Full text of "Ferns and fern allies of Canada"

See other formats

1*1

Agriculture
Canada

Ferns

and fern allies of Canada

Digitized by the Internet Archive

in 2011 with funding from

Agriculture and Agri-Food Canada - Agriculture et Agroalimentaire Canada

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

Fcrns

and fern allies of Canada

William J Cody

Biosystematics Research Centre, Ottawa, Ontario

Donald M. Britton

University of Guelph, Guelph, Ontario

Research Branch
Agriculture Canada

Publication 1829/E
1989

^Minister of Supply and Services Canada 1989

Available in Canada through

Authorized Bookstore Agents
and other bookstores

or by mail from

Canadian Government Publishing Centre
Supply and Services Canada
Ottawa, Canada Kl A 0S9

Cat. No. A53-1829/1989E
ISBN 0-660-13102-1

Price is subject to change without notice

Canadian Cataloguing in Publication Data

Cody, William J., 1922-

Ferns and fern allies of Canada

(Publication ; 1829)

Issued also in French under title: Les fougeres et

les plantes alliees du Canada.

Includes index.

Bibliography: p.

Cat. No. A53-1829/1989E

ISBN 0-660-13102-1

1. Ferns-Canada. 2. Ferns-Canada-Nomenclature.
I. Britton, Donald M. II. Canada. Agriculture
Canada. Research Branch. III. Title. IV. Series:
Publication (Canada. Agriculture Canada). English;
1829.

QK525.C6 1989 587'.3r0971 C89-099204-5

Staff Editor

Frances Smith

CONTENTS

Acknowledgements iv

Introduction 1

Addendum to the 13
Introduction, 1989

Key to the genera 15

Families and genera 19

1. LYCOPODIACEAE 19
1. Lycopodium 19

2. SELAGINELLACEAE 49
1. Selaginella 49

3. ISOETACEAE 61
1. Isoetes 61

4. EQUISETACEAE 73
1. Equisetum 73

5. OPHIOGLOSSACEAE 95

1. Ophioglossum 95

2. Botrychium 97

6. OSMUNDACEAE 124
1. Osmunda 124

7. SCHIZAEACEAE 130
1. Schizaea 130

8. HYMENOPHYLLACEAE 132
1. Mecodium 132

9. PTERIDACEAE 133

1. Dennstaedtia 133

2. Pteridium 135

3. Cheilanthes 138

4. Aspidotis 140

5. Pellaea 143

6. Crypto gramma 150

7. Pityrogramma 155

8. Adiantum 157

10. ASPIDIACEAE 165

1. Matteuccia 166

2. Onoclea 168

3. Woodsia 170

4. Polystichum 182

5. Dryopteris 205

6. Gymnocarpium 231

7. Thelypteris 239

8. Phegopteris 247

9. Cystopteris 253
10. Athyrium 263

11. BLECHNACEAE 273

1. Blechnum 273

2. Woodwardia 275

12. ASPLENIACEAE 281

1. Asplenium 281

2. Camptosorus 290

3. Phyllitis 292

13. POLYPODIACEAE 295

1. Poly podium 295

14. MARSILEACEAE 307

1. Marsilea 307

15. SALVINIACEAE 310

1. Azo/Za 310

Distribution maps 313
Glossary 393
References 401
Index 423

ACKNOWLEDGMKNTS

The helpful comments of S.G. Aiken and P.M. Catling, who
reviewed earlier versions of the manuscript, are gratefully
acknowledged. We are particularly appreciative of the comments of
T.M.C. Taylor, who took time from his retirement to read the
manuscript. Technical support in the preparation of the distribution
maps was provided by L.D. Black and W.A. Wojtas. The fine line
drawings were done by V. Fulford; her valuable contribution is
gratefully acknowledged. We are also indebted to the curators of the
various herbaria who kindly lent specimens for our study.

INTRODUCTION

The first treatment of Canadian ferns was undertaken by
Macoun and Burgess, who published Canadian Filicineae in 1884. As
these authors state, "Probably no form of growth throughout the
vegetable kingdom attracts more general attention than ferns, which,
while appealing strongly to the scientific tastes, have an equally
powerful claim upon the artistic. Their distribution over the whole
surface of the globe, with the exception of the sterile portions of the
polar regions, places at least some forms within the reach of everyone,
while, grow in what locality they may, there is none to which they do
not lend an added charm ...."

In 1889 George Lawson published The School Fern-Flora of
Canada as an appendix to Asa Gray's Botany for Young People and
Common Schools: How Plants Grow, A Simple Introduction to
Structural Botany, with a Popular Flora. Robert Campbell in 1898
and 1899 produced a series of fascicles on Canadian ferns in successive
issues of Canadian Horticultural Magazine. John Macoun (1890) also
listed the ferns and fern allies in his Catalogue of Canadian Plants.

Since these early works, the only treatments of all the ferns
known to occur in Canada have been brief accounts by Boivin (1968) in
Enumeration des plantes du Canada and Scoggan (1978) in The Flora
of Canada. Local treatments, some including the fern allies, have
appeared for Nova Scotia (Roland 1941); Quebec (Marie- Victorin 1923,
1925); Ottawa District (Cody 1978, 1980); and British Columbia
(T.M.C. Taylor 1963, 1970); these treatments have been most useful
for those regions. Other regional treatments have appeared in various
floras such as Calder and Taylor (1968) Flora of the Queen Charlotte
Islands, Part 1; Erskine (1961) The Plants of Prince Edward Island;
Fernald (1950) Gray's Manual of Botany, eighth edition; Gleason and
Cronquist (1963) Manual of the Vascular Plants of Northeastern
United States and Adjacent Canada; Hitchcock et al. (1969) Vascular
Plants of the Pacific Northwest; Moss (1959) Flora of Alberta; Scoggan
(1957) Flora of Manitoba; and various local lists, such as Soper (1963)
for Manitoulin Island. The present volume is thus the first in over 90
years to combine descriptive information on the various taxa with
comments on their relationships, habitats, and distributions. It is
hoped that the book will prove to be a useful tool, not only to
individuals taking a first look at these interesting plants but also to
the dedicated amateur and the professional botanist.

We have found that writing this book has been a singular
challenge. We are very aware that books such as this are written by
professionals who claim they are writing for amateurs! Our objectives
have been to bring together for the first time all the ferns and fern
allies (pteridophytes) of Canada; to make readily available references
to the literature for those who wish more information on certain taxa;
and to supply keys, descriptions, illustrations, and distributions for all

the species. We have tried to give the reader an appreciation of or
insight into the various approaches of recognizing and naming species.
In addition, we have tried to challenge the reader to find gaps in
distribution, to extend ranges, and to find rare hybrids. To this end we
have included the following:

• Information for correct identification, including easily used field
characters.

• Comments on whether a species is easily recognized and has a
discrete, limited range or whether much is still to be discovered
about the species.

• Comments concerning the aesthetic aspects, popular appeal, and
folklore of the plants, although space restrictions have been a
limiting factor in this regard.

• References to all the major essential literature.

• An assessment of the taxonomy that appears to be clear and
straightforward at this time and that which does not, keeping in
mind that we are aware that a great deal of work that is still
unpublished is currently being done on many of the species in this
book. Taxonomy is not static, and one should therefore expect
further change and interpretations of the species that are different
from those we have presented.

• An assessment of current trends in the taxonomy of the
pteridophytes, i.e., an extrapolation from the past to the present
and an indication of the direction in which we are moving.

Latin name

The Latin name, e.g., Onoclea sensibilis L., and a few relevant
synonyms are used, followed by an accepted common name or names
(e.g., sensitive fern). Although synonymy is of great interest to the
professional, it usually does not excite the amateur; we have therefore
attempted to keep the synonymy as short as possible. Most amateurs
are bewildered, or even annoyed, at what appears to be constant name
changing, e.g., Dryopteris carthusiana for D. spinulosa or Lycopodium
digitatum for L. flabelli forme. To increase the amateur's under-
standing and appreciation of some of the problems, we have included a
brief history of some names, as follows:

• Onoclea sensibilis L. 1753. The genus name Onoclea (initial capital
letter) is followed by the species name sensibilis (all in lower case).
The abbreviation for the author, L., stands for Linnaeus, the father
of binomial names (genus plus species), who described the plant and
named it in 1753. This name has remained constant, which is an
indication of the distinctiveness of the plant.

• Gymnocarpium dryopteris (L.) Newman. Linnaeus described the
common oak fern in 1753 as Polypodium dryopteris, which is
accordingly recognized as the basionym. There has been great
disagreement through the years as to which genus this species
belongs. It has been placed in Thelypteris, Phegopteris, Dryopteris,

Currania, and Carpogymnia, among others. It is now recognized as
a Gymnocarpium and is called G. dryopteris (L.) Newman, following
work published by Newman in 1851. Because Linnaeus is
recognized as the person who originally described the species, his
initial appears in parentheses before Newman's name (not all
authors' names are abbreviated).

Aspidotis densa (Brack.) Lellinger. This plant has been placed in
Cryptogramma, Cheilanthes, and Pellaea by various authorities. Its
affinities with those genera have been questioned and it was placed
in Aspidotis by Lellinger in 1968.

Dryopteris expansa (C. Presl) Fraser-Jenkins & Jermy. Extensive
studies on European populations of Dryopteris dilatata showed that
there were two species within this taxon, a diploid and a derived
allotetraploid. The ancestral diploid species was segregated as D.
assimilis S. Walker. It was later found that an earlier name,
Nephrodium expansum C. Presl, 1825, based on a specimen
collected by Haenke from Nootka Sound, existed and had priority.
Accordingly, D. assimilis became invalid and the current name D.
expansa was created in 1977.

Gymnocarpium jessoense (Koidz.) Koidz. ssp. paruulum Sarvela.
After studying all the gymnocarpiums of the world, Sarvela decided
that material from Alaska, the Yukon, and the Prairie Provinces,
which had for the last century been known as Gymnocarpium
robertianum (Hoffm.) Newman, was related to the Asian G.
jessoense. He selected a type collected in the Nahanni and described
our plant as a new subspecies. At this time, one can find treatments
that reduce G. robertianum to a mere variety of G. dryopteris, e.g.,
G. dryopteris (L.) Newman var. pumilum (DC.) Boivin. In addition,
many authors recognize both G. dryopteris and G. robertianum as
separate species. In our treatment, based on Sarvela, we recognize
a third species, G. jessoense, as well as G. dryopteris and G.
robertianum.

Subspecies and varieties

Canada's royal fern, Osmunda regalis, is closely related to the
European plant. Gray in 1856 described the new world material as
varietally distinct, e.g., O. regalis (L.) var. spectabilis (Willd.) Gray.
The story is similar for Equisetum hyemale L. The Canadian plant is
known as ssp. affine (Engelm.) Stone. The situation is less clear for
the ostrich fern, Matteuccia. Some researchers believe that Canada's
species is distinct and should be called M. pensyluanica, whereas
others treat our plant as a variety of the European M. struthiopteris
(L.) Todaro.

Some modern treatments would suggest that the above examples
describe problems of subspecies, not varieties. A subspecies has no
barrier to interbreeding with the species, but it has a distinct
morphology and a quite separate distribution, e.g., Adiantum

pedatum L. ssp. aleuticum (Rupr.) Calder & Taylor. An amateur
might ask why old varieties have not been logically and uniformly
changed to subspecies. Primarily, the reason for this has been to avoid
adding complexity to taxonomic literature. Also, the intent of the
original author is not always apparent, i.e., we do not always know
whether it can be safely assumed that a "variety" was in reality a
subspecies. In the current use of categories, variety is a lower rank
than subspecies and applies to a group of similar individuals within a
subspecies, e.g., Adiantum pedatum L. var. subpumilum W.H.
Wagner.

Taxonomists differ widely in their interpretation of variation in
populations. For example, Boivin (1968) recognized Dryopteris
austriaca without subspecies or varieties. In our opinion, this taxon
includes in Canada's flora D. expansa, D. Intermedia, D. campyloptera,
and D. carthusiana. After extensive studies on Botrychium, W.H.
Wagner recognizes B. oneidense and B. minganense as species, while
dismissing B. obliquum as a mere form of B. dissectum and the var.
europaeum of B. uirginianum as not meriting taxonomic rank.
Further experimental work should help to clarify some of the widely
divergent views that are currently held.

Common names

These vary widely in various parts of the world. Some are well
entrenched and difficult to change, even if they are not particularly
suitable. For example, ebony spleenwort is not ebony-colored, and
Wherry has called it the brown-stemmed spleenwort. The silvery
spleenwort is not a spleenwort (Asplenium) but an Athyrium. We
have not attempted to introduce major changes in long-term usage.

Description

The lengths of the frond are given; the divisions (if any) of the
blade are described; any scales, hairs, or glands are mentioned; and
the important fruiting bodies and indusia (if present) are described.
The descriptions are reasonably full, but at the same time are not of
the length found in such works as the Flora of the Pacific Northwest.
At the end of the technical description, certain useful field characters
are highlighted.

Cytology

The chromosome number and author of the report are given. An
attempt is made to stress reports based on Canadian material. Such
Canadian reports are indicated by an asterisk. We have attempted to
cite recent literature in the hope that later reports will cite previous

records. For those interested in this information, checking the
uniformity of opinion, e.g., Love et al. (1977), is extremely useful.
Most chromosome numbers for ferns have been determined from
meiotic studies, and so we have usually given the n or gametic
number. Where 2n numbers are given, we consider that they were
determined from somatic material, usually root tips. The designation
"m" = 2n indicates that there was premeiotic doubling of chromosome
number (see Manton 1950), and the species is apogamous.

Habitat

Habitat varies widely for even a single species. We have
attempted to give what we consider to be typical habitats. Because
some ferns are restricted to basic rocks and others to acidic rocks, we
have attempted to indicate these preferences.

Range

Range and occurrence in Canada are indicated primarily by dot
maps, but in some cases are highlighted by special comments when
the range is unusual. The complete distribution is indicated for North
America, and if the taxon is found on other continents, that fact is
briefly noted. The Canadian distribution is given from east to west,
followed by the United States distribution from east to west. For those
interested in visualizing distributions on a shaded map, Mickel (1979)
should be consulted.

Remarks

In the Remarks section we attempt to indicate whether the
species is clear-cut or not and whether there are problems in its
interpretation. Any special features of interest are mentioned.

Segregate species

The most casual observer of the pteridophyte flora of Canada
must be impressed and perhaps bewildered by the seeming
proliferation of a large number of "new" species, e.g., Dryopteris
expansa, Lycopodium digitatum, Gymnocarpium jessoense, and
Botrychium rugulosum, and might ask how this has occurred and
what the final outcome will be.

Segregate species, to a large extent, are the natural outcome of
applying the concept of biological species after surveying the variation
of populations. For example, if one compares the current treatment of
Dryopteris with that in Fernald (1950), Dryopteris spinulosa is now

composed of D. intermedia, D. expansa, D. campyloptera, and D.
carthusiana. One species is now four. The rationale behind this
splitting is clear. Dryopteris intermedia is a sexual diploid {n = 41)
with certain limits of variability that can be described, e.g., lacy,
subevergreen fronds with glandular indusia, and so on. In terms of
evolution, it is considered to be ancestral to the derived tetraploids D.
campyloptera and D. carthusiana. Similarly, D. expansa is another
sexual ancestral diploid that superficially is a segregate species in
Canada's flora, but if the plants in Europe and Japan are considered,
D. expansa is actually a composite ancestral diploid species because it
is extremely wide ranging (circumpolar). We can now include Japan,
British Columbia, eastern Canada, and northern Europe in the range
of this species. The tetraploids D. carthusiana (which has long been
known as D. spinulosa) and D. campyloptera have quite different
origins and distributions and are separate interbreeding populations,
and therefore are also species. The same situation has been repeated
in many other genera, e.g., Polystichum, Cystopteris, Polypodium, and
Isoetes, among others. It is an inevitable trend, resulting from modern
research techniques, which is only rarely balanced by the merging of
two species that were once considered separate, e.g., Asplenium
ruta-muraria and A. cryptolepis, Woodsia alpina and W. bellii. Critics
of segregate species have pointed out that more emphasis is placed on
differences between species than on similarities. This difference in
philosophy has led to the recognition of Pellaea glabella var. simplex.
This variety is certainly close in both morphology and evolution to P.
glabella var. glabella, and it is therefore more appropriately called
var. simplex rather than P. suksdorfiana. This change in name
recognizes the morphological differences between var. simplex and P.
glabella var. glabella. Another example is Matteuccia struthiopteris of
Europe versus M. pensylvanica of North America. If differences are
stressed, they may be considered separate species (although critical
experimental breeding data are absent), whereas if similarities are
stressed they belong to the same species with two varieties or
subspecies.

For many of these plants we do not have the critical evidence
from experimental crosses or from naturally occurring hybrids to
make an objective decision. In Botrychium, for example, the fall
Botrychium species (such as B. dissectum and B. multifidum) all have
n — 45. To date, artificial crosses have failed because of difficulties in
growing spores, and even culture of mature plants is difficult.
Consequently, we are left with a number of species that are known to
grow together. Some researchers have argued that the variation
exhibited is an example of the great variability of each species and
have recognized only Botrychium dissectum and B. multifidum as
species, whereas others, e.g., W.H. Wagner and Wherry, have stressed
that if the plants are growing in close proximity and hence identical
environments, then the variation seen is genetic, and they recognize
as many as five species. Similar problems arise with various species of
Lycopodium, e.g., L. complanatum, L. digitatum, and L. tristachyum.

One could argue in favor of only one collective species for all these — all
have the same chromosome number and all apparently can intercross
without leading to meiotic irregularities; or conversely one can
emphasize the large differences in morphology between extremes and
recognize three species.

It would seem to us that the final outcome will inevitably be the
recognition of additional species and the further partitioning of
variation. We will be able to say that the system is a more "natural"
classification, or one that emphasizes evolutionary units. Basic
diploids and derived tetraploids will be more clearly delineated.

The methods being used, which include chemical analysis,
comparative morphology, and scanning electron microscopy (SEM) of
spores, will continue to emphasize differences rather than similarities,
and so inevitably, the larger collective species, such as Athyrium
filix-femina and Cystopteris fragilis, will yield more segregates after
further study.

Cytology and biosystematics

Manton (1950) brought the methods and philosophy of
cytogenetic analysis, which she had used successfully on the
Cruciferae, to a study of pteridophytes. By means of artificial
hybridization techniques, analysis of naturally occurring hybrids, and
analysis of chromosome numbers, she showed that the evolution and
phylogeny of the pteridophytes could be greatly clarified. It is
impossible to summarize briefly all her results here, but it is
important to stress the impact of her work, and of those who followed
her, on our classification of the ferns and fern allies. She found, for
instance, that all the species of true Dryopteris have a basic
chromosome number of 41, which is an unusual number not easily
manipulated arithmetically because it is not divisible by an integer.
Within Dryopteris, Manton showed that there were derived
tetraploids, e.g., D. carthusiana and D. filix-mas, and that
furthermore, these species initially arose from interspecific
hybridization, followed by a doubling of the chromosome number, thus
restoring fertility, i.e., by allopolyploidy. This may be shown
schematically, as in Diagram 1.

In morphology, D. carthusiana is considered to be a blend of the
characteristics of its two diploid ancestors, and in breeding behavior,
it is a new, derived allotetraploid species. To take this scheme one
step further, we think that D. cristata is LLBB (2m = 164) and that it
crossed at one time with D. goldiana 2n = 82, which is GG. The
resulting sterile hybrid would be (L)(B)(G), which upon doubling of
chromosome number would give rise to the fertile allohexaploid as
shown in Diagram 2.

C\J
00

C\J

c
o

(/)

O
<D
Cl

>^
Q)

E
o

M—

CO
CD

E
o

o
o

-t— '
CD
CO

<D
■♦— '

_Q)
Q.

E
o
o

E
o

CD
CD

E
o

CO
CO

O
(D

D
O

o
o

0) CO

)r. CD

c
o

c
o
o

CC
Q.

c E

^ o

CD CO

O o

E E

CO O

CNJ
CO

CO
C\J 00

Sen

-—- c

-r^ CO

^ CO

X CO

"O +
CD

-^ CO

^ '^

O Q.

■D ^

.f^ "^

CO SI

(D .t^

E ^

O ctj

CO c:

O CTJ

E "55

*- -C:

o ^

Q--D

CO -

CO ._ Q_

II £ --5

CO '^ _

O CO

CO ^
.^ CO

CO CD
CD O
Q. C
CO CO

CD O

. CO
CO CO

Ih ^

o ±:

'<^ CD

E ^

CO CD

CD jQ

C= ■:
0 ll

^ c:

4- c\j

E
o

■g
'o

-♦— '

_o
lo

CO CO

Q. CO

c
o
o

O
CD
Cl
CO

•2

•c

a
o

C/3

.2
a

'S
"a

s:
■*^

O
C

The allohexaploid is what we call Dryopteris clintoniana, and a
close scrutiny of the plant indicates the influence of both D. cristata
and D. goldiana in its origin. ' The message is clear — it is not only the
chromosome number, but also the character of the sets of
chromosomes (genomes) that determine the final make-up of the
species. Patterns such as those above are now well known in
Asplenium, Cystopteris, Polypodium, and Polystichum, for example. It
would appear to be a common pattern of evolution for many species.

A different sort of pattern is seen in Pellaea. In P. glabella var.
nana the basic chromosome number is 29 and the taxon is sexual, but
in P. glabella var. glabella the somatic chromosome number is 116 and
the species is apogamous, i.e., the spores also have 116 chromosomes.
The long beech fern is similar in this regard; although the basic
chromosome number in Phegopteris is 30, P. connectilis has a somatic
chromosome number of 90 and the viable spores also have 90
chromosomes.

Not all speciation has involved polyploidy. Many of the
Botrychium species have n = 45 and yet some have evolved quite
different morphologies, e.g., B. lunaria versus B. multifidum. At
times, the degree of allopolyploidy is in doubt, or analysis may show
that we are dealing with an autopolyploid. For example, the diploid
hart's tongue (Phyllitis) in Europe versus tetraploid hart's tongue in
Canada are not markedly different in morphology. Similarly, diploid
and tetraploid Asplenium trichomanes are superficially very similar
in appearance. For amateurs to make such distinctions usually
demands more effort in identification than they may wish to expend.
They may be forced to measure spore sizes and stomatal sizes or even
to determine chromosome number.

Chromosome numbers have also been used as an aid to
determine systems of classification at higher levels, such as family.
More recent systems of classification have tended to group genera
with a basic chromosome number of 29 or 30 into an adiantoid group,
e.g., Adiantum, Cheilanthes, Cryptogram ma, and Pellaea (Lovis 1977).
These chromosome numbers are quite unlike those for Dryopteris (41),
Polystichum (41), Cystopteris (42), and Athyrium (40), which in turn
are markedly different from Polypodium, which has the unusual basic
number (X) of 37.

Cytology has been less useful for phylogeny with some groups.
Equisetum, for instance, appears to be completely uniform with n =
108. A different problem arises in Lycopodium. Here there are a
number of different basic numbers, which suggests that the genus is
unnatural or polyphyletic, and some of the species are extremely
difficult to analyze, e.g., L. lucidulum and L. selago.

Modern monographic treatment of genera now includes a whole
battery of experimental techniques (Britton 1974) such as cytology,
chromatography, electron microscopy, comparative anatomy, and
computer analysis. There is even some promise from DNA
hybridization studies and the analysis of isoenzymes. It is no wonder.

*(0

'<D

*-•

C\J

1 1

C\J

CO
CM

— ►■^

y—

► CO

y—

—1

^ CO

J=
J=

«M

o
s
.2

^>
'u

with so many methods of analysis, that the taxonomy of the
pteridophytes is slowly changing as the species themselves have
evolved through the years. The reader who is interested in these
aspects of evolution should consult the review articles in Jermy et al.
(1973), Lovis (1977), and Walker (1979).

Hybrids

Interspecific (and intergeneric) hybrids are well known in some
genera. Much work has been done on Asplenium hybrids (Lovis 1977;
W.H. Wagner 1954); Dryopteris (W.H. Wagner 1970); Equisetum
(Hauke 1963); Lycopodium (Wilce 1965); Polypodium (Shivas 1961);
Polystichum (D.H. Wagner 1979); and Phegopter is (Mulligan and Cody
1979). Knobloch (1976) has published a list of pteridophyte hybrids.

In some other genera, hybrids are either extremely rare or have
never been detected, e.g., Blechnum, Botrychium, Cryptogramma,
Osmunda, Pellaea, and Thelypteris.

Interspecific fern hybrids are usually characterized by
intermediate morphology, although so-called "one-way hybrids" are
not unknown. Even these, e.g., Dryopteris goldiana X D. intermedia
(Evans and Wagner 1964), may superficially resemble one parent (D.
intermedia), but may prove on analysis to have a blend of
characteristics from both parents. In ferns, one expects a high
incidence of sterility in these hybrids. For example, D. goldiana (GG)
X marginalis (MM) results in a hybrid with (G)(M). Because there is
no homology between the chromosomes at meiosis, one obtains 41 +
41 or 82 single chromosomes at meiosis, which separate very unevenly
and the resulting spores abort. On this basis, each resulting hybrid is
an end point that is not involved in further crosses. However, it is
apparent that some hybrids also produce quite numerous large,
spherical spores, a few of which are capable of germinating
(DeBenedictis 1969). It is thought that these spores contain all the
chromosomes of the parent hybrid plant and are able to germinate and
produce genotypes identical to the original hybrid. This would appear
to account for the abundance of a few hybrid combinations, e.g.,
Dryopteris X triploidea, D. X boottii, D. filix-mas X marginalis,
Gymnocarpium X intermedium, and G. dryopteris ssp. X
hrittonianum.

Finding and identifying hybrid pteridophytes is a challenge.
First, it is necessary to become thoroughly familiar with the parent
species in order to assess the full range of variability exhibited by
them. Then one should be prepared to examine carefully the spores for
abortion and, in some cases, to undertake cytology. It is no longer
acceptable to collect a fern with bizarre morphology and blithely pass
it off as some rare and exotic hybrid combination. A British "hybrid"
said to be Polypodium vulgare X Pteridium aquilinum reported in
1907 is not taken seriously today. There should be little difficulty in
finding such hybrids as Dryopteris X triploidea and D. X boottii

because they seem to be present wherever the parents are abundant
and are growing intermixed. On the other hand, the Osmunda hybrid
O. X ruggii Tryon would be a rare find indeed, even though the
prospective parents often grow in close proximity over literally
thousands of square kilometres.

Throughout the publication, hybrids placed within parentheses
are not known to occur in Canada.

Sequence of families, genera, and species

In the present work, the families and genera follow the
taxonomic sequence of Genera Filicum (Copeland 1947). This follows
the modern trend of separating the families Pteridaceae, Aspidiaceae,
Blechnaceae and Asplenianeae, which were at one time lumped under
the Polypodiaceae. Within genera, the species are placed in such an
order that similar species can be compared readily.

Distribution maps

It was not possible for the authors to visit all the herbaria in
Canada as well as the major ones in the United States and elsewhere.
Our objective was to depict the broad picture of distribution of the
species in Canada. At times we have included locations in Alaska and
Greenland when this has helped to show the distribution more clearly.
Thus, provincial or regional maps with more dots might give a slightly
different picture for a smaller area, e.g., the maps for rare plants of the
various provinces (Maher et al. 1979; White and Johnson 1980;
Douglas et al. 1981). Our maps were prepared initially from
specimens in the herbarium of the Biosystematics Research Centre,
Ottawa, Ont. (DAO) (Holmgren et al. 1981). These were
supplemented by records from 26 Ontario herbaria including National
Museum of Natural Sciences, Ottawa, Ont. (CAN); University of
Toronto, Toronto, Ont. (TRT); University of Guelph, Guelph, Ont.
(OAC); Queen's University, Kingston, Ont. (QK); and University of
Western Ontario, London, Ont. (UWO). Additional Quebec and
British Columbia records were obtained from the published works of
Rousseau (1974), Taylor (1970), Erskine (1961), and Roland and Smith
(1969), who relied heavily on herbaria of Laval University, Quebec,
Que. (QFA); University of Montreal, Montreal, Que. (MT); University
of British Columbia, Vancouver, B.C. (UBC); British Columbia
Provincial Museum, Victoria, B.C. (V); Acadia University, Wolfville,
N.S. (ACAD); and Nova Scotia Agricultural College, Truro, N.S.
(NSAC). Selected specimens have been borrowed from various
herbaria, e.g., University of Manitoba, Winnipeg, Man. (WIN), and
selected literature citations have been included as well, where they
are significant.

ADDENDUM TO THE INTRODUCTION, 1989

In our introduction, we stressed the current trend toward
segregate species. This trend is strongly apparent in recent studies on
Botrychium by W.H. Wagner of the University of Michigan. W.H.
Wagner and F.S. Wagner (1983a) have outlined their methods for
deciding whether the variation observed in populations is genetic or
environmental, and they give their rationale for their description of a
large number of new species of Botrychium in North America. Five
new species have been described recently (W.H. Wagner and F.S.
Wagner 1981, 19836), at least three others are not yet published, and
still others may be recognized after further study (W.H. Wagner and
F.S. Wagner 1983c). These authors have suggested that what we have
called B. boreale ssp. obtusilobum should be called B. pinnatum St.
John, and that the name B. hesperium (Maxon & Clausen) Wagner &
Lellinger should be applied to some populations in Cypress Hills and
in Waterton Lakes National Park.

A specimen of B. paradoxum W.H. Wagner (W.H. Wagner and
F.S. Wagner 1981) was cited from Waterton Lakes National Park. Dr.
Wagner reported finding additional specimens in the park in 1982.
Elsewhere it has been reported from Montana and British Columbia.
This plant is extraordinary in that it lacks a sterile blade. Its
relationship to other taxa in the genus is uncertain and needs further
study.

The key to the four groups of Botrychium subgenus Botrychium
given by W.H. Wagner and F.S. Wagner (1983a) is perhaps useful,
because it bridges the gap between the more classical treatment of
Clausen (1938) and the modern, segregation treatment followed by
W.H. Wagner and F.S. Wagner. It should be noted that the emphasis
is on the species category, at the expense of recognizing subspecies or
varieties of any kind. The four groups keyed out encompass six
widespread species: B. simplex, B. lunaria, B. minganense, B. boreale,
B. lanceolatum, and B. matricariae folium. F.S. Wagner (1983) states
that in western North America there are 13 species and five
interspecific hybrids of subgenus Botrychium. Of these 13 species,
nine are considered to be endemic in the West, two are in the B.
lunaria group, two in the B. simplex group, four in the B. lanceolatum
group, and B. paradoxum is in its own group. W.H. Wagner and F.S.
Wagner (1983a) give a key for five species in the B. lanceolatum group.
Of these five species, only B. echo is considered by them to be absent
from Canada. It is only known in Colorado, Utah, and Arizona.

W.H. Wagner and F.S. Wagner (1981) state that some specimens
annotated by W.H. Wagner to B. dusenii are now referable to B.
crenulatum W.H. Wagner.

However, until all the new entities are described by them, and all
the Canadian material has been restudied and the results of their field
studies are published, we are unable to state clearly how many species

are present in Canada. As an example, W.H. Wagner and F.S.
Wagner (1983c) suggest that there are as many as eight species of
Botrychium in the Cypress Hills of Saskatchewan, and 10 species of
the genus in Waterton Lakes National Park. In both cases, these are
much larger numbers of species than have been recognized for these
areas in the past.

It is interesting to note that R.C. Moran (1983) has produced
evidence that the plant we have called Cystopteris frag'ilis var.
mackayii should be treated as a species, C tenuis (Michx.) Desv., and
indeed this was suggested earlier by Lellinger (1981). It is
unfortunate that Moran did not see more Canadian material to give a
better impression of the northern distribution of this taxon on his
map.

Again, V.L. Harms (1983) has reported on the occurrence of
Athyrium filix-femina in Saskatchewan. He is of the opinion that both
var. sitchense (cyclosorum) and var. michauxii occur in that province.
Athyrium filix-femina is an extremely variable and difficult species to
understand. It was studied by Liew Fah Seong at the New York
Botanical Garden in the early 1970s, but to our knowledge the study
has not been published.

It is of interest, too, to note that the National Museum of Natural
Sciences, National Museums of Canada, has published two more
volumes in the series on rare plants (Bouchard et al. 1983; Hinds
1983). For Quebec, two fern allies and 26 ferns were considered to be
rare. They were not mapped. For New Brunswick, four fern allies and
nine ferns were considered to be rare. Distribution maps for the New
Brunswick taxa were included in that volume. Also, Argus and White
(1983) have mapped 14 species of ferns and fern allies for Ontario. It is
worth noting, however, that a large number of species are now
withdrawn from the original list of rare vascular plants of Ontario.

KEY TO THE GENERA

Stems jointed; nodes covered by sheaths composed of basally
united scarious leaves, otherwise leafless; sporangia borne on

inner surface of peltate scales of terminal spike-like cones

Equisetum p. 73

Stems not conspicuously jointed, bearing green leaves or leaf-like
fronds.

B. Leaves (fronds) small, entire or serrate, very numerous and

imbricated, or quill-like and crowning a short corm-like

stem; sporangia sessile or subsessile in leaf axils.

C. Stems elongate, covered with persistent small more or

less flattened leaves; plants creeping from rhizomes or

decumbent stems.

D. Leaves without a ligule; strobiles terete,

homosporous Lycopodium p. 19

D. Leaves ligulate; strobiles 4-sided; sporangia of
two kinds, microsporangia containing many
minute microspores (male) and macrosporangia
containing fewer and larger macrospores

(female) Selaginella p. 49

C. Stems short, thick, and corm-like, crowned by a rosette

of quill-like leaves; spores of two kinds

Isoetes p. 61

B. Leaves (fronds) usually pinnate or deeply lobed, not closely
or only slightly imbricated; sporangia naked, or in sori on
the backs or margins of sometimes specially adapted fronds
or their divisions, or in sporocarps.

E. Fronds tiny, bilobed, 2-ranked, floating free on the
surface of quiet water; sporangia in sporocarps borne

on the underside of the axis Azolla p. 310

E. Fronds larger, simple or divided, not bilobed, growing
from a persistent rhizome.

F. Plants aquatic, although sometimes becoming
stranded; fronds long-petioled, 4-foliate, floating
on the surface; sporangia in hard sporocarps

borne on or close to the rhizome

Marsilea p. 307

F. Plants typically terrestrial; fronds not 4-foliate;
sporangia not in sporocarps.

G. Fronds or portions of them conspicuously
dimorphic.

H. Sterile fronds linear-filiform, blade-
less, crowded on a short crown; fertile
fronds flliform, tipped by a few tiny
crowded flnger-like pinnae bearing the
sporangia Schizaea p. 130

1 1 . Sterile fronds with a distinct blade, not
crowded; fertile fronds not filiform and
tipped by a few tiny pinnae, but much
larger.

I. Sterile part of frond simple; fer-
tile part of frond a long-stalked
simple spike with two rows of

coherent sporangia

Ophioglossum p. 95

I. Sterile part of frond pinnately
divided one or more times.
J. Sporangia naked.

K. Fronds fleshy, single,
or sometimes two
growing from a
scarcely developed
rhizome and consisting
of a sterile lower part
and, when present, a
more or less upright
fertile panicle or spike;
sporangia 2-ranked . .
.... Botrychium p. 97
K. Fronds forming a more
or less dense crown at
the end of a stout
rhizome; sporangia not

2-ranked

Osmundap. 124

J. Sporangia partly or wholly
covered by the rolled-up
pinnules, forming globular
berry-like divisions of the
stiff fertile frond.
L. Fronds in vase-like
clumps; simple

pinnate fertile fronds
surrounded by tall
regularly pinnate
sterile ones; sterile
fronds oblong-

lanceolate; rachis not

winged

.... Matteuccia p. 166
L. Fronds solitary or
scattered along the
rhizome; sterile fronds
deltoid, coarsely pin-
natifid; rachis winged

apically; fertile fronds

bipinnate

Onocleap. 168

G. Fertile fronds or fertile portions of fronds
similar to the sterile; fertile fronds
sometimes longer but not hardened and
berry-like (fertile and infertile fronds of
Cryptogramma, Blechnum, and Aspidotis
are dimorphic, but of similar texture).
M. Fronds simple, commonly auricled at
the base.
N. Fronds long-caudate, sometimes

rooting at the tip

Camptosorus p. 290

N. Fronds oblong, not attenuate or

rooting at the tip

Phyllitis p. 292

M. Fronds variously divided.

O. Fronds small and delicate; blades

one cell thick

Mecodium p. 132

O. Fronds larger and coarser; blades
more than one cell thick.
P. Fronds covered beneath by a
conspicuous white to golden

yellow powder

Pityrogramma p. 155

P. Fronds not as above

Group I

A. Sori marginal; indusium formed entirely or in part by the
re volute margin of the frond.
B. Sori distinct, short, mostly not confluent.

C. Stipe and fronds glabrous Adiantum p. 157

C. Stipe and fronds glandular-hairy

Dennstaedtia p. 133

B. Sori usually confluent as a marginal band.

D. Fronds coarse, scattered along stout elongate and
forking rhizomes Pteridium p. 135

D. Fronds finer, tufted from a very short rhizome.

E. Segments of frond bead-like

Cheilanthes p. 138

E. Segments of frond not bead-like.

F. Pinnules and segments of frond jointed at
the base Pellaea p. 143

F. Pinnules and segments of frond not jointed
at the base.
G. Stipes herbaceous, green except at the

base Cryptogramma p 150

G. Stipes wiry, dark, and shiny

Aspidotis p. 140

A. Sori dorsal on the frond or, if marginal, the indusium not formed
by the revolute margin.
H. Sori elongate.

I. Indusia continuous, attached near the margins of the

pinnae Blechnum p. 273

I. Indusia not continuous.

J. Sori in chain-like rows, parallel to the midrib . . .

Woodwardia p. 275

J. Sori parallel to the oblique lateral veins.

K. Fronds to 1 m long, herbaceous; veins

reaching the margin .... Athyrium p. 263

K. Fronds smaller, to 40 cm long, evergreen or

herbaceous; veins not reaching the margin

Asplenium p. 281

H. Sori round or nearly so.
L. Indusia present.

M. Indusia segmented Woodsia p. 170

M. Indusia not segmented.

N. Indusium hood-shaped, attached by its base

on the side toward the midrib

Cystopteris p. 253

N. Indusium round or reniform.

O. Fronds scattered along a thin cord-like
rhizome (or tufted from a stout

rhizome in T. limbosperma)

Thelypteris p. 239

O. Fronds tufted at the end of a stout
rhizome
P. Indusium reniform or with a deep

sinus Dryopteris p. 205

P. Indusium round, without a deep

sinus Polystichum p. 182

L. Indusia absent.

Q. Fronds coriaceous, evergreen, simply pinnatifid

Polypodium p. 295

Q. Fronds deciduous, at least pinnate-pinnatifid.

R. Rhizome stout; fronds forming a crown ....

Athyrium p. 263

R. Rhizome cord-like.

S. Fronds more or less ternate

Gymnocarpium p. 231

S. Fronds pinnate-pinnatifid

Phegopteris p. 247

FAMILIES AND GENERA

1. LYCOPODIACEAE club- moss family

1. LycopodiumL. club-moss

Plants low, evergreen, coarsely moss-like, with simple to much
branched stems covered with simple, 1-nerved, 4- to many-ranked,
lanceolate or linear leaves. Sporangia in the axils of leaf-like
sporophylls similar to the vegetative leaves or segregated in a
terminal strobilus or cone. Spores numerous, yellow.

The genus Lycopodium has over 400 species throughout the
world and occurs on all the continents except Antarctica. There are a
large number of tropical species, many of which are epiphytes,
although we usually think of Canadian species as being especially
adapted to cool, moist, and northern habitats of often bleak and barren
places.

In the past 40 years the genus has been examined in some detail
(Beitel 19796), and several workers have been impressed by the
diversity of sporophyte morphology, gametophyte morphology (Bruce
1976), spore morphology (Wilce 1972), and chromosome numbers
(Love et al. 1977). This diversity has prompted European workers to
adopt four generic names {Lycopodium, Diphasiastrum, Lycopodiella,
and Huperzia) for their species (Jermy et al. 1978). Some researchers
have gone so far as to place Huperzia in a separate family,
Huperziaceae (Love et al. 1977), whereas others have attempted
groupings at a subgeneric or section level, noting that it is a great
advantage to have a genus name that is familiar to most workers and
typifies plants that can be recognized at a glance (Wilce 1972).

We have retained the name Lycopodium here for the 13 species
that we recognize as occurring in Canada, but at the same time we
have grouped the species into categories corresponding to the genera
recognized by others (Holub 1964, 1975).

A. Sporangia in the axils of leaf-like sporophylls.

B. Leaves sharply erose-serrulate at the apex, flat at the base;

leaves in alternating belts, some long, others short

12. /.. lucidulum

B. Leaves entire or nearly so, acuminate, plump, and hollow at
the base; all leaves essentially the same length

13. L. selago

A. Sporangia in the axils of modified terminal leafy-bracted
strobiles.

C. Sterile branches horizontal or arching; strobili with green
leaf-like bracts 11./.. inundatum

C. Sterile branches erect or ascending; strobili with firm
yellowish scale-like bracts.

D. Strobili sessile at the ends of leafy stems.
E. Aerial stems erect and tree-like.

F. Leaves of lower part of stem strongly
divergent 3. /.. dendroideum

F. Leaves of lower part of stem strongly

appressed to slightly divergent

4. /.. obscurum

E. Aerial stems tufted, bushy, or fan-like;
branchlets more or less flattened.

G. Plants to 13-25 cm; leaves 8-ranked,

6-10 mm, not fused to stem

2. L. annotinum

G. Plants short, less than 13 cm; leaves 4- to
5-ranked, scale-like, partly fused to stem.
H. Leaves mostly 4-ranked; leaves of the
upper and lower sides unlike the

marginal 8. L. alpinum

H. Leaves mostly 5-ranked; all the leaves

alike 9. /.. sitchense

D. Strobili peduncled.

I. Leaves linear-subulate with long soft hair-like

tips 1. L. clavatum

I. Leaves scale-like.

J. Sterile branchlets not compressed or
slightly compressed; leaves uniform and

usually in 4 rows 10. L. sabinifolium

J. Sterile branchlets flattened.

K. Constrictions between seasons'
growths conspicuous.
L. Upright growth habit straggly;

branchlets flat

5. /.- complanatum

L. Upright branches fastigiate;

branchlets roundish

7. 1. tristachyum

K. Constrictions between seasons'
growths usually only slightly pro-
nounced; branchlets of the branches

arching and fan-like

6. 1. digitatum

Lycopodium s.s. group

The leaves in this group are not fused to the stem along their
length, and the sporangia are borne in distinct cones or strobili. A
typical plant is L. clavatum, the common club-moss, staghorn
club-moss, or wolfs claw. This species has a rhizome on or near the
ground surface and unequally forked upright branches, suggesting

antlers of a stag or the paw of a wolf. Other species in this group are L.
annotinum (stiff club-moss), L. obscurum (ground-pine), and L.
dendroideum (round branched ground-pine). All have chromosome
numbers based on n = 34 (Love et al. 1977). The spore morphology is
shown accurately, and beautifully, by scanning electron microscopy,
and all the spores of the species in this group have a regular reticulum
of polygons, i.e., they are in a honeycomb pattern (Wilce 1972). It is,
however, a spore pattern similar to that seen in the L. complanatum
(Diphasiastrum) group.

1. Lycopodium clavatum L. var. clavatum

L. clavatum L. var. integerrimum Spring.

common club-moss
Fig. 1, habit. Mapl.

Stems elongated, horizontal on the surface of the ground,
forking, rooting at intervals; leaves uniform, but lower leaves turned
upward. Erect branches at first simple, becoming dichotomous; fertile
branches with a leafy-bracted peduncle bearing 2 to several sessile or
short-stalked strobiles. Leaves linear-subulate, incurved-spreading,
usually tipped with a soft white hair-like bristle. Bracts of strobili
yellow, fimbriate-erose, at least the lower with white filiform tips.

Cytology: n = 34 (Love and Love 1976).

Habitat: Dry woods and clearings.

Range: Circumpolar; in North America from Newfoundland to
British Columbia and Alaska, south to North Carolina, Michigan,
Minnesota, Idaho, and Washington.

Remarks: Mature fruiting plants present no problems in
identification. Young or sterile plants, however, may be confused with
L. annotinum. The extended soft, hair-like bristles on the leaves are
useful for discrimination.

1.1 Lycopodium clavatum L. var. monostachyon Hook. & Grev.

L. clavatum L. var. megastachyon Fern. 8l Bissell

L. clavatum L. var. brevispicatum Peck
Fig. 2 (a) habit; (6) portion of strobilus. Map 2.

Similar to var. clavatum, but with the leaves ordinarily
ascending or appressed and the cone single on a shorter peduncle.

Cytology: n = 34 (Love and Love 1966a).

Fig. 1 Lycopodium clavatum var. davatum; habit, 3/4 x

{^^<n

Fig. 2 Lycopodium clavatum var. monostachyon; (a) habit, Ix; (b) portion of
strobilus, 8x .

Habitat: Exposed situations, hilltops, alpine and subalpine regions
generally north of var. clauatum in North America (north of northern
Minnesota, Michigan, and northern New England).

Range: Greenland, Labrador, and Newfoundland to Alaska.

Remarks: This taxon, with 1 strobilus per peduncle and more
appressed leaves than for typical L. clauatum, has been considered a
separate species, L. lagopus (Laest.) Zinserl. ex Kuzen, as listed in
Czerepanov (1981). Others have variously treated the taxon as a
subspecies, variety, or form.

2. Lycopodium annotinum L.

bristly club-moss, stiff club-moss
Fig. 3 (a) habit; (6) portion of strobilus. Map 3.

Stems elongated, prostrate, mostly unbranched, rooting at
intervals; leaves uniform but the lower leaves turned upward. Erect
stems simple to forked several times, increasing annually to 20 cm or
more in height. Leaves 8-ranked, more or less stiff and hard,
linear-subulate to linear-oblanceolate, with a sharp spinule. Strobili
sessile at the ends of leafy stems.

Several intergrading varieties have been recognized: var.
annotinum, with leaves of erect stems 6-11 mm long, linear-lanceolate
or oblanceolate, coarsely toothed, spreading; var. acrifolium Fern.,
with leaves of erect stems 5.5-7.0 mm long, linear-subulate, spreading
or ascending; var. alpestre Hartm., with leaves of erect stems 2.5-6.0
mm long, linear-lanceolate to lance-attenuate, thick and hard,
dorsally convex, entire, and strongly ascending to appressed; and var.
pungens (La Pylaie) Desv., with leaves of erect stems 2.5-6.0 mm long,
lanceolate to lance-oblong, flat, obscurely serrate and strongly
ascending to tightly appressed.

Cytology: n = 34 (Love and Love 1966a) also for var. pungens.

Habitat: Moist woods and clearings, subalpine forests, and exposed
rocky and peaty habitats.

Range: Circumpolar; in North America from Greenland and
Labrador to Alaska, south to Virginia, Minnesota, and Oregon.

Remarks: The tightly appressed and strongly ascending entire leaves
of var. pungens seem distinctive from the typical variety, but
intergradations occur. There is the usual disparity of views as to
whether the varieties mentioned should be species, subspecies, or
forms. For example, Czerepanov (1981) includes var. alpestre and var.

Fig. 3 Lycopodium annotinum; (a) habit, 1 x ; (b) portion of strobilus, 6 x

pungens in the species L. dubium Zoega, whereas Love includes var.
pungens and the species L. dubium in ssp. alpestre Love and Love
(LoveetaL 1977).

Lycopodium annotinum is at times confused with L. lucidulum,
but the latter has no strobili and branches are equal in length.

3. Lycopodium dendroideum Michx.

L. obscurum L. var. dendroideum (Michx.) D.C. Eat.
round-branched ground-pine, tree club-moss
Fig. 4 (a) habit; (6) position of strobilus. Map 4.

Subterranean stems creeping, branching, and rhizome-like, with
broad scale-like leaves. Aerial stems upright, 10-30 cm high, simple
below, forking above, constricted between the seasons' growth. Lower
leaves strongly divergent; leaves of lateral branchlets in 2 dorsal, 2
ventral, and 2 lateral ranks; leaves strongly decurrent, the free part
linear-attenuate. Strobili sessile and terminal on the main axis, or
dominant branches and produced in the second, third, or fourth
growing season.

Lycopodium dendroideum may be quickly identified by grasping
the base of an aerial stem. This will feel distinctly prickly because of
the stiff divergent leaves.

Cytology: n - 34 (Love and Love 1976*).

Habitat: Woods and clearings.

Range: Labrador and Newfoundland to British Columbia and Alaska,
south to West Virginia, Michigan, Wisconsin, and Washington; Asia.

Remarks: Hickey (1977) considered this wide-ranging taxon to be a
good species and segregated it from L. obscurum. The latter used to be
considered as consisting of all flat-branched forms (Wherry 1961).
Now, however, L. obscurum var. obscurum is the flat-branched
variant, and var. isophyllum (equal-leaved ground-pine), with all 6
ranks of leaves of equal size, is a variant of the flat-branched species.

4. Lycopodium obscurum L. var. obscurum
ground-pine, tree club-moss

Fig. 5 (a) habit; (6) portion of branch. Map 5.

* Throughout the publication, an asterisk indicates that the count is
based on Canadian material.

Fig. 4 Lycopodium dendroideum; (a) habit, 2/3 x ; (b) portion of strobilus, 9 x .

Fig. 5 Lycopodium obscurum var. obscurum; (a) habit, 1/2 x ; (b) portion of branch,
5x.

Similar to L. dendroideum, from which it may be distinguished
by the strongly appressed to slightly divergent leaves on the lower
portion of the aerial shoot. Leaves of the lateral branchlets arranged
in 1 dorsal, 1 ventral, and 4 lateral ranks; leaves of ventral rank
linear-attenuate to long triangular, smaller than leaves of other
ranks; leaves of other ranks linear-acuminate to linear-acute.

Cytology: ai = 34 (Love and Love 1976*).

Habitat: Woodlands.

Range: Nova Scotia to Ontario, Michigan, and Wisconsin, south to
North Carolina, Tennessee, and Kentucky.

Remarks: Variety obscurum is flat-branched, having a single rank of
leaves that run along the upper surface of a branchlet and reduced
leaves along the ventral rank. This is a more southern and eastern
species than the widespread L. dendroideum.

4.1. Lycopodium obscurum L. var. isophyllum Hickey

equal-leaved ground-pine
Fig. 6, portion of branch. Map 6.

Similar to var. obscurum in that leaves of lower portion of stem
are strongly appressed to slightly divergent; leaves of branchlets are
all of equal size and linear-attenuate; all leaves lie in planes
tangential to the branchlet axis.

Habitat: Woodlands.

Range: Nova Scotia to Ontario and Minnesota, south to Tennessee
and Kentucky.

Remarks: Because this variety has been recognized only recently
(Hickey 1977), our knowledge of its distribution is still limited. Field
workers can make a contribution when they become familiar with the
three taxa, L. dendroideum, L. obscurum, and L. obscurum var.
Isophyllum.

Diphasiastrum group

This group of species was treated in a monograph by Wilce
(1965). Many of the species have flattened branches with 4 ranks of
scale-like leaves (1 dorsal, 2 lateral, and 1 ventral), which suggest the
name ground-cedar, but unfortunately this name has been used for
more than one species in the group. The species to be considered here

are the larger plants L. complanatum, L. digitatum, and L.
tristachyum, and the smaller and more mat-like L. alpinum, L.
sitchense, and L. sahinifolium. All have a chromosome number of n =
23 (Wilce 1965, but see Love et al. 1977). The spores of all the species
are reticulated {L. clavatum type, Wilce 1972) and the species
apparently can hybridize with one another. Presumably, these
hybrids produce fertile spores, because no meiotic irregularities occur
during meiosis. However, this extrapolation has not been tested
experimentally because of very low to nonexistent germination of
spores of even nonhybrid taxa. Interspecific hybridization in this
group is quite unlike that in Equisetum or the ferns, where hybrids
usually show lack of chromosomal homology between species.

5. Lycopodium complanatum L.

Diphasium complanatum (L.) Rothm.
Diphasiastrum complanatum (L.) Holub
flatbranch club-moss
Fig. 7 (a) habit; (6) portion of stobilus. Map 7.

Horizontal stems mostly below the surface of the ground; leaves
distant, scale-like. Upright stems to 30 cm high or higher, with
crowded or somewhat remotely forking branchlets. Branchlets
flattened, often strongly constricted between yearly growths, 2.0-4.0
mm wide. Leaves 4-ranked; lateral leaves usually appressed; leaves of
lower rank much reduced. Strobili mostly 1 or 2 on remotely bracted
peduncles.

Lycopodium complanatum is a familiar species in our boreal
woods. Field characters are surficial to buried rhizomes, wide
branches, and conspicuous annual constrictions. The irregular growth
pattern often gives the plant a rather irregular or straggly look, in
contrast to the extreme regularity of L. digitatum. The strobili are
also irregular in number (1-4) per peduncle, and the naked peduncles
seem very fine in relation to the size of the strobili.

Cytology: n = 23 (Hersey and Britton 1981*).

Habitat: Woodlands and clearings.

Range: Circumpolar; in North America from Greenland, Labrador,
and Newfoundland to Alaska, south to New England, Michigan,
Montana, Idaho, and Washington.

Remarks: There are several varieties to consider, and their
disposition depends on whether one views the variation as belonging
to the parental taxon or whether one considers the varieties to be, in
reality, hybrids. Wilce (1965) equated var. gartonis Boivin with var.
elongatum Vict., and they are now interpreted as L. complanatum X
tristachyum. Stunted northern forms called var. pseudoalpinum

Fig. 6 Lycopodium obscurum var. isophyllum; portion of branch, 5 x

Fig. 7 Lycopodium complanatum; (a) habit, 2/3 x ; (b) portion of strobilus, 10 x

F'arwell, var. montellii Kukkonen, and var. canadense Vict, are
considered by us to be northern ecotypes.

6. Lycopodium digitatum A. Braun

L. flabelliforme (P'ern.) Blanch.

L. complanatum L. var. flabelliforme Fern.

L. complanatum L. var. dillenianum Doll

Diphasiastrum digitatum (A. Braun) Holub

Diphasium flabelliforme (Fern.) Rothm.

crowfoot club-moss, running-pine
Fig. 8, habit. Map 8.

Stems horizontal, mostly on or near the surface of the ground;
leaves distant, scale-like. Upright stems to 30 cm high or higher, with
the branchlets of the branches arched and fan-like; constrictions
between annual growth not present or only slightly evident.
Branchlets 2.0-3.0 mm wide. Leaves 4-ranked; lateral leaves usually
spreading; lower leaves much reduced. Strobili mostly 3 or 4 on
remotely bracted peduncles; peduncle forking at one point.

Cytology: n = 23 (Hersey and Britton 1981*).

Habitat: Dry woods and clearings.

Range: Newfoundland to Ontario and Minnesota, south to New
England, Kentucky, and Iowa.

Remarks: Linnaeus described L. complanatum in 1753, and by 1814
L. tristachyum was recognized. Many authors now believe that var.
flabelliforme should be treated as a separate species, L. digitatum.
Typical material is quite distinctive. The stems are on or very near
the surface, the branchlets are very regular and fan-like, annual
constrictions are lacking, and the strobili are usually in groups of four
on long, naked peduncles.

The species is characteristic of sandy woods and clearings in
southeastern Canada, and is endemic in North America.

7. Lycopodium tristachyum Pursh

Diphasiastrum tristachyum (Pursh) Holub

ground-cedar
Fig. 9 (a) habit; (b) portion of strobilus. Map 9.

Horizontal stems usually deeply buried; leaves distant, scale-
like. Upright stems to 30 cm high or higher. Sterile branches ascend-
ing to loosely divergent, flattened, 1.0-1.5 cm wide. Leaves 4-ranked,
bluish green, lanceolate-subulate; lateral leaves appressed; lower
leaves somewhat smaller. Strobili 2-6 on leafy-bracted peduncles.

Fig. 8 Lycopodium digitatum; habit, 1 x

Fig. 9 Lycopodium tristachyum; (a) habit, 1/3 x ; (b) portion of strobilus, 6 x .

The vase-shaped and crowded branches, which are bluish green
and have whitish wax on their underside, give a striking appearance
to the sun forms. Shade forms are more diffusely branched, but the
branchlets are still more rounded than those of L. complanatum and L.
digitatum. Good field characters to observe are the annual
constrictions along the branches, and ventral and lateral leaves of the
same size and shape. The peduncles often branch and then branch
again (2-forked) to give rise to 4 strobili.

Cytology: n = 23 (Hersey and Britton 1981*).

Habitat: Dry, sometimes sandy woods and clearings.

Range: Newfoundland to Ontario, Michigan, and Minnesota, south
to West Virginia and Alabama.

Remarks: Variety laurentianum Vict, is considered by Wilce (1965)
to be L. X habereri House.

8. Lycopodium alpinum L.

Diphasiastrum alpinum (L.) Holub
alpine club-moss
Fig. 10 (a) habit; (6) leaves; (c) portion of strobilus. Map 10.

Stems elongate, horizontal, rooting at intervals, and bearing few
leaves; leaves distant, yellow, bract-like. Erect stems dichotomously
forked to 9 cm high. Sterile branchlets somewhat flattened. Leaves
4-ranked, dimorphic; dorsal leaves lanceolate-subulate, appressed,
adnate for about half their length; lateral leaves deltoid-ovate to
lanceolate, 4-5 mm long, adnate for about half their length, with the
free part spreading and incurved at the tip; ventral leaf shorter and
trowel-shaped. Strobili essentially sessile at the ends of branched
leafy peduncle-like stems.

This northern and alpine species is often confused with L.
sitchense and L. sabinifolium. The 4-ranked leaves, which are free
from the stem for about half their length, are distinct from the
5-ranked rounded branches of L. sitchense. The ventral trowel-shaped
leaves are unlike those of L. sabinifolium, which has pedunculate
strobili.

Cytology: n = 23-24 probably 23 (Love et al. 1977).

Habitat: Alpine and subalpine meadows and wooded alpine slopes.

Range: Circumpolar; in North America from Greenland to Alaska,
south to Newfoundland, eastern Quebec, Michigan, Washington, and
Montana.

Fig. 10 Lycopodium alpinum; (a) habit, 1 x ; (b) leaves, 12 x ; (c) portion of strobilus,
12x.

