550. Hieracium triste Willd. ex Sprengel

MORESBY ISLAND: In = 9u, Mt. Moresby, CT 36457; In = 9n,
Takakia Lake, CT 36281.

Material of H. albiflorum Hook, and H. triste from the Queen Charlotte
Islands was regular at all stages of meiosis. Old World species of this genus
are known to be apomictic. The regular meiosis in Queen Charlotte Islands
plants suggests that North American species of this genus are sexual. Further
cytological and breeding studies on New World species of Hieracium are
needed to determine the extent of apomixis present on this continent. Sokolov-
skaja (1963) reported 2/t = 18 for H. triste from Kamchatka.

Hypochaeris

551. *Hypochaeris radicata L.

GRAHAM ISLAND: 2/7 = 8, between Queen Charlotte City and Skide-
gate Village, CTJJiPi.

HYPOCHAERIS 117

This species has often been reported to be diploid, 2n = S (see Mulligan
1957, Reiser 1963, and Turner and King 1964).

Lapsana

552. ^Lapsana communis L.

GRAHAM ISLAND: 2n - 7„, Queen Charlotte City, CT35845.

Several chromosome numbers have been reported for this species,
2n = 12, 14 and 16 {see Sorsa 1963). All three chromosome races are known
to occur in Europe; however, plants with only 14 somatic chromosomes are
known from North America.

Leontodon

553. *Leontodon nudicaulis (L.) Merat. ssp. taraxacoides (Vill.) Shinz &
Thellung

MORESBY ISLAND: In = 8, Alliford Bay, CT 36091.

In a discussion of the cytology of Leontodon by Stebbins et al. (1953),
the numbers reported for L. nudicaulis (L.) Merat. are In = ^ and In = 10.
However, the count of 2/? = 10 obtained by Wulff (1937/?) on botanical
garden material may be an error. Stebbins et al. reported 2« = 8 for material
from California.

Matricaria

554. ^Matricaria matricarioides (Less.) Porter

GRAHAM ISLAND: 2n = 9„, Tlell, CT35917.

MORESBY ISLAND: 2n = 9„, Sandspit, CT 36038; /? = 9, Horn
Rock, CT36519.

The same chromosome number was previously reported for North Ameri-
can and European material by many workers (see Love and Love \96\a).

Petasites

555. Petasites nivalis Greene

Prenanthes

556. Prenanthes alata (Hook.) D. Dictr.

GRAHAM ISLAND: 2n = 16, Millar Creek, CST23455; 2n 8„,
Blackwater Creek, CT S3 5069; 2n - 16, N end of Long Inlet, CT36010.

118 COMPOSITAE

MORESBY ISLAND: 2n = 16, Kaisun, CT36532; In = 16, between
Cumshewa and Peel inlets, CT37458.

The base numbers proposed for this genus are x = 8 and 9 {see Bab-
cock et al. 1937). The Queen Charlotte Islands population is diploid, based
on X = 8.

Senecio

557. Senecio cymbalarioides Nutt. ssp. moresbiensis Calder & Taylor

MORESBY ISLAND: In = 45ii, Bigsby Inlet, CTS34887; 2n = ca. 90,
Takakia Lake, CT36323.

The meiotic examination of this material revealed regular meiosis with
45 bivalents formed at metaphase I and regular separation at telophase I and
II. The previously reported number of the more southern typical subspecies
from California and Washington material was n = 23 (Omduff et al. 1963).

558. Senecio newcombei Greene

GRAHAM ISLAND: n = 24, Jalun Lake, CT35665.

MORESBY ISLAND: 2n = 24ii, Bigsby Inlet, CTS34888; 2n = 24n,
Takakia Lake, CT 36271 .

This species, like Saxifraga taylori Calder & Savile, Isopyrum savilei
Calder & Taylor, and Ligusticum calderi Math. & Const., belongs to a small
group of endemics from the Queen Charlotte Islands that are related to more
southern North American floristic elements. Calder and Taylor in Part 1 state
that ''Senecio newcombei was included by Barkley (1962) in the polytypic
section Aurei Rydb., and though its relationships within this section are not
clearly understood it appears to be rather closely related to the monocephalus
S. ported Greene of Colorado and Oregon." Omduff et al. (1963) report
chromosome counts for a number of species of section Aurei and all these
species had a gametic chromosome number of 23 or 46. Ornduff et al. suggest
that the basic number for the Senecioneae is x = 10 and that haploid numbers
higher than this are to be regarded as polyploid or aneuploid derivatives. Our
material from Jalun Lake most often had the configuration 22ii + In- at
diakinesis and metaphase I of meiosis.