9. Lycopodium sitchense Rupr.

L. sabinifolium Willd. var. sitchense (Rupr.) Fern.
Diphasiastrum sitchense (Rupr.) Holub
Diphasium sitchense (Rupr.) Love & Love
Sitka club-moss
Fig. 11 (a) habit; (b) portion of strobilus. Map 11.

Stems elongate, horizontal, rooting at intervals, bearing distant,
yellowish, scale-like leaves. Erect stems dichotomously forked to
18 cm high. Sterile branchlets cylindrical. Leaves in 4 or, more often,
5 ranks, uniform, subulate, adnate for less than half their length, with
the free parts usually incurved at the tips. Strobili sessile on leafy
branches, not on naked peduncles.

Cytology: n = 23 (Love and Love 1976*).

Habitat: Alpine and subalpine barrens and wooded slopes.

Range: Greenland, Labrador, and Newfoundland to British
Columbia and Alaska, south to Maine, New Hampshire, Montana,
Washington, and Oregon.

10. Lycopodium sabinifolium Willd.
Diphasiastrum sabinifolium (Willd.) Holub
savin leaf club-moss

Fig. 12 (a) habit; (6) portion of strobilus. Map 12.

Stems elongate, horizontal, rooting at intervals, and bearing a
few distant yellowish bract-like leaves. Erect stems dichotomously
forked, up to 20 cm high. Sterile branchlets flattened. Leaves
4-ranked, linear-subulate, scarcely dimorphic; dorsal and ventral
leaves appressed and slightly adnate; lateral leaves slightly larger
and adnate for about half their length, with the free parts spreading
and incurved at the tip. Strobili on leafy-bracted peduncles 1-8 cm
long.

Cytology: n = 23 (Love and Love 1976*).

Habitat: Subalpine often dry and sandy woods and meadows.

Range: Labrador and Newfoundland to Algoma District, Ont., south
to Pennsylvania and Michigan.

Remarks: Beitel (1979a, 19796) stresses the indistinct strobilus base
and the "scattered sporophylls and sporangia straggling down naked
peduncles." The species is considered by Wilce (1965) and Beitel

Fig. 1 1 Lycopodium sitchense; (a) habit, 1 x ; (b) portion of strobilus, 7 x

r.)^ b

Fig. 12 Lycopodium sabinifolium; (a) habit, 1 x ; (b) strobilus, 7 x .

(1979/)) to be a hybrid between L. sitchense and L. tristachyum.
Variants (perhaps segregants) resemble the parental species in
appearance. The branchlets are flattened, with leaves in 4 ranks
(compare with L. sitchense, which is 5-ranked and has round
branchlets). In disturbed sites in Ontario, e.g., jack pine blowouts, one
can usually find both L. sahinifolium and L. sitchense in the same
location.

Lycopodiella group

The sporophyte is deciduous, except for the extreme tip of the
rhizome. The plants are small and creeping, and the most striking
attribute is the erect, fertile branches. The spores, known as the
rugulate type (Wilce 1972) and having shallow, rolling ridges, are
quite unlike the previous groups. The species have a chromosome
number based on n = 78 (Love et al. 1977). We have recognized only
one species in this complex for Canada, although we know from Beitel
(19796) that the genus has been studied by Bruce (1975), who
recognized additional species as well as a host of hybrids. Beitel
(19796) writes (quoting Bruce 1975) of two new tetraploid species
(which are still undescribed) in the Great Lakes region, so that we can
expect revisions to this group for Canada.

11. Lycopodium inundatum L. var. inundatum

Lepidotis inundata (L.) C. Borner

Lycopodiella inundata (L.) Holub

bog club-moss
Fig. 13 (a) habit; (b) portion of strobilus. Map 13.

Stems horizontal or arching, forking, rooting at intervals; leaves
linear-subulate, gradually long-acuminate, not adnate, spiralled in 8
or 10 ranks. Leaves on the underside of the stem twisted upwards.
Fertile stems upright, ascending, or slightly incurved, with leaves
similar to those of the sterile stems. Strobili single, sessile; strobilus
(6-10 mm wide) definitely wider than stem.

Cytology: m = 78 (Love and Love 1976*).

Habitat: Acid bogs, shores, damp sandy banks, and disturbed
situations.

Range: Labrador and Newfoundland to Ontario, south to Virginia,
north-central United States; northern Saskatchewan; Alaskan
Panhandle, south through British Columbia to Oregon and Idaho;
Eurasia.

Fig, 13 Lycopodium inundatum var. inundatum; (a) habit, 2/3 x ; (6) portion of
strobilus, 10 x .

11.1 Lycopodium inundatum L. var. bigelovii Tuckerm.
Fig. 14, habit. Map 14.

Differs from var. inundatum by its taller (up to 35 cm) fertile
stems and by its mostly ciliate-denticulate leaves. Peduncle leaves
and sporophylls tightly appressed. Strobilus narrow, 3-4(5) mm wide.

Cytology: n = 78 (W.H. Wagner et al. 1970) for L. appressum.

Habitat: Wet shores, bogs, and savannas.

Range: Newfoundland and Nova Scotia to Florida and Texas.

Remarks: According to Beitel (19796), following Bruce (1975), this
taxon belongs in a separate species, L. appressum (Chapman) Lloyd &
Underwood, the southern bog club-moss. Lycopodium appressum has
a coastal and lowland distribution in the Gulf and Atlantic states, and
Canadian plants would, in their view, be northern outliers. Beitel
(19796) includes Newfoundland in the distribution, and we are told
there are two undescribed tetraploid species in this complex, plus
several hybrids. The Canadian plants are in need of further study.

Huperzia group

The final group of lycopods to consider includes L. lucidulum and
L. selago, which have leaves in many ranks and lack the specialized
strobili that are characteristic of L. clavatum. The spores have small
pits (foveolate) and are quite different in size and appearance from
those of the other groups. The spores are triangular, with concave
sides and truncate angles (Wilce 1972). Both L. lucidulum and L.
selago s.l. are considered collective entities by some researchers
(Beitel 19796), and undoubtedly further segregate species will appear.
The cytology of this group is difficult because of many meiotic
irregularities (hybrid taxa?), small chromosomes, and large numbers
of chromosomes. "Determinations" of chromosome number are often
only estimates (Manton 1950).

12. Lycopodium lucidulum Michx.

Huperzia selago (L.) Bernh. ssp. lucidula (Michx.) Love & Love
shining club-moss
Fig. 15 (a) habit (6) portion of branch with sporangia. Map 15.

Stems ascending and sprawling, few-forked, to 40 cm long, leafy,
rooting towards the base from among the brown marcescent leaves.
Leaves mostly 6-ranked, 7-12 mm long, oblanceolate, spreading or
deflexed, acuminate, sharply erose-serrulate near the apex.

Fig. 14 Lycopodium inundatum var. bigelovii; habit, 1 x

u^-

Fig. 15 Lycopodium lucidulum; (a) habit, Ix; (6) portion of branch with
sporangia, 10 x.

alternating in bands; shorter leaves appearing early in the season
followed later by longer leaves. Stomates only on the lower surface.
Sporangia in the axils of the shorter leaves. Gemmae or reproductive
buds often borne in the upper leaf axils.

Lycopodium lucidulum may be distinguished from L. selago ssp.
patens by its erose-serrulate rather than entire leaves and by the
presence of stomates on the lower leaf surface only, rather than on
both surfaces.

Cytology: n = 61 (Beitel and Wagner 1982*).

Habitat: Cool moist woods.

Range: Newfoundland to Ontario, Minnesota, and Iowa, south to
South Carolina and Indiana.

Remarks: This is a characteristic species of rich boreal and hardwood
forests in eastern Canada. The undulating outline to the branches
and the toothed leaves of a dark and shiny green are good field
characters. The species seems distinct, until one considers L.
lucidulum var. occidentale or L. selago var. patens. The latter is
considered under L. selago, and the former is a sporadic form
(entire-leaved) in Canadian populations of L. lucidulum. Variety or
forma occidentale as it occurs in Canada should not be confused with
L. porophilum Lloyd & Underwood, even if listed in the synonymy of
that species. Lycopodium porophilum is not yet known from Canada
(compare with Wherry 1961), but is most frequent in central United
States (Wisconsin, Iowa, Ohio, and Pennsylvania), growing on acidic
sandstone cliffs and ledges. The color of the plant is markedly yellow
green towards the base, it has some undulations in shoot outline, and
the leaves are entire, linear-lanceolate, with parallel sides. Stomata
occur on both surfaces of the leaves. Lycopodium porophilum is
reported to hybridize frequently with L. lucidulum, and the hybrids
can rapidly reproduce by gemmae.

13. Lycopodium selago L. ssp. selago

L. selago L. var. appressum Desv.
Huperzia selago (L.) Bernh.
mountain club-moss
Fig. 16 (a) habit; (b) portion of strobilus. Map 16 (s.l.).

Horizontal stems short, leafy, rooting from among the
marcescent leaves; erect stems to 20 cm high or higher, branched
several times, usually near the base. Leaves yellow-green, crowded,
8- to 10-ranked, 3-8 mm in length, ovate-lanceolate, entire or nearly
so, acuminate, usually hollow at the base, and with stomates on both
surfaces. Sporangia in the axils of leaves produced early in the season

followed later by sterile leaves, thus appearing in bands. Gemmae or
reproductive buds often borne in the upper leaf axils.

Cytology: n = 132 (Love and Love 1966a).

Habitat: Arctic tundra species, south in the mountains, on barrens,
in bogs, and in cold woods.

Range: Circumpolar; in North America from Greenland to Alaska,
south to Virginia, Michigan, Wisconsin, Montana, and Washington.

Remarks: According to Beitel (19796), Lycopodium selago s.l. is a
complex of species, hybrids, and environmental forms. Because all the
Canadian material has not been compared and is inadequately known
at this time, we have fallen back on an older and simpler
interpretation of these plants, recognizing L. selago ssp. selago, ssp.
patens, and ssp. miyoshianum. Beitel (19796) would refer ssp. patens
to a "catch-all" taxon, i.e., a complex of a species, hybrids, and
environmental forms. Some specimens are perhaps hybrids of ssp.
selago and L. lucidulum. Undoubtedly, one can expect to see quite
different treatments of this species in the future. For the present, we
are following Calder and Taylor (1968) until new research is published
and has been carefully evaluated.

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

Differs from ssp. selago in its longer leaves, 8-12 mm,
lance-attenuate, reflexed or strongly divergent, and by its stems to
30 cm long or longer.

Habitat: Cold woods and rocky situations.

Range: Newfoundland to Manitoba and Wisconsin, south to New
England; British Columbia and Alaska.

13.2 Lycopodium selago L. ssp. miyoshianum (Makino)

Calder & Taylor
Fig. 17 (a) habit; (6) portion of strobilus.

Differs from ssp. selago by its leaves dark green, thin and
flexuous, narrowly lanceolate, about 6 mm long, strongly imbricated
but not appressed, and by its usually longer stems.

Habitat: Mountain slopes.

Range: British Columbia through Alaska to Japan, Korea, and
China.

Fig. 16 Lycopodium selago ssp. selago; (a) habit, 1/2 x; (b) portion of strobilus,
lOx.

Fig. 17 Lycopodium selago ssp. miyoshianum; (a) habit, ix; (b) portion of
strobilus, 3x.

Hybrids of Lycopodium

The occurrence of hybrids in Lycopodium is not subject to direct
experimental testing because of our inability to germinate and grow
spores in large numbers. A further difficulty is the varying concepts of
species within the genus. Some researchers would deny that hybrids
exist, whereas others believe that hybrids are quite frequent. Until
there is greater unanimity concerning the characteristics of the basic
species in our flora, the characteristics of the hybrids between these
species (or even their existence) will be uncertain. Accordingly, we
have dismissed from consideration hybrids in all the groups except
those in section Complanata or genus Diphasiastrum Holub. As
mentioned previously, they are unusual hybrids for pteridophytes, in
that few if any meiotic irregularities or lack of chromosomal homology
are observed (Hersey and Britton 1981), and it is assumed that the
spores are viable. The most frequently cited hybrids involve the
triangle of L. complanatum, L. digitatum, and L. tristachyum. The
most convincing ones are those whose parents differ rather widely in
morphology.

Wilce (1965) cited nine Canadian specimens of Lycopodium
complanatum X digitatum from Quebec and Ontario, but considered it
rare. The parents, however, do not appear very different, so that
hybrids are difficult to distinguish.

Wilce (1965) cited 24 specimens of Lycopodium complanatum X
tristachyum (Diphasiastrum X zeilleri (Rouy) Holub) for Canada, from
Northwest Territories, Saskatchewan, Manitoba, Ontario, Quebec,
Prince Edward Island, Newfoundland, and Labrador. It is supposedly
more common in Minnesota than is L. tristachyum. A number of our
collections have come from the Thunder Bay District, Ont.

The extremes of the species are distinctive, so that the hybrid L.
digitatum X tristachyum (Lycopodium X habereri House) seems
reasonably clear-cut. Wilce (1965) reported 16 collections for Canada
from Ontario, Quebec, and New Brunswick. The cytology was studied
by Hersey and Britton (1981).

Lycopodium alpinum X complanatum (L. X issleri (Rouy)
Lawalree) is known from a few localities in Europe, as well as from
Maine in North America (Wilce 1965). It should be looked for where
the ranges of the parents overlap.

A specimen of Lycopodium alpinum X sitchense from the Mealy
Mountains in southern Labrador has been seen. Wilce (1965) cited
two specimens for North America, one from Oregon and the other from
Washington.

According to Wilce (1965) the species L. sabinifolium arose from
a cross of L. sitchense with L. tristachyum. L. sabinifolium is
extremely variable, and variants approach the parents in morphology.
Accordingly, it is not possible to detect hybrids of L. sabinifolium with
either L. sitchense or L. tristachyum.

2. SELAGINELLACEAE spikemoss family

1. Se/ag/ne//a Beau V. spikemoss

Low, creeping plants with branching stems and few fine roots.
Leaves simple, imbricated, in 4 or 6 rows, with or without a bristle tip.
Some sporangia containing macrospores, others microspores, borne in
the axils of the leaf-like sporophylls of a terminal cone.

The genus Selaginella, in which several hundred species are
included, is the only genus in the family Selaginellaceae. These
species are widely distributed and are mostly tropical. About 37
species occur in North America north of the Mexican border. A few
species are cultivated as greenhouse plants, and one, S. lepidophylla,
the resurrection plant, is imported as a novelty. Members of the S.
rupestris section that grow in semiarid and subalpine regions are
adapted to being almost completely desiccated and revive a few hours
after moisture becomes available.

A. Leaves flat, not bristle-tipped.

B. Leaves dimorphic; sporophylls and leaves eciliate

2. S. apoda

B. Leaves uniform; sporophylls and leaves ciliate

1 . S. selaginoides

A. Leaves grooved on the back, bristle-tipped.

C. Leaves abruptly adnate to the stem and differing in color
from it 3. S. wallacei

C. Leaves decurrent on the sides of the stem.

D. Bristles of leaves about 1 mm long 5. S. densa

D. Bristles shorter.

E. Epiphytic; stems lax and freely branching; cones

hardly differentiated 4. S. oregana

E. Terrestrial; densely matted; cones distinct

F. Leaves tapering to the setae

6. S. rupestris

F. Leaves truncate at the apex, then setate . . .
7. S. sibirica

1. Selaginella selaginoides (L.) Link

Fig. 18 (a) habit; (6) portion of strobilus. Map 17.

Plants delicate, branching, forming small mats. Leaves uniform,
2-4 mm long, spreading-ascending, acute, ciliate. Fertile branches
upright, with lower leaves similar to those of the stem but becoming
larger upwards to form the sporophylls of a subcylindric spike.

The uniform, ciliate leaves and upright fertile branches readily
separate S. selaginoides from the only other herbaceous-leaved species
in our area, S. apoda.

Cytology: 2n = 18 (Love and Love 1976*).

Habitat: Moist banks and shores, bogs, and boggy woods.

Range: Circumpolar; in North America from Greenland and
Labrador to Alaska, south to New England, Michigan, Minnesota, and
southern British Columbia.

Remarks: This widespread circumpolar species, which occurs
northward nearly to the limit of trees, is often partly buried in mosses
or muskeg, and thus is easily overlooked. It is rare in Manitoba
(White and Johnson 1980) and Saskatchewan (Maher et al. 1979).

2. Selaginella apoda (L.) Fern.

S. eclipes Buck
Fig. 19 (a) habit; (6) portion of the branch bearing sporangia. Map 18.

Plants delicate, freely branching, matted. Leaves membranous,
4-ranked, the lateral 2 rows bluntish, oblong to oval, and spreading;
dorsal and ventral leaves pointed, smaller, and appressed. Spikes
sessile; sporophylls eciliate, similar to the foliage leaves.

The pale or whitish green eciliate heterophyllous leaves readily
set this species off from all others in our area.

Cytology: 2n = 18 (Love and Love 1976*).

Habitat: Wet woods, swamps, bogs, and shores.

Range: Southwestern Quebec and Ontario to Wisconsin, south to
Florida and Texas.

Remarks: Buck (1977) described a new species, S. eclipes, with a
range north of, but essentially adjacent to, the range he ascribed to S.
apoda. The characters used to differentiate S. eclipes from S. apoda
largely overlapped those of the latter, and the author therefore
suggested that it might prove to be better placed at a subspecific level.
Because this publication is not an appropriate vehicle for making such
a transfer, we have included S. eclipes in the synonymy of S. apoda. A
map depicting the distribution of S. eclipes and S. apoda in the
restricted sense is given by Buck (1977). Alston (1955) reported S.
apoda as occurring west to British Columbia, but in Canada the
species is found only in southwestern Quebec and Ontario.

Fig. 18 Selaginella selaginoides; (a) habit, 1 x ; (6) portion of strobilus, 8 x

Fig, 19 Selaginella apoda: (a) habit, 1 x ; (b) portion of branch bearing sporangia,
2x.

3. Selaginella wallacei Wieron.

S. montanensis Hieron.
Fig. 20 (a) habit; (6) leaves; (c) portion of strobilis. Map 19.

Main stems prostrate, sparsely rooted, forming loose mats.
Ascending branches numerous. Leaves abruptly adnate to the stem,
tightly appressed, more or less glaucous, oblong-linear, more or less
obtuse at the apex, grooved on the back, ciliate, about 3 mm long,
including the approximately 0.5 mm long scabrous seta. Sporophylls
ovate-deltoid, shorter than the leaves, ciliate. Setae nearly smooth.

S. wallacei may be distinguished from S. densa var. scopulorum,
the taxon it sometimes most closely resembles, by its remote branches
and its abruptly adnate rather than decurrent leaf bases.

Cytology: none.

Habitat: Open and shaded rocky slopes.

Range: Southern British Columbia and adjacent mountain slopes of
Alberta, south to northern California.

Remarks: Specimens from moist, shady situations have long stems
that form loose mats and somewhat distant leaves, whereas specimens
from drier, more open situations are more compact and the leaves are
closer together. The species is rare in Alberta (Argus and White
1978).

4. Selaginella oregana DC. Eat.

Fig. 21 (a) habit; (6) portion of strobilus. Map 20.

Stems long, lax, and freely branching. Leaves bright green,
loosely imbricate, ovate-triangular, adnate for about half their length,
eciliate or slightly ciliate towards the tip; setae short, green to
whitish. Spikes sessile, inconspicuous; sporophylls ovate, long-
acuminate, eciliate towards the tip.

According to R.M. Tryon (1955) S. oregana is the only species in
the S. rupestris complex that is commonly an epiphyte. The long
pendant branches are characteristic of the species.

Cytology: None.

Habitat: Usually epiphytic on such trees as Acer macrophyllum.

Range: West coast of Vancouver Island, British Columbia, south in
coastal United States to northern California.

ufy

/// M

Fig. 20 Selaginella wallacei; (a) habit, 1 x ; (b) leaves, 10 x ; (c) portion of strobilis,
5x.

Remarks: Selaginella oregana was recently collected at Barclay
Sound and Power River on the west coast of Vancouver Island; an
early collection by Scouler, labeled Observatory Inlet, is most likely
mislabeled as to locality.

5. Selaginella densa Rydb.

Fig. 22 (a) habit; (b) leaf; (c) portion of strobilus. Map 21.

Stems forming dense cushion mats. Leaves decurrent on the
sides of the stem, with under leaves longer than upper leaves on the
same portion of the stem; leaves grooved on the back, ciliate, about 3
mm long, including the approximately 1-mm-long scabrous seta.
Setae often forming a distinct brush at the tips of the branches.
Sporophylls ovate-deltoid, apiculate, ciliate.

On the Canadian prairies both S. densa and S. rupestris occur.
The former can usually be distinguished by the longer (about 1 mm
long) setae.

Cytology: 2n = 18 (Love and Love 1976*).

Habitat: Dry prairies.

Range: Southwestern Manitoba to southeastern British Columbia,
south to New Mexico and Arizona.

5.1 Selaginella densa Rydb. var. scopulorum (Maxon) Tryon

S. scopulorum Maxon
Fig. 23 (a) habit; (b) leaf; (c) portion of strobilus. Map 22.

Differs from var. densa by its sporophylls eciliate in the upper
part and by the slightly shorter setae of the leaves.

Habitat: Alpine rocky slopes and ridges.

Range: Southwestern Alberta and southern British Columbia, south
to Texas, Arizona, and northern California.

Remarks: R.M. Tryon (1955) deemed S. densa to be one of the most
complex species in the S. rupestris complex. He upheld three varieties
that freely intergrade, var. densa, var. scopulorum (Maxon) Tryon,
and var. standleyi (Maxon) Tryon. The last variety, which has the
apex of leaves predominantly or entirely truncate in profile, was
recorded from southwestern Alberta and British Columbia by R.M.
Tryon (1955), but we have been unable to assign any specimens there.

' a

Fig. 21 Selaginella oregana; (a) habit, 1/2 x ; (b) portion of strobilus, 2 1/2 x

Fig. 22 Selaginella densa; (a) habit, 1 3/5 x ; (b) leaf, 21 x ; (c) portion of strobilus,
3 3/5 X .

Fig. 23 Selaginella densa var. scopulorum; (a) habit, 2 1/2 x; (b) leaf, 40 x ;
(c) portion of strobilus, 7 x .

6. Selaginella rupestris (L.) Spring

Fig. 24 (a) habit; ib) leaves; (c) portion of strobilus. Map 23.

Prostrate stems forming open mats. Leaves decurrent on the
sides of the stem; leaves linear-lanceolate, about 2.8 mm long
(including the approximately 0.7-mm-long scabrous seta), grooved on
the back, ciliate. Sporophylls narrowly ovate-deltoid, apiculate,
ciliate, about as long as the leaves.

Selaginella rupestris and its allies were studied in detail by R.M.
Tryon (1955); S. rupestris most closely resembles S. densa var. densa,
from which it can most easily be distinguished by its radially
symmetrical leafy stem.

Cytology: 2m = 18 (Love and Love 1976*).

Habitat: Sand dunes and open or shaded, dry, often igneous rocky
bluffs.

Range: Greenland, Quebec, and New Brunswick to northern Alberta,
south to Georgia, Michigan, Kansas, and Oklahoma.

Remarks: According to R.M. Tryon (1955), S. rupestris is the only
species that is certainly apogamous; however, some populations have
four megaspores in a sporangium, and because microsporangia are
present in the strobilis, the plants are presumably sexual. The species
is rare in Nova Scotia (Maher et al. 1978).

7. Selaginella sibirica (Milde) Hieron.

Fig. 25 (a) habit; (b) leaves; (c) portion of strobilus. Map 24.

Stems forming small intricate mats. Leaves densely appressed-
ascending, decurrent on the sides of the stem; leaves linear-ligulate,
about 2.7 mm long (including the approximately 0.5-mm-long
scabrous seta), grooved on the back, subtruncate to truncate at the
apex. Sporophylls broadly ovate-deltoid, short-apiculate, ciliate,
shorter than the leaves.

Selaginella sibirica is perhaps closest in aspect to some forms of
S. densa, from which it can usually be distinguished by the milk-white
rather than lutescent setae and the intricate rather than discrete
branches.

Cytology: 2n = 18 (Zhukova and Petrovsky 1972); 2n = 20 (Johnson
and Packer 1968).

Habitat: Dry exposed rocks and ridges.

t/.?. c

Fig. 24 Selaginella rupestris; (a) habit, 1 x ; (b) leaves, 10 x ; (c) portion of strobilus,
lOx.

Fig. 25 Selaginella sibirica; (a) habit, 1 x ; (b) leaves, 12 x ; (c) portion of strobilus,
12x.

Range: Amphi-Beringian; in North America, Alaska to
northwestern District of Mackenzie.

Remarks: R.M. Tryon (1955) noted that although the North
American material is relatively uniform, there is a phase in Asia that
has longer tawny rather than milk-white setae.

3. ISOETACEAE quillwort family

1. IsoetesL. quillwort

Perennial usually aquatic herbs. Leaves superficially grass-like,
few to numerous from a lobed, corm-like rhizome. Sporangia in a
hollow at the expanded base of the leaves and more or less covered by
the thin edges of the hollow (velum), and with a small ligule situated
above. Spores dimorphic, numerous, variously ornamented;
megaspores (female) borne in megasporangia; microspores (male)
borne in microsporangia.

This is a technically difficult poorly known genus, with perhaps
100 species in the world. Pfeiffer (1922) discussed, in monographs, all
the species (approximately 60 at the time) in the genus, and Kott
(19806) studied eight species in northeastern North America. Even
with a battery of modern techniques, such as scanning electron
microscopy (SEM), cytology, and chemical analysis (chromatography)
(Kott and Britton 1982a), the species are difficult to identify and their
phylogeny is in doubt. It is possible that a controlled hybridization
program might be informative.

Simple vegetative characters such as length of leaves, rigidity of
leaves, and their color and shape, for example, are extremely variable,
depending greatly on habitat. Most systems of classification depend
almost entirely on megaspore and microspore size and ornamentation,
which results in the necessity of microscopy. The average field
pteridologist is not likely to be challenged by these difficult plants and
should perhaps be satisfied with a determination to genus. To
distinguish quillworts from other aquatic plants such as Eleocharis,
which often look somewhat similar, one should first apply the "thumb
test." After pulling up the plant, one can feel the swollen base or corm
when it is squeezed gently between the thumb and forefinger. When
the sporophylls (leaves) are removed singly, the sporangia at their
bases are evident.

Cytology has proven useful for classification. The basic
chromosome number (X) is 11, and diploids (2X) to decaploids (10 X)
are known in nature. Because spore size is correlated with ploidy
level, careful measurement of a sample of spores is often useful for
identification of dried material.

In eastern Canada there are seven species to consider. Two are
wide-ranging and frequent. Isoetes echinospora (2 X ) with small spiny
megaspores is found usually in shallow water, and /. macrospora
(10 X) with large ridged megaspores is usually in deep water. The
other species are much less widespread, and indeed some are
considered rare.

In western Canada, there are at least six species. Three are
wide-ranging and frequent: /. echinospora, with small spiny
megaspores and almost smooth microspores; /. maritima (4X), with
larger spores and spiny or papillate microspores; and /. occidentalis

(6X), which sometimes has been confused with /. lacustris of Europe.
The last is lOX and is undoubtedly closer to /. macrospora than to /.
occidentalis. The other three species are more localized in their
distribution and include the interesting terrestrial species /. nuttallii
(2X) and the amphibious species /. howellii (2X ). It is interesting to
note that although 12 Canadian species are treated here, there is no
species occurring in Canada with megaspores that have the regular,
neat honeycombs of/, engelmannii, which occurs in the eastern United
States as far north as New York State, and o( I.japonica, which occurs
in Japan.

A. Plants terrestrial; corms more or less 3-lobed; leaves trigonous;

megaspores smooth to spongy fibrillar 9. /. nuttallii

A. Plants aquatic; corms 2-lobed; leaves usually rounded.
B. Megaspores with sparse or dense spines.

C. Spines on megaspores long and acute, not reduced in
size near the equator; microspores smooth or with very
fine thread-like spines; across Canada and abundant
1 . /. echinospora

C. Spines on megaspores blunt, sometimes confluent into
ridges, reduced to small tubercles near equator;
microspores echinate with coarse pronounced spines;
British Columbia and Alberta 2. /. maritima

B. Megaspores not spiny, but with various types of ridges.

D. Megaspores with scattered or more or less confluent
low ridges or wrinkles.

E. Amphibious; hyaline wing-margins extending 1-
5 cm above the sporangium; ligule elongated
triangular 10. /. howellii

E. Submerged; hyaline wing-margins not extending
more than 1 cm above the sporangium; ligule
cordate 11./. bolanderi

D. Megaspores with more or less connected, distinct
ridges.

F. Megaspores with short, closely set, meandering
but not anastomosing ridges or mounds; ridges
and mounds minutely spiny 3. /. eatonii

F. Megaspores with ridges variously textured,
branched, or anastomosing.

G. Megaspores with ridges rounded, smooth,
and with a smooth unornamented band
encircling the distal side of the equatorial
ridge.

H. Microspores usually smooth to slightly
low-papillate, 37-45 pm in length ....

6. /. hieroglyphica

H. Microspores roughly echinate,

25-31 pm in length

5. /. acadiensis

G. Megaspores with anastomosing or branched

ridges that have rough or sharp crests;

equatorial zone variously ornamented.

I. Megaspores usually averaging over

600 pm; leaves usually dark green and

stiff 8. /. macrospora

I. Megaspores usually averaging less
than 600 pm.
J. Ligule cordate; megaspores

cream-colored

12. /. occidentalis

J. Ligule elongate; megaspores
white.

K. Megaspores with ridges on
the distal face forming
reticulations; microspores
roughened to smoothish . . .

7. /. tuckermanii

K. Megaspores with close
ridges not forming reticula-
tions; microspores papillate
4. /. riparia

1. Isoetesechinospora Dur.

I. muricata Dur.

/. echinospora Dur. var. muricata (Dur.) Engelm.

/. braunii Dur.

/. echinospora Dur. var. braunii Engelm.

/. muricata Dur. var. braunii (Engelm.) Reed

/. echinospora Dur. ssp. muricata (Dur.) Boivin var.
savilei Boivin
Fig. 26 (a) habit; (b) ventral side of leaf base showing sporangia and
ligule; (c) megaspore. Map 25.

Corm 2-lobed. Leaves 7-25 or more, usually erect, fine and soft,
bright green to yellowish green. Sporangium to 10 mm long and 3 mm
wide, unspotted or spotted. Velum covering one-quarter to
three-quarters of sporangium. Ligule deltoid to elongate, up to
2.5 mm long. Megaspores spherical, white, averaging 480 pm (350-
550) in diameter, covered with sparse to dense spines; microspores
kidney-shaped, averaging 26 pm (23-32) in length, usually smooth or
with fme thread-like spines under SEM.

Cytology: 2m = 22 (Kott and Britton 1980*; Britton and Ceska
unpublished*).

Habitat: Shallow water up to about 1 m in depth in ponds, lakes, and
slow-moving rivers.

Fig. 26 Isoetes echinospora; (a) habit, 3/4 x ; (b) ventral side of leaf base showing
sporangia and ligule, 4 x ; (c) megaspore, 60 x .

Range: Labrador and Newfoundland to Alaska, south to
Pennsylvania, Wisconsin, Colorado, and California.

Remarks: Isoetes echinospora is Canada's most abundant and
widespread species. It is often found in shallow water in sand or
gravel. The sharp straight leaves of some specimens are seen growing
intermixed with Lobelia dortmanna and Eriocaulon. The Canadian
species is sometimes considered distinct from the European /.
echinospora and is then called /. muricata. A similar species, /.
asiatica, is known from northern Japan and Sakhalin Island.

2. Isoetes mariti ma Under w.

I. macounii A.A. Eat.

/. echinospora var. maritima (Underw.) A.A. Eat.
/. beringensis Komarov
Fig. 27, megaspore. Map 26.

Corm 2-lobed. Leaves 8-15, erect, rigid, or somewhat recurved,
dark green, 2-5 cm long, 1.5 mm wide. Sporangia oval, 4 mm long,
2.5 mm wide, covered one-third to one-half by the velum. Ligule
small, inconspicuous. Megaspores spherical, white, 490-670 pm in
diameter, covered with spines that are rather blunt and sometimes
confluent into ridges or plates; spines reduced in size near the equator.
Microspores kidney-shaped, white, 30-36 pm in length, rough with
short sharp spines under SEM; light microscopy suggests papillose or
reticulated surface.

Cytology: 2n — 44 (Britton and Ceska unpublished*).

Habitat: Shallow water to about 1 m in lakes and estuaries.

Range: Alaska, British Columbia, Alberta, and south to Washington.

Remarks: Historically, Isoetes maritima was a plant related to tidal
flats because it was found by Macoun near Port Alberni, B.C., and
treated by Hulten (1968) as a coastal species. It is now considered
present in interior lakes also, as well as east of the Rockies near
Jasper.

3. /soetes eaton/7 Dodge

/. grauesii A.A. Eat.
Fig. 28, megaspore. Map 27.

Corm 2-lobed. Leaves 12-100, 8-45 cm long or longer, erect,
usually fine and soft, bright green to yellowish green. Sporangium to
12 mm long and 5 mm wide, unspotted to tan-colored. Velum covering
one-sixth to one-quarter of sporangium. Ligule 3.5 mm long, elongate.

Megaspores flattish, white, averaging 400 pm (320-530) in diameter
with closely set, short, meandering, spiny ridges or mounds.
Microspores rounded, averaging 23 pm (22-25) in length, roughened
to smooth.

Cytology: 2m = 22 (Kott and Britton 1980*).

Habitat: Ponds and slow rivers to 1 m in depth.

Range: In Canada, apparently isolated in the Severn River, Muskoka
District, and Simcoe County, Ont. (Kott and Bobbette 1980); in the
United States occurring in the New England states. New Jersey,
Pennsylvania, and New York.

4. Isoetes riparia E nge 1 m .

/. echinospora Dur. var. robusta Engelm.

/. braunii Dur. f. robusta (Engelm.) Reed

/. canadensis (Engelm.) A. A. Eat.
Fig. 29, megaspore. Map 28.

Corm 2-lobed. Leaves 5-35, 6-35 cm long or longer, usually
erect, fine, and lax, bright green to yellow-green. Sporangium to
7 mm long and 4 mm wide, unspotted or with horizontal streaks.
Velum covering one-quarter of sporangium. Ligule elongate to 3 mm
long. Megaspores spherical, averaging 540 pm (430-680) in diameter,
with widely or closely set long and branching or short and broken
ridges. Microspores kidney-shaped, averaging 31 pm (24-35) in
length, granular-textured and usually with spaced spine-tipped
tubercules.

Cytology: 2n = 44 (Kott and Britton 1980*).

Habitat: River shores, creeks, and tidal mud flats.

Range: Southern Quebec and southeastern Ontario southward
through Maine, Vermont, and eastern New York.

Remarks: The megaspores have characteristics that resemble both
those of /. echinospora and those of /. macrospora. At times, the
megaspores appear eroded, with rough projections that one could
mistake for spines, and at other times the broken ridges approach the
sculpture of/, macrospora.

5. Isoetes acadiensis Kott
Fig. 30, megaspore. Map 29.

Corm 2-lobed. Leaves 9-35 or more, 5-21 cm long, mostly
recurved, dark green and sometimes tinged with red. Sporangium to
5 mm long and 3 mm wide, unspotted or with a few brown spots.
Velum covering one-sixth to one-third of sporangium. Ligule
elongate, to 3 mm in length. Megaspores spherical, 400-570 pm in
diameter, with smooth rounded reticulating or branching ridges.
Microspores kidney-shaped, 25-30 pm in length, roughly echinate.

Cytology: 2n = 44 (Kott 1981).

Habitat: Shallow water along borders of lakes, ponds, and rivers.

Range: Newfoundland, Nova Scotia, and New Brunswick to Maine,
Massachusetts, and New Hampshire.

Remarks: This recently described species (Kott 1981) appears to
have a distribution and ecology similar to /. tuckermanii. It was
formerly included in /. hieroglyphica because of the similarity of
megaspore sculpturing.

6. Isoetes hieroglyphica A. A. Eat.

/. macrospora Dur. f. hieroglyphica (A. A. Eat.) Pfeiffer
Fig. 31, megaspore. Map 30.

Corm 2-lobed; leaves 7-15 or more, 5-11 cm long, erect or
recurved. Sporangium to 5 mm long and 3 mm wide, usually
unspotted. Velum covering one-third of sporangium. Megaspores
spherical, white, averaging 635 pm (580-700) in diameter, with low
rounded ridges forming a network. Microspores kidney-shaped,
averaging 40 pm (37-45) in length, smoothish to low papillate.

Cytology: Unknown.

Habitat: Lakes.

Range: Southwestern Quebec and adjacent New Brunswick, Maine,
and Wisconsin. Few collections are known, and knowledge of
distribution is incomplete.

7. Isoetes tuckermanii A. Br.
Fig. 32, megaspore. Map 31.

Corm 2-lobed. Leaves 10-45 or more, 4-25 cm long or
occasionally longer, erect or recurved, soft and fine, bright green, or
sometimes yellowish green. Sporangium to 5 mm long and 3 mm
wide, usually unspotted. Velum covering one-quarter or less of
sporangium. Ligule elongate, to 2 mm long. iMegaspores spherical,
white, averaging 518 pm (400-650) in diameter, with rough-crested
ridges forming a honeycomb. Microspores kidney-shaped, averaging
27 pm (24-33) in length, tuberculate to almost smooth.

Cytology: 2n = 44 (Kott and Britton 1980*).

Habitat: Shallow water of estuaries, slow-moving streams, lakes,
and ponds.

Range: Newfoundland, Nova Scotia, New Brunswick, and the St.
Lawrence estuary region of Quebec, south through the New England
states at least to Maryland.

Remarks: The soft and fine recurved leaves and the Atlantic coastal
plain distribution are typical of this tetraploid species. It is sometimes
difficult to distinguish it from /. macrospora. Without cytology, the
easiest way to separate /. macrospora from /. tuckermanii is to
measure carefully about 20 microspores with the use of a microscope.

8. Isoetes macrospora Dur.

/. heterospora A. A. Eat.
Fig. 33, megaspore. Map 32.

Corm 2-lobed. Leaves few to 70 or more, 3-17 cm long, stiff and
erect or with recurving tips, dark green. Sporangium to 5 mm long
and 4 mm wide, usually unspotted. Velum covering one-sixth to
one-quarter of sporangium. Ligule deltoid, to 2 mm long. Megaspores
spherical, white, averaging 640 pm (400-800) in diameter, with ridges
that form honeycomb-like areas. Microspores kidney-shaped,
averaging 42 pm (32-50) in length, with the surface having evenly
spaced blunt or rounded papillae.

The usually large, coarse, and stiff-leaved plants from water as
deep as 6 m are field aids for recognition of this eastern species.

Cytology: 2m = 110 (Kott and Britton 1980*).

Habitat: Usually in deep water of oligotrophic lakes in the
Precambrian Shield.

Range: Newfoundland, Nova Scotia, Quebec, and Ontario to
Minnesota, south through the Appalachian region of the United
States to Virginia.

9. Isoetes nuttallii A. Br.

/. suksdorfii Baker
Fig. 34, megaspore. Map 33.

Corms more or less 3-lobed. Leaves up to 60, 7-17 cm long,
3-angled, slender with conspicuous hyaline margins towards the base.
Sporangium conspicuous, about 5 mm long and 1.5 mm wide. Velum
completely covering the sporangium. Ligule small, triangular.
Megaspores 400-500 pm in diameter, densely fibrillar spongy or even
smooth. Microspores 28-31 pm long, spiny tuberculate.

Cytology: 2n = 22 (Britton and Ceska unpublished*).

Habitat: Usually terrestrial, on springy but not regularly inundated
terrain.

Range: Southern Vancouver Island, British Columbia, to California.

Remarks: This species has most of its distribution to the south of
Canada. It is associated with the limited zone where the madrona, or
Arbutus, tree grows in Canada. Growth starts in the fall, and the
plant is dormant by early summer in hot, dry weather.

10. Isoetes bowellii Engelm.

/. melanopoda Gay & Dur. var. californica A. A. Eat.
Fig. 35, megaspore. Map 34.

Corm 2-lobed. Leaves 5-28, up to 30 cm long, slender but tough,
with membranous margins at the base above the sporangia.
Sporangia about 6 mm long. Velum covering about one-third of
sporangium. Ligule narrow, elongated-triangular. Megaspores about
475 pm (420-610) in diameter, with inconspicuous anastomosing
wrinkles or slightly tuberculate ridges. Microspores about 27 pm
(25-30) in length, coarsely and roughly very spinulose.

Cytology: 2/i = 22 (Britton and Ceska unpublished*).

Habitat: Muddy shores and wet depressions, both in and out of water.

Range: In Canada known only from the vicinity of Kamloops, B.C.; in
the United States from Oregon to California, east to Montana and
Idaho.

Remarks: This species, like /. nuttallii, has only the fringe of its
distribution in Canada. It is closely related to, and perhaps conspecific
with, the wide-ranging species /. melanopoda Gay & Dur. (Taylor et
al. 1975).

11. Isoetes bolanderi Hngelm.
Fig. 36, megaspore. Map 35.

Corm 2-lobed. Leaves up to 20 in number, up to 15 cm long,
slender and soft. Sporangium about 4 mm long. Velum covering
about one-third of sporangium. Ligule small, cordate. Megaspores
white, sometimes bluish, about 370 pm (350-390) in diameter, with
very low tuberculate ridges or only wrinkles. Microspores about
27 pm (25-30) in length, obscurely fine spinulose.

Aids for identification are the slender and soft leaves, which are
light yellow green, and a centre of distribution largely in the alpine
lakes of Colorado, Montana, and Wyoming.

Cytology: 2m = 22 (Britton and Ceska unpublished*).

Habitat: Lakes and ponds in often deep water, often at high
altitudes.

Range: Southwestern British Columbia and Waterton Lakes, Alta.,
south to California, Wyoming, and Arizona.

Remarks: At this time, /. bolanderi is known from very few localities
in Canada close to the United States border.

12. Isoetes occidentalis Henderson

/. lacustris L. var. paupercula Engelm.

/. paupercula (Engelm.) A. A. Eat.

/. piperi A. A. Eat.

/. flettii (A.A. Eat.) Pfeiffer
Fig. 37, megaspore. Map 36.

Corm 2-lobed. Leaves 10-30 or more, 5-20 cm long, more or less
rigid, dark green. Sporangium almost orbicular, 5-6 mm in diameter.
Velum covering one-quarter to one-third of the sporangium. Ligule
short-triangular. Megaspores 500-700 pm in diameter, cream-colored
or white, with sharp ridges and crests, and sometimes tuberculate or
almost smooth. Microspores 36-43 pm long, papillose.

Cytology: 2/i = 66 (Britton and Ceska unpublished*).

Habitat: Ponds and lakes.

Range: Coastal Alaska, British Columbia, south to California and
Colorado, at low elevations. Frequent on Vancouver Island and in
lakes around the Eraser Valley.

Fig. 27 Isoetes maritima; megaspore, 45 x .

Fig. 28 Isoetes eatonii; megaspore, 75 x .

Fig. 29 Isoetes riparia; megaspore, 50 x .

Fig. 30 Isoetes acadiensis; megaspore, 55 x .

Fig. 3 1 Isoetes hieroglyphica; megaspore, 45 x

Fig. 32 Isoetes tuckermanii; megaspore, 50 x .

Fig. 33 Isoetes macrospora; megaspore, 45 x .

Fig. 34 Isoetes nuttallii; megaspore, 60 x .

Fig. 35 Isoetes howellii; megaspore, 60 x .

Fig. 36 Isoetes bolanderi; megaspore, 75 x .

Fig. 37 Isoetes occidentalism megaspore, 50 x .

Remarks: The rigid, dark green leaves, often with a reddish base,
have suggested to some previous workers an association with the
European /. lacustris. Isoetes occidentalis is a hexaploid, not a
decaploid, and the megaspores are very variable. The almost smooth,
chalk white, fragile (easily cracked) megaspores of some collections
are distinctive, but unfortunately they can range to almost spiny
megaspores (as seen in a variant once known as /. flettii) or they can
have rounded protuberances (as in the /. piperi type). The large
microspores with characteristic papillae are more uniform.

4. EQUISETACEAE horsetail family

1. EquisetumL. horsetail

Rhizomatous perennials. Stems rush-like, jointed, sometimes
hollow, branched or unbranched; internodes of stems commonly ridged
longitudinally, with stomata in rows or bands in the grooves and with
ridges bearing siliceous tubercules or bands. Leaves small, whorled,
fused into nodal sheaths. Spores green, spherical, wrapped with 4
elaters, and borne in sporangia on sporophylls in cones. Cones
terminal on vegetative stems, or occasionally on branches, or in some
species on specialized precocious shoots that lack chlorophyll.

The genus Equisetum is the only genus in the family
Equisetaceae. It is mostly cool north-temperate in distribution.
According to Hauke (1978), there are 15 species, eight in the subgenus
Equisetum, and seven in the subgenus Hippochaete. Ten species are
known to occur in Canada, all but one of which, E. laevigatum, occur
also outside North America. All species have the same chromosome
number, n = 108. Sterile hybrids, a few of them widespread, occur
within the two subgenera.

A. Stems unbranched.

B. Fertile stems green.

C. Stomata in bands or scattered in the grooves; stems
annual; cones not apiculate.
D. Central cavity four-fifths the diameter of the

stem; sheaths with 15-20 dark brown teeth

1. E. fluviatile

D. Central cavity about one-sixth the diameter of
the stem; sheaths with 10 or fewer white-
margined teeth 2. E. palustre

C. Stomata in two lines in each groove; stems perennial
or mostly annual (E. laeuigatum); cones apiculate or
obtuse (E. laeuigatum).

E. Sheaths with 3 (4) teeth; stems lacking a central
cavity 9. E. scirpoides

E. Sheaths with 4 or more teeth; stems with a
central cavity.
F. Teeth few, not articulated at the base,

persistent 10. E. variegatum

F. Teeth numerous, articulated at the base.

G. Stems annual, soft; sheaths with black

bands at the apex only

8. E. laevigatum

G. Stems perennial, firm; sheaths becom-
ing black-banded at the base and apex

I.E. hyemale ssp. affine

B. Fertile stems not green.

H. Coning stems fleshy, lacking stomata, withering after
sporulation.
I. Sheaths with more than 14 teeth; cones 4-8 cm

long 3. f. telmateia

I. Sheaths with less than 14 teeth; cones 2-4 cm

long 4. £. arvense

H. Coning stems not fleshy, with stomata, and becoming
green and branched after sporulation.
J. Sheaths chestnut brown, flaring upwards, with
teeth cohering in several broad lobes; branches

usually branched again 5. E. sylvaticum

J. Sheaths green, rather tight, with teeth white-
margined, free or nearly so; branches usually

unbranched 6. £. pratense

A. Stems branched.

K. Sterile stems 0.5-3.0 m tall (British Columbia)

3. f. telmateia

K. Sterile stems to 0.6 m tall (widespread).

L. Fertile and sterile stems similar, green; first internode
of the primary branches (if present) equaling or mostly
shorter than the stem sheath; coning in summer.
M. Central cavity about one-sixth the diameter of

the stem 2. E. palustre

M. Central cavity about four-fifths the diameter of

the stem I.E. fluviatile

L. Fertile and sterile stems not alike; first internode of
the primary branches considerably longer than the
stem sheath; coning in spring.

N. Stem sheath teeth chestnut brown, papery

5. E. sylvaticum

N. Stem teeth dark, stiff.

O. Branches spreading, with teeth of their

sheaths deltoid 6. E. pratense

O. Branches ascending, with teeth of their
sheaths lance-attenuate .... 4. E. arvense

1. Equisetum fluviatile L.

E. limosum L.
water horsetail
Fig. 38 (a) fertile branch; (b) sterile branch; (c) node. Map 37.

Stems up to 1 m long or longer, but usually shorter, 3-8 mm
thick, annual, single, but often forming dense stands from branching
smooth light brown rhizomes. Central cavity four-fifths or more the
diameter of the stem; vallecular cavities absent; 10-30 smooth ridges
present; stomata in a broad band in each groove; sheaths tightly
appressed, with 15-20 teeth; teeth dark brown, narrow, acuminate,

^"^

Fig. 38 Equisetum fluviatile; (a) fertile stem, 1/4 x ; (b) sterile stem, 1/4 x ; (c) node,
4x.

persistent. Plant unbranched, or with branches occurring
sporadically, or verticillate. Branches up to 15 cm long, hollow, with
4-6 ridges, with the first internode shorter than the stem sheath and
with the teeth narrowly pointed. Cones up to 2.5 cm long, yellow to
brown, obtuse, peduncled, deciduous, shedding spores from May to
August.

The water horsetail may be distinguished from all other species
o{ Equisetum in Canada by the soft annual stems in which the central
cavity is about four-fifths the diameter of the stem. The stems collapse
readily when squeezed, because of the thin walls.

Cytology: n = 108 (Love and Love 1976*).

Habitat: Quiet, shallow water of rivers and lakes, wet shores, swales,
and ditches.

Range: Circumpolar; in North America from Labrador to Alaska,
south to New England, Virginia, Indiana, Wyoming, and Oregon.

Remarks: The form with simple stems or with merely a few scattered
branches has been described as f. linaeanum (Doll) Broun. This is
what Linnaeus, who believed it to be a distinct species, called E.
limosum (Fernald 1950). The species often forms extensive stands in
shallow, slow-moving water.

2. Equisetum palustre L.

marsh horsetail
Fig. 39 (a) sterile stem; (6) fertile stem; (c) node. Map 38.

Stems annual, 20-80 cm long, 1-3 mm thick, erect, solitary or
clustered, and growing from shiny, black to brown, occasionally
tuber-bearing rhizomes. Central cavity one-sixth to one-third the
diameter of the stem; vallecular cavities about the same size as the
central cavity and alternating with the 5-10 prominently angled
smooth or rough ridges; stomata in a single wide band in the valley;
sheaths green with the teeth long, narrow, black, scarious-margined.
Branches (sometimes few to none) spreading in regular whorls from
the middle nodes, with the first internode shorter than the subtending
stem sheath; sheaths with 5-6 teeth similar to the stem teeth but with
less obvious scarious margins. Cones 1-3.5 cm long, not apiculate,
deciduous, peduncled at the end of the main stems. Spores shed from
June to August.

Points by which this species may be distinguished from the
somewhat similar E. aruense are discussed under that species. The
small central cavity causes the stem to feel firm when squeezed.

Cytology: n = 108 (Taylor and Mulligan 1968*).

Fig. 39 Equisetum palustre; (a) sterile stem, 1/3 x ; (b) fertile stem, 1/3 x ; (c) node,
3x.

Habitat. Wet woods and meadows, shores, and shallow waters.

Range: Circumpolar; in North America from Newfoundland to
Alaska south to New York, Minnesota, Idaho, and California.

Remarks: Marsh horsetail usually occurs in wetter situations than
does field horsetail, with which it might be confused. It has been
reported as being poisonous to horses in Europe (Bottarelli 1968;
Richter 1961).

3. Equisetum telmateia Ehrh. ssp. braunii (Milde) Hauke

giant horsetail
Fig. 40 (a) fertile stem; (6) sterile stem. Map 39.

Stems annual, of two kinds, sterile and fertile, both erect, mostly
solitary from felted tuber-bearing rhizomes. Sterile stems to 2 m long
or longer, 0.5-2.0 cm thick; central cavity two-thirds to three-quarters
the diameter of the stem, and prominent vallecular cavities
alternating with 14-30 somewhat scabrous ridges; internodes whitish,
lacking stomata; sheaths pale below, dark above, with two-keeled
teeth; teeth long-attenuate, broadly hyaline-margined, united in
groups of two or three. Branches whorled, solid, with 4 or rarely 5
grooved scabrous ridges; stomata in bands on each side of the valleys.
Fertile stems unbranched, lacking chlorophyll, generally shorter than
the sterile stems but thicker and fleshy, with longer sheaths and
longer cohering teeth; fertile stems normally withering and dying
after spores are shed but occasionally persisting and becoming
branched. Cones to 7 cm or more long, shedding spores in April and
May.

Giant horsetail can usually be recognized easily by its size alone;
small plants can be distinguished from E. aruense by the larger, looser
sheaths, which have 2-ribbed teeth.

Cytology: n — 108 (ssp. telmateia, Sorsa 1965).

Habitat: Swamps and low wet places by lakes and streams.

Range: Subspecies telmateia is found in Europe, North Africa, and
western Asia; ssp. braunii occurs along the Pacific coast of North
America from Kodiak Island, Alaska, to California; disjunct in
Keewenaw County, Mich., but this inland station remains to be
rediscovered.

Remarks: Equisetum telmateia ssp. braunii is usually found near the
Pacific coast, but it has been collected at Penticton and observed at
Kelowna in the Okanagan Valley, B.C. The species was collected by

Fig. 40 Equisetum telmateia ssp. braunii; (a) fertile stem, 1/3 x ; (b) sterile stem,
1/3 X.

Farwell far inland on the Keewenaw Peninsula of northern Michigan
in 1880 and 1895 (Billington 1952). It has not been found there since
and may have been extirpated.

4. Equisetum arvense L.

E. boreale Bongard

E. calderi Boivin

field horsetail
Fig. 41 (a) fertile stem; (6) sterile stem; (c) habit of reduced arctic form;
id) branch node; (e) root nodules. Map 40.

Stems of two kinds, sterile and fertile, annual, growing from
dark brown to black, hairy, occasionally tuber-bearing rhizomes.
Sterile stems upright to 50 cm long or longer, to prostrate or diffusely
branched, 1.5-5 mm thick; central cavity one-third to two-thirds the
diameter of the stem, and large vallecular cavities alternating with
the 4-14 ridges; silica in dots on the ridges; stomata in 2 broad bands
in the valleys; sheaths with 4-14 short narrow dark scarious-
margined teeth and occasionally cohering in pairs. Branches solid,
whorled, spreading or ascending, mostly unbranched, 3- or 4-angled;
teeth lance-attenuate; first internode longer than the subtending stem
sheath. Fertile stems lacking chlorophyll, precocious and fleshy,
withering and dying after spores are shed, generally shorter than the
sterile stems; sheaths 0.5-2.5 mm long, with 8-12 brown, scarious-
margined, persistent, distinct or partly united teeth; terminal cone
long-peduncled, not apiculate. Spores shedding from late March to
mid May or later, depending on latitude, altitude, and season.

Sterile stems of E. arvense are perhaps most frequently confused
with E. pratense and E. palustre. Equisetum pratense is more delicate
in aspect, and the stems are whitish green; also, the teeth are deltoid
rather than lance-attenuate as in E. arvense. In E. arvense the first
internode of the branches is longer than the subtending teeth,
whereas in E. palustre the first internode is shorter than the
subtending teeth. This species is very variable and many forms have
been described; these forms, however, are the result of exposure,
peculiar ecology, season, or damage, and they do not warrant
taxonomic status (Hauke 1966).

Cytology: n = 108 (Love and Love 1976*).

Habitat: Damp open woods, low open ground and meadows, roadside
fill, and embankments, often where the surface is dry and sandy; but
even in apparently xeric situations the rhizome system can be found
penetrating saturated soil. In arctic situations it may grow on
shattered limestone or in pockets of soil with permafrost close to the
surface.

Range: Circumpolar; in North America from Greenland to Alaska,
south to Georgia, Alabama, Texas, and California.

<Z1

^ c

Fig. 4 1 Equisetum arvense; (a) fertile stem, 1/2 x ; (b) sterile stem, 1/2 x ; (c) habit of
reduced arctic form, 1/2 x ; (d) branch node, 1/2 x ; (e) root nodules, 1/2 x ,

Remarks: This species has been studied in great detail by Hauke
(1966, 1978) and others. Cody and Wagner (1981) published a review
of field horsetail in the series Biology of Canadian Weeds. The plant is
a serious weed in low-lying pastures and in some crops, and has been
reported as poisonous to livestock in Canada (Gussow 1912). Others
(Pohl 1955; Rapp 1954) have more recently reviewed the reports of
toxicity in the genus Equisetum. Field horsetail is difficult to
eradicate by cultivation because of the deeply buried rhizome that
continues to send up new branches.

5. Equisetum sylvaticum L.

wood horsetail
Fig. 42 (a) sterile and fertile stems; (6) node; (c) portion of strobilus.
Map 41.

Stems annual, of two kinds, sterile and fertile, both erect, mostly
solitary, from shiny, light brown, smooth, hairy, occasionally
tuber-bearing rhizomes. Sterile stems up to 70 cm in length and 1.5-
3 mm thick; central cavity one-half to two-thirds the diameter of the
stem, and prominent vallecular cavities alternating with the 10-18
ridges; silica tubercules in 2 rows on the ridges; stomata borne in 2
bands in the valleys; sheaths loosely inflated, with the reddish brown
papery teeth persistent, usually united into 3 or 4 groups; branches
whorled, arched, appearing lacy from secondary branches; branches
with 3 or 4 (rarely 5) ridges, with stomata in a single line on either
side of the valleys, and with teeth narrow, pointed, and spreading.
Fertile stems at first unbranched and lacking chlorophyll, precocious
and fleshy, becoming green and branched after the spores have been
released; sheaths and teeth usually larger than in sterile stems.
Peduncled cones to 3 cm in length, blunt, deciduous, shedding spores
in April and May, or later at higher latitudes.

The wood horsetail is readily recognized by its secondary
branches, which give the plant a lacy appearance, and by the loosely
inflated, reddish brown sheaths.

Cytology: n = 108 (Love and Love 1976*).

Habitat: Moist open woods, wet banks, swamps, and meadows.

Range: Circumpolar; in North America from southern Greenland
and Labrador to Alaska, south to Washington, Montana, Michigan,
and Virginia.

Remarks: This is a very variable species in which many varieties and
forms, which have little taxonomic significance, have been described
(Hauke 1978).

Fig. 42 Equisetum sylvaticum; (a) sterile and fertile stems, 1/2 x; (b) node, 5 x ;
(c) portionof strobilus, 5x.

6. Equisetum pratense Ehrh
meadow horsetail

Fig. 43 (a) sterile and fertile stems; (6) node. Map 42.

Stems annual, of two kinds, sterile and fertile, both upright,
mostly solitary from dull black rhizomes. Sterile stems whitish green,
to 50 cm in length and 1-3 mm thick; central cavity from one-sixth to
one-third the diameter of the stem, and small vallecular cavities
alternating with the 8-18 ridges; silica spicules long, thin, in 3 rows
on the ridges of the middle and upper internodes; stomata in 2 bands
in the valleys; sheaths pale, with the teeth narrow, persistent,
white-margined, and dark-centred; branches whorled, horizontal to
drooping, with 3 ridges; deltoid teeth slightly incurved, with thin
white margins. Fertile stems apparently not common, at first
unbranched and lacking chlorophyll, precocious and fleshy, becoming
green and branched after the spores are shed; sheaths and teeth about
twice as long as those of the sterile stems; peduncled cones to 2.5 cm in
length, blunt, deciduous, shedding spores from late April to early July.

This species is perhaps most easily confused with E. arvense, the
field horsetail, from which it can be separated by its more delicate
aspect, the whitish green stems that possess thin silica spicules on the
ridges of the middle and upper internodes, and the deltoid, rather than
lance-attenuate, teeth on the branches.

Cytology: n = 108 (Love and Love 1961).

Habitat: Moist woods or meadows, in sun or partial shade.

Range: Circumpolar; in North America from Newfoundland and
Labrador to Alaska, south to Montana, Michigan, and New York.

Remarks: Equisetum pratense is apparently more common in
northwestern Canada, where it sometimes forms dense stands on open
wooded floodplains of rivers and streams.

7. Equisetum hyemale L. ssp. affine (Engelm.) Stone
E. prealtum Raf.

Hippochaete hyemalis (L.) Bruhin ssp. affinis (Engelm.) Holub
scouring-rush
Fig. 44 (a) sterile and fertile stems; (6) node. Map 43.

Stems perennial, up to 1.2 m long or longer, but much shorter
throughout most of the Canadian range, 0.3-1.0 cm thick, upright,
usually unbranched, single or several together from a thick, dark
brown, dull, rough rhizome. Central cavity three-quarters or more the
diameter of the stem; small vallecular cavities alternating with 14-50

Fig. 43 Equisetum pratense; (a) sterile and fertile stems, 1/2 x ; (b) node, 5 x

Fig, 44 Equisetum hyemale ssp. affine; (a) sterile and fertile stems, 1/2 x ; (b) node,
2x.

ridges. Ridges broad, flat, or rounded, with prominent cross-bands to
double rows of tubercules; stomata in 2 lines, one on each side of the
grooves; sheaths constricted at the base, the same color as the stem
when young, but soon developing dark bands at the base and summit,
with the part between base and summit white or ashy gray; teeth
lanceolate, usually promptly deciduous, dark brown with broad
scarious margins. Cones to 2 cm in length when expanded, yellow to
black, apiculate, short-peduncled, shedding spores from June to
September, or persisting unopened till the following spring.

The scouring-rush is perhaps most easily confused with the
smooth scouring-rush, from which it can usually be distinguished by
its perennial rather than annual stems and by its sheaths, which are
only slightly expanded upward and are black-banded at the base and
apex, rather than flaring, with the black bands occurring at the apex
only.

Cytology: n = 108 (Cody and Mulligan 1982*).

Habitat: Sandy and gravelly river terraces, lakeshores, old fields,
railway embankments, and roadsides.

Range: Equisetum hyemale s. 1. is circumpolar, the ssp. affine North
American, from Newfoundland to southern Alaska, south to Texas
and New Mexico.

Remarks: This species may form extensive, dense stands on sandy
slopes; Muenscher (1955) considered it to be a weed. Early Canadian
settlers used the rough, silica-encrusted stems of scouring-rush to
clean pots and pans, hence the common name; the stems have also
been used for honing the reeds of woodwind musical instruments.
Hauke (1963) treated the subgenus Hippochaete, to which this and the
following species belong, in a monograph.

8. Equisetum laevigatum A. Br.

E. kansanum Schaffn.
Hippochaete laevigata (A. Br.) Farwell
smooth scouring-rush
Fig. 45 (a) fertile stems; {b) node. Map 44.

Stems annual, up to 1 m or more in length, but usually shorter in
our range, 2-7 mm thick, soft, upright, usually unbranched, single or
several together from a thick, dull, rough, dark brown rhizome.
Central cavity two-thirds to three-quarters the diameter of the stem;
small vallecular cavities alternating with 14-26 ridges; ridges
rounded, smooth or with cross-b^nds of silica; stomata in two lines,
one on each side of the groove; sheaths constricted at the base, flaring
towards the top, the same color as the stem except for a narrow black

Fig. 45 Equisetum laevigatum; (a) fertile stems, 1/2 x ; (b) node, 5 x

band at the apex, or in old stems the lower becoming girdled with
brown; teeth lanceolate-subulate, promptly deciduous, dark with
scarious margins. Branches resulting from injury smaller and
rougher than stems, retaining their white teeth. Cones up to 2 cm in
length when expanded, yellow to brown, blunt or only slightly
apiculate, short-peduncled, shedding spores from mid May to July.

See under E. hyemale for characters used to differentiate that
species from E. laeuigatum.

Cytology: n = 108 (Love and Love 1976*).

Habitat: Moist or dry sandy river terraces and banks, meadows, and
prairies.

Range: North American, from southern Ontario to southern interior
British Columbia, south to northern Mexico.

Remarks: Hauke (1961) has summarized the problems related to the
various forms of this taxon and the hybrid between it and E. hyemale
ssp. affine (E. X ferrissii). Equisetum laeuigatum is the only species of
Equisetum endemic in North America.

9. Equisetum scirpoides Michx.

Hippochaete scirpoides (Michx.) Farwell

dwarf scouring-rush
Fig. 46 (a) habit; (6) strobilus; (c) node. Map 45.

Stems perennial, 3-20 cm long or longer, slender, 0.5-1.0 mm
thick, usually unbranched, ascending or prostrate, arched-recurving
and flexuous, caespitose from fine branching rhizomes. Centre of stem
solid, with 3 or rarely 4 vallecular cavities alternating with the deeply
grooved ridges; stomata in single lines on either side of the ridge;
silica rosettes in lines on the crests of the ridges; sheaths green below,
black above, loose, with 3 or rarely 4 deltoid teeth; teeth
scarious-margined, subpersistent, but their subulate tips usually soon
breaking off. Cones small, 2-3 mm long, apiculate, black, shedding
spores in July or August, or persisting unopened until the following
spring.

E. scirpoides can be separated from E. uariegatum, with which it
might be confused, by its usually 3 rather than 4 teeth, and by its
flexuous stems, which are solid rather than hollow.

Cytology: n = 108 (Love and Love 1976*).

Habitat: Tundra, mossy places and woods, the stems often partly
buried in humus.

Fig. 46 Equisetum scirpoides; (a) habit, 1/2 x ; (b) strobilus, 5 x ; (c) node, 5 x

Range: Circumpolar; in North America from Labrador to Alaska,
south to New England, Washington, and Illinois.

Remarks: This is a common species in the northern boreal forest, but
is often overlooked because it is more or less buried in mosses and
humus.

10. Equisetum variegatum Schleich. ssp. variegatum

Hippochaete uariegata (Schleich.) Bruhin

variegated horsetail
Fig. 47 (a) habit; (b) node. Map 46.

Stems 6-50 cm long, 0.5-3.0 mm thick, evergreen, usually
unbranched, ascending, tufted from smooth branching rhizomes.
Central cavity one-third to two-thirds the diameter of the stem; large
vallecular cavities alternating with 3-12 furrowed ridges; stomata in
single lines on each side of the ridge; silica tubercules in two lines on
ridges, separated by the furrow; sheaths green at the base, black
above, slightly spreading; teeth lanceolate to lance-deltoid, obtuse,
persistent, with or without filiform tips, and with a brown central
portion and wide white margins. Cones 5-10 mm long, apiculate,
shedding spores in July or August or more often persisting unopened
until the following spring.

The variegated horsetail may be distinguished from the dwarf
scouring-rush by its straighter, more upright stems and by the
characters mentioned under the latter species.

Cytology: n = 108 (Love and Love 1976*).

Habitat: Tundra, moist sand, river banks, and meadows.

Range: Circumpolar; in North America from Greenland to Alaska,
south to Oregon, Utah, Michigan, New York, and New England.

Remarks: This circumpolar species, like E. aruense, is found north of
the treeline as far north as northern Ellesmere Island, where it is
quite reduced in size. Equisetum variegatum ssp. variegatum is rare in
Nova Scotia (Maher et al. 1978).

10.1 Equisetum variegatum Schleich. ssp. alaskanum (A. A. Eat.)

Hulten
Fig. 48 (a) habit; (6) node. Map 47.

Can be distinguished by its more robust stature and by its teeth
incurved, completely black, or at most narrowly white-margined, with
the black covering part or most of the sheath.

Fig. 47 Equisetum variegatum ssp. variegatum; (a) habit, 1/2 x ; (b) node, 4 x

Fig. 48 Equisetum variegatum ssp. alaskanum; (a) habit, 1/2 x ; (b) node, 5 x

Habitat: Found in habitats similar to those of ssp. variegatum.

Range: Alaska through southwestern Yukon, south along the British
Columbia coast to Vancouver Island.

Hybrids of Equisetum

Hybrids may occur spontaneously between various species in
subgenus Equisetum and in subgenus Hippochaete. They may be
recognized on the basis of their intermediacy between the parents and
by the presence of abortive spores. Some are found in nature much
more frequently than are others. Such hybrids will spread
vegetatively by fragmentation of the rhizomes. All the hybrids that
follow have been produced under experimental conditions in petri
dishes, but not all are known to occur in Canada (Duckett 1979).

E. aruense X fluuiatile (E. X litorale Kuhlewein) occurs
frequently in southern Quebec and southern Ontario, and rarely in
British Columbia.

E. arvense X telmateia {E. X dubium Dostal) occurs rarely in
Czechoslovakia.

E. arvense X palustre {E. X rothmaleri C.N. Page) occurs rarely
in Scotland.

E. arvense X pratense {E. X suecicum Rothm.) occurs in Europe.

E. fluviatile X palustre occurs in Scotland.

E. palustre X telmateia [E. X font-queri Rothm.) occurs rarely in
British Columbia.

E. pratense X sylvaticum (E. X mildanum Rothm.) occurs in
Europe.

E. hyemale ssp. affine X laevigatum {E. X ferrissii Clute, E.
hyemale var. intermedium A. A. Eat.) occurs occasionally from Quebec
to British Columbia.

E. hyemale ssp. affine X variegatum (E. X trachyodon A. Braun,
E. hyemale wa^r.jesupi (A. A. Eat.) Vict., E. variegatum var. jesupi A. A.
Eat.) occurs frequently from western Newfoundland to British
Columbia.

E. laevigatum X variegatum (E. X nelsonii (A. A. Eat.) Schaffn.)
is common in the Great Lakes region, especially adjacent to the Lakes.
Manitoulin Island was given special mention by Hauke (1963).

E. scirpoides X variegatum {E. X arcticum Rothm.) may be
represented by a specimen in DAO from the Richardson Mountains,
District of Mackenzie.

5. OPHIOGLOSSACEAE adder's-tongue family

Herbs perennial, more or less succulent. Rhizome short, bearing
one or more stalked or sessile fronds and a fertile spike or panicle.
Sporangia naked, bivalvate, producing thick-walled spores.
Gametophyte subterranean, usually without chlorophyll and
associated with an endophytic mycorrhiza.

A. Sporangia cohering in a simple spike; fronds simple, entire,
usually one; veins reticulate 1. Ophioglossum

A. Sporangia separate in a pinnate, a compound, or rarely, a simple

spike; sterile segments of fronds not simple; veins free

2. Botrycbium

1. Ophioglossum L. adder's-tongue

1. Ophioglossum vulgatum L. var. pseudopodum (Blake) Farw.

adder's-tongue
Fig. 49 (a) fronds; (b) sporangia. Map 48.

Sporophyte bearing usually 1 frond, 15-25 cm long, from an erect
rootstock. Sterile segment sessile, glabrous, entire, attached near the
middle, varying in shape from broadly lanceolate, to ovate, to
oblanceolate, 4.0-9.5 cm long, 1.5-3.0 cm wide. Fertile segment a
simple stalked spike bearing two rows of cohering sporangia.

This is a very distinctive plant, if one can find a specimen. They
tend to be hidden in grass or under sensitive ferns or are associated
with plants that have leaves that are confusingly similar in general
shape, e.g., plantains, trout lilies, or even the orchid Pogonia. Quite
often the search involves a hands-and-knees approach, near ground
level.

Cytology: n = 480 (Mulligan and Cody 1969*).

Habitat: Moist humus-rich depressions, wet meadows, and some-
times grassy hillsides and high, dry, sunny locations, or even bogs.

Range: Ophioglossum vulgatum s.l. is circumpolar; var.
pseudopodum, occurs from Nova Scotia to Ontario, Vancouver Island,
and Washington, south to Virginia, Arizona, and Mexico; var.
pycnostichum Fern, is found in the United States from Michigan
(Wagner 1971) eastward to Ohio, and south to Florida and Alabama
(Cranfill 1980); and var. alaskanum (E.G. Britt.) Christens is known
only from Kodiak Island, Alaska.

Fig. 49 Ophioglossum vulgatum var. pseudopodum; (a) fronds, 1/2 x ;
(b) sporangia, 2 x .

Remarks: The extremely high chromosome numbers in
Ophioglossum have been noted by many biologists. Ophioglossum
reticulatum from India has n = 630, and consequently every body cell
has 1260 chromosomes. However, number of chromosomes and
amounts of DNA are only two factors to consider; Selaginella apoda or
S. densa, which have /i = 9, seem to have as many morphological
characters as O. uulgatum, which has n = 480.

Love recognizes var. pseudopodum as one species, O. pusillum
Raf., and var. pycnostichum as another species, O. pycnostichum
(Fern.) Love & Love. The latter has n — ca. 630 (Love and Love 1976).
We find it difficult to evaluate the chromosome count of 2/1 = ca. 1260
for a collection of O. pycnostichum from Prince Edward Island, near
Souris, because we have not seen any herbarium material of var.
pycnostichum from Canada. The matter is further confused by the fact
that var. pycnostichum is noted for spores that are smaller (44 pm)
(Wagner 1971; Cranfill 1980) and chromosomes that are fewer in
number {n — 250-260) (Cranfill 1980) than those of var. pseudopodum
(spores, 50-54 pm) (Wagner 1971) (chromosomes, n — 480) (see above).

2. Botrychium Sw. moonwort, grape fern

Sporophyte bearing 1 to several fronds from an erect unbranched
rootstock. Roots thick and fieshy. Sterile segment sessile or stalked,
in Canadian species pinnately or palmately once to many times
decompound; venation dichotomous, open. Fertile segment stalked;
spike simple (rarely) to pinnately compound. Sporangia naked and
distinct, borne laterally on its branches.

The genus Botrychium is a small one of approximately 40 species.
There are about 20 species in North America, and we have recognized
13 in Canada's fiora. The only really common and familiar species in
Canada is B. virginianum, the rattlesnake fern, which is in a separate
section of the genus; some researchers have even placed it in a
separate genus, e.g., Osmundopteris, Japanobotrychium, or Botrypus.
It has a chromosome number of n — 92 and is not based on x = 45, as
are Canada's other species.

The genus was treated in a monograph by Clausen (1938) and
has been extensively studied by W.H. Wagner at the University of
Michigan for over 30 years. Clausen's approach to taxonomy is quite
different from Wagner's. Clausen attempted to choose a central type
and then group the variants as subspecies or varieties around this
type. Wagner, on the other hand, has attempted to separate out the
genetically distinct species and downplay the environmental forms.
The genus is one that has resisted the usual biosystematic approach.
Many of the species are basic diploids, and hybrids are considered
extremely rare. Experimental crosses are not yet feasible, because the
spores can be germinated only in low numbers and no plants have
been raised to maturity. Spore sizes have been useful diagnostically,
but spore morphology has not, and the chemotaxonomists have yet to

produce a diagnostic scheme for us. The plants are even difficult to
transplant, and we are certain that crop scientists must be amused at
our inability to separate out environmental components either by
controlled growth conditions or by field-plot techniques.

W.H. Wagner (19606) made use of mass collections, relying on
the dictum that several species of^ Botrychium often grow at the same
location. Unfortunately, this has left us with two divergent
viewpoints, one represented by workers who are impressed with the
large range of variation exhibited by one species, including variants of
different ages, and the other by workers who stress the large number
of species at one location. Herbarium material can also be frustrating.
It is not unusual to have six or more small plants on a sheet that the
collector has identified as one species (e.g., B. lunaria), but no notes of
the area covered by the collector or of his impressions regarding which
specimen was the most common type present. Various annotators
subsequently identified each of the plants as B. minganense, B.
simplex, or B. matricariifolium, for example — several species for one
sheet. W.H. Wagner and Lord (1956) and W.H. Wagner (19606) have
carefully examined Botrychium species for new taxonomic characters,
such as vernation, color of the roots, and folding of the leaf, among
others, but the nature of the plants is such that judgments are still
more subjective than one would wish. We are lacking clear objective
criteria for specific identification. Some workers continue to
emphasize differences and others similarities. Perhaps we can all
agree with Wherry (1961) when he writes about the small species,
"The search for these tiny plants, and the correct naming of them after
collection, is recommended to every fern student as a rewarding
undertaking." We trust fern students will not become frustrated by
our inability to describe clearly the limits of variation of each taxon!

A. Sterile blades usually large, up to 20 cm wide or wider at the

base, deltoid, tripinnate; lamina usually thin and papery

1. B. virginianum

A. Sterile blades various, usually leathery or fleshy.

B. Blade evergreen; plants appearing in summer and spores
maturing in autumn.

C. Segments of blade all of about the same size and shape,
ovate, obovate, rhomboid, or oblong; chief terminal
segments not elongate.

D. Sterile blades very fleshy; margins slightly
hyaline; ultimate segments somewhat acute or

obtuse and either crenate or entire

5. B. multifidum

D. Sterile blades membranous; margins not hyaline;
ultimate segments somewhat acute, usually

serrate, rarely entire 6. B. rugulosum

C. Segments of blade not all of the same size and shape;
chief terminal divisions usually elongate and little
divided.

E. Chief terminal divisions of blade broad and

rounded; blades normally winter green

4. B. oneidense

E. Chief terminal divisions of blade usually
narrower and acute or somewhat acute; blades
becoming bronze-colored in late autumn.
F. Divisions of blade deeply and finely lacerate

or divided 2. B. dissectum

F. Divisions of blade not deeply lacerate or

divided 3. B. obliquum

B. Blade deciduous; plants appearing in spring and spores soon
maturing.

G. Sterile blade deltoid, usually sessile

13. 8. lanceolatum

G. Sterile blade oblong or ovate, sessile or stalked.

H. Blades variously divided, usually bipinnate in
mature specimens.

I. Blade either palmately or pinnately
divided, ovate or broadly ovate-oblong,

usually sessile 12. B. boreale

I. Blade usually pinnately, but sometimes

ternately divided, commonly oblong,

usually sessile .... 11. B. matricariifolium

H. Blades simple or once pinnate; basal divisions

sometimes divided again, thus appearing

ternate.

J. Sterile blade short, less than 4 cm long,
stalked; divisions often remote, but at times
imbricate, with at most 3 pairs, obliquely
ovate to obovate to oblong, seldom

flabellate, often dissimilar in shape

10. B. simplex

J. Sterile blade 5-20 cm long, often imbricate;
divisions at times remote, over 3 pairs (to
10), shapes various.

K. Pinnae 3-6 pairs, bluish green, fan-
shaped to lunate, standing straight out
from the rachis, and abruptly reduced
to irregular segments at blade tip. . . .

IB. lunaria

K. Pinnae 6-10 pairs, yellowish green,

rhomboid to fan-shaped, ascending

towards the rachis, and uniformly

reduced at the blade tip.

L. Blade short, oblong-ovate,

inserted above the middle; small

divisions flabellate, often

crenate; plants less than 10 cm

tall 9. 8. dusenii

L. Blade much longer than wide,
inserted below the middle, entire
to incised; plants to 20 cm tall . .
S. B. minganense

1. Botrychium virginianum (L.) Sw. var. virginianum

rattlesnake fern
Fig. 50 (a) frond; (6) pinnule. Map 49.

Fronds erect, 50 cm long or longer, glabrous or nearly so,
deciduous. Blades broadly deltoid, sessile, attached above the middle,
bipinnate to tripinnate; ultimate segments oblong-lanceolate, toothed,
membranous or slightly fleshy. Fertile segment pinnately compound.

This is a large, fairly common species that has a broad
distribution across Canada. The highly divided blade and
characteristic fertile segment make identification simple.

Cytology: n = 92 (Britton 1953*; Love and Love 1976). This number
is a departure from the x — 45 found in the other species.

Habitat: Usually in dry to somewhat moist deciduous woodlands, but
occasionally in wet cedar woods and boggy areas.

Range: Newfoundland to British Columbia, south to Florida and
California.

Remarks: The common name comes from the fanciful similarity in
appearance of the cluster of unopened sporangia to the rattles on the
tail of a rattlesnake.

1.1 Botrychium virginianum (L.) Sw. var. europaeum Angstr.

B. virginianum (L.) Sw. var. laurentianum Butters
Fig. 51, frond. Map 50.

Similar to var. virginianum but usually smaller and stiffer.
Blade leathery; ultimate segments less toothed, often crowded, and
overlapping.

Habitat: Thickets and damp often coniferous woods.

Range: More northern than var. virginianum; Labrador to Alaska,
south to the northern United States; northern Eurasia.

Remarks: This variety would seem to be somewhat clinal and may
not merit taxonomic rank; compare with Dryopteris fragrans var.
remotiuscula and Woodsia alpina iW . bellii).

100

Fig. 50 Botrychium virginianum var. virginianum; (a) frond, 1/3 x ; (b) pinnule,
2 1/2 X .

101

Fig. 51 Botrychium virginianum var. europaeum; frond, 1/2 x

102

The variety is, however, quite striking when one compares the
far northern plants with those in the south. It is rare in the District of
Mackenzie (Cody 1979) and the Yukon (Douglas et al. 1981).

Species 2-6 comprise a group of five species that are sometimes
called the "fall botrychiums." W.H. Wagner (1959, 19606, 1962) and
W.H. Wagner and Rawlings (1962) have considered this group of
Botrychium species in the subgenus Sceptridium in some detail.
Wagner recognizes four species in this group for northeastern United
States and adjacent Canada, as follows: Botrychium dissectum
(including f. obliquum), B. multifidum, B. oneidense, and B.
rugulosum. We are following Wherry (1961) in recognizing five
species, whereas earlier workers, e.g., Clausen (1938) have recognized
as few as two species, B. dissectum and B. multifidum. All our species
are basic diploids with x = m = 45.

2. Botrychium dissectum Spreng.

cut-leaved grape fern
Fig. 52, frond. Map 51.

Fronds up to 27 cm long; stem and blade less coriaceous than B.
multifidum. Blades long-petioled, triangular, ternate, attached at or
near the base; ultimate divisions of blade cut in linear segments;
segments more or less notched at the apex. Fertile segment
paniculate. Spores mature from September to November.

Cytology: n = 45 (Britton 1953*; W.H. Wagner 19606).

Habitat: Sterile hilltops, dry pastures, dry woodlands, and grassy
banks.

Range: New Brunswick to Ontario and Minnesota, south to North
Carolina, Tennessee, and Missouri.

Remarks: Extremely dissected and lacerated plants appear almost
skeletonized and are very distinctive. The fronds are often bronze or
even turn reddish in late fall. This species occurs in southeastern
Canada.

3. Botrychium obliquum Muhl.

B. dissectum Spreng. var. obliquum (Muhl.) Clute

B. dissectum Spreng. f. obliquum (Muhl.) Fern.
Fig. 53, frond. Map 52.

Fronds 30 cm long or longer; stem and blade somewhat
coriaceous. Blades triangular, ternate, attached at or near the base;

103

Fig. 52 Botrychium dissectum; frond, 1/2 x

104

Fig. 53 Botrychium obliquum; frond, 1/2 x .

105

ultimate segments of blade elongate, somewhat acute, somewhat
divided below; margins of segments entire or inconspicuously crenate.
Fertile segment paniculate. Spores mature from September to
November.

Cytology: n = 45 (Britton 1953*).

Habitat: Sterile fields, dry pastures, meadows, thickets, dry
woodlands, and rich swampy woods.

Range: Nova Scotia to Ontario, Wisconsin, and Iowa, south to South
Carolina, Georgia, and Louisiana.

Remarks: This taxon, which often grows intermixed with B.
dissectum, is considered a mere form by W.H. Wagner (1961). For
many field workers, it is much more easily identified than the others
and may occur in large numbers. The genetic relationships between
B. obliquum and B. dissectum are still obscure (see also comments by
Wherry 1961).

4. Botrychium oneidense (Gilbert) House

B. dissectum Spreng. var. oneidense (Gilbert) Farw.
B. dissectum Spreng. f. oneidense (Gilbert) Clute

Fig. 54 (a) sterile frond; (6) fertile frond; (c) sporangia. Map 53.

Fronds 40 cm long or longer; stem and blade somewhat
coriaceous. Blades triangular, ternately decompound, little divided,
attached at or near the base; chief terminal segments of blade broadly
ovate and obtuse. Fertile segment paniculate. Spores mature in
September and October.

Cytology: n = 45 (W.H. Wagner 1955)

Habitat: Rich moist woodland.

Range: New Brunswick to Ontario and Minnesota, south to North
Carolina, Ohio, and Indiana.

Remarks: This species has had a checkered career. It has been
variously treated as a variety, a form, or even a hybrid of both B.
dissectum and B. multifidum. It was studied in some detail by W.H.
Wagner (1961) and is considered a good species. The broader, more
rounded divisions and the more shady habitat are characteristic of the
species.

Botrychium oneidense appears to be a rare plant in Canadian
flora. We have seen very few good colonies of it. Most plants are with-
out fertile segments, and many of the collections seem to be occasional
plants that are selected from larger colonies of B. obliquum. Some
specimens of B. multifidum also closely resemble B. oneidense.

106

Fig. 54 Botrychium oneidense; (a) sterile frond, 3/5 x ; (b) fertile frond, 3/5 x ;
(c) sporangia, 5 x .

107

5. Botrychium multifidum (Gmel.) Rupr

leathery grape fern
Fig. 55, frond. Map 54 (s.l.).

Fronds up to 20 cm long; stem and blade coriaceous. Blade
evergreen, long-petioled, ternate, attached near the base of the plant;
ultimate segments of blade crowded, sometimes imbricate, varying
from flabellate to ovate, but more or less the same size, obtuse or
somewhat acute. Fertile segment paniculate. Spores mature in
August and September.

Cytology: n = 45 (W.H. Wagner 1955, 19606; Taylor and Mulligan
1968*).

Habitat: Grassy hillsides, sterile fields, exposed meadows, and sandy
open places.

Range: Labrador and Newfoundland to British Columbia and
southern District of Mackenzie, south to New York, Minnesota, and
Wisconsin; northern Eurasia.

Remarks: The species is rare in the District of Mackenzie (Cody
1979) and Saskatchewan (Maher et al. 1979).

5.1 Botrychium multifidum (Gmel.) Rupr. var. intermedium

(D.C. Eat.)Farw.
B. silaifolium Presl
Fig. 56, frond.

Similar to var. multifidum, but taller. Blades much larger, up to
20 cm wide and 15 cm long; ultimate segments of blade not so closely
crowded, varying from oblong to obovate or ovate and usually
somewhat crenate. Spores mature in August and September.

These larger plants with many divisions of the blade are
strikingly distinctive in their bright yellow green coloration.

Habitat: Open fields, pastures, dry hillsides, borders of woods, and
sandy places.

Range: Southern Labrador and Nova Scotia to British Columbia and
Alaska, south to New York, Pennsylvania, Michigan, Montana, and
California.

6. Botrychium rugulosum W.H. Wagner

Botrychium ternatum Am. auth.
Fig. 57 frond. Map 55.

108

Fig. 55 Botrychium multifidum; frond, 1 x

109

Fig. 56 Botrychium multifidum var. intermedium; frond, 1/2 x

110

Fronds 25 cm long or longer, thin and membranous. Blades
inserted at the base, ternate, with the three major divisions stalked;
ultimate segments of blade all about the same size, ovate to oblong,
acutish, serrate or entire, and concave in the living state. Fertile
segment paniculate. Spores mature from August to October.

Cytology: n = 45 (W.H. Wagner 19606).

Habitat: Swampy woods, brushy fields, and wooded stream banks.

Range: Western Quebec and Ontario south and west to New York,
Michigan, Indiana, and Iowa; Asia.

Remarks: W.H. Wagner (W.H. Wagner and F.S. Wagner 19826) has
recently separated this taxon from the closely related Japanese B.
ternatum (Thumb.) Swartz. Botrychium rugulosum has been sought in
Canada at some length. We have found no good colonies of the species,
although we can occasionally find a few plants that match Wagner's
descriptions (W.H. Wagner 1959, 19606). We appreciate his
confirming identifications of these collections. We consider the species
to be extremely rare in Canada's fiora.

The remaining species of this genus produce their fertile spikes
in spring and early summer.

7. Botrychium lunaria (L.) Sw.

moonwort
Fig. 58, frond. Map 56.

Fronds up to 25 cm long, somewhat leathery. Blades more or less
oblong, sessile, inserted at or below the middle, pinnate; segments of
blade opposite, fiabellate and often overlapping, with margins entire
or somewhat incised. Fertile segment racemose or paniculate. Spores
mature from June to August.

Cytology: n = 45 (W.H. Wagner and Lord 1956; Love and Love
1976*).

Habitat: Open, turfy, or gravelly slopes, shores, and meadows,
usually on basic soils.

Range: Circumpolar; in boreal North America from Greenland,
Labrador, and Newfoundland to British Columbia and Alaska, south
to Maine, Michigan, Wyoming, Colorado, and California.

Remarks: The moonwort has a long and illustrious history in early
herbals. We are told that the "seeds" can make one invisible or can be

111

Fig. 57 Botrychium rugulosum; frond, 1/2 x

112

used to unlock doors. It is more common northward in Canada and is
an early quest for the field worker to see and photograph. The fully
mature plants are distinctive, but the species should be identified with
care when it is compared with B. minganense (W.H. Wagner and Lord
1956). Moonwort is rare in the District of Mackenzie (Cody 1979),
Nova Scotia (Maher et al. 1978), and Saskatchewan (Maher et al.
1979).

8. Botrychium minganense Vict.

Botrychium lunaria (L.) Sw. var. minganense (Vict.) Dole
Fig. 59, frond. Map 57.

Fronds up to 30 cm long, somewhat membranous. Blades
narrowly oblong, sessile or nearly so, inserted below the middle, pin-
nate or occasionally pinnate-pinnatifid at the base; segments of blades
opposite, obovate, rhomboidal or oblong, frequently incised, remote.
Fertile segment paniculate. Spores mature in July and August.

Botrychium minganense can be distinguished from B. lunaria by
its yellowish green hue and by its trough-shaped sterile segments,
which are ascending rather than at right angles to the stalk and which
rarely overlap with each other. (W.H. Wagner and Lord 1956).

Cytology: n = 90 (W.H. Wagner and Lord 1956; Love and Love
1976).

Habitat: Marly meadows and open alpine areas.

Range: Southern Labrador to Hudson Bay, Alaska, and British
Columbia south to New York, Michigan, Montana, and California.

Remarks: This tetraploid species of uncertain origin, other than as a
derivative of B. lunaria, is usually present at B. lunaria localities. It
has been given various ranks, from form to species, and was originally
described by one of Canada's eminent botanists, Frere Marie- Victorin,
from material collected on the Mingan Islands in the lower St.
Lawrence River. It is rare in the District of Mackenzie (Cody 1979).

9 Botrychium dusenii (Christ) Alst.
Fig. 60, frond. Map 58.

Fronds up to 13 cm long; stem and blade deciduous. Blades short,
oblong-ovate, inserted above the middle, sessile or nearly so, pinnate;
segments of blade usually remote, about as wide as long, often
spatulate and distinctly petioled, sometimes cuneate, often crenate.
Fertile segment pinnate, or the lowermost pinnae again pinnate.
Spores mature from July to early September.

113

Fig. 58 Botrychium lunaria; frond, 2/3 x

(\-

Fig. 59 Botrychium minganense; frond, 4/5 x

114

Cytology: n — 45 (W.H. Wagner and Lord 1956). From material that
had previously been considered to be western B. minganense.

Habitat: Mountains and western plains.

Range: District of Mackenzie, Alberta, and British Columbia, south
to California and Arizona; South America.

Remarks: W.H. Wagner, after a trip to a scientific meeting in
Edmonton, said that the limited amount of western material available
of B. minganense, B. lunaria, B. simplex, and B. matricariifolium
should be investigated and that a species thought to be restricted to
South America, B. dusenii (Reeves 1977), closely matched specimens
from western Canada. W.H. Wagner took up his own challenge and
has recently reported (W.H. Wagner and F.S. Wagner 1981) further
studies on western moonworts. He says, "Plants we previously
identified as the South American B. dusenii (Christ) Alston from
various western states may prove to be variations of B. crenulatum"
We are also told, "More detailed reports on all of these plants will be
made in the future." Accordingly, our treatment here is tentative,
pending further study of these interesting, diminutive plants.

10. Botrychium simplex E. Hitchc. var. simplex
least grape fern

Fronds up to 16 cm long, rather fleshy. Blades simple, lobed or
pinnately divided, inserted at the base or towards the middle;
segments of blade oblong, rhomboid or reniform, and usually
overlapping, with the basal segments occasionally pinnatifid. Fertile
segment simple or compound. Spores mature in late May and June.

Cytology: n = 45 (W.H. Wagner 1955).

Habitat: Pastures, meadows, lakeshores, and gravelly slopes.

Range: Newfoundland to Ontario and British Columbia, south to
Pennsylvania, Wisconsin, Colorado, and California; Europe and
Japan.

Remarks: The least grape fern is indeed a small species and is
therefore often overlooked in the field. Unfortunately, young or small
individuals of other species often resemble this one. Tryon et al.
(1953) say that to distinguish this species from the very small B.
matricariifolium may require measuring the size of spores — B.
simplex (35-50 pm) and B. matricariifolium (25-35 pm). Individuals
with a few sporangia on the edge of the blade occur quite often. This is

115

also true for B. minganense. This species is rare in Nova Scotia
(Maheretal. 1978).

10.1 Botrychium simplex E. Hitchc. var. tenebrosum (A. A. Eat.)

Clausen
Fig. 61, frond. Map59(s.l.).

Similar to var. simplex, but usually very slender and taller.
Blade not sessile, inserted above the middle, simple or lobed; lobes
subopposite, obovate-oblong.

Habitat: Deep w^oods and the borders of swamps.

Range: New Brunswick to British Columbia, south to Pennsylvania,
Michigan, Minnesota, and Washington.

Remarks: W.H. Wagner (19606) has suggested that this is a
"questionable" variety. It does however, look quite distinct from var.
simplex, being long-attenuate, with a high insertion of the very small
blade. The habitat of this variety, which is in deep cedar swamps,
seems quite unlike that of var. simplex, which is open pastures and
meadows.

1 1 . Botrychium matricariaefolium A. Br.

matricary grape fern
Fig. 62, frond. Map 60.

Fronds 28 cm long or longer, membranous to fleshy. Blades
narrowly deltoid to ovate, short-stalked, inserted above the middle,
pinnatifid to bipinnate-pinnatifid; segments of blade blunt and
usually toothed. Fertile segment paniculate. Spores mature in June
and July.

This species is somewhat larger than B. simplex and seems to
occur more frequently than some of the other species. The shape of the
blade is variable (deltoid to ovate) but it is stalked, and the toothed
segments are distinctive (compare lanceolatum).

Cytology: m = 90 (W.H. Wagner 1955; W.H. Wagner and Lord 1956;
Love and Love 1976*) This is a tetraploid species of uncertain origin.

Habitat: In acid soil in old sandy and sterile fields, dry wooded
slopes, rocky woods, moist cedar woods, and rich swamps.

Range: Southern Labrador and Newfoundland to southern British
Columbia, south to Pennsylvania, Minnesota, and Idaho.

116

Fig. 60 Botrychium dusenii; frond, 1 x

!\-

Fig. 61 Botrychium simplex var. tenebrosum; frond, 1 x

117

Fig. 62 Botrychium matricariifolium; frond, 1/2 x

118

Remarks: The species is distinctive for a lesser Botrychium, although
specimens with deltoid blades and a more fleshy nature (found in
northern, open sites) mimic B. boreale, and the very small plants look
like B. simplex var. simplex. Botrychium matricariifolium is rare in
Manitoba (White and Johnson 1980) and Saskatchewan (Maher et al.
1979).

12. Botrychium boreale Milde ssp. boreale
Fig. 63, frond. Map 61.

Fronds erect, up to 26 cm long, stout and fleshy. Blades
pinnately divided, sessile, inserted above the middle; primary
divisions of blades palmately lobed or crenate, acute at the apex.
Fertile segment simple or paniculate. Spores mature in June and
July.

Cytology: /i = 90 (W.H. Wagner 1963, without variety).

Habitat: Grassy and rocky slopes and alpine meadows.

Range: Circumpolar, but in North America known only in Greenland
and Alaska.

12.1 Botrychium boreale Milde ssp. obtusilobum (Rupr.) Clausen

B. boreale Milde var. obtusilobum (Rupr.) Broun
Fig. 64 (a) frond; (b) sporangia. Map 62.

Similar to ssp. boreale, but with the divisions oblong and obtuse
at the apex.

Habitat: Grassy and rocky alpine slopes and meadows, extending
down into open deciduous and evergreen woodland.

Range: Western Alberta and interior British Columbia, south to
Washington and Montana, and north through the Yukon and Alaska
to eastern Siberia.

Remarks: This is the western counterpart of the eastern B.
matricariifolium, but usually lacks the short blade stalk, is more often
deltoid, and has the fleshy characteristics of so many of the smaller
Botrychium species (Reeves 1977). It is rare in the District of
Mackenzie (Cody 1979), the Yukon (Douglas et al. 1981), and Alberta
(Argus and White 1978).

119

Fig. 63 Botrychium boreale ssp. boreale; frond, 3/4 x

120

Fig. 64 Botrychium boreale ssp. obtusilobum; (a) frond, 1/2 x ; (b) sporangia, 7 x .

121

13. Botrychium lanceolatum (Gmel.) Angstr. var. lanceolatum

lance- leaved grape fern
Fig. 65, frond. Map 63.

Fronds 20 cm long or longer, stout and fleshy. Blades sessile,
broadly deltoid, inserted near the summit; segments of blades
lanceolate, pinnatifid. Fertile segment paniculate. Spores mature in
July and August.

The sessile triangular blade with sharp toothing (but not as
acute as in taxon 13.1) is characteristic of the species (Reeves 1977).
See also remarks under B. matricariifolium.

Cytology: n = 45 (W.H. Wagner 1963; Love and Love 1976).

Habitat: Alpine meadows, dry slopes, sandy open places, and
swampy forests.

Range: Greenland, south through Newfoundland, eastern Quebec,
and northern Maine; Aleutian Islands, south through British
Columbia to Washington, Wyoming, and Colorado; Eurasia.

Remarks: This species is rare in the Yukon (Douglas et al. 1981) and
Saskatchewan (Maher et al. 1979).

13.1 Botrychium lanceolatum (Gmel.) Angstr. var.
angustisegmentum Pease & Moore
B. angustisegmentum (Pease & Moore) Fernald
Fig. 66, frond. Map 64.

Differs from var. lanceolatum in being lax and membranous,
with the divisions of the blade narrow and more acute.

Cytology: m =: 45 (W.H. Wagner 1955; Love and Love 1976*).

Habitat: Shaded woodlands, edges of swamps, along streams, and
occasionally in open fields.

Range: Newfoundland to southern Ontario and Michigan, south to
Pennsylvania and Ohio.

Remarks: This more southern variety was sufficiently distinct for
Fernald to give it specific status. The blade is much thinner or less
fleshy than that of var. lanceolatum, and the divisions are narrower
and very sharp-pointed. It is a dark but bright green, rather than the
bluish green of B. matricariifolium. It is a rare plant in Ontario, and
some of the records we have for the urbanized south of that province
now represent extirpated plants. B. lanceolatum var.
angustisegmentum is rare in Nova Scotia (Maher et al. 1978) and
Ontario (Argus and White 1977).

122

Fig. 65 Botrychium lanceolatum
var. lanceolatum; frond, 1/2 x .

Fig. 66 Botrychium lanceolatum
var. angustisegmentum; frond, 2/3 x

123

6. OSMUNDACEAE Howering fern family

1. Osmunda L. flowering fern

Tall ferns of marshy places, frequently in large clumps; fertile
fronds surrounded by sterile ones. Rootstocks creeping or suberect.
Fronds with stipes winged at the base. Blades with free, usually
forked veins extending to the margins. Sporangia naked, large,
globose, bivalved, borne on modified, contracted pinnae.

The genus Osmunda, which contains 12 species, is one of three
genera in the family Osmundaceae. The species occur from Canada's
temperate region south to tropical swamps. We have three species in
Canada.

A. Fronds bipinnate, some of them fertile at the tip; pinnules finely

toothed 1. O. regalis var. spectabilis

A. Fronds pinnate; sterile pinnae deeply pinnatifid; lobes usually
entire.
B. Fertile fronds with fertile pinnae near the middle; no tufts

of wool at the base of the pinnae of sterile fronds

2. O. claytoniana

B. Fertile and sterile fronds separate; pinnae of sterile fronds
with a tuft of wool in the axils 3. O. cinnamomea

1. Osmunda regalis L. var. spectabilis (Willd.) Gray

royal fern
Fig. 67 (a) frond; (b) sporangia. Map 65.

Fronds up to 1 m long and 25 cm wide, bipinnate. Pinnules
oblong to lance-oblong, up to 6 cm long, sessile, subentire to finely
toothed, rounded to the base; fertile pinnules contracted, borne at the
tip of the frond.

Forma anomala (Farw.) Harris has the normally fertile part of
the frond intermixed with sterile pinnae, and some of the normally
sterile pinnae more or less fertile.

The royal fern may be recognized by the clumps of large doubly
pinnate fronds. The fertile pinnules are contracted at the tips of the
fertile fronds.

Cytology: n = 22 (Cody and Mulligan 1982*).

Habitat: Swamps, low lying woods, wet marshy meadows, and cedar
bogs.

124

Fig. 67 Osmunda regalis var. spectabilis; (a) frond, 1/3 x ; (b) sporangia, 3 x .

125

Range: Variety spectahilis is found in eastern North America,
Newfoundland to Rainy River District, Ont. (not to Saskatchewan as
given in some manuals), south to Florida, Alabama, Mississippi,
Louisiana, and Texas; var. regalis occurs in Eurasia.

Remarks: Three other subspecies are known to occur in Europe and
Asia. All have the same chromosome number, as do all the members
of the family Osmundaceae counted to date.

The range "to Sask." given in Fernald (1950) is probably based on
the reports in Macoun (1890) "on Muskeg Island, Lake Winnipeg (J.M.
Macoun). Through Canada westward to the Saskatchewan (Eaton)."
No specimens have been found to substantiate these early reports, and
no recent collections have been seen from west of the Rainy River
District, Ont.

2. Osmunda claytoniana L.

interrupted fern
Fig. 68 (a) frond; (6) sporangia. Map 66.

Fronds up to 1.2 m long, 15-25 cm wide or wider, pinnate-
pinnatifld. Sterile pinnae oblong-lanceolate; pinnules elliptic-oblong
to oblong-oval, blunt; lower pinnules 1.3-1.8 cm long; young pinnae
and rachis with a rusty wool, promptly glabrous. Fertile frond with 3-
5 pairs of dark brown contracted fertile pinnae situated at about the
middle.

The contracted fertile pinnae in the middle of the fertile fronds
give the interrupted fern its name.

Cytology: n = 22 (Britton 1964*).

Habitat: Moist wooded slopes, swamp margins, and open thickets.

Range: Eastern North America, Newfoundland to southeastern
Manitoba, south to Georgia, Kentucky, and Arkansas.

Remarks: Another subspecies, ssp. uestita (Wahl.) Love and Love,
occurs in Asia. It has the same chromosome number as ssp.
claytoniana.

3. Osmunda cinnamomea L.

cinnamon fern
Fig. 69 (a) fertile frond; (b) sterile frond; (c) sporangia. Map 67.

Sterile fronds up to 1.2 m long, 15-20 cm wide or wider, similar to
O. claytoniana, but with a tuft of wool at the base of each of the
linear-lanceolate pinnae. Fertile fronds shorter than the sterile, the

126

Fig. 68 Osmunda daytoniana; (a) frond, 1/4 x ; (b) sporangia, 4 x

127

Fig. 69 Osmunda cinnamomea; (a) fertile frond, 1/4 x ; (b) sterile frond, 1/4 x ;
(c) sporangia, 2 x .

128

dark brown pinnae contracted, withering after the spores are cast.
Immature sterile and fertile fronds covered with a thick rusty wool,
still partly present on the fertile fronds even at maturity.

Forma frondosa (T. & G.) Britt. has the fertile frond partly leafy
and the fertile and sterile pinnae variously mixed.

The cinnamon fern can quickly be recognized by the densely
woolly cinnamon-colored fertile fronds, which quickly shrivel after the
spores are shed.

Cytology: n = 22 (Britton 1964*).

Habitat: Low ground, thickets, and wet marshy woods.

Range: In eastern North America from Newfoundland to Rainy
River District, Ont., south to the Gulf States and New Mexico.

Remarks: An Asiatic subspecies, ssp. asiatica (Fern.) Hulten, is
found in eastern Asia. Both subspecies have the same chromosome
number.

Hybrids of Osmunda

In spite of the fact that our species of Osmunda frequently occur
side by side, only one hybrid, 0. X ruggii Tryon (0. claytoniana X
regalis) has yet been found, and that only twice (W.H. Wagner et al.
1978). This hybrid is not known in Canada.

129

7 SCHIZAEACP]AE

1. Schizaea Sm.

1. Schizaea pusilla Pursh

curly-grass
Fig. 70 (a) fronds; (6) fertile pinnae. Map 68.

Fronds dimorphic, forming dense tufts. Sterile fronds slenderly
linear, spiraling and curling, 8 cm or more in length. Fertile fronds
erect, very slender, up to 12 cm in length, with 3-8 tiny crowded pairs
of pinnae at the tip folded together and thus looking one-sided.
Sporangia pear-shaped, bivalvate, in a double row along the vein on
the back of the pinnae.

This species is so insignificant that only the keenest observers
can find its grass-like fronds, which are hidden among other
vegetation. The tiny infolded pinnae at the tip of the fertile fronds are
unlike any other fern in Canada.

Cytology: n = 103 (W.H. Wagner 1963).

Habitat: Damp peaty and sandy depressions, sphagnum bogs, and
low, mossy, open woods and crevices along shores, both on tablelands
and lowlands; easily overlooked among other vegetation.

Range: Newfoundland and Nova Scotia and on the Pine Barrens of
New Jersey. There is a specimen in the University of Toronto
Herbarium collected by E.A. Moxley reputedly from Sauble Beach,
Bruce County, Ont. However, various botanists have searched for it
there without success, and the locality is questionable. The site is now
a cottage area, and so if the plant did grow there it has been
extirpated.

Remarks: Schizaea is one of four genera of the family Schizaeaceae.
The approximately 160 species in the family are mostly tropical in
distribution. The climbing fern, Lygodium palmatum, which is rare
from Florida to New Hampshire in the eastern United States, is one of
this family.

130

Fig. 70 Schizaea pusilla; (a) fronds, 1 x ; (b) fertile pinnae, 4 x .

131

8 HYMEN0PHYLLACP:AP] filmy fern family

1. Mecodium Copeland filmy fern

1 . Mecodium wrightii (van den Bosch) Cope land

Hymenophyllum wrightii van den Bosch
Fig. 71 (a) fronds; (6) pinnule with sporangia. Map 69.

Fronds up to 7 cm long, arising along the extensively creeping
and branched thread-like rhizome. Stipes thread-like, blackish, with
a tuft of hair-like scales at the base and narrowly decurrent from the
basal pinnae. Blades 3-5 cm long, pinnate; pinnae pinnatifid through
3 or 4 dichotomous branchings; ultimate segments few, linear, blunt,
very delicate, pale green and almost translucent so that the dark
dichotomously branched veins are very conspicuous. Sori at the ends
of the veinlets. Indusia bivalvate to the base.

This delicate fern could be easily passed over in its moist, mossy
habitat. The dark, dichotomously branched veins are conspicuous in
the pale green almost translucent pinnae.

Cytology: x = m = 27(?) (Manton and Vida 1968); 42(?) (Tatuno and
Takei 1969).

Habitat: Forming mats on shaded cliff faces, boulders, and bases of
trees.

Range: In North America from the Alaskan Panhandle, south to
Vancouver Island, and apparently of very local occurrence; also in
Japan and Korea.

Remarks: This is the only filmy fern known in Canada. It was first
found at Dawson Inlet on the west coast of Graham Island in the
Queen Charlotte Islands by H. Persson while he was collecting
mosses. Its disjunct distribution in North America, from the main
range in Japan and Korea, was discussed by Iwatsuki (1961). W.
Schofield, also while searching for mosses, has since discovered the
gametophyte on nearby Chaatl Island and on the British Columbia
mainland near Prince Rupert. The gametophytes bear deciduous
marginal cell-masses, which allow the gametophyte to reproduce
asexually. Thus they may be found in quantity where the sporophyte
generation is absent. A detailed description of the habitat is given by
Calder and Taylor (1968). Taylor (1967) described the gametophyte
and reported its occurrence in British Columbia, and Cordes and
Krajina (1968) reported finding male gametophytes on old bark or
decaying wood of Sitka spruce at three sites on the west coast of
Vancouver Island.

132

9 PTERIDACEAE

Ferns delicate to coarse, deciduous, or evergreen. Fronds pinnate
to decompound. Sori marginal, protected by the indusium, which
opens toward the margin, or by the reflexed margins of the pinnae, or
borne along the veins and lacking an indusium.

^ A large family of mainly terrestrial ferns, comprising over 60
genera. Some tropical and subtropical members are arborescent.

A. Fronds covered beneath by a conspicuous white or golden yellow
powder; sori borne along the veins and lacking an indusium ....

7. Pityrogramma

A. Fronds without a white or golden yellow powder; sori marginal.
B. Sori usually confluent as a marginal band.

C. Fronds coarse, scattered, from stout elongate and

forking rhizomes 2. Pteridium

C. Fronds tufted from a very short rhizome.

D. Segments of frond bead-like .... 3. Cheilanthes
D. Segments of frond not bead-like

E. Pinnules and segments of frond jointed at

the base 5. Pellaea

E. Pinnules and segments of frond not jointed
at the base.
F. Stipes herbaceous, green except at the

base 6. Cryptogramma

F. Stipes wiry, dark and shiny

4. Aspidotis

B. Sori distinct, short, mostly not confluent

G. Stipe and fronds glabrous 8. Adiantum

G. Stipe and fronds glandular-hairy

1. Dennstaedtia

1. Dennstaedtia Bernh.

1 . Dennstaedtia punctilobula (Michx.) Moore

Dicksonia pilosiuscula Willd.

Dicksonia punctilobula (Michx.) A. Gray

hay-scented fern
Fig. 72 (a) frond and rhizome; (b) pinnule with sporangia. Map 70.

Fronds 30-70 cm long or longer, arising from slender, naked,
freely creeping and forking rhizomes. Stipes pale brown, lustrous,
chaffless. Rachis and under surface of the blades minutely glandular-
pubescent. Blades lanceolate, bipinnate; pinnae lanceolate; pinnules
pinnatifid with toothed lobes. Sori minute, round, situated on the
upper margin of the underside of the lobes. Indusia cup-shaped.

133

Fig. 71 Mecodium wrightii; (a) fronds, 1/2 x ; (b) pinnule with sporangia, 10 x

Fig. 72 Dennstaedtia punctilobula; (a) frond and rhizome, 1/4 x ; (b) pinnule with
sporangia, 7 x .

134

The hay-scented fern has sometimes been confused with the lady
fern and bulblet fern. It can be distinguished by the characteristic
odor when crushed, the lustrous brown stipes, the glandular hairs on
the rachis, which can readily be seen when held up to the light, the
rather hard round sori, and its occurrence in large patches of single
fronds arising from the underground rhizome.

Cytology: m = 34 (Britton 1964*); n = ca. 33 (Cody and Mulligan
1982*).

Habitat: In gently sloping well-drained light sandy soils and around
rock piles in clearings, open woods, pastures, and old fields. It is a
weed of lowbush blueberry fields, rough pastures, and old fields in the
eastern part of its range.

Range: In North America, from southern Newfoundland (where it is
rare) and Nova Scotia to Ontario, south to Georgia and Arkansas.

Remarks: R.M. Tryon (1960) recognized 11 species of Dennstaedtia in
the Americas, which, with the exception of D. punctilobala, are
essentially tropical in range. A review of D. punctilobula in Canada
was published in the series The Biology of Canadian Weeds (Cody et
al. 1977).

2. Pteridium Scop, bracken

1. Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw.

P. latiusculum (Desv.) Hieron.

P. aquilinum (L.) Kuhn var. champlainense Boivin

bracken
Fig. 73 (a) frond; (6) fertile pinnule; (c) venation. Map 71.

Coarse fronds 30-70 cm or more long, in extensive colonies from
creeping and forking hairy rhizomes. Stipes longer or shorter than the
blades. Blades triangular, usually ternate, 30-50 cm wide, bipinnate-
pinnatifid to tripinnate-pinnatifid; lower pinnules more or less
pinnatifid; upper pinnules entire; ultimate divisions very numerous,
oblong to linear, glabrous or slightly pubescent beneath, with revolute
margins. Sporangia borne in marginal sori on the undersurface of the
pinnules; sporangia covered by a mostly continuous false outer
indusium formed by the revolute margin and by a minute often nearly
obsolete hyaline inner indusium.

Plants that grow in the shade tend to have ternate fronds, with
the rachis bent so that the blade is presented to the available light,
whereas plants that grow in the open tend to be upright and stiff, with
shorter ascending pinnae that are twisted at right angles to the rachis.

135

Fig. 73 Pteridium aquilinum var. latiusculum; (a) frond, 1/4 x ; (b) fertile pinnule,
3 X ; (c) venation, 3 x .

136

Cytology: n = 52 (Britton 1953*, Cody and Mulligan 1982*).

Habitat: Occurs as a weed in pastures, on grassy slopes in abandoned
fields, in burnt-over areas, in damp or more often dry, usually sterile
soil, on open slopes, and in open woods and thickets. This is probably
the most common Canadian fern.

Range: Worldwide distribution; var. latiusculum widespread in
Canada from Newfoundland to eastern Manitoba and rare in eastern
British Columbia and the foothills of the Rocky Mountains in Alberta.
To the south, extending to North Carolina, Tennessee, Missouri, and
Oklahoma.

1.1 Pteridium aquilinum (L.) Kuhn var. pubescens Underw.
P. aquilinum (L.) Kuhn var. lanuginosum (Bong.) Fern.
Fig. 74, undersurface of pinnule. Map 72.

Fronds usually longer than var. latiusculum, 1.5 m long or
longer. Blade broadly triangular, but rarely ternate. Lower surface of
pinnules more or less densely villous or villous-puberulent. Inner
indusium ciliate and sometimes also pubescent.

Cytology: n = 62 (Love et al. 1971).

Habitat: Moist to dry woods, clearings, open slopes, and roadsides.

Range: British Columbia and southwestern Alberta, south through
Washington and Oregon to California and Utah.

Remarks: R.M. Tryon (1941) treated the worldwide genus Pteridium
as comprising a single species, P. aquilinum, made up of two
subspecies, each with several varieties. Australian botanists,
however, now treat their plant as a separate species, P. esculentum
(Forst.) Diels, and similarly South American botanists treat their
plant as P. arachnoideum (Kaulf.) Maxon.

Material from the Bruce Peninsula, Ont., which R.M. Tryon
(1941) called var. pubescens, was later included by Boivin (1952) in his
var. champlainense. Because of the great variation in frond shape and
indument in the eastern var. latiusculum, dependent largely on
shading, we have included this material in var. latiusculum. A review
of Pteridium aquilinum in Canada was published in the series The
Biology of Canadian Weeds (Cody and Crompton 1975).

In early spring the young fronds o( Pteridium are highly valued
as a green vegetable, particularly in Japan. However, recent studies
have shown that this species is carcinogenic in rats, is responsible for
enzootic bovine hematuria in cattle, and causes vitamin Bi
avitaminosis in horses and other nonruminants. It is therefore not
recommended for human consumption.

137

3. CheilanthesSw. lip fern

Small evergreen ferns of dry rocky regions. Rhizomes short and
much branched to longer and simple, bearing numerous slender,
brown to blackish, hyaline scarious-margined scales. Segments small
and bead-like. Sori marginal, often confluent, covered by the inrolled
margin of the pinnule.

This is a genus comprising over 100 species, widely distributed
around the world in mainly arid regions. The two species that occur in
Alberta and British Columbia are readily distinguishable from other
ferns in the region by the small bead-like segments of their pinnae.

A. Blade tomentose below, thinly villous above; lacking scales ....

I. C feci

A. Blade villous and scaly below, glabrous above

2. C gracillima

1. Cheilanthesfeei Moore

slender lip fern
Fig. 75 (a) fronds; (6) upper surface of pinnule; (c) lower surface of
pinnule. Map 73.

Fronds 5-20 cm long, tufted from a short and much branched
rhizome. Stipes 3-10 cm long, dark purplish brown, with a few
scarious-margined scales at the base and tawny multicellular hairs
above. Blades 2-10 cm long, linear-oblong to ovate, tripinnate.
Pinnae deltoid to ovate-oblong; rachis and lower side densely
tomentose with pale brown hairs; upper surface with soft whitish
hairs; ultimate segments small and rounded. Margins of segments
somewhat inrolled but not covering the mature sporangia, which
cover the entire lower surface.

Cheilanthes feei can be distinguished from C. gracillima by the
absence of scales on the lower surface and thinly villous upper surface
of the pinnules.

Cytology: 2n = 87 (Knobloch 1967). An apogamous triploid species.

Habitat: Crevices of limestone or calcareous cliffs.

Range: Southern British Columbia and southwestern Alberta, south
and east in the United States to California and Illinois.

Remarks: In British Columbia this species is restricted to two
limestone regions, where it is quite common. It is rare in the
mountains of western Alberta (Argus and White 1978).

138

Fig. 74 Pteridium aquilinum var. pubescens; undersurface of pinnule, 3 x .

Fig. 75 Cheilanthes feei; (a) fronds, 4/5 x; (b) upper surface of pinnules, 1/3 x;
(c) lower surface of pinnules, 4 x .