559. Senecio pseudo-arnica Less.

GRAHAM ISLAND: 2n = 20„, Sangan River, CT35596; 2n = ca. 40,
about 2 mi S of Rose Spit, CT35910.

Three previous counts on this species have revealed 2n = 40 (Afzelius
1949, Sokolovskaja 1963, and Arano 1964). A single count of 2n = 36 was
reported by Zhukova (1964) on cultivated material.

SENECIO 119

560. *Senecio sylvaticus L.

GRAHAM ISLAND: 2n = 20„, W of Queen Charlotte City, CT36936.
MORESBY ISLAND: 2n = 20n, Dass Pt., CTS35024.

In a recent review of the chromosome numbers in the Senecioneae, Om-
duff et al. (1963) reported n = 20 from introduced plants of Washington.
Afzelius (1924) gave the same chromosome number for European material.

561. Senecio triangularis Hook.

MORESBY ISLAND: n = 40, Bigsby Inlet, CTS34886; 2n = ca. 80,
Takakia Lake, CT37545.

The recent study by Omduff et al. (1963) revealed that there are two
ploidy levels in this species. They reported that plants from Whatcom County,
Washington, and Nevada County, California, were n = 20. Packer (1964)
obtained the same number for plants from Jasper National Park in Alberta.
Plants with n = 40 were reported by Omduff et al. from Clallam County,
Washington, and Clackamas County, Oregon. A single count oi n = 20 was
given for plants from western North America by Afzelius (1949).

562. * Senecio vulgaris L.

GRAHAM ISLAND: 2n = 40, Masset Spit, CST22635; 2n = 20„,
Masset Spit, CTS34725.

MORESBY ISLAND: 2n = 20„, Sandspit, CT 36027; 2n = 40, Cres-
cent Inlet, CT37478.

This widely distributed adventive has been counted numerous times from
many regions of Europe and North America and all have n ^ 20 or 2n ^ 40
(see Mulligan 1 96 1 , and Omduff etal. 1963).

SOLIDAGO

563. Solidago canadensis L.

GRAHAM ISLAND: 2n = 36, Tlell, CT35934.

The base number of x = 9 has been proposed by Beaudry and Chabot
(1957). Many diploid counts of this species have been given for eastern North
American plants by Beaudry and Chabot (1957, 1959). Raven et al. (1960)
have reported numerous diploid counts, but one hexaploid count of n = 27
was noted on a subspecies of .S\ canadensis from Oregon. Our count from the
Queen Charlotte Islands represents the first tetraploid reported for S. cana-
densis. It is apparent that several ploidy levels exist in the western races of

120 COMPOSITAE

this widely distributed polymorphic species. Cytological investigations of a
similar nature to those conducted by Beaudry on eastern North American
populations should be conducted on the species in the Pacific Northwest. Per-
haps such studies could provide some insight for the taxonomy of this species,
as it has not been clearly analyzed for the western populations (see discussion
of taxonomy by Calder and Taylor in Part 1 ) .

564. Solidago multiradiata Ait.

SONCHUS

565. *Sonchus asper (L.) Hill

GRAHAM ISLAND: 2n = 9ii, between Queen Charlotte City and
Skidegate Village, CTS34780.

All previous counts reported for this species are either n = 9 or 2n = 18
(see Love and Love 1961a. and Koul 1964).

Tanacetum

566. Tanacetum huronense Nutt.

GRAHAM ISLAND: 2n = 54, Tlell, CST23248; 2n = 27„, Tlell,
CTS35078; 2n = llu, mouth of Sangan River, CT35598.

The base number for this genus \s x = 9 and the Queen Charlotte Islands
material forms part of a hexaploid population.

567. *Tanacetum vulgare L.

GRAHAM ISLAND: 2n = 9„, west of Queen Charlotte City, CT36923.

The diploid number 2n = 18 has been consistently reported in many
studies of European and Asiatic material (see Love and Love 1961 a, and
Sokolovskaja and Strelkova 1962).

Taraxacum

568. *Taraxacum officinale Weber in Wiggers

GRAHAM ISLAND: 2n = 24, Honna River, CT37394.
MORESBY ISLAND: 2n = 24, Sandspit, CT 36041.

The chromosome number 2n = 24 has often been reported on European
and North American material of this species (see Love and Love 1961a).
Fiimkranz (1960) also obtained the numbers 2n = 16, 18, 24, 32, 34, 36,
and 37 for Taraxacum officinale.

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INDEX

Two typefaces are used:

Boldface — Plant entity found on the Queen Charlotte Islands and page reference to
the page on which its chromosome number is reported.