139

2. Cheilanthes gracillima DC. Eat.

lace fern
Fig. 76 (a) fronds; (6) sterile pinnule; (c) fertile pinnule. Map 74.

Fronds 5-25 cm long or longer, densely tufted from the short,
much branched rhizome. Stipes dark brown, glabrous, or with
scattered long hairs and narrow scales. Blades bipinnate, linear to
oblong-lanceolate, 3-10 cm long; rachis and costae frequently villous-
puberulent, with long narrow brown scales. Pinnae lance-oblong;
pinnules oblong to oval, woolly below, green and glabrous or with a
few stellate hairs above, and with broadly inrolled margins. Mature
sporangia covering the entire lower surface and extruding from
between the folded margins.

Cheilanthes gracillima may be distinguished from C feei by the
scaly undersurface and glabrous upper surface of the pinnules.

Cytology: None.

Habitat: Dry crevices of cliffs and rocky slopes of usually igneous
rocks.

Range: Southern British Columbia and southwestern Alberta to
California, eastward to Idaho, Montana, and Nevada.

Remarks: In British Columbia C. gracillima is found across the
extreme southern part of the province. In Alberta it is known only in
Waterton Lakes National Park, where it is very rare (Argus and
White 1978).

4. Aspidotis (Nutt. ex Hook. & Bak.) Copel.

1. Aspidotis densa (Brack.) Lellinger

Cheilanthes siliquosa Maxon

C. densa (Brack.) St. John

Pellaea densa (Brack.) Hook.

Cryptogramma densa (Brack.) Die Is

Indian's-dream
Fig. 77 (a) fertile and sterile fronds; (b) fertile pinnule. Map 75.

Fronds dimorphic, 30 cm long or longer, densely tufted from a
short, much branched, chestnut-brown, scaly rhizome. Stipes wiry,
chestnut brown, lustrous, glabrous, much longer than the blades.
Fertile blades tripinnate, 2-6 cm long, 1-4 cm wide, broadly ovate;
pinnules narrowly linear, mucronate. Indusium marginal, thin,
continuous, erose-denticulate. Sterile fronds usually much shorter
than the fertile (fewer and often lacking), with segments smaller,
somewhat broader, and sharply toothed or incised.

140

Fig. 76 Cheilanthes gracillima; (a) fronds, 3/4 x ; (b) sterile pinnule, 3 x ; (c) fertile
pinnule, 3 x .

141

Fig. 77 Aspidotis densa; (a) fertile and sterile fronds, 3/4 x ; (b) fertile pinnule,
lOx.

142

This small fern can usually be separated from species in the
genera Cheilanthes, Pellaea, and Cryptogramma, in which it has been
included in the past, by its tripinnate fertile fronds with narrowly
linear mucronate pinnules and usually by the absence of sterile
fronds.

Cytology: n = 30 (W.H. Wagner 1963; Smith 1975).

Habitat: Exposed cliff crevices and rocky or talus slopes.

Range: Southern British Columbia, south and east to Montana,
Wyoming, Utah, and California; disjunct to Gaspe, Megantic, and
Wolfe counties of Quebec, where it is rare. A specimen collected by
H.M. Ami labeled "rocky hillsides of Guelph dolomite, Durham, Ont.
(CAN)" may represent a mislabeling because the species is not
otherwise known from Ontario, although it has been searched for by
numerous fern enthusiasts in the vicinity of Durham.

Remarks: As noted above, this species has been unsatisfactorily
placed at various times in the genera Pellaea, Cheilanthes, and
Cryptogramma. A recent study by Lellinger (1968) has placed it in the
genus Aspidotis, together with two other western American species
and one from Africa.

5. Pellaea Link cliff-brake

Small tufted plants from compact rootstocks. Fronds firm; stipes
and rachises wiry; pinnae gray green; veins free. Sori marginal and
confluent under the inrolled and altered margin of the fertile
pinnules.

The genus Pellaea section Pellaea was treated in a monograph by
A.F. Tryon in 1957, and cytotaxonomic studies were reported by A.F.
Tryon and Britton (1958). The genus is medium-sized and comprises
about 80 species, but has only 16 species in the section Pellaea. The
basic chromosome number (x) is 29, and both sexual and apogamous
species and varieties are known. The centre of diversity for the section
Pellaea is in the southwestern United States and Mexico, and
Canadian species are definitely outliers from the centres of
distribution. We have only two species, P. atropurpurea and
P. glabella, although the latter has two varieties (some would main-
tain that we have four distinct species — see the Introduction). All
Canadian plants have a similar appearance, a rather gray green color
that blends well with the limestone rock crevices and ledges with
which they are associated. The apogamous development of some of the
species is an adaptation to xerophytic habitats (A.F. Tryon 1968).

143

A. Fronds dimorphic; stipe and rachis scurfy, with appressed
pubescence 1. P. atropurpurea

A. Fronds monomorphic; stipe and rachis glabrous or with a few
spreading hairs 2. P. glabella

1. Pellaea atropurpurea (L.) Link

purple cliff-brake
Fig. 78 (a) sterile and fertile fronds; (6) fertile pinnule. Map 76.

Fronds dimorphic; fertile frond 10-35 cm long, 3.5-8 cm wide,
longer than the sterile frond. Stipes and rachis dark purple brown,
dull, pubescent, with more or less appressed hairs. Pinnae rigid,
evergreen, bluish green, simple above, bipinnate below; fertile pinnae
linear to oblong or narrowly ovate, with the lower pinnules stalked;
sterile pinnules ovate-oblong. Sori situated around the margins of the
fertile pinnules. Inrolled margin of pinnule forming the indusium.

This species looks somewhat like P. glabella but may be
distinguished from it by the usually taller, more upright habit, with
fertile fronds that are more divided, that are darker blue green to olive
green, and that have markedly hairy stipes and rachis.

Cytology: "m" = 2n
apogamous triploid.

87 (Rigby 1973*). This species is an

Habitat: Dry, steep, exposed, limestone rock slopes or cliffs,
limestone paving, and tops of large talus boulders.

Range: Southern Quebec (Britton et al. 1967; Brunton 1972; Brunton
and Lafontaine 1974), southern Ontario (Britton and Rigby 1968;
Soper 1963), Lake Athabaska, southwestern Alberta (Brunton 1979),
and adjacent southeastern British Columbia, south to Florida and
Arizona. Distribution of Pellaea in Canada is given by Rigby and
Britton (1970). For the whole distribution of the species see A.F.
Tryon(1972).

Remarks: This species and P. glabella are often confused. Pellaea
atropurpurea is a rare plant in the Canadian flora and occurs at widely
separated locations. The Lake Athabasca station is amazingly distant
from the centre of distribution in the southwestern United States
(A.F. Tryon 1972). Pellaea atropurpurea is rare in Ontario (Argus and
White 1977), Saskatchewan (Maher et al. 1979), and Alberta (Argus
and White 1978).

144

f^ a

Fig. 78 Pellaea atropurpurea; (a) sterile and fertile fronds, 2/3 x ; (b) fertile pinna,
3x.

145

2. Pellaea glabella Mett. var glabella

P. atropurpurea (L.) Link var. bushii Mackenzie
smooth clifT-brake
Fig. 79 (a) fronds (6) fertile pinnule. Map 77.

Fronds similar, 10-25 cm long or longer, usually shorter than
those of P. atropurpurea, open and spreading out beyond the rock face.
Stipes and rachis dark reddish brown, smooth, and lustrous. Pinnae
rigid, evergreen, bluish green, simple above, pinnate below; basal
pinnae persistent, with stalk and rachis up to 5.0 cm long. Pinnules
sessile or nearly so, oblong-lanceolate. Sori situated around the
margins of the fertile pinnules; inrolled margin forming the indusium.
Spores 32 per sporangium.

The smooth cliff-brake is a distinctive species of high, steep
limestone cliffs. It appears from small, tight, crevices and blends well
with the background. Quite often there are no other ferns or
vegetation associated with it. Because the stipes are dark reddish
brown, smooth, and shiny, the species is sometimes misidentified as
purple cliff-brake.

Cytology: "n" =: 2m = 116 (Britton 1953, Rigby 1973*). This taxon is
an apogamous tetraploid, but see W.H. Wagner et al. (1965) for
Missouri.

Habitat: Crevices of dry, sometimes partly shaded, limestone cliffs.

Range: Southwestern Quebec and southern Ontario (Britton and
Rigby 1968; Soper 1963), south to Tennessee and Texas. (Rigby and
Britton 1970; Brunton and Lafontaine 1974).

Remarks: Some researchers consider the plant quite "unfernlike."
The tough, evergreen leaves with few divisions and distinctive
coloration are reminiscent of some of the Old World pteridophytes,
which also can invade the masonry efforts, castles, and stone walls.

2.1 Pellaea glabella Mett. ex Kuhn var. nana (Richards.) Cody
P. glabella Mett. ex Kuhn var. occidentalis (E. Nels.) Butters
P. occidentalis (E. Nels.) Rydb.
P. pumila Rydb.

Fig. 80 (a) fronds; (6) fertile pinnae. Map 78.

Differs from var. glabella by its thin, brittle, and golden brown
stipes, forming dense thickly clumped "pincushions" flush with the
rock face, and by its mitten-shaped sessile pinnae perpendicular to the
rachis. Spores 64 per sporangium.

The smaller size of the plants and their distribution are aids for
rapid identification.

146

Fig. 79 Pellaea glabella var. glabella; (a) fronds, 1 x ; (b) fertile pinna, 1 1/2 x

Fig. 80 Pellaea glabella var. nana; (a) fronds, 1 x ; (b) fertile pinnae, 1 1/2 x .

147

Cytology: n = 29 (A.F. Tryon and Britton 1958). This is a basic
sexual diploid taxon.

Habitat: Dry, exposed crevices of limestone cliffs.

Range: Manitoba to southwestern District of Mackenzie and
southwestern Alberta (Brunton 1979), south to Wyoming and South
Dakota.

Remarks: Some researchers maintain that this basic diploid taxon
should be given specific rank. That view stresses the differences
between var. nana and var. glabella. At the same time, var. nana has
many similarities to var. glabella, and they therefore must be closely
allied genetically. We have followed A.F. Tryon (1957) and accepted
the latter position. Variety nana is rare in the District of Mackenzie
(Cody 1979), Manitoba (White and Johnson 1980), and Saskatchewan
(Maheretal. 1979).

2.2 Pellaea glabella Mett. ex Kuhn var. simplex (E. Nels.) Butters
P. atropurpurea (L.) Link var. simplex (Butters) Morton
P. suksdorfiana Butters

Fig. 81 (a) fronds; {b) fertile pinnules. Map 79.

Differs from var. glabella by its usually withered basal pinnae
with stalk and rachis up to 1.0 cm long. Stipes sturdy, reddish brown
to brown purple, rarely sparsely pubescent; old stipes conspicuous and
often silver in color. Pinnae oblong-lanceolate, petioled, acute to the
rachis. Spores 32 per sporangium.

Cytology: m = 2n = 116 (A.F. Tryon and Britton 1958). An
apogamous tetraploid taxon.

Habitat: Crevices of shaded cool, east- or north-facing calcareous
cliffs, often overlooking water.

Range: Southwestern Alberta (Brunton 1979) and southern interior
British Columbia, south to Washington, Utah, Arizona, New Mexico,
and Colorado.

Remarks: Some researchers recognize var. simplex as P.
suksdorfiana. The remarks under P. glabella var. nana are pertinent
here. For var. simplex, however, the usual definition of a biological
species does not apply because it is apogamous.

148

Fig. 81 Pellaea glabella var. simplex; (a) fronds, 1 x ; (b) fertile pinna, 2 x

149

6. Cryptogramma R. Br.

Small rock ferns with dimorphic fronds, from short much
branched or slender elongate rhizomes. Blades glabrous, evergreen,
or deciduous; veins free. Sori marginal, covered by a continuous
indusium formed by the reflexed margin.

This is a small genus of widespread distribution in boreal and
alpine situations. The South American and Himalayan
representatives are closely related to C. crispa. They are found in
rocky habitats.

A. Fronds scattered on an elongate slender rhizome, deciduous ....

I.e. stelleri

A. Fronds densely tufted from a short much branched rhizome,

evergreen 2. C crispa

1. Cryptogramma stelleri (Gmel.) Prantl
slender cliff-brake

Fig. 82 (a) sterile and fertile fronds; (6) fertile pinnules. Map 80.

Fronds dimorphic, scattered along the horizontal rhizome.
Sterile fronds almost flaccid, 3-10 cm long; stipes pale to purplish;
blades ovate to ovate-deltoid, bipinnate; pinnules oblong, ovate, or
obovate flabelliform. Fertile fronds stiffer than sterile fronds, 9-21 cm
long; pinnules lanceolate to oblong. Sori situated around margins of
fertile pinnules; inrolled margin forming a false indusium.

The slender cliff-brake may be recognized by its delicate fronds,
which are scattered along an elongate slender rhizome. The fronds
may easily be overlooked, particularly as they turn brown later in the
season.

Cytology: n = 30 (Britton 1964*; Cody and Mulligan 1982*). This is
a basic diploid species.

Habitat: Moist, shaded, usually calcareous crevices and cliffs.

Range: Circumpolar; in North America from Alaska to
Newfoundland and Labrador, south into the northern United States,
but interrupted in distribution because of habitat limitations.

Remarks: This species is rare in the District of Mackenzie (Cody
1979) and Nova Scotia (Maher et al. 1978).

2. Cryptogramma crispa (L.) R. Br. var. acrostichoides (R. Br.)

C.B. Clarke
C acrostichoides R. Br.

150

Fig. 82 Cryptogramma stelleri; (a) sterile and fertile fronds, Ix; (b) fertile
pinnules, 3 x .

151

mountain-parsley; parsley fern
Fig. 83 (a) sterile and fertile fronds; (6) fertile pinnule. Map 81.

Fronds dimorphic, winter green, densely clustered from short-
creeping mostly ascending rhizomes. Sterile fronds up to 15 cm long;
stipes straw-colored; blades ovate to ovate-lanceolate, bipinnate-
pinnatifid; pinnae short-petioled; ultimate segments thick, ovate,
oblong, or obovate, obtuse, with crenate or toothed margins. Fertile
fronds standing stiffly above the sterile, with fewer linear-oblong
entire segments; margins of segments broadly reflexed often to the
midrib, but opening as the sporangia mature. Sori eventually
covering the surface of the fertile pinnule.

Mountain-parsley may be recognized by its densely bunched,
crisp, glabrous, winter green fronds, which usually stand out from the
dull, rocky substrate.

Cytology: n = 30 (R.L. Taylor and Mulligan 1968*; Cody and
Mulligan 1982*).

Habitat: Crevices, ledges, and talus slopes and in pockets of organic
soil in the Precambrian region.

Range: In western North America from southern Alaska, through
the mountains of British Columbia and western Alberta, south to
California and New Mexico, across the Precambrian Shield to
northwestern Ontario, northeastern Minnesota, and Isle Royale,
Mich. (Marquis and Voss 1981); also in Kamchatka.

2. 1 Cryptogramma crispa (L.) R. Br. var. sitchensis (Rupr.) C. Chr.
Fig. 84 (a) sterile and fertile fronds; (6) sterile pinnules. Map 82.

Similar to var. acrostichoides, but with the sterile fronds broadly
triangular, finely dissected; ultimate segments obovate.

Habitat: In the Mackenzie Mountains, N.W.T., on limestone talus
and moraines.

Range: Southern Alaska to the Mackenzie Mountains, south into
northern British Columbia.

Remarks: The Eurasian var. crispa is tetraploid. It tends to have
softer sterile leaves that are often more dissected, and the rhizome
scales are uniformly brown. Variety acrostichoides is rare in Ontario
(Argus and White 1977). A report by Macoun (1890) of its occurrence
at McLeod's Harbour on Manitoulin Island was refuted by Soper
(1963).

152

Fig. 83 Cryptogramma crispa var. acrostichoides; (a) sterile and fertile fronds,
2/3 X ; (b) fertile pinnule, 5 x .

153

Fig. 84 Cryptogramma crispa var. sitchensis; (a) sterile and fertile fronds, 1/2 x ;
(b) sterile pinnules, 5 x .

154

Calder and Taylor (1968) consider that variety sitchensis is a
"weak segregate hardly worthy of recognition," whereas others, e.g.
Love (in Love et aL 1977) treat var. acrostichoides as a species,
because its morphology and chromosome number are distinct from C.
crispa of Europe. It is true that var. acrostichoides is a basic diploid
entity, but we would like to see a thorough biosystematic study that
compares all three varieties before definite taxonomic conclusions are
reached.

Variety sitchensis is rare in the District of Mackenzie (Cody
1979).

7. Pityrogramma Link

1 . Pityrogramma triangularis ( Kaulf . ) Maxon

Gymnogramma triangularis Kaulf.

goldback fern
Fig. 85 (a) fronds; (6) portion of undersurface of pinna. Map 83.

Fronds to 30 cm long or longer, tufted from the thickish some-
what ascending rhizome. Stipes much longer than the blade, stiff and
wiry, lustrous dark brown, glabrous except at the base. Blade deltoid,
pinnate; lowest pinnae pinnate, with the two lower first pinnules
longer than the rest; the remaining pinnae pinnatifid; segments blunt,
coriaceous; margins narrowly revolute; upper surface glabrous; lower
surface with a white or yellowish waxy powder. Sporangia borne
along the veins, confluent in age. Indusium lacking.

The white or yellowish waxy powder on the undersurface of the
deltoid blades is characteristic.

Cytology: n = 30, 60 (Alt and Grant 1960).

Habitat: Open to partly shaded rocky slopes and crevices.

Range: Southwestern coastal British Columbia, south to California,
and inland to Arizona, Nevada, and Utah. This very distinctive
species seems to be restricted to the floristic region in Canada where
madrona (Arbutus), Canada's only broad-leaved evergreen tree, grows
(see also Isoetes nuttallii).

Remarks: Pityrogramma triangularis belongs to a genus of about 15
species of small, mainly tropical ferns. Alt and Grant (1960) have
shown that there are both diploids and tetraploids of P. triangularis
from north to south in California, with the tetraploid occurring near
the coast. No discernible characters with which the two races can be
separated have as yet been detected. Chromosome counts have
apparently not been made on plants from north of California, and so
we have no knowledge of which race occurs in British Columbia.

155

Fig. 85 Pityrogramma triangularis; (a) fronds, 2/3 x ; (b) portion of undersurface of
pinna, 5 x .

156

According to T.M.C. Taylor (1970), the fronds curl up in dry
weather, showing their characteristic light-colored undersurface,
which has a whitish or yellowish powder on it.

8. Adiantum L. maidenhair fern

Delicate ferns. Fronds produced in rows from slender creeping
rhizomes. Veins free-forking. Sori oblong, borne along the upper
margin of the pinnules; each sorus covered by an indusium that arises
from the inrolled margin.

The genus Adiantum is worldwide in distribution and numbers
over 200 species. Both our species have been used in horticulture, and
in Canada, A. pedatum does well in shaded spots in the fern lover's
garden.

A. Frond with a simple main zigzag rachis continuing the arching to
pendulous stipe 1.4. capillus-veneris

A. Frond palmately forking at the summit of the upright stipe

2. A. pedatum

1 . Adiantum capillus- veneris L.

Venus'-hair fern
Fig. 86 (a) fronds; (b) pinnules with sporangia. Map 84.

Fronds to 40 cm long or longer, often pendulous from a slender
elongate rhizome. Stipe lustrous blackish brown, continuing into a
zigzag rachis. Blade ovate-lanceolate, 2-3 pinnate at the base, to
simply pinnate above. Pinnules rhombic-ovate with irregularly
jagged lobes; veins flabellate, forking from the base. Sori oblong to
lunate on the outer margins of the pinnules, which are inrolled to form
an indusium.

The zigzag rachis, which is simply pinnate above, readily
separates this species from the maidenhair fern, A. pedatum.

Cytology: m = 30 (Britton 1953).

Habitat: Runnelsof hot springs.

Range: Circumpolar: Primarily in warm temperate regions on wet
cliffs and seeps, extending northward to Virginia, Missouri, Colorado,
Utah, and California, and introduced farther north in sewers and as a
weed in greenhouses. In Canada known only from Fairmont Hot
Springs in British Columbia, where it is in danger of being extirpated.

Remarks: This species is rare and endangered in British Columbia.

157

Fig. 86 Adiantum capillus-veneris; (a) fronds, 2/3 x; (b) pinnules with sporangia,
5x.

158

2. Adiantum pedatum L. ssp. pedatum

maidenhair fern
Fig. 87 (a) frond; (6) pinnule with sporangia. Map 85.

Fronds 30-55 cm long, in colonies arising from horizontal
rhizomes. Stipes lustrous purple brown, forking at the summit into
two arching rachises, each of which is divided several times into
spreading divisions, thus forming a semicircular blade 15-35 cm wide
or wider. Pinnules short-stalked, obliquely triangular oblong;
terminal pinnule fan-shaped; main vein along the lower margin;
upper margin cleft, with lobes thus formed blunt. Sori elongate, borne
on the upper margins of the lobes of the pinnules. Indusium formed by
the inrolled margin.

The usually arching and palmately divided lustrous purple
brown rachises and the fan-shaped pinnules with the main vein along
the lower margin set this fern apart from all others in Canada.

Cytology: n = 29 (Britton 1953*, Cody and Mulligan 1982*).

Habitat: Wooded, sometimes rocky slopes in humus-rich soil.

Range: Nova Scotia to Ontario, south in the United States to
Georgia, Alabama, Mississippi, Louisiana, and Oklahoma.

2. 1 Adiantum pedatum L. ssp. aleuticum (Rupr.) Calder & Taylor

A. pedatum L. var. rangiferinum Burgess
Fig. 88 (a) frond; (6) pinnule with sporangia. Map 86.

Differs from ssp. pedatum in the branches strongly ascending
rather than widely divergent. Compared with ssp. pedatum, pinnae
usually fewer and pinnules (10) 12-20 (23) mm long, more deeply
lobed, with their tips acute rather than rounded and with the sinuses
between the lobes usually broader.

Cytology: n = 29 (R.L. Taylor and Mulligan 1968*).

Habitat: In usually shaded humus-rich soil on ledges and in rocky
woods from sea level to the treeline.

Range: Western Alberta, British Columbia, and Alaska, south in the
United States to California.

2.2 Adiantum pedatum L. var. subpumilum W.H Wagner
Fig. 89, frond. Map 87.

159

Fig. 87 Adiantum pedatum ssp. pedatum; (a) frond, 1/3 x; (b) pinnule with
sporangia, 3x.

160

Fig. 88 Adiantum pedatum ssp. aleuticum; (a) frond, 1/3 x; (b) pinnule with
sporangia, 5 x .

161

A dwarf variety that differs in its smaller stature and smaller
imbricate pinnules. Pinnules with fewer vein forkings; vein forkings
occurring in the distal parts of the pinnules.

Cytology: n = 29 (Wagner and Boydston 1978).

Habitat: Wet, exposed cliffs on exposed metamorphic coastal rocks.

Range: In nature known only in the Brooks Peninsula, northwest
Vancouver Island, B.C. (type locality); also known in cultivation but
from unknown sources (W.H. Wagner and Boydston 1978).

2.3 Adiantum pedatum L. ssp. calderi Cody

Fig. 90, frond. Map 88.

Differs from ssp. aleuticum, with which it has been associated, by
its generally shorter stature, stiffly crowded stipes, bluish green
glaucus fronds, consistently smaller pinnules (middle pinnules 7-12
(17) mm long), and conspicuous indusia.

Cytology: n = 29 (Cody and Mulligan 1982*).

Habitat: Serpentine and dolomite talus slopes, tablelands, and rocky
woods.

Range: Western Newfoundland, Gaspe Peninsula, Eastern
Townships of Quebec, and adjacent northern Vermont, and disjunct to
serpentines in northern California and Washington (Cody 1982).

Remarks: A large form with deeply lacerate pinnules up to 2.5 cm
long was described from British Columbia as var. rangiferinum by
Burgess (1886). This has been included in the synonymy of ssp.
aleuticum.

162

Fig. 89 Adiantum pedatum var. subpumilum; frond, 2/3 x

163

Fig. 90 Adiantum pedatum ssp. calderi; fronds, 1/2 x

164

10. ASPIDIACEAE

Small to large ferns. Fronds pinnate to decompound, forming a
crown at the top of a stout rhizome or singly along a fine rhizome. Sori
dorsal, usually roundish, but sometimes somewhat elongate.
Indusium opening on one side, peltate or absent.

Aspidiaceae is a large family of mainly tropical and subtropical
terrestrial ferns comprising over 60 genera.

A. Sporangia partly or wholly covered by the rolled-up pinnules;
pinnules forming globular berry-like divisions of the stiff fertile
frond.
B. Fronds in vase-like clumps; simple pinnate fertile fronds

surrounded by tall regularly pinnate sterile ones

1. Matteuccia

B. Fronds solitary or scattered along the rhizome; sterile

fronds coarsely pinnatifid; fertile fronds bipinnate

2. Onoclea

A. Sporangia not in hard rolled-up berry-like divisions.

C. Sori round or nearly so.
D. Indusia present.

E. Indusia segmented 3. Woodsia

E. Indusia not segmented.

F. Indusium hood-shaped, attached by its base

on the side toward the midrib

9. Cystopteris

F. Indusium round, reniform, or elongate.

G. Fronds scattered along a thin cord-like
rhizome (or tufted from a stout

rhizome in T. limbosperma)

7. Thelypteris

G. Fronds tufted or forming a crown at
the end of a stout rhizome.
H. Sori elongate, often curved over
the ends of the veins; indusia

attached on one side

10. Athyrium

H. Sori round.

I. Indusia reniform or with a
deep sinus.
J. Veins reaching the

margin

7. Thelypteris

limbosperma
J. Veins not reaching the

margin

5. Dryopteris

165

I. Indusia round, without a

deep sinus

4. Polystichum

D. Indusia absent.

K. Fronds in a crown at the end of a stout rhizome ..

10. Athyrium

K. Fronds singly along a cord-like rhizome.

L. Fronds more or less ternate

6. Gymnocarpium

L. Fronds pinnate-pinnatifid

8. Phegopteris

C. Sori elongate or horseshoe-shaped 10. Athyrium

1. A/fa tte licc/a Todaro

1. Matteuccia struthiopteris (L.) Todaro var. pensylvanica (Willd.)
Morton
Pteretis pensylvanica (Willd.) Fern.
P. nodulosa (Michx.) Nieuwl.
Onoclea struthiopteris (L.) Hoffm. var. pensylvanica (Willd.)

Boivin
ostrich fern
Fig. 91 (a) upper portion of sterile frond; (b) fertile frond; (c) portion of
fertile pinna. Map 89.

Fronds dimorphic, forming a crown at the end of the stout widely
creeping and forking rhizome. Sterile fronds up to 1 .2 m long or longer
and 12-24 cm wide, pinnate-pinnatifid, abruptly narrowed to the base;
pinnae broadly linear, acuminate; pinnules oblong, bluntish. Fertile
fronds much shorter than the sterile, persistent over winter; pinnae
greenish, becoming dark brown at maturity. Veins free, not forked, on
both sterile and fertile pinnae. Sori borne on the margins of the
shallowly lobed, tightly inrolled, and pod-like pinnae.

A full crown of fronds is somewhat reminiscent of a large
headdress. Sterile fronds might be mistaken for Osmunda
cinnamomea, but can readily be distinguished from it by the shape of
the blade, which tapers to the base and is sharply cut off at the tip.

Cytology: n = 40 (Britton 1953*).

Habitat: Damp shady places, roadside ditches, and floodplains of
streams and rivers.

Range: Matteuccia struthiopteris s.l. circumpolar; var. pensylvanica
from Newfoundland to British Columbia, southwestern District of
Mackenzie, southeastern Yukon Territory and Alaska, south to
Virginia, Ohio, Indiana, Illinois, Missouri, and South Dakota.

166

Fig. 91 Matteuccia struthiopteris var. pensylvanica; (a) upper portion of sterile
frond, 1/3 x ; (b) fertile frond, 1/3 x ; (c) portion of fertile pinna, 2 x .

167

Remarks: Morton (1950) quotes E.T. Wherry as calling the ostrich
fern "our most renamed fern." It has been variously placed in Onoclea,
Struthiopteris, Pteretis, and Matteuccia. Fernald (1945) maintained
that the North American fern was specifically distinct from the
European plant. The differences, as pointed out by Morton (1950) are
slight, and the varietal level is more satisfactory. The young
fiddleheads of this fern are used as a vegetable, particularly in the
Maritime Provinces. They may be prepared fresh, or preserved by
either freezing or canning. Cruise (1972) reported that the removal of
young fronds of the ostrich fern seemed to cause spore-bearing fronds
to appear earlier and in greater numbers. The species is common in
eastern Canada but rare west of Manitoba. It is rare in the Yukon
(Douglas etal. 1981).

2. On odea L.

1. Onoclea sens i bills L.

sensitive fern
Fig. 92 (a) sterile and fertile fronds; (b) portion of fertile pinna.
Map 90.

Fronds dimorphic, borne several together on slender creeping
rhizomes. Sterile fronds up to 80 cm long; blades 12-30 cm long, 15-
30 cm wide, broadest at the base, pinnate at the base, pinnatifid above;
rachis winged, with the wing becoming broader toward the tip; pinnae
wavy-margined or coarsely toothed. Fertile fronds persistent over the
winter, shorter than the sterile; pinnules greenish, becoming blackish
at maturity, modified and inrolled to form berry-like structures.
Veins free on the fertile fronds and netted on the sterile fronds. Sori
borne within the tightly inrolled, berry-like pinnules.

Forma obtusilobata (Schkukr) Gilbert has intermediate fronds
between the normal fertile phase and the normal sterile phase; it
occurs with the typical form and may be the result of damage to young
fronds.

Where Woodwardia areolata occurs in Nova Scotia, it might
possibly be confused with the sensitive fern. The latter can, however,
be readily distinguished by its entire rather than minutely serrate
pinnae margins and by its basal pinnae, which are subopposite rather
than alternate.

Cytology: n = SI (Cody and Mulligan 1982*).

Habitat: Forms large patches in low places in woodlands, wet
meadows, and roadside ditches. In ditches and meadows it often
reaches proportions that make it an undesirable weed.

168

Fig. 92 Onoclea sensibilis; (a) sterile and fertile fronds, 1/3 x ; (b) portion of fertile
pinna, 2 x .

169

Range: Eastern North America, southern Labrador and
Newfoundland to southeastern Manitoba, south to Florida, Louisiana,
and Texas; also in eastern Asia.

Remarks: The common name, sensitive fern, is derived from the fact
that the fronds, although coarse, are sensitive and blacken when
touched by the first frost.

3. Woodsia R. Br. woodsia

Small tufted ferns with free veins arising from compact
rootstocks. Indusium of thread-like or plate-like segments, attached
below, and more or less arched over the round sori.

The genus Woodsia is a medium-sized genus of perhaps 40
species in the world and was treated in a monograph by Brown (1964).
In Canada there are only five, six, or seven species, depending on
which authority you consult. Amateurs find the species difUcult to
recognize because some plants superficially resemble the abundant
Cystopteris fragilis s.l. (fragile fern), and many keys stress soral
characters such as the indusium, which in mature specimens may be
lost or damaged. To distinguish Woodsia from Cystopteris one should
check to see if the indusium is attached below the sorus (Woodsia) or is
hooded (Cystopteris), i.e., attached at one side and arching over the
sorus. Failing this, one can compare the stipes by holding them up to
the light in the field. They are opaque in Woodsia and translucent in
Cystopteris. The venation is useful also. In the former the veins are
less distinct and appear to stop short of the margin, whereas in the
latter, they are distinct right to the margin. Also, in Woodsia^ one
expects to find either even or uneven stubble from the remains of old
stipe bases.

The species in the genus fall naturally into two groups (see key
under A). In the first group, there is an articulation point towards the
base of the stipe, and when old fronds drop off, an even stubble is left.
The other group lacks these joints and is the uneven stubble group.

We have three species in the first group: two basic diploids, W.
glabella and W. iluensis, and a derived tetraploid, W. alpina.

The second group consists of two basic ancestral diploids that are
abundant in Western Canada, W. scopulina and W. oregana, and two
tetraploids, W. oregana var. cathcartiana and W. obtusa. The last
occurs fairly frequently southeast of Canada but has very few stations
in Canada. In this respect, it is analogous to Asplenium platyneuron.

There are also a number of hybrids known, both within each
group and between the groups, but on the whole they have not been
studied extensively with the use of modern experimental methods.

A. Stipes jointed at the base, with persistent bases appearing about
the same length.

170

B. Fronds delicate, glabrous; stipes and rachises green or

stramineous; rachises chaffless 1. W. glabella

B. Fronds more or less firm; stipes and rachises brown;
rachises chaffy, at least towards the base.
C. Fronds hairy and usually chaffy below; stipes usually

very chaffy 2. W. ilvensis

C. Fronds glabrous or glabrate, chaffless; stipes chaffless

or with a few deciduous scales 3. W. alpina

A. Stipes not jointed at the base, with persistent broken bases of
various lengths.
D. Pinnae and rachis bearing glands (which often stain drying

papers yellow) and white articulate hairs

5. W. scopulina

D. Pinnae and rachis with or without a fine glandular
pubescence and lacking white articulate hairs.
E. Pinnules broadly rounded; indusia of a few broad

segments 6. W. obtusa

E. Pinnules slightly lobed or finely toothed; indusia of
narrow and thread-like segments 4. W. oregana

1 . Woodsia glabella R. Br.

smooth woodsia
Fig. 93 (a) fronds; (6) fertile pinnae. Map 91.

Fronds to 16 cm long or longer, 1.5 cm wide, linear to linear-
lanceolate; pinnae thin-membranous, suborbicular to ovate, toothed or
lobed, glabrous. Stipes jointed near the base, usually with chaff only
below the joint. Sori distinct or confluent. Indusia composed of 5-8
ciliate-like segments.

This small and attractive species, with its green rachis and stipe,
is not likely to be confused with the other species of Woodsia. It is,
however, sometimes confused with Asplenium uiride (see under that
species).

Cytology: n = 39 (Britton 1964*; Cody and Mulligan 1982*); n = 38
(Love and Love 1976*).

Habitat: In moss or humus among rocks or in protected, cool, moist
calcareous crevices.

Range: Circumpolar; in North America from Greenland and
Newfoundland to Alaska south to the Gaspe Peninsula, Que., New
York, Ontario, Minnesota, and northern British Columbia.

Remarks: This basic diploid ancestral species would seem to have the
same chromosome number here as in Europe. It is rare in Manitoba
(White and Johnson 1980), Nova Scotia (Maher et al. 1978), Ontario
(Argus and White 1977), Saskatchewan (Maher et al. 1979), and
Alberta (Argus and White 1978).

171

Fig. 93 Woodsia glabella; (a) fronds, 1 x ; (b) fertile pinnae, 5 x

172

2 . Woods i a ilvensis ( L . ) R . B r .

rusty woodsia
Fig. 94 (a) fronds; (6) fertile pinna; (c) sorus. Map 92.

Fronds 5-25 cm long or longer, 2-3 cm wide, oblong-lanceolate,
pinnate-pinnatifid to bipinnate; pinnae oblong-lanceolate; margins of
the segments crenate and usually somewhat inrolled. Stipes jointed,
with the old stipe-bases persistent. Rachis and undersurface of the
blade usually brown-chaffy. Sori round, numerous, and close together
on the undersurface. Indusia of up to 20 long ciliate-like segments.

The rusty woodsia is noted for being both scaly and glandular. It
is one of the most abundant ferns on the cliffs and talus slopes north of
Lake Superior. In eastern Canada, it is certainly the species that the
amateur is likely to see first in large numbers. The species is so rare
in Great Britain that almost every plant known has been tabulated.

Cytology: n = 41 (Cody and Mulligan 1982*).

Habitat: Dry, often exposed, usually acid rocks and crevices of cliff
faces.

Range: Circumpolar; in North America from Greenland to Alaska,
south to North Carolina, Michigan, Illinois, Banff, Alta., and central
British Columbia.

Remarks: The species is quite variable in size, form, and degree of
chafliness. At times, plants that grow in the shade look quite unlike
those from exposed sites. It is rare in the Yukon (Douglas et al. 1981)
and Alberta (Argus and White 1978).

3. Woodsia alpina (Bolton) S.F. Gray

W. bellii (Lawson) A.E. Porsild

northern woodsia
Fig. 95 (a) fronds; (6) fertile pinna; (c) sorus. Map 93.

Fronds up to 15 cm long, 0.5-2.5 cm broad, linear to oblong-
lanceolate; pinnae suborbicular to oblong or lanceolate, crenate to
pinnatifid, flat, glabrous, and with no chaff. Stipes without chaff or
somewhat scaly, jointed near the base. Sori separate or confluent and
occurring near the margins. Indusia of ciliated plate-like lobes.

Woodsia alpina is considered to be the derived allotetraploid
from a cross of W. glabella X ilvensis. It is a variable species that
looks like a more robust W. glabella with shining brown to purple-
colored stipes and thicker blades. The middle pinnae are 2- to 3-lobed,
whereas in W. ilvensis they are 3- to 6-lobed.

Cytology: n = 82 (Love and Love 1976*).

173

Fig. 94 Woodsia ilvensis; (a) fronds, 2/3 x ; (b) fertile pinna, 4 x ; (c) sorus, 10 x

174

Fig. 95 Woodsia alpina; (a) fronds, 1 x ; (b) fertile pinna, 4 x ; (c) sorus, 10 x .

175

Habitat: Rock crevices and rock screes, usually on calcareous or
nonacid rocks.

Range: Circumpolar; in North America from Greenland to Alaska,
south to New York, Ontario, Michigan, and Minnesota. The known
distribution was extended to southern Ontario by Catling (1975).

Remarks: Lawson (1864) described some material from the Gaspe as
varietally distinct from W. alpina in Scotland. Porsild (1945) raised
this variety to specific rank as W. bellii. We are following Brown
(1964), who followed R.M. Tryon (1948), in not recognizing these
plants (which have less chaffy, more delicate, larger fronds) other than
to indicate that they seem to be expressions of a less rigorous climate
in the south. The situation is analogous to that of var. remotiuscula of
Dryopteris fragrans. Woodsia alpina is rare in Manitoba (White and
Johnson 1980), Nova Scotia (Maher et al. 1978), and Ontario (Argus
and White 1977).

4. Woodsia oregana D.C. Eat.

Oregon woodsia
Fig. 96 (a) fronds; (6) fertile pinna; (c) sorus. Map 94.

Fronds 10-30 cm long or longer, 1.0-3.5 cm wide. Blades linear-
lanceolate. Pinnae opposite, remote, triangular-oblong. Pinnules
oblong, blunt, with marginal crenulate-serrate teeth often inrolled.
Stipes not jointed. Rachis dark brown at the base, becoming straw-
colored above, glabrous or somewhat finely glandular, usually without
scales. Sori round, medial. Indusia of narrow and threadlike
segments.

This western species looks somewhat like W. iluensis but is
usually without scales and is a characteristic plant of calcareous
rather than more acid substrates. Unlike W. iluensis it belongs to the
group with uneven stubble.

Cytology: n = 38 (Brown 1964; Cody and Mulligan 1982*). Basic
diploid species.

Habitat: More or less protected crevices of calcareous ledges and
cliffs.

Range: Gaspe, Que., Ottawa District, Algonquin Park, upper Great
Lakes, Alberta and British Columbia, south to Oklahoma and New
Mexico.

176

Fig. 96 Woodsia oregana; (a) fronds, 1 x ; (b) fertile pinna, 5 x ; (c) sorus, 10 x

177

Remarks: Brown (1964) believed that the tetraploid taxon
cathcartiana had a very small distribution in only two counties along
the St. Croix River in Minnesota and Wisconsin, so that W. oregana
var. oregana extended from British Columbia to the Gaspe. Our
interpretation is quite different. We know the material in Canada
from Manitoulin Island belongs to the taxon cathcartiana. Also,
Ontario material is highly glandular and was at one time referred to
W. pusilla var. cathcartiana (T.M.C. Taylor 1947). The few collections
from the Prairie Provinces (Cody and Lafontaine 1975) and the
material from eastern Canada are in need of further study. We prefer
at this time to restrict the name W. oregana var. oregana to the
western diploid, and var. cathcartiana (Robins.) Morton to the
tetraploid, which is highly glandular. Plants in the Great Lakes
region, and presumably those farther east, belong to var. cathcartiana,
but this should be investigated. Woodsia oregana is rare in Manitoba
(White and Johnson 1980), Ontario (Argus and White 1977), and
Saskatchewan (Maheretal. 1979).

5. Woodsia scopulina DC. Eat.

W. oregana D.C. Eat. var. lyallii (Hook.) Boivin

W. appalachiana T.M.C. Taylor

Rocky Mountain woodsia
Fig. 97 (a) fronds; (6) fertile pinna. Map 95.

Fronds to 40 cm long or longer, 1.5-8 cm wide. Pinnae oblong-
lanceolate to ovate; pinnules oblong, denticulate; rachis and blade
with scattered white multicellular hairs mixed with the glandular
pubescence (which often stains drying papers yellow). Sori round,
near the margins. Indusia composed of flat plate-like segments mostly
hidden under the sori.

When it is in suitable sites, the Rocky Mountain woodsia is
usually a larger plant than the Oregon woodsia. The nonjointed,
shiny, chestnut-colored stipes are characteristic. On the rachis and
blade are prominent, white, articulated scales, which are diagnostic.

Cytology: n = 38 (R.L. Taylor and Brockman 1966*; Cody and
Mulligan 1982*).

Habitat: Among rocks and in crevices of cliffs (usually calcareous).

Range: Gaspe County, Que., Algonquin Park and Thunder Bay Dis-
trict, Ont., western Alberta, British Columbia, southern Yukon and
Alaska, south to Arkansas, Tennessee, California, and New Mexico.

178

Fig. 97 Woodsia scopulina; (a) fronds, 1/2 x ; (b) fertile pinnule, 3 x

179

Remarks: T.M.C. Taylor (1947) described the plants in the
Appalachians as W. appalachiana and noted that the indusial
segments were broader and the rhizome scales narrower than those in
typical W. scopulina. We are following Brown (1964) in not
recognizing this species, although we know of no recent
comprehensive study that compares the eastern Canadian plants with
those in the Appalachians, and in turn with the western plants. Most
authorities consider W. scopulina to be a very distinctive species
(Brown 1964). Boivin (1966) referred it to a variety of W. oregana.
Woodsia scopulina is rare in the Yukon (Douglas et al. 1981),
Manitoba (White and Johnson 1980), Ontario (Argus and White 1977),
and Saskatchewan (Maher et al. 1979).

6. Woods/a obtusa(Spreng.)Torr.

blunt-lobed woodsia
Fig. 98 (a) fronds; (6) fertile pinna. Map 96.

Fronds 10-30 cm long, 2-10 cm wide. Blades broadly lanceolate,
pinnate; pinnae mostly remote; lower pinnae triangular; median and
upper pinnae ovate-lanceolate to oblong, pinnatifid, or pinnate at the
base. Pinnules oblong, obtuse. Stipes not jointed. Rachis straw-
colored, glandular-pubescent. Sori round, near the margins. Indusia
covering the sori, later splitting into several jagged lobes.

Woodsia obtusa is an erect rather robust species. In Canada it is
highly restricted in distribution (Britton 1977; Lafontaine 1973) and
is a talus species rather than a cliff species. In aspect it looks
somewhat like Cystopteris fragilis, with which it often grows. The
stifTer aspect and the glands and scales on the axes and veins are good
field characters (see also Woodsia compared with Cystopteris under
comments on the genus Woodsia).

Cytology: m = 76 (Brown 1964). A tetraploid species. Brown (1964)
is uncertain as to its origin, but considers it to be clearly related to
W. oregana.

Habitat: Shaded ledges and rocky slopes.

Range: Southwestern Quebec and southern Ontario, south to
Georgia, Alabama, and Texas.

Remarks: The Ontario habitats seem disturbed and might indicate
that the species is a recent arrival. It is rare in Ontario (Argus and
White 1977) and certainly rare in Quebec.

180

Fig. 98 Woodsia obtusa; (a) fronds, 1/2 x ; (6) fertile pinna, 4 x

181

Hybrids of Woods i a

Within the group, three species have an even stubble. One might
expect three hybrids. The basic cross of the two diploids would be V^.
glabella X iluensis or W. X tryonis Boivin; it was collected at Silver
Islet, Thunder Bay District, Ont. {Tryon and Faber 4962) (see R.M.
Tryon 1948). This plant was sterile and had the morphology of W.
alpina, as expected. Unfortunately, no cytology is known for the
hybrid combination (one would expect all unpaired chromosomes), and
Brown (1964) cited Tryon and Faber 4962 under W. X gracilis.

There should also be two backcrosses of W. alpina to its two
parents. Woodsia alpina X iluensis is W. X gracilis (Lawson) Butters
and is reported from Thunder Cape, Thunder Bay District, Ont., and
Riviere du Loup, Que., by Brown (1964). Woodsia alpina X glabella
was reported by Soper and Maycock (1963) from Algoma District, Ont.
Their cytology is unknown, but they should be sterile triploids.

Hybrids between the even and uneven stubble groups are known.
The most often cited hybrid is W. X abbeae Butters, which was
considered to be one of the following: W. iluensis X scopulina by R.M.
Tryon (1948); possibly W. iluensis X oregana by Hagenah (1963); and
more specifically, W. iluensis X oregana var. cathcartiana by W.H.
Wagner and F.S. Wagner (unpublished). It was studied cytologically
and was said to be triploid. We consider W. confusa Taylor and W.
oregana var. squammosa Boivin to be synonyms of W. X abbeae.

The type oiW. X maxonii Tryon {W. oregana X scopulina) Tryon
(1948) was collected on Sleeping Giant, Thunder Bay District, Ont.

Woodsia hybrids are in need of further study with the use of
modern experimental methods.

4. Polystichum Roth

Ferns rather large, tufted, evergreen, and leather-textured, with
usually scaly stipes, arising from short stout chaffy rhizomes. Sori
round. Indusia round, attached at the center.

The four species and all their hybrids known in Europe have
been extensively analyzed cytogenetically (Lovis 1977). The
chromosome numbers of over 75 species in the world are known, which
is perhaps a little over half the total number of species. The genus is a
large one, almost as large as Dryopteris, and is quite complex in
western North America. In fact, one can make an analogy between
the complexities of Dryopteris in eastern North America and the
relative dearth of species in the west, with the reverse situation in
Polystichum. In each case, derived tetraploids are known, and in each
case, there is even a hexaploid. Indeed, in Polystichum outside
Canada, octaploids are known (Lovis 1977). Another parallel for the
amateur is the recognition here of a large number of biological species
that have been delineated from the cytogenetic analyses. Our present
understanding of the western Polystichum species is presented by

182

D.H. Wagner (1979) and will be briefly outlined. After extensive
studies, Wagner recognized P. imbricans as an important basic species
rather than as a variety of P. munitum. Accordingly, for the smaller
talus and cliff species we have the phylogenetic schemes shown in
Diagram 3.

There are still some questions regarding these schemes.
Polystichum braunii and P. andersonii are often in the same ecological
niche in British Columbia, and although D.H. Wagner (1979) is
convinced that they are quite distinct in morphology and have entirely
separate origins, others find them quite difficult to identify easily.
D.H. Wagner (1979) also believes that he has a good lead from an old
herbarium specimen to And the ancestral diploid species that we have
called species W. Yet another problem is the relationship of P.
lemmonii to P. mohrioides, although again D.H. Wagner (1979) is
quite convinced that they are separate entities. The latter species has
an amazing range, right down the Andes chain to southern South
America.

Accordingly, there are 10 species in western Canada of which
three, P. californicum, P. lemmonii, and P. scopulinum, are of rare to
very rare status. Polystichum kruckebergii and P. setigerum are of
local occurrence in British Columbia, so that it was quite possible for
the junior author to spend 6 months in British Columbia traveling
around Vancouver and Victoria and see only P. munitum and P.
imbricans of the 10 species recorded for the province.

In eastern Canada there are only four species: P. braunii, P.
lonchitis, P. scopulinum, and P. acrostichoides. The most common and
familiar of these is P. acrostichoides, a characteristic plant of maple
and beech woods. It is a basic diploid species that as yet has not been
implicated as an ancestral diploid species in the evolution of the
polyploids.

The genus Polystichum is noted for hybridization, and so if one is
in a region where there are a number of species growing together,
there is a good opportunity for interspecific hybrids to be present.
W.H. Wagner (19736) studied a number of hybrids at one locality in
Washington.

A. Sori borne on reduced upper pinnae 1. P. acrostichoides

A. Sori borne on the backs of unmodified pinnae.

B. Fronds pinnate; pinnae entire, denticulate, or serrate,
spinulose.

C. Pinnae mostly oblong-lanceolate, progressively
reduced towards the base, with the lowest pinnae
subtriangular to broadly trowel-shaped symmetrical;

spinulose tipsof teeth of pinnae spreading

2. P. lonchitis

C. Pinnae linear-attenuate, not much reduced below,
with the lowest pinnae ovate to lanceolate-falcate,
auriculate, asymmetrical; spinulose tips of teeth of
pinnae incurved.

183

D. Stipe and rachis persistently chaffy; pinnae
acuminate, cuneate at the base; indusium ciliate

7. P. munitum

D. Stipe and rachis often naked; pinnae cuspidate or
apiculate, oblique at the base; indusium entire to

sharply toothed 4. P. imbricans

B. Fronds bipinnatifid or bipinnate; pinnae spinulose or not.

E. Pinnae not at all spinulose 3. P. lemmonii

E. Pinnae apiculate to spinulose.

F. Fronds bipinnate; pinnules distinct, sessile or

petiolate 10. P. braunii

F. Fronds bipinnatifid; pinnules adnate to the costa.
G. Pinnae with conspicuous filiform scales on
the lower surface.

H. Fronds with a proliferous bud on the
rachis about one-third of the way down

from the tip 8. P. andersonii

H. Fronds lacking a proliferous bud.

I. Pinnae incised to the costa;

pinnules slightly toothed

11. P. setigerum

I. Pinnae not incised to the costa . .

9. P. californicum

G. Pinnae lacking filiform scales.

J. Pinnae acute at the apex; teeth coarse,

spreading 5. P. kruckebergii

J. Pinnae obtuse at the apex,
occasionally cuspidate; teeth fine,
incurved 6. P. scopulinum

1. Polystichum acrostichoides (Michx.) Schott

Christmas fern
Fig. 99 (a) sterile and fertile fronds; (6) portion of fertile pinna with
immature sori; (c) portion of fertile pinna with mature sori. Map 97.

Fronds 35-65 cm long. Stipes and rachis chaffy. Blades
lanceolate, 7-12 cm wide or wider, simply pinnate; pinnae oblong to
lanceolate, acute or sometimes bluntish at the tip, auricled at the base
on the upper side; margins serrulate-bristly. Sori borne on reduced
upper pinnae distinct or more often confluent.

Forma incisum (Gray) Gilbert has the pinnae coarsely toothed,
the fertile pinnae usually less reduced, and the sori usually less
confluent.

This medium-sized species with dark green, subevergreen fronds
is sufficiently striking to be soon familiar to all amateurs walking in
maple-beech woods in southeastern Canada. The first impression is a

184

.Cj

:5>

O."

en ^

,(0

■5J

.■t^

.^iX

-55

$ o

.<^ Q

ex.

^ *

-^ -1

:t:

ex:-'

.■».j

^ o

C CD
-1^

Cl.

*><. ^

^ m

^ *

^.1

.9> 5

cx:

X "

<M X

(£>

^ X

."2 E

Q. C/)

o '^

CL <1>

o £

P^ o

r~-- ro c

^ c .9

r- TO

.t^ E o

> O) ^

O CD —

_l CO Q.

- CD E

CO ^ ._

■D CT3 en

o .9 en

■a"g CD

CT3 CO CD

.^ CD

^ E

■D O

a; o
^ E

_ o

O JD

E _ CD

CT3 "o "^

1- ^ -!

CD O)
.5 C

.^ 5^ o

i: "-^ ~

-^ CD 1^

^ ^K

^ -^

jz - O

'^ -^ Q.

2^ OJ >^

CO w O

CO .5^ §-

CD o —

E g CO

^ CO o

CD C £

-c x: c

en o <D

< Q. :t

CD y (D

° TO ■^

Z -D 2

Q
c

CO
05

3
■♦3

u
.0

c
o

185

Fig. 99 Polystichum acrostichoides; (a) sterile and fertile fronds, 1/3 x ; (b) portion
of fertile pinna with immature sori, 5 x ; (c) portion of fertile pinna with mature
sori, 5 X .

186

darker green, tougher Boston fern that grows scattered about in open
woods. The Boston fern is a tropical genus, Nephrolepis, but there is a
superficial similarity in aspect.

Cytology: n = 41 (Britton 1953*; Cody and Mulligan 1982*).

Habitat: Rich woods and humus-rich rocky slopes.

Range: Nova Scotia to southern Ontario, south to northern Florida
and eastern Texas. The forma incisum may be found through the
range of the species.

Remarks: There are many described forms (Weatherby 1936
describes seven of them) that are considered to be ecological variants
or, in some cases, mutations. The observation of variation in the leaf
form of the species is an interesting hobby for the amateur if a limited
number of fern species are available near at hand.

2. Polystichum lonchitis (L.) Roth

Aspidium lonchitis (L.) Sw.

Dryopteris lonchitis (L.) O. Kuntze

Holly fern
Fig. 100 (a) fertile frond; (6) pinna with mature sori; (c) sorus;
(d) undersurface of sterile pinna. Map 98.

Fronds 10-60 cm long. Stipes very short, chaffy. Blades linear to
narrowly linear-oblanceolate, acuminate, tapering to the base.
Middle and upper pinnae oblong-lanceolate, falcate; bases of pinnae
auriculate above and cuneate below; basal pinnae deltoid, often very
small, equilateral, serrate-dentate; teeth spreading-spinulose. Sori
round, in two rows, occurring midway between the midvein and the
margin. Indusium entire.

Field characters are the medium to small size of the plants, the
short pinnae, the extremely short stipe, and the pinnae towards the
base reduced to small triangular auricles. The species is aptly called
the holly fern because of its shiny, lustrous green, tough blades.

Cytology: n = 41 (Britton 1964; Cody and Mulligan 1982*, eastern
Canada; R.L. Taylor and Mulligan 1968*, western Canada).

Habitat: Limestone cliffs, moist rocky slopes, talus slopes, and
occasionally coniferous woods.

. / .

Range: Circumpolar; in North America in Greenland, western
Newfoundland, Cape Breton Island, Gaspe Peninsula, central
Quebec-Labrador (Waterway and Lei 1982), Bruce Peninsula and

187

Fig. 100 Polystichum lonchitis; (a) fertile frond, 1/2 x; (b) pinna with mature sori,
4 X ; (c) sorus, 15 x ; (d) lower surface of sterile pinna, 4 x .

188

Algoma District, Ont., Keweenaw County, Mich., southwestern
Alberta (Brunton 1978), British Columbia, Yukon Territory to Kenai
Peninsula, Alaska, south in the western United States to Colorado,
Utah, and southern California.

Remarks: This characteristic fern of cool, northern, limestone
habitats has a broad distribution. In Ontario it is often a companion
plant for Hart's-tongue (Soper 1954). The species is rare in the Yukon
(Douglas et al. 1981), Nova Scotia (Maher et al. 1978), and Ontario
(Argus and White 1977).

3. Polystichum lemmonii Under w.

P. mohrioides (Bory) Presl var. lemmonii (Underw.) Fern.

P. mohrioides auth. non (Bory) Presl
Fig. 101 (a) frond; (6) fertile pinna. Map 99.

Fronds 15-35 cm long or longer, densely clustered from a short
ascending rhizome. Stipes glandular, puberulent, and very chaffy at
the base. Blades linear to narrowly lance-oblong; pinnae deeply
pinnatifid or the lower pinnae pinnate; ultimate segments oval,
obtuse, crenate, or crenately lobed; lobes lacking mucronate or
spinulose tips. Sori on the middle and upper pinnae, towards the base
of the pinnules. Indusia large, entire, or obscurely erose-toothed.

This species is confined to ultramafic rocks (Kruckeberg 1964).
In comparison with P. imbricans, it is a small species, with fronds less
than 30 cm long. The pinnules are rounded and overlapping and lack
spines. Both the upper and lower epidermis have unicellular glands.

Cytology: n = 41 (W.H. Wagner 19736). This is a basic ancestral
species with genomes LeLe.

Habitat: Open serpentine and asbestos subalpine slopes.

Range: In Canada known only in the Okanagan Divide in southern
British Columbia; in the United States, from northern Washington to
northern California.

Remarks: This is a rare species in all of western North America and
the distribution barely extends into Canada. We are following D.H.
Wagner (1979) in considering this species distinct from P. mohrioides.
The latter is known from the Andes south to Chile.

4. Polystichum imbricans (DC. Eat.) D.H. Wagner

P. munitum (Kaulf ) Presl var. imbricans (D.C. Eat.) Maxon
Fig. 102 (a) frond; (6) fertile pinna. Map 100.

189

^ N

Fig. 101 Polystichum lemmonii; (a) frond, 1/2 x ; (b) fertile pinna, 2 x

190

Fig. 102 Polystichum imbricans; (a) frond, 1/3 x ; (b) fertile pinna, 3 x

191

Fronds similar to P. munitum, linear-lanceolate, but shorter, up
to 60 cm long. Stipes less chaffy than P. munitum, with the upper part
and rachis smooth or nearly so. Pinnae usually overlapping and
folded inward, lanceolate, 2-4 cm long or longer, auriculate above,
abruptly tapering to the spinulose tip; teeth incurved, spinulose. Sori
midway between the margin and midvein. Indusia entire.

Plants are generally smaller than those of P. munitum and have
crowded ascending pinnae, stiffly erect habit, flat pinnae or pinnae
slightly cupped on the upper side; the upper surface of the pinna is
perpendicular to the rachis. D.H. Wagner (1979) has studied P.
imbricans extensively and recommends using a combination of
characters for absolute identification. He lists eight characters in
order of reliability for exact determination.

Cytology: n = 41 (Taylor and Lang 1963*). This is a basic ancestral
species with genomes II.

Habitat: Usually in the open in rock crevices, clearings, and dry
rocky coniferous woods.

Range: Southern coastal British Columbia, south to southern
California.

Remarks: In making determinations, make sure you use technical
characters, because plants growing in shady, moist situations
superficially mimic P. munitum. Conversely, P. munitum in drier,
more exposed locations tends to look like P. imbricans.

5. Polystichum kruckebergii W . H . Wagner

Kruckeberg's holly fern
Fig. 103(a) fronds; (6) fertile pinna. Map 101.

Fronds up to 30 cm long, few together, tufted from a small stout
erect rhizome. Stipe short, scaly. Blade linear-lanceolate; pinnae
overlapping, ovate-triangular, conspicuously spreading, toothed; teeth
tips cartilaginous; larger pinnae frequently with one or more pairs of
basal pinnules. Sori borne in two rows on the backs of the pinnae on
the upper half of the frond, becoming confluent. Indusia with entire
wavy margins.

Polystichum kruckebergii was described by W.H. Wagner
(1966a), and the type chosen was a collection from near Lillooet in
southwestern British Columbia. W.H. Wagner (1966a) carefully
delineates it from P. scopulinum, noting that the former is usually
smaller, with shorter and less oblong pinnae; in a median pinna the
number of teeth per side is approximately 6 rather than 12 (8-25), as
in P. scopulinum. The margins of P. kruckebergii are markedly
bristly, and the pinna tips are more pointed.

192

Fig. 103 Polystichum kruckebergii; (a) fronds, 1/2 x ; (b) fertile pinna, 10 x

193

Cytology: n = S2 (W.FI. Wagner 19736). Derived tetraploid
LoLoLeLe from P. lonchitis and P. lemmonii (W.H. Wagner 19736).

Habitat: Subalpine cliffs and talus slopes.

Range: Central British Columbia (A.L. Kruckeberg 1982) to
northern California, east to Idaho and Utah.

Remarks: This is a rare species that should be looked for on
ultramafic rocks. Its known distribution in Canada is very limited.

6. Polystichum scopulinum (DC. Eat.) Maxon

P. mohrioides (Bory) Presl var. scopulinum (D.C. Eat.) Fern.
Aspidium aculeatum (L.) Roth var. scopulinum D.C. Eat.
crag holly fern

Fig. 104 (a) fronds; (6) fertile pinna. Map 102.

Fronds 15-40 cm long, densely tufted from short erect or decum-
bent scaly rhizomes. Stipes densely chaffy at the base; scales sparse
and deciduous above. Blades narrowly lanceolate, slightly tapered to
the base and tip; pinnae usually folded inwards and upwards,
deltoid-ovate to deltoid-oblong, pinnately lobed, especially towards the
base; teeth with a cartilaginous tip. Sori borne on the middle and
upper pinnae, in two median rows. Indusia thin, erose-dentate.

The species is intermediate between P. imbricans and P.
lemmonii. Characteristics from the former include the folding inward
and upward of the pinnae and the leathery to fleshy texture of the
blade. The pinnae are oblong and have about 12 short but distinct
cartilaginous teeth per side.

Cytology: n = S2 (W.H. Wagner 19736). A derived tetraploid of
constitution II LeLe from P. imbricans and P. lemmonii (D.H. Wagner
1979), not P. munitum and P. mohrioides (W.H. Wagner 19736).

Habitat: Crevices of cliffs and rocky slopes, often of ultramafic, or at
least basic, rocks.

Range: In western North America from southernmost British
Columbia to southern California, Idaho, and Utah; disjunct in the
Gaspe Peninsula, Que., and western Newfoundland.

Remarks: Polystichum scopulinum has a much wider distribution
than its presumed parents. The disjunct station on Mont-Albert, Que.,
has been a noted topic through the years. The species should be looked
for when on serpentine rocks.

194

Fig. 104 Polystkhum scopulinum; (a) fronds, 2/3 x ; (b) fertile pinna, 2 1/2 x .

195

7. Polystichum munitum (Kaulf.) Presl

sword fern
Fig. 105 (a) frond; (6) fertile pinna. Map 103.

Fronds 20-150 cm long, forming a stiffly erect crown at the stout
woody scaly rhizome. Stipes densely chaffy. Blade linear-lanceolate,
short acuminate, pinnate; pinnae linear-attenuate, auriculate at the
base above, cuneate below, sharply serrate. Sori large, situated
midway between the margin and the midvein. Indusium fimbriate-
margined.

Large plants (over 1 m) present no identification problems. The
fronds are once pinnate and the pinnae are sharply serrate. The
sharply serrate pinnae are a variable feature, and some plants have
deeply serrate or even incised pinnae (Calder and Taylor 1968). There
is no difficulty in seeing impressive colonies of this species in easily
accessible localities in the coastal forests of British Columbia, e.g.
Stanley Park, Vancouver, and Pacific Rim National Park. Subalpine
plants superficially resemble P. lonchitis.

Cytology: n ^ 41 (W.H. Wagner 19736; Cody and Mulligan 1982*).
This is an important basic ancestral diploid species MM.

Habitat: Moist coniferous woods and shaded slopes; particularly
common along roadside clearings in southern coastal British
Columbia.

Range: Alaskan Panhandle, south near the coast to Baja California,
Mexico; inland in southern British Columbia and to northern Idaho
and northwestern Montana.

Remarks: This species is a striking component of the western coastal
forests.

8. Polystichum andersonii Hopkins

P. hraunii (Spenner) Fee ssp. andersonii (Hopkins) Calder
& Taylor

Anderson's holly fern
Fig. 106 (a) frond; {h) fertile pinna. Map 104.

Fronds to 1 m long or longer, usually with a proliferous scaly bud
near the apex. Stipes about one-fifth the length of the frond, persis-
tently chaffy. Blades lanceolate, narrowed towards the base; pinnae
oblong-lanceolate; lowermost pinnae subtriangular, deeply cut to the
costa, but with segments rarely undercut; segments with spinulose
teeth; both surfaces with filiform scales; rachis and costa with broader
scales. Sori 1-8 on the lateral segments of the middle and upper
pinnae. Indusia erose-dentate; teeth gland-tipped.

196

Fig. 105 Polystichum munitum; (a) frond, 1/2 x ; (b) fertile pinna, 3 x

197

'»^-

Fig. 106 Polystichum andersonii; (a) frond, 1/2 x ; (b) fertile pinna, 3 x

198

This species has often been referred to subspecific or varietal
status under P. braunii. Taylor (1970) suggests that it may prove to be
"only a geographical variant."

Polystichum andersonii is identified by the presence of one or
more proliferous buds on the rachis, and the basal distal or upper pin-
nules on the pinnae are longer than the adjacent ones. D.H. Wagner
(1979) considers the species to be quite distinct from P. braunii and
postulates that it is an allotetraploid, with one parent being P.
munitum and the other an undescribed species to which we have
referred as Species W in Scheme B. W.H. Wagner (19736) postulated
quite a different origin for P. andersonii, saying it seemed to be too far
removed from P. munitum to have that species as an ancestor.

Cytology: n — %2 (Taylor and Lang 1963*). Postulated genomes
MMWW.

Habitat: Moist woods and shaded rocky slopes in the mountains.

Range: Alaskan Panhandle, south to Oregon, Idaho, and Montana.

Remarks: An analysis that compares hybrids of P. andersonii with
those of P. braunii, as well as with those of other species, would clarify
the origin of P. andersonii. The origin of the species, as presented by
D.H. Wagner (1979), is still hypothetical.

9. Polystichum califomicum (D.C. Eat.) Diels

Aspidium californicum D.C. Eat.

California holly fern
Fig. 107 (a) frond; (b) fertile pinna. Map 105.

Fronds 40-75 cm long from the erect rhizome. Stipes about
one-third the length of the frond and chaffy, especially towards the
base; upper part of stipe becoming naked. Blades linear-lanceolate to
lanceolate, little narrowed towards the base; pinnae deeply cut, with
segments often slightly undercut, overlapping, and toothed in the
upper part; teeth with short ascending or incurved spinulose tips and
with filiform hairs below and along the costa above. Basal pinnule on
the upper side of the pinnae usually somewhat enlarged. Sori in two
rows on the segments of the middle and upper pinnae. Indusia large,
ciliate.

Scheme B of Diagram 3 suggests that P. munitum crossed with P.
dudleyi (not in Canada), and the hybrid gave rise to P. californicum
(W.H. Wagner 19736). D.H. Wagner (1979) postulated that P. dudleyi
might have crossed with P. imbricans to give rise to another
allotetraploid now included in P. californicum. He rejected the second
hypothesis at that time because all the P. californicum plants that he
studied were uniform with the chemical analysis that he used.

199

Fig. 107 Polystichum californicum; (a) frond, 1/2 x ; (b) fertile pinna, 1 1/2 x

200

Recognition of the smaller, northern forms of this species
presents great difficulty. If a colony of plants can be found that key
out to this species, it will be necessary to use all the methods we now
have available to analyze the plants (see remarks that follow).

Cytology: n = S2 (W.H. Wagner 19736). Derived allotetraploid
MMDD.

Habitat: Lowland coastal forests of Canada.

Range: Known in Canada only in Texada Island, B.C.; in the United
States south in the mountains to central California.

Remarks: This species was not included in Canada's flora by T.M.C.
Taylor (1970). Identification problems are certainly apparent when
D.H. Wagner (1979) refers to "recent misidentiiications of northern
populations from B.C., Washington and California of this species as P.
scopulinum.'' Further study is required. Inclusion of this species in
the Canadian flora rests on one old specimen from Texada Island —
Anderson 666 (V).

10. Po/yst/chum braun/7 (Spenner) Fee

P. braunii (Spenner) Fee var. purshii Fern.
Braun's holly fern
Fig. 108 (a) frond; (6) portion of fertile pinnule; (c) sorus. Map 106.

Fronds to 1 m long forming a crown at the end of the stout
ascending rhizome. Stipe about one-sixth the length of the frond,
persistently chaffy. Blades broadly lanceolate, narrowed at the base;
rachis with persistent dense chaff; pinnae slenderly lanceolate; middle
and upper pinnae gradually tapering, with the lower straight-sided
and abruptly tapering to the apex; pinnae generally once pinnate;
pinnules petiolate or rarely slightly decurrent, narrowly ovate to
trapezoid-oblong, obtuse, nearly rectangular at the base and slightly
auricled on the upper side, sharply serrate with incurved bristle-
tipped teeth. Sori in two rows near the midrib. Indusia often erose.

This large and handsome species, with fully bipinnate blades and
bristle-tipped teeth, is readily identified, provided it can be
distinguished from P. andersonii and P. setigerum. The lack of
proliferous buds and of enlarged proximal pinnules on the basal
pinnae is useful in this regard.

Cytology: n = S2 (Taylor and Lang 1963; R.L. Taylor and Mulligan
1968*, western Canada; Cody and Mulligan 1982,* eastern Canada).
The ancestral genomes of the European plants have not been
identified. Levis (1977) considers it to be a segmental allotetraploid

201

Fig. 108 Polystichum braunii; (a) frond, 1/3 x ; (b) portion of fertile pinnule, 4 x ; (c)
sorus, 20x.

202

and gives it the formula BBBB (the origin of B is obscure, but it is
presumably an ancestral diploid, P. braunii). We have tentatively
designated it as XXYY, although the origins of X and Y are obscure.

Habitat: Rich woods and shaded talus slopes.

Range: Circumpolar; in North America from Labrador and
Newfoundland to Thunder Bay District, Ont., south to Pennsylvania
and Michigan; western British Columbia north to the Kenai
Peninsula and Kodiak Island, Alaska.

Remarks: We are in agreement with D.H. Wagner (1979) that
recognition of a var. or ssp. purshii, based on plants with an increased
proportion of broad to filiform laminar scales (Calder and Taylor
1968a), serves no useful purpose and should be dropped. It is not
possible to designate the plants in western Canada as either all var.
braunii or as all var. purshii. Polystichum braunii is rare in Ontario.
(Argus and White 1977).

1 1 . Polystichum setigerum (Presl) Presl

P. alaskense Maxon

P. braunii (Spenner) Fee ssp. alaskense (Maxon) Calder & Taylor

Alaskan holly fern
Fig. 109 (a) frond; (b) portion of fertile pinna. Map 107.

Fronds 1 m long or longer. Stipes about one-fifth the length of
the frond, persistently chaffy. Blades lanceolate, narrowed at the
base; lower pinnae often deflexed; pinnae oblong-lanceolate,
pinnatifid, with segments undercut; teeth bristle-tipped; pinnae scaly
on both surfaces, with filiform scales and broader scales on the rachis
and the costa. Sori in two rows on the pinnules of the upper half of the
frond. Indusia ciliate.

Polystichum setigerum is still another entity in Scheme B that
looks very like P. braunii and P. andersonii — so much so, that Calder
and Taylor (1968a) treated it as ssp. alaskense (Maxon) Calder &
Taylor. Because it is a derived hexaploid, it is now treated as a full
species, P. setigerum, based on a plant collected by Thaddaeus Haenke
in 1791 at Nootka Sound, B.C. Maxon in 1918 named this species P.
alaskense. D.H. Wagner (1979) states that the best features for
determining this species are the lack of a proliferous bud (a P.
andersonii feature), a degree of cutting or incision of the pinnae
(similar to that of P. braunii), and the enlarged proximal pinnules on
the basal pinnae (as in P. andersonii).

Cytology: n = 123 (D.H. Wagner 1979*). Perhaps derived from P.
munitum and P . braunii.

203

Fig. 109 Polystichum setigerum; (a) frond, 1/3 x ; (b) portion of fertile pinna, 3 x

204

Habitat: Lowland coastal forests in dense woods and on shaded rocky
slopes; at times growing with P. andersonii and P. braunii.

Range: Alaskan Panhandle, south to southern British Columbia;
disjunct to Attu Island in the Aleutian Islands.

Remarks: It is unlikely that the last word has been written on all the
confusing species in Scheme B of Diagram 3. D.H. Wagner (1979)
admits that P. lonchitis rather than P. munitum might be a possible
parent and that P. andersonii is also a possibility rather than P.
braunii. Only by an extensive cytogenetic analysis of hybrids, with
the use of the modern arsenal of SEM and chemistry, will we be able to
solve this problem.

Hybrids of Polystichum

Polystichum X hagenahii Cody (P. acrostichoides X lonchitis) is
known only from the type locality. Cape Crocker Indian Reserve,
Bruce County, Ont.

Specimens of P. braunii X acrostichoides have been seen from
Inverness County, N.S., and Waterloo, Que.

Polystichum munitum X imbricans is known from Mount
Newton, Vancouver Island, B.C. (D.H. Wagner 1979).

W.H. Wagner (19736) discusses a large number of sterile hybrids
that he has studied in the western United States, many of which could
occur in Canada. Still other hybrids have been reported in the
literature (Knobloch 1976).

5. Dryop ter/s Adans. wood fern

Usually large (one species is small) ferns with fronds arising in
clusters from stout creeping or erect rootstocks. Stipes continuous
with the rootstock, not jointed. Blades bipinnatifld or pinnate to
nearly bipinnate, glabrous, or somewhat pubescent. Indusium
roundish reniform, attached in the centre, covering the rounded sori.
Veins usually free, simple, or forked.

The genus has been extensively studied cytologically in Europe
starting with Manton (1950), followed by her graduate student S.
Walker (1961) and Walker's graduate student, M. Gibby (Gibby and
Walker 1977). In North America, Britton (1953) and Wagner (1970)
have studied Dryopteris in some detail. Our current understanding of
the evolution of the species in eastern North America is summarized
in Wagner (1970) and Lovis (1977). Widen in Finland has studied the
chemistry of the phloroglucinols of the various species of the world,
which has been useful in our understanding of the relationships
within the genus (see review by von Euw et al. 1980). The external

205

morphology of the spores has been examined by SEM (Britton 1972a,
19726; Britton and Jermy 1974).

In the phylogenetic scheme (see Diagram 4) there is still one
ancestral genome (B) missing (S of W.H. Wagner 1970); otherwise the
species seem quite well analyzed from a few artificial hybrids and
from many natural ones.

Of the seven extant diploids, only three are known that have not
participated further in evolution. These are Dryopteris arguta, D.
fragrans, and D. marginalis. The others are ancestral to the derived
alloploids. Although D. ludouiciana is confined to southeastern
United States and so is not part of our flora, its influence is considered
to be present in the origin of D. cristata and from this, in turn, of D.
clintoniana. The evolution of the species can be shown schematically
as in Diagram 5.

Readers familiar with previous treatments of this group can
appreciate that problems in identification have arisen regarding the
so-called "Z). spinulosa complex", i.e., those species to the left of species
B. Other problems have arisen in the past between D. cristata and D.
clintoniana, two species to the right of species B. At this time, there
are still problems in distinguishing between D. expansa and D.
campyloptera, and some researchers are impressed by the differences
between D. expansa in eastern Canada compared with D. expansa in
western Canada. Carlson and W.H. Wagner (1982) have recently
compared the distributions of the North American members of this
genus.

A. Blades usually small, copiously scaly on the under surface; old
fronds or their bases forming a conspicuously persistent curled

tuft at the base of the plant; indusia large, glandular

2. D. fragrans

A. Blades usually large, scales few or absent.

B. Sori marginal or nearly so; blade leathery, grayish green,
paler beneath; ultimate segments of pinnae round-lobed . . .

8. D. marginalis

B. Sori medial to submedial; pinnae with sharp-toothed
segments.
C. Basal pinnules on basal pinnae sessile or adnate.

D. Fronds dimorphic; the sterile fronds shorter and
more lax; pinnae of fertile fronds often in a

nearly horizontal position 10. D. cristata

D. Sterile and fertile fronds similar; pinnae of the
fertile fronds in the same plane as the blade or
nearly so.
E. Blade broadest near the middle; stipe much

shorter than the blade 7. D. filix-mas

E. Blade broadest or nearly so at the base;
stipe longer than the blade.

206

F. Stipe up to half the length of the blade;

teeth of pinnules spine-like

1. D. arguta

F. Stipe about as long as the blade; teeth

of pinnules not spine-like.

G. Blade reduced rather gradually

to the apex; pinnae broadly

triangular to long-triangular,

broadest at the base

1. D. clintoniana

G. Blade abruptly reduced to an
acuminate apex; pinnae narrowly
lanceolate to narrowly oblong-
lanceolate, broadest at the

middle 9. O. goldiana

C. Basal pinnules on basal pinnae stalked.

H. Indusia and blade (especially at the base of the

pinnae) definitely to densely glandular

3. D. intermedia

H. Indusia glabrous; blade usually glabrous,

occasionally slightly glandular.

I. Lower basal pinnule on each basal pinna

closer to the second upper pinnule than to

the basal upper one

J. Blades ovate-triangular, arching,

short-stiped 5. D. campyloptera

J. Blades broadly triangular to broadly
oblong, nearly upright, long-stiped . . .

4. D. expansa

I. Lower basal pinnule on each basal pinna
closer to the upper basal pinnule than to the
second upper one 6. D. carthusiana

1 . Dryopteris arguta (Kaulf.) Maxon

Aspidium rigidum Am. auth.

D. rigida (Sw.) A. Gray var. arguta (Kaulf.) Underw.

coastal shield fern
Fig. 110 (a) frond; (6) fertile pinnule. Map 108.

Fronds up to 70 cm long, evergreen, tufted from the short-
creeping stout rhizome. Stipe stout, up to half the length of the blade,
scaly. Blade widest towards the base, twice pinnate; pinnae oblong-
lanceolate, long-acuminate; pinnules oblong, mostly rounded-obtuse,
serrate to pinnately incised; veinlets spreading, all ending in salient
often cartilaginous spine-like teeth. Sori large, medial. Indusia pale
greenish yellow, glabrous, but with somewhat glandular margins.

207

CD
■D
CD
C
CD

c^-

"D
CO

CD
"D
CD
CD

sz
c

CD
O

(/)

—

(—

E
o

c
_l

LLI
LU

CD
CD

O
O
O

CO
CD
CJ

■*->
^-

CQ
CD

U-

>—t

J5
CD

cts

.CD

.CO
CD

.CD

CD
.CD

CtJ

■ CXJ

Ct3

CD
CD

CO
.0)

O
CD

CD
O

»-^
CD
O

■♦«^

CD
.CO

c::i

CO
(D

CD
Q.

"en

' — ^

O
CD
Q.

■D

CX)

II
c:

id (6X)
n = 123

£S

T-

o -

-^ CD

■D

E
o

5 ^

O
Q.

cnT

CX)

C
C\J

CD II

g CN

"co

^
^

"co

■D

C\J

208

t>«

1-H

^..^

■^

■0

CJ5

CtJ

-0

.Co

-C

1-H

§ O

.to

■^ ^

c *

D.cli
BBLL

(

Q
n

1-H

CtJ

»H

•5

€

"rt

<tj

«j

^
.^

a
o

«z

«T3

■

«tj

Ct3

e«

— _— ^^^^— ^— ^— ^-^

209

Fig. 1 10 Dryopteris arguta; (a) fronds, 1/4 x ; (b) fertile pinnule, 3 x

210

This fern might be confused with D. filix-mas; however, it differs
from that species by having the blade widest towards the base and the
presence of spine-like teeth on the pinnules. Also, on the West Coast it
is more strongly evergreen thanD. filix-mas.

Cytology: n = 41 (W.H. Wagner and Chen 1964).

Habitat: Deep humus among broken rocks and in rocky woods along
the coast.

Range: Apparently limited in Canada to the southeast coast of
Vancouver Island and the islands of the Gulf of Georgia, B.C.; south in
the United States to California and inland to Arizona.

Remarks: This is a rare fern in Canada. We have seen a large colony
near Nanaimo, B.C., which must have been established for a long
time. It seemed surprising, when we searched the area, that
superficially similar habitats did not harbor this species. This lack of
aggression is of course one reason for its rarity.

2. Dryopteris fragrans (L.) Schott

D. fragrans (L.) Schott var. remotiuscula Komarov

Aspidium fragrans (L.) Sw.

Thelypteris fragrans (L.) Nieuwl.

fragrant cliff fern
Fig. Ill (a) frond; (b) portion of fertile pinnule. Map 109.

Fronds up to 30 cm long or longer, forming a spreading or
ascending crown from a stout rhizome; old fronds curled, shriveled,
and persistent. Stipes 1-15 cm long, glandular, and chaffy. Blades
coriaceous, tapering from the middle to the base and apex; pinnae
overlapping and often inrolled, densely chaffy with brown to reddish
scales; pinnae oblong-lanceolate, pinnately incised or crenate;
rachises and pinnae glandular. Indusia large and often overlapping,
whitish, becoming brown, with their margins often ragged.

Dryopteris fragrans is always a pleasant surprise to anyone
visiting a rocky environment. When climbing talus slopes or skirting
cliffs, one is rewarded by seeing this fern emerging from a crevice or
from under a talus boulder. At times, it might be mistaken for
Woodsia ilvensis, but the tell-tale curled dead fronds hanging below
the plant make for an easy field check, even from some distance.

Cytology: n = 41 (Britton and Soper 1966*; T.M.C. Taylor and Lang
1963*). This species is not ancestral to any of our other species.

Habitat: Cliffs and talus slopes (often somewhat calcareous).

211

Fig. 1 1 1 Dryopteris fragrans; (a) frond, 2/3 x ; (b) portion of fertile pinnule, 3 x

212

Range: Circumpolar; in North America from Greenland to Alaska,
south to Newfoundland, New York, Wisconsin, Minnesota, and
northern British Columbia. Extremely abundant near Lake Superior.

Remarks: We consider the southern var. remotiuscula Komarov,
which is larger, more lax, and has more distant pinnae, to be a
response to the longer growing season in the southern part of its
range. This expression is clinal, with no clear demarcation
geographically. We prefer to ignore the variety. Dryopteris fragrans
is rare in Nova Scotia (Maher et al. 1978) and Alberta (Argus and
White 1978).

3. Dryopteris intermedia (Muhl.) A. Gray

D. spinulosa (O.F. Muell.) Watt var. intermedia (Muhl.) Underw.

Aspidium spinulosum (O.F. Muell.) Sw. var. intermedium
(Muhl.) D.C. Eat.

evergreen wood fern
Fig. 112 (a) frond; (6) fertile pinnule. Map 110.

Fronds up to 70 cm long or longer, winter green, forming a crown
at the end of the stout rhizome. Stipe scaly, particularly towards the
base, and one-quarter to one-third the length of the frond. Blade
oblong-ovate to lanceolate, more or less acuminate, twice pinnate-
pinnatifid, usually glandular, particularly near the bases of the
pinnae; pinnae at right angles to the rachis, lanceolate to triangular-
ovate; inner lower pinnules on the basal pinnae usually shorter than
the others. Indusium glandular.

Good field characters are the extremely lacy appearance, the
deep bluish green of the subevergreen fronds and, more particularly,
the short inner lower pinnules and the glandularity.

Cytology: n = 41 (Britton and Soper 1966*). This is an important
ancestral diploid species that has contributed genomes to D.
campyloptera andZ). carthusiana. Dryopteris intermedia has the same
genomes and the same phloroglucinol chemistry as two Old World
species, D. azorica (only in the Azores) and D. maderensis (Gibby and
Walker 1977).

Habitat: Moist woods, swamps, and bogs.

Range: Newfoundland to Ontario, west to Minnesota, south in the
United States to North Carolina, Tennessee, and Alabama.

Remarks: This extremely attractive plant is a characteristic species
of eastern North America. It is an important species for interspecific
hybrids, giving rise to two particularly abundant hybrids, D. X
triploidea and D. X boottii.

213

Fig. 1 12 Dryopteris intermedia; (a) frond, 1/2 x ; (6) fertile pinnule, 3 x

214

4. Dryopteris expansa (Presl) Fraser-Jenkins & Jermy

D. assimilis S. Walker

D. dilatata Am. auth. pro parte

D. austriaca Am. auth. pro parte

northern wood fern
Fig. 113 (a) basal portion of frond; (6) portion of fertile pinnule.
Map 111.

Fronds to 1 m long, winter green in the west, forming a large
more or less upright crown at the end of the stout erect or ascending
chaffy rhizome. Stipes usually shorter than the blade, with brownish
often dark-centred ovate-lanceolate scales. Blades broadly triangular
to ovate or broadly oblong, abruptly acuminate, twice pinnate-
pinnatifid to tripinnate; pinnae short-stalked, acuminate; basal
pinnae broadly ovate or triangular, inequilateral; lower basal pinnule
on each basal pinna closer to the second upper pinnule than to the
basal upper one. Ultimate segments of pinnae serrate; teeth
mucronate. Sori medial. Indusia glabrous, with some populations
finely glandular.

Field recognition in western Canada is simplified because this is
a common large plant with a lacy distinct aspect, and there are few
places where any other species could be confused with it. In eastern
Canada, it is another matter. There the plant is quite variable in
aspect and often it looks quite like D. campyloptera. Typically, D.
expansa is more erect, and the superior pinnules next to the rachis do
not overlap the rachis as much as in D. campyloptera. Also, the petiole
is often longer and the blade is usually more elongated, i.e., less
triangular than in D. campyloptera.

Cytology: n = 41 (Britton and Widen 1974*; Mulligan and Cody
1968*). Ancestral diploid, part parent to D. campyloptera.

Habitat: Cool moist woods and thickets.

Range: Circumpolar with gaps; southern Greenland, Labrador, and
northern Newfoundland to Algoma, Thunder Bay, and Rainy River
districts, Ont., western Alberta, British Columbia, Yukon, and
Alaska.

Remarks: This is a difficult plant to identify when it is found where
the distributions of D. expansa and D. campyloptera overlap.
Amateurs are likely to be unhappy with decisions that lean so heavily
on cytology. They could make a contribution here by comparing the
two species, one in the Lake Superior basin and the other on Prince
Edward Island for example, and by pointing out useful field characters
to make the separation simpler. The species is rare in the District of
Mackenzie (Cody 1979).

215

Fig. 1 13 Dryopteris expansa; (a) basal portion of frond, 1/3 x ; (b) portion of fertile
pinnule, 5x.

216

5. Dryopteris campyloptera Clarkson

D. spinulosa (O.F. Muell.) Watt. var. americana (Fisch.) Fern.
D. austriaca Am. auth. pro parte

Appalachian mountain wood-fern or eastern spreading
wood-fern
Fig. 114, basal pinna. Map 112.

Fronds to 65 cm long, deciduous, forming an arching crown at the
end of the stout chaffy rhizome. Stipes shorter than the blade; scales
light brown, attenuate. Blade not glandular, ovate to ovate-
triangular, tripinnatifid, or with the basal pinnae sometimes
tripinnate; pinnae short-stalked, broadly lanceolate, attenuate; basal
pinnae triangular, with the basal upper and lower pinnules remote
and the inferior 2-4 times as long as the superior; ultimate segments
oblong, obtuse, sharply toothed or cleft; teeth spinulose-tipped. Sori
medial. Indusia glabrous or rarely with a few glands.

This fern has the laciness of D. intermedia and the width and
stature of D. expansa. The species is most easily identified by the long
basal pinnules next to the stipe on the lowermost pinnae, which are
often remote from the superior pinnules opposite, and by its somewhat
triangular blade, short petiole, and spreading habit (less erect).
Unfortunately, some individuals intergrade annoyingly with the
ancestral parent, D. expansa (especially in the northern part of the
range of Z). campyloptera) ^ and with luxuriant plants of D. intermedia.

Cytology: n = 82 (Britton and Widen 1974*). Genomes EEII from D.
expansa and D. intermedia (Gibby and Walker 1977).

Habitat: Cool, rocky woodlands at sea level in the north, but
restricted to higher elevations in the south of its range.

Range: Eastern North America; southern Labrador, Newfoundland,
Nova Scotia, Prince Edward Island, New Brunswick, and southern
Quebec (not positively identified in Ontario as yet), south in the
United States in the Appalachian region, to Tennessee and North
Carolina.

Remarks: This large and graceful species is most abundant in
Canada in the cool maple and yellow birch woods of the Laurentian
Mountains north of Montreal, Gaspe Peninsula (base of Mont-Albert),
Que., Cape Breton, N.S., and sheltered valleys of Newfoundland.

6. Dryopteris carthusiana (Vill.) HP Fuchs
D. spinulosa (O.F. Muell.) Watt
Thelypteris spinulosa {O.F. Muell.) Nieuwl.
Aspidium spinulosum (O.F. Muell.) Sw.
spinulose wood fern

Fig. 115 (a) frond; (b) portion of fertile pinnule. Map 113.

217

Fig. 1 14 Dryopteris campyloptera; basal pinna, 4/5 x

218

Fig. 115 Dry opteriscarthuslana; {a) irond, 1/3 x ;(b) portion of fertile pinnule, 3 x

219

Fronds 30-80 cm long, forming a crown at the top of a stout
ascending rhizome. Stipes with ovate brown scales, particularly near
the base. Blades lanceolate, 10-20 cm wide, bipinnate or bipinnate-
pinnatifid; pinnules oblong, with spine-tipped teeth; blade and rachis
not glandular; inner lower pinnule of basal pinnae usually longer than
the next one to it. Sori round. Indusia not glandular.

The well-known species Dryopteris spinulosa, now with a
changed name (D. carthusiana) because of the rules of priority, has a
much reduced variation when such taxa as D. intermedia, D. X
triploidea, D. X uliginosa, and D. campyloptera are removed from con-
sideration. When it is compared with D. intermedia^ D. carthusiana is
less lacy and the blade is paler, more yellow green, and has less
divergent teeth. The fronds and indusia should be almost completely
devoid of glands.

Cytology: n = 82 (Britton and Soper 1966*; Cody and Mulligan
1982*). Genomes IIBB.

Habitat: Moist to wet woodlands, thickets, and streambanks.

Range: Circumpolar; in North America from Labrador to locally in
British Columbia, south in the United States to northern South
Carolina, Kentucky, Arkansas, and Missouri, west to Montana, Idaho,
and Washington.

Remarks: The populations in western Canada are rather small and
widely separated. If the broad distribution of this species in the world
is taken into consideration, the species must be an ancient
allotetraploid, and so its place of origin is obscure. We cannot state
whether it is of New World or Old World origin. It is rare in the
District of Mackenzie (Cody 1979).

7. Dryopteris filix-mas (L.) Schott

Aspidium filix-mas (L.) Sw.

Thelypteris filix-mas (L.) Nieuwl.

male fern
Fig. 116 (a) frond; (6) portion of fertile pinna. Map 114.

Fronds up to 1 m long or longer, forming a crown from a stout
ascending scaly rhizome. Stipe usually short, thickly covered with
long-attenuate pale brown scales and shorter setiform scales. Blades
lanceolate to lance-oblong, narrowed towards the base, acuminate,
dark green above; pinnae lance-linear; lower pinnae short and more
ovate-lanceolate; pinnules oblong, obtuse, crenate, or serrate. Sori
medial, usually only on the lower three-quarters of the pinnules on the
upper half of the frond. Indusia glabrous.

220

Fig. 1 16 Dryopteris filix-mas; (a) frond, 1/3 x ; (b) fertile pinnules, 7 x

221

The double taper to the fronds and the vegetative growth of the
plant, which produces a confused crown or patch, are characteristics of
this species. Some researchers note a superficial resemblance to D.
marginalis, but the sori are not submarginal and the plant is much
less leathery.

Cytology: fi = 82 (Britton and Soper 1966*). The European plants
have an ancestry from two diploids, D. oreades (abbreuiata) and D.
caucus ica.

Habitat: Rich woods and rocky slopes of valleys (chiefly on limestone
in eastern Canada).

Range: Circumpolar; in North America from southern Greenland,
western Newfoundland, Cape Breton Island, N.S., Gaspe Peninsula,
Que., Bruce, Grey, and Simcoe counties and Michipicoten Island in
Lake Superior, Ont., northern Saskatchewan, Waterton Lakes
National Park, Alta., and southern British Columbia, south in the
United States to Maine, Vermont, Michigan (rare in northeastern
United States), California, Arizona, and Texas, and more widespread
in the western mountains.

Remarks: The chemistry of the phloroglucinols, spores, and cytology
are all reasonably similar to those for the species in Europe. The
North American species would seem to be part of the broader
distribution of the Eurasian one. The fact that D. filix-mas crosses so
readily with D. marginalis suggests that these two species share a
very ancient relationship, i.e., perhaps there was a common ancestor
of both D. oreades and D. marginalis. Dryopteris filix-mas is rare in
Ontario (Argus and White 1977) and Alberta (Argus and White 1978).

8 . Dryopteris marginalis ( L . ) Gray

Thelypteris marginalis (L.) Nieuwl.

marginal shield fern
Fig. 117 (a) frond; (6) fertile pinnule. Map 115.

Fronds 25-60 cm long or longer, crowded to form a crown on the
stout ascending rhizome; lower part of the stipe covered with thin,
light brown lance-linear scales. Blades 9-20 cm wide or wider, dark
green above, gray green below, leathery, lanceolate to oblong-ovate,
bipinnate; pinnae lanceolate; pinnules oblong, entire to deeply lobed.
Sori situated near the margin. Indusia smooth, whitish, becoming
light brown at maturity.

The marginal or leathery wood fern is extremely familiar to
amateurs in eastern Canada. The leathery or spongy character of the
subevergreen fronds and the submarginal sori are easy and reliable
field characters.

222

Fig. 117 Dryopteris marginalis; (a) frond, 1/3 x ; (b) fertile pinnule, 4 x

223

Cytology: fi = 41 (Britton and Soper 1966*; Cody and Mulligan
1982*). This distinctive diploid species is not an ancestor of any of our
other species.

Habitat: Rocky woods and shaded ledges and occasionally in
swamps.

Range: Newfoundland, Gaspe, Que., Nova Scotia to Ontario, west to
Wisconsin, south in the United States to Georgia, Alabama, Arkansas,
Oklahoma, and Kansas.

Remarks: A characteristic species of open woods in eastern Canada,
it can flourish in somewhat drier locations than its relatives, although
it often grows intermixed with another frequent easterner, D.
intermedia. There are some named forms that are more dissected than
the typical plant.

9. Dryopteris goldiana (Hook.) Gray

Thelypteris goldiana (Hook.) Nieuwl.

Goldie's fern
Fig. 118 (a) frond; (b) fertile pinnule. Map 116.

Fronds up to 1 m long, crowded at the top of the stout ascending
rhizome; lower part of the stipe covered with dark brown to blackish
lance-acuminate scales. Blades ovate-lanceolate, 20-40 cm wide,
pinnate-pinnatifid; pinnae broadly oblong-lanceolate; pinnules
linear-oblong, usually crenulate or serrated on the margins. Sori
round, situated near the midrib.

Goldie's fern has long been considered "one of the very finest and
largest of the species in the Eastern States, being surpassed in these
respects only by the osmundas and the ostrich fern" (Eaton 1879). The
sides of the blade are parallel, and the blade narrows to an apex rather
abruptly, so that amateurs refer to it as "being choked in the head."
Some collectors have commented on the play of dark to bright green
when a clump is viewed from a short distance. Mature specimens are
easy to identify.

Cytology: n = 41 (Britton and Soper 1966*; Cody and Mulligan
1982*). Ancestral diploid, considered a part parent to both D. celsa
andZ). clintoniana.

Habitat: Ravines in rich, moist woods and bordering swampy woods.

Range: New Brunswick, southwestern Quebec and southern Ontario,
south in the United States to North Carolina, Kentucky, Missouri,
and Minnesota.

224

Fig. 1 18 Dryopteris goldiana; (a) frond, 1/3 x ; (b) fertile pinnule, 1 1/2 x .

225

Remarks: This fern was found by John Goldie near Montreal in 1818
and was described and named by Hooker. Goldie was on a field trip
that included walking from Montreal to Niagara Falls, then to
Pittsburg, and back to Montreal. Dryopteris goldiana is rare in
Ontario (Argus and White 1977).

10. Dryopteris cristata (L.) Gray

Thelypteris cristata (L.) Nieuwl.

crested wood fern
Fig. 119 (a) fronds; (6) portion of fertile pinna. Map 117.

Fronds 25-70 cm long, forming a crown at the top of the stout
ascending rhizome; fertile frond longer than the sterile frond. Stipes
with pale brown ovate-lanceolate scales. Blades linear-oblong to
narrowly lance-oblong, 6-15 cm wide, pinnate-pinnatifid; basal
pinnae short, triangular. Pinnae of fertile fronds turned at right
angles to the rachis. Pinnules oblong, obtuse, serrate. Sori round,
situated midway between the margin and midvein. Indusia glabrous.

Typical characteristics are the extremely narrow upright fertile
fronds, with pinnae that can be perpendicular to the ground, giving a
Venetian blind effect. Intergradations with D. clintoniana (in
southern Ontario and southwestern Quebec) are usually hybrids or
are poorly developed plants of that species. The narrow, glossy, sterile
leaves with much reduced basal pinnae are features to note and they
distinguish it fromZ). clintoniana.

Cytology: n = S2 (Britton and Soper 1966*; Cody and Mulligan
1982*). Genomes LLBB.

Habitat: Thickets and wet woods to boggy or swampy open ground.

Range: Newfoundland to southeastern British Columbia, south in
the United States to North Carolina, Tennessee, disjunct in Nebraska,
and local in Idaho, Montana, and southeastern British Columbia;
Europe.

Remarks: A characteristic species, often occurring in small numbers
in Alnus thickets and sphagnum edges of lakes. It rarely makes solid
patches as doesD. intermedia or some of the other species of the genus.
Dryopteris cristata is rare in Alberta (Argus and White 1978).

226

Fig. 1 19 Dryopteris cristata; (a) fronds, 1/3 x ; (b) portion of fertile pinna, 1 1/2 x .

227

1 1 . Dryopteris clintoniana (DC. Eat.) Dowell

Dryopteris cristata (L.) Gray var. clintoniana (D.C. Eat.)

Underw.
Thelypteris cristata (L.) Nieuwl. var. clintoniana (D.C. Eat.)

Weath.
Clinton's wood fern
Fig. 120 (a) frond; (6) fertile pinnules. Map 118.

Fronds 30-80 cm long or longer, forming a crown at the top of the
stout ascending rhizome; fertile and sterile fronds similar. Stipes
scaly at the base; scales darker and shining at the middle. Blades
lanceolate, up to 20 cm wide, pinnate-pinnatifid; basal pinnae little
reduced, gradually acuminate at the apex; pinnae oblong-lanceolate,
acuminate; segments united by a narrow wing, oblong, obtuse,
incurved serrate or biserrate, with subspinulose teeth. Sori medial.
Indusia glabrous.

The long and relatively broad fronds without dimorphism are
characteristic of this species. Difficulties in identification arise
mainly from poorly developed plants. Individual fronds on a plant
vary a great deal, and often there will be only one or two large fronds
present.

Cytology: n ^ 123 (Britton and Soper 1966*). Genomes LLBBGG.
Considered to have arisen from a cross of D. cristata X goldiana.

Habitat: Swamps and rich wet woods.

Range: New Brunswick, southern Quebec, and southern Ontario,
south in the United States to Maine, Pennsylvania, New Jersey, Ohio,
and northwest Indiana.

Remarks: The species is common only in a limited area of central
southeastern Canada — in Ontario, south of the Precambrian Shield.
Canadian plants, which are on the northern edge of the distribution of
the species, are smaller and have fewer segments per pinna than those
from farther south.

Hybrids of Dryopteris

There are some well-known interspecific hybrids in Dryopteris
(Montgomery 1982), and as many as 31 different hybrid combinations
have been mentioned in the literature for temperate North America.
In Canada, we know of no hybrids of the western D. arguta, but even
so, we are left with 10 species that in theory could give rise to 9-8-7-
6-5-4-3-2-1 = 45 separate hybrids. The number known in Canada is
much less than that, approximately 16 or 17, and commonly occurring
hybrids are very few.

228

Fig. 120 Dryopteris clintoniana; (a) frond, 1/3 x ; (b) fertile pinnules, 1 1/3 x .

229

In morphology, hybrids usually have aborted spores and possess
characteristics from each parent.

Since D. marginalis is such a distinctive species, with its
leathery fronds, submarginal sori, and deep bluish green color,
hybrids with this species are most easily recognized and will be
considered first.

D. campyloptera X marginalis occurs very rarely in Virginia and
Pennsylvania.

D. carthusiana X marginalis (D. X pittsfordensis Slosson) occurs
rarely in Ontario, New England, south to West Virginia, Michigan,
and Wisconsin.

D. clintoniana X marginalis [D. X burgessii Boivin) occurs
infrequently in Quebec, Ontario, New Hampshire to Michigan, south
to New Jersey and Pennsylvania.

D. cristata X marginalis (D. X slossonae Wherry) occurs rarely
in New Brunswick, Ontario, west to Wisconsin, south to Virginia and
Ohio.

D. expansa X marginalis occurs rarely in Michigan.

D, filix-mas X marginalis is abundant within the range of D.
filix-mas in Ontario, Vermont, New York, and Michigan.

D. fragrans X marginalis (D. X algonquinensis Britton) is
known only from the type locality in Algonquin Park, Ont.

D. goldiana X marginalis {D. X neo-wherryi Wagner) occurs
rarely in Ontario, New England, south to West Virginia and North
Carolina, west to Illinois and Arizona.

D. intermedia X marginalis occurs rarely in Ontario, Vermont,
south to Virginia, Indiana, and Michigan.

Another good hybridizer is D. intermedia, which is noted for its
subevergreen fronds, lacy texture, and very particularly the glandular
indusia. Hybrids with this species are glandular and show some
influence of the mentioned characteristics of finely dissected blades
and subevergreenness.

D. campyloptera X intermedia occurs very rarely in Virginia,
Pennsylvania, and North Carolina.

D. carthusiana X intermedia (D. X triploidea Wherry) is
Canada's most common hybrid, occurring from Nova Scotia to
northwestern Ontario (R.M. Tryon and Britton 1966). It is also found
in New England, south to North Carolina, west to Kentucky and
Minnesota.

D. clintoniana X intermedia (D. X dowellii Wherry) occurs
frequently in Ontario, New Hampshire to Michigan, south to New
Jersey and Pennsylvania.

D. cristata X intermedia {D. X boottii (Tuckerm.) Underw.) is
probably Canada's second most frequent hybrid, occurring from
Newfoundland to Ontario. It is also found in Wisconsin, south to
Virginia, West Virginia, and Tennessee.

D. expansa X intermedia occurs rarely in Michigan.

230

D. fragrans X intermedia is cited by R.M. Tryon (1942). It was
rejected by W.H. Wagner and Chen (1965) and is reported only from
Sibley Peninsula, Ont.

D. ftlix-mas X intermedia is unknown but should be searched for
where the ranges coincide.

D. goldiana X intermedia extends from New England to
Michigan and Ohio.

Since 11 of the 16 crosses in Canada have been considered, the
other five will be shown without all the other possibilities, as follows.

D. clintoniana crosses:

D. carthusiana X clintoniana {D. X benedictii (Farw.) Wherry).
Occurs rarely in Ontario, New Hampshire, and Vermont, south to
Virginia and west to Michigan.

D. clintoniana X cristata. Occurs frequently in Ontario, New
England to Michigan, south to Pennsylvania.

D. clintoniana X goldiana. Occurs rarely in Ontario, New
Jersey, New York, Pennsylvania, Michigan, and doubtfully in
Tennessee.

Miscellaneous:

D. carthusiana X cristata {D. X uliginosa (A. Br.) Druce).
Occurs rarely in Ontario, Maine to Virginia and West Virginia, west
to Minnesota and North Dakota.

D. campyloptera X expansa. Rare; known only from Gaspe West
County, Que.

D. carthusiana X goldiana. Very rare; Vermont(?).

D. filix-mas X goldiana. Rare; reported from Vermont.
Note: In the United States there are also the D. celsa hybrids, so that
the total is approximately 30-31 of known or reported hybrids.

6. Gymnocarpium Newm. oak fern

Small ferns with fronds delicate, glabrous, or glandular, arising
singly from slender rootstocks. Sori round. Indusium absent. Veins
free, simple, or forking.

This is a small genus of perhaps fewer than 10 species, with its
greatest diversity in Asia. Sarvela (1978) lists 17 species names, and
in his synopsis and key arrives at six species for the genus. These
have been considered previously under a very large number of generic
names, e.g., Polypodium, Dryopteris, Lastrea, Phegopteris, Thelypteris,
Currania, and Carpogymnia, to mention some. This is a clear
indication of the uncertain affinity of the genus. There seems general
acceptance now of the genus name Gymnocarpium.

In the Canadian flora there is one common species, G. dryopteris,
the oak fern, and two others that are much less frequent, G.
robertianum, the limestone oak fern, and G. jessoense ssp. paruulum,
the Nahanni oak fern.

231

A. Blades membranous, with the two lower divisions nearly as long

as the terminal one; rachis essentially glabrous

1 . 6. dryopteris

A. Blades firm and somewhat stiff, with the two lower divisions

about half as long as the terminal one; rachis at least at the

junction of the second and third pinna pair densely glandular.

B. Proximal basal pinnules of the lowermost pair of pinnae

usually much longer than the corresponding upper pinnule;

lobes or pinnules in central part of the basal pinnae at right

angles to the rachis; upper surface of blade moderately

glandular; rachis and lower surface of blade densely

glandular 3. 6. robertianum

B. Proximal basal pinnules of the lowermost pair of pinnae
usually only slightly longer than the corresponding upper
pinnules; lobes or pinnules in central part of the basal
pinnae oblique to the rachis, or curved; upper surface of
blade glabrous; rachis and lower surface of blade

moderately densely glandular

2. 6. jessoense ssp. parvulum

1. Gymnocarpium dryopteris (L.) Newm. ssp. dryopteris

Dryopteris disjuncta Am. auth.

D. linnaeana C. Chr.

Thelypteris dryopteris (L.) Slosson

Carpogymnia dryopteris (L.) Love & Love

oak fern
Fig. 121 (a) frond; (6) fertile pinnule. Map 119.

Fronds up to 30 cm long or longer, arising singly from a slender
forking blackish rhizome. Blades glabrous, or almost so, triangular,
ternate; three divisions pinnate-pinnatifid; pinnules oblong, blunt.
Sori small, situated near the margin.

The oak fern is fairly common in Canada from the Atlantic to the
far West. Its small, delicate, triangular blades horizontal to the
ground and its bright lime to yellow green color are distinctive. Some
researchers see it as a miniature bracken, but bracken is too coarse
and its tissue too thick for such a comparison. The oak fern is
particularly striking in early spring, when the trees are beginning to
leaf out. The fronds unfold early in this species and they are smooth or
with only an occasional gland.

Cytology: n = 80 (Britton 1953*; Cody and Mulligan 1982*) eastern
Canada; ca. 4X (R.L. Taylor and Brockman 1966*), British Columbia,
which is the same chromosome number for this taxon as in Europe.

Habitat: Cool rocky woods, swamp margins, and shaded slopes.

232

Fig. 121 Gymnocarpium dryopteris ssp. dryopteris; (a) frond» 1/3 x; (b) fertile
pinnule, 1 1/2 x.

233

Range: Circumpolar; in North America from Newfoundland to
British Columbia, the Yukon and Alaska, south to Virginia,
Michigan, and Wisconsin.

Remarks: The species makes attractive patches in moist hollows in
open woods and is generally abundant over most of forested Canada,
particularly in boreal woods.

1 . 1 Gymnocarpium dryopteris (L.) Newm. ssp. disjunctum (Rupr.)
Sarvela

Dryopteris disjuncta (Rupr.) Morton

western oak fern
Map 120.

Differs from ssp. dryopteris in being more robust (to 50 cm long).
Blades tripinnate rather than bipinnate.

This subspecies has few characters to delineate it from ssp.
dryopteris, other than those given above. It is a basic diploid entity,
and if the definition of a biological species is strictly applied, ssp.
disjunctum should be accorded specific rank. We agree with W.H.
Wagner (19666) that this would be a mistake because of our lack of
knowledge of the small-spored plants in Alaska and neighboring
USSR. Also, it should be noted that both northern populations of this
subspecies and those at higher altitudes would then agree with the
description of ssp. dryopteris.

Cytology: n = 40 (R.L. Taylor and Mulligan 1968*). This is a basic
diploid entity, x = 40.

Habitat: Moist woods and rocky slopes.

Range: British Columbia, Washington, Oregon, Idaho, and Alaska;
Sakhalin Island and Kamchatka.

Remarks: Plants that grow under optimum conditions such as those
in the MacMillan Memorial Grove, Vancouver Island, B.C., are easy to
identify as to subspecies. Plants from higher elevations are quite
another matter. One sometimes hears the generalization that
polyploids are larger. In this case, the basic diploid is larger.

2. Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvulum Sarvela

G. continentale (Petrov) Pojak

Nahanni oak fern
Fig. 122 (a) frond; (b) fertile pinnule. Map 121.

234

^F^

= b

Fig. 122 Gymnocarpium jessoense ssp. parvulum; (a) frond, 1/2 x; (b) fertile
pinnule, 3 x.

235

Fronds up to 30 cm long, arising singly from a slender forking
blackish rhizome. Blades glandular, narrowly triangular, bipinnate-
pinnatifid; proximal basal pinnules usually only slightly longer than
the corresponding upper pinnules; lobes of pinnules of central part of
basal pinnae oblique to the rachis or curved. Sori small, situated near
the margin.

This species and subspecies are new to Canada's flora. The
subspecies was described by Sarvela (1978), and the type chosen came
from below Virginia Falls, in Nahanni National Park, District of
Mackenzie.

The blade and rachis are glandular, and so previous workers
have associated this subspecies with G. robertianum. It differs from
that species in being a smaller, more slender species of cool, moist,
calcareous cliffs. The pinnae are usually curved upwards, and the
pinnules have a definite curve outwards, rather than being
perpendicular to their axis (Sarvela et al. 1981). Subspecies jessoense
has a widespread distribution, entirely in Eurasia (Sarvela 1978).

Cytology: n = SO (Sarvela et al. 1981*).

Habitat: Limestone or basic rock cliffs and moist, rocky woods.

Range: Upper Great Lakes in Ontario to British Columbia and
Alaska, south to Minnesota and Wisconsin; northern Eurasia. The
records for this species in Atlantic Canada are from very old sheets
and should be verified from new collections.

Remarks: We have called this new subspecies the Nahanni oak fern,
to highlight the fact that the type was collected there and because it
has a generally northern distribution on cool, moist, calcareous sites.

3. Gymnocarpium robertianum (Hoffm.) Newm.

G. dryopteris (L.) Newm. var. pumilum (DC.) Boivin

Dryopteris robertiana (Hoffm.) C. Chr.

Carpogymnia robertiana (Hoffm.) Love & Love

limestone oak fern
Fig. 123, frond. Map 122.

Fronds up to 40 cm long, arising singly from a slender blackish
rhizome. Blades glandular, triangular, bipinnate-pinnatifid;
proximal basal pinnules of the lowermost pair of pinnae usually much
longer than the corresponding upper pinnule; lobes or pinnules in the
central part of the basal pinnae at right angles to the rachis. Sori
small, situated near the margin.

This species has been recognized for a long time and, for North
America at least, has always been considered a rare plant. Boivin
(1962) almost alone believes that it does not merit specific rank. The
pronounced glandularity of the blades, including the upper surface,
and the long triangular shape of the blade with the pinnules at right
angles are features to notice.

236

Fig. 123 Gymnocarpium robertianum; frond, 1/2 x

237

Cytology: n = 80 (Sarvela et al. 1981*).

Habitat: Moist calcareous ledges, limestone paving, cliffs, and rocky
woods.

Range: Newfoundland to Ontario, south to Minnesota; Europe.

Remarks: This species was already rare in North America before G.
jessoense ssp. paruulum was segregated; consequently, even fewer
localities are known now for this species and they are all in eastern
Canada. Rare in Ontario (Argus and White 1977).

Hybrids of Gymnocarpium

W.H. Wagner (19666) was the first to draw attention to "the
apparent cross of G. dryopteris and G. robertianum" and he named
this new apomictic species G. heterosporum. The plants were triploid.
Sarvela (1978) gave this species the hybrid designation G. X hetero-
sporum W.H. Wagner and said the cross was G. jessoense X robertia-
num. Sarvela (1978) also described a new hybrid, G. X intermedium
(G. dryopteris X jessoense). Studies by Pryer (1981) indicate that G.
X intermedium is a frequent hybrid at sites where G. jessoense ssp.
paruulum grows, but instead of being tetraploid as one would expect, it
is triploid. Accordingly, she concluded that one parent was G.
dryopteris ssp. disjunctum (2x) and the other was G. jessoense ssp.
paruulum (4x). All Gymnocarpium hybrids are recognized by their
mostly aborted spores. However, few large spherical spores that do
germinate are produced, and we believe that the hybrids are able to
propagate themselves apomictically in this way, although no one has
raised mature plants yet. If we are right, it should be noted that these
hybrids are then much more difficult to study than Dryopteris hybrids,
because they could have been formed far away from their present
location and at quite a different time. Gymnocarpium X intermedium
is found in very large colonies north of Lake Superior. It can greatly
outnumber G. jessoense ssp. paruulum at some locations.

Another hybrid of Gymnocarpium was described by Sarvela
(1980) as G. dryopteris ssp. X brittonianum. This is considered to be
G. dryopteris ssp. disjunctum X ssp. dryopteris. It is an easy hybrid to
identify and seems to be of frequent occurrence. Its hybrid nature is
evident again by its mostly aborted spores, and it has a larger stature
than ssp. dryopteris. The parents are glabrous, as is the hybrid.
Again, Pryer (1981) believes that the few large spores that are present
are able to perpetuate this hybrid as an apomictic species. It has been
identified as occurring from coast to coast in Canada (Sarvela 1980;
Pryer 1981).

One additional hybrid might be mentioned, G. X achriosporum
Sarvela (G. dryopteris X robertianum) . Hybrids of G. robertianum
seem to be of rare occurrence. The type is referable to Swedish

238

material, and only two collections in North America have been
ascribed to this combination. These are from Chicoutimi and Gaspe,
both in Quebec (Sarvela 1981). No material has been examined
cytologically, but since both parents are tetraploid, the hybrid should
be tetraploid.

Gymnocarpium X heterosporum is known only from the type
material in North America, but was ascribed also to one locality in
Finland (Sarvela 1978). It is not known in Canada (Sarvela 1980).

7. Tbelypteris Schmidel

Small to medium-sized ferns with more or less pubescent fronds
arising from a slender (stout in T. limbosperma) rhizome. Veins free,
simple, or forking. Sori small, round. Indusia reniform or horseshoe-
shaped, attached at the sinus.

It is difficult to estimate the number of species in the genus
Thelypteris at this time because of the large number of segregate
genera that are being recognized in both the Old World and the New
World. The number in the world with affinities to this genus is large
(approximately 800). We have recognized five species in the genus for
Canada. Other researchers might include these species in as many as
four genera. The five Canadian species, however, have features in
common that can be identified by the amateur. They all form upright
colonies or patches rather than neat individual "shuttlecocks," and
none is as finely divided as Dennstaedtia punctilobula or Athyrium
filix-femina, for example. All, except T. limbosperma, are roughly in
the same size range of 20-60 cm and all are deciduous. In general, the
blades are quite soft and thin in texture, unless in the full sun, and the
plants wilt rapidly when picked.

There is one species in western Canada, T. nevadensis, another
in both the West and isolated in the East, T. limbosperma, and three
in the East, T. noueboracensis, T. palustris, and T. simulata.

An indication of the diversity of Canada's five species is
illustrated by their chromosome numbers, which range from 27 to 64.
In other words, the situation is quite unlike the one for Dryopteris,
where the basic diploids are all derived from the number 41.

A. Lower pinnae gradually decreasing in size; lowermost pinnae

often very much decreased.

B. Rhizome short and stout 1. T. limbosperma

B. Rhizome slender and elongate.

C. Pinnules ciliate (eastern) 3. T. noveboracensis

C. Pinnules not ciliate (western) 2. 7". nevadensis

A. Lower pinnae only slightly if at all smaller.

D. Fronds dimorphic; lateral veins of pinnules of sterile fronds
mostly forking; glandular dots on undersurface of pinnules
absent 4. T. palustris var. pubescens

239

D. Fronds similar; lateral veins of pinnules of sterile fronds not
forked; glandular dots on undersurface of pinnules present
5. T. simulata

1. Thelypteris limbosperma (All.) HP. Fuchs
T. oreopteris (Ehrh.) Slosson
Dryopteris oreopteris (Ehrh.) Maxon
Oreopteris limbosperma (All.) Holub
mountain fern

Fig. 124 (a) frond; (6) fertile pinnule; (c) sorus. Map 123.

Fronds up to 1 m in length, tufted at the end of a short thick more
or less ascending rhizome. Stipes and rachis scaly. Blades lanceolate,
elongate, abruptly acuminate; lower pinnae reduced, pinnate-
pinnatifid, triangular; middle and upper pinnae linear-lanceolate,
tapering at the tip; pinnules oblong, obliquely set, blunt or subacute,
entire or somewhat wavy margined; margins of pinnules slightly
inrolled. Sori situated near the margins. Indusia glandular.