Roman — Plant entity not found in the flora but referred to in a cytological discus-
sion, page reference to Queen Charlotte taxon that has not been counted, and page refer-
ence to a count that has been reported for a taxon not found on the Islands.

Abronia, 58

latifolia, 58
Achillea, 111

borealis, 1 12

millefolium, 111
Aconitum, 63

delphinifolium, 63
Actaea, 63

rubra

ssp. arguta, 63
Adiantum, 13

pedatum

ssp. aleuticum, 13
Agropyron, 21

repens, 21
Agrostis, 21

aequivalvis, 21

borealis, 22

exarata, 22

gigantea, 22

pallens, 22

palustris, 22

scabra, 22

thurberiana, 21, 23
Aira

caryophyllea, 23

praecox, 23
Allium, 46

schoenoprasum, 46
AInus. 52

crispa, 52

ssp. sinuata, 52

rubra, 53
Alopecurus, 23

geniculatus, 23
Amelanchier, 79

alnifolia, 79

florida, 79
Ammophila, 24

arenaria, 24
Amsinckia, 101

spectabilis, 101

Anaphalis, 1 12

margaritacea, 112
Andromeda, 96

polifolia, 96
Anemone, 63

multifida, 63

narcissiflora

ssp. alaskana, 63

parviflora, 63
Angelica, 93

lucida, 93
Anthemis, 1 12

cotula, 112
Anthoxanthum, 24

odoratum, 24
Apargidium, 112

boreale, 112
Aphanes, 80

occidentalis, 80
Aquilegia, 64

formosa, 64

truncata, 64
Arabis, 67

glabra, 67

hirsuta, 68

ssp. eschscholtziana, 68

lyrata, 68

ssp. kamchatica, 68
ARACEAE, 42
ARALIACEAE, 93
Arceuthobium, 53

campy lopodum, 53
Arctium, 1 12

minus, 112
Arctostaphylos, 96

uva-ursi, 96
Arenaria, 59

lateriflora, 59

peploides

ssp. major, 59

slricla

ssp. macra, 59

140

INDEX

Arnica, 1 13

amplcxicaulis, 1 13
latifolia, 113

var. gracilis, 1 13

puberla, 1 13
Artemisia, 1 13

arctica, 113
Aruncus, 80

Sylvester, 80
Asplenium, 14

trichomanes, 14

viride, 14
Aster, 113

subspicatus, 113
Athyrium, 14

filix-femina

ssp. cyclosorum, 14
var. pectinata, 14
Atriplex, 57

hastata, 57

patula, 57

ssp. obtusa, 57
Avena, 24

fatua, 24

Barbarea, 68

ortboceras, 68
Bellis, 114

percnnis, 114
BETULACEAE, 52
Blechnum, 14

spicant, 14
BORAGINACEAE, 101
Botrychium, 13

lunaria

ssp. lunaria, 13
ssp. minganense, 13

multifidum, 13

ssp. silaifolium, 13
Brassica, 69

campestris, 69
Bromus, 24

mollis, 24

pacificus, 24

sitchensis, 25

Cakile, 69

californica, 69

edentula, 69

lacustris, 69

maritima, 69
Calamagrostis, 25

canadensis, 25

crassiglumis, 25

nutkaensis, 25

purpurascens, 26
ssp. tasuensis, 25

CALLITRICHACEAE, 88
Callitriche, 88

heterophylla

ssp. bolanderi, 88
Caltha, 64

biflora, 64

leptosepala, 64

palustris

ssp. asariflora, 64
Calypso, 49

bulbosa

ssp. occidentalis, 49
Camelina, 69

sativa, 69
Campanula, 1 1 1

alaskana, 111

lasiocarpa, 1 1 1

rotundifolia, 1 1 1
CAMPANULACEAE, HI
CARPIFOLIACEAE, 111
Capsella, 69

bursa-pastoris, 69
Cardamine, 70

angulata, 70

bellidifolia, 70

occidentalis, 70

oligosperma, 70, 71

pensylvanica, 71

umbellata, 70, 71
Carex, 35

anthoxantbea, 35

arcta, 35

arenicola

ssp. pansa, 35

brevicaulis, 36

brunnescens, 36

biixbaumii, 36

canescens, 36

ssp. arctaeformis, 36
ssp. canescens, 36

circinata, 36

cusickii, 37

deweyana, 37

ssp. leptopoda, 37

disperma, 37

enanderi, 37

exsiccata, 37

glareosa, 37

gmelinii, 37

kelloggii, 37

laeviculmis, 37

leptalea

ssp. pacifica, 38

livida, 38

lyngbyei, 38

macloviana

ssp. pachystachya, 38

INDEX

141

macrocephala, 38

macrocheata, 38

mertensii, 39

nigricans, 39

obnupta, 39

pachystachya, 38

pauciflora, 39

phyllomanica, 39

physocarpa, 39

pluriflora, 39

pyrenaica

ssp. micropoda, 39

scirpoides, 40

sitchensis, 40

stylosa, 40

tracyi, 40

viridula, 40
CARYOPHYLLACEAE, 59
Cassiope, 96

lycopodioides

ssp. cristapilosa, 96

martensiana, 96

stellariana, 96
Castilleja, 103

hyetophila, 103

parviflora, 103

unalaschensis, 103
Cerastium, 59

arvense, 59

fisherianum, 60

tomentosum, 60

viscosum, 60

vulgafum, 60
Chamaecyparis, 18

nootkatensis, 18
CHENOPODIACEAE, 57
Chenopodium, 57

album, 57
Chrysanthemum, 114

ircutianum, 1 14

leucanthemum, 114
Cinna, 26

latifolia, 26
Circaea, 90

aipina

ssp. aipina, 90
ssp. pacifica, 90
Cirsium, 1 14

arvense, 114

brevistylum, 1 14

vulgare, 1 14
Cladothamnus, 96

pyrolaeflorus, 96
Cochlearis, 71

officinalis

ssp. oblongifolia, 71
Coilinsia, 104

parviflora, 104

COMPOSITAE, 111
Conioselinum, 93

chinense, 93

pacificum, 93
CONVOLVULACEAE, 100
Convolvulus, 100

sepiuin, 100

soldanella, 100
Coptis, 46

asplenifolia, 64

trifolia, 64
Corallorhiza, 49

innata, 49

maculata

ssp. mertensiana, 49

trifida, 49
CORNACEAE, 95
Cornus, 95

canadensis, 95

intermedia, 95

stolonifera, 95

unalaschkensis, 95
Cotula, 114

coronopifolia, 114
CRASSULACEAE, 74
Crataegus, 80

douglasii, 80

monogyna, 80
Crepis, 1 15

capillaris, 115
CRUCIFERAE, 67
Cryptogramma, 14

crispa

ssp. acrosfichoides, 14
var. acrostichoides, 15
CUPRESSACEAE, 18
Cynosurus, 26

cristatus, 26
CYPERACEAE, 35
Cystoperis, 15

fragilis, 15
Cytisus, 85

scoparius, 85

Dactylis, 26
glomerata, 26

Danthonia, 26

californica, 26

intermedia, 27
Deschampsia, 27

caespitosa, 27

ssp. beringensis, 27

elongata, 27
Descurainia, 72

sophia, 72

142

INDEX

Digitalis. 104

purpurea, 104
Dodecatheon, 97

jcffreyi, 97

ssp. jefFreyi, 98

pulchellum, 98
Douglasia, 98

laevigata

ssp. ciliolata, 98
Draba, 72

hyperborea, 72

lonchocarpa, 72

ssp. kamtschatica, 72
Drosera, 74

rotundifolia, 74
DROSERACEAE, 74
Dryopteris, 15

austriaca, 15

Eleocharis, 40

acicularis, 40

kamtschatica, 40

macrostachya, 40

palustris, 41

obtusa, 41
Elodea, 21

canadensis, 21
X Elymordeum

schaakianum, 28
Elymus, 28

glaucus, 28

ssp. glaucus, 28
ssp. virescens, 28

hirsutus, 28

hirsutus x Hordeum
brachyantherum, 28

mollis, 28
EMPETRACEAE, 89
Empetrum, 89

nigrum, 89
Epjlobium 91

anagallidifolium, 91

angustifolium, 91

ssp. circumvagum, 91

brevistylum, 91

delicatum, 91

glandulosum, 91

latifolium, 92

minutum, 92

palustre, 92

platyphyllum, 91
EQUISETACEAE, 12
Equisetum, 12

arvense, 12

fluviatile, 12

hyemale

ssp. affine, 12

hyemale x E. variegatum, 12

palustre, 12

telmateia, 12

variegatum

ssp. alaskanum, 12
ERICACEAE, 96
Erigeron, 1 15

humilis, 115

peregrinus, 115
Eriophorum, 41

angustifolium, 41

chamissonis, 41

chamissonis X E. russeolum, 41
Erysimum, 72

chieranthoides, 72

Fauria, 99

crista-galli, 99

japonica, 100
Festuca, 29

arundinacea, 29

dertonensis, 29

elatior, 29

megalura, 29

myuros, 29

occidentalis, 29

prolifera, 30

rubra, 30

subulata, 30
Fragaria, 80

chiloensis

ssp. lucida, 80