Thelypteris limbosperm,a was transferred to Oreopteris
limbosperma by Holub (1969). We are aware of the shuffling of the
genera and of the new interpretations that are occurring (Holtum
1971). For Canada's limited flora, however, it seems more sensible to
group our few species into a common genus at this time.

This is a taller species than the others, with leaf margins often
recurved and finely hyaline-papillose. It is aromatic when crushed,
and therefore in Great Britain it is referred to as the lemon-scented
fern. (Jermyetal. 1978)

Cytology: n = 34 (T.M.C. Taylor and Lang 1963*) as in Europe.

Habitat: Margins of creeks and runnels in rocky woods, outcrops, and
crevices of cliffs to at least 700 m.

Range: Coastal in Alaska and British Columbia, but occurs inland in
the Cascade Mountains, Wash.; disjunct in Gros Morne National Park
in western Newfoundland (Bouchard and Hay 1976); Eurasia.

Remarks: The discovery of this fern in Gros Morne National Park,
Nfld., was most interesting, even if it poses a problem regarding
whether it is a western disjunct species there or whether it has its
affinities with the populations in Europe (Bouchard et al. 1977).

2. Thelypteris nevadensis (Baker) Clute
T. oregana (C. Chr.) St. John
Dryopteris nevadensis (Baker) Underw.

Fig. 125 (a) fronds; (b) fertile pinna. Map 124.

240

Fig. 124 Thelypteris limbosperma; (a) frond, 1/3 x ; (b) fertile pinnule, 8 x ; (c) sorus,

8x.

241

Q/1 -1

^a^mi

xJXJ J-^'-^^^

Fig. 125 Thelypteris nevadensis; (a) fronds, 1/2 x ; (b) fertile pinna, 2 x

242

Fronds 20-60 cm long or longer, tufted at the end of the slender,
horizontal rhizome. Blades elliptic-lanceolate, 5-12 cm wide, long
attenuate; lower pinnae much reduced, pinnate-pinnatifid; pinnae
linear to linear-lanceolate, acuminate to caudate, somewhat hairy on
the midveins; pinnules oblong, blunt, obliquely set, entire or slightly
toothed, resin-dotted on the lower surface. Sori round, situated near
the middle. Indusia horseshoe-shaped, glandular.

This species can readily be distinguished from the only other
western species, T. limbosperma, by its more delicate fronds and
slender rhizome.

Cytology: n = 21 (T.M.C. Taylor and Lang 1963 as n = 26-27*; A.R.
Smith 1971; A.F. Tryon and R.M. Tryon 1974).

Habitat: Rocky banks of streams.

Range: In British Columbia known in a single locality (Sooke River,
Vancouver Island), southward in the foothills and middle altitudes to
central California.

Remarks: This species is very closely related to T. noueboracensis. A
comparative study that included the Asiatic vicariads (A.F. Tryon and
R.M. Tryon 1974), together with attempts at hybridization, would be
very interesting. It was placed in Parathelypteris by Ching (1963).

3. Thelypteris noveboracensis (L.) Nieuwl.

Dryopteris noueboracensis (L.) Gray

New York fern
Fig. 126 (a) fronds; (6) fertile pinnule. Map 125.

Fronds 25-55 cm long, arising from a slender rhizome. Blades
elliptic to elliptic-lanceolate, 9-15 cm wide; lower pinnae very
reduced, pinnate-pinnatifid; pinnae oblong to oblong-lanceolate,
somewhat hairy on the rachis and veins; pinnules oblong, somewhat
blunt. Sori round, situated near the margin. Indusia glandular-
ciliate.

This is a characteristic fern of sandy, acid glades and roadsides in
southeastern Canada. The light green patches are soon easy to
identify by noting the double taper to the blades of a frond. The last
characteristic should be used with caution, however, because some
small plants of the lady fern have this same general shape. The New
York fern has round sori, whereas those oi Athyrium filix-femina are
elongated.

Cytology: n ^ 21 (Britton 1964*; A.F. Tryon and R.M. Tryon 1973,
1974).

243

Fig. 126 Thelypteris noveboracensis; (a) fronds, 1/3 x ; (b) fertile pinna, 5 x

244

Habitat: Moist woods, thickets, and swamps, chiefly in moderately
acid soil.

Range: Eastern North America, Newfoundland to Ontario,
Michigan, and Illinois, south to Georgia, Alabama, Mississippi, and
Arkansas.

Remarks: For most ferns (Osmunda excepted), this species has a low
chromosome number. It is the lowest number for the thelypterids. See
also remarks under T. nevadensis.

4. Thelypteris palustris Schott var. pubescens (Lawson) Fern.

T. thelypterioides sensu Holub

Dryopteris thelypteris (L.) Gray var. pubescens (Lawson) Nakai

marsh fern
Fig. 127 (a) fronds; (6) fertile pinna. Map 126.

Fronds 20-60 cm long or longer, arising from an elongate
rhizome. Blades lanceolate, 7-15 cm wide, pinnate-pinnatifid; pinnae
linear-lanceolate; pinnules oblong, blunt, dimorphic. Fertile fronds
usually longer than the sterile fronds; pinnules somewhat thicker and
inrolled. Rachis and blade minutely pubescent and sometimes
glandular. Sori round, situated about halfway between the margin
and midvein, at maturity partly covered by the inrolled margin.
Lateral veins of pinnules of sterile fronds mostly forking.

Cytology: ;i = 35 (Britton 1953*; A.F. Tryon and R.M. Tryon 1973).
This is the same chromosome number as for the species in Europe.

Habitat: Marshes, swamps, wet thickets, bog margins, and ditches.

Range: Eastern North America, southern Newfoundland to south-
eastern Manitoba, south to Georgia, Tennessee, and Oklahoma,
Japan, and northeastern Asia; var. palustris occurs in Eurasia.

Remarks: This is a common fern in much of southeastern Canada. It
is not a particularly graceful species and is so soft and fragile that it is
often rather windblown and distorted — so much so, that some
researchers consider it either weed-like or an immature form of a
larger species. The reflexed margins of the segments of the blades of
the fertile fronds suggested to early writers the shapes of snuff boxes.

The marsh fern is suffering the vicissitudes of nomenclatural
changes. If the Canadian plant is considered as a variety or subspecies
of the species in Europe, it may be considered as var. pubescens
(Lawson) Fernald. Recently, workers in Europe have taken up the
name T. thelypterioides (Michx.) Holub, but A.F. Tryon et al. (1980)

245

Fig. 127 Thelypteris palustris var. pubescens; (a) fronds, 1/2 x ; (b) fertile pinna, 3 x

246

say that this is a mistake, and that the familiar name T. palustris
should be used. Variety pubescens is rare in Newfoundland and
Manitoba.

5. Thelypteris simulata (Davenp.) Nieuwl.

Dryopteris simulata Davenp.

Massachusetts fern
Fig. 128 (a) frond; (b) fertile pinnule. Map 127.

Fronds 20-50 cm long or longer, arising from an elongate
rhizome. Blades oblong-lanceolate, 7-15 cm wide, tapering at the tip,
pinnate-pinnatifid; pinnae oblong-lanceolate; fertile pinnae long-
acuminate; pinnules oblong, obtuse, flat, or sometimes slightly
inrolled. Upper surface of pinnules strigose. Indusia and lower
surface of the pinnules glandular dotted.

This species is sufficiently similar to both the New York fern and
the marsh fern to cause identification problems. In the Massachusetts
fern the lower pinnae are noticeably narrowed at their base next to the
rachis (pinched in) and are usually retrorse (angled down). The veins
in both the fertile and sterile fronds are unbranched. The marsh fern
has veins in the sterile fronds mostly forking.

Cytology: n = 64 (A.F. Tryon and R.M. Tryon 1973).

Habitat: Moist woods and boggy thickets in intensely acid situations
often in association with Sphagnum.

Range: Eastern North America from southwestern Nova Scotia,
south to Virginia, and extending locally inland to southwestern
Quebec, southeastern Ontario, central New York, and Maryland; also
disjunct in the Driftless Area of Wisconsin (mapped by A.F. Tryon and
R.M. Tryon 1973).

Remarks: At one time T. simulata was considered as a possible
hybrid derivative of T. noveboracensis and T. palustris, but as pointed
out by A.F. Tryon and R.M. Tryon (1973), there are vicariads in Asia,
and all the species of the world of the same general affinity should be
studied and carefully compared. Love and Love (1976) created the
genus Wagner iopteris for this species, but this seems premature.
Thelypteris simulata is rare in Quebec and Ontario.

8. Phegopteris Fee

Small to medium-sized ferns with more or less pubescent fronds
arising singly from elongate, slender, horizontal, scaly rhizomes.
Veins free, simple, or forking. Sori small, round. Indusium absent.

247

Fig. 128 Thelypteris simulata; (a) frond, 1/3 x ; (b) fertile pinnule, 3 x

248

This genus has progressed from being part oi Dryopteris s.L, in
Christensen's time, to a Thelypteris, and now to the segregate genus
Phegopteris, which comprises only three or four species in the world.
The basic chromosome number x is 30.

A. Wings of rachis extending down to the lowest pinna-like
divisions; blades broadly triangular 1. P. hexagonoptera

A. Wings of rachis not extending down to the lowest pinnae; blades
narrowly triangular; lowest pair of pinnae usually projected
downward and forward 2. P. connect! I is

1 . Pbegopteris hexagonoptera (Michx.) Fee

Dryopteris hexagonoptera (Michx.) Christens.

Thelypteris hexagonoptera (Michx.) Weatherby

broad beech fern
Fig. 129 (a) frond; (6) fertile pinnule. Map 128.

Fronds 30-60 cm long or longer. Blades broadly triangular, 15-
30 cm wide or wider, about as broad as long, tapering to the top,
pinnate-pinnatifid; middle and upper pinna-like divisions lanceolate;
lower pinna-like divisions unequally ovate to lanceolate-ovate, not
projected forward; all divisions connected by a wing; segments,
particularly those of the lower pinnae, often deeply pinnatifid. Stipe
naked except at the base; rachis not chaffy or with almost colorless
scales. Rachis and veins minutely glandular puberulent. Sori small,
near the margin.

Although the other species in Canada is called P. connectilis, it is
P. hexagonoptera that has all the divisions of the blade connected to
the rachis, including the basal pair. In shape, the blade is more
broadly triangular in the broad, or southern, beech fern than in the
long beech fern. The shape of the basal segments is unlike that in P.
connectilis, being widest in the middle and lobed again rather than
entire.

Cytology: n = 30 (Mulligan and Cody 1979*). A sexual basic diploid.

Habitat: Rich often rocky woods and wooded slopes.

Range: Southwestern Quebec and southern Ontario, south to Florida
and Texas.

Remarks: This species is very local even within its Canadian distri-
bution. It would seem to be perhaps Canada's only fern with a strictly
Carolinian distribution, compared with the distribution of Sassafras,
or black walnut, although some researchers might question even that
observation for the Quebec populations. Phegopteris hexagonoptera is
rare in Ontario (Argus and White 1977) and Quebec (Vincent 1981).

249

Fig. 129 Phegopteris bexagonoptera; (a) frond, 1/3 x ; (6) fertile pinnule, 3 x

250

2. Phegopteris connectilis (Michx.) Watt

P. polypodioides Fee

Dryopteris phegopteris (L.) Christens.

Thelypter is phegopteris (L.) Slosson

long beech fern
Fig. 130 (a) frond; (6) portion of fertile pinnule. Map 129.

Fronds 15-35 cm long or longer. Blades triangular, longer than
broad, 8-20 cm long or longer, 6-16 cm wide or wider, pinnate-
pinnatiiid, tapering to the tip; pinna-like divisions lance-acuminate,
with all but the lower pair (usually projected downward and forward)
connected by a wing. Pinnules oblong, rounded at the tip, more or less
hairy on both faces and on the rachis. Stipe and rachis with brown
scales. Sori small, situated near the margin.

Our more common representative, the long, or northern, beech
fern, was known as P. polypodoides for a number of years, but is now
called P. connectilis (Michx.) Watt. This species is known from coast to
coast in Canada and is soon familiar to amateurs in our northern
woods. The distinctive triangular blades with only the base pair of
pinna-segments free, and these projecting downward and forward, are
good field characters.

Cytology: "n" = 2n = 90 (Mulligan and Cody 1979*). The species
reproduces apogamously, and the chromosome number in the spores is
identical to that of the somatic cells of the mature plant. Because the
basic chromosome number for the genus is 30, this species is a triploid.
There are reports for a diploid sexual race of limited range in Japan
(Mulligan and Cody 1979), although for most of the distribution there
and elsewhere in the world, the number is 2n — 90.

Habitat: In Ontario often in soil on banks of rivulets or creeks. In
Newfoundland and British Columbia on moist rocky hillsides,
appearing from under large boulders or on moist rocky ledges.

Range: Circumpolar; in North America, Greenland, Labrador, and
Newfoundland to Alaska, south to North Carolina, Michigan, Iowa,
and Oregon.

Remarks: This is an attractive species of interesting form. It is not
as abundant as some species, but is of broad occurrence. Phegopteris
connectilis is rare in the District of Mackenzie (Cody 1979), the Yukon
(Douglas et al. 1981), Manitoba (White and Johnson 1980), Saskat-
chewan (Maher et al. 1979), and Alberta (Argus and White 1978).

Hybrids of Phegopteris

Mulligan et al. (1972) reported the occurrence in Quebec of a
hybrid between P. connectilis and P. hexagonoptera. Mulligan and

251

Fig. 130 Phegopteris connectilis; (a) frond, 2/3 x ; (b) portion of fertile pinnule, 6 x

252

Cody (1979) reported the chromosome numbers of 10 collections of this
hybrid from six different localities in Quebec, New Brunswick, and
Nova Scotia. The plants were tetraploid, based on x = 30, and had "m"
— 2n = 120 chromosomes, i.e., the spores and the somatic cells both
had 120 chromosomes, and the plants were apogamous and fertile.
They elected to refer this entity to P. connectilis because the tetraploid
hybrid was "not, however, always distinguishable from triploid plants
of P. connectilis."

9. Cys top ten's Bernh. bladder fern

Delicate medium-sized ferns, with bipinnate to tripinnate or
ternate fronds arising from short creeping rhizomes. Veins free.
Indusium hoodshaped, thin, and withering, attached at one side and
arching over the rounded sori.

The genus Cystopteris occurs worldwide and is a common
element of temperate floras in both the northern and southern
hemispheres. Blasdell (1963), who studied the species of the world,
recognized 10 species, five varieties, and six hybrids. The genus is
noted for much variation and cytogenetic complexity, with much
polyploidy (Lovis 1977). The basic chromosome number is 42, and
diploids to octaploids are known.

In the Canadian flora there is the highly distinctive northern
and alpine C montana; C. bulbifera, a basic diploid of southeastern
and central North America; the very widespread and abundant C
fragilis with its variety, mackayii; a diploid segregate species of C
fragilis, now known as C protrusa; and a derived allohexaploid
species, C laurentiana. We are recognizing five species, but because of
all the biosystematic work on this genus that is in progress, it seems
inevitable that further segregate species will be recognized. These
will be derived from the "C. fragilis complex."

A. Fronds ternate 1. C montana

A. Fronds bipinnate to tripinnate.

B. Fronds lanceolate and usually long-attenuate, often
bearing bulblets beneath; veins mostly ending in a notch . .

2. C bulbifera

B. Fronds lanceolate only, without bulblets.

C. Fronds scattered along a creeping rhizome

3. C protrusa

C. Fronds tufted from a short creeping rhizome.

D. Indusium glandular; veins ending both in teeth

and sinuses 5. C laurentiana

D. Indusium not glandular; veins usually ending in
the teeth 4. C fragilis

253

1. Cystopteris montana (Lam.) Bernh.
mountain bladder fern

P'ig. 131 (a) frond; (6) fertile pinnule. Map 130.

Fronds 40 cm long or longer, arising singly from a widely
creeping slender rhizome. Stipes usually longer than the blade,
sparsely chaffy. Blades ternate, broadly deltoid-ovate; two lower
divisions somewhat narrower and shorter than the upper; pinnae
bipinnate to tripinnate; ultimate segments ovate, rounded, often cleft
at the apex. Veins ending in the sinuses between minute teeth.
Similar in aspect to Gymnocarpium dryopteris, but more finely
dissected.

Cytology: n - S4 (Britton 1964*). This is a tetraploid and has the
same number as reported for Europe. Blasdell (1963) suggests from
spore sizes that diploids exist.

Habitat: Cool moist woods and rocky slopes, mainly in calcareous
places.

Range: Circumpolar; in North America in southern Greenland,
Quebec, north shore of Lake Superior, Alaska, British Columbia,
southwestern District of Mackenzie and western Alberta, and the
mountains of Colorado.

Remarks: This is a most attractive species. In general aspect it
might remind one of an oak fern, but the degree of dissection of the
blade is so fine that it is sometimes called a lace fern. It is widely
distributed, but of rare occurrence, and is considered a find when one
finally sees it. Cystopteris montana is rare in the District of
Mackenzie (Cody 1979), Ontario (Argus and White 1977), and
Saskatchewan (Maheretal. 1979).

2. Cystopteris bulbifera (L.) Bernh.
Filix bulbifera (L.) Und.
bulblet fern

Fig. 132 (a) frond; (6) fertile pinna. Map 131.

Fronds 30-80 cm long or longer, from a short stout rhizome.
Blades lanceolate and usually long-attenuate, 6-15 cm wide or wider
at the base; sterile blades usually shorter, bipinnate; pinnules oblong,
obtuse, pinnatifid to lobed. Veins mostly ending in a notch or sinus.
Dark green bulblets often borne on the underside of the rachis and
pinnules. Indusium minutely glandular.

Key field characters are the fine dissections, the veins ending in
notches, the bulblets on the axes, and the glandular indusia. It is a
distinctive species.

254

Fig. 131 Cystopteris montana; (a) frond, 1/3 x ; (b) fertile pinnule, 7 x .

255

Fig. 132 Cystopteris bulbifera; (a) frond, 1/3 x ; (6) fertile pinnule, 8 x

256

Cytology: n = 42 (Britton 1953*). This is a basic diploid species.

Habitat: Chiefly on calcareous rocks, in shaded ravines, and in moist
woods.

Range: Newfoundland to Ontario, Minnesota, and South Dakota,
south to Georgia and Texas. Doubtfully recorded from southeastern
Manitoba (Scoggan 1957).

Remarks: The long, graceful, arching fronds of mature plants are
distinctive. Smaller, shorter, and more triangular fronds must be
identified with care. In early spring, the bright maroon-colored stipes
are striking.

3. Cystopteris protrusa (Weath.) Blasdell

C. fragilis (L.) Bernh. var. protrusa Weath.
Fig. 133, sterile and fertile fronds. Map 132.

Fronds 20-45 cm long or longer, scattered along a creeping
rhizome; rhizome projecting beyond the current year's fronds. Stipes
greenish, or straw-colored, or pale brown. Blades lanceolate, 13-25 cm
long, 5-10 cm wide; sterile blades usually shorter, bipinnate; pinnules
sharply toothed, ovate-lanceolate to deltoid-ovate; lower pinnules
tapering to a stalk-like base. Veins mostly ending in a tooth or on the
unnotched margin. Indusium up to 0.5 mm long, shallowly or not at
all toothed at the apex.

Cystopteris protrusa may be distinguished from C fragilis var.
mackayii, with which it might be confused, by the long internodes on
the rhizome, the greenish or straw-colored stipes, the softer and more
easily wilting blade, which is more ample and more feathery, and the
lower pinnules, which taper to a stalk-like base.

Cytology: n = 42 (Britton unpublished*).

Habitat: In the shade of deciduous trees on rich river-bottom
benches.

Range: In Canada known from only two stations in southwestern-
most Ontario (Carolinian Zone); Ontario to Wisconsin, south to
Georgia and Mississippi. Further investigation of similar habitats
should yield more stations in Ontario.

Remarks: This basic diploid species has only recently been identified
in the Canadian flora, although its occurrence in northern Michigan
(W.H. Wagner and Hagenah 1956) had suggested that it should be
present in southwestern Ontario in similar sites.

257

Fig. 133 Cystopteris protrusa; sterile and fertile fronds, 1/2 x

258

4. Cystopteris fragilis (L.) Bernh. var. fragilis

Filix fragilis (L.) Und.

fragile fern
Fig. 134, frond. Map 133.

Fronds 10-35 cm long or longer, tufted from short creeping
rhizomes. Blades lanceolate, 3-8 cm wide or wider near the base,
bipinnate; pinnae pinnatifid to lobed, and at least the basal pinnules
varying from orbicular to triangular and rounded to the base. Veins
mostly ending in a tooth or on the unnotched margin. Indusium up to
1 mm long and more or less cleft at the apex.

A taxon in which the spores are rugose rather than echinate has
been called C. dickieana Sim or has been treated as a subspecies, a
variety, or a form of C. fragilis. This taxon is in need of further
investigation (Hagenah 1961).

Field characters are the translucent stipe, the veiny thin blades
with the veins going to the very tips of the teeth, and a smooth rachis
and indusium.

This highly variable species is known to have at least two levels
of ploidy (4jc and 6x) after C. protrusa (2x) has been segregated. There
is sufficient interest in this wide-ranging taxon, together with a great
deal of biosystematic work in progress, that one can expect further
segregate species to be described (Lovis 1977).

Cytology: n = 84 (Britton 1953*, eastern; Cody and Mulligan 1982*,
western). Both tetraploids and hexaploids are known in Europe.

Habitat: Sheltered crevices in cliffs, moist banks, and wooded talus
slopes.

Range: Circumpolar; in North America from Greenland to Alaska,
south to Virginia, Texas, and California.

Remarks: This is an abundant, attractive species, which one expects
to find on most moist, shady cliffs. It is present on both acidic and
basic rocks.

4.1 Cystopteris fragilis (L.) Bernh. var. mackayii Lawson
Fig. 135 (a) frond; (b) portion of fertile pinna. Map 134.

Similar to var. fragilis, but with the pinnules oblong to nearly
lanceolate and evenly wedge-shaped at the base, and the indusium
about 0.5 mm long and shallowly or not at all toothed at the apex.

This quite frequent variety of C. fragilis has a rather constant
morphology. Blasdell (1963) relegated the variety to a Cystopteris
diaphana X fragilis complex, thereby eliminating C fragilis from

259

Fig. 134 Cystopteris fragilis var. fragilis; fronds, 2/3 x

260

Fig. 135 Cystoptehs fragilis var. mackayii; (a) fronds, 2/3 x ; (b) portion of fertile
pinna, 4 x .

261

Newfoundland, New Brunswick, and Nova Scotia. A new
interpretation has recently appeared that will require a great deal of
biosystematic study to validate. Lellinger (1981) suggests that this
taxon is C. tenuis (Michx.) Desv., which has arisen from a cross of C.
protrusa (2x) X reeuesiana (2x), the latter a new species from Utah and
Arizona (Lellinger 1981).

Cytology: n = 84 (Britton unpublished*).

Habitat: Found in habitats similar to var. fragilis, but more often on
banks, on rotted logs, and in moist glades on soil.

Range: Nova Scotia to Ontario and Michigan, south to Virginia and
Missouri. In Canada it is a more southern and eastern variety than is
C. fragilis var. fragilis.

5. Cystopteris laurentiana (Weath.) Blasdell

C. fragilis (L.) Bernh. var. laurentiana Weath.

Laurentian fragile fern
Fig. 136, frond. Map 135.

Fronds tufted from a short creeping rhizome, up to 60 cm long or
longer. Stipes light brown to red-tinged. Blades ovate-attenuate, up
to 34 cm long and 13 cm wide; sterile blades usually shorter,
tripinnatifid; pinnules with the veins ending both in teeth and
emarginations. Indusium up to 1 mm in diameter, minutely
glandular.

This species combines the attributes of its presumed parents, C.
fragilis var. fragilis (4x) and C. bulbifera (2x). It is usually an upright,
vigorous plant of greater stature than a small or medium-sized C.
fragilis. The veins go both to the points and to the sinuses, the indusia
are glandular, and the spores are larger than those of either parent.

Cytology: n = 126 (Britton 1974*).

Habitat: Calcareous rock or slopes.

Range: Newfoundland, Nova Scotia, New Brunswick, Quebec, and
Ontario, west to Wisconsin. Reported as highly localized.

Remarks: This species of hybrid origin is comparable to Dryopteris
clintoniana and Polystichum setigerum, which are also considered to
be allohexaploids. The spores are large, regular, and freely produced.
The chromosome number and the morphology of the plants are
consistent with the presumed parentage.

Cystopteris laurentiana is usually considered a rare plant, but too
often is merely lumped into C. fragilis s.l. We found it to be quite

262

abundant on Manitoulin Island, Ont. It was reported by Argus and
White (1977) as rare in Ontario and by Maher et al. (1978) as rare in
Nova Scotia.

Hybrids of Cystopteris

We know of no confirmed hybrids for Canada, after C.
laurentiana and some of BlasdelTs C diaphana complexes are
removed from consideration. Now that C. protrusa has been confirmed
for the Canadian flora, C. bulbifera X protrusa is a further possibility
(Cranfill 1980), as are also various interspecific crosses such as C.
fragilis X montana, known in central Europe but not in North
America.

10. Athyhum Roth

Rather large ferns with large rootstocks; fronds pinnate to
tripinnatifid. Veins either simple or somewhat forked. Sori curved or
straight, borne along the veins. Indusium attached on one side of the
sorus or lacking.

This is a large genus of about 300 species, mainly of tropical
distribution. At one time all the species were included in the genus
Asplenium. More recently, there has been an attempt to split off such
small genera as Homalosorus and Lunathyrium (Love et al. 1977),
both of which have the same basic number, x — 40, as Athyrium.
Pending further study, we are following Copeland (1947) in
maintaining all our species in the genus Athyrium.

A. Fronds pinnate or bipinnatifid.

B. Fronds pinnate 3. A. pycnocarpon

B. Fronds deeply bipinnatifid 4. A. thelypterioides

A. Fronds bipinnate to tripinnatifid.

C. Ultimate segments of frond broad and close together;

indusium curved or horseshoe-shaped

2. A. filix-femina

C. Ultimate segments of frond very narrow and distant;

indusium lacking, with sori roundish

I. A. alpestre ssp. americanum

1 . Athyrium alpestre (Hoppe) Ry lands ssp. americanum (Butters)

Lellinger

A. alpestre (Hoppe) Rylands var. americanum Butters

A. alpestre (Hoppe) Rylands var. gaspense Fern.

A. distentifolium Tausch. ssp. americanum (Butters) Hulten

A. distentifolium Tausch. var. americanum (Butters) Boivin

A. americanum (Butters) Maxon
Fig. 137 (a) frond; (6) fertile pinnule. Map 136.

263

Fig. 136 Cystopteris laurentiana; frond, 1/3 x

264

4^1^

Fig. 137 Athyrium alpestre ssp. americanum; (a) frond, 1/2 x ; (b) fertile pinnule,
5x.

265

Fronds up to 80 cm long, forming large clumps from stout erect or
somewhat decumbent scaly rhizomes. Stipe short, sparsely scaly.
Blades glabrous, subcoriaceous, linear to oblong-lanceolate,
acuminate, twice pinnate-pinnatifid; pinnae narrowly deltoid,
gradually acuminate; pinnules oblong-lanceolate to narrowly
triangular; ultimate segments narrow and distant. Sori roundish,
lacking indusia.

The subalpine-alpine habitat of this species, its narrow and more
distant pinnules, and the absence of an indusium separate this fern
from the more common and widespread lowland A. filix-femina.

Cytology: n = 40 (Taylor and Lang 1963*).

Habitat: Moist, open, rocky subalpine slopes and alpine meadows.

Range: The species is circumpolar; ssp. americanum occurs in
Greenland, Newfoundland, and Gaspe Peninsula, Que., and in
western North America from southeastern Alaska through western
and southern British Columbia to California, Nevada, and Colorado.

Remarks: This is a wide-ranging species, comprising several
subspecies. Fernald (1928) described the plant from the Tabletop
Range, in the Gaspe Peninsula, as var. gaspense, but more recent
authors have included it in the North American ssp. americanum
(Scoggan 1978).

2. Athyrium filix-femina (L.) Roth

lady fern
Fig. 138 (a) frond; (6) fertile pinnule. Map 137 (var. cyclosorum).
Fig. 139 (a) frond; (6) fertile pinnule. Map 138 (var. michauxii).

Fronds up to 2 m long, tufted and erect-spreading from stout
chaffy erect or ascending rhizomes. Stipes brittle, scaly near the base.
Blades narrowly to broadly lanceolate, bipinnate to tripinnate; pinnae
lanceolate, acuminate to attenuate; pinnules somewhat lobed to
deeply toothed, blunt, or acute at the tip. Sori oblong to
horseshoe-shaped; indusia often toothed, ciliate, and attached by their
inner side to a veinlet.

Athyrium filix-femina is a very variable fern throughout its
circumpolar range. In western North America var. cyclosorum
(Ledeb.) Moore (var. sitchense Rupr.) is a tall coarse fern with
subcoriaceous fronds, and its ultimate segments have broad, bluntish
teeth. In eastern North America var. michauxii (Spreng.) Farw. is
shorter and more delicate; on the basis of differences in the frond, the
following forms of var. michauxii might be recognized, but in all cases
they gradate to one another (Butters 1917).

266

Fig. 138 Athyrium filix-femina var. cydosorum; (a) frond, 1/3 x ; (b) fertile pinnule,
3x.

267

Fig. 139 Athyrium filix-femina var. michauxii; (a) frond, 1/3 x; (b) fertile pinnule,
3x.

268

Forma michauxii has fronds dimorphic; fertile fronds contracted.
Pinnae 5-12 cm long. Pinnules 7-12 mm long, rounded, and only
shallowly lobed. Sori usually confluent at maturity.

Forma elatius (Link) Clute has fronds dimorphic; fertile fronds
contracted. Pinnae 10-20 cm long. Pinnules 12-25 mm long,
pinnatifid, acutish. Lower sori often strongly curved or
horseshoe-shaped; sori usually becoming confluent at maturity.

Forma rubellum (Gilbert) Farw. has fronds not dimorphic, larger
than the two preceding forms. Pinnules strongly toothed or pinnatifid.
Sori separate at maturity.

The lady fern might possibly be confused with some segregates of
the Dryopteris carthusiana complex, from which it can readily be
distinguished by its elongate, sometimes curved (rather than round)
sori, which are covered by an indusium attached on one side.

Cytology: var. cyclosorum, n = 40 (Mulligan and Cody 1968*); var.
michauxii, n = 40 (Britton 1953*; Cody and Mulligan 1982*).

Habitat: Meadows to open thickets, moist woods, and even swamps.

Range: The species is circumpolar; var. cyclosorum occurs in Alaska,
south to California; var. michauxii occurs in Labrador and
Newfoundland to northern Saskatchewan, south to Pennsylvania,
Ohio, Wisconsin, and Iowa.

Remarks: This species can be almost weedy in low-lying overgrown
meadows and roadside ditches. The var. sitchense, a synonym of var.
cyclosorum, was recorded by Fernald (1950) from northern
Newfoundland and the Gaspe Peninsula, Que. The various forms of
these plants are here included in var. michauxii. The group is in need
of further study.

3. Athyrium pycnocarpon (Spreng.) Tidestr.

A. angustifolium (Michx.) Milde

Diplazium pycnocarpon (Spreng.) Brown

Homalosorus pycnocarpon (Spreng.) Small ex Pichi Sermolli

narrow-leaved spleenwort
Fig. 140 (a) sterile and fertile fronds; (6) portion of fertile pinna;
(c) venation of pinna. Map 139.

Fronds up to 80 cm long or longer, forming a crown at the end of a
stout horizontal rhizome. Sterile blades lanceolate, 8-16 cm wide,
simply pinnate; pinnae long-acuminate, rounded to truncate at the
base, membranous; fertile pinnae lance-linear. Sori linear, situated
on the veins in crowded rows between the midrib and the margin.
Indusium opening along one side.

269

Fig. 140 Athyrium pycnocarpon; (a) sterile and fertile fronds, 1/3 x ; (b) portion of
fertile pinna, 4 x ; (c) venation of pinna, 2 x ,

270

The sterile fronds of the narrow-leaved spleenwort are somewhat
similar to the Christmas fern, but are much more delicate and thus
easily differentiated from the evergreen fronds of the latter.

Cytology: ;i = 40 (Britton 1964*; Cody and Mulligan 1982*).

Habitat: Deep, rich, moist woods and ravines.

Range: Southern Quebec and Ontario, south and west to Georgia,
Alabama, Minnesota, Iowa, and Kansas.

Remarks: This species is recorded as rare in Ontario by Argus and
White (1977), and it certainly is rare in Quebec also.

4. Athyhum thelypterioides (Michx.) Desv.

A. acrostic hoides (Sw.) Diels

Diplazium thelypterioides (Michx.) Presl

D. acrostic hoides (Sw.) Butters

Lunathyrium acrostichoides (Sw.) Ching.

silvery spleenwort
Fig. 141 (a) frond; (6) fertile pinnule. Map 140.

Fronds up to 100 cm long or longer, forming a crown at the end of
the horizontal rhizome. Blades lanceolate to elliptic-lanceolate, 8-
22 cm wide, pinnate-pinnatifid; pinnae long-tapering; segments
oblong, blunt, finely toothed. Sori straight or slightly curved, situated
on the veins between the midrib and margin. Indusia becoming
silvery at maturity, opening on one side or, if double, opening on both
sides.

The silvery spleenwort can readily be distinguished by the
straight or slightly curved sori, which toward maturity fill the
underside of the fertile fronds and become silvery in color.

Cytology: n = 40 (Britton 1964*; Cody and Mulligan 1982*).

Habitat: Rich woods, stream banks, shaded slopes, and rarely in open
thickets.

Range: Nova Scotia to southern Ontario, south and west to Georgia,
Alabama, Michigan, Wisconsin, and Minnesota.

Remarks: Athyrium thelypterioides is often associated with other
ferns of rich woodlands, such as Dryopteris goldiana. A report by
Macoun (1890) of this species growing at Current River, Lake
Superior (Thunder Bay, Ont.) was repeated by Scoggan (1978), but no
substantiating specimen has been found, nor have recent collectors
gathered it in that area.

271

Fig. 141 Athyrium thelypterioides; (a) frond, 1/3 x ; (b) fertile pinnule, 5 x

272

11. BLECHNACEAE

Ferns coarse, tufted from a short creeping rhizome or scattered
along a widely creeping rhizome. Fronds similar or dimorphic,
pinnatifid, pinnate, or bipinnatifid. Veins anastomosing to form
costal or costular areolae, and then free to the margin or forming
additional areolae. Sori elongate. Indusium opening on the costal
side.

A small family of terrestrial or occasionally climbing ferns.

A. Fronds evergreen, dimorphic, pinnate 1. Blechnum

A. Fronds deciduous, dimorphic or similar, pinnatifid or
bipinnatifid 2. Woodwardia

1. Blechnum L.

1. Blechnum spicant (L.) Roth

B. spicant (L.) Roth ssp. nipponicum (Kunze) Love & Love

B. doodioides Hook.

Lomaria spicant (L.) Desv.

Struthiopteris spicant (L.) Weiss

deer fern
Fig. 142 (a) sterile and fertile fronds; (6) portion of fertile pinna.
Map 141.

Fronds dimorphic, tufted from a short creeping rhizome. Sterile
fronds evergreen, 10-40 cm long, spreading and appressed to the
ground; stipes short; blades linear-oblanceolate, pinnate; pinnae
oblong or linear-oblong, blunt or somewhat pointed, becoming much
reduced towards the base. Fertile fronds fewer, upright, deciduous,
much longer than the sterile; pinnae linear, narrower than the sterile.
Veins simple or branched. Sporangia confluent, parallel to the midrib.
Indusium continuous, brown-hyaline, attached close to the margin.

The genus Blechnum numbers about 200 species, mainly of
south-temperate and tropical distribution. The species are both
terrestrial and epiphytic. Only one species, B. spicant, of wide
distribution, is represented in our area. As stated by Hitchcock et al.
(1969), the American plants tend to be a little more robust than the
European ones, but are scarcely separable taxonomically. Love and
Love (19666, 1968) did, however, refer the plants of the Pacific region
to ssp. nipponicum. If these were treated at specific levels, then the
name to be applied would probably be B. doodioides, the type of which
was collected in British Columbia.

Cytology: m = 34 (Cody and Mulligan 1982*).

273

Fig. 142 Blechnum spicant; (a) sterile and fertile fronds, 1/3 x ; (b) portion of fertile
pinna, 3x.

274

Habitat: Wet woods and clearings near the Pacific coast, and also
inland in the Revelstoke region.

Range: Interruptedly circumpolar; in North America from coastal
Alaska to California.

Remarks: A common fern in coastal British Columbia, B. spicant is
found in the same habitat as Polystichum munitum. In the Queen
Charlotte Islands, it is found in almost every habitat, from sea level to
the tree line (Calder and Taylor 1968).

2. Woodwardia Sm.

Coarse ferns, with horizontal widely creeping or stout ascending
rhizomes. Fronds similar or dimorphic, pinnatifid or bipinnatifid.
Veins anastomosing to form costal or costular areolae, then free to the
margin or forming additional areolae. Sori linear or oblong, parallel
to the midveins, borne along the veinlets, which form the outer side of
the first row of areolae. Indusium persistent, opening on the side
adjacent to the midrib.

The species of this small genus are found in bogs and wet shady
places. Some authors (Cranfill 1980) have separated Woodwardia
areolata from the genus as a monotypic species in the genus
Lorinseria, based on the marked dimorphism and other characters. In
a treatment of the Florida chain ferns, Lucansky (1981) states that
comparative anatomical data support the placement in the genus
Woodwardia of the three species found in that state, W. virginica, W.
radicans, and W. areolata. Cody (1963) reported on the genus in
Canada.

A. Fronds dimorphic, pinnatifid 1. W. areolata

A. Sterile and fertile fronds similar or nearly so, pinnate-pinnatifid.
B. Fronds 0.7-1.3 m long; pinnules 0.6-1.5 cm long; sori

usually confluent when mature 3. W. virginica

B. Fronds 1-2 m long; pinnules 2-6 cm long; sori usually
separate at maturity 2. W. fimbriata

1 . Woodwardia areolata ( L. ) Moore

Lorinseria areolata (L.) Presl

W. angustifoliaJ.E. Smith

netted chain fern
Fig. 143 (a) sterile and fertile fronds; (6) portion of fertile pinna;
(c) venation. Map 142.

Fronds dimorphic, pinnatifid, arising from a slender branching
rhizome. Sterile fronds 10-70 cm long; stipes greenish yellow; blades

275

Fig. 143 Woodwardia areolata; (a) sterile and fertile fronds, 1/2 x ; (b) portion of
fertile pinna, 2 x ; (c) venation, 2 x .

276

oblong-lanceolate to ovate, 7-35 cm long or longer; lanceolate
divisions united at the base by a broad wing, with margins finely
toothed; venation in several rows of areolae, then open to the margin.
Fertile fronds taller than the sterile; stipes darker; blades narrower;
divisions narrowly linear and almost distinct. Sori in a single row on
each side of the secondary midrib.

The sterile fronds of this species might at first glance be confused
with those of the common and sometimes weedy Onoclea sensibilis,
sensitive fern. Woodwardia areolata can easily be distinguished from
that species by its minutely serrate rather than entire margins and by
its basal pinnae, which are alternate rather than subopposite.

Cytology: n = 35 (Cody and Mulligan 1982*).

Habitat: Mediacid situations along streams and among cobblestones
on beaches.

Range: In Canada known only in Yarmouth, Shelbourne, and
Queens counties in Nova Scotia; in the United States on or near the
coastal plain to Florida and Texas and sparingly inland to Missouri
and Oklahoma.

Remarks: Woodwardia areolata is rare in Nova Scotia (Maher et al.
1968).

2. Woodwardia fimbriata J.E. Smith

W. chamissoi Brack.

W. paradoxa Wright

W. radicans (L.) Smith var. americana Hook.
Fig. 144 (a) portion of frond; (b) fertile pinnule. Map 143.

Fronds up to 70 cm long or longer (longer to the south),
evergreen, forming a crown at the end of the stout and widely creeping
or more or less erect rhizome. Stipes equaling the blades,
conspicuously chaffy at the base. Blades linear-oblong to oblong-
lanceolate, pinnate-pinnatifid, acuminate at the tip and narrowed
towards the base; pinnae linear-oblong, obliquely pinnatifid; ultimate
segments narrowly triangular to linear, acuminate; venation in a
single row of areolae on either side of the midvein, then free and
sometimes forking to the sharply and closely serrulate teeth. Sori in
shallow pits in the areolae on either side of the secondary midrib.

The elongated sori on either side of the secondary midrib and the
tall stature of this fern readily set it off from other fern species in
British Columbia.

Cytology: Manton and Sledge (1954) reported a count of n = 34 based
on material from Ceylon (Sri Lanka), but it is rather doubtful if the
species was W. fimbriata.

277

Fig. 144 Woodwardia fimbriata; (a) portion of frond, 1/3 x ; (b) fertile pinnule, 1 x

278

Habitat: Damp and boggy woodland and banks.

Range: In Canada known only from coastal British Columbia:
Lasqueti Island, Texada Island, and Saanich Arm on Vancouver
Island; south in the United States to California and Arizona.

Remarks: This species will undoubtedly be placed on the list of rare
plants of British Columbia.

3. Woodwardia virginica (L.) Sm.

Anchistea virginica (L.) Presl

Virginian chain fern
Fig. 145 (a) portion of frond; (6) fertile pinnules. Map 144.

Fronds 60-100 cm long, scattered along the creeping rhizome.
Stipes long, lustrous. Blades oblong-lanceolate, 10-30 cm wide,
pinnate-pinnatifid; pinnae linear-lanceolate; pinnules oblong, obtuse,
with finely serrulate margins. Veins united to form a single series of
areolae next to the midrib of both the pinnae and the pinnules, then
free to the margin. Sori oblong, usually becoming confluent at
maturity, one to each areole.

The distinctive chains of sori on the areolae adjacent to the
midrib set this fern off from all other fern species in eastern Canada.

Cytology: n = 35 (Britton 1964*).

Habitat: Swampy woods, boggy shores, cobbly lakeshores, and peat
bogs.

Range: Eastern North America, Nova Scotia to Ontario, south to
Florida and Texas.

Remarks: Woodwardia virginica was originally placed in the genus
Blechnum by Linnaeus. Presl (1851) later transferred it to the genus
Anchistea, where some researchers still prefer to retain it, although
only the glandular indusia and the presence of a single row of areoles
distinguish this species from other species of Woodwardia.

279

Fig. 145 Woodwardia virginica; (a) portion of frond, 1/3 x; (b) fertile pinnules,
1 1/2 X .

280

12 ASPLENIACEAE

Mostly small ferns with fronds firm, simple, pinnate, or
bipinnate. Veins free or forking. Sori elongate, occurring along the
veinlets. Indusia attached to the veinlets.

The family consists of 600-700 species worldwide and is
dominated by the very large genus Asplenium. In all, there are fewer
than 14 genera, and the other genera have very few species.
Asplenium has been extensively studied in Europe, the United States,
and New Zealand, and chromosome numbers for over 140 species are
known. Lovis, who followed Manton at Leeds, has made many
artificial crosses and, together with Meyer in Berlin, has analyzed the
European species in great detail (Lovis 1977; see also Reichstein
1981). There are 38 European species of which 18 are tetraploids and
of these, 11 are genomic or alloploids and six are considered to be auto-
ploids (Lovis 1977). Evans (1970) reviewed studies on Asplenium,
with special emphasis on those in the southern Appalachians. Ten
species and 10 hybrids are included in the discussion, and
Camptosorus is such an integral part of the crossing diagram and is
involved in so many hybrid combinations that it was evident that it
should be considered as an entire-leaved Asplenium rather than a
separate genus. Recent work from Europe (Jermy et al. 1978) has
included the genera Phyllitis and Ceterach in Asplenium as well.

In the Canadian flora, there are only six species to consider, but
these are all attractive species that are always admired when seen in
the wild. Most are found in rock crevices and usually in shady and
mossy places, so that they create a very pleasing tapestry of different
shades of green.

A. Fronds simple, commonly auricled at the base.

B. Fronds long-caudate, sometimes rooting at the tip

2. Camptosorus

B. Fronds oblong, not attenuate or rooting at the tip

3. Phyllitis

A. Fronds pinnate or bipinnate 1. Asplenium

1 . Asplenium L.

Small ferns, usually of rocky places, from small compact
rhizomes. Veins free. Sori linear, oblique. Indusium usually
membranous, attached lengthwise along one side of the sorus.

A very large genus with a broad distribution on all continents.
In our flora we have only four species of distinctive morphology. One,
A. viride, is a northerner, and the other three are cool temperate
plants. Asplenium platyneuron has almost all its distribution south of
Canada, and A. ruta-muraria has very few stations in Canada.

281

Our species are either diploid or tetraploid, but polyploidy is
impressive in some species in the world. Twelve-ploid and 16-ploid
species are known based on x = 36 (Lovis 1977).

A. Fronds dimorphic; fertile fronds upright, much taller than the

spreading sterile ones 1. A. platyneuron

A. Fronds similar.

B. Rachis purplish black, lustrous 3. A. trichomanes

B. Rachis green.

C. Blade linear, simply pinnate 4. A. viride

C. Blade deltoid-ovate, bipinnate .... 2. A. ruta-muraria

1. Asplenium platyneuron (L.) Oakes

A. ebeneum Ait.

ebony spleenwort
Fig. 146 (a) sterile and fertile fronds; {b) fertile pinnules. Map 145.

Fronds dimorphic, tufted from a short rhizome. Fertile fronds
stiff and upright, 20-40 cm long, 2.5-4.0 cm wide, gradually tapering
to the base; pinnae linear-oblong or basal pinnae triangular, auricled,
widely separated and alternated; rachis lustrous, chestnut purple.
Sterile fronds shorter, spreading, and prostrate, with oblong
approximate pinnae. Sori linear-oblong, situated on the veins, nearer
the midvein than the margin.

This species is easily identified by the stiff, upright fertile fronds.
It is not often associated with the other species of Asplenium in our
flora, and so the probability of hybridization is reduced.

Cytology: m = 36 (W.H. Wagner 1973a).

Habitat: In partial shade in open woods, grown-over areas and
clearings, often in moss or in very shallow soil over rocks.

Range: Southwestern Quebec and southern Ontario, south to Florida
and Texas and west to Iowa and Kansas in recent years.

Remarks: W.H. Wagner and Johnson (1981) have documented the
recent spread of A. platyneuron in parts of Ontario and adjacent
Quebec and have provided a wealth of biological data on the species.
These recently produced plants are found in very atypical habitats and
may prove to be short-lived. We have seen small colonies in young
pine plantations on sand; in beds of Equisetum hyemale on moist
sandy slopes, i.e., acidic locations; and on rocks and in moss on
limestone paving, both presumably of high pH. The species was
considered to be rare in Ontario by Argus and White (1977), but now
seems to be too widespread to be so regarded.

282

Fig. 146 Asplenium platyneuron; (a) sterile and fertile fronds, 1/2 x ; (b) fertile
pinnules»3 x .

283

2. Asplenium ruta-muraria L.

A. cryptolepis P^ern.
wall-rue
Fig. 147 (a) fronds; (6) fertile pinnules. Map 146.

Fronds 3-9 cm long, tufted from a short rhizome. Stipes naked or
minutely scaly at the base, green. Blades deltoid-ovate, 1-6 cm long,
bipinnate; pinnae and pinnules mostly alternate, petioled; ultimate
segments rhombic or obovate, long-cuneate at the base, with the
broadly rounded apex crenately toothed. Veins flabellate, simple or
forked. Sori few, linear-oblong, usually not confluent in age.

This is a very diminutive fern that is rarely abundant in our
area. One is fortunate to locate a small plant after inspecting literally
thousands of large dolomitic talus boulders without success.

Cytology: n = 72 (W.H. Wagner and F.S. Wagner 1966). This has
the same chromosome number as the European plants. Lovis (1977)
says that ssp. ruta-muraria is an autotetraploid of ssp. dolomiticum
(n = 36) in Europe.

Habitat: Sunny or shaded crevices of limestone cliffs.

Range: Manitoulin Island and Bruce Peninsula, Ont., southern
Quebec, in the United States from Vermont to Michigan, south to
Alabama and Missouri.

Remarks: The distribution in Ontario is considered in detail by Soper
(1955). The species is on the rare plant list for Ontario (Argus and
White 1977). This is such a common and abundant plant of stone
walls in Great Britain that it is a surprise for field workers from
Europe to find that the plant is considered such a rarity in Canada.
Frere Louis-Alphonse in 1951, 1952, and 1953 collected plants from
"crevasses dans un rocher" (crevices in a rock) at Baie Missisquoi in
southern Quebec near the United States border (specimens at MT). B.
Boivin has annotated these specimens "var. ruta-muraria et
vraisemblement plante," (var. ruta-muraria and presumably planted)
but in view of the fact that the plant is known in nearby Vermont, this
seems doubtful.

Variety cryptolepis (Fern.) Massey was considered to differ from
the European ssp. ruta-muraria by hidden rhizome scales (Fernald
1928). We are in agreement with Wherry (1961) that the variety
should be dropped.

284

Fig. 147 Asplenium ruta-muraria; (a) fronds, 1 x ; (b) fertile pinnules, 3 x .

285

3. Asplenium trichomanes L.

maidenhair spleenwort
Fig. 148 (a) fronds; (6) fertile pinnae. Map 147.

Fronds 6-20 cm long or longer, forming a dense tuft from a
compact rhizome. Stipe and rachis purple brown; old rachises
persistent. Blades linear, pinnate; pinnae usually opposite or
subopposite, oval, rounded to cuneate at the inequilateral base and
slightly toothed on the sides and at the blunt apex. Sori linear,
situated on the veins between the midrib and the margin.

Cytology: n = 36 (Britton 1964*; Cody and Mulligan 1982*); n = 72
(Britton 1953*; Cody and Mulligan 1982*).

Habitat: Sheltered rock crevices.

Range: Circumpolar; in North America from western Newfoundland
to Ontario, British Columbia, and south to Georgia, Alabama,
Arkansas, Oklahoma, and Arizona.

Remarks: This extremely attractive species has received a great deal
of attention from the Leeds school (Lovis 1977). Early work showed
that there were both diploids and tetraploids in the complex, the
former on more acidic rocks than the latter. Extensive analysis
showed that the tetraploid was an autotetraploid, and names were
selected to indicate this relationship. The basic diploid became
Asplenium trichomanes ssp. trichomanes and the autotetraploid A.
trichomanes ssp. quadrivalens D.E. Meyer emend Lovis. Later,
another diploid was found in Europe on limestone and was described
as A. trichomanes ssp. inexpectans Lovis. This work has been followed
with interest in North America, although authors have been reluctant
to utilize the subspecific names from Europe without further study of
North American plants. It has been known since 1953 (Britton 1953)
that both diploids and tetraploids occur in North America and that soil
preferences exist for these (diploid on granite and tetraploid on
limestone). The complexity of the situation was highlighted by D.H.
Wagner and W.H. Wagner (1966), who reported a colony in Virginia
consisting of one diploid, three triploids, and 85 tetraploids, and no
superficial differences in morphology.

For those who wish to delineate ssp. trichomanes from ssp.
quadrivalens, one should study the note by Lovis in the Atlas of Ferns
of the British Isles (Jermy et al. 1978), Jermy and Page (1980), and
Moran(1982).

Arriving at names for a simple recognition of diploids and
tetraploids in North America has been greatly complicated by Love
(Love et al. 1977), who decided that Asplenium trichomanes L. ssp.
trichomanes is the tetraploid, not the diploid, as treated by Lovis

286

Fig. 148 Asplenium trichomanes; (a) fronds, 1/2 x ; (b) fertile pinnae, 3 x .

287

(1964), and that the diploid should be known as A. melanocaulon
Willd. It would be a great pity if the taxonomic rules forced us to use
two entirely different specific names for plants that have so many
features in common. We side with Lovis (1964) "that when a Linnean
name covers a species complex, this name should be retained by that
segregate with which Linnaeus would have been most familiar, and
which would therefore most likely have represented to him the most
typical form of his species."

4. Asplenium viride H uds .

green spleenwort
Fig. 149 (a) fronds; (6) fertile pinnae. Map 148.

Fronds 2-14 cm long, tufted from a short rhizome. Stipes
darkened below, green above. Rachis green. Blade linear to linear-
lanceolate, pinnate; pinnae rounded or rhomboid-ovate, crenate. Sori
elongate, borne near the indistinct midrib, becoming confluent at
maturity.

This northern species, with its bright green stipes, poses no
identification problems unless extremely small, poorly developed
fronds from subarctic sites are encountered and are subsequently
confused with Woodsia glabella. One should check for an elongate
sorus, with the indusium attached on one side, for positive
identification of A. viride.

Cytology: m = 36 (Taylor and Lang 1963*; Britton 1964*).

Habitat: Among talus and in usually protected crevices of limestone
or basic rocks in shady locations.

Range: Circumpolar; in North America from Newfoundland to
Alaska, south to New York, Colorado, Utah, and Washington.

Remarks: A distinctive species usually less common on the Niagara
Escarpment than is Asplenium trie ho manes, with which it is some-
times associated. One might expect hybrids to be frequent in localities
where the two species grow intermixed, but although that hybrid is
reported as a very rare one in Europe, none are known for North
America. Considered to be rare in Ontario by Argus and White (1977).

Hybrids of Asplenium

Since there are four species of Asplenium in Canada and two
levels of ploidy (2x and 4jc), the theoretical expectations with five
entities and no barriers to hybridization are (4—3-2-1), or 10 possible
primary crosses. Five of these are known in nature or have been

288

Fig. 149 Asplenium viride; (a) fronds, 1 x ; (b) fertile pinnae, 3 x

289

produced artificially and are recorded in the literature (Knobloch
1976). Reichstein (1981) considers four known in Europe. We know of
none in Canada, although five A. trichomanes hybrids are known in
the United States and all are very rare (Moran 1982).

What was once considered even more unusual — intergeneric
crosses between Camptosorus and various species of Asp/enmm — are
well known in the United States. In theory, each of the ^ive taxa of
Asplenium could cross with Camptosorus, and so there would be five
possible combinations (Knobloch 1976). None has been found in
Canada so far.

It is worth noting that one can see evolution in action when one
compares the primary hybrid of the two diploids, Asplenium
platyneuron (PP) X Camptosorus rhizophyllus (RR). The result is a
sterile diploid (P)(R) known as Scott's spleenwort, or Asplenium X
ebenoides. If an occasional spore should be produced with all the
chromosomes of this hybrid, it gives rise after fertilization to a new
fertile allotetraploid with the constitution PPRR and would be known
as A. ebenoides, a derived species.

Excluded report

Asplenium marinum L.

Lawson (1889) stated, "There are Nova Scotian specimens in the
Kew Herbarium, referred to in Hooker & Baker's Synopsis Filicum,
second edition, 1883, but this fern has not recently been found in
Canada. It grows around the shores of Western Europe, and extends
from Orkney, the British Isles, Canaries and Azores, to St. Vincent
and South Brazil." A search of the herbaria at Kew, Edinburgh, and
Glasgow failed to turn up the specimens. Presumably they have been
revised to some other species.

2. Camptosorus Link

1 . Camptosorus rhizophyllus ( L. ) Link

Asplenium rhizophyllum L.

walking fern
Fig. 150 (a) fronds; (6) portion of fertile frond. Map 149.

Fronds 5-30 cm long or longer, clustered at the end of the erect or
ascending scaly rhizome. Blades evergreen, entire, 1-3 cm wide at the
cordate or auriculate base, usually tapering to a long caudate tip.
Veins reticulate. Sori elongate, scattered along the veins. Indusium
attached on one side of the sorus. Tips of the arching blades often
rooting to form new plants, hence the name walking fern.

290

Fig. 150 Camptosorus rhizophyllus; (a) fronds, 1/2 x; (b) portion of fertile frond,

2 1/2 X .

291

This fern is most distinctive and is much admired by rock garden
enthusiasts. European visitors are always eager to see the species in
the wild and are often initially disappointed in its stature, expecting it
to be as long as a hart's-tongue.

Cytology: n = 36 (W.H. Wagner et al. 1970).

Habitat: Damp, sheltered, often mossy rocks and stony banks,

preferring limestone.

Range: Southwestern Quebec and southern Ontario to Minnesota,
south in the United States to Georgia and Mississippi, and west to
Oklahoma and Kansas.

Remarks: It is impossible to refute the argument that this fern is in
reality an entire-leaved Asplenium, based on its propensity to cross
with Asplenium species. We have retained it here largely for
sentimental reasons, because it is well known by this name and to
highlight the fact that it has a counterpart in Asia, C. sibiricus Rupr.
These represent a pair of interesting species, treated as subspecies by
Love (Love et al. 1977), and are the only members of the genus.

3. Phyllitis Hill

1. Phyllitis scolopendrium (L.) Newm. var. americanum Fern.

Phyllitis fernaldiana Love

Phyllitis japonica Kom. ssp. americana (Fern.) Love &, Love

Scolopendrium uulgare auth.

hart's-tongue
Fig. 151 (a) fronds; (b) portion of fertile frond. Map 150.

Fronds 15-40 cm long or longer, from a short caudex; blade
simple, oblong-lingulate or strap-like, deeply cordate-auriculate at the
base, and tapered to a point at the tip. Stipe short, clothed with
narrow, curling, long-caudate scales. Veins free, forking. Sori
narrowly oblong, nearly at right angles to the midrib, and located on
either side of adjacent veinlets so that the sori appear to have double
indusia opening along the middle.

This is a very striking fern that reminds one of tropical plants —
to some it is even reminiscent of banana leaves. It is an exciting
experience to see a large number of these plants on the rich, cool,
shady limestone talus of the Niagara Escarpment. Identification is
easy, but there is some disagreement as to what name should be
applied to North American plants because of their cytology. Canada's
plants are tetraploid in = 72, Britton 1953), as are those of Japan,
whereas the European ones are diploid. Emmott (1964) made a
number of crosses between European, American, and Japanese plants,

292

Fig. 151 Phyllitis scolopendrium var. americana; (a) fronds, 1/3 x; (b) portion of
fertile frond, 4 x .