ssp. pacifica, 80
Franseria, 1 15

chamissonis, 115
Fritillaria, 46

camschatcensis, 46
ssp. alpina, 46

Galeopsis, 102

tetrahit, 102
Galium, 108

aparine, 108

kamtschaticum, 108

trifidum, 108

triflorum, 109
Gaullheria, 96

shallon, 96
Gentiana, 99

douglasiana, 99

platypetala, 99
GENTIANACEAE, 99
Gentianella, 99

amarella

ssp. acuta, 99
GERANIACEAE, 88

dissectum, 88

INDEX

143

molle, 88
richardsonii, 88

Geum, 81

calthifolium, 81
macrophyllum, 81
schoiieldii, 81

Glaux, 98

maritima, 98

ssp. obtusifolia, 98
Glehnia, 93

littoralis

ssp. leiocarpa, 93
Glyceria, 30

occidentalis, 30
Gnaphalium, 115

uliginosum, 115
Goodyera, 49

oblongifolia, 49
GRAMINEAE, 21
Grindelia, 116

integrifolia, 116

stricta, 116
Gymnocarpium, 15

dryopteris, 15

Habenaria, 50

chorisiana, 50

dilatata, 50

saccata, 50

unalascensis

ssp. maritima, 50
ssp. unalascensis, 50
HALORAGIDACEAE, 92
Heracleum, 94

lanatum, 94
Hesperis, 72

matronalis, 72
Heuchera, 75

chlorantha, 75

glabra, 75
Hieracium, 1 16

albiflorum, 116

aurantiacum, 116

triste, 116
Hierochloe, 31

odorata, 31
HIPPURIDACEAE, 92
Hippuris, 92

montana, 92

vulgaris, 92
Holcus, 31

lanatus, 31
Honckenya

peploides, 59
Hordeum, 31

brachyantherum, 31
Humulus, 53

lupulus, 53
HYDROCHARITACEAE, 21
HYDROPHYLLACEAE, 100
HYMENOPHYLLACEAE, 13
Hypochaeris, 1 16

radicata, 116
Hypopitys, 95

monotropa, 95

IRIDACEAE, 49
ISOETACEAE, 13
Isoetes, 13

echinospora

ssp. muricata, 13
Isopyrum, 65

savilei, 65

JUNCACEAE, 42
JUNCAGINACEAE, 20
Juncus, 42
albescens. 45
alpinus, 42

ssp. nodulosus, 42
arcticus, 42

ssp. ater, 43

ssp. sitchensis, 42
articulatus, 43
bufonius, 43
covillei, 44
drummondii, 43
effusus, 43

var. brunneus, 43

var. gracilis, 43

var. pacificus, 43
ensifolius, 43
falcatus, 44
filiformis, 44
leseurii, 44
mertensianus, 44
oreganus, 44
tenuis, 44
triglumis, 45
Juniperus, 18
communis, 18

Kalmia, 96
polifolia, 96

LABIATAE. 102
Lappula, 101

echinata, 101
Lapsana, 1 17

communis, 1 17
Lathyriis. 85

japonicus, 85

littoralis, 85

ochrolcucus, 85

palustris, 85

144

INDEX

Ledum, 96

palustre

ssp. j^roenlandicum, 96
LEGUMINOSAE, 85
LENTIBULARIACEAE, 107
Leontodon, 1 17

nudicaulis, 1 17

ssp. taraxacoides, 117
Lepidium, 73

canipestre, 73

densiflorum, 73
Leptarrhena, 75

pyrolifolia, 75
Ligusticum, 94

calderi, 94

filiciniim, 94

scoticum

ssp. hultenii, 94
Lilaeopsis, 94

occidentalis, 94
LILIACEAE, 46
LINACEAE, 88
Linnaea, 109

borealis, 109

ssp. longiflora, 109
Linum, 88

bienne, 88
Listera, 50

caurina, 50

cordata, 51
Lithospermum, 101

officinale, 101
Lloydia, 47

serotina

ssp. flava, 47
Lobelia, 1 1 1

dortmanna, 111
Loiseleuria, 96

procumbens, 96
Lolium, 31

perenne, 31
Lonicera, 109

etrusca, 109

involucrata, 109
LORANTHACEAE, 53
Luetkea, 81

pectinata, 81
Lupinus, 85

littoralis, 85

nootkatensis, 86
Luzula, 45

mukiflora, 45
ssp. comosa, 45

parviffora, 46
LYCOPODIACEAE, 12
Lycopodium, 12

annotinum, 12

clavafum, 12
coniplanafum, 12
iniindatum, 12
obscurum, 12

sabinaefolium

ssp. sitchense, 12
selago

ssp. miyoshianum, 13
ssp. patens, 13
ssp. selago, 12
Lycopus, 102

uniflorus, 102
Lysichiton, 42
americanum, 42
camtschatcense, 42
Lysimachia, 98
punctata, 98