293

but the degree of relatedness between these widely separated
populations was not clear from her results. Love, initially impressed
by the fact that the North American plants were tetraploid, suggested
the name Phyllitis fernaldiana Love, but later believed that the two
tetraploids should be called P. japonica Kom., with the North
American known as P. japonica ssp. americana (Fern.) Love and Love.
Further experimental study is needed, and so we have retained the
familiar name Phyllitis scolopendrium, even though we are aware
that some Europeans now refer this species to Asplenium rather than
to Phyllitis.

Cytology: n = 12 (Cody and Mulligan 1982*).

Habitat: Rich rocky woodland adjacent to the Niagara Escarpment,
often associated with Polystichum lonchitis.

Range: In Ontario known from Bruce, Dufferin, Gray, Halton, Peel,
Simcoe, and Welland counties. It may have been introduced in
Welland County. An early report from Woodstock, N.B., has not been
substantiated by recent collections and may also have represented an
introduction. In the United States, hart's-tongue is known only from a
few widely scattered localities: central New York, Tennessee, and
Michigan.

Remarks: Because North American plants are tetraploid, it is now
possible to reinvestigate plants in isolated sinkholes, such as those in
Tennessee and Alabama, to determine if the plants are native or were
introduced from Europe. A small colony of small plants discovered on
western Vancouver Island consists of diploid plants. These are
considered to have been introduced.

Most of the extant North American plants occur in a small area
of Ontario. A special effort should be made to protect them. It is a
mistake to move them to gardens, because they are adapted to very
special requirements of substrate, drainage, moisture, and degree of
shade, which cannot be duplicated in the average garden. The species
is rare in all of North America, including Canada and Ontario (Argus
and White 1977).

294

13 POLYPODIACEAE

1. Polypodium L.

Small to medium-sized ferns from widely creeping branching
chaffy rhizomes. Fronds jointed to the rhizome, evergreen, not
reduced toward the base. Veins once or twice branched or
anastomosing. Sori round or oval, naked, in rows on either side of the
midrib, with or without glandular paraphyses among the sporangia.

The genus Polypodium has many tropical representatives and
over 100 species in the world. The basic chromosome number (x) for
almost all of these is firmly established as 37, although there are very
few species with either 36 or 35. Ploidy levels range from 2x to 6x in
the genus (Lovis 1977). In north-temperate regions, there are few
species to consider, but these have proved to be quite complex. In
Britain, P. uulgare s.l. has proved to be a complex of three levels of
ploidy: diploid, tetraploid, and hexaploid, each with its own
morphological and ecological characteristics. These are now known as
P. australe, P. uulgare s.s., and P. interjectum, respectively (T.G.
Walker 1979). Interestingly, P. uulgare, which is proven as an
allotetraploid, does not have the genomes of P. australe in its
constitution, and it was necessary to look elsewhere for the diploid
progenitors of this long-established species. Cytogenetic analysis has
suggested that two of our species, P. glycyrrhiza and the diploid P.
uirginianum, might be the ancestors of the European P. uulgare.
Accordingly, in spite of attempts through the years to keep P.
uirginianum in North America clearly separate from P. uulgare in
Europe, it would seem that the P. uulgare complex must be approached
on a worldwide basis. Polypodium fauriei Christ in Asia should not be
forgotten.

In eastern Canada, there is only P. uirginianum to consider,
although it is clear that there are both diploids and tetraploids to
delineate and a very common sterile triploid hybrid that arises from
hybridization between these two.

In western Canada, there are four other species to consider. One,
P. scouleri x = n = 37, is most distinctive in our flora, and is
apparently quite remote from the ancestry of the others. There are
two other basic diploids in the West, the widespread coastal P.
glycyrrhiza (GG) and the more montane P. amorphum (AA). There is
also a presumptive allotetraploid, P. hesperium (AAGG), considered to
have arisen from the doubled hybrid of P. amorphum and P.
glycyrrhiza (Lang 1971).

These polypodies are most often found on rock surfaces, e.g.,
talus boulders, rocky ledges, and cliffs, although in the coastal forests
of British Columbia, P. glycyrrhiza may be seen quite far up on trunks
and branches of maple trees, suggesting epiphytes of the tropics. The
leathery evergreen fronds of all our species, with little dissection of
the blade, make it easy for the amateur to identify the genus.

295

A. Blade stiffly coriaceous; veins anastomosing 1. P. scouleri

A. Blade herbaceous to membranous; veins free.

B. Segments of fronds usually 3 cm long or longer, with the

tips acute to attenuate 2. P. glycyrrhiza

B. Segments of fronds usually less than 3 cm long, with the
tips obtuse to acute.

C. Sori round; rhizome scales with a darker central
stripe.
D. Blades abruptly tipped; pinnae oblong, blunt or

somewhat acute (eastern and northern)

5. P. virginianum

D. Blades tapering to the tip; pinnae somewhat oval

(western British Columbia)

3. P. amorphum

4. P. hesperium

C. Sori oval; rhizome scales concolorous

1 . Poly podium scouleri Hook. & Grev.
Fig. 152, fronds. Map 151.

Fronds 15-40 cm long or longer, from a sparsely scaly stout
rhizome. Stipes stiff, shorter than the blade, and with a few deciduous
scales. Blades thick and leathery, deltoid-ovate, pinnate to pinnatifid;
segments linear to linear-oblong, rounded at the apex. Rachis and
midveins of pinnae with deciduous scales. Veins anastomosing. Sori
round, large, near the midvein, and situated on the upper pinnae.

The very thick and leathery fronds, with blunt divisions and
lobes, make this species easy to identify at a glance.

Cytology: n = 37 (Cody and Mulligan 1982*).

Habitat: Near the coast and often reached by the salt spray, on
banks, cliffs, and tree trunks.

Range: Southern British Columbia, south to southern California.

Remarks: This species has others that resemble it in California and
is part of the P. californicum complex. Various hybrids are known in
that region.

2. Polypodium glycyrrhiza DC. Eat.
P. uulgare L. var. occidentale Hook.

P. vulgare L. ssp. occidentale (Hook.) Hulten
licorice fern
Fig. 153 (a) fronds; (6) fertile pinna. Map 152.

296

Fig. 152 Poly podium scouleri; fronds, 1/3 x .

297

Fig. 153 Poly podium glycyrrhiza; (a) frond, 1/2 x ; (b) fertile pinna, 1 1/2 x

298

Fronds up to 60 cm long or longer from the relatively thick
licorice-tasting rhizome. Scales light brown or straw-colored, cordate
to peltate, ovate, often with a capillary tip. Blades oblong to ovate;
segments oblong-attenuate, with acute to acuminate tips, with finely
serrate margins, and pubescent along the midveins on the lower
surface. Sori on the upper segments round or occasionally somewhat
oval, about equidistant between the margin and midvein. Paraphyses
absent.

The long, acute tips to the segments and the licorice taste of the
rhizome are useful field characters.

Cytology: n = 37 (Lang 1971*). A basic diploid ancestral species.

Habitat: On tree trunks, mossy logs, and moist banks at low
elevations near the coast.

Range: Kamchatka, the Aleutian Islands, and through the Alaskan
Panhandle, south to central California.

Remarks: This basic diploid species forms great sheets over rocky
banks. It is conspicuous along several highways in coastal British
Columbia.

3. Poly podium amorphum Suksdorf

P. montense F.A. Lang
Fig. 154, fronds. Map 153.

Fronds up to 30 cm long from a thin acrid rhizome. Scales dark
brown to chestnut, usually with a darker centre, narrowly ovate, often
constricted near the base and with a long capillary tip. Blades oblong,
up to 20 cm in length; segments oblong to obovate, with obtuse or
rarely acute tips; margins entire to crenulate. Veins free. Sori round,
near the margins of the segments, and rarely with a few glandular
paraphyses.

This basic ancestral diploid species was segregated from P.
hesperium, by Lang (1969), as P. montense Lang. Morton (1970)
pointed out that P. amorphum Suks., even if monstrous in form, had
priority; it apparently was extirpated at the type locality.
Unfortunately, T.M.C. Taylor (1970) has P. amorphum listed as a
synonym of P. hesperium Maxon, whereas now P. amorphum is the
accepted name for what was P. montense.

This species is more montane than P. glycyrrhiza, has rounded
segments, round sori, and an acrid-tasting rhizome. It resembles
closely some of the plants of the tetraploids of P. uirginianum.

Cytology: n = 37 (Lang 1969, 1971*). A basic diploid ancestral
species.

299

Fig. 154 Polypodium amorphum; fronds, 1/2 x

300

Fig. 155 Poly podium hesperium; (a) fronds, 2/3 x ; (b) fertile pinna» 1 1/2 x ,

301

Habitat: Rock crevices in the mountains.

Range: Coastal Mountains in British Columbia, south to Oregon and
the Sierra Nevada, Calif.

Remarks: The rhizome is acrid to the taste.

4. Polypodium hesperium Maxon

P. uulgare L. var. columbianum Gilbert
P. uulgare L. ssp. columbianum (Gilbert) Hulten
Fig. 155 (a) fronds; (b) fertile pinna. Map 154.

Fronds up to 35 cm long or longer from a rather thick, licorice-
tasting rhizome. Scales chestnut brown, lance-ovate, more or less
crenate-serrate. Blades oblong, up to 20 cm in length; segments
oblong, obtuse to acute at the tip; margins entire to serrate. Sori on
the upper segments oval, about equidistant between the margin and
midvein. Paraphyses common.

This species is considered to be an allotetraploid derivative of P.
amorphum and P. glycyrrhiza. From the former, it has the markedly
rounded lobes of the segments and from the latter it has inherited the
licorice taste of the rhizomes. The medial sori, which are oval, are
good field characters.

Cytology: n = 14 (Lang 1971*). Allotetraploid of constitution
AAGG.

Habitat: Rocky slopes and crevices.

Range: Southern British Columbia, south to Arizona, New Mexico,
and Baja California, Mexico, and east to the Black Hills of South
Dakota.

Remarks: Although Lang had two of the three hybrid combinations
needed to analyze this species (P. amorphum X hesperium and P.
glycyrrhiza X hesperium), and the morphological data are fairly
consistent, the direct proof of the allotetraploid origin of P. hesperium
is lacking (Lovis 1977). Artificial crosses that take 6-8 years to
produce fertile plants or the analysis of a natural hybrid of P.
amorphum X glycyrrhiza are needed for certain proof. Polypodium
hesperium is rare in Alberta (Argus and White 1978).

5. Polypodium virginianum L.

P. uulgare L. ssp. uirginianum (L.) Hulten

rock polypody
Fig. 156, fronds (diploid). Fig 157 (a) fronds; (6) portion of fertile pinna
(tetraploid). Map 155 (s.l.).

302

Fig. 156 Poly podium virginianum; fronds (diploid), 1/3 x

303

# b

Fig. 157 Polypodium virginianum; (a) fronds, 2/3 x; (b) portion of fertile pinna
(tetraploid), 2 x .

304

Fronds up to 35 cm long from a creeping somewhat acrid
rhizome. Scales often with a dark central stripe, deeply cordate.
Blades oblong-lanceolate; segments linear-oblong to deltoid, entire to
remotely dentate, blunt or acutish at the tip. Veins free. Sori round,
midway between the midvein and margin, and occurring on the upper
segments. Paraphyses present.

In eastern Canada Polypodium uirginianum is easily identified
by its small evergreen fronds of distinctive morphology and its usually
large colonies on rocky talus, capping boulders, climbing rocky slopes,
and ledges. The species is more complex than a casual glance would
indicate. It is known that there are both diploids in — 37) and
tetraploids {n — 74) in eastern Canada, and that the triploid hybrid is
quite common (Evans 1970). The diploids have blades that are more
deltoid in shape, with acute apices of the segments, and the tetraploids
have a more oblong blade (narrowed at the base), with rounded apices
of the segments. There are a number of other morphological
differences that are considered in detail by Kott and Britton (19826)
and Cranfill (1980).

Cytology: n = 37 and 74 (Britton 1953*; Love and Love 1976*; Cody
and Mulligan 1982*).

Habitat: In shallow humus on rocks, in crevices, on woodland banks,
and rarely on mossy stumps and in crotches of trees.

Range: Newfoundland to central Alaska and northeastern British
Columbia, south in the east to Georgia, Alabama, Tennessee, and
Arkansas. The tetraploid is more widespread in Canada to the north
and west, whereas the diploid is more abundant in the southeast.
Most plants in Ontario are tetraploids. The diploid has a distribution
in Canada similar to that of Dryopteris campyloptera.

Remarks: Love and Love (1977) named the tetraploid P. vinlandicum
without lectotypification of P. uirginianum L. The type of the former
is given as P. uirginianum L. var. americanum Hooker. It is
unfortunate to designate such old, poorly studied material as the type
for a new species, when we would like to know as much as possible
about the type material, e.g., chromosome number, number of
paraphyses, spore morphology, and chromatography. This is
impossible with old herbarium specimens of uncertain provenance,
which sometimes contain mixtures.

The frequent triploid hybrid, with mostly aborted spores, is
thought to be able to reproduce by occasional spores that have an
unreduced chromosome number (Evans 1970).

It is premature to describe a new species before P. uirginianum is
lectotypified and before the relationship of the tetraploid to the diploid
race as well as to the diploid P. amorphum is clarified by experimental

305

means. Polypodium uirginianum is rare in the Yukon (Douglas et al.
1981).

Hybrids of Polypodium

Polypodium amorphum X glycyrrhiza has not yet been reported
in Canada.

Polypodium glycyrrhiza X hesperium has been reported in three
localities in British Columbia: Kaske Creek, approximately 100 km
east of Prince Rupert; Alexandra Bridge, Eraser River; and Green
River, Pemberton.

Polypodium amorphum X hesperium has been reported in
Alexandra Bridge, Fraser River, B.C.

The triploid hybrid between the two cytotypes of P. uirginianum
is frequent in eastern Canada (Kott and Britton 19826). It is more
widespread in Ontario than is the diploid.

306

14 MARSILEACEAE

Marsileaceae, which contains only three genera, is characterized
by the presence of sporocarps. It is not closely related to other fern
families, except perhaps Salvineaceae. About 75 species, mainly of
Old World distribution, are in the genus Marsilea.

1. MarsileaL. water-clover

Plants aquatic, perennial, herbaceous, submersed or emersed,
growing from widely creeping rhizomes. Leaves alternate, long-
petioled; blade divided into four clover-like leaflets. Sori embedded in
a gelatinous sheath in sporocarps; sporocarps hard, ovoid,
pedunculate, two-loculate, bearing 2 more or less conspicuous teeth
near the base. Microspores numerous, but megaspores only one to a
sporangium.

As is apparent by the common name, the four-foliate blade of this
genus mimics some members of the family Leguminosae.

A. Leaflets broadly obovate-cuneate, glabrous; sporocarps often in
pairs borne on a long stalk arising from the stipe above its base
2. M. quadrifolia

A. Leaflets broadly cuneate, sparsely appressed-pubescent;
sporocarps borne singly on short stalks from the rhizome or base
of the stipe 1. M. vestita

1. Marsilea vestita Hook. & Grev.

M. mucronata A.Br.
hairy pepperwort
Fig. 158, fronds and sporocarp. Map 156.

Leaves with petioles up to 10 cm long or longer, tufted or
scattered from an elongate rhizome. Leaflets to 1 cm long, broadly
cuneate, sparsely appressed pubescent. Sporocarps borne singly on
short stalks from the rhizome or from the base of the stipe.

Cytology: No recent reports seen.

Habitat: Shallow ponds, ditches, sloughs, marshy places, and quiet
streams, often stranded later in the season.

Range: Southern Saskatchewan to southern British Columbia, south
to California, Texas, and Arkansas.

Remarks: The species is rare in British Columbia. It was collected at
the Indian reserve in the vicinity of Kamloops, B.C., by John Macoun

307

in 1898, during his explorations of western Canada (CAN). T.C.
Brayshaw found it there again in 1948 (UBC). It could not be located
there when the senior author searched for it in 1976. A small stand is
also known at Goose Lake near Vernon, B.C. The hard sporocarps
may allow the species to survive dry periods. Indeed, Bloom (1955,
1961) has found that as long as the sporocarps remain dry, they could
be autoclaved for up to 15 minutes and still show excellent
"germination." Scoggan (1978) stated that there is no confirmation for
the report by Burman (1909) of this species occurring in Manitoba.
There is, however, a specimen in WIN labeled "Western Manitoba,"
which presumably is the basis of the report, but the exact locality, date
of collection, and collector are unknown. The species should be
searched for in western Manitoba.

2. Marsilea quadrifolia L.

water-clover
Fig. 159 (a) fronds and sporocarps; (b) venation. Map 157.

More robust than M. uestita. Leaves with petioles up to 20 cm
long, scattered from the elongated rhizome. Leaflets up to 2 cm long,
obovate-cuneate, glabrous. Sporocarps often in pairs, borne on a long
stalk arising from the stipe above its base.

Cytology: n = 20 (Mehra and Loyal 1959).

Habitat: Shallow water of slow-moving streams.

Range: In Ontario, only in Nanticoke Creek in Haldimand and
Norfolk counties and Mississauga in Peel County; in the United States
from New England to Iowa and Kentucky; naturalized in Europe.

Remarks: This species has been sold for use in pools and aquaria
whence it escapes to slow-moving streams and ponds.

308

Fig. 158 Marsilea vestita; fronds and sporocarp, 1/2 x .

Fig. 159 Marsilea quadrifolia; (a) fronds and sporocarps, 1/2 x ; (6) venation, 2 x

309

15 SALVINIACEAE

Salviniaceae is a family of only two genera, Azolla and Saluinia;
it is sometimes separated into two families, with a total of about 16
species that occur primarily in tropical regions around the world.

1. Azolla Lam. mosquito fern

Ferns small, annual, aquatic, free-floating, with unbranched
thread-like roots. Plants compact, dichotomously branched, forming
small mats. Leaves usually crowded, two-lobed; upper lobe green or
often reddish later in the season; lower lobe usually larger than the
upper, only one cell thick, and mostly without chlorophyll. Sporocarps
in pairs in the leaf axils, each enclosed in an indusium.
Microsporocarps containing numerous microsporangia, each of which
produces masses of microspores, which when released exhibit peculiar
barb-tipped hairs (glochidia). Megasporocarps smaller, acorn-shaped,
each containing a single megasporangium with a single megaspore.

The New World species of Azolla were treated by Svenson (1944);
species of Azolla are often difficult to distinguish because of the
absence of sporocarps and the need to use a microscope to examine the
glochidia; Svenson's report of A. filiculoides for Alaska was not taken
upbyHulten(1967).

A. Plants less than 1 cm in diameter; leaves about 0.5 mm long;

glochidia without cross walls 1. A. caroliniana

A. Plants 1 cm in diameter or larger; leaves about 0.7 mm long or

longer; glochidia with cross walls 2. A. mexicana

1 . Azolla caroliniana Willd.
Map 158.

Plants 0.5-1.0 cm in diameter. Upper leaf-lobes 0.5-0.6 mm
long, smooth, not closely imbricate. Glochidia without cross walls.
Megaspores unknown.

Cytology: n = 24 (Tschermak-Woess and Dolezal-Janisch 1959).

Habitat: Quiet waters.

Range: Eastern United States.

Remarks: A collection by Judge Logie in 1862 from Hamilton Beach,
Ont., is now in the National Museum of Natural Sciences, Ottawa
(Lawson Herbarium). The species has not been collected in that area

310

Fig. 160 Azolla mexicana; crowded leaves and sporocarps, 9 x

311

since and has probably been extirpated. Soper (1949) noted that this
Azolla had been collected in 1934 from the American side of the
Niagara frontier region, and Pursh (1814) reported it from Lake
Ontario.

2. Azolla mexicana Presl

A. caroliniana auth. non Willd.

A. filiculoides sensu Scoggan (1978)
Fig. 160, crowded leaves and sporocarps. Map 159.

Plants 1.0-1.5 cm in diameter. Leaves crowded and overlapping;
upper leaf-lobes 0.7 mm long, papillose. Glochidia many-septate.
Lower part of megaspores pitted.

Cytology: None.

Habitat: Ponds and slow-moving streams.

Range: Southern British Columbia, south to northern South
America, east to Missouri, Illinois, and Wisconsin.

Remarks: In British Columbia known only in the area adjacent to
Sicamous and Salmon Arm, where it was collected as recently as 1976.
Scoggan (1978) was of the opinion that the species was introduced in
this area.

312

DISTRIBUTION MAPS

y-^

^^ •

Map 1 Lycopodium da va turn var. davatum

. . . \ .4

Sag

Map 2 Lycopodium davatum var. monostachyon

313

•\=i:>

"T • • • .#

'*!:'«9K'tS:*<-..

M_-^?J^<

fnn

ra>fi!^-

-^^

^. ^ -^-^ --v-^ *>'^ ^^->^

^ •^

_-*• • • • T^-i..

/»^v

Map 3 Lycopodium annotinum

.v._. ^ --^-h-,-,4-4

Map 4 Lycopodium dendroideum

314

./~^

IjU '

^ il\ ■

Map 5 Lycopodium obscurum var. obscurum

Map 6 Lycopodium obscurum var. isophyllum

315

Map 7 Lycopodium complanatum

?^W*5'"'^^^

Map 8 Lycopodium digitatum

316

Map 9 Lycopodium tristachyum

Map 10 Lycopodium alpinum

317

s )

*'*v^,.

Map 1 1 Lycopodium sitchense

""^^

Map 12 Lycopodium sabinifolium

318

Map 13 Lycopodium inundatum var. inundatum

Map 14 Lycopodium inundatum var. bigelovii

319

ry<tr^

- ; ^ A!3%^^j"x? '^ >' ■ ^. >>.^

PIT- Vc<-

Map 15 Lycopodium lucidulum

-^-^■^

•-., .^%

• •^

^•^■••;^-

Map 16 Lycopodium selago ssp. selago

320

Map 17 Selaginella selaginoides

Map 18 Selaginella apoda

321

Map 19 Selaginella wallacei

Map 20 Selaginella oregana

322

Map 21 Selaginella densa

Map 22 Selaginella densa var. scopulorum

323

^^-A

S^^.'

^■^]\^ ^^'^^^

01' ,''

1 ■=-^''

■ ■^ l3- -*V '^- -^■'

i;-^

C? 0<^ ^,. b>:^

--J:?^^?/-^*'

Map 23 Selaginella rupestris

V^ ill. 5

Map 24 Selaginella sibirica

324

Map 25 Isoetes echinospora

Map 26 Isoetes maritima

325

Map 27 Isoetes eatonii

Map 28 Isoetes riparia

326

Map 29 Isoetes acadiensis

Map 30 Isoetes hieroglyphica

327

Map 3 1 Isoetes tuckermanii

Map 32 Isoetes macrospora

328

Map 33 Isoetes nuttallii

Map 34 Isoetes howellii

329

Map 3 5 Isoe tes bola nderi

Map 36 Isoetes occidentalis

330

>^-.- .f|«£¥).

C '

• ^%^H,

"I.

' r^

'i^^'^iW^':-^?'

Map 37 Equisetum fluviatile

'^':<s

I v^^

■^.i.

■T--^^^:^:^^*^.

Map 38 Equisetum palustre

331

Map 39 Equisetum telmateia ssp. braunii

•-%•

•>.

-■4,^^^v-""'.i^.>-?^,-i

■^•:* . ^-/.:^;^vv ^-^ r ^.t/ .^ ^W ^-^ --^

Map 40 Equisetum arvense

332

'V.

-: ,^

5^ '^'^^C c?S

Map 4 1 Equisetum sylvaticum

Map 42 Equisetum pratense

333

V )

'\r^

Map 43 Equisetum hyemale ssp. affine

7\' .'-^::

V-.. >

1 .0 ,-,'■

^^^.

^- — 'XTi

^'y>^^.'"'^-

■ TV ^"^J^-^^ i

Map 44 Equisetum laevigatum

334

Map 45 Equisetum scirpoides

^^M^fSS?^

►--D'

■■•*

-'>^^-

II'.

Map 46 Equisetum variegatum ssp. variegatum

335

.:?k^^.

-M

Map 47 Equisetum variegatum ssp. alaskanum

{^f"2ix|V/f^V

V ^

S^^,

f^-^

. M

'^.., '^^^v. X

y,*^' -^^'>(ll. '-'^"^^^

\(w

)e^

Map 48 Ophioglossum vulgatum var. pseudopodum

336

"^tf

'V/

Map 49 Botrychium virginianum var. virginianum

Map 50 Botrychium virginianum var. europaeum

337

'X/'^

'-Xr

'V. 'iqj

)

* ^ / . J,

-*kc»»

Map 51 Botrychium dissectum

Map 52 Botrychium obliquum

338

Map 53 Botrychium oneidense

^i>.

^1. \

Map 54 Botrychium multifidum (s. I.)

339

Map 55 Botrychium rugulosum

'"^C^
^?-.

=5^-^

\ •

^ - tL-p^'%7^ ' -"^->*'^'

""■^^^jvj-

Map 56 Botrychium lunaria

340

Map 57 Botrychium minganense

Map 58 Botrychium dusenii

341

Map 59 Botrychium simplex (s. I.)

Map 60 Botrychium matricariifolium

342

/y ''lS\!^ ^ y^ V 1"? v-'vv

^"^^^ -^'C aC ^ ..-..

V v>^

Map 6 1 Botrychium boreale ssp. boreale

Map 62 Botrychium boreale ssp. obtusilobum

343

•u-(

. — < v., ->•

•.."'^%

/^>^ ^V(^-, v""^- — li^^

I"v

-v, ,' '. />^T.

f: ^ -n;^ '.

,/ ^

Map 63 Botrychium lanceolatum var. lanceolatum

' \^^ h^-A'Ji ( ^\?!i -.

Map 64 Botrychium lanceolatum var. angustisegmentum

344

Map 65 Osmunda regalis var. spectabilis

Map 66 Osmunda claytoniana

345

V*.^.^«KX§S**

^"0«.f

Map 67 Osmunda cinnamomea

-v/-^)^

Map 68 Schizaea pusilla

346

Map 69 Mecodium wrightii

-,/^ ^

"V,, yx>

Map 70 Dennstaedtia punctilobula

347

Map 7 1 Pteridium aquilinum var. latiusculum

Map 72 Pteridium aquilinum var. pubescens

348

Map 73 Cheilanthes feel

iMap 74 Cheilanthes gracillima

349

^=3^.K:,-f^

" 0 Ixs.£^

^v.^"■-^^

Map 75 Aspidotis densa

Map 76 Pellaea atropurpurea

350

Map 77 Pellaea glabella var. glabella

-i.rrn(i^

"<4.

"^i<L

•"V,

Map 78 Pellaea glabella var. nana

351

Map 79 Pellaea glabella var. simplex

Map 80 Cryptogramma stelleri

352

Map 8 1 Cryptogramma crispa var . acrostichoides

^^'u^m^

Map 82 Cryptogramma crispa var. sitcherisis

353

'"^"^■^■'^

Map 83 Pityrogramma triangularis

Map 84 Adiantum capillus-veneris

354

Map 85 Adiantum pedatum ssp. pedatum

Map 86 Adiantum pedatum ssp. aleuticum

355

Map 87 Adiantum pedatum var. subpumilum

Map 88 Adiantum pedatum ssp. calderi

356

Map 89 Matteuccia struthiopteris var. pensylvanica

Map 90 Onoclea sensibilis

357

^-^

M ap 9 1 Woods ia glabella

Map 92 Woodsia ilvensis

358

Map 93 Woods ia alpina

Map 94 Woodsia oregana

359

Map 95 Woodsia scopulina

Map 96 Woodsia obtusa

360

Map 97 Polystichum acrostichoides

:>'rV

v/^"4

Map 98 Dryopteris lonchitis

361

Map 99 Polystichum lemmonii

Map 100 Polystichum imbricans

362

Map 101 Polystichum kruckebergii

-^x^

r-^-^^M'^-WK

Map 102 Polystichum scopulinum

363

X.'

.'^.

:^^

"i ' ' -?-<, ^ /»«*^

7 .. 7J

T-<

' ;

"%r^

• *5'

■:.^.r),

Map 103 Polystichum munitum

■^»*«v',-etfK'^is:--.

Map 104 Polystichum andersonii

364

Map 105 Polystichum californicum

'"'V;/'"-'"\g9

V :jf

M ap 1 06 Polystich um bra unii

365

^^'3^m^, ""-^

Map 107 Polystichum setigerum

Map 108 Dryopteris arguta

366

■^w^

Map 109 Dryopteris fragrans

Map 1 1 0 Dryopteris intermedia

367

Map 111 Dryopteris expansa

Map 112 Dryopteris campyloptera

368

Map 113 Dryopteriscartbusiana

Map 1 14 Dryopteris filix-mas

369

-s^^"^^

Map 115 Dryopteris marginalis

Map 116 Dryopteris goldiana

370

Map 117 Dryopteris cristata

Map 118 Dryopteris clintoniana

371

-vx^

Map 119 Gymnocarpium dryopteris ssp. dryopteris

Map 1 20 Gymnocarpium dryopteris ssp. disjunctum

372

Map 121 Gymnocarpium jessoense ssp. parvulum

Map 122 Gymnocarpium robertianum

373

Map 123 Thelypteris limbosperma

Map 124 Thelypteris nevadensis

374

■m

■A.

Map 125 Thelypteris noveboracensis

Map 1 26 Thelypteris palustris var. pubescens

375

tvT^

■--^..>-^

-t- v

u».^^

Map 127 Thelypteris simulata

Map 128 Phegopteris hexagonoptera

376

Map 129 Phegopteris connectilis

Map 130 Cystopteris montana

377

Map 131 Cystopteris bulbifera

Map 132 Cystopteris protrusa

378

Map 133 Cystopteris fragilis var. fragilis

M^p 134 Cystopteris fragilis var. mackayii

379

Map 135 Cystopteris laurentiana

Map 136 Athyrium alpestre ssp. americanum

380

Map 137 Athyrium filix-femina var. cyclosorum

*rv

Map 138 Athyrium filix-femina var. michauxii

381

Map 139 Athyrium pycnocarpon

Map 140 Athyrium thelypterioides

382

'^■■^s._Aa^

Map 1 4 1 Blechnum spicant

Map 142 Woodwardia areolata

383

Map 143 Woodwardia fimbriata

Map 144 Woodwardia virginica

384

Map 145 Asplenium platyneuron

Map 146 Asplenium ruta-muraria

385

Map 147 Asplenium trichomanes

Map 148 Asplenium viride

386

Map 149 Camptosorus rhizophyllus

Map 1 50 Phyllitis scolopendrium var, americana

387

■\y-^

Map 151 Poly podium scouleri

Map 1 52 Poly podium glycyrrhiza

388

Map 153 Polypodium amorphum

Map 154 Polypodium hesperium

389

'^•^v.'*-^

Map 155 Poly podium virginianum (s. 1.)

Map 156 Marsilea vestita

390

^. ~H^ %

Map 157 Marsilea quadrifolia

Map 158 Azolla caroliniana

391

Map 159 Azolla mexicana

392

GLOSSARY

abaxial On the side of an organ away from the axis; dorsal.
acrid Sharp and harsh or unpleasantly pungent in taste or odor.
acuminate Tapering to a slender point.
adaxial Toward the axis; ventral.

I adnata Grown together or attached; applied only to unlike organs,
as stipules adnate to the petiole.
allopolyploidy The doubling or higher multiplication of chromo-
some sets from various species or genera either spontaneously or
' experimentally induced (allotetraploid, allohexaploid).

anastomosing Connecting by crossveins and forming a network.

annual Ofone year's duration.

antheridium In cryptogams the organ corresponding to an anther.

apiculate Ending abruptly in a small, usually sharp tip.

apogamous Developed without fertilization; parthenogenetic.

appressed Lying close to or parallel to an organ, as hairs appressed
to a leaf or leaves appressed to a stem.

arborescent Of large size and more or less tree-like, but without the

clear distinction of a single trunk.
archegonium The organ in the higher cryptogams that corresponds

to a pistil in the flowering plants.

areole A small space marked out upon or beneath a surface.

articulate Jointed; having nodes, joints, or places where separation

may naturally take place.
ascending Growing obliquely upward (of stems); directed obliquely

forward with respect to the organ to which they are attached (of

parts of a plant).

attenuate Gradually tapering to a very slender point.

auricle A small, ear-shaped projecting lobe or appendage at the base
of an organ.

auriculate Having an auricle.

autopolyploidy The presence of more than two (diploidy) of the
monoploid chromosome sets characteristic of the species.

axil The angle formed between any two organs.

bipinnate Doubly or twice pinnate.

bivalvate Having two valves.

blade The expanded part of a frond.

bract A more-or-less modified leaf subtending a flower, or belonging
to an inflorescence, or sometimes appearing cauline.

bristle A stiff hair, or any slender body, that may be likened to a
hog's bristle.

393

bristly Provided with bristles.

caespitose Growing in dense tufts; usually applied only to small
plants.

caudate Having a slender tail-like terminal appendage.

chaff A small thin scale or bract that becomes dry and membranous.

chaffy Having or resembling chaff.

ciliate Having marginal hairs.

clinal Series of changes in form; a gradient of biotypes along an envi-
ronmental transition.

concolorous Uniform in color.

confluent Flowing or running together.

continuous Marked by uninterrupted extension in space, time, or

sequence.
cordate Heart-shaped; sometimes applied to whole organs, but

more often to the base only.

coriaceous Leathery in texture.

corm The enlarged fleshy base of a stem, bulb-like but solid; a solid
bulb.

costa A rib; a midrib or mid nerve.

crenate Dentate with teeth much rounded.

crenulate Finely crenate.

cuneate Wedge-shaped; narrowly triangular with the acute angle
pointed downward.

cuspidate Tipped with a cusp or a sharp and firm point.

cytotype Any variety (race) of a species whose chromosome comple-
ment differs quantitatively or qualitatively from the standard
complement of that species.

deciduous Falling after completion of the normal function; not
evergreen.

decompound More than once compound or divided.

decumbent Prostrate at base, either erect or ascending elsewhere.

decurrent Extending downward from the point of intersection.

deflexed Bent or turned abruptly downward.

deltoid Broadly triangular.

dentate Toothed along the margin, the apex of each tooth sharp and

directed outward.
denticulate Minutely dentate.
dichotomous Forking more or less regularly into branches of about

equal size.
dimorphic Occurring in two forms.
diploid With 2n chromosomes per cell.
distinct Separate; not united; evident.
divergent Inclining away from each other.

394

dorsal Located on or pertaining to the back or outer surface of an
organ.

ecilate Without cilia.

echinate Provided with prickles.

elater Appendage of spores of horsetails, formed from the outermost
wall layer, coiling and uncoiling as air is dry or moist, possibly
assisting in spore dispersal.

ellipse A regular oval.

elliptical Oval in outline; having narrowed to rounded ends and
being widest at or about the middle; of, relating to, or shaped like
an ellipse.

emersed Standing out of or rising above a surface (as of a fluid).

endophyte A plant living within another plant.

entire With a continuous, unbroken margin.

epiphyte A plant growing attached to another plant, but not
parasitic.

arose Irregularly cut or toothed along the margin.

extirpate To destroy completely; eradicate.

falcate Scythe-shaped; curved and flat, tapering gradually.

fastigiate Erect and close together.

fertile Capable of normal reproductive functions.

fibrillose With fine fibers.

filiform Thread-like; long, slender, and terete.

fimbriate Fringed.

flabellate, flabelliform Fan-shaped or broadly wedge-shaped.

flaccid Flabby; lacking in stiffness.

flexuous Curved alternately in opposite directions.

frond The expanded leaf-like portion of a fern, including stipe,

rachis, and pinnae.
gametophyte In the life cycle, the generation in which sexual

organs are produced.
gemma A bud or body analogous to a bud by which some plants

propagate themselves.
genome The basic chromosome set of an organism.
glabrous Lacking pubescence; smooth.
glandular Containing or bearing glands.
glaucous Gray, grayish green, or bluish green, with a thin coat of

fine removable particles that are often waxy in texture; covered

or whitened with a bloom.

globose Spherical or nearly so.

herbaceous Without a persistent woody stem above ground; dying
back to the ground at the end of the growing season; leaf-like in
color and texture.

395

hyaline Transparent or translucent.

hybrid Produced by dissimilar parents; a cross-breed of two species.

hybridization The production of a hybrid.

imbricate Overlapping, either vertically or spirally, where the

lower piece covers the base of the next higher.
indusium The covering of the sorus.

internode The portion of a stem or other structure between two
nodes.

interspecific Between two different species.

isoenzyme A phase of an enzyme (a protein that even in low concen-
tration speeds up, enables, or controls chemical reactions in
living organisms without being used up in the reactions).

lacerate Having an irregularly jagged margin; irregularly cut as if
torn.

lamina A sheet or plate; the flat, expanded portion of a structure
such as a leaf or petal.

lanceolate Shaped like a lance head, much longer than wide and
widest below the middle.

lateral Situated on or arising from the side of an organ.

ligulate Having a ligule; having the nature of a ligule.

ligule In Isoetes, a small triangular or elongate delicate tissue ex-
tending slightly above the sporangium.

linear Narrow and elongate, with parallel sides.

loculate Having or divided into loculi.

locule A cavity or one of the cavities within an ovary, a fruit, or an
anther.

lunate Of the shape of a half- moon or crescent.

macrospore The larger kind of spore in Selaginellaceae and Isoetes,

and in other genera.
marcescent Withering and persistent.
medial Being or occurring in the middle.
megasporangium, macrosporangium Sporangium within which

megaspores are formed. In flowering plants known as the ovary.
megaspore or macrospore The larger of the two kinds of spores

produced by heterosporous ferns; the first cell of a female

gametophyte generation of these plants and of seed plants.
meiosis The reduction divisions that result in the production of four

cells from a single one, the number of chromosomes per cell being

reduced from 2n to n.
meiotic Characterized by meiosis.
membranous Thin and pliable, as an ordinary leaf, in contrast to

chartaceous, coriaceous, or succulent.
microsporangium Sporangium within which microspores are

formed. In flowering plants, the pollen sac.

396

microspore The smaller kind of spore in Selaginellaceae and

Isoetes, and in other genera.
mucro A short and small abrupt tip.
mucronate Tipped with a mucro.

mycorrhiza An association of a fungus with the root of a higher
plant.

node The place upon a stem that normally bears a leaf or whorl of
leaves; the solid constriction in the culm of a grass; a knot-like or
knob-like enlargement.

oblanceolate Lanceolate with the broadest part above the middle.

oblique Unequal-sided; slanting.

oblong Two to three times longer than broad and with nearly paral-
lel sides.

obtuse Blunt or rounded at the end.

orbicular Essentially circular.

ovate Egg-shaped; having an outline like that of an egg, with the
broader end basal.

palmate Having three or more lobes, nerves, leaflets, or branches
arising from one point; digitate.

panicle A loose, irregularly compound inflorescence with pedicel-
late flowers.

paniculate Arranged in a panicle.

papillose Bearing minute nipple-shaped projections.

paraphyses Filaments of sterile cells among sporangia.

pedunculate Born upon a peduncle.

peltate Shield-shaped and attached to the support by the lower
surface.

pendulous Hanging or drooping.

perennial A plant that continues its growth from year to year.

petiolate Having a petiole.

petiole The basal stalk-like portion of an ordinary leaf, in contrast
with the expanded blade; the support of a leaf.

pinna One of the primary divisions of a pinnate or pinnately com-
pound frond.

pinnate Compound; having branches, lobes, or leaflets arranged on
two sides of a common rachis.

pinnatitid Pinnately cleft.

pinnule A secondary pinna.

polyphyletic Of individuals derived in the course of evolution from

two interbreeding populations or phyletic stocks.
precocious Appearing or developing very early.

proliferating, proliferous Producing buds and plantlets from
leaves or as other offshoots.

397

prostrate Lying flat upon the ground.

prothallus A cellular, usually flat, thallus-like growth, resulting
from the germination of a spore, upon which sexual organs and
eventually new plants are developed.

puberulent Minutely or sparsely pubescent with scarcely elongate
hairs.

pubescent Bearing hairs on the surface.

raceme A simple inflorescence of pediceled flowers upon a common
more or less elongated axis.

racemose In racemes; or resembling a raceme.

rachis The upper part of the petiole, bearing the pinnae and contin-
uous with the stipe.

recurved Curved downward or backward.

reflexed Abruptly bent downward or backward.

reniform Kidney-shaped; wider than long, rounded in general out-
line, and with a wide basal sinus.
reticulate In the form of a network.

revolute Rolled backward, so that the upper surface of the organ is
exposed and the lower side more or less concealed.

rhizome An underground usually horizontal stem; a rootstock.

rhombic Having the outline of an equilateral parallelogram.

rhomboid A solid with a rhombic outline.

rhomboidal Having the shape of a rhomboid.

salient Something that projects outward or upward from its sur-
roundings.

scabrous Rough to the touch, owing to the structure of the epi-
dermes or the pressure of short, stiff hairs.

scarious Thin, dry, and membranous; not green.

scurfy Covered with scale-like or bran-like particles.

serrate Toothed along the margin, the apex of each tooth sharp
(compare crenate) and directed forward (compare dentate).

serrulate Finely serrate.
sessile Without a stalk of any kind.
seta A bristle.
setiform Like a bristle.
siliceous Composed of or abounding in silica.
sinus The cleft or recess between two lobes.

sorus (pi. sori) A heap or cluster of sporangia bearing the spores.
spatulate Shaped like a spatula; maintaining its width or somewhat
broadened toward the rounded summit; spoon-shaped.

spicule A minute, slender, pointed, usually hard body.

spinule A small spine.

spinulose Bearing small spines over the surface.

398

sporangium The globular organ in which the spores are produced.
spore An asexual reproductive cell that germinates into a prothal-
lus, which in turn gives rise to sexual reproduction.

sporocarp The fruit cases of certain cryptogams that contain spor-
angia or spores.

sporophyll A specialized organ for the production of spores in
sporangia.

sporophyte In the life cycle, the generation in which spores are
produced.

sporulation The formation of spores.

stellate, stelliform Star-shaped.

stipe The lower part of the petiole, which does not bear pinnae.

stoma (pi. stomata) A minute orifice or mouth-like opening between
two guard cells in the epidermis, particularly on the lower
surface of the leaves, through which gaseous interchange
between the atmosphere and the intercellular spaces of the
parenchyma is effected.

stramineous Straw-colored.

strigose Having appressed, sharp, straight, and stiff hairs pointing
in the same direction.

strobile An inflorescence resembling a spruce or fir cone, partly

made up of imbricated bracts or scales.
sub- (prefix) Slightly; more or less; somewhat.
subcylindrical Slightly or somewhat cylindrical.
submersed Growing, or adapted to growing, under water.
subulate Awl-shaped.
succulent Juicy; fleshy.

sympatric With areas of distribution that coincide or overlap.
tangential Of, relating to, or of the nature of a tangent.
ternate Arranged in threes.
tetraploid With 4m chromosomes per cell.
tomentose Woolly, with an indument of crooked, matted hairs.
trapezoid A quadrilateral having only two sides parallel.
trigonous Three-angled.
tripinnate Three times pinnate.
triploid With 3m chromosomes per cell.
truncate Ending abruptly, as if cut off.
tuberculate Bearing small processes or tubercules.

tubercule A small tuber or tuber-like (not necessarily subterran-
ean) body, often formed as the result of a symbiotic relation of
organisms.

ultramafic Rock types in which the elemental composition is largely
silicates of iron and magnesium.

399

vallecula Applied to the grooves in the intervals between the
ridges, as in the stems oi Eqaisetum. Vallecular: pertaining to
such grooves.

velum The membranous indusium in Isoetes.

ventral Belonging to the anterior or inner face of an organ, as op-
posed to dorsal; adaxial.
verticillate Arranged in a whorl.

vicariad One of two or more related organisms that occur in similar
environments but in distinct and often widely separated areas.

villous Covered densely with fine long hairs but not matted.

whorl Leaves or other plant parts arranged in a circle around the
stem.

xeric Characterized by, relating to, or requiring only a small
amount of moisture.

400

REFERENCES

Alston, A. H.G. 1955. The heterophyllous Selaginellae of continental
North America. Bull. Br. Mus. (Nat. Hist.) Bot. l(8):221-274.

Alt, K.S.; Grant, V. 1960. Cytotaxonomic observations on the gold-
backfern. Brittonia 12:153-70.

Argus, G.W.; White, D.J. 1977. The rare vascular plants of Ontario.
Syllogeus 14:1-63.

Argus, G.W.; White, D.J. 1978. The rare vascular plants of Alberta.
Syllogeus 17:1-46.

Argus, G.W.; White, D.J. 1983. Atlas of the rare vascular plants of
Ontario, Part 2. National Museum of Natural Sciences, Ottawa,
Ont.

Barclay-Estrup, P.; Hess, G.V. 1974. Adder's tongue fern, Op/iio^/os-
sum uulgatum L. in northwestern Ontario. Can. Field-Nat.
88:217-219.

Beitel, J.M. 1979a. The clubmosses Lycopodium sitchense and L. sab-
sinifolium in the Upper Great Lakes region. Mich. Bot. 18:3-13.

Beitel, J.M. 19796. Clubmosses {Lycopodium) in North America.
Fiddlehead Forum 6:1-8.

Beitel, J.M.; Wagner, F.S. 1982. The chromosomes of Lycopodium
lucidulum. Am. Fern J. 72:33-35.

Billington, C. 1952. Ferns of Michigan. Cranbrook Inst. Sci. Bull. 32.
240 pp.

Blasdell, R.F. 1963. A monographic study of the fern genus Cjs^op-
teris. Mem. Torrey Bot. Club 21:1-102.

Bloom, W. 1955. Comparative viability of sporocarps of Marsilea
quadrifolia in relation to age. Trans. 111. State Acad. Sci. 47:72-
76.

Bloom, W. 1961. Heat resistance of sporocarps of Marsilea quadri-
folia. Am. Fern J. 51:95-97.

Boivin, B. 1952. Two variations of P^ericf mm a<7wi/mum. Am. Fern J.
42:131-133.

Boivin, B. 1960a. Centurie de plantes canadiennes III. Nat. Can.
(Que.) 87:25-49.

Boivin,B. 19606. Anew Equisetum. Am. FernJ. 50:107-109.

Boivin, B. 1962. Etudes pteridologiques. II: Gymnocarpium New-
man. Bull. Soc. Bot. Fr. 109:127-128.

Boivin, B. 1966a. Etudes pteridologiques. Ill: Variations du Wood-
siaoregana. Bull. Soc. Bot. Fr. 113:407-409.

401

Boivin, B. 19666. Notes sur les Lycopodium du Canada. Nat. Can.
(Que.) 93:355-359.

Boivin, B. 1968. Enumeration des plantes du Canada. Provan-
cheria 6.

Bottarelli, F. 1968. On poisoning: Pteris aquilina, Equisetum aruense
and E.palustre. Veterinaria (Milano) 17:308-322.

Bouchard, A.; Barabe, D.; Hay, S. 1977. An isolated colony of Oreop-
teris limbosperma (All.) Holub in Gros Morne National Park,
Newfoundland, Canada. Nat. Can. (Que.) 104:239-244.

Bouchard, A.; Barabe, D.; Dumais, M.; Hay, S. 1983. The rare vascu-
lar plants of Quebec. Syllogeus 48: 1-75.

Bouchard, A.; Hay, S.G. 1974. Addition a la flore de Terre-Neuve:
Lycopodium alpinum L. Nat. Can. (Que.) 101:803.

Bouchard, A.; Hay, S.G. 1976. Thelypteris limbosperma in eastern
North America. Rhodora 78:552-553.

Braun, M. 1938. Index to North American ferns. Published by the
author, Orleans, Mass. 217 pp.

Britton, D.M. 1953. Chromosome studies in ferns. Am. J. Bot.
40:575-583.

Britton, D.M. 1964. Chromosome numbers of ferns in Ontario. Can.
J. Bot. 42:1349-1354.

Britton, D.M. 1965. Hybrid wood ferns in Ontario. Mich. Bot. 4:3-9.

Britton, D.M. 1967. Diploid Dryopteris dilatata from Quebec.
Rhodora 69:1-4.

Britton, D.M. 1968. The spores of four species of spinulose wood ferns
{Dryopteris) in eastern North America. Rhodora 70:340-347.

Britton, D.M. 1972a. Spore ornamentation in the Dryopteris spinu-
/osa complex. Can. J. Bot. 50:1617-1621.

Britton, D.M. 19726. The spores of Dryopteris clintoniana and its
relatives. Can. J. Bot. 50:2027-2029.

Britton, D.M. 1972c. Spinulose wood ferns, Dryopteris, in western
North America. Can. Field-Nat. 86:241-247.

Britton, D.M. 1974. The significance of chromosome numbers in
ferns. Ann. Mo. Bot. Gard. 61:310-317.

Britton, D.M. 1976. The distribution of Dryopteris spinulosa and its
relatives in eastern Canada. Am. Fern J. 66:69-74.

Britton, D.M. 1977. The fern Woodsia obtusa (Spreng.) Torrey in
Ontario. Can. Field-Nat. 91:84-85.

Britton, D.M.; Jermy, A.C. 1974. The spores of Dryopteris filix-mas
and related taxa in North America. Can. J. Bot. 52:1923-1926.

402

Britton, D.M.; Legault, A.; Rigby, S.J. 1967. Pellaea atropurpurea
(L.) Link and Pellaea glabella Mett in Quebec. Nat. Can. (Que.)
94:761-763.

Britton, D.M.; Rigby, S.J. 1968. In search of the purple cliff-brake.
Ont. Nat. 5:5-7, 12.

Britton, D.M.; Soper, J.H. 1966. The cytology and distribution of
Dryopteris species in Ontario. Can. J. Bot. 44:63-78.

Britton, D.M.; Widen, C.-J. 1974. Chemotaxonomic studies on
Dryopteris from Quebec and eastern North America. Can. J. Bot.
52:627-638.

Britton, D.M.; Widen, C.-J.; Brunton, D.F.;Keddy, P.A. 1975. Anew
hybrid woodfern, Dryopteris X algonquinensis D.M. Britton,
from Algonquin Park, Ontario. Can. Field-Nat. 89:163-171.

Brown, D.F.M. 1964. A monographic study of the fern genus
Woodsia. Nova Hedwigia 16:1-154.

Bruce, J.G. 1975. Systematics and morphology of subgenus Lepidotis
of the genus Lycopodium (Lycopodiaceae). Ph.D. thesis,
University of Michigan, Ann Arbor, Mich.

Bruce, J.G. 1976. Gametophytes and subgeneric concepts in Lycopo-
dium. Am. J. Bot. 63:919-924.

Brunton, D.F. 1972. More slender cliff-brake in the Ottawa District.
Trail & Landscape 6(3): 92-93.

Brunton, D.F. 1978. The holly fern in Alberta. Blue Jay 36:82-83.

Brunton, D.F. 1979. Taxonomy, distribution, and ecology of the cliff-
brake ferns {Pellaea: Polypodiaceae) in Alberta. Can. Field-Nat.
93:288-295.

Brunton, D.F.; Lafontaine, J.D. 1974. The distribution of Pellaea in
Quebec and eastern Ontario. Nat. Can. (Que.) 101:937-939.

Buck, W.R. 1977. A new species oi Selaginella in the S. apoda com-
plex. Can. J. Bot. 55:366-371.

Burgess, T.J.W. 1886. Recent additions to Canadian Filicineae, with
new stations for some of the species previously recorded. Trans.
Roy. Soc. Can., Sect. IV:9-18.

Burman, W.A. 1909. Flora of Manitoba. Pages 156-182 in British
Association for the Advancement of Science: A handbook to
Winnipeg and the province of Manitoba. Winnipeg, Man.

Butters, F.K. 1917. The genus Athyrium and the North American
ferns allied to Athyrium filix-femina. Rhodora 19: 170-207.

Calder,J.A.;Taylor, R.L. 1968. Flora of the Queen Charlotte Islands,
Part 1. Can. Dep. Agr. Res. Branch Monogr. 4(1). 659 pp.

Campbell, C.A.; Britton, D.M. 1977. Pteridophytes of the Regional
Municipality of Waterloo, Ontario. Can. Field-Nat. 91:262-268.

403

Campbell, R. 1898-1899. Canadian ferns. Can. Hort. Mag. 2:81-87,
131-137, 157-165, 183-192, 209-218, 235-244, 261-270.

Carlson, T.M.; Wagner, W.H. 1982. The North American distribu-
tions of the genus Dryopteris. Contrib. Univ. Mich. Herb.
15:141-162.

Catling, P.M. 1975. Alpine woodsia {Woodsia alpina (Bolton) S.F.
Gray) in southeastern Ontario. Can. Field-Nat. 89:177-178.

Ching, R.C. 1963. A reclassification of the family Thelypteridaceae
from the mainland of Asia. Acta Phytotaxon. Sin. 8:289-335.

Clausen, R.T. 1938. A monograph of the Ophioglossaceae. Mem.
Torrey Bot. Club 19(2):1-177.

Cody, W.J. 1963. Woodwardia in CanadsL. Am. Fern J. 53:17-27.

Cody, W.J. 1978. Ferns of the Ottawa District (rev. ed.). Agric. Can.
Publ.974. 111pp.

Cody, W.J. 1979. Vascular plants of restricted range in the continen-
tal Northwest Territories. Syllogeus 23; 1-57.

Cody, W.J. 1980. Fougeres du district d'Ottawa. Agric. Can. Publ.
974. 112 pp.

Cody, W.J. 1982. Adiantum pedatum ssp. calderi, a new subspecies in
northeastern North America. Rhodora 85:93-96.

Cody, W.J. ; Crompton, C.W. 1975. The biology of Canadian weeds.
15: Pteridium aquilinum (L.) Kuhn. Can. J. Plant Sci. 55:1059-
1072.

Cody, W.J.; Hall, I.V.; Crompton, C.W. 1977. The biology of Cana-
dian weeds. 26: Dennstaedtia punctilobula (Michx.) Moore. Can.
J. Plant Sci. 57:1159-1168.

Cody, W.J.; Lafontaine, J.D. 1975. The fern genus Woodsia in Mani-
toba. Can. Field-Nat. 89:66-69.

Cody, W.J.; Mulligan, G.A. 1982. Chromosome numbers of some
Canadian ferns and fern allies. Nat. Can. (Que.) 109:273-275.

Cody, W.J.; Wagner, V. 1981. The biology of Canadian Weeds. 49:
EquisetumarvenseL. Can. J. Plant Sci. 61:123-133.

Copeland, E.B. 1947. Genera Filicum. Chronica Botanica Co.,
Waltham, Mass. 247 pp.

Cordes, L.D.; Krajina, V.J. 1968. Mecodium wrightii on Vancouver
Island. Am. Fern J. 58:181.

Cranfill, R. 1980. Ferns and fern allies of Kentucky. Kentucky
Nature Preserves Commission, Scientific and Technical Series,
No. 1. 284 pp.

Cruise, J. 1972. Spring harvest. Ont. Nat. 11:18-23.

404

Czerepanov, S.K. 1981. Plantae Vasculares URSS. Nauka, Lenin-
grad. 509 pp.

DeBenedictis, V.M.M. 1969. Apomixis in ferns with special reference
to sterile hybrids. Ph.D. dissertation, University of Michigan,
Ann Arbor, Mich. 203 pp.

Douglas, G.W.; Argus, G.W.; Dickson, H.L.; Brunton, D.F. 1981. The
rare vascular plants of the Yukon. Syllogeus 28:1-61.

Duckett, J.G. 1979. An experimental study of the reproductive biol-
ogy and hybridization in the European and North American
species of Equisetum. Bot. J. Linn. Soc. 79:205-229.

Eaton, D.C. 1879. The ferns of North America, Vol. 1. Cassino,
Salem, Mass. 352 pp.

Eifert, V.S.; Metcalfe, B. 1963. Native ferns of eastern North
America. Federation of Ontario Naturalists, Toronto, Ont.
64 pp.

Emmott, J.L 1964. A cytogenetic investigation in a Phyllitis-
Asplenium complex. New Phytol. 63:306-318.

Erskine, D.S. 1961. The plants of Prince Edward Island. Agric. Can.
Publ. 1088. 270 pp.

Evans, A.M. 1970. A review of systematic studies of the pterido-
phytes of the southern Appalachians. Pages 117-146 in Holt,
P.C. ed. The distributional history of the biota of the southern
Appalachians, Part II. Flora Research Division, Monogr. 2.
Virginia Polytechnic Institute and State University, Blacksburg,
Va.

Evans, A.M.; Wagner, W.H. 1964. Dryopteris goldiana X intermedia:
A natural woodfern cross of noteworthy morphology. Rhodora
66:255-266.

Fernald, M.L. 1928. The American representatives of Asplenium
ruta-muraria. Rhodora 30:37-43.

Fernald, M.L. 1945. Botanical specialties of the Seward Forest and
adjacentareasof southeastern Virginia. Rhodora 47:93-142.

Fernald, M.L. 1950. Gray's manual of botany (8th ed.). American
Book Co., New York. 1632 pp.

Fraser-Jenkins, C.R. 1980. Nomenclatural notes on Dryopteris. 4.
Taxon 29:607-612.

Gibby, M. 1977. The origin of Dryopteris campyloptera. Can. J. Bot.
55:1419-1428.

Gibby, M.; Walker, S. 1977. Further cytogenetic studies and reap-
praisal of the diploid ancestry in the Dryopteris carthusiana
complex. Fern Gaz. 11:315-324.

405

Gleason, H.A.; Cronquist, A. 1963. Manual of the vascular plants of
northeastern United States and adjacent Canada. Van Nostrand
Co., Princeton, N.J. 810 pp.

Greenwood, E.W. 1967. Mass occurrences of the fern Ophioglossum
uulgatum in the Ottawa District, Ontario. Can. Field-Nat.
81:186-188.

Gussow, H.T. 1912. Field horsetail {Equisetum aruense L.) Pages
210-211 m Agric. Can. Exp. Farm Rep. Ottawa, Ont.

Hagenah, D.J. 1961. Spore studies in the genus Cystopteris. I: The
distribution of Cystopteris with non-spiny spores in North
America. Rhodora 63:181-193.

Hagenah, D.J. 1963. Pteridophytes of the Huron Mountains. Mich.
Bot. 2:73-93.

Harms, V.L. 1978. Athyrium filix-femina new to Saiskaitchew3.n. Am.
Fern J. 68:119-120.

Harms, V.L. 1983. The lady fern, Athyrium filix-femina, in Saskat-
chewan. Am. Fern J. 73:117—21.

Hauke, R.L. 1961. The smooth scouring rush and its complexities.
Am. Fern J. 50:185-193.

Hauke, R.L. 1963. A taxonomic monograph of the genus Equisetum
subgenus Hippochaete. Beih. Nova Hedwigia 8:1-123.