Maianthemum, 47

dilatatum, 47
Malaxis, 51

paludosa, 51
Matricaria, 117

matricarioides, 117
Mecodium, 13

wrightii, 13
Melica, 31

subulata, 31
Melilotus, 86

alba, 86

officinalis, 86
Mentha, 102

arvensis, 102

citrata, 103
MENYANTHACEAE, 99
Menyanthes, 100

trifoliata, 100
Menziesia, 97

ferruginea, 97
Mertensia, 101

maritima, 101
Mimulus, 104

guttatus, 104

ssp. guttatus, 104
ssp. haidensis, 104
Mitella, 76

pentandra, 76
Moneses, 96

uniflora

ssp. reticulata, 96
Montia, 58

fontana, 58

parvifolia, 58

sibirica, 58
MORACEAE, 53
Myosotis, 101

arvensis, 101

INDEX

145

laxa, 102
scorpioides, 102

Myrica, 52

gale, 52
MYRICACEAE, 52
Myriophyllum, 92

spicatum, 92

Neslia, 73

paniculata, 73
Nuphar, 62

luteum, 63

ssp. polysepalum, 62
NYCTAGINACEAE, 58
NYMPHAEACEAE, 62

Oenanthe, 94

sarmentosa, 94
ONAGRACEAE, 90
OPHIOGLOSSACEAE, 13
Oplopanax, 93

horridus, 93
ORCHIDACEAE, 49
Osmorhiza, 95

cbilensis, 95

purpurea, 95
Oxyria, 53

digyna, 53
Oxytropis, 86

campestris, 86

Parnassia, 76

fimbriata, 76
Pedicularis, 105

lanata, 105

oederi, 105

ornithorhyncha, 105

pennellii

ssp. insularis, 105

verticellata, 105
Petasites, 1 17

nivalis, 117
Phalaris, 32

arundinacea, 32
Phleum, 32

alpinum, 32

pratcnse, 32
Phyllodoce, 97

glanduliflora, 97
Phyllospadix, 19

scouleri, 19
Picea, 17

sitchensis, 17
PINACEAE, 17
Pinguicula, 107

vHlosa, 107

vulgaris

ssp. niacroccras, 107

Pinus, 17

conforta, 17
PLANTAGINACEAE, 107
Plantago, 107

lanceolata, 107

macrocarpa, 107

major, 107

var. asiatica, 108

maritima, 108
Plectritis, 110

congesta, 110

macrocera, 1 10
Poa, 32

annua, 32

confinis, 32

douglasii

ssp. macrantha, 32

interior, 33

laxiflora, 33

leptocoma, 33

pratensis, 33

stenantba, 33

trivialis, 33
POLEMONIACEAE, 100
Polemonium, 100

pulcherrimum, 100
POLYGONACEAE, 53
Polygonum, 54

arenastrum, 54

aviculare, 54

convolvulus, 54

cuspidatum, 55

fowleri, 54

persicaria, 55

polystachyum, 55

scabrum, 55

vivparum, 55
POLYPODIACEAE, 13
Polypodium, 15

scouleri, 15

vulgare, 16

ssp. columbianum, 16
ssp. occidentale, 15
Polystichum, 16

braunii, 16

ssp. alaskense, 16
ssp. andersonii, 16
ssp. pursbii, 16

lonchitis, 16

munitum, 16
PORTULACACEAE, 58
Potamogeton, 19

alpinus

ssp. tenuifolius, 19

berchtoldii

ssp. bcrcbfoldii, 19
ssp. tcnuissimus, 19

146

INDEX

cpihydrus

ssp. nuttallii, 19

fluitans, 20

^ramincus, 19

nafans, 20

nodosus, 20

pectinatus, 20

pusillus, 19

richardsonii, 20

robbinsii, 20
Potentilla, 81

anserina, 82
ssp. pacifica, 82

crantzii, 82

pacifica, 81

palustris, 82

villosa, 82
Prenanthes, 1 17

alata, 117
PRIMULACEAE, 97
Prunella, 103

vulgaris, 103
Pteridium, 16

aquilinum

ssp. aquilinum

var. pubescens, 16
Puccinellia, 33

borealis, 33

interior, 34

nutkaensis, 34

pumila, 34
Pyrola, 96

secunda, 96
PYROLACEAE, 95
Pyrus 82

fusca, 82

RANUNCULACEAE, 63
Ranunculus, 65

acris, 65

aquatilis, 65

circinatus, 65

cooleyae, 65

eschscholtzii, 66

flammula, 66

var. filiformis, 66
var. flammula, 66
var. ovalis, 66

hyperboreus, 66

occidentalis, 67

ssp. occidentalis, 66

orthorhynchus, 67

pygmaeus, 67

repcns, 67

uncinatus, 67
Raphanus, 73

raphanistrum, 73
Rhinanthus, 105

crista-galli, 105

Rhynchospora, 41

alba, 41
Ribes, 76

bracteosum, 76

lacustre, 76

laxiflorum, 76
Romanzoffia, 100

sitchensis, 100
Rorippa, 73

islandica, 73
Rosa, 82

nutkana, 82
ROSACEAE, 79
RUBIACEAE, 108
Rubus, 82

chamaemorus, 82

laciniatus, 82

parviflorus, 83

pedatus, 83

procerus, 83

spectabilis, 83

ursinus, 83
Rumex, 55

acetosella, 55

angiocarpus, 56

crispus, 56

graminifolius, 56

obtusifolius, 56

occidentalis, 56

tenuifolius, 56

transitorius, 56
Ruppia, 20

maritima, 20

spiralis, 20

Sagina, 60
maxima, 60
procumbens, 60

SALICACEAE, 51
Salicornia, 57
pacifica, 57
Salix, 51

hookeriana, 51
lasiandra, 52
reticulata

ssp. glabellicarpa, 52
scouleriana, 52
sitchensis, 52
Sambucus, 1 10
racemosa

ssp. pubcns, 110
Sanguisorba, 83
canadensis, 83

ssp. latifolia, 83
menziesii, 83
officinalis, 84

ssp. microcephala, 84

INDEX

147

Sanicula, 95

crassicaulis, 95
Saxifraga, 77

caespitosa, 77

ferruginea, 77

lyallii

ssp. hultenii, 77

mertensiana, 77

odontoloma, 77

oppositifolia, 78

punctata, 78

ssp. carlottae, 78
ssp. cascadensis, 78
ssp. nelsonjana, 78
ssp. porsildiana, 78

taylori, 78

tolmiei, 79
SAXIFRAGACEAE, 75
Scirpus, 41

atrocinctus, 41

cernuus, 41

cespitosus, 41

lacustris

ssp. glaucus, 42

palustris, 41

sylvaticus, 42
ssp. digynus, 42
SCROPHULARIACEAE, 103
Sedum, 74

divergens, 74

roseum, 75
Selaginella, 13

selaginoides, 13

wallacei, 13
SELAGINELLACEAE, 13
Senecio, 118

cymbalarioides

ssp. moresbiensis, 118

newcombei, 118

porteri, 1 18

pseudo-arnica, 118

sylvaticus, 119

triangularis, 119

vulgaris, 119
Sherardia, 109

arvensis, 109
Sibbaldia, 84

procumbens, 84
Silene, 61

acaulis

ssp. subacaulescens, 61

noctiflora, 61
Sinapis, 73

arvensis, 73
Sisymbrium, 73

altissimum, 73

officinale, 74
Sisyrinchium, 49

littorale, 49

montanum, 49
Solidago, 119

canadensis, 119

inultiradiata, 120
Sonchus, 120

aspcr, 120
Sorbus, 84

aucuparia, 84

sitchensis

ssp. grayi, 84
ssp. sitchensis, 84
SPARC ANIACEAE, i:
Sparganium, 18

hyperboreum, 18

minimum 18

simplex, 19
Spergula, 61

arvensis, 61
Spergularia, 61

canadensis, 61
Spiraea, 84

douglasii, 85

ssp. douglasii, 85
ssp. menziesii, 84

menziesii, 85
Spiranthes, 51

romanzoffiana, 51
Stachys, 103

cooleyae, 103
Stellaria, 61

calycantha, 61

crispa, 62

graminea, 62

humifusa, 62

longipes, 62

media, 62
Streptopus, 47

amplexifolius, 47

roseus

ssp. curvipes, 48

streptopoides
ssp. brevipes, 48
Suaeda, 58

depressa, 58

maritima, 58
Subularia, 74

aquatica

ssp. americana, 74
ssp. aquatica, 74

monticola, 74
Swertia, 99

perennis, 99
Symphoricarpos, 110

albus, 110

rivularis, 1 10

Tanacetum, 120

148

INDEX

hiironcnsc, 120
vul^are, 120

Taraxacum, 120

officinale, 120
TAXACEAE, 17
Taxus, 17

brevifolia, 17
Tellima, 79

grandiflora, 79
Thalictruni, 67

alpinum, 67
Thelypteris, 17

oreopteris, 17

pliegopteris, 17
Thlaspi, 74

arvense, 74
Thuja, 18

pllcata, 18
Tiarella, 79

trifoliata, 79

ssp. unifoliata, 79

unifoliata, 79
Tofieldia, 48

glutinosa

ssp. brevistyla, 48
ssp. glutinosa, 48
Tolmiea, 79

menziesii, 79
Torreyochloa, 34

pauciflora, 34
Trientalis, 98

europaea, 98
Trifolium, 86

dubium, 86

pratense, 86

repens, 86

wormskjoldii, 87
Triglochin, 20

concinnum, 21

debile, 21

elatum, 21

maritimum, 20

palustre, 21
Trisetum, 34

cernuum, 34

ssp. canescens, 35

spicatum, 35
Tsuga, 17

heterophylla, 17