Hauke, R.L. 1966. A systematic study of Equisetum aruense Beih.
Nova Hedwigia 13:81-109.

Hauke, R.L. 1978. A taxonomic monograph of E(jatse^wm subgenus
Equisetum. Beih. Nova Hedwigia 30:385-455.

Hersey, R.E.; Britton, D.M. 1981. A cytological study of three species
and a hybrid taxon of Lycopodium (Section complanata) in
Ontario. Can. J. Genet. Cytol. 23:497-504.

Hickey, R.J. 1977. The Lycopodium obscurum complex in North
America. Am. Fern J. 67:45-48.

Hickey, R.J.; Beitel, J.M. 1979. A name change for Lycopodium
flabelliforme. Rhodora 81:137-140.

Hinds, H.R. 1983. The rare vascular plants of New Brunswick.
Syllogeus 50:1-38.

Hitchcock, C.L.; Cronquist, A.; Ownbey, M.; Thompson, J.W. 1969.
Vascular plants of the Pacific Northwest, Part 1. University of
Washington Press, Seattle, Wash. 914 pp.

Holmgren, P.K.; Keuken, W.; Schofield, E.K. 1981. Index Herbario-
rum, Part I (7th ed.). Regnum Veg. 106:1-452.

Holtum, R.E. 1971. Studies in the family Thelypteridaceae. Ill: A
new system of genera in the Old World. Blumea 19:2-52.

406

Holub, J. 1964. Lycopodiella, novy rod radu Lycopodiales. Preslia
(Prague) 36:16-22.

Holub, J. 1969. Oreopteris, a new genus of the family Thelypteri-
daceae. Folia Geobot. Phytotaxon. 4:33-53.

Holub, J. 1975. Diphasiastrum, a new genus in Lycopodiaceae.
Preslia (Prague) 47:97-110.

Hulten, E. 1967. Comments on the flora of Alaska and Yukon. Ark.
Bot. 7(1):1-147.

Hulten, E. 1968. Flora of Alaska and neighboring territories.
Stanford University Press, Stanford, Calif. 1008 pp.

Iwatsuki, I. 1961. The occurrence o^ Mecodium wrightii in Canada.
Am. Fern J. 51:141-144.

Jermy, A.C.; Arnold, H.R.; Farrell, L.; Perring, F.H. 1978. Atlas of
ferns of the British Isles. Botanical Society of the British Isles
and the British Pteridological Society, London. 1-101.

Jermy, A.C.; Crabbe, J. A.; Thomas, B.A. 1973. The phylogeny and
classification of the ferns. Bot. J. Linn. Soc. 67 (Suppl. 1). 284 pp.

Jermy, A.C.; Page, C.N. 1980. Additional field characters separating
the subspecies of Asplenium trichomanes in Britain. Fern Gaz.
12:112-113.

Johnson, A. W.; Packer, J. G. 1968. Chromosome numbers in the flora
of Ogotoruk Creek, N.W. Alaska. Bot. Not. 121:403-456.

Knobloch, I.W. 1967. Chromosome numbers in Cheilanthes, Notho-
laena, Llauea and Polypodium. Am. J. Bot. 54:461-464.

Knobloch, I.W. 1976. Pteridophyte hybrids. Mich. State Univ. Mus.
Biol. Ser. 5(4):277-352.

Kott, L. 1980a. Polystichum hraunii in Waterloo County. Ont. Field
Biol. 34:47-48.

Kott, L. 19806. The taxonomy and biology of the genus Isoetes L. in
northeastern North America. Ph.D. thesis. University of
Guelph, Guelph, Ont. 234 pp.

Kott, L.S. 1981. Isoetes acadiensis, a new species from eastern North
America. Can. J. Bot. 59:2592-2594.

Kott, L.S. ; Bobbette, R.W.S. 1980. /soe^es ea^onii, a quillwort new for
Canada. Can. Field-Nat. 94:163-166.

Kott, L.S.; Britton, D.M. 1980. Chromosome numbers for Isoetes in
northeastern North America. Can. J. Bot. 58:980-984.

Kott, L.S. ; Britton, D.M. 1982a. Comparison of chromatographic spot
patterns of some North American Isoetes species. Am. Fern J.
72:15-18.

407

Kott, L.S.; Britton, D.M. 1982/). A comparative study of sporophyte
morphology of the three cytotypes o\ Poly podium uirginianum in
Ontario. Can. J. Bot. 60:1360-1370.

Kruckeberg, A.L. 1982. Noteworthy collections, British Columbia.
Madrono 29:271.

Kruckeberg, A.R. 1964. Ferns associated with ultramafic rocks in
the Pacific Northwest. Am. Fern J. 54:113-126.

Lafontaine, J.D. 1973. Range extension of the blunt-lobed woodsia,
Woodsia obtusa (Spreng.) Torr. (Polypodiaceae) in Canada. Can.
Field-Nat. 87:56.

Lafontaine, J.D.; Brunton, D.F. 1972. The purple cliff-brake, Pellaea
atropurpurea (L.) Link, in western Quebec. Can. Field-Nat.
86:297-298.

Lang, F. A. 1969. A new name for a species of Po/jpodmm from north-
western North America. Madrono 20:53-60.

Lang, F.A. 1971. The Poly podium uulgare complex in the Pacific
Northwest. Madrono 21:235-254.

Lawson, G. 1864. Synopsis of Canadian ferns and filicoid plants.
Edinb. New Philos. J. n.s. 19:273-290.

Lawson, G. 1889. The school fern-flora of Canada. Pages 221-251 in
A. Gray, Botany for young people and common schools: How
plants grow, a simple introduction to structural botany with a
popular flora. Mackinlay, Halifax, N.S.

Legault, A.; Blais, V. 1968. Le Cheilanthes siliquosa Maxon dans le
nord-est Americain. Nat. Can. (Que.) 95:307-316.

Lellinger, D.B. 1968. A note on Aspidotis. Am. Fern J. 58:140-141.

Lellinger, D.B. 1981. Notes on North American ferns. Am. Fern J.
71:90-94.

Love, A.; Love, D. 1961. Some chromosome numbers of Icelandic
ferns and fern-allies. Am. Fern J. 51:127-128.

Love, A.; Love, D. 1966a. Cytotaxonomy of the alpine vascular plants
of Mount Washington. Univ. Colo. Stud. Ser. Biol. 24:1-74.

Love, A.; Love, D. 19666. The variation of Blechnum spicant. Bot.
Tidsskr. 62:186-196.

Love, A.; Love, D. 1968. Cytotaxonomy of B/ec/inwm spicant Collect.
Bot. (Bare.) 7:665-676.

Love, A.; Love, D. 1976. Pages 483-500 in A. Love, lOPB chromosome
number reports LIII. Taxon 25:483-500.

Love, A.; Love, D. 1977. New combinations in ferns. Taxon
26:324-326.

Love, A.; Love, D.; Kapoor, B.M. 1971. Cytotaxonomy of a century of
Rocky Mountain orophytes. Arc. Alp. Res. 3:139-165.

408

Love, A.; Love, D.; Pichi Sermolli, R.E.G. 1977. Cytotaxonomical
atlas of the Pteridophyta. Cramer, Vaduz, Liechtenstein. 398 pp.

Lovis, J.D. 1964. The taxonomy of Asp/e/iium Mc/iomanes in Europe.
Br. Fern Gaz. 9:147-160.

Lovis, J.D. 1977. Evolutionary patterns and processes in ferns. Adv.
Bot. Res. 4:229-415.

Lovis, J.D. ; Melzer, H.; Reichstein, T. 1965. Asplenium X adulterini-
forme hybr. nov. = diploides Asplenium trichomanes L. X A.
viride Hudson. Bauhinia 2(3):231-237.

Lucansky, T.W. 1981. Chain ferns of Florida. Am. Fern J.
71:101-108.

Macoun, J. 1890. Catalogue of Canadian plants. Part V: Acrogens.
Brown, Montreal, Que.

Macoun, J.; Burgess, T.J.W. 1884. Canadian Filicineae. Trans. R.
Soc. Can., Sect. IV: 163-226.

Maher,R.V.; Argus, G.W; Harms, V.L.; Hudson, J. H. 1979. The rare
vascular plants of Saskatchewan. Syllogeus 20:1-55.

Maher, R.V., White, D.J.; Argus, G.W.; Keddy, P.A. 1978. The rare
vascular plants of Nova Scotia. Syllogeus 18:1-37.

Manton, I. 1950. Problems of cytology and evolution in the Pterido-
phyta. Cambridge University Press, Cambridge, England.
316 pp.

Manton, I.; Sledge, W.A. 1954. Observations on the cytology and tax-
onomy of the pteridophyte flora of Ceylon. Philos. Trans. R. Soc.
Lond. Biol. Sci. 238:127-185.

Manton, L; Vida, G. 1968. Cytology of the fern flora of Tristan da
Cunha. Proc. R. Soc. Lond. Biol. Sci. 170:361-379.

Marie- Victorin, Frere. 1923. Les Filicinees du Quebec. Contrib. Inst.
Bot. Univ. Montreal 2:1-98.

Marie- Victorin, Frere. 1925. Les Lycopodinees du Quebec et leurs
formes mineures. Contrib. Inst. Bot. Univ. Montreal 3:1-117.

Marquis, R.J.; Voss, E.G. 1981. Distributions of some western North
American plants disjunct in the Great Lakes region. Mich. Bot.
20:53-82.

McCord, D.R. 1864. Notes on the habitats and varieties of some
Canadian ferns. Can. Nat. Geol. 1:354-362.

Mehra, P.N.; Loyal, D.S. 1959. Cytological studies in Marsilea with
particular reference to Marsilea minuta L. Res. Bull. Panjab
Univ. Sci. 10:357-374.

Mickel, J.T. 1979. How to know the ferns and fern allies. Brown,
Dubuque, Iowa. 229 pp.

409

Montgomery, J. D. 1982. Drjop/eris in North America. Part II: The
hybrids. Fiddlehead Forum 9:23-30.

Montgomery, J. D.; Paulton, E.M. 1981. Dryop^eris in North America.
Fiddlehead Forum 8:25-31.

Moran, R.C. 1981. X Asplenosorus shawneensis, a new natural hy-
brid between Asplenium trichomanes and Camptosorus
rhizophyllus. Am. Fern J. 71:85-89.

Moran, R.C. 1982. The Asplenium trichomanes complex in the
United States and adjacent Canada. Am. Fern J. 72:5-11.

Moran, R.C. 1983. Cystopteris tenuis (Michx.) Desv.: A poorly under-
stood species. Castanea 48(3):218-223.

Morton, C. V. 1950. Notes on the ferns of the eastern United States.
Am. Fern J. 40:213-225,241-252.

Morton, C.V. 1970. Recent fern literature. Am. Fern J. 60:126-127.

Moss, E.H. 1959. Flora of Alberta. University of Toronto Press,
Toronto, Ont. 546 pp.

Muenscher, W.C. 1955. Weeds (2nd ed.). Macmillan, New York.
577 pp.

Mulligan, G.A.; Cinq-Mars, L.; Cody, W.J. 1972. Natural inter-
specific hybridization between sexual and apogamous species of
the beech fern genus Phegopteris Fee. Can. J. Bot. 50:1295-1300.

Mulligan, G.A.; Cody, W.J. 1968. Pages 285-288 in A. Love, lOPB
chromosome number reports XVII. Taxon 17.

Mulligan, G.A.; Cody, W.J. 1969. The highest chromosome number
known to occur in a North American plant. Can. Field-Nat.
83:277-278.

Mulligan, G. A.; Cody, W.J. 1979. Chromosome numbers in Canadian
Phegopteris. Can. J. Bot. 57:1815-1819.

Ogden, E.B. 1948. The ferns of Maine. University of Maine Studies,
Second Series, 62. 128 pp.

Parnis, E.M. Ferns of the Montreal region. McGill University Herb-
arium, Sainte-Anne de Bellevue, Que. 11 pp.

Pfeiffer, N. 1922. Monograph on the Isoetaceae. Trans. Acad. Sci.
(St. Louis) 9(2):79-233.

Pohl, R. 1955. Toxicity of ferns and Equisetum. Am. Fern J.
45:95-97.

Porsild, A.E. 1945. The so-called Woodsia alpina in North America.
Rhodora 47:145-148.

Presl, K.B. 1851. Epimeliae Botanicae. Abh. K. Boehm. Gesell.
Wiss. 5(6):361-624.

410

Pryer, K.M. 1981. Systematic studies in the genus Gymnocarpium
Newm. in North America. M.Sc. thesis, University of Guelph,
Guelph, Ont. 166 pp.

Pursh, F. 1814. Flora Americae septentrionalis. White, Cochrane,
London. 2 vols. 751 pp.

Rapp,W.F. 1954. The toxicity of Eqaise^^/m. Am. Fern J. 44:148-154.

Reed, C.F. 1954. Index Marsileata and Salviniata. Bol. Soc. Broteri-
ana 28:5-61.

Reeves, T. 1977. The genus Botrychium (Ophioglossaceae) in
Arizona. Am. Fern J. 67:33-39.

Reichstein, T. 1981. Hybrids in European Aspleniaceae (Pterido-
phyta). Rot. Helvitica 91:89-139.

Reznicek, T.A. 1972. Woodsia scopulina in Algonquin Park, Nipis-
sing District, Ontario. Can. Field-Nat. 86:368-369.

Richter, H.E. 1961. Equisetum palustre poisoning in horses. Wien.
tieraerztl. Monatsschr. 48:761-762.

Rigby, S.J. 1973. Chromosome pairing in obligately apogamous
ferns: Pellaea atropurpurea and Pellaea glabella var. glabella.
Rhodora 75:122-131.

Rigby, S.J., Britton, D.M. 1970. The distribution of Pellaea in
Canada. Can. Field-Nat. 84:137-144.

Roland, A.E. 1941. The ferns of Nova Scotia. Proc. N.S. Inst. Sci.
20(3):64-120.

Roland, A.E. ; Smith, E.C. 1969. The flora of Nova Scotia. Nova
Scotia Museum, Halifax, N.S.

Rousseau, C. 1974. Geographie floristique du Quebec/Labrador. Les
Presses de TUniversite Laval, Quebec, Que. 799 pp.

Sarvela, J. 1978. A synopsis of the fern genus Gjmn-ocarpmm. Ann.
Bot. Fenn. 15:101-106.

Sarvela, J. 1980. Gymnocarpium hybrids from Canada and Alaska.
Ann. Bot. Fenn. 17:292-295.

Sarvela, J.; Britton, D.M.; Pryer, K. 1981. Studies on the Gymnocar-
pium robertianum complex in North America. Rhodora 83:421-
431.

Scoggan, H.J. 1957. Flora of Manitoba. Natl. Mus. Can. Bull. 140.
619 pp.

Scoggan, H.J. 1978. The flora of Canada. Natl. Mus. Nat. Sc. Publ.
Bot. 7(2):93-545.

Shivas, M.G. 1961. Contributions to the cytology and taxonomy of
species of Polypodium in Europe and America. I: Cytology. J.
Linn. Soc. Lond. Bot. 58:13-25.

411

Smith, A.R. 1971. Chromosome numbers of some New World species
o^Thelypteris. Brittonia 23:354—360.

Smith, A.R. 1975. The California species of Aspidotis. Madrono
23:15-24.

Soper, J.H. 1949. The vascular plants of southern Ontario. Depart-
ment of Botany, University of Toronto, Toronto, Ont. 95 pp.

Soper, J.H. 1954. The hart's-tongue fern in Ontario. Am. Fern J.
44:129-147.

Soper, J.H. 1955. Asplenium cryptolepis on Manitoulin Island. Am.
Fern J. 45:97-104.

Soper, J.H. 1963. Ferns of Manitoulin Island, Ontario. Am. Fern J.
53:28-40,71-81,109-123.

Soper, J.H. 1964. The slender clifTbrake. Ont. Nat. 2:20-23.

Soper, J.H.; Maycock, P.F. 1963. A community of arctic-alpine plants
on the east shore of Lake Superior. Can. J.Bot. 41:183-198.

Soper, J.H.; Rao, S. 1958. Isoetes in eastern Canada. Am. Fern J.
48:97-102.

Strasburger, E. 1907. Apogamie bei Marsi/ea. Flora 97:123-191.

Svenson, H.K. 1944. The New World species of Azolla. Am. Fern J.
34:69-84.

Tatuno, S.; Takei, M. 1969. Karyological studies in Hymenophyl-
laceae. I: Chromosomes of the genus Hymenophyllum and
Mecodium in Japan. Bot. Mag. Tokyo 82:121-129.

Taylor, R.L.; Brockman, R.P. 1966. Chromosome numbers of some
western Canadian plants. Can. J. Bot. 44:1093-1103.

Taylor, R.L.; Mulligan, G.A. 1968. Flora of the Queen Charlotte
Islands, Part 2. Can. Dep. Agr. Res. Branch Monogr. 4(2) 148 pp.

Taylor, T.M.C. 1947. New species and combinations in Woodsia,
section Per rinia. Am. Fern J. 37:84-88.

Taylor, T.M.C. 1963. The ferns and fern-allies of British Columbia.
B.C. Prov. Mus. Nat. Hist. Handb. 12. 172 pp.

Taylor, T.M.C. 1967. Mecodium wrightii in British Columbia and
Alaska. Am. Fern J. 57:1-6.

Taylor, T.M.C. 1970. Pacific Northwest ferns and their allies. Uni-
versity of Toronto Press, Toronto, Ont. 247 pp.

Taylor, T.M.C; Lang, FA. 1963. Chromosome counts in some British
Columbia ferns. Am. Fern J. 53:123-126.

Taylor, W.C.; Mohlenbrock, R.H.; Murphy, J. 1975. The spores and
taxonomy of Isoetes butleri and /. melanopoda. Am. Fern J.
65:33-38.

412

Tryon, A.F. 1957. A revision of the fern genus Pellaea section
Pellaea. Ann. Mo. Bot. Gard. 44:125-193.

Tryon, A.F. 1968. Comparisons of sexual and apogamous races in the
fern genus Pellaea. Rhodora 70:1-24.

Tryon, A.F. 1972. Spores, chromosomes and relations of the fern
Pellaea atropurpurea. Rhodora 74:220-241.

Tryon, A.F.; Britton, D.M. 1958. Cytotaxonomic studies of the fern
genus PeZZaea. Evolution 12:137-145.

Tryon, A.F.; Tryon, R. 1973. Thelypteris in northeastern North
America. Am. Fern J. 63:65-76.

Tryon, A.F.; Tryon, R. 1974. Geographic patterns in temperate
American ferns and some relationships in Thelypteris. Am. Fern
J. 64:99-104.

Tryon, A.F.; Tryon, R.; Badre, F. 1980. Classification, spores and
nomenclature of the marsh fern. Rhodora 82:461-474.

Tryon, R.M. 1941. Revision of the genus Pteridium. Rhodora
43:1-31,37-67.

Tryon, R.M. 1942. A new Drjop^eris hybrid. Am. Fern J. 32:81-85.

Tryon, R.M. 1948. Some woodsias from the north shore of Lake
Superior. Am. Fern J. 38:159-170.

Tryon, R.M. 1955. Selaginella rupestris and its allies. Ann. Mo. Bot.
Gard. 42:1-99.

Tryon, R.M. 1960. A review of the genus DcnnsZaed^ia in America.
Contrib. Gray Herb. Harv. Univ. 187:23-52.

Tryon, R.M.; Britton, D.M. 1966. A study of variation in the cyto-
types of Dryopteris spinulosa. Rhodora 68:59-92.

Tryon, R.M.; Fassett, N.C.; Dunlop, D.W.; Diemer, M.E. 1953. The
ferns and fern allies of Wisconsin (2nd ed.). University of
Wisconsin, Madison, Wise. 158 pp.

Tschermak-Woess, E.; Dolezal-Janisch, R. 1959. Uber die karyolo-
gische anatomie einiger Pteridophyten sowie auffallende unter-
schiede in kernvolumen bei Cyrtomium falcatum. Oesterr. Bot.
Z. 106:315-324.

Vincent, G. 1981. P/iegopZeris /lexa^o/iopZera, espece rare et menacee.
Bull. Soc. Anim. Jard. Inst. Bot. 6:2-24.

von Euw, J.; Lounasmaa, M.; Reichstein, T.; Widen, C. -J. 1980.
Chemotaxonomy in Dryopteris and related fern genera. Review
andevaluationof analytical methods. Stud. Geobot. 1:275-311.

Wagner, D.H. 1979. Systematics of Polystichum in western North
America north of Mexico. Pteridologia 1:1-64,

Wagner, W.H. 1954. Reticulate evolution in the Appalachian asplen-
iums. Evolution 8:103-118.

413

Wagner, W.H. 1955. Cytotaxonomic observations on North Ameri-
can ferns. Rhodora 57:219-240.

Wagner, W.H. 1959. American grapeferns resembling Botrychium
ternatum. Am. Fern J. 49:97-103.

Wagner, W.H. 1960a. Periodicity and pigmentation in Botrychium
subg. sceptridium in the northeastern United States. Bull.
Torrey Bot. Club 87:303-325.

Wagner, W.H. 19606. Evergreen grapeferns and the meanings of
infraspecific categories as used in North American
pteridophytes. Am. Fern J. 50:32-45.

Wagner, W.H. 1961. Roots and taxonomic difference between Botry-
chium oneidense and B. dissectum. Rhodora 63:164-175.

Wagner, W.H. 1962. Plant compactness and leaf production in Botry-
chium multifidum "ssp. typicum" and "forma dentatum.'* Am.
Fern J. 52:1-18.

Wagner, W.H. 1963. A biosystematic study of United States ferns.
Am. Fern J. 53:1-16.

Wagner, W.H. 1966a. Two new species of ferns from the United
States. Am. Fern J. 56:3-17.

Wagner, W.H. 19666. New data on North American oak ferns, Gym-
nocarpium. Rhodora 68:121-138.

Wagner, W.H. 1970. Evolution of Dry opter is in relation to the Ap-
palachians. Pages 147-192 in P.C. Holt, ed. Distributional
history of the biota of the southern Appalachians. Part II: Flora.
Research Division, Monogr. 2. Virginia Polytechnic Institute
and State University, Blacksburg, Va.

Wagner, W.H. 1971. The southeastern adder's-tongue, Ophioglossum
vulgatum var. pycnostichum, found for the first time in
Michigan. Mich. Bot. 10:67-74.

Wagner, W.H. 1973a. Asplenium montanum X platy neuron: A new
primary member of the Appalachian spleenwort complex for
Crowder's Mountain, N.C. J. Elisha Mitchell Sci. Soc. 89:218-
223.

Wagner, W.H. 19736. Reticulation of holly ferns (Polystichum) in the
western United States and adjacent Canada. Am. Fern J. 63:99-
115.

Wagner, W.H.; Boydston, K.E. 1978. A dwarf coastal variety of maid-
enhair fern, Acfian^ampec/a^Mm. Can. J. Bot. 56:1726-1729.

Wagner, W.H.; Chen, K.L. 1964. Pages 99-110 in A. Love and O.T.
Solbrig, lOPB chromosome number reports I. Taxon 13.

Wagner, W.H.; Chen, K.L. 1965. Abortion of spores and sporangia as
a tool in the detection of Dryopteris hybrids. Am. Fern J. 55:9-
29.

414

Wagner, W.H.; Farrar, D.R.; Chen, K.L. 1965. A new sexual form of
Pellaea glabella var. glabella from Missouri. Am. Fern J. 55:171-
178.

Wagner, W.H.; Farrar, D.R.;McAlpine,B.W. 1970. Pteridology of the
Highlands Biological Station area, southern Appalachians. J.
Elisha Mitchell Sci. Soc. 86:1-27.

Wagner, W.H.; Hagenah, D.J. 1956. A diploid variety in the Cystop-
teris fragilis complex. Rhodora 58:79-87.

Wagner, W.H.; Johnson, D.M. 1981. Natural history of the ebony
spleenwort, Asplenium platyneuron (Aspleniaceae), in the Great
Lakes area. Can. Field-Nat. 95:156-166.

Wagner, W.H.; Lord, L.P. 1956. The morphological and cytological
distinctness of Botrychium minganense and B. lunaria in
Michigan. Bull. Torrey Bot. Club 83:261-280.

Wagner, W.H.; Rawlings, D.E. 1962. A sampling o^ Botrychium subg.
Sceptridium in the vicinity of Leonardtown, St. Mary's Co., Md.
Castanea 27:132-142.

Wagner, W.H.; Wagner, F.S. 1966. Pteridophytes of the Mountain
Lake area, Giles Co. Virginia: Biosystematic studies 1964-1965.
Castanea 31:121-140.

Wagner, W.H.; Wagner, F.S. 1981. New species of moonworts, Bo^rj-
chium subg. Botrychium (Ophioglossaceae), from North America.
Am. Fern J. 71:20-30.

Wagner, W.H.; Wagner, F.S. 1982a. The taxonomy of Dryopteris X
poyseri Wherry. Mich. Bot. 21:75-88.

Wagner, W.H.; Wagner, F.S. 19826. Botrychium rugulosum (Ophio-
glossaceae): A newly recognized species of evergreen grapefern
in the Great Lakes area of North America. Contrib. Univ. Mich.
Herb. 15:315-324.

Wagner, W.H.; Wagner, F.S. 1983a. Genus communities as a sys-
tematic tool in the study of New World Botrychium
(Ophioglossaceae). Taxon 32:51-63.

Wagner, W.H.; Wagner, F.S. 19836. The moonworts of the Rocky
Mountains; Botrychium hesperium and a new species formerly
confused with it. Am. Fern J. 73:53-62.

Wagner, W.H.; Wagner, F.S. 1983c. Western Canada. Botrychium
Newsl. No. 4. University of Michigan Herbarium, Ann Arbor,
Mich. 10 pp.

Wagner, W.H.; Wagner, F.S.; Miller, C.N., Jr.; Wagner, D.H. 1978.
New observations on the royal fern hybrid, Osmunda X ruggii.
Rhodora 80:92-106.

Wagner, F.S. 1983. The Botrychium lanceolatum group in western
North America (Abstr.). Am. J. Bot. 70(5):95.

415

Walker, S. 1961. Cytogenetic studies in the Dryopteris spinulosa
complex, II. Am. J. Bot. 48:607-614.

Walker, T.G. 1979. The cytogenetics of ferns. Pages 229-415 in A. F.
Dyer. The experimental biology of ferns. Academic Press,
London.

Waterway, M.J.; Lei, T.T. 1982. Polystichum lonchitis in central
Quebec-Labrador. Am. Fern J. 72:85-87.

Weatherby, C.A. 1936. A list of varieties and forms of the ferns of
eastern North America. Am. Fern J. 26:97-98.

Wherry, E.T., Jr. 1961. The fern guide. Doubleday, Garden City,
N.Y. 318 pp.

White, D.J.; Johnson, K.L. 1980. The rare vascular plants of Mani-
toba. Syllogeus 27:1-52.

Widen, C.-J.; Britton, D.M. 1971. A chromatographic and cytological
study of Dryopteris dilatata in North America and eastern Asia.
Can.J. Bot. 49:247-258.

Wilce, J.H. 1965. Section Comp/amito of the genus Ljcopoc/mm.
Nova Hedwigia 19:1-233.

Wilce, J.H. 1972. Lycopod spores, I: General spore patterns and the
generic segregates of Ljcopodmm. Am. Fern J. 62:65-79.

Zhukova, P.G.; Petrovski, V.V. 1972. Kromosomnye chisla neko-
torykh tsvetkovykh rastenii ostrova Vrangelia, II. Bot. Zh.
(Leningr.) 57:554-567.

Addenda to References

Work on this publication was completed in 1983. Since then,
some important publications on ferns and fern allies have been
published. Among them are the following:

Alverson, E.R. 1988. Biosystematics of North American parsley
ferns, Cryptogramma (Adiantaceae). Am. J. Bot. 75:136-137
(Abstr.).

Argus, G.W.; Pryer, K.M.; White, D.J.; Keddy, C.J., eds. 1982-1987.
Atlas of the rare vascular plants of Ontario. Four parts.
National Museum of Natural Sciences, Ottawa, Ont.

Barrington, D.S. 1986. The morphology and cytology oi Polystichum
X potteri hybr. nov. (= P. acrostichoides X P. braunii). Rhodora
88:297-313.

Beitel, J. 1986. The Huperzia selago (Lycopodiaceae) complex in the
Pacific Northwest. Am. J. Bot. 73(5):733-734.

Blondeau, M.; Cayouette, J. (1987). Extensions d'aire dans la flore
vasculaire du Nouveau-Quebec. Nat. Can. (Que). 114:117-126.

416

Bouchard, A.; Barabe, D.; Bergeron, Y.; Dumais, M.; Hay, S. 1985. La
phytogeographie des plantes vasculaires rares du Quebec. Nat.
Can. (Que.) 112:283-300.

Bouchard, A.; Barabe, D.; Dumais, M.; Hay, S. 1983. The rare vascu-
lar plants of Quebec. Syllogeus 48: 1-75.

Britton, D.M. 1984a. Biosystematic studies on pteridophytes in
Canada: Progress and problems. Pages 543-560 in Grant, W.F.,
ed. Plant Biosystematics. Academic Press, Don Mills, Ont.
674 pp.

Britton, D.M. 19846. Checklist of Ontario pteridophytes. Part I:
Fern allies. Plant Press 2(4):95-99.

Britton, D.M. 1985. Checklist of Ontario pteridophytes. Part II:
Ferns. Plant Press 3(1): 14-23.

Britton, D.M.; Anderson, A.B. 1986. The ferns of Manitoulin Island:
Notes and a new record. Plant Press 4:60-61.

Britton, D.M.; Catling, P.M.; Norris, J.; Varga, S. (in press). Isoetes
engelmannii an addition to the flora of Canada. Can. Field-Nat.

Britton, D.M.; Stewart, W.G.; Cody, W.J. 1985. Cystopteris protrusa,
creeping fragile fern, an addition to the flora of Canada. Can.
Field-Nat. 99(3):380-382.

Brunton, D.F. 1986a. Status of the southern maidenhair Fern,
Adiantum capillus -veneris (Adiantaceae), in Canada. Can.
Field-Nat. 100:404-408.

Brunton, D.F. 19866. Status of the mosquito fern, Azolla mexicana
(Salviniaceae), in Canada. Can. Field-Nat. 100(3):409-413.

Bryan, F.A.; Soltis, D.E. 1987. Electrophoretic evidence for allopoly-
ploidy in the fern Polypodium uirginianum. Syst. Bot. 1 2(4): 553-
561.

Catling, P.M. 1985. Notes on the occurrence, ecology and identifica-
tion of the Massachusetts fern, Thelypteris simulata, in Ontario.
Can. Field-Nat. 99(3):300-307.

Catling, P.M.; Erskine, D.S.; MacLaren, R.B. 1985. The plants of
Prince Edward Island [D.S. Erskine] with new records,
nomenclatural changes, and corrections and deletions. Agric.
Can. Publ. 1798. 272 pp.

Cayouette, J. 1984. Additions et extensions d'aire dans la flore vascu-
laire du Nouveau-Quebec. Nat. Can. (Que.) 111:263-274.

Ceska, A. 1986. An annotated list of rare and uncommon plants of the
Victoria area. Victoria Nat. 43(5): 1-14.

Cody, W.J. 1983. Adiantum pedatum ssp. calderi, a new subspecies in
northeastern North America. Rhodora 85:93-96.

417

Cody, W.J.; Britton, DM. 1984. Polystichum lemmonii, a rock shield-
^ fern new to British Columbia. Can. Field-Nat. 98(3):375.

Cody, W.J.; Britton, D.M. 1985. Male fern, Dryopteris filix-mas, a
phytogeographically important discovery in northern
Saskatchewan. Can. Field-Nat. 99(1): 101-102.

Cody, W.J.; Schueler, F.W. 1988. A second record of the mosquito
fern, Azolla caroliniana, in Ontario. Can. Field-Nat.
102:545-546.

Cranfill, R.; Britton, D.M. 1983. Typification within the Poly podium
uirginianum complex (Polypodiaceae). Taxon 32(4):557-560.

Crist, K.C.; Farrar, D.R. 1983. Genetic load and long-distance disper-
sal in Asplenium platy neuron. Can. J. Bot. 61(6): 1809-1814.

Derrick, L.N.; Jermy, A.C.; Paul, A.M. 1987. Checklist of European
Pteridophytes. Sommerfeltia 6:1-94.

Deshaye, J.; Morisset, P. 1985. La flore vasculaire du lac a I'Eau
Claire, Nouveau-Quebec. Provancheria 18:1-52.

Fraser-Jenkins, C.R. 1986. A classification of the genus Dryopteris
(Pteridophyta: Dryopteridaceae). Bull. Br. Mus. (Nat. Hist.) Bot.
Ser. 14(3):183-218.

Gastony, G.J. 1986. Electrophoretic evidence for the origin of fern
species by unreduced spores. Am. J. Bot. 73(11): 1563-1569.

Gawler, S.C. 1983. Note on Adiantum pedatum L. ssp. calderi Cody.
Rhodora 85:389-390.

Goltz, J.P.; Britton, D.M.; Whiting, R.E. 1984. Phegopteris hexa-
gonoptera (Michx.) Fee (southern or broad beech fern) discovered
in the district of Muskoka. Plant Press 2(2):39-40.

Haufler, C.H. 1985. Pteridophyte evolutionary biology: The electro-
phoretic approach. Proc. R. Soc. Edinb. Sect. B (Biol.) 86B:315-
323.

Haufler, C.H; Soltis, D.E. 1986. Genetic evidence suggests that homo-
sporous ferns with high chromosome numbers are diploid. Proc.
Nat. Acad. Sci. USA 83:4389-4393.

Haufler, C.H.; Windham, M.D. 1988. Exploring the origin of Poly-
podium vulgare, again. Am. J. Bot. 75:139. (Abstr.).

Haufler, C.H. ; Windham, M.D.; Britton, D.M. ; Robinson, S.J. 1985.
Triploidy and its evolutionary significance in Cystopteris
protrusa. Can. J. Bot. 63(10): 1855-1863.

Hickey,J.R. 1986. /soe^es megaspore surface morphology: Nomencla-
ture, variation and systematic importance. Am. Fern J. 76:1-16.

Hinds, H.R. 1983. The rare vascular plants of New Brunswick.
Syllogeus 50:1-38.

Hinds, H.R. 1986. Flora of New Brunswick. Primrose Press, Fred-
ericton, N.B. 460 pp.

418

Holtum, R.E. 1981. The genus Oreopteris (Thelypteridaceae). Kew
Bull. 36(2):223-226.

Knobloch, I.W.; Gibby, M.; Fraser-Jenkins, C. 1984. Recent advances
in our knowledge of pteridophyte hybrids. Taxon 33(2):256-270.

Kott, L.S. 1982. A comparative study of spore germination of some
Isoetes species of northeastern North America. Can. J. Bot.
60(9):1679-1687.

Kott, L.S.; Britton, D.M. 1982. A comparative study of sporophyte
morphology of the three cytotypes oi Poly podium uirginianum in
Ontario. Can. J. Bot. 60(8):1360-1370.

Kott, L.S.; Britton, D.M. 1983. Spore morphology and taxonomy of
Isoetes in northeastern North America. Can. J. Bot.
61(12):3140-3163.

Lavoie, G. 1984. Contribution a la connaissance de la flore vasculaire
de la Moyenne-et-Basse-Cote-Nord, Quebec/Labrador.
Provancheria 17:1-149.

Lellinger, D.B. 1985. A field manual of the ferns and fern allies of the
United States and Canada. Smithsonian Institution Press,
Washington, D.C. 389 pp.

Markham, K.R.; Moore, N.A.; Given, D.R. 1983. Phytochemical reap-
praisal of taxonomic subdivisions of Lycopodium (Pteridophyta-
Lycopodiaceae) based on flavonoid glycoside distribution. N.Z. J.
Bot. 21:113-120.

McNeill, J.; Pryer, K. 1985. The status and typification oi Phegopteris
and Gymnocarpium. Taxon 34(1): 136-156.

Mills, A. 1985. Revised status of the holly fern, Polystichum lonchitis,
in Ontario. Can. Field-Nat. 99(2):252-254.

Mokry, F.; Rasbach, H.; Reichstein, T. 1986. Asplenium adulterinum
Milde ssp. presolanense subsp. nova (Aspleniaceae,
Pteridophyta). Bot. Helv. 96(1):7-18.

Moran, R.C. 1983a. Cystopteris tenuis (Michx.) Desv.: A poorly
understood species. Castanea 48:218-223.

Moran, R.C. 19836. Cystopteris X wagneri: A new naturally occur-
ring hybrid between C. X tennesseensis and C. tenuis. Castanea
48:224-229.

Morton, J.K.; Venn, J.M. 1984. The flora of Manitoulin Island and
the adjacent islands of Lake Huron, Georgian Bay and the North
Channel. Department of Biology, University of Waterloo,
Waterloo, Ont. 181pp.

Moss, E.H. 1983. Flora of Alberta. 2nd ed. Revised by Packer, J. G.
University of Toronto Press, Toronto, Ont. 687 pp.

419

Munro, D. 1988. A disjunct station q{ Asplenium ruta-muraria with
Pellaea atropurpurea and Pellaea glabella in eastern Ontario.
Am. F^ernJ. 78(4):136-138.

Packer, J.G.; Bradley, C.E. 1984. A checklist of the rare vascular
plants in Alberta. Prov. Mus. Alberta, Nat. Hist. Occ. Pap. No. 5.
112 pp.

Paris, C. 1986. A biosystematic investigation of the Adiantum
pedatum complex in eastern North America. Am. J. Bot.
73(5):738-739.

Paris, C.A; Windham, M.D. 1988. A biosystematic investigation of
the Adiantum pedatum complex in eastern North America. Syst.
Bot. 13(2):240-255.

Perkins, S.K.; Peters, G.A.; Lumpkin, T.A.; Calvert, H.E. 1985. Scan-
ning electron microscopy of perine architecture as a taxonomic
tool in the genus Azolla Lamarck. Scanning Electron Microsc.
4:1719-1734.

Prange, R.K.; Von Aderkas, P. 1985. The biological flora of Canada 6.
Matteuccia struthiopteris (L.) Todaro, Ostrich Fern. Can.
Field-Nat. 99(4):517-532.

Pryer, K.M.; Britton, D.M. 1983. Spore studies in the genus Gymno-
carpium. Can. J. Bot. 61(2):377-388.

Pryer, K.M.; Britton, D.M.; McNeill, J. 1983. A numerical analysis of
chromatographic profiles in North American taxa of the fern
genus Gymnocarpium. Can. J. Bot. 61(10):2592-2602.

Pryer, K.M.; Windham, M.D. 1988. A re-examination of Gymno-
carpium dryopteris (L.) Newman in North America. Am. J. Bot.
75:142. (Abstr.).

Soltis, D.E. 1986. Genetic evidence for diploidy in Equisetum. Am. J.
Bot. 73(5):740.

Soltis, D.E; Soltis, P.S. 1987a. Breeding system of the fern Dryopteris
expansa: Evidence for mixed mating. Am. J. Bot. 74(4):504-509.

Soltis, D.E.; Soltis, P.S. 19876. Polyploidy and breeding systems in
homosporous pteridophyta: A reevaluation. Am. Nat. 130:219-
232.

Soltis, P.S.; Soltis, D.E. 1987. Population structure and estimates of
gene flow in the homosporous fern Polystichum munitum.
Evolution 41(3):620-629.

Straley, G.B.; Taylor, R.L.; Douglas, G.W. 1985. The rare vascular
plants of British Columbia. Syllogeus 59:1-165.

Taylor, W.C; Luebke, N.T. 1986a. Germinating spores and growing
sporelings of aquatic Isoetes. Am. Fern J. 76:21-24.

420

Taylor, W.C.; Luebke, N.T. 19866. Interspecific hybridization and the
taxonomy of North American Isoetes. Am. J. Bot. 73:740-741.
(Abstr.).

Taylor, W.C; Luebke, N.T.; Smith, M.B. 1985. Speciation and hybrid-
ization in North American Isoetes. Proc. R. Soc. Edinb. Sect. B
(Biol.) 86B:259-263.

Tryon, R. 1986. The biogeography of species, with special reference to
ferns. Bot. Rev. 52(2): 117-156.

Vincent, G. 1981. P hegopter is hexagonoptera, espece rare etmena.cee.
Bull. Soc. Anim. Jard. Inst. Bot. 6(2):2-24.

von Aderkas, P. 1983. Studies of gametophytes of Matteucia
struthiopteris (ostrich fern) in nature and in culture. Can. J. Bot.
61(12):3267-3270.

von Aderkas, P. 1984. Economic history of ostrich fern, Matteuccia
struthiopteris, the edible fiddlehead. Econ. Bot. 38(1): 14-23.

Wagner, F.S. 1987. Evidence for the origin of the hybrid cliff fern,
Woodsia Xabheae (Aspleniaceae: Athyrioideae). Syst. Bot.
12(1):116-124.

Wagner, F.S. 1988a. Moonworts recently discovered in the Great
Lakes Area. Fiddlehead Forum 15(l):2-3.

Wagner, F.S. 19886. Natural hybrids between woodferns, Dryopteris,
and holly ferns, Polystichum. Am. J. Bot. 75:143-144. (Abstr.).

Wagner, W.H.; Rouleau, E. 1984. A western holly fern, Polystichum
X scopulinum in Newfoundland. Am. Fern J. 74:33-36.

Wagner, W.H.; Wagner, F.S. 1983a. Genus communities as a syste-
matic tool in the study of New World Botrychium
(Ophioglossaceae). Taxon 32(l):51-63.

Wagner, W.H.; Wagner, F.S. 19836. Two moonworts of the Rocky
Mountains: Botrychium hesperium and a new species formerly
confused with it. Am. Fern J. 73:53-62.

Wagner, W.H.; Wagner, F.S. 1984. Western Canada. Botrychium
Newsl. No. 4. University of Michigan Herbarium, Ann Arbor,
Mich. 10 pp.

Wagner, W.H.; Wagner, F.S. 1986. Three new species of moonworts
(Botrychium subg. Botrychium) endemic in western North
America. Am. Fern J. 76:33-47.

Wagner, W.H.; Wagner, F.S. 1988. Detecting Bo^rj'c/imm hybrids in
the Lake Superior Region. Mich. Bot. 27(3): 75-80.

Wagner, W.H.; Wagner, F.S.;Haufler,C.; Emerson, J. K. 1984. Anew
nothospecies of moonwort (Ophioglossaceae, Botrychium). Can.
J. Bot. 62:629-634.

421

Waterway, M.J. 1986. A reevaluation of Lycopodium porophilum and
its relationship to L. lucidulum (Lycopodiaceae). Syst. Bot.
ll(2):263-276.

Webber, J.M. 1984. Marsilea quadrifolia (four leaf waterfern) in
Ontario. Plant Press 2(3):68-70.

Werth, C.R.; Guttman, S.I.; Eshbaugh, W.H. 1985. Recurring origins
of allopolyploid species in Asplenium. Science 228:731-733.

Widen, C.-J.; Britton, D.M. 1985. Phloroglucinol derivatives of
Dryopteris tokyoensis and the missing genome in D. cristata and
D. car thus iana (Dry opteridiaceae). Ann. Bot. Fenn. 22:213-218.

Wollenweber, E.; Dietz, V.H.; Shilling, G.; Favre-Bonvin, J.; Smith,
D.M. 1985. Flavonoids from chemotypes of the goldback fern,
Pityrogramma triangularis. Phytochemistry (Oxf.) 24:965-971.

422

INDEX

Accepted names that are part of
the Canadian flora are set in
bold italics; synonyms are set in
light italics.

adder's-tongue 95

~ family 95
Adiantum 157

capillus-veneris 157

pedatum ssp. aleuticum 159

peda turn s sp . ca Ideri 162

pedatum ssp. pedatum 159

pedatum var.

rangiferinum 159

pedatum var.

subpumilum 159
Anchistea

uirginica 279
Aspidiaceae 165
Aspidium

aculeatum var.
scopulinum 194

caiifornicum 199

fdix-mas 220

fragrans 211

lonchitis 187

rigidum 207

spinulosum 217

spinulosum var.
intermedium 213
Aspidotis 140

densa 140
ASPLENIACEAE 281
Asplenium 281

cryptolepis 284

ebeneum 282

ebenoides 290

marinum 290

melanocaulon 288

platyneuron 282

rhizophyllum 290

ruta-muraria 284

ruta-muraria var.
cryptolepis 284

ruta-muraria ssp.
dolomiticum 284

trichomanes 286

trichomanes ssp.

inexpectans 286
trichomanes ssp.

quadrivalens 286
wr/de 288
Athyrium 263

acrostichoides 271
alpestre ssp.

americanum 263
alpestre var.

americanum 263
alpestre var. gaspense 263
americanum 263
angustifolium 269
distentifolium ssp.

americanum 263
distentifolium var.

americanum 263
filix-femina 266
filix-femina var.

cyclosorum 266
filix-femina var.

michauxii 266
filix-femina var.

sitchense 266
pycnocarpon 269
thelypterioides 271
>Azo//a 310

caroliniana 310
filiculoides 312
mexicana 312

BLECHNACEAE 273
Blechnum 273

doodioides 273

spicant 273

spicant var. nipponicum 273
Botrychium 97

angustisegmentum 122

boreale ssp. boreale 119

boreale ssp.

obtusilobum 119

boreale var. obtusilobum 119

crenulatum 115

dissectum 103

423

dissectum^. obliquum 103
dissectum L oneidense 106
dissectum var. obliquum 103
dissectum var.

oneidense 106
dusenii 113
lanceolatum var.

angustisegmentum 122
lanceolatum var.

lanceolatum 122
I una ha 111

lunaria war. minganense 113
matricariaefolium 116
minganense 113
multifidum 108
multifidum var

/n term edium 108
obliquum 103
106
108
108

oneidense
rugulosum

silaifolium
simplex var. simplex
simplex var.

tenebrosum 116
ternatum 108, 111
virginianum var.

europaeum 100
virginianum var.

laurentianum 100
virginianum var.

virginianum 100
Botrypus 97
bracken 135

Camptosorus 290
rhizophyllus 290

sibiricus 292
Carpogymnia 231

dryopteris 232

robertiana 236
cedar

ground- 32
Ceterach 281
Cheilanthes 138

densa 140

Zee/ 138

gracillima 140

siliquosa 140
cliff-brake 143

115

purple 144

slender 150

smooth 146
cliff fern, fragrant 211
club-moss 19

alpine 35

bog 40

bristly 24

common 21

crowfoot 32

family 19

flatbranch 30

mountain 45

savin leaf 37

shining 42

Sitka 37

stiff 24

tree 26
Cryptogramma 150

acrostichoides 150

crispa var.

acrostichoides 150

crispa var. s/'tc/i ens/s 152

densa 140

ste//er/ 150
curly-grass 130
Currania 231
Cystopteris 253

bulbifera 254

diaphana 263

dickieana 259

fragilis var. fragilis 259

fragilis var. laurentiana 262

fragilis var. mackayii 259

fragilis var. protrusa 257

laurentiana 262

montana 254

protrusa 257

reevesiana 262

tenuis 262

Dennstaedtia 133
punctilobula 133

Dtc^soAiia

pilosiuscula 133

punctilobula 133
Diphasiastrum

alpinum 35

complanatum 30

424

digitatum 32

sabinifolium 37

sitchense 37

tristachyum 32

X zeilleri 48
Diphasium

complanatum 30

flabelliforme 32

sitchense 37
Diplazium

acrostichoides 271

pycnocarpon 269

thelypterioides 271
Dryopteris 205

abbreuiata 222

X algonquinensis 230

arguta 207

assimilis 215

austriaca 215,217

azorica 213

X benedictii 231

X boon/7 230

X burgessii 230

campy loptera 217

carthusiana 217

caucasica 222

ce/sa 231

clintoniana 228

cristata 226

cristata var. clintoniana 228

dilatata 215

disjuncta 232, 234

X dowellii 230

expansa 215

filix-mas 220

fragrans 211

fragrans var.

remotiuscula 211

goldiana 224

hexagonoptera 249

intermedia 213

linnaeana 232

lonchitis 187

ludouiciana 206

maderensis 213

marginalis 222

X neo-wherryi 230

neuadensis 240

noueboracensis 243

oreades 222
oreopteris 240
phegopteris 251
X pittsfordensis 230
rigida var. arguta 207
robertiana 236
simulata 247
X slossonae 230
spinulosa 217
spinulosa var.

americana 217
spinulosa var.

intermedia 213
thelypteris var.

pubescens 245
X triploidea 230
X uliginosa 231

EQUISETACEAE 73
Equisetum 73

X arcticum 94

arvense 80

boreale 80
calderi 80
X dubium 94
X ferrissii 94
fluviatile 74
fluuiatile f. linaeanum 76
X font-queri 94
hyemale ssp. affine 84
hyemale var.

intermedium 94
hyemale war. jesupi 94
kansanum 87
laevigatum 87
limosum 74
X litorale 94
X mildeanum 94
X nelsonii 94
palustre 76
pra tense 84
prealtum 84
X rothmaleri 94
scirpoides 89
X suecicum 94
sylvaticum 82
telmateia ssp. braunii 78
X trachyodon 94

425

variegatum ssp.

alaskanum 91
variegatum ssp.

variegatum 91

variegatum war. jesupi 94
fern

Alaskan holly 203
Anderson's holly 196
bladder 253
Braun's holly 201
broad beech 249
bulblet 254
California holly 199
Christmas 184
cinnamon 126
coastal shield 207
crag holly 194
deer 273
filmy 132
flowering 124
fragile 259
goldback 155
Goldie's 224
hay-scented 133
holly 187
interrupted 126
Kruckeberg's holly 192
lace 140
lady 266

Laurentian fragile 262
licorice 296
limestone oak 236
long beech 251
maidenhair 157, 159
male 220

marginal shield 222
marsh 245
Massachusetts 247
mosquito 310
mountain 240
mountain bladder 254
Nahannioak 234
netted chain 275
New York 243
oak 231,232
ostrich 166
parsley 152
rattlesnake 100
royal 124

sensitive 168
sword 196
Venus'-hair 157
Virginian chain 279
walking 290
western oak 234
Filix

bulbifera 254
fragilis 259

grape fern 97

branching

cut-leaved 103

lance-leaved 122

least 115

leathery 108

matricary 116
ground-pine 26

equal-leaved 29

round-branched 26
Gymnocarpium 231

X achriosporum 238

continentale 234

dryopteris ssp. X
brittonianum 238

dryopteris ssp.
disjunctum 234

dryopteris ssp.
dryopteris 232

dryopteris var. pumilum 236

X heterosporum 238

X intermedium 238

jessoense ssp. parvulum 234

robertianum 236
Gymnogramma

triangularis 155

hart's-tongue 292
Hippochaete

hyemalis ssp. afftnis 84

laevigata 87

scirpoides 89

variegata 91
Homalosorus pycnocarpon 269
horsetail 73

~ family 73

field 80

giant 78

marsh 76

426

meadow 84

variegated 91

water 74

wood 82
Huperzia

selago 45

selago ssp. lucidula 42
HYMENOPHYLLACEAE 132
Hymenophyllum

wrightii 132

Indian's-dream 140
ISOETACEAE 61
Isoetes 61
acadiensis 67
beringensis 65
bolanderi 70
braunii 63
braunii f. robusta 66
canadensis 66
eatonil 65
echinospora 63
echinospora var, braunii 63
echinospora var.

maritima 65
echinospora var.

muricata 63
echinospora var. robusta 66
echinospora var. savilei 63
engelmanii 62
/ZeWii 70
gravesii 65
heterospora 68
hieroglyphica 67
howellii 69
japonica 62
lacustris 72

lacustris var. paupercula 70
macounii 65
macrospora 68
macrospora f.

hieroglyphica 67
maritima 65
melanopoda 69
melanopoda var.

californica 69
muricata 63
muricata var. braunii 63
nuttallii 69

occidentalis 70
paupercula 70
piperi 70
riparia 66
suksdorfii 69
tuckermanii 67

Japanobotrychium 97

Lastrea 231
Lepidotis

inundata 40
lip fern 138

slender 138
Lomaria

spicant 273
Lori/iseria

areolata 275
Lunathyrium

acrostichoides 271
LYCOPODIACEAE 19
Lycopodiella

inundata 40
Lycopodium 19
alpinum 35
annotinum 24
annotinum var.

acrifolium 24
annotinum var. alpestre 24
annotinum var. pungens 24
appressum 42
clavatum var.

brevispicatum 21
clavatum var. clavatum 21
clavatum var.

integerrimum 21
clavatum var.

megastachyon 21
clavatum var.

monostachyon 21

complanatum 30

complanatum var.

canadense 32

complanatum var.

dillenianum 32
complanatum var.
elongatum 30
complanatum var.
flabelliforme 32

427

complanatum var.

gartonis 30
complanatum var.

montelLii 32
complanatum var.

pseudoalpinum 30
dendroideum 26
digitatum 32
dubium 26
flabelliforme 32
X habereri 48
inundatum var.

bigelovii 42
inundatum var.

inundatum 40
X issleri 48
lagopus 24
lucidulum 42
lucidulum var.

occidentale 45
obscurum var.

dendroideum 26
obscurum var.

isophyllum 29
obscurum var. obscurum 26
porophilum 45
sabinifolium 37
sabinifolium var.

sitchense 37
selago var. appressum 45
selago ssp. miyoshianum 46
selago ssp. patens 46
selago ssp. selago 45
sitchense 37
tristachyum 32
tristachyum var.

laurentianum 35

Lygodium palmatum 130
MARSILEACEAE 307
Marsilea 307

mucronata 307

quadrifolia 308

vest/ta 307
yWatteucc/a 166

struthiopteris var.
pensylvanica 166
Mecodium 132

wrightii 132

moonwort 97, 111
mountain-parsley 152

Nephrolepis 1 87

Onoclea 168
sensibilis 1 68
sensibilis f.

obtusilobata 168

struthiopteris var.
pensylvanica 166
OPHIOGLOSSACEAE 95
Ophioglossum 95

pusillum 97

pychnostichum 97

reticulatum 97

uulgatum var. alaskanum 95

vulgatum var.

pseudopodum 95

vulgatum var.

pycnostichum 95
Oreopteris limbosperma 240
Osmunda 124

cinnamomea 126

cinnamomea f.
frondosa 129

cinnnamomea ssp.
asiatica 129

claytoniana 126

regal is f. anomala 124

regalis ssp. vestita 126

X rag^ii 129

regalis var. spectabilis 124
OSMUNDACEAE 124
Osmundopteris 97

Pellaea 143

atropurpurea 144

atropurpurea var. bushii 146
atropurpurea var.

simplex 148
densa 140

glabella var. glabella 146
glabella var. nana 146
glabella var.

occidentalis 146
glabella var. simplex 148
occidentalis 146
pumila 146

428

suksdorfiana 148
pepperwort

hairy 307
Phegopteris 247

connect! I is 251

hexagonoptera 249

polypodioides 251
Phyllitis 292

fernaldiana 292

japonica ssp. americana 292

scolopendrium var.
americanum 292
pine

ground- 26

running- 32
Pityrogramma 155

triangularis 155
POLYPODIACEAE 295
Polypodium 295

amorphum 299

australe 295

fauriei 295

glycyrrhiza 296

hesperium 302

interjectum 295

montense 299

scouleri 296

uinlandicum 305

virginianum 302

uirginianum var.
americanum 305

vulgare ssp.

columbianum 302

vulgare ssp. occidentale 296

vulgare ssp.

virginianum 302

vulgare var.

columbianum 302

vulgare var. occidentale 296
polypody

rock 302
Polystichum 182

acrostichoides 184

acrostichoides f.
incisum 184
' alaskense 203

andersonii 196

braunii 201

braunii ssp. alaskense 203

braunii ssp. andersonii 196
braunii ssp. purshii 201
californicum 199
dudleyi 199
X hagenahii 205
imbricans 1 89
kruckebergii 192
lemmonii 189
lonchitis 187
mohrioides 189
mohrioides var.

lemmonii 189
mohrioides var.

scopulinum 194
munitum 196

munitum var. imbricans 189
scopulinum 194
setigerum 203
PTERIDACEAE 133
Pteridium 135
aquilinum var.

champlainense 135
aquilinum var.

lanuginosum 137
aquilinum var.

latiusculum 135
aquilinum var.

pubescens 137
arachnoideum 137
esculentum 137
latiusculum 135

nodulosa 166
pensylvanica 166

quillwort 61
~ family 61

SALVINIACEAE 310
Sceptridium 103
SCHIZAEACEAE 130
Schizaea 130

pus/7/a 130
Scolopendrium

vulgare 292
scouring-rush 84

dwarf 89

smooth 87
SELAGINELLACEAE 49

429

Selaginella 49
apoda 50
densa 54
densa var. scopulorum 54

eclipes 50

montane ns is 52

oregana 52

rupestris 57

scopulorum 54

selaginoides 49

sibirica 57

wallacei 52
spikemoss

~ family 49
spleenwort

ebony 282

green 288

maidenhair 286

narrow-leaved 269

silvery 271
Struthiopteris

spicant 273

Thelypteris 239

cristata 226

cristata var. clintoniana 228

dryopteris 232

filix-mas 220

fragrans 211

goldiana 224

hexagonoptera 249

limbosperma 240

marginalis 222

nevadensis 240

noveboracensis 243

oregana 240

oreopteris 240

palustris var. pubescens 245

phegopteris 251

simulata 247

spinulosa 217

thelypterioides 245
Wagneriopteris 247
wall-rue 284

water-clover 307,308
wood fern 205

Appalachian Mountain 217

Clinton's 228

crested 226

eastern spreading 217

evergreen 213

northern 215

spinulose 217
Woods/a 170

X abbeae 182

a/p/na 173

appalachiana 178

6eZ/ii 173

confusa 182

glabella 171

X gracilis 182

ilvensis 173

X maxonii 182

obtusa 180

oregana 176

oregana var.

cathcartiana 178

oregana var. lyallii 178

oregana var.

squammosa 182

pusillavar. cathcartiana 178

scopulina 178

X tryonis 182
woodsia 170

blunt-lobed 180

northern 173

Oregon 176

Rocky Mountain 178

rusty 173

smooth 171
Woodwardia 275

angustifolia 275

areolata 275

chamissoi 277

fimbriata 277

paradoxa 277

radicans var. americana 277

virginica 279

430

Canada

Internet Archive Audio

Featured

Top

Images

Featured

Top

Software

Featured

Top

Books

Featured

Top

Video

Featured

Top

Mobile Apps

Browser Extensions

Archive-It Subscription

Save Page Now

Full text of "Ferns and fern allies of Canada"

See other formats