mertensiana, 18

Ulex, 87

europaeus, 87
UMBELLIFERAE, 93
Urtica, 53

dioica, 53
URTICACEAE, 53
Urticularia, 107

intermedia, 107
minor, 107
vulgaris, 107

Vaccinium, 97

alaskense, 97

caespitosuni, 97

ovalifolium, 97

oxycoccus, 97

parvifolium, 97

scoparium, 97

uliginosum, 97

vitis-idaea

ssp. minus, 97
Vahlodea, 35

atropurpurea

ssp. paramushirensis, 35
Valeriana, 1 1 1

sitchensis, 111
VALERIAN ACEAE, 110
Veratrum, 48

eschscholtzii, 48

viride, 49
Veronica, 105

alpina, 106

americana, 105

arvensis, 106

filiformis, 106

peregrina

ssp. xalpensis, 106

scutellata, 106

serpyllifolia, 106

wormskjoldii, 106
Viburnum, 110

edule, 110
Vicia, 87

cracca, 87

gigantea, 87

sativa, 87

villosa, 88
VIOLACEAE, 89
Viola, 89

adunca, 89

biflora, 89

ssp. carlottae, 89

crassa, 89

glabella, 89

kamtschadalorum, 90

langsdorfFii, 90

palustris, 90
Vulpia

major, 29

myuros, 29
var. major, 29

ZOSTER ACEAE, 19
Zostera, 20
marina, 20

The type in this book is Times New Roman with italic, text in 10 pt. and 8 pt., tables in 6,
7 and 8 pt., and titles in 12 pt. and 18 pt. Captions were set in 7 and 8 pt. 20th Century
medium with italic. The volume was lithographed on Imperial Offset Enamel paper made
by Rolland Paper Company Limited, and was bound in Winterbottom Art Buckram. The
complete manufacture of the book was done by The Bryant Press Limited, Toronto, for
the Queen's Printer, Ottawa.

ROY L. TAYLOR, the senior author of this
volume, was born in Olds, Alberta, in 1932.
He attended school and university in his
native province and taught school there
for four years before moving to Montreal to
specialize in botany at Sir George Williams
University and later at McGill. He has
a doctorate from the University of
California at Berkeley and has also held the
position of Associate in Botany at that
university. Since 1962, Dr. Taylor has been
on the staff of the Plant Research Institute
in Ottawa, where he is now Chief of the
Taxonomy Section. Besides making frequent
field trips over large areas of the
Cordilleran region and publishing the
results of his surveys, Dr. Taylor is active
in promoting a closer association of biolo-
gists in Canada through his work in the
Canadian Botanical Association and the
Biological Council of Canada. He has
recently been appointed to the Editorial
Committee for the Flora North America
Project, initiated in 1967.

GERALD A. MULLIGAN has spent most
of his life in Ottawa. He was born here in
1928, spent his childhood here, and upon
graduation from MacDonald College of
McGill University with B.Sc. in biology in
1952, he returned to join the Weed
Investigation Section of the Canada
Department of Agriculture. He joined the
Plant Research Institute when it was formed
in 1961 and immediately initiated cyto-
taxonomic studies on the Cruciferae of
Canada and adjacent areas. In recent years
he has traveled widely in Canada and the
United States to conduct botanical surveys,
and in 1964 he participated in the Genetics
of Colonizing Species Symposium at
Asilomar, California. His many scientific
papers on the cytotaxonomy and chromo-
some numbers of weeds have made him an
international authority on that group
of plants. His present research is concen-
trated on cvtotaxonomic studies of the
family Cruciferae.